Masarykova univerzita
Přírodovědecká fakulta
Ľubomír VIDLIČKA
Vybrané aspekty z etológie a taxonómie švábov
(Blattaria)
Habilitačná práca
v odbore Zoológia
Bratislava 2015
Bibliografická identifikácia
Meno a priezvisko autora: Ľubomír Vidlička
Názov dizertačnej práce:
Vybrané aspekty z etológie a taxonómie švábov (Blattaria)
Názov dizertačnej práce po anglicky:
Selected aspects of ethology and taxonomy of cockroaches (Blattaria)
Odbor: Zoológia
Rok obhajoby: 2016
Kľúčové slová v slovenčine:
etológia, taxonómia, paleontológia, Blattaria, Nocticolidae, Blaberidae, Blattellidae
Kľúčové slová v angličtine:
ethology, taxonomy, paleontology, Blattaria, Nocticolidae, Blaberidae, Blattellidae
Prehlásenie
Prehlasujem, že som prácu napísal sám s použitím patričných literárnych zdrojov, ktoré sú
v práci všetky riadne citované.
V Bratislave, 20.11.2015 Ľubomír Vidlička
Poďakovanie
Na tomto mieste si dovoľujem poďakovať sa všetkým, ktorí ma na mojej dlhej ceste za
poznaním švábov sprevádzali alebo správnym spôsobom ovplyvnili. V prvom rade je to
RNDr. Milan Kozánek, CSc., u ktorého som ako mladý ašpirant začínal a po prvýkrát som
prišiel do styku s masovo chovanými švábmi (Nauphoeta cinerea) na vedecké účely. Od
tohto okamihu som už šváby nikdy neopustil i keď môj záujem o ne sa vyvíjal rôznymi
smermi.
Ďalej musím na tomto mieste poďakovať všetkým bývalým kolegom z Ústavu
experimentálnej fytopatológie a entomológie SAV v Ivanke pri Dunaji a taktiež súčasným
kolegom z Ústavu zoológie SAV v Bratislave za ich trpezlivosť s mojimi obľúbenými
chovancami, ktorí sa viac-menej pravidelne vydávali v nočných hodinách na prieskum
ústavu a ráno spôsobovali u nežnejšieho pohlavia zdesenie.
Za dlhoročnú a plodnú spoluprácu ďakujem aj RNDr. Petrovi Vršanskému, PhD. z
Geologického ústavu SAV, ktorý sa síce prednostne venuje fosílnym švábom, ale našiel som
v ňom spriaznenú dušu aj pri živých predstaviteľoch tohto radu hmyzu.
Poďakovanie patrí aj Prof. Ing. Jaroslavovi Holušovi, Ph.D. (Česká zemědělská univerzita
v Praze) a Doc. RNDr. Petrovi Kočárkovi, Ph.D. (Přírodovědecká fakulta Ostravská
univerzita v Ostravě) za príkladnú spoluprácu pri tvorbe monografie o ortopteroidnom
hmyze Českej a Slovenskej republiky, kde zapadli aj „moje“ šváby.
V neposlednom rade patrí moje poďakovanie mojej manželke, že mala pre moju záľubu
pochopenie pri mojich neskorých príchodoch z práce, pri mimoeurópskych expedíciách, ako
aj počas dovoleniek venovaných zberom švábov.
3
Abstrakt
Predložená práca pojednáva o niektorých aspektoch etológie a taxonómie zástupcov radu
švábov (Blattaria). Je založená na súbore 17 vedeckých článkov (väčšina z nich je zahrnutá
v databáze WOS) a dvoch monografiách, ktoré majú vzťah k riešenej problematike.
Obsah práce je rozdelený do 4 sekcií. Prvá sekcia „Etológia švábov“ pojednáva o
sexuálnom správaní švábov druhu Nauphoeta cinerea počas dvorenia a párenia. Párenie je
veľmi zložitý proces pozostávajúci zo 4 fáz. V každej fáze prebiehajú iné deje: i) vytvorenie
pevného genitálneho spojenia, ii) kompletizácia a presun spermatofóru v tele samčeka, iii)
prenos spermatofóru do burzy copulatrix samičky, iv) ukončenie párenia a rozpojenie
genitálií. Jednotlivé fázy sú presne riadené nervovou sústavou, sprostredkované aj zmenami
koncentrácie histamínu; počas dvorenia sú riadené hlavne mozgom (nadhltanové ganglium)
a vo fáze kopulácie najmä posledným (šiestym) abdominálnym gangliom.
Druhá sekcia „Rozšírenie a taxonómia švábov v Európe a na Slovensku“ sa skladá z
troch častí. Prvá časť pojednáva o šváboch z rodu Ectobius, ktorý je v Európe zastúpený 35
druhmi. Tie sú však rozšírené veľmi nerovnomerne. Väčšinu tvoria endemitné druhy s
veľmi malými areálmi rozšírenia, hlavne v južnej Európe. V podmienkach strednej Európy
a Slovenska majú širšie zastúpenie iba druhy E. lapponicus, E. sylvestris a E. erythronotus.
V posledných rokoch môžeme pozorovať prenikanie druhu E. vittiventris z južného
Švajčiarska do oblasti strednej Európy a aj na územie Slovenska. Druh má podobné nároky
ako naše natívne druhy. Ťažko si hľadá v prírode voľnú niku a tak často preniká do ľudských
príbytkov. Druhá časť tejto sekcie sa zaoberá skupinou maculata z rodu Phyllodromica.
Rod Phyllodromica je najväčší európsky rod s množstvom endemitných druhov v centrálnej
Európe, vrátane územia Slovenska, Maďarska a Rumunska.
Posledná časť v tejto sekcii hovorí o šváboch zo skupiny megerlei z rodu Phyllodromica.
Túto skupinu tvoria zatiaľ iba tri druhy, z ktorých iba Ph. megerlei má širšie rozšírenie.
Ďalšie druhy z tejto skupiny sa dajú očakávať hlavne v štátoch východného Stredomoria
(Chorvátsko, Albánsko, Macedónsko, Čierna Hora, Srbsko) a západne od Čierneho mora
(Bulharsko, Rumunsko).
Tretia sekcia „Šváby juhovýchodnej Ázie a Južnej Ameriky“ je zameraná na vybrané
endemitné rody žijúce v oblastiach s najväčšou druhovou bohatosťou švábov na svete. V
časti o JV Ázii sú detailne spracované informácie o šváboch z rodov Caeparia,
Chorisoserrata a Spelaeoblatta. Južnú Ameriku reprezentuje rod Macrophyllodromia, ktorý
v súčasnosti združuje 12 druhov. Táto časť poukazuje na potrebu urýchleného výskumu v
oblastiach tropických pralesov, ktoré sú veľmi rýchlo devastované a miznú.
Posledná sekcia „Fosílne šváby“ je pohľadom do života švábov za posledných 300
miliónov rokov. Paleontológovia prinášajú takmer denne nové informácie o fosílnych
druhoch, ktoré menia náš pohľad na evolúciu tejto skupiny. Výrazné zmeny nastali aj v
paleoetológii švábov. Pozornosť v poslednej dobe vzbudili hlavne dva objavy: šváby ako
čističi po dinosauroch a objav švába z „európskeho“ rodu Ectobius v amerických eocénnych
usadeninách.
4
Abstract
The present habilitation thesis deals with some aspects of ethology and taxonomy focusing
on representatives of the order cockroaches (Blattaria). The thesis is based on the collection
of 17 scientific papers (most of them included in the Web of Sciences) and 2 monographs,
which are related to the thesis main topic.
The content of thesis is divided into four sections. The first section “Ethology of
cockroaches” deals with a sexual behaviour of cockroach species Nauphoeta cinerea during
courtship and copulation. Mating is a very complex process comprising 4 phases. In each
phase, different actions are involved: i) forming stable genital connection, ii) assembling
and movement of spermatophore in the male body, iii) transferring of spermatophore into
female bursa copulatrix and iv) finishing of copulation and disconnection of genitalia.
All phases are exactly controlled by nervous system mediated by the changes of histamine
concentration; during courtship is controlled mainly by brain (supraesophageal ganglion)
and in the phase of copulation controlled mainly by the last (sixth) abdominal ganglion.
The second section “Distribution and taxonomy of cockroaches in Europe and Slovakia”
has three parts. First part deals with cockroach genus Ectobius that is represented by 35
species. These species are unequal distributed. The most of them are endemic species with
small distribution areas mainly in South Europe. In central Europe including Slovakia, the
species E. lapponicus, E. sylvestris and E. erythronotus are wide distributed. In the last
years, we observe infiltration of species E. vittiventris from South Switzerland into the area
of central Europe including Slovakia. This species has the same food request as our native
species. It often occupies human habitats due to non-existing free niche. The second part
deals with maculata-group of genus Phyllodromica. Phyllodromica is the most dominant
European genus with numbers of endemic species in central Europe including Slovakia,
Hungary and Romania.
The last part deals with cockroaches from megerlei-group of Phyllodromica. This group
consists of three species of which only Ph. megerlei is wide distributed. Other species from
this group are expected to be distributed mainly in the states of east Mediterranean (Croatia,
Albania, Macedonia, Monte Negro and Serbia) and westward from Black sea (Bulgaria and
Romania).
The third section “The cockroaches of south-east Asia and South America” focuses on
selected endemic genera living in the area with the highest species richness worldwide. In
the part of south-east Asia, detailed information about cockroaches from the genera
Caeparia, Chorisoserrata and Spelaeoblatta can be found. South American cockroaches are
represented by genus Macrophyllodromia consisting of 12 species. This section highlights
the urgent needs of taxonomic research in the area of tropical forest which are nowadays
massively devastated.
The last section “Fossil cockroaches” is view to the life of cockroaches over the past 300
million years. The paleontologists bring daily new information about fossil species that are
changing are view on the evolution of this group. Significant changes are also described in
paleoethology of cockroaches. In last days, more attention is paid on two observations:
cockroaches as a cleaner after dinosaurs and cockroaches from “European” genus Ectobius
found in American Eocene sediments
5
Obsah
1. Úvod ................................................................................................................................. 7
2. Štruktúra práce a jej zameranie ........................................................................................ 8
2.1. Tematické okruhy .......................................................................................................... 8
3. Etológia švábov ................................................................................................................ 9
3.1. Dvorenie a párenie ........................................................................................................ 9
3.2. Prehľad príspevkov autora k poznaniu etológie švábov ............................................. 13
4. Rozšírenie a taxonómia švábov v Európe a na Slovensku ............................................ 14
4.1. Rod Ectobius – rozšírenie a biológia .......................................................................... 14
4.2. Rod Phyllodromica – skupina druhov maculata ........................................................ 16
4.3. Rod Phyllodromica – skupina druhov megerlei ......................................................... 20
4.4. Prehľad príspevkov autora k poznaniu švábov na Slovensku a v Európe ................. 22
5. Šváby juhovýchodnej Ázie a Južnej Ameriky ............................................................... 24
5.1. Šváby z rodu Caeparia Stål, 1877 (Blaberidae: Panesthiinae) .................................. 24
5.2. Šváby z rodu Chorisoserra (Blattellidae: Pseudophyllodrominae) ............................ 27
5.3. Šváby z rodu Spelaeoblatta Bolívar, 1897 (Noticolidae) ........................................... 29
5.4. Šváby z rodu Macrophyllodromia Saussure & Zehnter, 1893 (Blattellidae) ............. 31
5.5. Prehľad príspevkov autora k poznaniu švábov juhovýchodnej Ázie
a Južnej Ameriky ......................................................................................................... 34
6. Fosílne šváby (Blaberidae, Blattellidae, Ectobiidae) ..................................................... 35
6.1. Prehľad príspevkov autora k poznaniu fosílnych švábov
(Blaberidae, Blattellidae, Ectobiidae) ......................................................................... 37
7. Použitá literatúra ............................................................................................................ 38
Prehľad príloh..................................................................................................................... 48
6
1. Úvod
Predložená habilitačná práca sa zaoberá etológiou, taxonómiou a fylogenézou radu švábov
(Blattaria), ktorý tvorí spolu s modlivkami (Mantodea), koníkmi, kobylkami, svrčkami
(Orthoptera), ucholakmi (Dermaptera), pakobylkami (Phasmatodea), svrčkovcami
(Grylloblattodea) a nedávno opísaným radom Mantophasmatodea (KLASS et al. 2002)
skupinu ortopteroidného hmyzu.
Šváby ako rad sú tak u nás, ako aj vo svete, pomerne zanedbávaná skupina, čo je zrejme
dané malým počtom druhov, ktoré tento rad zahŕňa. Na druhej strane však sú šváby
ideálnymi laboratórnymi živočíchmi. Rýchlo a bezproblémovo sa množia, na malom
priestore je ich možné dochovať stovky. Preto sa stali laboratórnou „myšou“ pri výskume
bezstavovcov. Keď som koncom 80-tych rokov minulého storočia začínal svoju kariéru, bol
to práve výskum fyziológie bezstavovcov, zhodou okolností švábov. Príprava pokusov si
vyžadovala dôkladné poznanie ich pohlavného správania (pozri časť 3). Táto prvá
skúsenosť spojená s výskumom švábov predurčila moju celoživotnú dráhu.
Od etológie tropických laboratórnych druhov sa časom záujem presunul aj na naše
autochtónne druhy švábov. V tej dobe bolo zo Slovenska známych len 5 druhov (MAŘAN &
ČEJCHAN 1977) a z celej Európy asi 80 druhov švábov. O ich rozšírení či sezónnej
dynamike sa vedelo veľmi málo a nakoniec sa aj existujúce determinačné kľúče ukázali ako
nespoľahlivé. Vedomosti o šváboch bolo treba získavať takmer od začiatku. Na Slovensku
sa faunistikou či taxonómiou švábov nezaoberal žiadny špecialista a v Českej republike sa
im v tej dobe venovali iba dvaja odborníci, obaja však iba popri koníkoch a kobylkách.
Štúdium začalo spracovaním muzeálnych zbierok v slovenských a českých múzeách
(VIDLIČKA & MAJZLAN 1992, VIDLIČKA & HOLUŠA 1999). Dôkladné zoznámenie sa s
literatúrou a všetkými dostupnými dátami vyústilo v spolupráci s vtedy tiež mladými
kolegami z Čiech do publikovania nového zoznamu druhov (KOČÁREK et al 1999) ako aj
bibliografie ortopteroidného hmyzu (HOLUŠA et al. 1999). Okruh záujmu sa postupne
rozšíril aj na územie Maďarska a Rumunska. Spracované boli šváby celej Karpatskej kotliny
(VIDLIČKA & SZIRÁKY 1997), Švajčiarska (VIDLIČKA & REZBÁNYAI-RESER 2005),
čiastočne Chorvátska (VIDLIČKA & OZIMEC 2011). Popri tom boli objavené a opísané aj pre
vedu nové druhy z Maďarska, Rumunska, Bulharska (pozri časť 4). Nazhromaždené údaje
vyústili do monografického spracovania Fauny švábov Slovenska (VIDLIČKA 2001) a
vďaka možnosti ponúknutej kolegami z Českej republiky aj ilustrovaného kľúča pre Českú
Republika a Slovensko (KOČÁREK et al. 2005).
Záujem o taxonómiu švábov zákonite viedol k rozšíreniu záujmového územia na teritóriá
bohatšie na faunu švábov – juhovýchodnú Áziu a Južnú Ameriku (pozri časť 5). V týchto
dvoch oblastiach žijú tri štvrtiny všetkých známych druhov švábov. Na rozdiel od Európy
sú tu početné malé endemické rody. Ich identifikácia je pomerne náročná, odborníkov na
šváby je na svete iba pár. Zároveň sú to oblasti postihnuté rýchle vzrastajúcim počtom
obyvateľstva a v ruka v ruke s tým aj devastáciou prírody nebývalých rozmerov.
Úzka spolupráca s paleontológom, ktorý sa veľmi úspešne etabloval pri výskume fosílnych
švábov, vyústila okrem taxonomickej problematiky aj do výskumu ich paleoetológie (pozri
časť 6), čím sa kruh záujmov akoby uzavrel a znovu sa vrátil tam kde začal.
7
2. Štruktúra práce a jej zameranie
Habilitačná práca pozostáva z úvodnej časti, v ktorej sú stručne načrtnuté štyri záujmové
okruhy, ktoré autor rieši alebo v minulosti riešil a zo súboru 17 vedeckých článkov
publikovaných v impaktovaných (12) ako aj neimpaktovaných recenzovaných (5)
časopisoch a dvoch monografií, ktoré majú vzťah k problematike vybraných tematických
okruhov. Pripojené publikované práce sú iba výberom prác autora, keďže autor sa venuje
okrem švábov (Blattaria) aj sieťokrídlovcom (Neuroptera) a príležitostne aj iným radom
hmyzu (Insecta). Pripojené práce sú zároveň aj reprezentatívnym výberom z prác autora
venovaných výskumu švábov z rokov 1993-2015.
Úvodná časť práce slúžia na zoznámenie sa s tým, čo bolo v danej problematike známe
do doby autorovho príspevku, s výsledkami autora a prípadne aj s tým, čo sa v tejto oblasti
zistilo odvtedy dodnes.
2.1. Tematické okruhy práce
a) Etológia švábov – dvorenie a párenie
b) Rozšírenie a taxonómia švábov v strednej Európe a na Slovensku
b1) Rod Ectobius – rozšírenie a biológia
b2) Rod Phyllodromica – skupina druhov maculata
b3) Rod Phyllodromica – skupina druhov megerlei
c) Šváby v juhovýchodnej Ázii a Južnej Amerike
c1) Šváby z rodu Caeparia Stål, 1877 (Blaberidae: Panesthiinae)
c2) Šváby z rodu Chorisoserra (Blattellidae: Pseudophyllodromidae)
c3) Šváby z rodu Spelaeoblatta Bolívar, 1897 (Noticolidae)
c4) Šváby z rodu Macrophyllodromia Saussure & Zehnter, 1893 (Blattellidae)
d) Fosílne šváby (Blaberidae, Blattellidae, Ectobiidae)
8
3. Etológia švábov
3.1. Dvorenie a párenie
Záujem o detailný priebeh správania sa švábov počas dvorenia a párenia možno datovať do
50-tych rokov minulého storočia. Prelomovou bola práca dvojice amerických švábológov
Louisa M. Rotha a Edwina R. Willisa o reprodukčnej biológii švábov (ROTH & WILLIS 1954).
Predkopulačné správanie švábov vykazuje u celej skupiny určité známky uniformity
(ROTH & WILLIS 1954, ROTH & DATEO 1966; ROTH & BARTH 1967, ROTH 1969, SIMON
& BARTH 1977a,b). Priebeh párenia sa však
môže v určitých detailoch líšiť v závislosti
od umiestnenia tergálnych žliaz samčeka. V
tergálnych žľazách sa tvorí sexuálny
(afrodiziakálny) feromón. Podľa spôsobu
akým samček láka samičku a zahajuje
kopuláciu rozlišujeme tri typy páriaceho
správania (A, B a C) (SRENG 1993). Párenie je najlepšie preskúmané na laboratórne
chovaných druhoch švábov. Jedným z najčastejšie chovaných druhov je cirkumtropicky
rozšírený druh Nauphoeta cinerea (Blaberidae: Oxyhaloinae). Z viacerých príčin práve on
plní úlohu „laboratórnej myši“ pri výskume
bezstavovcov. Spôsobom dvorenia a párenia je šváb
Nauphoeta cinerea typickým predstaviteľom
základného typu A.
Keď sa k sebe priblížia samček a samička švába
N. cinerea (obr. 1), ktorí sú pripravení na párenie,
začne samček vylučovať prchavé látky krátkeho
dosahu a na povrchu samičky sa objavia kontaktné
feromóny, ktoré umožňujú druhovú identifikáciu,
rozpoznanie pohlavia a zároveň, v prípade stretnutia
správneho druhu a páru, uľahčujú aj koordináciu dvorenia samčeka samičke. Využívaniu
pohlavných feromónov švábmi sa dlhodobo venoval Coby Schal (SCHAL 1982, SCHAL &
BELL 1984, 1985, SCHAL et al. 1990, 1991) a tému prednedávnom komplexne spracoval
(GEMENO & SCHAL 2004). Feromóny neslúžia len pri výbere partnera a na spustenie
samčieho dvorenia, ale zároveň môžu regulovať aj viaceré fyziologické procesy. U N.
cinerea môžu samčie feromóny ovplyvniť dĺžku
života samičky, počet potomkov, pomer ich pohlavia
a rýchlosť ich vývinu v plodovom vaku (MOORE et al.
2001, 2002, 2003).
Pri krátkom tykadlovom kontakte overí samček
fyziologickú pripravenosť samičky na párenie (obr.
2). V prípade samičkinej fertility zahajuje samček
fázu dvorenia. Dvihne tegminy a krídla do takmer
kolmej polohy k telu, čím odokryje tergity na svojom
chrbte (obr. 3) a zároveň umožní lepšie pôsobenie
9
Ľubomír Vidlička Habilitačná práca
Obr. 1. Stretnutie jedincov rôzneho pohlavia.
Obr. 2. Tykadlový kontakt.
Obr. 3. Dvorenie samčeka samičke.
feromónu produkovaného tergálnymi žľazami. Následne sa ku samičke natočí tak, aby
mohla zozadu vyliezť na jeho chrbát. Ak samička nereaguje na takúto ponuku snaží sa
samček podsunúť bruško pod jej hlavu čo
najtesnejšie.
Fertilná samička zvyčajne reaguje na ponuku
samčeka postupným vyliezaním na jeho chrbát
(obr. 4). Súčasne ústnymi orgánmi „olizuje“
produkt samčích tergálnych žliaz (obr. 5).
Vyliezanie zvyčajne končí dotykom samičkinej
hlavy o dvihnuté krídla samčeka.
Doba, ktorú strávi samička olizovaním
sekrétov stačí samčekovi na to, aby vystrčil
genitálny hák (ľavá faloméra), zasunul ho
pomedzi samičie genitálie, a tak vytvoril dostatočne pevné spojenie so samičkou (obr. 6).
Hákom spoľahlivo pripojený samček sa začne pohybovať dopredu a skladať krídla a
tegminy na bruško, čím prinúti samičku zliezť bokom z jeho chrbta (obr. 7) a otočiť sa o 180
stupňov (obr. 8). Tým sa koniec samčieho
bruška pretočí a následne ho samček musí
dostať do prirodzenej polohy bez toho, aby
bolo prerušené ukotvenie pomocou háku. Až
po tomto pomerne náročnom úkone sa obaja
v páre dostávajú do správnej kopulačne
pozície a môže začať vlastná kopulácia (obr.
9). Zaujatú polohu nemení pár až do
skončenia kopulácie (VIDLIČKA 2001, BELL
et al. 2007).
Proces kopulácie môže trvať rôzne dlho. Dĺžka kopulácie je samozrejme medzi druhmi
variabilná, ale mení sa aj v rámci druhu. Jej dĺžku ovplyvňuje viacero faktorov. V prvom
rade je to vek samčeka, ale vplyv má aj doba uplynutá od jeho posledného párenie a jeho
sociálny status. Reálna dĺžka párenia sa dá zistiť iba odstránením vplyvu všetkých týchto
faktorov. To znamená, že pozorovania z prírody sú ťažko interpretovateľné. Najdlhšie
párenia pozorované v prírodných podmienkach sa týkajú juhoamerického druhu Xestoblatta
hamata (Blattellidae), kde párenie v dažďovom pralese trvalo až 5 hodín (SCHAL & BELL
1982) a austrálskeho druhu Polyzosteria limbata (Blattidae, Polyzosterinae), kde párenie
prebiehalo za denného svetla a páry niekedy zostávali spojené viac ako 24 hodín
(MACKERRAS 1965).
Zhodou okolností sa Nauphoeta cinerea pári
najkratšie z doteraz študovaných druhov. Za
dodržania štandardných podmienok (použitie 14
dní starých panenských samčekov chovaných už od
posledného nymfálneho instaru v kolektíve nýmf
rovnakého pohlavia pri teplote 251°C) je doba
jeho párenia prekvapivo stabilná – 12,5 minúty
(N=100, SE=0,9 – VIDLIČKA & HUČKOVÁ 1993).
10
Obr. 4. Vyliezanie samičky na chrbát
dvoriaceho samčeka.
Obr. 5. Olizovanie samčích sekrétov.
Obr. 6. Prichytenie samičky hákom.
Ľubomír Vidlička Habilitačná práca
Pri pozmenených podmienkach môže prvé párenie trvať aj 17 minút (ROTH 1964). Druhé
párenie s nedostatočným odstupom môže trvať podstatne dlhšie (1008 min.) a pri treťom
párení v rade až 1412 minúty (ROTH 1964).
Pozornosť bola zvyčajne venovaná dĺžke
párenia a jeho úspešnosti, prípadne vlastnostiam
potomstva, ale už menej vlastnému priebehu
kopulácie. Telo samčekov počas kopulácie pulzuje
v nepravidelných intervaloch. U Nauphoeta cinerea
pulzuje telo samčeka prvé 4 minúty kopulácie
približne 15x za minútu, ďalšie 4 minúty frekvencia
klesá pod 10 pulzov za minútu, v 9. minúte sa počet
pulzov prudko zväčšuje až na 20 za minútu, potom
frekvencia pulzov bruška samčeka prudko klesá
pod 10 pulzov za minútu (obr. 10) a v poslednej
minúte párenia sa fakticky takmer úplne zastavuje
(VIDLIČKA & HUČKOVÁ 1993).
Experimentálne prerušovanie kopulácie v
pravidelných intervaloch (po 4, 6, 8, 10 minútach od zaujatia páriacej polohy) prispelo k
pochopeniu dejov v priebehu kopulácie. Vo väčšine prípadov mal samček po 4 minútach
párenia penis vysunutý, podobne to bolo
aj po 8 minútach párenia. V oboch
prípadoch došlo v deviatej minúte k
vysunutiu spermatoforu a jeho
upevneniu k podkladu. Pri prerušení
kopulácie po 10. minúte bol penis tiež
vysunutý a zostal tak až do 13. minúty
(podobne to bolo aj v predchádzajúcich
dvoch prípadoch), a potom bol
zatiahnutý. Na základe zistených zmien
v rýchlosti pulzácie a tiež na základe výsledkov z prerušovania kopulácie bol priebeh
kopulácie rozdelený do štyroch fáz: i. tvorba genitálneho kontaktu, vysunutie penisu (1.-4.
minúta), ii. tvorba, kompletizácia a presun spermatoforu v tele samčeka (5.-8. minúta), iii.
prenos spermatoforu do kopulačnej komôrky (bursa copulatrix) samičky (9. minúta), iv.
prerušenie genitálneho kontaktu, zasunutie penisu a uvoľnenie upevňovacieho háku z tela
samičky (10.-13. minúta) (VIDLIČKA & HUČKOVÁ 1993).
Spermie po skončení párenia nemigrujú zo samčieho spermatoforu umiestneného v
kopulačnej komôrke do samičej spermatéky okamžite. Spermatofor obsahuje nepohyblivé,
stočené spermie, ktoré sa vystierajú a
začínajú sa aktívne pohybovať až asi
po 2 hodinách (VIDLIČKA &
HUČKOVÁ 1993). Spermie sú
aktivované výlučkami spermatéky
(KHALIFA 1950).
11
Obr. 7. Bočné zliezanie samičky zo samčeka.
Obr. 8. Natáčanie samičky do páriacej polohy.
Obr. 9. Kopulácia v lineárnej polohe.
Ľubomír Vidlička Habilitačná práca
Keďže niektoré výskumy ukázali, že úroveň histamínu ako neurotransmiteru v hmyzích
CNS sa mení v odozve na rôzne stresové podnety (KOZÁNEK et al. 1985, TAKÁČ et al.
1990), zvažovala sa aj jeho úloha v reprodukcii švábov Nauphoeta cinerea (HUČKOVÁ et al.
1992).
Priebeh koncentračných zmien histamínu v mozgu (supraoesofageálnom gangliu)
Nauphoeta cinerea počas párenia bol u samčekov a samičiek veľmi podobný (obr. 10). V
čase dvorenia, keď samička vylezie na bruško samčeka a na začiatku kopulácie sa úroveň
histamínu zvýšila len mierne. Tento nárast bol trochu prudší v mozgu samičiek oproti
mozgu samčekov. V 5. minúte kopulácie sa koncentrácie histamínu u oboch pohlaví výrazne
zvýšili a dosiahli asi 3-krát vyššie hodnoty, než boli pozorované u kontrolných nepárených
skupín. V 10. minúte kopulácie sa znížila hladina histamínu a bola približne na rovnakej
úrovni ako na začiatku kopulácie, avšak u samčekov bola signifikantne vyššia v porovnaní
s kontrolnou skupinou. Po 2 hodinách bola hladina histamínu u oboch pohlaví takmer
rovnaká ako v kontrolných skupinách alebo na začiatku kopulácie.
V 6. abdominálnom gangliu boli zmeny koncentrácie histamínu výrazne odlišné u
samčekov a samičiek (obr. 10). Na
začiatku kopulácie bola síce u
oboch pohlaví takmer rovnaká ako
u kontrolnej skupiny, ale potom (v
5. a 10. minúte párenia) bol u
samčekov zaznamenaný prudký
nárast koncentrácie. U samičiek
zostala koncentrácia histamínu
porovnateľná s kontrolnou
skupinou. Dve hodiny po ukončení
kopulácie boli hodnoty histamínu
nižšie ako u kontrolných skupín. To
naznačuje úlohu 6. brušného
ganglia a histamínu práve pri
dotvorení spermatoforu v brušku
samčeka a pri jeho presune a
upevnení v kopulačnej komôrke
samičky (KOZÁNEK et al. 1992,
HUČKOVÁ et al. 1994). HINTZEPODUFAL
& VETTER (1996)
poukázali na funkciu juvenilného
hormónu pri párení švába Blaptica
dubia. Významnú úlohu hral hlavne
v prvej fáze dvorenia – tykadlový
kontakt.
Ako už bolo naznačené, samčekovi po párení trvá príprava nového spermatoforu (hlavne
jeho naplnenie spermiami) značnú dobu. Preto trvajú opakované párenia výrazne dlhšie.
Samčekovia švába N. cinerea, ktorí sa opätovne párili skôr ako za 24 hodín, mali
obmedzený počet spermií a produkovali menej potomkov než panenskí samčekovia a tiež
12
Obr. 10. Priebeh kontrakcií bruška u samčeka a zmeny
hladiny histamínu v mozgu (plná čiara)
a v 6. abdominálnom gangliu (bodkovaná čiara)
u samčekov (modrá čiara) a samičiek (červená čiara)
Nauphoeta cinerea počas 1. kopulácie.
Ľubomír Vidlička Habilitačná práca
samčekovia, ktorí mali päťdňovú dobu na zotavenie medzi dvoma kopuláciami (MONTROSE
et al. 2004). Ukázalo sa, že samičky Nauphoeta cinerea sú schopné rozlíšiť samčekov, ktorí
sa ešte nepárili od takých, ktorí sa už párili. Ak majú možnosť, vyberajú si panenských
samčekov radšej ako samčekov s nedostatkom spermií (HARRIS & MOORE 2005).
U druhu Schultesia nitor (Blaberidae: Zetoborinae) bolo pozorované, že samčekovia sa
stávajú pohlavne receptívnymi až pár dní po imaginálnom zvlečení. Pred týmto časom nie
sú schopní párenia. Samičky sa stávajú fertilnými takmer okamžite po imaginálnom
zvlečení, skôr ako sú plne zafarbené a sklerotizované (tenerálne samičky). U tohto druhu je
larválny vývoj u samčekov kratší ako u samičiek, čiže tento druh je protandrický. Samičky
sa správajú monandricky (pária sa iba raz), zatiaľ čo samčekovia sú polygynní (ak majú
príležitosť pária sa viackrát). Samičky u tohto druhu produkujú iba jednu znášku a krátko
po vyliahnutí potomstva hynú. Aj u tohto druhu sa ukázalo, že ak majú samičky možnosť
voľby, uprednostňujú panenských samčekov pred už skúsenejšími samčekmi (MONCEAU &
VAN BAAREN 2012).
Samičky ovoviviparného druhu Nauphoeta cinerea vynosia v priemere 6 ooték. Po
oplodnení strácajú fertilitu. Keď porodia potomstvo chránia nymfy počas prvého instaru
(MOORE & MOORE 2001). Až potom nadobúdajú opäť fertilitu a sú ochotné sa znova páriť
(ROTH 1964).
3.2. Prehľad príspevkov autora k poznaniu etológie švábov
(hnedou farbou sú uvedené príspevky, ktoré sú súčasťou habilitačnej práce)
KOZÁNEK, M., TAKÁČ, P., VIDLIČKA, Ľ. 1990. Concentration changes of some
monoamines and steroids during courtship and copulation of cockroach Nauphoeta
cinerea. Invertebrate Reproduction & Development 18(1-2): 120.
TAKÁČ, P., KOZÁNEK, M., VIDLIČKA, Ľ. 1990. Circadian changes of some vertebrate-type
hormones in cockroach Nauphoeta cinerea. Invertebrate Reproduction & Development
18(1-2): 130.
HUČKOVÁ, A., KOZÁNEK, M., VIDLIČKA, Ľ., TAKÁČ, P. 1992. Histamine distribution in the
nervous system of the cockroach Nauphoeta cinerea (Blattodea, Panchloridae) and its
changes during development, pp. 129-134. In: Advances in regulation of insect
reproduction. Institute of Entomology Czech Academy Sciences, České Budějovice.
ISBN 80-901250-0-X
VIDLIČKA, Ľ., HUČKOVÁ, A. 1993. Mating of the cockroach Nauphoeta cinerea (Blattodea:
Blaberidae): I. Copulatory behaviour. Entomological Problems 24(2): 69-73. (Príloha č. 1)
HUČKOVÁ, A., VIDLIČKA, Ľ., KOZÁNEK, M. 1994. Mating of the cockroach Nauphoeta
cinerea (Blattodea: Blaberidae): II. Histamine changes during courtship and copulation.
Biologia 49(5): 691-695. (Príloha č. 2)
VIDLIČKA, Ľ. 2001. Blattaria – šváby; Mantodea – modlivky (Insecta: Orthopteroidea).
Veda, Bratislava, 169 pp. (Fauna Slovenska) ISBN 80-224-0640-6 (Príloha č. 18)
13
Ľubomír Vidlička Habilitačná práca
4. Rozšírenie a taxonómia švábov v Európe a na Slovensku
Šváby (Blattaria) sú prevažne tropickou skupinou zahŕňajúcou okolo 5000 druhov. Z tohto
množstva sa v Európe vyskytuje len okolo 120 druhov čo znamená, že v Európe je
najmenšia druhová pestrosť švábov zo všetkých svetadielov (samozrejme okrem
Antarktídy).
Napriek pomerne malému množstvu druhov je švábologická literatúra zaoberajúca sa
územím Európy nepomerne bohatšia ako na iných svetadieloch. Viaceré Európske druhy
majú dosť široké rozšírenie a sú značne variabilné, čo spôsobilo, že boli opísané aj
niekoľkokrát. Majú množstvo synoným, niektoré druhy aj viac ako desať. Situácia sa preto
stala krajne neprehľadnou a paradoxne, určenie niektorých európskych druhov (hlavne
balkánskych) je takmer nemožné.
Z územia Európy je známych z natívnych druhov iba 6 rodov švábov – Ectobius,
Phyllodromica, Loboptera, Planuncus, Capraiellus, Polyphaga a Hemelytroblatta. Z rodu
Polyphaga zasahuje do Európy iba 1 druh – P. aegyptiaca. Rod Hemelytroblatta je
zastúpený 5 druhmi zasahujúcimi okrajovo do juhovýchodnej Európy. Celý rod sa
vyskytuje prevažne v severnej Afrike a na Blízkom Východe. Podobne rod Loboptera, ktorý
zahŕňa 32 druhov, je z Európy známy iba 6 druhmi, z ktorých iba jeden je širšie rozšírený v
Stredomorí – Loboptera decipiens. Ostatných 5 druhov je známych iba zo Španielska a boli
opísané pomerne nedávno (BOHN 1990) odčlenením od druhu Loboptera decipiens. Zvyšok
druhov je endemických na Kanárskych ostrovoch, Azorách, Madeire a v Maroku. Výnimku
predstavujú dva druhy vyskytujúce sa v Kamerune (L. loboptera Princis, 1962) a v Saudskej
Arábii (L. isolata Grandcolas, 1994).
Rody Ectobius a Phyllodromica majú európske až eurázijské (palearktické) rozšírenie.
Viaceré druhy zasahujú aj do Afriky. V rode Ectobius evidujeme v súčasnosti 67 druhov, z
ktorých 35 je rozšírených v Európe, pri rode Phyllodromica je to 60 z 99 známych druhov.
Rod Capraiellus vznikol odčlenením troch druhov z rodu Ectobius do samostatného
podrodu (HARZ 1976), ktorý bol prednedávnom povýšený na samostatný rod (BOHN et al.
2013). Všetky tri druhy rodu Capraiellus sa vyskytujú v Európe. Rod Planuncus bol tiež
zriadený iba pred dvomi rokmi (BOHN et al. 2013) a boli sem preradené 4 druhy z rodu
Ectobius a 9 druhov z rodu Phyllodromica. Až 11 z nich žije v Európe. Oba tieto prípady
naznačujú dosť neprehľadnú situáciu v rozlišovaní rodov Ectobius a Phyllodromica, kde
bola donedávna hlavným kritériom dĺžka tegmín.
4.1. Rod Ectobius – rozšírenie a biológia
Zo spomínaných 35 európskych druhov rodu Ectobius žije 27
na Apeninskom polostrove a na blízkych ostrovoch. Mnoho z
nich je tam endemických, hlavne na Sicílii, Sardínii a ďalších
menších ostrovoch. Hlbšie do kontinentálnej Európy preniká
iba pár druhov. Na území Slovenska boli dlho známe iba 3
druhy – E. sylvestris (obr. 11), E. lapponicus a E. erythronotus.
Status a tiež rozšírenie štvrtého druhu E. lucidus sú nejasné
(BOHN 1989).
14
Ľubomír Vidlička Habilitačná práca
Obr. 11. Ectobius sylvestris
(VIDLIČKA 2001).
Vďaka otepľovaniu klímy pribudol na našom území približne od roku 2010 nový druh
švába pôvodom z južnej (podalpskej) časti Švajčiarska – Ectobius vittiventris. Keďže je
teplomilnejší ako naše druhy a má podobné teritoriálne nároky, udomácniť sa mu darí iba
vo väčších mestách, zvyčajne v trávnikoch, parkoch a záhradách pri domoch. Na Slovensku
bol zatiaľ zistený iba v Bratislave, jeho prenikanie do ľudských príbytkov bolo iba
sporadické (VIDLIČKA 2014).
V roku 2015 však začal do
príbytkov prenikať masovo,
hlavne na jeseň. Svojim
správaním (na rozdiel od
synantropného druhu Blattella
germanica sa neskrýva cez deň
pred svetlom) vyvolal v
postihnutých domácnostiach
zdesenie a u dezinsekčných
firiem zmätok (nepubl. údaje).
Šírenie tohto druhu do roku
2014 je zobrazené na obrázku 12.
Výskyt a rozšírenie
švábikov z rodu Ectobius ako
aj blízko príbuzného rodu Phyllodromica v Poľsku spracoval BAZYLUK (1977). V tom
istom roku bol spracovaný aj zoznam Československých druhov švábov (MAŘAN &
ČEJCHAN 1977). V tej dobe však neboli zo Slovenska ešte známe žiadne endemické druhy.
Na Slovensku spracovali zoznam druhov švábov ako aj ich rozšírenie VIDLIČKA a
MAJZLAN (1992). Následne bola spracovaná bibliografia literatúry o šváboch v Českej
Republike a na Slovensku (HOLUŠA et al. 1999) a na jej základe aj zoznam druhov pre
Čechy, Moravu a Slovensko (KOČÁREK et al. 1999). Priebežne boli spracované doplnkové
informácie o výskyte švábov v rôznych oblastiach Slovenska: Muránska planina (VIDLIČKA
1997), Devínska Kobyla (VIDLIČKA 2005), ostrov Kopáč (VIDLIČKA 2007), Jurský Šúr
(FEDOR et al. 2010),
Východné Slovensko
(TOMKO et al. 2013).
Detailné poznanie
rozšírenia švábov v celej
Karpatskej kotline (zahrnuté
boli aj celé Karpaty) (obr.
13) je založené na štúdiu
bohatých muzeálnych
materiálov z viacerých
múzeí. Pre každý druh bola
vypracovaná aj distribučná
mapa (VIDLIČKA &
SZIRÁKI 1997).
15
Ľubomír Vidlička Habilitačná práca
Obr. 12. Mapa pôvodného rozšírenia švábika Ectobius vittiventris
a smery jeho šírenia po roku 1985.
Podľa VIDLIČKU (2014).
Obr. 13. Rozšírenie druhu Ectobius lapponicus v Karpatskej kotline.
Podľa VIDLIČKU & SZIRÁKIHO (1997) – upravené.
Neskôr bolo podobne spracovaných takmer 800 exemplárov švábov z 25 rokov
trvajúcich zberov (1975-2000) zo Švajčiarska. Zistených bolo 10 druhov z rodu Ectobius
(VIDLIČKA & REZBANYAI-RESER 2005). Počas spracovávania bol zistený aj výskyt nového
druhu švába (Ectobius supramontanus Bohn, 2004). Preto bola práca publikovaná až po
zverejnení jeho opisu (BOHN 2004).
Začaté bolo aj pomerne náročné spracovávanie balkánskej fauny švábov. Existuje
zoznam švábov bývalej Juhoslávie (US & MATVEJEV 1967), tento však treba inovovať pre
jednotlivé nástupnícke štáty a hlavne podľa najnovších zmien v taxonómii. Spracovanie
balkánskeho komplexu druhov Ectobius erythronotus (KARAMAN & KARAMAN 1987) do
danej problematiky väčšiu prehľadnosť neprinieslo, skôr naopak. Najnovší výskum sa začal
v oblasti pohoria Biokovo (Chorvátsko), kde bolo v rozmedzí 900-1650 m n.m. zistených
predbežne 5 druhov švábov (VIDLIČKA & OZIMEC 2011). Ďalší balkánsky materiál však
zatiaľ čaká na komplexnú revíziu.
O biológii švábov z rodu Ectobius sa zatiaľ toho veľa nevie. Ectobius sylvestris sa v
južných Alpách dá zastihnúť až vo výškach okolo 2400 m (BEIER 1974). U nás bol
pozorovaný v Belianskych Tatrách do 1700 m (CHLÁDEK 1986). Druhy tohto rodu sa živia
hlavne organickým detritom. Cez deň môžeme samčekov často zastihnúť na kvetoch rastlín,
najmä na sedmokráskach (Bellis sp.) a omanoch (Inula sp.), kde sa kŕmia peľom rastlín
(VIDLIČKA 2001). Nymfy a samičky sú aktívne hlavne v noci (DREISIG 1971).
V prírode (na Slovensku) sa švábiky hôrne (Ectobius sylvestris) vyskytujú od apríla do
októbra (VIDLIČKA 1993, 2012) s vrcholom výskytu imág v polovici apríla až začiatkom mája
(VIDLIČKA 1993, 2013). V Nemecku v jelšovom lese bola u toho istého druhu zaznamenaná
najväčšia migrácia po kmeňoch stromov v druhej polovici mája (GHARAGJEDAGHI 1994).
Imága švábika tmavoštíteho (Ectobius lapponicus) na východe Čiech dosiahli maximum v
druhej polovici júna, respektíve až v júli (HOLUŠA & KOČÁREK 2000). Vrcholy výskytu
imág korešpondujú aj s obdobiami znášok ooték (VIDLIČKA 1993) a zároveň aj s výskytom
parazitoida Brachygaster minutus (VIDLIČKA 1998). Nymfy sa liahnu počas júla a zimujú
ako štvrtý nymfálny instar (BROWN 1973a, b, VIDLIČKA 2012), zriedkavo ako druhý alebo
tretí instar (BROWN 1983). Zimovanie prebieha najčastejšie v zemi, v trsoch tráv, ale aj v
opustených hniezdach vtákov (VIDLIČKA 1999).
4.2. Rod Phyllodromica – skupina druhov maculata
Je zaujímavé, že tak ako v južnej Európe majú sklony
vytvárať množstvo endemitov zástupcovia z rodu Ectobius,
tak v strednej Európe, kde je rod Ectobius zastúpený veľmi
chudobne, plnia túto funkciu druhy rodu Phyllodromica,
hlavne zo skupiny maculata. Typový druh Phyllodromica
maculata (obr. 14) má veľmi široké rozšírenie od Nemecka po
JZ Ukrajinu a od stredného Poľska až po severné Chorvátsko
a Rumunsko. Jeho sfarbenie je veľmi variabilné a preto
vznikli v minulosti problémy pri delení tohto druhu na variety,
poddruhy resp. druhy. Novo opisované druhy majú spravidla
malé areály rozšírenia a sú endemitmi na daných územiach.
16
Ľubomír Vidlička Habilitačná práca
Obr. 14. Phyllodromica maculata
(VIDLIČKA 2001).
Viaceré endemické druhy boli opísané priamo zo Slovenska alebo susedného Maďarska.
Prvými boli Phyllodromica harzi Chládek, 1977 zo Slovenského krasu a Phyllodromica
chladeki Harz, 1977 z Muránskej planiny (CHLÁDEK & HARZ 1977). Druh Phyllodromica
hungarica Vidlička, 1993 bol pôvodne opísaný z pohoria Bükk na severe Maďarska
(VIDLIČKA 1993), ale neskôr bol zistený aj na južnom Slovensku (VIDLIČKA 2001, BOHN
& CHLÁDEK 2011, TOMKO et al. 2013). Ďalší nový druh Phyllodromica dobsiki (dobšiki)
Chládek, 1996 bol opísaný z Muránskej planiny, z rezervácie Suché doly pri Tisovci
(CHLÁDEK 1996). Neskôr bol tento druh synonymizovaný s druhom Ph. hungarica (BOHN
& CHLÁDEK 2011).
Zo Slovenska bol opísaný aj poddruh Phyllodromica maculata marani (mařani) Chládek
& Harz, 1980 s výskytom na strednom a východnom Slovensku, v Poľsku a Rumunsku. Od
nominálneho poddruhu sa odlišoval morfológiou ootéky (CHLÁDEK & HARZ 1980).
Zobrazená ootéka pochádzala zo Slovenského krasu. Ako vyzerajú imága švába sa v práci
neuvádza, ale autor tvrdil, že sú zhodné s exemplármi z Brašova (Rumunsko) spomínanými
v práci RAMMEHO (1951) ako poddruh Hololampra maculata schäfferi. Aj keď Ramme
vyslovene hovorí o poddruhu, Chládek ich hodnotí ako melanickú formu schaefferi s tým,
že medzi formami existujú prechody. To bol dôvod vytvorenia nového mena pre tento
poddruh. Zo Slovenska známe jedince daného poddruhu a literatúru k nim sa vzťahujúcu
neuvádza. VIDLIČKA a SZIRÁKI (1997) poukázali na to, že tento poddruh je iba synonymom
Ph. maculata schaefferi známeho zo Slovenského krasu už 150 rokov (FRIVALDSZKY 1867)
a z Maďarska 120 rokov (CHYZER 1897). Následne CHLÁDEK (1998) povýšil nový poddruh
bez udania dôvodu na nový druh (a omylom uviedol opačné poradie autorov ako v pôvodnej
práci – Harz et Chládek, 1980). Druh bol opätovne synonymizovaný s Ph. maculata
schaefferi (VIDLIČKA 2001). Ak by bol uznaný jeho druhový status, meno by malo byť
Phyllodromica schaefferi. Status tohto druhu (resp. poddruhu) je značne komplikovaný a
zaslúži si krátke vysvetlenie.
Druh bol (ako Blatta qvarta) pôvodne „opísaný“ (zobrazený na ilustrácii) v diele
prírodovedca Jacoba Chritiana Schaeffera (*1718-†1790) pojednávajúcom o hmyze z
okolia nemeckého mesta Regensburg (SCHAEFFER 1769). Pôvodný názov bol uznaný za
neplatný z dôvodu, že meno bolo vlastne poradovým číslom a šváb samotný bol iba
zobrazený na obrázku. Na Schaefferovu počesť dostal od Johanna Augusta Ephraima
Goezeho (*1731-†1793) druh nové meno Blatta Schaefferi. GOEZE (1778) sa odvolal na
Schaefferov obrázok a stručne druh charakterizoval. O typovej lokalite sa nezmienil. Neskôr
Johann Friedrich Gmelin (*1748-†1804) publikoval podobný stručný opis v lipskom
vydaní Linného „Systema Naturae“ (GMELIN 1789), ktoré sa stalo veľmi populárne. V práci
neuviedol, že použil meno vytvorené Goezem. Vlastný problém vznikol, keď Johann
Christian Daniel von Schreber (*1739-†1810) opísal z Nemecka druh Blatta maculata
(teraz Ph. maculata) (SCHREBER 1781). Druh bol veľmi podobný druhu Blatta Schaefferi
len mal svetlejšie tegminy s menšími čiernymi škvrnami. Gottlieb August Wilhelm
Herrich-Schäffer (*1799-†1874) zaradil Gmelinom použité meno medzi synonymá druhu
Blatta maculata, čo bolo z hľadiska princípu priority opodstatnené (HERRICH-SCHÄFFER
1840). Leopold Heinrich Fischer (*1817-†1886) vo svojom diele „Orthoptera Europaea“
s odvolaním sa na Gmelinom použité meno „Blatta Schaefferi“ zaradil tmavé exempláre ako
varietu druhu Blatta maculata (FISCHER 1853). Keďže sa odvolal na Gmelinov opis bolo to
17
Ľubomír Vidlička Habilitačná práca
z hľadiska časovej priority mien opäť logické riešenie. Pridal obsiahly opis, avšak nevyriešil
rozšírenie, keďže spomína iba jedince z Nemecka a Rakúska. Keď uhorský entomológ Ján
Frivaldský (*1822 Rajec-†1895 Budapešť) našiel v Zádielskej doline tmavé exempláre z
rodu Phyllodromica zhodné s Gmelinovým aj Fischerovým opisom, začal pre ne používať
meno Aphlebia maculata var. Schäfferi (FRIVALDSKY 1867). Rod Aphlebia Brunner von
Wattenwyl, 1865 je synonymom staršieho rodového mena Phyllodromica Fieber, 1853.
Varietu na poddruh povýšil Richard Ebner, keď použil pre tmavé jedince z Brašova
(Rumunsko) názov Hololampra maculata schaefferi (EBNER 1930). Rod Hololampra
Saussure, 1864 je tiež synonymom rodu Phyllodromica. Obaja (Frivaldský aj Ebner) použili
názov „schaefferi“ v podradenom postavení k menu „maculata“, čo znamená, že ho
považovali za mladší. Vychádzali teda z Gmelinovho opisu, ktorý je mladší ako Schäfferov
opis druhu Blatta maculata. Pôvodný Goezeho starší opis nepoznali. Nanešťastie sa
ukázalo, že takáto tmavá varieta resp. poddruh druhu Ph. maculata nežije ani v okolí
Regensburgu ani v Nemecku vôbec. Z Nemecka sú známe iba klasicky svetlo sfarbené
jedince zodpovedajúce opisu druhu Phyllodromica maculata.
Lokalita Regensburg v Nemecku evidentne nemohla byť typovou lokalitou tmavých
jedincov zodpovedajúcich Goezeho, Gmelinovmu či Fischerovmu opisu. Prvou
publikovanou lokalitou zodpovedajúcou spomínaným opisom bola Zádielská dolina na
Slovensku (FRIVALDSKÝ 1867). Preto bola Zádielská dolina dodatočne určená ako terra
typica pre Phyllodromica maculata schaefferi v zmysle Gmelinovho opisu (VIDLIČKA
2001). Profesor Bohn z Nemecka chcel riešiť vzniknutú situáciu synonymizovaním názvu
„schaefferi“ s druhom Ph. maculata (pers. comm.). Po upozornení, že meno „shaefferi“ je
staršie ako „maculata“ bolo meno Blatta schaefferi klasifikované ako „nomen dubium“ a
ponechaný bol iba názov Ph. maculata (BOHN & CHLÁDEK 2011). Avšak okolnosť, že druh
Ph. schaefferi sa nevyskytuje na lokalite, ktorá mu bola pôvodne určená, nie je dôvod na
zrušenie mena. Okrem toho sa Goeze ani Gmelin na lokalitu neodvolávali, takže
Regensburg nebol oficiálnou typovou lokalitou. Napriek tomu zostalo pre tmavé jedince z
okolia Zádielu meno Ph. marani, pre časť ďalších tmavo sfarbených jedincov zo stredného
Slovenska a severného Maďarska bol vytvorený druh Phyllodromica latipennis Bohn &
Chládek, 2011 (obr. 15) a pre tmavé exempláre zo Sedmohradska bol tiež vytvorený nový
druh Phyllodromica variabilis Bohn, 2011 in BOHN & CHLÁDEK 2011). Tento stav je
oficiálne doteraz platný, i
keď praktické rozlišovanie
medzi novo opísanými
druhmi je vďaka ich
variabilite dosť náročné a
neisté. Vzhľadovo
podobná Phyllodromica
hungarica je od ostatných
spomenutých druhov
ľahko odlíšiteľná kvôli
výrazne zahnutým krídlam
(VIDLIČKA 1993).
18
Ľubomír Vidlička Habilitačná práca
Obr. 15. Rozšírenie švábov zo skupiny maculata rodu Phyllodromica na
Slovensku, v Maďarsku a Rakúsku.
V práci BOHNA a CHLÁDKA (2011) je podrobne spracované rozšírenie všetkých druhov
zo skupiny maculata v oblastiach ich výskytu. V práci chýba spracovanie rozšírenia Ph.
maculata na Morave a v Čechách. Viaceré údaje sa nachádzajú v Chládkových starších i
novších prácach (CHLÁDEK 1965,
1998, 2006).
Podrobne bolo spracované
rozšírenie tohto druhu v Českej
republike v práci VIDLIČKU a
HOLUŠU (1999). Druh sa
pravdepodobne vyskytuje na
vhodných lokalitách po celej
republike (KOČÁREK et al. 2005),
ani jedným smerom tu nedosahuje
hranicu rozšírenia (obr. 16). Je
známy aj zo všetkých okolitých
štátov.
O rozšírení druhu Phyllodromica maculata v Poľsku pojednávajú staršie práce
BAZYLUKA (1976, 1977). V Poľsku, podobne ako aj v susednom Nemecku, prebieha
severná hranica rozšírenia druhu približne na 53° severnej zemepisnej šírky (obr. 17).
Prvé zmienky o šváboch zo
skupiny maculata z Rumunska
publikoval prírodovedec a
entomológ slovenského pôvodu
Oto Herman (*1835-†1914).
Zmienil sa o výskyte druhu
Phyllodromica maculata z okolia
Kluža (Cluj, SZ Rumunsko), ale
či sa jednalo o tmavú formu
neuvádza (HERMAN 1871). Prvý
kto konkrétne písal o tmavej
forme (Aphlebia maculata var.
Schäfferi) bol entomológ Karl
Brunner von Wattenwyl (*1823†1914)
(BRUNNER v. W. 1882).
Určiť v súčasnosti o ktorý
konkrétny druh išlo sa naisto
nedá, keďže v súčasnosti je z
Rumunska známych viacej
druhov zo skupiny maculata.
Okrem už spomenutého Bohnom opísaného druhu Phyllodromica variabilis, rozšíreného
v centrálnej časti Rumunska, tu žijú aj ďalšie dva endemické druhy zo skupiny maculata.
Prvým je už dávnejšie opísaná Phyllodromica transylvanica Vidlička, 1994 s pomerne
19
Ľubomír Vidlička Habilitačná práca
Obr. 16. Rozšírenie švába Phyllodromica maculata
v Českej Republike.
Podľa VIDLIČKU a HOLUŠU (1999) - doplnené.
Obr. 17. Rozšírenie švába Phyllodromica maculata v Poľsku.
Podľa BAZYLUKA (1977) – upravené.
širokým rozšírením v Sedmohradskej
oblasti (Transylvánia). Momentálne
je areál jeho výskytu ohraničený
Karpatmi (VIDLIČKA 1994, BOHN &
CHLÁDEK 2011). Tento druh nebol
zatiaľ nájdený v Maďarsku, ale jeho
výskyt na východe Maďarska je
pravdepodobný, podobne ako aj na
severe Srbska. Ďalším druhom je
Phyllodromica halterisignata Bohn,
2011 (in BOHN & CHLÁDEK 2011).
Zatiaľ je známy iba z jednej lokality
na SZ Rumunska (obr. 18).
4.3. Rod Phyllodromica – skupina druhov megerlei
Okrem skupiny druhov maculata z rodu Phyllodromica, je
veľmi dobre definovanou aj skupina druhov megerlei.
Skupina je charakteristická veľmi komplikovaným sieťovaným
vzorom na tegminách a zároveň silne skrátenými tegminami
u samičiek. Typovým druhom je Phyllodromica megerlei
(obr. 19). Ani opis tohto druhu sa nezaobišiel bez problémov.
Druh pôvodne opísal Toussaint de Charpentier (*1779†1847).
Charpentier bol nemecký geológ. V rokoch 1811-
1828 zastával funkciu Sliezskeho vrchného banského
odborníka vo Wroclawe (Wratislaviae). Amatérsky sa na
pomerne slušnej úrovni venoval aj entomológii. V roku 1825
mu vo Wroclawe vyšlo dielo „Horae entomologicae“ o
európskom hmyze, v ktorom najväčšiu pozornosť venoval ortopteroidnému hmyzu
(Dermaptera, Blattaria, Mantodea, Orthoptera). V práci spomína 14 druhov švábov (vtedy
všetky v rode Blatta), z ktorých dva sú novo opísané (CHARPENTIER 1825). Prvým je Blatta
limbata z Lusitanie (oblasť zaberajúca časť Portugalska a Španielska), ktorý bol neskôr
synonymizovaný s druhom Loboptera decipiens (Germar, 1817). Tento druh sa dokonca v
práci tiež spomína (ako Blatta decipiens). Druhým opísaným druhom bola Blatta punctata
z Rakúska. Opis druhu je v Charpentierovej práci prisúdený rakúskemu entomológovi
Johannovi Carlovi Megerle von Mühlfeld (*1775-†1840). Až o 28 rokov si český
entomológ (nemeckého pôvodu) Franz Xaver Fieber (*1807-†1872) všimol, že meno
Blatta punctata použil Johann Friedrich Gustav von Eschscholtz (*1793-†1831) už v
roku 1822 pri opise iného druhu švába (terajšia Diploptera punctata) (ESCHSCHOLTZ 1822).
Preto na počesť objaviteľa premenoval druh z Rakúska na Phyllodromica megerlei. Keďže
Fieber zaradil tento druh do ním novo opísaného rodu, stal sa druh zároveň typovým
druhom rodu Phyllodromica (FIEBER 1853).
20
Ľubomír Vidlička Habilitačná práca
Obr. 18. Rozšírenie švábov zo skupiny maculata rodu
Phyllodromica v Rumunsku.
Podľa BOHNA & CHLÁDKA (2011) – upravené.
Obr. 19. Phyllodromica megerlei
(VIDLIČKA 2001).
Phyllodromica megerlei je
síce druh s pomerne širokým
areálom výskytu (obr. 20), ale na
lokalitách sa vyskytuje vždy iba v
neveľkých počtoch. Svojim
výzorom (husto škvrnitými
tegminami) je dosť odlišný od
väčšiny druhov, ktoré boli neskôr
do tohto rodu zaradené.
Rakúsky ortopterológ Richard
Ebner (*1885-†1961), ktorý sa
venoval ortopteroidnému hmyzu
Malej Ázie si všimol, že
exempláre Ph. megerlei vyskytujúce sa v Turecku sú odlišné od jedincov z Európy, takže
ich spomenul ako varietu, ale meno jej nepridelil (EBNER 1919). Významný ruský
entomológ Grigorij Jakovlevič Bej-Bienko (*1903-†1971) v rámci edície „Fauna SSSR“
spracoval skupinu švábov (Blattaria). Aj on si uvedomil odlišnosť ázijských (Tureckých a
Sýrskych) exemplárov. Na základe žlto-oranžovej škvrny na štíte (na rozdiel od čiernej
škvrny u typickej formy) opísal nový poddruh Phyllodromica megerlei asiatica (BEYBIENKO
1950). Pri opise mal k dispozícii 7 exemplárov zo 6 lokalít, ale všetko to boli
samičky, takže opísal iba samičku. Keď nemecký ortopterológ Willy Adolf Theodor
Ramme (*1887-†1953) spracovával ortopteroidný hmyz južnej Európy a Malej Ázie nemal
ešte k dispozícii Bej-Bienkovu „Faunu“. Mal však k dispozícii samčeka švába z Adany
(Turecko) s oranžovou škvrnou na štíte a tak ho opísal ako Hololampra punctata f.
erythronota (RAMME 1951). Táto forma bola o pár rokov synonymizovaná s BejBienkovým
poddruhom (KARABAG 1958). Až takmer po 50-tich rokoch bol na základe
odlišností v tvare tergálnej žľazy, stilusu a genitálneho háku povýšený tento poddruh na
samostatný druh Phyllodromica asiatica (VIDLIČKA & MAJZLAN 1997). Zároveň bol na
základe viacerých exemplárov pochádzajúcich z Bulharska, z lokality Vlas neďaleko
Nesebaru opísaný druh podobný Ph. megerlei, len výrazne menší. Druh dostal meno
Phyllodromica pulcherrima (VIDLIČKA & MAJZLAN 1997). Neskôr bol tento druh zberaný
na viacerých lokalitách v Bulharsku (G. Hristov, pers. comm.). V súčasnosti sú tieto tri
druhy – Ph. megerlei, Ph.
asiatica a Ph. pulcherrima (obr.
21), jedinými zástupcami
skupiny druhov megerlei v rode
Phyllodromica. Rozšírenie Ph.
megerlei bolo spracované v
rámci Karpatskej kotliny
(VIDLIČKA & SZIRÁKI 1997).
Najnovšie údaje o výskyte na
východnom Slovensku sú
uvedené v práci TOMKU et al.
(2013).
Ľubomír Vidlička Habilitačná práca
21
Obr. 20. Rozšírenie švábov zo skupiny megerlei rodu
Phyllodromica v Európe a Ázii.
Podľa VIDLIČKU & MAJZLANA (1997) – upravené.
Obr. 21. Phyllodromica pulcherrima ♂ a ♀
(VIDLIČKA & MAJZLAN 1997).
4.4. Prehľad príspevkov autora k poznaniu švábov na Slovensku a v Európe
(hnedou farbou sú uvedené príspevky, ktoré sú súčasťou habilitačnej práce)
VIDLIČKA, Ľ., MAJZLAN, O. 1992. Survey and geographical distribution of native
cockroach species (Blattaria: Blattellidae: Ectobiinae) in Slovakia. Entomologické
problémy 23: 21-29.
VIDLIČKA, Ľ. 1993. Seasonal dynamycs of vertical migration and distribution of cockroach
Ectobius sylvestris (Blattaria: Blattellidae: Ectobiinae) in oak forest. Biologia 48(2): 163-
166. (Príloha č. 3)
VIDLIČKA, Ľ. 1993. Phyllodromica hungarica sp.nov., a new cockroach species from
Hungary (Insecta: Blattodea: Blattellidae: Ectobiinae). Entomological Problems 24(1):
63-68. (Príloha č. 4)
VIDLIČKA, Ľ. 1994. Phyllodromica transylvanica sp. nov., a new cockroach species from
Romania and key of the maculata-group of Phyllodromica in central Europe.
Entomological Problems 25(2): 55-62. (Príloha č. 5)
VIDLIČKA, Ľ. 1997. Výskum švábov na Muránskej planine = Research on native cockroach
species in the Muránska planina Mts. Výskum a ochrana prírody Muránskej planiny
1997: 89-92.
VIDLIČKA, Ľ., MAJZLAN, O. 1997. Revision of the megerlei - group of the cockroach genus
Phyllodromica Fieber (Blattaria: Blattellidae, Ectobiinae). Entomologica Scandinavica
28: 163-173. (Príloha č. 6)
VIDLIČKA, Ľ., SZIRÁKI, Gy. 1997. The native cockroaches (Blattaria) in the Carpathian
Basin. Folia Entomologica Hungarica 58: 187-220.
VIDLIČKA, Ľ. 1998. Seasonal flight pattern of the evaniid wasp Brachygaster minutus
(Hymenoptera: Evaniidae) - parasitoid of cockroach egg cases. Entomofauna Carpathica
10: 65-69.
HOLUŠA, J., KOČÁREK, P., VIDLIČKA, Ľ. 1999. Bibliography to the fauna of Blattaria,
Mantodea, Orthoptera and Dermaptera of the Czech and Slovak Republics. Articulata
14(2): 145-176.
KOČÁREK, P., HOLUŠA, J., VIDLIČKA, Ľ. 1999. Check-list of Blattaria, Mantodea,
Orthoptera and Dermaptera of the Czech and Slovak Republics. Articulata 14(2): 177-184.
VIDLIČKA, Ľ. 1999. Šváby (Blattaria) v hniezdach vtákov. Folia faunistica Slovaca 4: 41-43.
VIDLIČKA, Ľ., HOLUŠA, J. 1999. Rusec plamatý Phyllodromica maculata maculata
(Schreber, 1781) (Blattodea: Ectobiidae: Ectobiinae) na Moravě a v Čechách
[Phyllodromica maculata maculata (Schreber, 1781) (Blattodea: Ectobiidae: Ectobiinae)
in Moravia and Bohemia]. Sborník Přírodovědného klubu v Uherském Hradišti 4: 107-
114.
Ľubomír Vidlička Habilitačná práca
22
VIDLIČKA, Ľ. 2001. Blattaria – šváby; Mantodea – modlivky: (Insecta: Orthopteroidea).
Veda SAV, Bratislava, 169 pp. (Fauna Slovenska) ISBN 80-224-0640-6 (Príloha č. 18)
KOČÁREK, P., HOLUŠA, J., VIDLIČKA, Ľ. 2005: Blattaria, Mantodea, Orthoptera &
Dermaptera of the Czech and Slovak Republics. Illustrated key 3. Blattaria, Mantodea,
Orthoptera & Dermaptera České a Slovenské republiky. Ilustrovaný klíč 3. Kabourek,
Zlín, 349 pp. ISBN 80-86447-05-7 (Príloha č. 19)
VIDLIČKA, Ľ. 2005. Šváby (Blattaria) a modlivky (Mantodea), pp. 62-63. In: Fauna
Devínskej Kobyly. Asociácia priemyslu a ochrany prírody, Bratislava. ISBN 80-968217-
1-7
VIDLIČKA, Ľ., REZBANYAI-RESER, L. 2005. Neuere Angaben zur Schabenfauna der
Schweiz (Blattaria, Blattellidae: Ectobius). Entomologische Berichte Luzern 53: 123-134.
VIDLIČKA, Ľ. 2007. Šváby (Blattaria) a ich parazitoidy (Hymenoptera: Evaniidae) na
ostrove Kopáč (Bratislava-Podunajské Biskupice), pp. 113-118. In: Príroda ostrova
Kopáč. Fytoterapia OZ pri Pedagogickej fakulte UK, Bratislava. ISBN 978-80-969718-
7-9
FEDOR, P., VIDLIČKA, Ľ., MAJZLAN, O., VARGA, L. 2010. Ucholaky (Dermaptera),
modlivky (Mantodea) a šváby (Blattaria) PR Šúr, pp. 127-133. In: Majzlan, O., Vidlička,
Ľ. (eds). Príroda rezervácie Šúr. Ústav zoológie SAV, Bratislava. ISBN 978-80-970326-
0-9.
VIDLIČKA, Ľ., OZIMEC, R. 2011. Research of cockroaches (Blattaria) in PP Biokovo - first
preliminary results [Istraživanje žohara (Blattaria) u PP Biokovo - prvi preliminarni
rezultati], pp. 33-34. In: Protrka, K., Škrabić, H. & Srzić, S. (Eds) Znanstveno – stručni
skup “Biokovo na razmeđi milenija: razvoj parka prirode u 21. stoljeću“ [Scientific and
professional meeting "Biokovo at the turn of the millennium: the development of Nature
Park in the 21st century”]. Park prirode Biokovo, Makarska. ISBN 978-953-56909-0-0
VIDLIČKA, Ľ. 2012. Sezónne zmeny vo výskyte švábika hôrneho (Ectobius sylvestris)
(Blattaria) v Martinskom lese (Podunajská rovina), pp. 115-120. In: Fedor, P., Vidlička,
Ľ. (eds) Príroda Martinského lesa (vybrané kapitoly). Ústav zoológie SAV, Bratislava.
TOMKO, P., VIDLIČKA, Ľ., MOCK, A. 2013. Nové údaje k rozšíreniu švábov (Blattodea) a
ucholakov (Dermaptera) na východnom Slovensku. Folia faunistica Slovaca 18(1): 47-53.
VIDLIČKA, Ľ. 2014. Ectobius vittiventris – nový šváb (Blattaria) pre faunu Slovenska.
Entomofauna Carpathica 26(1): 33-40.
Ľubomír Vidlička Habilitačná práca
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5. Šváby juhovýchodnej Ázie a Južnej Ameriky
Juhovýchodná Ázia (orientálna faunistická oblasť) a Južná Amerika (neotropická
faunistická oblasť) sú dve na šváby najbohatšie oblasti s množstvom endemických druhov
a rodov. Štyri z nich si bližšie spomenieme.
5.1. Šváby z rodu Caeparia Stål, 1877 (Blaberidae: Panesthiinae)
1. C. saussurii (Wood-Mason, 1876) – India, Bután, Bangladéš
2. C. crenulata (Bruijning, 1948) – Indonézia (Sumatra), Malajzia
3. C. donskoffi Roth, 1979 – Vietnam, Laos, Thajsko
4. C. kaltenbachi Roth, 1979 – Thajsko, Laos
5. C. sausai Vidlička, 1999 – Laos
Všetky druhy rodu Caeparia sú pomerne veľké šváby, ktoré majú dobre vyvinuté
tegminy aj krídla. Nebolo u nich pozorované ohryzenie tegmín typické pre príslušníkov
príbuzného rodu Panesthia, ako aj iných príslušníkov
podčeľade Panesthiinae. Všetky druhy rodu Caeparia majú
dvojfarebné tegminy, čo je tiež v rámci podčeľade
nezvyklé.
História rodu Caeparia sa začala odvíjať od omylu.
Svetoznámy švajčiarsky ortopterológ Henri Louis
Frederic de Saussure (*1829-†1905) opísal v roku 1863 z
Číny švába Panesthia mandarinea (obr. 24). Pohlavie
švába neuviedol, lebo opisovanému jedincovi chýbalo
bruško (SAUSSURE 1863). O pár rokov neskôr sa mu
dostala do rúk samička švába z východnej Indie. Priradil ju
k tomu istému druhu, urobil jej pomerne obsiahly opis a v
úvode doplnil poznámku, že v predchádzajúcom článku opísal samčeka (SAUSSURE 1869).
Anglický zoológ James Wood-Mason (*1846-†1893) po skončení štúdia v Oxforde
odišiel v roku 1869 do Indie a získal zamestnanie v Indickom múzeu v Kalkate. Dostal
obsiahlu zbierku hmyzu zo štátu Johor v Malajzii s početnou sériou exemplárov Panesthia
mandarinea. Pri ich štúdiu si všimol, že oba jedince z tohto druhu, ktoré opísal Saussure sú
samičky, ale nepatria k tomu istému druhu. Samičke z Indie, ktorá bola opísaná neskôr,
vytvoril v roku 1876 nové meno Panesthia Saussurii. Do tohto druhu zaradil aj exemplár
práve sa zvliekajúceho samčeka zo štátu Sikkim ležiaceho v Himalájach na severovýchode
Indie (WOOD-MASON 1876).
O rok neskôr publikoval švédsky ortopterológ Carl Stål (*1833-†1878) prácu s opisom
viacerých druhov švábov z Filipín a medzi nimi bol aj druh Panesthia saussurii (iný ako bol
druh rovnako pomenovaný Wood-Masonom). Zároveň v rode Panesthia stanovil nový
podrod Caeparia a zaradil do neho druh Panesthia mandarinea opísaný Saussurem.
Nenapísal z ktorej Saussureho práce vychádzal, ale na základe diagnostických znakov
podrodu je evidentné, že sa jednalo o samičku z Indie opísanú v roku 1869 (STÅL 1877).
Ľubomír Vidlička Habilitačná práca
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Obr. 24. Panesthia mandarinea
(SAUSSURE 1863).
V sérii omylov pokračoval aj Karl Brunner von Wattenwyl (*1823-†1914).
V obsiahlej revízii ortopteroidného hmyzu, ktorú v roku 1893 vydal v Ženeve, povýšil
Stålom vytvorený podrod Caeparia na samostatný rod s typovým druhom Caeparia
mandarinea Saussure z lokality Teinzò (BRUNNER DE WATTENWYL 1893). Ktorý druh v
skutočnosti myslel nie je isté, ale keďže sa odvoláva na Stålov opis supraanálnej
platničky, tak pravdepodobne sa tiež jednalo o exemplár z Indie. Udávaná lokalita Teinzò
(asi Teinzô) sa nachádza na severe Mjanmarska, neďaleko od Indie. Brunner uvádza v
zozname použitých prác 6 Wood-Masonových článkov (3 o Phasmatodea a 3 o Mantodea),
ale jeho prácu o Panesthia mandarinea a P. saussurii nepoznal (alebo prinajmenšom
nepoužil).
Približne v tom istom čase SAUSSURE (1895) spracovával revíziu Panesthinov. Na
rozdiel od svojich predchodcov inkriminovaný článok Wood-Masona poznal a poznal aj
práce Ståla a Brunnera von Wattenwyl o rode Caeparia. Preto poopravil Brunnerov údaj a
za typový druh rodu určil Caeparia saussurei (upravil chybne vytvorené meno Saussurii na
Saussurei, omylom však uviedol, že pôvodné meno bolo Sausseuri) (SAUSSURE 1895).
Takže meno Panesthia mandarinea zostalo pre druh opísaný v roku 1863. William Forsell
Kirby (*1844-†1912) vo svojom synonymickom katalógu ortopteroidného hmyzu uvádza
v rode Caeparia len druh C. Saussurii Wood-Mason a pri rode Panesthia aj druh P.
Saussurii Stål (KIRBY 1904).
Situáciu opäť skomplikoval americký ortopterológ Andrew Nelson Caudell (*1872†1936).
Domnieval sa, že typom rodu Caeparia je druh Panesthia mandarinea opísaná
Saussurem v roku 1863 (a nie tá z roku 1869). Preto pre druh Panesthia Saussurii stanovený
Wood-Masonom v roku 1876 vytvoril nový rod Neocaeparia (CAUDELL 1924).
Vzniknutý problém sa podujal vyriešiť najvýznamnejší americký švábológ Louis
Marcus Roth (*1918-†2003). Na základe článku 70(a) pravidiel medzinárodnej
zoologickej nomenklatúry sa obrátil na Medzinárodnú komisiu. Keďže Stål zle identifikoval
druh „mandarinea“ ako typový druh rodu Caeparia, navrhol Roth komisii akceptovať
možnosť (i) článku 70(a) a designovať za typový druh rodu Caeparia druh Panesthia
saussurii Wood-Mason (ROTH & GURNEY 1983). Toto riešenie bolo prijaté, keďže
korešpondovalo aj s prácami SAUSSUREHO (1895) a KIRBYHO (1904). ROTH (1979) na
základe tohto rozhodnutia pri revízii rodu Caeparia opätovne stanovil za typový druh rodu
Panestia sausurii Wood-Mason [=Panesthia mandarinea Saussure, 1869 (nec 1863)].
Zároveň v zhode s PRINCISOM (1950) synonymizoval rod Neocaeparia Caudell, 1924 s
rodom Caeparia.
Zaujímavý je aj osud druhu Panesthia mandarinea Saussure, 1863. V roku 1932 sa
nemecký ortopterológ Karl Richard Hanitsch (*1860-†1940) zaoberal zbierkami švábov
Dr. Odoarda Beccariho a Dr. Elia Modiglianiho zo Sumatry (HANITSCH 1932). Na základe
Modiglianim chytených exemplárov druhu Panesthia transversa zistil, že tento druh je
totožný s jedincom opísaným Saussurem v roku 1863 pod menom Panesthia mandarinea.
Druh Panesthia transversa opísal Burmeister už v roku 1838. Preto Hanitsch názov
Panesthia mandarinea synonymizoval s názvom Panesthia transversa. Z názvu Panesthia
mandarinae, ktorý bol použitý dvakrát pre dva rôzne druhy tak nezostal nakoniec platný ani
jeden.
Ľubomír Vidlička Habilitačná práca
25
Všetky doteraz uvedené údaje sa týkali len jediného druhu rodu Caeparia, typového
druhu C. saussurii. Až v roku 1948 opísal holandský entomológ Conrad Friedrich Albert
Bruijning (*1919-†2004) druh Neoceparia crenulata podľa samčeka chyteného na Mount
Kerinci (v orig. Mount Indrapura) v pohorí Barisan na západe ostrova Sumatra (Indonézia).
ROTH (1979a) revidoval aj samčeka tohto druhu z pevninskej časti Malajzie, z lokality
Tapah Perak. BRUIJNING (1948) sa pri zaradení do rodu Neocaeparia pridŕžal Caudellovej
práce z roku 1924. Do rodu Caeparia tento druh presunul až PRINCIS (1965).
Na opis ďalších druhov z rodu Caeparia bolo treba počkať ďalších 31 rokov. Roth začal
v roku 1977 vydávať rozsiahlu celosvetovú revíziu podčeľade Panesthinae. Celkovo vydal
4 monografické práce (ROTH 1977, 1979a, b, 1982). V druhom diele spracoval aj rod
Caeparia. Tu pri druhu Caeparia saussurii dôkladne charakterizoval obe pohlavia a tiež
pridal ďalšie lokality výskytu z Indie, Bangladéša, Bhutánu a Laosu. SAUSSUREM (1895)
spomenutý výskyt tohto druhu na lokalite Malacca (JZ Malajzia) Roth spochybnil (podobne
ako pred ním aj PRINCIS (1965)), lebo z okolia tejto lokality determinoval len druh
Caeparia crenulata, ktorý však v dobe vyjdenia Saussureho práce nebol ešte opísaný.
V práci, v ktorej ROTH (1979a)
revidoval rod Caeparia, zároveň
opísal aj dva nové druhy z tohto
rodu: C. donskoffi (pomenovaný
podľa odborníka na Orthoptera
Michela Donskoffa z Muséum
national d’Histoire naturelle v
Paríži) a C. kaltenbachi
(pomenovaný podľa Dr. Alfreda
Kaltenbacha (*1920-†2005) z
Naturhistorisches Museum vo
Viedni). U druhu Caeparia donskoffi bola pôvodne opísaná iba samička (holotyp z Ko-Tichi
z Južného Vietnamu, paratypy z Tonkinu zo severného Vietnamu) (ROTH 1979a). Samček
bol opísaný až dodatočne zo severného Vietnamu a zo
severného Laosu (VIDLIČKA 1999) a samička bola
zistená aj v Thajsku (VIDLIČKA 1999).
Druh Caeparia kaltenbachi bol opísaný tiež iba na
základe samičiek. Holotyp pochádza zo severu Thajska
(pohorie Khun Tan) a paratyp zo severného Laosu
(Muong Pek) (ROTH 1979a). Samček nie je doteraz
známy.
ROTH (1982) uverejnil opis nepomenovanej nymfy z
Thajska, ktorá by mohla patriť k jednému z ním
opísaných druhov z rodu Caeparia. Farebným vzorom
sa síce odlišuje od imág C. donskoffi (obr. 25) aj C.
kaltenbachi, ale zadný okraj supraanálnej platničky
(dôležitý pri determinácii) je podobný ako u C.
donskoffi.
Ľubomír Vidlička Habilitačná práca
26
Obr. 25. Samička druhu Caeparia donskoffi
(VIDLIČKA 1999).
Obr. 26. Caeparia sausai ♀
(VIDLIČKA 1999).
Zatiaľ posledný, piaty druh z
rodu Caeparia bol opísaný v
roku 1999 na základe samičky
zo severného Laosu (provincia
Attapu, Bolaven Plateau, 15 km
JV od Ban Houaykong).
Pomenovaný bol Caeparia
sausai (obr. 26) na počesť
slovenského amatérskeho
entomológa Ondreja Šaušu,
ktorý tento druh objavil
(VIDLIČKA 1999). Rozšírenie
druhov rodu Caeparia ukazuje
obrázok 27.
5.2. Šváby z rodu Chorisoserra (Blattellidae: Pseudophyllodrominae)
1. Ch. sagitaria (Hanitsch, 1927) – Vietnam
2. Ch. apicalis (Hanitsch, 1929) – Indonézia (Sumatra), východné Borneo
3. Ch. jendeki Vidlička, 2002 – Laos
4. Ch. biceps Wang, Zhang, Feng, 2006 – Čína
5. Ch. brevicaudata Wu, Wang, 2011 - Čína
V roku 1927 Hanitsch spracoval zbierku švábov z južného Annamu (terajší stredný
Vietnam). Jedným z 19 opísaných nových druhov bol aj druh Chorisoneura sagitaria. Opis
urobil na základe samčeka z Langbian Peak (provincia Lam Dong) (HANITSCH 1927). Rod
Chorisoneura vytvoril Brunner von Wattewyl v roku 1865. Je to veľký rod, v súčasnosti s
90 druhmi. V čase keď Hanitsch publikoval opis druhu Chorisoneura sagitaria obsahoval
tento rod už 55 druhov. Na novoopísanom druhu bolo zaujímavé, že bol nájdený v
orientálnej oblasti. Takmer všetky ostatné druhy tohto rodu pochádzali z neotropickej
oblasti. Výnimkou boli vtedy 3 druhy z Tajvanu, Číny a Kambodže.
Ďalší druh z orientálnej oblasti opísal Hanitsch zo zbierky švábov švédskeho zoológa
Erica Georga Mjöberga (*1882-†1938). Zbierku nazhromaždil Mjöberg počas expedície
na Sumatre v rokoch 1919-1922. Hanitschom opísaný druh Chorisoneura apicalis pochádzal
z lokalít Medan a Boeloe-Tjina (HANITSCH 1929). Jedince z rodu Chorisoneura majú dosť
charakteristický vzhľad a tak správnosť zaradenia týchto dvoch Hanitschom opísaných
druhov, svojim rozšírením anomálnych, nikto dlhú dobu nespochybňoval. Dokonca ani
jeden z najväčších švábológov sveta Lotiš Kārlis Aleksandrs Princis (*1893-†1978), autor
zatiaľ neprekonaného 8 dielneho celosvetového katalógu švábov z rokov 1965-1972. V roku
1950 Princis spracovával zbierku švábov z orientálnej oblasti uloženú v entomologickom múzeu
Univerzity v Lunde (Švédsko). V zbierke našiel aj druh, ktorý určil ako Chorisoneura
apicalis. Jedinec pochádzal z východného Bornea (PRINCIS 1950). Na zaradení sa nič
nezmenilo ani pri vydaní prvého dielu spomínaného katalógu švábov (PRINCIS 1965).
Ľubomír Vidlička Habilitačná práca
27
Obr. 27. Rozšírenie švábov z rodu Caeparia
v juhovýchodnej Ázii.
V roku 1998 Roth revidoval zástupcov rodu Chorisoneura.
Všimol si určité odlišnosti dvoch orientálnych druhov z rodu
Chorisoneura od neotropických druhov, ale tiež aj odlišnosti
druhov z Číny, Tajvanu a Kamerunu. Preto pre Hanitschom
opísané druhy vytvoril nový rod Chorisoserrata a pre ostatné
odlišné druhy vytvoril rody Chorisoneurodes a Sorineuchora
(ROTH 1998).
Tretí druh do rodu Chorisoserrata, Ch. jendeki (obr. 28),
pribudol až v roku 2002. Opísaný bol na základe dvoch
samčekov a jednej samičky z Laosu (provincie Bolikhamxai a
Luang Namtha). Od predchádzajúcich druhov sa okrem iného
líšil aj prítomnosťou nezreteľnej tergálnej žľazy na siedmom
abdominálnom tergu (VIDLIČKA 2002). Druh dostal meno podľa
slovenského entomológa Eduarda Jendeka, nálezcu holotypu.
Čínsky entomológovia Zong-Qing Wang, Yan-Ning Zhang
a Ping-Zhang Feng z Čínskej akadémie poľnohospodárskych
vied opísali na základe 6 jedincov (4 ♂♂, 2 ♀♀) pochádzajúcich
z provincie Hainan (ostrov Chaj-nan) a z autonómnej oblasti
Guangxi Zhuang (Kuang-si, JV Čína) nový druh Chorisoserrata
biceps. Od ostatných príslušníkov rodu sa odlišuje dlhou,
zakrútenou mediánnou falomérou a zúbkami na konci prídavnej
mediánnej faloméry (WANG et al 2006).
Ke-Liang Wu a Zong-Qing
Wang objavili v južnej Číne, v
provincii Yunnan (Jün-nan)
samčeka a samičku z rodu
Chorisoserrata. Samčie
genitálie obsahovali dlhú, úzku,
na konci guľovito zhrubnutú
mediánnu faloméru a prídavnú
mediánnu faloméru zakončenú
v tvare chĺpkov štetca. Druh
pomenovali Chorisoserrata
brevicaudata (WU & WANG
2011). Oba čínske druhy sú bez
špecializácie na 7 abdominálnom
tergite.
Ľubomír Vidlička Habilitačná práca
28
Obr. 29. Rozšírenie švábov z rodu Chorisoserrata
v juhovýchodnej Ázii.
Obr. 28. Chorisoserrata
jendeki - štít a tegmina
(VIDLIČKA 2002).
5.3. Šváby z rodu Spelaeoblatta Bolívar, 1897 (Noticolidae)
1. S. gestroi Bolívar, 1897 – Mjanmarsko (Barma)
2. S. thamfaranga Roth, 1994 – Thajsko
3. S. myugei Vidlička, Vršanský & Shcherbakov, 2003 – Thajsko
4. S. thailandica Vidlička, Vršanský & Shcherbakov, 2003 – Thajsko
V roku 1892 spracovával španielsky ortopterológ Ignacio Bolívar y Urrutia (*1850†1944)
zbierku švábov, ktoré nazberal v marci a apríli 1890 francúzsky arachnológ Eugène
Simon (*1848-†1924) v jaskyniach na ostrove Luzon (Filipíny). Materiál pochádzal z
dvoch provincií – provincia Manila a provincia Morong. Boli to prvé naozaj jaskynné šváby
prispôsobené plne životu v tomto netypickom prostredí – malé, dlhonohé, depigmentované,
so silne redukovanými očami resp. bez očí (BOLÍVAR 1892). Bolívar opísal na ich základe
nový rod Nocticola, ktorý zaradil do čeľade Blattidae. Zároveň v rode opísal dva druhy Nocticola
simoni (na počesť zberateľa, na základe samčeka a samičky z jaskyne San Mateo,
prov. Manila) a Nocticola caeca (na základe samičky z jaskyne Antipodo, prov. Morong).
Samičky nemali vyvinuté tegminy ani krídla.
Známy taliansky cestovateľ, zoológ, maliar a zberateľ prírodnín Leonardo Fea (*1852†1903)
sa v roku 1871 stal asistentom na oddelení entomológie v Museo Civico di Storia
Naturele di Genova v Janove. V roku 1885 sa vydal na cestu do Barmy, kde zotrval 4 roky
(1885-1889). Nachytané živočíchy pravidelne odosielal do svojho domovského múzea. V
decembri 1887 sa dostal do oblasti vrchu Carin medzi riekami Sittang a Saluin. Tu sa mu
podarilo v januári 1888 v jaskyni Jaddò (Ya-dò) uloviť samičku jaskynného švába a tiež ho
odoslal do múzea v Janove. Riaditeľom múzea bol v tej dobe Raffaello Gestro (*1845†1936).
Ten spracoval skupinu Coleoptera a zistenia publikoval v muzeálnom časopise
(GESTRO 1888, 1891). Zvyšok materiálu distribuoval na určenie špecialistom na dané
skupiny.
Švába zaslal na determináciu Bolívarovi. Na prvý
pohľad bola zaslaná samička dosť podobná jedincom z
rodu Nocticola, ale bola väčšia a hlavne mala vyvinuté,
aj keď silne redukované, tegminy. Na základe tejto
samičky opísal Bolívar nový rod jaskynných švábov
Spelaeoblatta. Nový druh nazval Spelaeoblatta Gestroi
(na počesť Raffaella Gestroa) (obr. 30) (BOLÍVAR 1897).
Rod Spelaeoblatta zostal dlhú dobu monotypický.
George C. McGavin1
sa špecializuje na výskum
suchozemských článkonožcov v tropických lesoch a
jaskyniach. V apríli až máji 1992 sa zúčastnil expedície
usporiadanej CSCA (Combined Services Caving
Association) nazvanej „Exercise Tham Farang“ do
Ľubomír Vidlička Habilitačná práca
29
Obr. 30. Spelaeoblatta gestroi ♀
(BOLÍVAR 1897).
1
McGavin nie je špecialistom na šváby, zaoberá sa viac popularizačnou činnosťou. Má vlastnú reláciu v BBC
a napísal viacero populárnych knižiek o hmyze. Jednu z nich – „Insects, spiders and other terrestrial
arthropods“ (MCGAVIN 2000) som mal česť v roku 2000 preložiť pre vydavateľstvo IKAR do slovenčiny
(spolu s L. Rollerom).
Thajska. Cieľom expedície bolo zhromaždiť a identifikovať kavernikolné živočíchy
jaskynných systémov v provincii Kanchanaburi. V jaskynnom systéme Tham Nam Farang
bolo nájdené aj väčšie množstvo samčekov, samičiek aj nýmf jaskynných švábov
neznámeho druhu. Preto McGavin požiadal o pomoc s identifikáciou amerického experta na
šváby Louisa M. Rotha. Šváby boli identifikované ako nový druh z rodu Spelaeoblatta.
Roth pomenoval tento druh Spelaeoblatta thamfaranga na počesť expedície. Thajské slová
„Tham Farang“ znamenajú „jaskynný cudzinec“. Na základe tohto materiálu bol po
prvýkrát charakterizovaný aj samček z tohto rodu. Na abdomene majú samčekovia vyvinutú
pohlavnú tergálnu žľazu. Prítomnosť tergálnej žľazy na 2 a 3 tergite samčekov je dosť
vzácny úkaz medzi švábmi. V práci bola tiež podrobnejšie opísaná aj samička. U tohto
druhu na rozdiel od S. gestroi mali samičky vyvinuté facetové oči (ROTH & MCGAVIN
1993).
Ruská expedícia skúmajúca jaskyne severného Thajska nazbierala v marci 1997 okrem
iného aj neveľký materiál jaskynných švábov. Slovenská expedícia na Ďaleký východ do
Primorského (Ussurijského) kraja (Vidlička, Vršanský,
Roller – jún 1997) prevzala nazberané šváby na
determináciu. Ako sa ukázalo materiál obsahoval
minimálne tri druhy jaskynných švábov, z ktorých dva
druhy boli zaradené do rodu Spelaeoblatta. Opisy oboch
vyšli v roku 2003. Prvý dostal meno Spelaeoblatta
myugei (obr. 31) po svojom zberateľovi N. Myugem (Н.
Мюге), ktorý zbieral šváby v jaskyni Tham Pha Mon
(provincia Mae Hong Son, región Nam Lang). Druhý
bol pomenovaný Spelaeoblatta thailandica podľa
krajiny pôvodu a opísaný bol z jaskyne Red Cliff
(provincia Mae Hong Son, región Nam Lang). Oba
druhy sa veľmi nápadnými a v rade švábov úplne
unikátnymi výbežkami na tergitoch bruška výrazne líšili
od predchádzajúcich dvoch druhov a preto bola pre nich
vytvorená samostatná skupina druhov myugei
(VIDLIČKA et al. 2003).
Ľubomír Vidlička Habilitačná práca
30
Obr. 31. Spelaeoblatta myugei ♂
(VIDLIČKA et al. 2003).
5.4. Šváby z rodu Macrophyllodromia Saussure & Zehnter, 1893 (Blattellidae)
1. M. maximiliani (Saussure, 1873) – Mexiko, Guatemala, Honduras, Kostarika, Panama
2. M. splendida Hebard, 1920 – Panama
3. M. nigrigena Hebard, 1926 – Francúzska Guiana, Guyana, Bolívia (?)
4. M. ecuadorana Rocha e Silva, 1962 – Ekvádor, Honduras (?)
5. M. panamae Rocha e Silva, 1962 – Panama
6. M. lanceolata Lopes & Oliveira, 2006 – Brazília
7. M. multipunctata Lopes & Oliveira, 2006 – Brazília
8. M. amabile Anisyutkin, 2007 – Ekvádor
9. M. nobile Anisyutkin, 2007 – Ekvádor
10. M. beccalonii Anisyutkin, 2012 – Ekvádor
11. M. rufidula Anisyutkin, 2012 – Mexiko
12. M. onorei Vidlička, 2013 – Ekvádor
Švajčiarsky ortopterológ Henri Louis Frederic de Saussure (*1829-†1905) sa výskumu
ortopterofauny strednej Ameriky venoval dlhodobo. Už v roku 1854 podnikol spolu s
ďalším švajčiarskym prírodovedcom Adrienom Jeanom Louisom Françoiscom de
Sumichrast (*1828-†1882) expedíciu do „Západnej Indie“ (Karibské ostrovy) a do Mexika.
Chytali hmyz a posielali ho do ženevského múzea. Saussure sa v roku 1856 z Mexika vrátil
do Ženevy a začal nazberaný materiál spracovávať.
V roku 1873 spracoval Saussure rozsiahlu zbierku švábov a modliviek, medzi ktorým
bolo aj niekoľko exemplárov z Mexika. O ich
pôvode sa bližšie nezmieňuje, ale nie všetky
pochádzali z jeho vlastných zberov. Jedného zo
švábov, charakteristického dvomi nápadnými
čiernymi pozdĺžnymi pruhmi na pronóte,
pomenoval Pseudophyllodromia Maximiliani
(obr. 32). Holotyp bol poškodený, chýbalo mu
bruško, takže pohlavie neuvádza (SAUSSURE
1873). Dá sa predpokladať, že druh dostal meno
po Ferdinandovi Maximiliánovi Jozefovi
Habsburskom (*1832-†1867), ktorý bol ako
Maximilián I. v rokoch 1861-1867 mexickým
panovníkom.
Saussure a jeho spolupracovník Leo Zehntner (*1864-†1961), asistent v Natural History
Museum v Ženeve, vydali v roku 1893 v rámci edície „Biologia Centrali Americana“
rozsiahle dielo o šváboch a modlivkách Strednej Ameriky. V rámci rodu
Pseudophyllodromia uvádzajú dva druhy – P. venosa a P. maximiliani, oba z Mexika.
Vzhľadom na odlišnú žilnatinu na tegminách a krídlach týchto druhov ako aj odlišné
otŕnenie stehien zaradili druh P. venosa (=Euphyllodromia angustata (Latreille, 1811)) do
nominátneho podrodu Pseudophyllodromia a druh P. maximiliani do novo vytvoreného
podrodu Macrophyllodromia (SAUSSURE & ZEHNTNER 1893).
Ľubomír Vidlička Habilitačná práca
31
Obr. 32. Macrophyllodromia maximiliani
(SAUSSURE 1873).
Robert Walter Campbel Shelford (*1850-†1944) zistil, že príslušníci rodu
Pseudophyllodromia Brunner von Wattenwyl, 1865, ktoré sú rozšírené v indomalajskej
(orientálnej) oblasti sa odlišujú od druhov pochádzajúcich z juhoamerickej oblasti, preto
vytvoril pre druhy z tohto rodu z juhoamerickej oblasti nový podrod Euphyllodromia. Do
podrodu Euphyllodromia bol zaradený aj druh P. venosa a zároveň bol synonymizovaný so
starším druhom Euphyllodromia angustata (SHELFORD 1908). Saussurem a Zehntnerom
stanovený podrod Macrophyllodromia bol následne povýšený na rod s jediným druhom M.
maximiliani (SHELFORD 1908).
Rodák z poľskej časti Sliezska Eugene Amandus Schwarz (*1844-†928) emigroval v
roku 1872 do USA. Tu prešiel viacerými zamestnaniami až sa stal kurátorom pre chrobáky
(Coleoptera) v United States National Museum vo Washingtone D.C. (súčasť Smithsonian
Institution). Odtiaľ podnikol viacero dlhších expedícií do Panamy, Guatemaly, Mexika a na
Kubu. Šváby, ktoré počas expedícií nazbieral, spracoval americký ortopterológ Morgan
Hebard (*1887-†1946). Hebard sa tiež zúčastnil v roku 1913 kratšej expedície do oblasti
Panamského prieplavu a zberal hmyz. V roku 1920 spracoval Hebard šváby Panamy v
monografickom diele (HEBARD 1920). Medzi švábmi sa nachádzal aj samček zo zberov E.
A. Schwarza z roku 1911, ktorý bol veľmi podobný jedincovi, ktorého opísal Saussure z
Mexika. Hebard ho opísal pod názvom Macrophyllodromia splendida. Keďže mal k
dispozícii celého jedinca, opísal rod aj druh obšírne, vrátane genitálií (HEBARD 1920).
V ďalších rokoch sa Hebard pustil aj do
spracovania švábov z Francúzskej Guiany. Mal
vlastnú kolekciu a zároveň spracoval aj materiál od
iných Amerických prírodovedcov a zberateľov.
Tentoraz natrafil na väčšiu sadu (3 ♂♂, 3 ♀♀ a 2
nymfy) jedincov z rodu Macrophyllodromia. Ich
genitálie sa značne líšili od predchádzajúceho druhu,
takže opísal nový druh Macrophyllodromia nigrigena
(obr. 33). Jeden samček a dve samičky pochádzali z
Hebardovej zbierky z osady a francúzskeho
trestaneckého tábora Saint Laurent du Maroni
(Francúzska Guiana) a zvyšok jedincov nazberal v
roku 1922 americký prírodovedec a entomológ
Charles William Beebe (*1877-†1962) na lokalite
Kartabo (oblasť Bartica, Britská Guyana).
Samčekovia mali veľmi komplikovaným spôsobom
utvárané výbežky na subgenitálnej platničke
(HEBARD 1926). Práve tieto výbežky sú dobrým
determinačným znakom.
Isolda Rocha e Silva Albuquerque (*1935)
začala už vo svojich 15-tich rokoch pracovať ako
praktikantka v Národnom múzeu Brazílskej
univerzity, kde v roku 1952 začala budovať zbierku
neotropických švábov, ktorá existuje dodnes. V roku
1960 sa stala praktikantkou v United States National
Ľubomír Vidlička Habilitačná práca
32
Obr. 33. Macrophyllodromia nigrigena ♂,
habitus a subgenitálna platnička
(HEBARD 1926).
Museum u Dr. Gurneyho a pokračovala u Dr. Rehna v Academy of Natural Sciences vo
Filadelfii. Počas pobytu v USA preštudovala zbierky neotropických švábov z viacerých
múzeí. Po návrate z USA sa zamestnala v Museu Nacional v Rio de Janeiro. Tu publikovala
prácu revidujúcu rod Macrophylloromia. Preštudovala 5 ♂♂ a 4♀♀ z druhu M. maximiliani
z Hondurasu, Guatemaly, Kostariky a Panamy. V zbierke USNM objavila aj samčeka z
Panamy, ktorý mal odlišne tvarovanú subgenitálnu platničku ako M. maximiliani. Opísala
ho pod menom M. panamae. Holotyp pochádzal z ostrova Barro Colorado v Panamskom
prieplave (ROCHA E SILVA 1962). Uloviť sa ho podarilo Jamesovi Zetekovi (*1886-1†959),
synovi českých emigrantov v USA. V roku 1911 odcestoval Zetek do Panamy, kde sa ako
entomológ stal základňovým riaditeľom Canal Zone Biological Area (CZBA). Základňa
bola umiestnená na ostrova Barro Colorado. Zetek sa tu venoval hlavne výskumu termitov,
ale zberal aj iné druhy hmyzu, okrem iného aj šváby.
Pri štúdiu švábov v U.S. National Museum sa Rocha e Silva dostala aj ku švábom, ktoré
boli do USA dovezené so zásielkami banánov z Hondurasu (?) a Ekvádoru. Na ich základe
opísala druh Macrophyllodromia ecuadorana. Holotyp (♂) chytený v San Diegu pochádzal
z Guayaquilu a allotyp (♀) chytený v Kalifornii pochádzal tiež z Ekvádoru. Okrem toho
preštudovala ďalších 7 paratypov chytených na banánoch pochádzajúcich z Ekvádoru a v
jednom prípade možno z Hondurasu (ROCHA E SILVA 1962). Na priložených obrázkoch
zrejme zobrazila holotyp a allotyp. Je však možné (a vzhľadom na rôzny pôvod aj veľmi
pravdepodobné), že všetky jedince označené ako paratypy nepatrili k rovnakému druhu. Z
druhu Macrophyllodromia nigrigena preštudovala samičku chytenú na lokalite Tumupasa
(Bolívia), čo tiež nie je (vzhľadom na umiestnenie lokality) pravdepodobne úplne
spoľahlivá determinácia.
Skôr ako v roku 1988 odišla Rocha e
Silva na dôchodok, vychovala si
niekoľko následníkov. Jej študentmi
boli aj Sonia Maria Lopes a Edivar
Heeren de Oliveira, ktorí momentálne
tiež pracujú v Museu Nacional,
Universisade do Rio de Janeiro v
Brazílii a zaoberajú sa taxonómiou
švábov. Zo zbierok múzea opísali dva
druhy (oba druhy podľa samčekov) z
rodu Macrophyllodromia: M. lanceolata
a M. multipunctata zo štátu Acre
ležiaceho na západe Brazílie (LOPES &
OLIVEIRA 2006).
Zo zvyšných 5 doteraz opísaných
druhov z rodu Macrophyllodromia
pochádzajú 4 z Ekvádoru. Väčšinu
opísal ruský entomológ Leonid
Anisyutkin podľa jedincov
nazberaných jeho kolegami: M. amabile
(1♂ a 1♀) z okolia vodopádu San
Ľubomír Vidlička Habilitačná práca
33
Obr. 34. Rozšírenie rodu Macrophyllodromia.
Podľa VIDLIČKU (2013a) – upravené a doplnené.
Rafael na Rio Coca a M. nobile (1♂) z okolia jazera Lago Grande na Rio Cuyabeno, oba
druhy z Ekvádoru (ANISYUTKIN 2007) a druhy M. beccalonii (1♂) z provincie Napo
(Ekvádor) a M. rufidula (1♂) z Veracruz (Mexiko)
(ANISYUTKIN 2012). Anisyutkin mal k dispozícii aj
paratyp druhu M. ecuadorana, avšak podľa opisu sa
zdá, že v tomto prípade sa jedná o iný druh. Výskyt
jednotlivých druhov zobrazuje obrázok 34.
Posledná spomínaná Anisyutkinova práca má síce
vročenie 2012, ale vyšla s oneskorením až v roku
2013. Preto nemohla byť zohľadnená v revízii rodu
Macrophyllodromia (VIDLIČKA 2013a). V rámci tejto
revízie bol opísaný nový druh M. onorei podľa
samčeka (obr. 35) z rezervácie Otongachi (okolie
Santo Domingo de los Colorados) (VIDLIČKA 2013a).
Paratyp M. ecuadorana študovaný Anisyutkinom za
zdá byť práve z tohto novo opísaného druhu. Novo
opísaný druh nebol zahrnutý do check-listu
ekvádorských druhov švábov, ktorý vyšiel súbežne s
opisom (VIDLIČKA 2013b).
5.5. Prehľad príspevkov autora k poznaniu švábov juhovýchodnej Ázie a Južnej Ameriky
(hnedou farbou sú uvedené príspevky, ktoré sú súčasťou habilitačnej práce)
VIDLIČKA, Ľ. 1999. Caeparia sausai sp.nov. from Laos, and description of the male
Caeparia donskoffi (Blattaria: Blaberidae: Panesthiinae). Entomological Problems 30(2):
1-5. (Príloha č. 7)
VIDLIČKA, Ľ. 2002. The new cockroach species from the genus Chorisoserrata from Laos
(Blattaria: Blattellidae: Pseudophyllodromiinae). Entomological Problems 32(2): 145-
147. (Príloha č. 8)
VIDLIČKA, Ľ., VRŠANSKÝ, P., SHCHERBAKOV, D.E. 2003. Two new troglobitic cockroach
species of the genus Spelaeoblatta (Blattaria: Nocticolidae) from North Thailand.
Journal of Natural History 37(1): 107-114. (Príloha č. 9)
VIDLIČKA, Ľ. 2013a. New species of Macrophyllodromia (Blattaria, Blattellidae) from
Ecuador and a key to males of the genus. Zootaxa 3635(2): 185-193. (Príloha č. 10)
VIDLIČKA, Ľ. 2013b. Cockroaches (Blattaria) of Ecuador—checklist and history of
research. Zootaxa 3599(5): 401-445. (Príloha č. 11)
Ľubomír Vidlička Habilitačná práca
34
Obr. 35. Macrophyllodromia onorei ♂
(VIDLIČKA 2013a).
6. Fosílne šváby (Blaberidae, Blattellidae, Ectobiidae)
Šváby (Blattaria) vznikli v karbóne v stupni Bashkirian (pred 323,20,4 až 315,20,2
miliónmi rokov). Najstaršie nálezy pochádzajú z Quilianshan v Číne (ZHANG et al. 2012,
GUO et al. 2012). Typické mezozoické čeľade boli odvodené z Phyloblattidae pri P/T hranici
a kmeň recentných čeľadí (a tiež všetkých modliviek a termitov) možno vysledovať z
druhohornej čeľade Liberiblattinidae (VRŠANSKÝ 2010, VRŠANSKÝ et al. 2012a).
Najstarším fosílnym záznamom o druhoch švábov recentnej čeľade je ectobiid (blattellid)
Piniblattella vitimica (VISHNIAKOVA 1964) z neskorej kriedy (VRŠANSKÝ 1997). Súčasné
rody boli známe až od skorého eocénu (ARCHIBALD & MATHEWES 2000) a moderná fauna
švábov pochádza z obdobia okolo paleocénno-eocénneho teplotného maxima (PETM;
VRŠANSKÝ et al. 2011, 2012b).
V triase, jure a kriede (druhohory; pred 2510,4 až
65,50,3 miliónmi rokov) žili na Zemi obrovské
terestrické plazy – dinosaury. Predpokladá sa, že tieto
obrie plazy živiace sa rastlinnou potravou produkovali
úmerne veľké množstvo exkrementov. V súčasnosti sú
exkrementy odstraňované (rozkladané) hlavne
koprofágnym hmyzom. V dobe dinosaurov však bol
takýto hmyz zriedkavý. Pri štúdiu švábov z libanonského
jantáru vznikla hypotéza, či by tými chýbajúcimi
„čističmi po dinosauroch“ nemohli byť šváby.
Libanonský jantár spred 125 miliónov rokov uchoval aj
nymfu švába zatiaľ neznámeho druhu z vyhynutej čeľade
Blattulidae. Nymfa je dokonale zachovaná (obr. 36) a
zachovali sa aj jej skamenené exkrementy – koprolity. V
týchto koprolitoch boli objavené synchrotrónovou
mikrotomografiou zvyšky dreva s hladkými hranami,
ktorých zdrojom by mohli byť práve výkaly bylinožravcov. Takže šváby sa pravdepodobne
živili exkrementami (obr. 37) dinosaurov a napomáhali ich biodegradácii (VRŠANSKÝ et al.
2013a). Šváby živiace sa exkrementami poznáme aj zo súčasnosti, najčastejšie je to guáno
po netopieroch (ROTH & WILLIS 1960, BELL et
al. 2007, CHRISTOFFERSEN & DE ASSIS 2013).
Podľa databázy fosílneho hmyzu (EDNA) je
známych z paleocénu (pred 65,50,3 až 55,80,2
miliónmi rokov) 177 druhov hmyzu. Najviac sú
to chrobáky (44), dvojkrídlovce a hemipterný
hmyz (po 28 druhov). Šváby sa v tejto databáze
donedávna vôbec nevyskytovali. Prvé dva druhy
švábov z tohto obdobia boli opísané z
danianských (paleocénnych) sedimentov
pochádzajúcich z ruského Ďalekého východu
(VRŠANSKÝ et al. 2013b). Orientálny
(indomalajský) rod Morphna zahŕňa 12
Ľubomír Vidlička Habilitačná práca
35
Obr. 36. Nymfa švába
z čeľade Blattulidae
(VRŠANSKÝ et al. 2013a).
Obr. 37. Koprofágne šváby
v druhohorách (HTTPS://WWW.PHACTUAL.COM/9-
CREEPY-COCKROACH-FACTS-YOU-DIDNT-WANT-TO-
KNOW/).
recentných druhov. Morphna paleo (obr. 38) bola
opísaná na základe zachovanej fosílnej tegminy
(VRŠANSKÝ et al. 2013b). Rod Ergaula obsahuje
7 recentných druhov, až na jednu výnimku tiež s
rozšírením v orientálnej oblasti. Jeden druh
(Ergaula atica) je známy z eocénu Izraela
(ANISYUTKIN et al. 2008) a novoopísaná
Ergaula stonebut je paleocénneho veku
(VRŠANSKÝ et al 2013b).
Chiapas, štát na najjužnejšom výbežku Mexika, je známy výskytom jantáru, pre ktorý je
príznačné, že pomerne často obsahuje zvyšky rastlín, príležitostne huby, vzácne ulitníky,
pavúky, kôrovce, pomerne hojný je hmyz a dajú sa objaviť aj obojživelníky, plazy, vtáčie
perie a srsť cicavcov (POINAR 2003, ENGEL 2004, SOLORZANO KRAEMER 2007, VEGA et
al. 2009a, b).
Chiapaský jantár vznikol v dolnom miocéne (pred
23,030,05 až 15,970,05 miliónmi rokov). Z hmyzu
zachovaného v tomto jantári bolo doteraz opísaných 235
druhov. Druhy, ktoré sa v tomto jantári vyskytujú, patria
často do recentných rodov. To je aj prípad švába nájdeného
v tomto jantári. Patrí do rodu Supella, kde patrí 10
recentných druhov. Súčasné druhy sú výskytom
obmedzené na africký kontinent. Supella miocenica (obr.
39) z chiapaského jantáru je sesterským druhom ku druhu
S. mirabilis, ktorý je široko rozšírený v štátoch rovníkovej
Afriky. Rozdiel je iba v tvare stredovej škvrny na pronóte
a vo veľkosti tela (VRŠANSKÝ et al. 2011). Výskyt rodu
Supella v miocénnej Amerike naznačuje niekdajší
kozmopolitný výskyt tohto rodu, ktorý sa zachoval len v Afrike.
Eocénne (pred 55,80,2 až 33,90,1 miliónmi rokov) sedimenty rieky Green River v
americkom Colorade sú bohaté na fosílie švábov. Z 11 tu zistených rodov švábov (17
druhov), až 9 rodov patrí aj k recentnej faune. Rod Blattella môžeme považovať za
historicky starší, bol zaznamenaný už v druhohornej kriede (VRŠANSKÝ 2008). Ostatné
recentné rody švábov sa objavujú až na
začiatku eocénu a majú moderný charakter. K
rodom, ktoré sa vyvinuli počas eocénu alebo
tesne pred, patrí aj rod Cariblattoides. Rod v
súčasnosti zahŕňa 13 recentných druhov
rozšírených v Strednej a Južnej Amerike.
Cariblattoides labandeirai (obr. 40) bol v
skorom eocéne pri Green River bežným
druhom. Našlo sa až 13 fosílií patriacich
tomuto druhu.
Prekvapivý bol objav švába z rodu Ectobius
v eocénnych sedimentoch rieky Green River
Ľubomír Vidlička Habilitačná práca
36
Obr. 38. Tegmina švába Morphna paleo
(VRŠANSKÝ et al. 2013b).
Obr. 39. Supella miocenica
(VRŠANSKÝ et al. 2011).
Obr. 39. Supella miocenica
(VRŠANSKÝ et al. 2011).
spred 49 miliónov rokov. Šváby rodu Ectobius sú v súčasnosti obmedzené výskytom na
Euráziu a Afriku (palearktická oblasť). Opísaný bol druh Ectobius kohlsi (obr. 41). Jeho
nález v Severnej Amerike (nearktická oblasť) indikuje kozmopolitné rozšírenie tohto rodu
počas stredného paleogénu a jeho neskoršie vyhynutie (VRŠANSKÝ et al. 2014).
Zástupcovia rodu sa reintrodukovali na Americký
kontinent v 20. storočí (CHANDLER 1985, 1992,
HOEBEKE & NICKLE 1981, HOEBEKE & CARTER
2010).
Zo stredného eocénu (pred 47,8 až 38
miliónmi rokov) pochádzajú fosílie zberané pri
Flathead River v Montane (USA). Našiel sa
medzi nimi aj šváb z rodu Latiblattella. Recentné
druhy rodu Latiblattella (18 druhov) sú rozšírené
v Strednej Amerike a v Karibiku. Latiblattella
avita predstavuje prvý fosílny nález druhu z tohto
rodu (GREENWALT & VIDLIČKA 2015).
6.1. Prehľad príspevkov autora k poznaniu fosílnych švábov
(Blaberidae, Blattellidae, Ectobiidae)
(hnedou farbou sú uvedené príspevky, ktoré sú súčasťou habilitačnej práce)
VRŠANSKÝ, P., CIFUENTES-RUIZ, P., VIDLIČKA, Ľ., ČIAMPOR, F., VEGA, F.J. 2011. AfroAsian
cockroach from Chiapas amber and the lost Tertiary American entomofauna.
Geologica Carpathica 62: 463-475. (Príloha č. 12)
VRŠANSKÝ , P., VIDLIČKA, Ľ., ČIAMPOR, F., MARSH, F. 2012b: Derived, still living
cockroach genus Cariblattoides (Blattida: Blattellidae) from the Eocene sediments of
Green River in Colorado, USA. Insect Science 19: 143–152. (Príloha č. 13)
VRŠANSKÝ , P., VAN DE KAMP, T., AZAR, D., PROKIN, A., VIDLIČKA, Ľ., VAGOVIČ, P.
2013a. Cockroaches Probably Cleaned Up after Dinosaurs. PLoS ONE 8(12): e80560.
doi:10.1371/journal. pone.0080560 (Príloha č. 14)
VRŠANSKÝ , P., VIDLIČKA, Ľ., BARNA, P., BUGDAEVA, E., MARKEVICH, V. 2013b:
Paleocene origin of the cockroach families Blaberidae and Corydiidae: Evidence from
Amur River region of Russia. Zootaxa 3635(2): 117-126. (Príloha č. 15)
VRŠANSKÝ , P., ORUŽINSKÝ, R., BARNA, P., VIDLIČKA, Ľ. & LABANDEIRA, C.C. 2014.
Native Ectobius (Blattaria: Ectobiidae) from the Early Eocene Green River Formation of
Colorado and Its Reintroduction to North America 49 Million Years Later. Annals of the
Entomological Society of America 107(1): 28-36. (Príloha č. 16)
GREENWALT, D.E., VIDLIČKA, Ľ. 2015. Latiblattella avita sp. nov. (Blattaria: Ectobiidae)
from the Eocene Kishenehn Formation, Montana, USA. Palaeontologia Electronica
18.1.16A: 1-9. (Príloha č. 17)
Ľubomír Vidlička Habilitačná práca
37
Obr. 41. Ectobius kohlsi
(VRŠANSKÝ et al. 2014).
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doi:10.1371/journal. pone.0080560
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Ľubomír Vidlička Habilitačná práca
47
Prehľad príloh
(Publikácie autora, ktoré tvoria súčasť habilitačnej práce.)
Tematický okruh 1: Etológia švábov
Príloha 1: VIDLIČKA, Ľ., HUČKOVÁ, A. 1993. Mating of the cockroach Nauphoeta cinerea
(Blattodea: Blaberidae): I. Copulatory behaviour. Entomological Problems
24(2): 69-73.
Príloha 2: HUČKOVÁ, A., VIDLIČKA, Ľ., KOZÁNEK, M. 1994. Mating of the cockroach
Nauphoeta cinerea (Blattodea: Blaberidae): II. Histamine changes during
courtship and copulation. Biologia 49(5): 691-695.
Tematický okruh 2: Rozšírenie a taxonómia švábov v strednej Európe a na Slovensku
Príloha 3: VIDLIČKA, Ľ. 1993. Seasonal dynamycs of vertical migration and distribution
of cockroach Ectobius sylvestris (Blattaria: Blattellidae: Ectobiinae) in oak
forest. Biologia 48(2): 163-166.
Príloha 4: VIDLIČKA, Ľ. 1993. Phyllodromica hungarica sp.nov., a new cockroach species
from Hungary (Insecta: Blattodea: Blattellidae: Ectobiinae). Entomological
Problems 24(1): 63-68.
Príloha 5: VIDLIČKA, Ľ. 1994. Phyllodromica transylvanica sp. nov., a new cockroach
species from Romania and key of the maculata-group of Phyllodromica in
central Europe. Entomological Problems 25(2): 55-62.
Príloha 6: VIDLIČKA, Ľ., MAJZLAN, O. 1997. Revision of the megerlei - group of the
cockroach genus Phyllodromica Fieber (Blattaria: Blattellidae, Ectobiinae).
Entomologica Scandinavica 28: 163-173.
Tematický okruh 3: Šváby juhovýchodnej Ázie a Južnej Ameriky
Príloha 7: VIDLIČKA, Ľ. 1999. Caeparia sausai sp.nov. from Laos, and description of the
male Caeparia donskoffi (Blattaria: Blaberidae: Panesthiinae). Entomological
Problems 30(2): 1-5.
Príloha 8: VIDLIČKA, Ľ. 2002. The new cockroach species from the genus Chorisoserrata
from Laos (Blattaria: Blattellidae: Pseudophyllodromiinae). Entomological
Problems 32(2): 145-147.
Príloha 9: VIDLIČKA, Ľ., VRŠANSKÝ, P., SHCHERBAKOV, D.E. 2003. Two new
troglobitic cockroach species of the genus Spelaeoblatta (Blattaria:
Nocticolidae) from North Thailand. Journal of Natural History 37(1): 107-114.
Ľubomír Vidlička Habilitačná práca
48
Príloha 10: VIDLIČKA, Ľ. 2013a. New species of Macrophyllodromia (Blattaria,
Blattellidae) from Ecuador and a key to males of the genus. Zootaxa 3635(2):
185-193.
Príloha 11: VIDLIČKA, Ľ. 2013b. Cockroaches (Blattaria) of Ecuador—checklist and
history of research. Zootaxa 3599(5): 401-445.
Tematický okruh 4: Fosílne šváby (Blaberidae, Blattellidae, Ectobiidae)
Príloha 12: VRŠANSKÝ, P., CIFUENTES-RUIZ, P., VIDLIČKA, Ľ., ČIAMPOR, F., VEGA,
F.J. 2011. Afro-Asian cockroach from Chiapas amber and the lost Tertiary
American entomofauna. Geologica Carpathica 62: 463-475.
Príloha 13: VRŠANSKÝ, P., VIDLIČKA, Ľ., ČIAMPOR, F., MARSH, F. 2012b. Derived, still
living cockroach genus Cariblattoides (Blattida: Blattellidae) from the Eocene
sediments of Green River in Colorado, USA. Insect Science 19: 143–152.
Príloha 14: VRŠANSKÝ, P., VAN DE KAMP, T., AZAR, D., PROKIN, A., VIDLIČKA, Ľ.,
VAGOVIČ, P. 2013a. Cockroaches Probably Cleaned Up after Dinosaurs. PLoS
ONE 8(12): e80560. doi:10.1371/journal. pone.0080560
Príloha 15: VRŠANSKÝ, P., VIDLIČKA, Ľ., BARNA, P., BUGDAEVA, E., MARKEVICH, V.
2013b. Paleocene origin of the cockroach families Blaberidae and Corydiidae:
Evidence from Amur River region of Russia. Zootaxa 3635(2): 117-126.
Príloha 16: VRŠANSKÝ, P., ORUŽINSKÝ, R., BARNA, P., VIDLIČKA, Ľ. & LABANDEIRA,
C.C. 2014. Native Ectobius (Blattaria: Ectobiidae) from the Early Eocene
Green River Formation of Colorado and Its Reintroduction to North America
49 Million Years Later. Annals of the Entomological Society of America
107(1): 28-36.
Príloha 17: GREENWALT, D.E., VIDLIČKA, Ľ. 2015. Latiblattella avita sp. nov. (Blattaria:
Ectobiidae) from the Eocene Kishenehn Formation, Montana, USA.
Palaeontologia Electronica 18.1.16A: 1-9.
Monografie
Príloha 18: VIDLIČKA, Ľ. 2001. Blattaria – šváby; Mantodea – modlivky: (Insecta:
Orthopteroidea). 1. vyd., Veda, Bratislava, 169 pp. (Fauna Slovenska) ISBN
80-224-0640-6
Príloha 19: KOČÁREK, P., HOLUŠA, J., VIDLIČKA, Ľ. 2005. Blattaria, Mantodea,
Orthoptera & Dermaptera of the Czech and Slovak Republics. Illustrated key
3. Blattaria, Mantodea, Orthoptera & Dermaptera České a Slovenské
republiky. Ilustrovaný klíč 3. Kabourek, Zlín, 349 pp. ISBN 80-86447-05-7
Ľubomír Vidlička Habilitačná práca
49
Príloha č. 1
VIDLIČKA, Ľ., HUČKOVÁ, A. 1993.
Mating of the cockroach Nauphoeta cinerea (Blattodea: Blaberidae):
I. Copulatory behaviour.
Entomological Problems 24(2): 69-73.
Entomol. Probl. , 24(2): 69 - 73 , 1993. ISSN 0071-0792
Mating of the cockroach Nauphoeta
cinerea (Blattodea: Blaberidae)
I. Copulatory behaviour
lubomlr VIDLICKA, Alica HUCKOVA
Institute of Zoology and Ekosozology, Slovak Academy of Sciences,
Dubravska cesta 9, 842 06 Bratislava, Slovakia
Abstract. The copulatory behaviour of cockroaches Nauphoeta cinerea was observed in details. Four
phases of copulation were identified. The durations of copulation and the duration of other copulatory
events are under defined environmental conditions relatively fixed. Cockroach, copulatory behaviour,
Nauphoeta , Blattodea, etology.
Introduction and overview
Sexual and especially mating behaviour of the cockroach Nauphoeta cinerea have
been observed by many authors (ROTH & WILLIS, 1954; ROTH, 1964; ROTH & BARTH, 1964;
ROTH & DATEO, 1966; FUKUI & TAKAHASHI, 1980; 1983 a,b; TAKAHASHI & FUKUI, 1980; 1983;
1991 ; SRENG, 1979, 1984, 1985, 1990, 1992, 1993). A distinct characteristic of N. cinerea
males is their calling behaviour. Raising the flattened and elongated abdomen the males
expose their sternal glands. From them a volatile sex pheromone, seducin (SRENG,
1990), attracting receptive females from a long distance is emitted. This calling posture
is identical to the aggressive one, described by EWING (1967). However, SRENG (1990)
suggest that the calling posture resembles the posture occupied by dominant males
after winning a combat. That is why dominant males copulate more frequently than
subordinate ones (SCHAL & BELL, 1983; MOORE, 1989; BREED, SMITH & GALL, 1980).
Males recognize the difference between sexes by contact chemoreception, particularly
via antennal contact with the antennae and body of the other individuals.
Intermale contact usually results in agonistic behaviour and characteristic antennal
fencing . Aggressive or subordinate status of cockroach male can by identified on the
basis of the antennaI fencing pattern (EWING, 1967). Mature males are producing a male
pheromone (nauphoetin) in the cuticular wax. This pheromone suppresses the
wing-raising activity caused by hydrocarbons (FUKUI & TAKAHASHI, 1983a; TAKAHASHI &
FUKUI, 1983), but it does not cause aggressive behaviour (SIRUGUE et ai., 1992). SIRUGUE et
al. (1992) suggested the existence of another contact pheromone that triggers the male
agonistic behaviour via antennal contact.
Males in presence of mature females normally display courtship behaviour. During
courtship the male raises his wings and tegmina. The chemical factor (wing-raising
pheromone) respons ible for the wing-raising is present mainly in the hydrocarbon
fraction of the cuticular wax of both sexes (FUKUI & TAKAHASHI, 1983). The receptive
female is attracted to the male's back. She mounts and feeds (licks) on the male's tergal
glands secretion - aphrodisiac sex pheromone (SRENG, 1990), or seducin (ROTH & DATEO,
1966; HARTMAN & ROTH, 1967). The feeding behaviour keeps the female in a proper
position for a period sufficient for bringing her genitalia into the contact with those of the
male. [Type A pattern of mating behaviour - female in upper position (SRENG, 1993)]. The
© Entomological Problems
69
sex pheromone serves as an attractant as well as arrestant (ROTH & DATEO, 1966). As
long as the female feeds on the tergal glands secretion, the male moves backward,
pushes his abdomen telescopically under the female's abdomen and grasps her abdominal
tip with a retractable hook, i.e. right phallomere (R2) (McKITTRICK, 1964). If the
female is willing to mate she allows the contact with the hook. In case of rejecting the
copulation attempt she dismounts from the male and prevents the genital contact
(MOORE & BREED, 1986). Following several unsuccessful mating attempts the male lowers
his tegmina and wings. Standing in a distance of about 2 cm from the female he pumps
and elongates his abdomen and emits a faint but audible sound (HARTMAN & ROTH,1967).
On the other hand, SIRUGUE at al. (1992) observed that if the female did not respond to
the wing-raising of the male, the male resumed the calling posture again, followed by
wing-raising . The females generally cease to respond to the male sex pheromone about
2 days after mating.
When the genital contact with the receptive female was successful, the pair
resumes in the typical false-linear position (end-to-end position) (SRENG, 1990; FUKUI &
TAKAHASHI, 1983b; ROTH & DATEO, 1966) and the copulation starts. During copulation
a spermatophore with non-motile sperm is transferred and firmly cemented into the
genital chamber. Its presence inhibits the receptivity of females. Two to three days after
mating the spermathecae (receptaculum seminis) are densely filled with motile spermatozoa.
The empty spermatophore is extruded by the female several days after mating
(ROTH, 1964).
This paper reports the results of a study of the behaviour of cockroaches during
copulation with special reference to the copulatory behaviour of males.
Material and methods
Observations were made on the cockroach Nauphoeta cinerea . Animals were reared at 29±1 ·C,
4S±1% RH and a 12L:12D light:dark cycle. Food (turkey "grower" diet) and water were provided ad libitum.
Cockroaches were selected from stock colonies maintained at the Institute of Experimental
Phytopathology & Entomology of the Slovak Academy of Sciences. Individuals of both sexes were selected
at the last-instar nymph stage and kept in unisexual groups to last ecdysis. The time of adult ecdysis
was recorded. About 10 adults were placed together in a 13 x 8 x 6 cm plexiglas container with a plexiglas
cover (the same container was used for the observation of mating).
Ethological observations were carried out during the light period. The mature cockroaches mated
14±1 days after the last moult. In the mating container only one male and one female were placed. The
durations of different events of the copulatory behaviour were recorded. Direct observations were used in
most cases. Some mating interactions (10% of observations) were recorded on a videotape using a Sony
HVC-4000 P camera and Sony SL-C9E video cassette recorder and analysed at low speed.
For the mating interruption experiment N. cinerea females were manually separated from their
partners after being in copula for various times.
Results
The mean duration of the mating [the time from the resuming the end-to-end
position to the release of the right phallomere (R2) from the female's body) was 12.5 min
(N=100, SE=0.9).
On the basis of the above mentioned results we interrupted the copulation manually
in minute 4, 6, 8 and 10. When the copulation was interrupted in minute 4 (N=15),
the male's right phallomere stayed pushed out from the male genital chamber and in
minute 9 the spermatophore was extruded. After interrupting the copulation the male
sometimes retracted the right phallomere into the body but in minute 9 the phallomere
was pushed out again and the spermathophore was extruded. In minute 13 the
phallomere was retracted into the male genital chamber.
70
When the copulation was interrupted in minute 6 (N = 15) or 8 (N = 15) the events
were the same as mentioned above.
In case the copulation was interrupted in minute 10 (N = 15), the phallomere stayed
pushed out to minute 13 and then it was retracted. The spermatophore was not
extruded.
The spermatophore contained non-motile and twisted sperm. The diameter
of spirals was 15 - 16 iJm. 2 hours later the sperm began to be active and was about
250 - 350 iJm in length. The frequency of male abdomen contractions and corresponding
movement of tegmina and wings during mating were analysed from videorecordings
using slow motion. The mean frequency of the abdomen contractions was 132 (N=1 0,
SE= 9.1). The strength and frequency of contractions varied throughout mating (fig. 1).
c
0
';:0
u
~
+-'
C
0
u
0
~
OJ
.D
E
::::>
c
30 r---------------------------------------------------------,
25
~o
I 5
I 0
5
2 3 4 5 6 7 8 9101112.13
minute
Fig. 1. Mean number (±SE) of male abdomen contractions during copulation, N=10.
On the basis of our results and previous data the following 4 phases of copulation
can by identified:
Phase 1 Minute 1 to 4 - establishment of the genital contact;
Phase 2 Minute 5 to 8 - completion of the spermatophore, preparation
of the spermatophore transfer;
Phase 3 Minute 9 to 10 - transferring the spermatophore into the female's
bursa;
Phase 4 Minute 11 to 13 - spermatophore firmly inserted and cemented
in female's bursa.
71
Discussion
Different durations of copulation in cockroaches have been reported by different
authors. ROTH (1964) has found mean duration of copulation 17±1 min at 28±2°C, while
MOORE & BREED (1986) 9.5 min at 25-27'C. The closest results to the data reported here
have been found by MOORE (1990 a,b) (11 to 11 .9 min). All above mentioned data were
determined for the initial mating. When males mated 2 or 3 times consecutively, the
average time spent in copula was 100±8 min and 141 ±2 min, respectively (ROTH, 1964).
Dominant males copulated significantly longer than subordinate ones (MOORE & BREED,
1986; MOORE, 1990). The presence of dominant males inhibited the calling posture in
subordinate individuals (SIRUGUE et aI. , 1992). Despite of variability in the duration of
copulation in various experiments it is very important to note that this value is very
constant under the same conditions. The durations of copulation and copulatory events
are fi xed. It was not possible to influence the copulatory events by manual separation of
the mating pair. Timing of spermatophore release and phallomere retraction was the
same as if the female was present.
In the subfamily Oxyhaloinae (Blaberidae) male genitalia are uniform. The hook (R2)
is on the right side (McKITTRICK, 1964; ROTH, 1971). In the family Blattidae (Blatta
orientalis, Periplaneta americana) the hooked titillator of the left phallomere is the first
phallic organ which fastens to the ovipositor to achieve the connection (BAO & ROBINSON,
1990). GUPTA (1947) on the basis of the observation of P. americana mating behaviour
suggested that the titillator was important to keep the female 's vestibulum open and the
right phallomere was the main clasping organ to hold the valvae of the female genitalia
during copulation. According to BAO & ROBINSON (1990) the right phallomere in
B. orientalis keeps the first pair of valvae opened, so that the genital chamber is
exposed . The male gonopore projects into the female genital chamber where the
spermathecal sac and the female gonopore are located. This mechanism probable
explains the fact that phallomere is always extended during the spermatophore release
even in absence of female.
ROTH (1964) was the first who interrupted the copulation of the cockroach
N. cinerea manually and observed the mechanism of the loss of female's receptivity. He
separated females from their partners after having been in copula for 10-12 min (the
average time of copulation was 17 min). In this phase, three different stages of
spermatophore transfer could be identified. In the first case the spermatophores were
not transfered, in the second case the spermatophores were transfered but not cemented
and in the third case they were transfered and firmly cemented. As the duration
of copulation in our experiments was 12.5 min , the events recorded in minutes 7.5-9
could correspond to those observed by ROTH (1964) in minutes 10-12 (at the duration of
copulation 17 min). The timing of behavioural events in phases 2 and 3 explains
variability in termination of spermatophore transfer described by ROTH (1964).
References
BAO, N. & ROBINSON, W.H., 1990: Morphology and mating configuration of genitalia of the oriental
cockroach Blatta orientalis L. (Blattodea: Blattidae). Proc. Entomol. Soc. Wash., 92(3): 416-421.
BREED, M.O, SMITH, S.K. & GALL, B.G ., 1980: Systems of mate selection in a cockroach species with male
dominance hierarchies. Anim. Behav., 28: 130-134.
EWING, L.S., 1967: Fighting and death from stress in a cockroach. Science, 155: 1035-1036.
FUKUI, M. & TAKAHASHI, S., 1980: Studies on the mating behavior of the cockroach Nauphoeta cinerea Olivier
1. Sex discrimination by males. Appl. Ent. Zool., 15(1): 20-26.
FUKUI, M. & TAKAHASHI, S., 1983a: Studies on the mating behavior of the cockroach Nauphoeta cinerea
(Olivier) (Oictyoptera: Blaberidae) III. Isolation and identification of intermale recognition pheromone.
Appl. Ent. Zool., 18(3): 351-356.
72
FUKUI, M. & TAKAHASHI, S., 1983b: Studies on the mating behavior of the cockroach, Nauphoeta cinerea
(Olivier) (Dictyoptera: Blaberidae). Mem. Coli. Agric., Kyoto Univ., 122: 25-36.
FUKUI, M. & TAKAHASHI S., 1991 : The possibility of sex and nymph discrimination by the male cockroach,
Nauphoeta cinerea. J. Ethol., 9(2): 57-66.
GUPTA, P.D., 1947: On copulation and insemination in the cockroach, Periplaneta americana (Linn.). Proc.
Nat. Inst. Science, India. 13: 65-71 .
HARTMAN, H.B. & ROTH, L.M., 1967: Stridulation by the cockroach Nauphoeta cinerea during courtship
behavior. J. Insect Physiol., 13: 579-586.
McKITTRICK, F.A., 1964: Evolutionary studies of cockroaches. Mem. Cornell Univ. N. Y. Agric. Exp. Stat.,
389: 1-197.
MOORE, A.J., 1989: Sexual selection in Nauphoeta cinerea: Inherited mating preference? Behavior Genetics,
19(5): 717-724.
MOORE, A.J. , 1990: Sexual selection and the genetics of pheromonally mediated social behavior in
Nauphoeta cinerea (Dictyoptera: Blaberidae). Entomol. Gener., 15 (2): 133-147.
MOORE, A.J., 1990: The inheritance of social dominance, mating behaviour and attractiveness to mates in
male Nauphoeta cinerea. Anim. Behav., 39: 388-397.
MOORE, A.J. & BREEO, M.D., 1986: Mate assessment in a cockroach, Nauphoeta cinerea. Anim. Behav.,
34: 1160-1165.
ROTH, L.M., 1964: Control of reproduction in female cockroaches with special reference to Nauphoeta
cinerea -I. First pre-oviposition period. J. Insect Physio!., 10: 915-945.
ROTH, L.M., 1971 : The male genitalia of Blattaria. VI. Blaberidae: Oxyhaloinae. Psyche, 78(1-2): 84- 106.
ROTH, L.M. &BARTH, R.H., 1964: The control of sexual receptivity in female cockroaches. J. !nsect Physio!.,
10: 965-975.
ROTH, L.M. & DATEO, G.P., 1966: A sex pheromone produced by males of the cockroach Nauphoeta
cinerea. J. !nsect Physiol., 12: 255-265.
ROTH, L.M. & WILLIS, E.R., 1954: The reproduction of cockroaches. Smithson. Misc. Coli., 122: 1-49.
SCHAL, C. &BELL, W.J. , 1983: Determinants of dominant- subordinate interactions in male of the cockroach
Nauphoeta cinerea. BioI. Behav., 8: 117- 139.
S,RUGUE, D., BONNARD, 0. , LE QUERE, J.-L., FARINE, J.-P. & Brossut, R., 1992: 2-methylthiazolidine and
4-ethylguaiacol , male sex pheromone components of the cockroach Nauphoeta cinerea (Dictyoptera,
Blaberidae): a reinvestigation. J. Chern. Ecol., 18(2): 2261-2276.
SRENG, L., 1979: Pheromones et comportement sexuel chez Nauphoeta cinerea (Olivier)(lnsecte, Dictyoptere).
C.R. Acad. Sci., 289: 687-690.
SRENG, L., 1984: Morphology of the sternal and tergal glands producting the sexual pheromones and the
aphrodisiacs among the cockroaches of the subfamily Oxyhaloinae. J. Morpho!., 182: 279-294 .
SRENG, L., 1985: Ultrastructure of the glands producing sex pheromones of the male Nauphoeta cinerea
(Insecta, Dictyoptera). Zoomorphology, 105: 133-142.
SRENG, L., 1990: Seducin, male sex pheromone of the cockroach Nauphoeta cinerea: isolation, identification,
and bioassay. J. Chern. Ecol., 16(10): 2899-2912.
SRENG, L., 1992: The evolution of cockroach mating behaviors correlated with sex pheromone glands. In
BILLEN, J. (ed.), Biology and Evolution of Social Insects, Leuven University Press, Leuven, Belgium, pp.
223-226.
SRENG, L., 1993: Cockroach mating behaviors, sex pheromones, and abdominal glands (Dictyoptera:
Blaberidae). J. Insect Behav., 6(6): 715-735.
TAKAHASHI, S. & FUKUI, M. , 1980: Studies on the mating behavior of the cockroach Nauphoeta cinerea Olivier
2. Wing-raising stimulant in cuticular wax. App!. Ent. Zool. , 15(2): 159-166.
TAKAHASHI, S. & FUKUI, M., 1983: Studies on the mating behavior of the cockroach Nauphoeta cinerea
(Olivier) (Dictyoptera: Blaberidae) IV. Synthesis and biological activity of nauphoetin and related
compounds. Appl. Ent. Zoo!., 18(3): 357-360.
73
Príloha č. 2
HUČKOVÁ, A., VIDLIČKA, Ľ., KOZÁNEK, M. 1994.
Mating of the cockroach Nauphoeta cinerea (Blattodea: Blaberidae):
II. Histamine changes during courtship and copulation.
Biologia 49(5): 691-695.
Biologia, Bratislava, 49/5: 691- 695, 1994
Mating of the cockroach Nauphoeta cinerea
(Blattodea: Blaberidae)
II. Histamine changes during courtship and copulation
Alica H UCKOVA, Lubomir VIDLICKA , Milan KOZANEK
Institute of Zoology and Ecosozology, Slovak Academy of Sciences, Dubravska cesta 9, SK- 842 06
Bratislava, Slovakia
HUCKOVA , A., VIDLICKA, L., KOZANEK , M. , Mating of the cockroach
NaupilOeta cinerea (Blattodea: Blaberidae) II. Histamine changes during
courtship and copulation.- Biologia, Bratislava, 49: 691- 695, 1994; ISSN
0006- 3088.
The concentrations of biogenic amine histamine were measured in the supraoesophageal
and the 6th abdominal ganglia of the cockroach Nauphoeta cinerea
during courtship and copulation . In both sexes the highest concentration
of histamine in their brains was found in the 5th minute of copulation. In
the last abdominal ganglion of males a sharp increase in histamine level was
recorded in the 5th and 10th minutes, at the time of spermatophore t ransfer
into the female's bursa copulatrix. It suggests the direct involvement of
histamine in the regulation of this process during copulation.
Key words: Cockroach Nauphoeta cinerea, mating behaviour, histamine, central
nervous syst.em, radioenzymatic assay.
Introduction
Biogenic amines are distributed throughout the
nervous systems of both vertebrates and invertebrates
and are generally considered to function
as transmitters, neuromodulators and neurohormones
(GERSCHENFELD, 1973; KLEMM, 1976;
EVANS, 1980; ORCHARD, 1982). There is good
evidence that these compounds play a key role
in moduhtting behaviour and autonomic neural
functions (MURDOCK , 1971; DAVID , VERRON,
1982; MERCER, MENZEL, 1982; MERCER,
ERBER, 1983), they control the circadian rhytmus
(MUSZYNSKA-PYTEL, CYMBOROWSKI, 1978;
PREE, RUTSCHKE , 1983) and are involved in the
insect's reaction to various stress stimuli (DAVENPORT,
EVANS , 1984; KOZA NEK et al. , 1986).
Histamine (HA) has been increasingly implicated
as a putative neurotransmitter in insect visual
systems (HARDIE, 1988; NASSEL et al., 1988;
SIMM ONS, HARDIE, 1988). However, some other
investigations have also shown that the HA level in
insect CNS changes in response to various stress
stimuli (KOZANEK et al. , 1985; HUCKOVA et al.,
1992) and in addition, possible role for HA in reproduction
of cockroach Naupboeta cinerea has
also been suggested (HucKovA et al., 1992).
In a previous study we described the ethological
aspect of the mating behaviour of N. cinerea
(VIDLICKA, HucKovA, 1993). The copulatory act
alone was divided in 4 phases with strict duration
according to the main internal physiological events
e.g. spermatophore completion and its transfer
into the female's bursa.
In this paper we report the changes in HA
concentrations in the brain and the 6th abdominal
ganglion of N. cinerea in relation to the main
copulatory events.
691
'2
!0
i5..
C>
~
(5
E
.s
CII
c:
'E
~
:c:
120
100
80
60
40
20
A ** - males
+females
O~-'---r---'----r---.---.----r~
<?:-<:- <?:-<:- <?:-<:"
. ~. "I:)'
Copulat i on
12
10
'2
S 6
ea.
Cl
.E.(5
6E
.s
CII
c:
'E
.l!!.. 4
:.;:
2
0
B -males
+females
* *
I
~<S i:>~ <?:-<:- <?:-<:- <?:-<:- ",' ",'
o~ ~<o
,.~ ~
Ci CpoS ". ~ . 1:)'
'It~"o
"Copulation
Fig.!. Histamine concentration changes in the brain (A) and last abdominal ganglion (B) of the cockroach
Nauphoeta cinerea during courtship and copulation. Each point represents the mean of 6 animals ± S.E.M. *
(P < 0.05) , ** (P < 0.01) .
M a teria l and meth o d s
Cockroaches were reared at temperature 29 ± 1ac,
relative humidity 50 ± 10% and 12 hours reversible
light/ dark cycle. A semisyntetic food for young
turkeys and water were provided ad lib.
Experimental animals were chosen from the stock
colony just after the last eclosion and separated into
the unisexual groups. For courtship and copulation experiments
ten-day-old males and females divided in 7
groups were used: 1 - control unmated individuals, 2
- females and males during courtship, 3, 4, 5 - couples
in the 1st, 5th and 10th minutes of copulation, 6
- cockroaches just after copulation, 7 - individuals 2
hours after copulation. For courtship and copulation
experiments each couple was placed in a plastic box
8 x 12 x 6 cm. Couples were killed by quick freezing
in a mixture of dry ice and ethanol. Supraoesohpageal
and the 6th abdominal ganglia were dissected on cooled
Petri dish and homogenized in 60- 150 1-'1of solution A
(0.2 mol.l- 1 sodium phosphate buffer pH 7.9:1 mol.l- 1
sodium hydroxide). The homogenates were centrifuged
at 5,000 g for 20 minutes at 4 ac. 10 1-'1 of the supernatant
was transferred into cooled conical vials and
the histamine content was determined radioenzymatically
according to BROWNSTEIN et al. (1974) . In all
692
experiments 0.5 ng of histamine-dihydrochloride in 10
1-'1of 0.1 M HCI was used as an internal standard. The
radioactivity of samples was measured by means of a
Packard scintillation counter.
Protein content was measured in 10 1-'1 of homogenate
according to LOWRY et al. (1951) using
bovine serum albumine as a standard. The data were
analyzed using Student 's t-test.
R esu lts
The pattern of the concentration changes of histamine
(HA) in the supraoesophageal ganglion of
N. cinerea males and females during mating was
very similar (Fig. 1A). In the time of courtship,
when the female climbs the male's abdomen, and
in the beginning of copulation HA level rose only
slightly, this increase was a bit sharper in the
female's than in the male's brain. In the 5th
minute of copulation HA concentration increased
significantly in both sexes (P < 0.01) , reaching
about 3-times higher amounts than were observed
in control unmated groups. In the 10th minute
of copulation the HA level decreased and was ap-
proximately at the same level as at the beginning
of copulation, however, in the males it was still
significantly higher in comparison with the control
group. A slight fall in HA concentration was
also found just after copulation, i.e. in the 13th
minute. After 2 hours the HA level in both sexes
was almost the same as in the control groups or at
the beginning of copulation.
In the last abdominal ganglion (AG) the concentration
of HA during courtship and at the
beginning of copulation was about the same as
in the control groups (Fig. 1B). Then a sharp
rise in concentration was observed in the 5th and
10th minutes of copulation in the male's 6th AG
(P < 0.05) . In terminal ganglion of females the
level of HA also increased reaching the highest
value in the 10th minute, but this rise was not so
distinct as in males. Then a continual decrease of
HA concentration was found in both sexes, and 2
hours after copulation the values were lower than
in the control groups.
Discussion
The critical age for N. cinerea female receptivity
is 3- 4 days (ROTH, 1962). According to SCHAL
et a1. (1984) the preoviposition period for this
cockroach is about 13 days. A virgin females that
has oviposited will not mate as long as an ootheca
is present in uterus. Males may begin to mate
and produce spermatophre when 2- 3 days old. In
respect to this, the ten-day-old individuals we used
in our experiment were at the suitable age.
In cockroaches the neurosecretory system in
the brain serves to release directly copulatory behaviour
(ROTH, BARTH, 1964). Biogenic amines
were investigated in relation to several aspects of
reproductive cycle, e.g. oviposition (HENTSCHEL,
1972, 1975), ootheca production and juvenile hormone
secretion (THOMPSON et a1. , 1990; PASTOR
et a1. , 1991). A great number of investigations
concerning insect reproduction were focused on
the effect of biogenic amines and other active substances
on the musculature of insect reproductory
organs (SOBEK et a1. , 1986; ORCHARD, LANGE,
1987; STOYA, PENZLIN , 1988). It has been shown
that the biogenic amine octopamine (OA) plays a
physiological role in the control of locust oviducts,
which receive innervation from two octopaminergic
dorsal unpaired median neurones located in the
7th abdominal ganglion (LANGE, ORCHARD, 1984;
ORCHARD, LANGE, 1985). OA reversibly reduced
the amplitude of neurally-evoked contractions in
a dose-dependent manner and caused a relaxation
in basal tonus (ORCHARD, LANGE, 1986).
Up to now few investigations have been carried
out on the relationship between biogenic
amine levels and mating behaviour. HODSON
and WRIGHT (1963) reported that application of
monoamines to Periplaneta americana females resulted
in contraction in the abdomen, wing rising
and behaviour similar to that during oviposition.
KozANEK et a1. (1990) investigated OA changes
in the brain of Nauphoeta cinerea during copulation.
They found that the level of this amine
increased during courtship, but during copulation
and in resting cockroaches it decreased.
In our previous research we found significant
cyclic changes in histamine concentration in the
brain and 6th AG of N. cinerea during its adult life
(HUCKOVA et a1., 1992). A possible involvement
of HA in the regulation of the reproductive cycle
in this cockroach was suggested. In this work we
present a relationship between physiological events
during mating of N. cinerea and HA concentration
in their eNS.
The male in the presence of a receptive female
showed characteristic mating behaviour. During
the courtship he palpated with his antennae the
back of the female who then climbed the male's
abdomen and fed on the tergal gland secretes. At
that time the concentration of HA in the brain increased
in both sexes. In the 5th minute of copulation,
i.e. in the 2nd phase, which is characterised
by completion of spermatophore and its transfer
(VIDLICKA, HUCKOVA , 1993), we found a sharp
increase of the HA level in the males' as well as
the females ' brain. Interestingly, at the same time
a significant increase of HA was also observed in
the males' terminal ganglion. According to our
previous ethological observations, the frequency
and strength of the male's abdominal contractions,
which varied throughout mating, was highest in
the 9th minute (VIDLICKA , HUCKOVA , 1993). By
manual interruption of copulating pairs it was revealed
that just in this phase (9th and 10th min)
spermatophore transfer into the female's bursa
took place. In resting couples HA concentration
in the brain stayed approximately at the same
level, while in the 6th AG continually decreased.
Thus, our findings of synchronised HA changes in
the nervous tissue of copulating cockroaches (at
first increase in the brain then in the 6th AG)
supported by ethological observations, allow us to
suppose that this monoamine is probably involved
in regulation of the copulatory process.
To define the precise role of HA in mating
behaviour a complex of electrophysiological, immunocytochemical
and neurophysiological investigations
is needed.
693
Acknowledgements
The authors are grateful to the Slovak Grant agency
for science (grant No 2100994) for partially supporting
this work.
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ORCHARD, I. , LANGE, A. B. , 1987: Cockroach
oviducts: the presence and release of octopamine
and proctolin.- J. Insect Physio!. 33: 26.5- 268.
PASTOR, D., ARTIGAS , F., MARTINEZ, E., BELLES ,
X. , 1991: Brain levels of 5-hydroxytryptamine and
dopamine in Blattela germanica adult females during
the first gonadotrophic cycle.- Biogenic Amines
8 : 109- 114.
PREE, J. , RUTSCHKE, E ., 1983: Zur circadian Rhytmik
des Dopamingehaltes im Gehirn von Periplaneta
americana L.- Zool. Jb. Physiol. 8 7: 455-
460.
ROTH , L. M. , 1964: Control of reproduction in female
cockroaches wit h special reference to Nauphoeta
cinerea - I. First preoviposition period.- J. Insect
Physiol. 10: 915- 945.
ROTH , L. M., BARTH , R., 1964: Control of sexual receptivity
in female cockroaches.- J. Insect Physiol.
10: 965- 975.
SCHAL, C., GAUTIER, J.-y' , BELL, W . J. , 1984: Behavioural
ecology of cockroaches.- BioI. Rev. 59:
209- 254.
SIMMONS , P. J., HARDIE, R. C., 1988: Evidence that
histamine is a neurotransmitter of photoreceptors
in t he locust ocellus.- J . expo BioI. 138: 205- 209.
STOYA , G. , PENZLlN , H., 1988: The effect of different
putative transmitters on spontaneous phasic contractions
of cockroach oviduct.- Zool. Jb. Physiol.
92: 301- 311.
THOMPSON, C. S., YAGI, K. J., CHEN, Z. F., TOBE,
S. S. , 1990: The effects of octopamine on juvenile
hormon biosynthesis, electrophysiology, and
cAMP content of the corpora allata of the cockroach
Diploptera punctata.- J. compo Physiol. B
160 : 241- 249.
VIDLICKA, L .. HUCKOYA , A ., 1993: Mating of the
cockroach Nauphoeta cinerea (Blattodea: Blaberidae)
1. Copulatory behaviour.- Ent. Prabl. 24 (2):
69- 73.
Received May 24, 1994
Accepted June 20, 1994
695
Príloha č. 3
VIDLIČKA, Ľ. 1993.
Seasonal dynamycs of vertical migration and distribution of cockroach
Ectobius sylvestris (Blattaria: Blattellidae: Ectobiinae) in oak forest.
Biologia 48(2): 163-166.
Biol6gia. Bratislava. 48/2: 163 - 166. 1993
Seasonal dynamycs of vertical migration and distribution
of cockroach Ectobius sylvestris (Blattaria: Blattellidae: Ectobiinae)
in oak forest
[,UBOM fR VIDLlCKA·
Department of Zoology. Faculty of Natural Sciences. Comenius University, Bratislava, Slo vakia
VIDLI CKA. L.. Seasonal dynam ics of vertical migrati on and distribution of cockroach
Ectobius sylvestris (B/altaria: Blauellida e: EClObiillac ) in oak forest. Biologia, Brat islava.
48: 163-166.1993.
In 1985-1 986 theseasonaldynami cs of vertica lmigrati on and distribution ofcokroaches
on oa k stems in the Sta te Nature Reseve Lindava (Western Slovakia) was studied.
Distribution of apterous larvae. brachyptero us females and winged males of Ectobius
sylvestriswas observed by means of traps placed in the height of 0.75 m and 4.20-5.20 m.
Migration of Ectobius sylvestris towards tree tops was recorded fro m May to October.
Key words: Ectobius sylvestris, seasonal dynamics. migration . vertical dist ribution.
Species of the genus Ectobius are typical representatives
of cockroaches in temperate climate. They are
abundant in European forests and moorlands (Ma R.
VAN, 1972). Rest ing sites are situated under dead
leaves in the litter, under loose bark or at the foot of
grass. Litter, bushes ferns und grasses are used as
foraging sites (GAUTIER, DELEPORTE, RIVAULT.
1988).
BEY-BIENKO (1%9) investigated the biological
cycle and population density of Ectobius duskei in
grassland on Russian steppe. Seasonality of Ectobius
panzeri has been studied by BROWN (1952). BROWN
(1973) provided information on overwintering stages
of Ectobiuslapponicusand developmental strategies
in Ectobius pallidus (BROWN, 1980). MORVAN'S
(1972) study of four species of Ectobius (E. lapponiCUS,
lividus, panzeri and sylvestris) dealt with their
biologicalcycle. DREISIG (1971) informed about diurnal
migration and vertical distribution of Ectobius
lapponicus. PATOCKA et al. (1%2) recorded the
occurrence of Ectobius lapponicusin tree tops.
Ectobius sylvestris (PODA, 1761) (Blat/aria: Blattellidae)
occurs according to MORVAN ( 1972) along
the borders between forest and mesophil ous neathlands,
dry heathlands and edges of peat bogs. It is
a semivoltine, SOlitaryand oviparous species. This species
has a bisannual development and 5 larval instars.
It overwinters in the 3rd to 4th larval stage.
The aim of this work is to supplement the knowledge
about larval and adult vert ical distribution and
upward migration of Ectobius sylvestris on oak tree
stems.
Material and methods
1. Study area
Our study area was situated in Lind ava State Nature Reserve,
Lindava Fores t (altitude 199-225 m, 80-100 yrs, canopy
cover 85 %) near the village Budmeri ce, map square code
7670, Western Slovakia . The sta nd was evalua ted as
·Present address:
Institute of Experimental Phytopathology and Entomology, Slovak Acad emy of Sciences. 900 28 Ivan ka pri Dunaji,
Slovakia
163
ex·/S traps
2 1
........__._._ .._..__..
18
15
12
9
6
3
0
V. VI.
_ Adulls-Sm
_ Nymphs-Sm
months
(198S)
Fig. 1. Seasonal dynamics of migrat ion of larval and adult cockroach Ectobius sylvestris on oak tree sterns in 1985. Thc traps
were placed in the height of about 5 m.
ex·/S traps
45
40
35
30
25
20
15
10
5
0
_ Adults -Sm
_ Nymphs-Sm
Fig. 2. Seasonal dynamics of migration of larval and adult cockroach Ectobius $)'Ivesrris on oa k tree stems in 1986. The tra ps
were placed in the height of about 5 m.
164
a variant with Impatiens parviflora into the association Poa
nemoralis-« Quercetwn dalechampii (MAlZlAN,1986).
2. Sampling design and methods
Cockroaches were collected by means of tree stem traps
described by FuNKE (1971) and modified by VIDUCKA
(1989). The function of the traps is based on the principle
of positive phototaxis, negative geotropism and silhouette
orientation of insects. Insects were entrapping to the trap
automatically and continuously. The traps were placed on
the stems of oak trees, Quercus dalechampi:
The investigations were performed during 1985 and
1986. In 1985 five traps were installed on the trunks, placed
in the height of 4.20 - 5.20 m. In 1986we used the same five
traps as in 1985 plus an additional sixth one placed in the
height of 0.75 m.
Traps were operating continuously from April to October
1985 and from April to December 1986. The traps were
emptied in regular 2-week intervals, 14 catches in 1985 and
18 catches in 1986. The insects collected in the traps were
killed and fixed in a mixture of 9 mmorl
picric acid and 68
mmorJ
nitric acid and stored in 75 % ethanol.
Material of cockroacheswas collected within the framework
solution of the research work VI-4-2110 at Department
of Zoology.
ex./S traps
40
35
30
25
20
15
10
5
0
Results and disccussion
1. Seasonal dynamics of migration
Our observations in 1985 and 1986 showed that the
larvae of Ectobius sylvestris migrated on oak tree
stems from the beginning of May to the middle of
September (Fig. 1)and from the middle ofMay to the
biginning of October (Fig. 2), respectively. We recorded
the maximum ofupward migration oflarvae in the
end ofJune and in the beginning of August 1985and
in the end of August 1986.In June 1986the increase
of migration was lessexpressive than in 1985. MOR·
VAN (1972) observed larval occurrence in natural habitat
from the middle of March to November.lt is
interresting that we did not record the migration of
larvae in October and November. According to MOR.
VAN (1972) 3rd and 4th instar larvae were abundant
in the nature in thesemonths. This may haveresulted
from unsuitable microclimatic conditions and from
the shortage of food on tree stems in these months.
PATOCKA et at (1962) found out that cockroaches
found their food in tops of trees.
_ Adults-O.75m
_ Ny-nphs-O.75m
Fig. 3. Seasonal dynamics of migration of larval and adult cockroach Ectobius sylvestris on oak tree stems in 1986.The trap
wasplaced in the height 0.75 m.
165
We recorded the adult cockroaches migration
from the middle of May to the middle of July (males
and females) and in October (only females). In May
and June 1985 the migration of adults was very high.
All these data suggest that the dynamics of migration
isevidentlyconnected with the dynamics of occurrence,
microelimaticconditions and presence/absence of food.
2 Vertical distribution
We found out the first time that larval stages and
brachypterous females of the cockroach Ectobius
sylvestris migrate on the stems into the tops of trees.
In both years we observed a much higher migration of
apterous larvae (1985-63 %, 1986-95 % ) than that of
brachypterous females and winged males.
We have not found differences between migration
up to the height 0.75 m and up to the height of
5 m (per 1 trap) (fig. 2,3). It is evident that apterous
larvae, adult females and partly males use the stems of
trees to migrate into the tree tops. This data differ
from data about brachypterous females of Parcoblatta
(GORTON, 1980). Males of this genus are good fliers
and are active in tree tops and bushes, however most
brachypterous females remain on or near the ground.
References
BEY-BIENKO, G. Y., 1969: Ectobius duskci Adel. as a characteristic
inhabitant of steppes in Ihe USSR (Blattoptera),
Soc. Enl. lIal., 28: 123-128.
1"
BROW:-l, E.B., 1952: Observations on Ihe life history of the
cockroach Ectobius panzeri Stephes (Onh., Blattidacs:
Entomologist's mon oMag., M : 20'J-212.
BROW:-;. V.I<.. 1973: Aspects of Ihe reproductive biology of
three species of Ectobius (f)icl)'oplcra: Blal/idac). Em,
exp. & appl.. 16: 213-222.
BROWN, V.I<.. 19R(): Developmental strategies in Ectobius
pallidus (f)iclyoptt:ra: Blauidaev . Int. J. Invert. Rcprod..
2: 85-93.
DREISIG, II., 1971: Diurnal activity in the dusky cockroach,
Ectobius lapponicus L. (II/al/odea). Ent, Scand., 2: 132-
138.
FlINKE, W., 1971: Food and energy turnover of leaf-eating
insects and their infl uence on primary production. Ecol.
Stud.. Anal. Synt., 2: 81-93.
GRAlITIER, J.Y, DELEPORll'. P., RIVAlILT, C.. I'JAA: Rclntionships
between ecology and social behavior in cockroach.
In: The ecol ogy of social behavior, Academic
Press, Inc.. p.335-35I.
GORTOS. R.E.. 1980: A com parativc ecological study of the
wood cockroaches in northeastern Kansas, Univ . K:IRS<I S
Sci. Bull., 52: 221-230.
MAlZtAN, 0 .. 1986: Sirukllira a dynamika artropodocenoz
vo vztahu k ochrane dubovych porastov, Biol6gia, Bralislava,
41: %3-969.
MORVAN, R.. I'J72: ElUde du cycle biologique de quatre
especes d 'Ectobius (BIOI/aria: B1al/d/idac) de la region
de Paimpont (Ille-ct-Vilaine). Bull. Soc. Scient, Brctagne,
47: 257-274.
PAroCKA, J., CAPEK, M.. CHARVAT, K., 1962: Prispevok
k poznaniu korunovcj fauny ~Iankono!cov na duhoch
Slovcnska, predovsetkym so zretclom na rad Lepidoptera.
Biol. prace, Bratislava, 8: 154.
VIDUCKA, r.... 19R9: Poufit ic stromovych [otoeklektorov
a umcleho kmella na zisCovanie migracie chrobakov po
kmelloch stromov. Biologia, Bratislava, 44: 941-952.
Received November I, 1991
Príloha č. 4
VIDLIČKA, Ľ. 1993.
Phyllodromica hungarica sp.nov., a new cockroach species from Hungary
(Insecta: Blattodea: Blattellidae: Ectobiinae).
Entomological Problems 24(1): 63-68.
Ph.yl.to-drorndca hungarica sp.nov., a ne'W
cockroach species front Hungary
(Insecta: Blattodea: Blattellidae: Ectobiinae)
t.ubornlr VIDLICKA
Institute of Experimental Phytopathology and Entomology,
Slovak Academy of Sciences, 900 28 Ivanka pri Dunaji,
Slovakia
Abstract: Phyllodromica hungarica sp. nov., a new cockroach species from Hungary (Insecta: Blattodea:Blattellidae:
Ectobiinae). Entomol. Probl., 24(1):63 - 68, Bratislava, 1993 . ISSN 0071-0792.
A new cockroach species, Phyllodromica hungarica sp. nOY. (Blattellidae) is described. It has been
found only on the terr itory of northern Hungary (BOkk mountains).
Key words: cockroach, Phyllodromica, Hungary, systematics, taxonomy.
Introduction
The genus Phyllodromica FIEBER covers about 80 species wh ich are distributed in
Europe (about 40 species), West Asia, North Africa, the Azores, the Canary Islands and
Madeira (HARZ, 1976).
Studying the material of cockroaches from Hungary deposited in the Zoological
Department of the Hungarian Natural History Museum in Budapest (HNHMB) some
males and females of a new Phyllodromica species were found and described.
This new species is distributed practically on the same zoogeographical area as
Phyllodromica harzi CHLADEK, 1977 and Phyllodromica maculata marani CHLADEK,
HARZ, 1980, wh ich were described from SE-Slovakia.
Description of the species
Phyllodromica hungarica sp. nov.
(Figs . 1-11 ,14,15)
Material examined: Holotype: Male, Hungary: Bukk National Park, Szarvask6, Tardoshegy'
- 350m above sea-level, 25.IX.1981, Adam, HNHMB.
Paratypes: 5f, Bukk N.P., Szarvask6, Tardos-hegy' - 350m above sea - level, 25 .IX.1981 ,
Adam, HNHMB; 1f, Hungary: Bukk NationalPark, Szilvasvarad, Bacs6-v6Igy", 4.V1.1986,
Steinmann, H., HNHMB; 1m, Hungary: Bukk N.P., Fels6tarkany, L6k-v6Igy", 6.V1.1984 ,
Steinmann, H., HNHMB.
hegy = mountain; v61gy = valley
Description : ~: Head (Fig.1) with few setae, interocular width larger than the
ocular width (IW/OW index = 1.7-1 .8). (The interocular width (IW) and ocular width (OW)
were measured at the top of the head.) Pronotum with short, th in setae, widest near the
hind margin (Fig.2). Front femur Type B (Princis, 1965); tarsal claws asymmetrical, arolia
C Entomolog ical Problems
63
well developed. Tegmina (Fig.3), fully developed but reaching only the end of the seventh
abdominal tergum; veins reduced but rad ial vein distinct. Hind wings (Fig.4) strongly
reduced, distal ends bent to the middle at a 1350
angle. Seventh abdominal tergum
strongly modified, separated into two lobes which are directed towards each other.
Glandular pit of tergum 7 (Fig. 11) is hollowed beyond the margins of the opening. The
hollows form a large, bilobed pouch. The footprint-like lobes are joined in the middle with
a narrow strip. The anterior parts of lobes are evidently wider than the posterior parts.
Unlike the species of the beatica group of Phyl/odromica, the pouch of Ph. hungarica sp.
nov. lacks bristles (BOHN, 1992). Subgenital plate asymmetrical (Fig. 9). A single short
stylus with few erect bristles is present only on the left side. Endophallic apodeme
(L2vm) at the anterior end broadened. Near the posterior end of the endophallic
apodeme there is the 'helmet sclerite ' covered with small scales which was also found in
the species of the sylvestris group of Ectobius STEPHENS (BOHN, 1989). Extensible and
retractable hook (left phallomere; L3 of MCKITIRICK, 1964; Fig. 10) on the left side. Cleft
sclerite (R2) and small R3 apodeme on the right side. Cerci composed of 9 segments,
ventral side th in and dorsal side thick pubescent.
Coloration. Head black or dark brown except for pale labrum. Antennae with dark
brown pedicel, the remaining segments pale. Disk of pronotum broad, in the front
angularly rounded, brown, sharply set off from the transparent lateral and anterior
borders. Tegmina brownish-black with yellowish spots (Fig.3). Abdominal terga dark
brown, sterna entirely black. Legs brown with yellow bristles.
Measurements (mm). Body lenght 6.2 - 8.7 ; pronotum lenght x width 2.3 x 3.5;
tegmina lenght x width 6.0 x 2.5; wing lenght 1.7.
Female. Head with few setae, interocular width greater than ocular width (lW/OW index
= 1.6-1 .7). Pronotum as in male. Tegmina (Fig.5) shortened, slightly apically truncated.
Hind wings (Fig.6) reduced, distal end straight or slightly bent to the middle. Genital
structures as in figs. 14,15.
Coloration. Head dark brown except for the pale labrum and the lower part of the
clypeus. Antennae pale brown. Pronotum translucent white, the disk of pronotum dark
brown. Tegmina yellow with big black spots (Fig.5). Hind wings translucent, apex of
wings dark. Abdominal terga and sterna black with white lateral and hind margins . Legs
and bristles yellow.
Measurements (mm). Body lenght 5.7-8.4; pronotum lenght x width 2.3 x 3.5 ;
tegmina lenght x width 3.3 x 2.4 ; wings lenght 1.6.
Etymology: Named after the country, where the type locality is situated.
Geografical distribution: This species is found in the northern part of Hungary.
Systematical remarks: Phyl/odromica hungarica sp. nov. ranks among the species with
the phallomeric "helmet sclerite" and one small stylus. Into this group belong
Phyl/odromica carpentana and relatives, Ph. chopardi, Ph. I/orentae, Ph. maculata, Ph.
pal/ida and relatives, Ph. panteli, Ph. subaptera and the Sylvestris Group of Ectobius
(BOHN, 1989). It is possible to include here also Ph. harzi CHLADEK and Ph. chladeki
HARZ.
64
1 2
3
5
4
6
7
8
Fig s. 1-4. Phyllodrom ica hungarica sp , nov, - male: (1) Head . (2) Pronotum. (3) Teg mina. (4) Hind wing .
Figs. 5-6. Phyllodromica hungarica sp . nov, - female: (5) Tegmina. (6) Hind wing.
Figs. 7-8. Phyllodrimica maculata marani: (7) Male hind wing . (8) Female hind wi ng. Sc ale in mm.
65
12
u
•
•
9
10
13
0.5
Figs. 9-10. Phyllodromica hungarica sp, nov. - male: (9) Subgenital plate (sg; dorsal view) with 2 anterior
apophyses (a), endophaliic apodeme of left phaliomere (ea), helmet sclerite (h), cleft sclerite (cs), apodeme
of right phaliomere (R3; hook removed) and stylus (5). (10) Hook (posterior end) .
Figs. 11-13. Male tergite 7 with gland: (11) Ph. hungarica sp. nov. (12) Ph. maculata marani. (13) Ph. harzi.
Scale in mm.
66
14
Figs. 14-15. Phy/lodromica hungarica sp. nov . - female: Gen ital structures of female. Dorsal complex supraanal
plate (sa), cerci (c). paraprocts (pp). paratergites (pt), valves (v). arm of first valvifer (vt), posterior
lobes of valvi fer II (pi), laterosternite (I). Scale in mm .
Taxonomical remarks and distinctive characteristics: The present new species is similar
to Ph. maculata marani (CHLADEK & HARZ, 1980) and Ph. harzi (CHLADEK & HARZ,
1977). It can be distinguished from Ph. maculata marani especially by the form and
co loration of the rud imentary hind wings (see figs. 4 and 7, 6 and 8) and by the form of
the tergal gland (see figs. 11 and 12). Ph. hungarica sp. nov. differs from Ph. harzi by
tegmi na co loration (Figs . 3,4; see also CHLADEK & HARZ, 1977) and also by the form of
the tergal gland (see figs . 11 and 13) and the hook.
Acknowledgements
I thank Dr. Gy6rgy SZIRAKI from the Zoological Department of the Hungarian
Natural History Museum in Budapest for lending me the cockroach specimens deposited
in th is museum and Mrs BLAHUTIAKOvA for making photographs.
References
BOHN, H., 1989: Revision of the Syivestris Group of Ect obius Stephens in Europe
(Blattaria:Blattellidae) . Ent. scand., 20: 317-342 .
BOHN, H., 1992: Rev ision of the beatica-group of Phyllodromica in Spain (Blattaria:
Blattellidae: Ectobinae) . Ent.scand., 23: 319-345.
CHLADEK, F. & HARZ, K., 1977: Zwei neue Phy llodromica-Arten aus der Siowakei.
Articulata, Bd .l. , 4: 21-24.
CHLADEK, F. & HARZ, K., 1980: Zur Variabilitat der Oothek von Phyllodromica maculata
(Schreb.)(Blattoptera). Articulata, Bd .I., 16: 176-178.
HARZ, K., 1976: Blattoptera. In HARZ & KALTENBACH: Die Orthopteren Europas - The
Orthoptera of Europe 3: 169-305. Dr. W.Junk N.V., The Hague.
67
MCKITIRICK, F.A., 1964: Evolutionary studies of cockroaches. Mem. Cornell Univ.
agric. Exp . Stn. , 389: 1-197.
PRINCIS, K., 1965: Ordnung Blattaria (Schaben). In BEIER : BestimmungsbOcher zur
Bodenfauna Europas. 3: 1-50
Suhrn: Phyllodromica hungarlca sp. nov., novy druh l'ivaba z Mad'arska (Insecta: Blattodea:
Blattellidae: Ectobilnae).
V praci je poplsany novy druh svaba, Phyllodromica hungarica sp .nov. z uzernia severneho
Madarska. Druh bol popfsany na zaklade exernplarov ulo zenych v Zoo log ickom oddeleni Madarskeho
pr frodovedneho rnuzea v Budapest].
Manuscript rec eived : 16.2.1993
68
Príloha č. 5
VIDLIČKA, Ľ. 1994.
Phyllodromica transylvanica sp. nov., a new cockroach species from Romania
and key of the maculata-group of Phyllodromica in central Europe.
Entomological Problems 25(2): 55-62.
Entomol. Probl., 25(2): 55-62, 1994. ISSN 0071-0792
Phyllodrornica transylvanica sp. nov.,
a ne-w cockroach species fro:rn Ro:rnania
and key of the rnaculata-group
of Phyllodrornica in central Europe
(B l a t t a r i a : Blattellidae: Ectobiinae)
l ubomir VIDUCKA
Institute of zoology and ecosozology, Slovak Academy of Sciences ,
Dubravska cesta 9, 842 06 Bratislava , Slovakia
Abstract. A new cockroach species from Romania, Phyl/odromica transylvanica . sp.nov. (Blattellidae)
belonging to the maculata-group of the Phyl/odromica , is described. This group includes also some others
species from central Europe, Le. Ph. maculata (SCHREBER), Ph. chladeki H ARZ, Ph. harzi CHLAOEKand Ph. hungarica
VIOUCKA. A key to the maculata-group in central Europe is given.
Key words. Cockroach, Phyl/odromica , Romania, systematic, taxonomy.
Introduction
BEy-BIENKO (1950) divided the species of the genus Phyllodromica into 4 subgenera and
subgenus Phyllodromica into 5 groups (1. marginata, 2. megerlei, 3. maculata, 4. brevipennis,
5. turanica) . He presented some characteristic features for the maculata-group: forewings of
males fully developed; females with shortened forewings; the costo-radial area of forewings
with distinct , more or less regular oblique veins; abdomen of the males strongly specialised;
6th abdominal tergite strongly depressed; 7th tergite elongated ; anterior part of the 7th tergite
with big angled elevation; furrow at the posterior part dividing the tergite into two lobes;
hypandrium with extraordinarily small stylus.
BEy-BIENKO (1950) includes in this group apart from one central European species (Ph.
maculata SCHREBER, 1781) 6 other species come from Crimea and Caucasus - Ph. adusta
(FISCHER OE WALOHEIM, 1846), Ph. polita KRAUSS, 1888, Ph. retowskii (KRAUSS, 1888), Ph.
schelkovnikovi BURR, 1913, Ph. kiritschenkoi BEy-BIENKO, 1948 and Ph. euxina BEy-BIENKO,
1950.
CHLAoEK & HARZ (1977) described two other species (Ph. harzi CHLAoEK, 1977 and Ph.
chladeki HARZ, 1977) from Slovakia classifie d within the maculata-group. The species Ph.
hungarica VIDLICKA, 1993, which also belongs to this group, was found in north-eastern
Hungary (VIOLICKA, 1993). In this paper a representative of the maculata -group, Ph. transylvanica
sp.nov. from Romania is described. Till now, the species found in Hungary and
Romania have probably been confused with Ph. maculata .
CHL'o'OEK & HARZ (1980) divided Ph. maculata into two subspecies - Ph. maculata
maculata (SCHREBER, 1781) and Ph. maculata marani CHLAoEK & HARZ, 1980. It is possible ,
however, that the species Ph. maculata included a complex of closely related species . In this
paper a key to the identification of the central European species of the maculata-group is
given.
© Entomological Problems
55
Description of the species
Phyllodromica transylvanica sp. nov.
(Figs 1A-C, 3E-H, 5C-D)
Material examined. Holotype, 0 , Romania, Cluj, 1.7.1966, leg. B. Kis, (terminalia on slide, Hungarian Natural
History Museum, Budapest , Hungary).
Paratype: 2 , Romania, Cluj, 1.7.1966, leg. B.Kis.
Description: Male: Head with few thin setae , interocular width larger than the ocular width
(lW/OW index = 1.9-2.0). Pronotum with short, thin setae , widest near the hind margin . Front
femur of type B (BEY-BIENKO, 1950) ; tarsal claws weakly asymmetrical, arolia well developed.
Forewings fully developed, distal end rounded , radial vein and veins in coste-radial area well
developed (Fig. 3E). Hindwings strongly reduced , distal end pointed (Fig. 3F). Posterior part of
6th tergite in the centre strongly depressed (Figs 1A, B). Posterior border of 7th abdom inal
tergite with deep angular excision , anterior part connected with rounded rampart (Figs 1A, 5C)
in the centre. Hollows of tergal gland form a large, bilobed pouch (Fig. 5D). Subgenital plate
asymmetrical. Small hill-like stylus with a few erected bristles well developed on the left side of
the subgenital plate. Right stylus strongly reduced (Fig. 1C). Endophallic apodeme (L2vm)
broadened at the end. Helmet sclerite , cleft sclerite (R2) and R3 apodeme present on the right
side, retractable hook (L3; Fig. 1D) on the left side. Cerci composed of 9 segments.
Colouration. Head dark brown or black except pale labrum . Antennae brown. Pronotum with
broad , dark brown or black disk, lateral and anterior margins transparent. Forewings brown
with yellowish anterior marg in (Fig. 3E). Hindw ings pale. Abdominal tergites dark brown ,
sternites black. Legs brown with yellow spines .
Size (mm). Body length 8; pronotum length x width 1.9 x 3.3; forew ing length x width 6 x 2.25;
hindwing length 1.5.
Female. Head with few setae , interocular width larger than ocular width (IW\OW index =
1.8-1 .9). Pronotum as in male. Forewings (Fig. 3G) shortened, not reaching the end of the 3rd
tergite. Hindwings reduced, narrow, pointed (Fig. 3H). Front femur of type B.
Colouration. Head dark brown except pale labrum. Antennae yellowish brown to brown . Disk
of pronotum dark brown , margins of the pronotum white , translucent. Legs yellowish brown to
brown. Forewings yellow with large dark spot. Hindwings pale. Abdominal terg ites and
sternites dark with white lateral and hind margins.
Size (mm). Body length 5.8-8.2; pronotum length x width 1.9 x 3.6; forewings length x width
3.2 x 2.4; hindwings length 1.5.
Etymology. Named after the part of Romania where the new species has been found .
Geographical distribution. Found only in the surroundings of Cluj in Roman ia.
Systematical remarks. The maculata -group is best characterised by the morphology of tergal
gland and bilobed pouch of the male (Figs 4, 5). The shape of the tergal gland and the bilobed
pouch is very similar in Ph. maculata maculata , Ph. maculata marani and Ph. hungarica. The
structure of the tergal gland of Ph. chladeki and Ph. transylvanica is somewhat different. In
these species the 7th tergite is not fully divided into two lobes. The anterior parts of the tergite
7 remain connected. Ph. harzi is a trans ition between the two groups - 7th tergite is divided
into two lobes, but in the anterior and posterior parts there is a connection between the lobes.
Key for the identification of males of the central European
species of the maculata - group
1. Apex of the hindwings clearly bent to the middle (Figs 2J, 3J) 2
- Hindwings more or less straight, the apex not bent 3
56
lA
_ . _ . ~ .
....~~~~..-.....
-"/.i;"'!"' " ' . ..•.,-,;,;~>-
OC~~"
0.5
o
Fig. 1. Ph. trensylvenice, male. (A. B) abdomen with gland opening. (A) dorsal view. (B) ventral view. (t5-t8)
tergites 5-8; (e) subgenital plate with stylus (s): (D) hook - posterior end. Scale in mm.
2. Dark spot in the apical third of the forewings (Fig. 31); hindwings suddenly narrow in the
apical third, black only at the tip (Fig. 3J); tergal gland as in Figs 5E, F. Occurrence: Slovakia
- Slovensky Kras mountains Ph. harzi CHLAoEK, 1977
- Dark spot nearly extending over the whole length of the forewings (Fig. 21); hindwings not
suddenly narrowed, apical third dark (Fig. 2J); tergal gland as in Figs 4E, F. Occurrence:
Hungary - BOkkmountains and surroundings Ph. hungarica VIDLICKA, 1993
3. Tergite 7 in the middle of the anterior border not divided into two lobes 4
- Tergite 7 separated into two lobes 5
4. Narrow dark spot, at most slightly surpassing the apical half of the forewings (Fig. 3A),
forewings tricoloured (yellow-brown-black); the hindwings almost entirely dark (Fig. 3B);
tergal gland as in Figs 5A, B. Occurrence: Slovakia - Muranska planina plateau .
......................................................................................................... Ph. chladeki HARZ, 1977
57
0.25oc0.251.0
0.25~
GF
1.0'----"'
0.25L0.25'-------'
J1.0L------..I
Fig. 2. Forewings (A, C, E, G, I, K) and hindwings (B, 0 , F, H, J, L) of the Phyllodromica species. (A-D) Ph.
maculata maran i, (A, B) male, (C, D) female; (E- H) Ph. maculata maculata, (E, F) male, (G, H) female; (I-L) Ph.
hungarica, (I, J) male, (K, L) female. Scale in mm.
58
D1.0~
cB
0.25HGF
0.251.0K0.25~
J
Fig. 3. Forewings (A, C, E, G, I, K) and hindwings (B, 0 , F, H, J, L) of the Phyllodromica species. (A-D) Ph.
chladeki, (A, B) male, (C, D) female; (E-H) Ph. transylvanica, (E, F) mele, (G, H) female: (I-L) Ph. harzi, (I, J)
male, (K, L) female. Scale in mm.
59
4A
c
Fig. 4. Tergites 7 of males with tergal glands. (A, C, E) dorsal view, (B, 0, F) ventral view; (A, B) Ph. maculata
marani; (C, 0) Ph. maculata maculata ; (E, F) Ph. hungarica. Same scale (in mm) for (A- F).
60
SA 0.5
B
Fig. 5. Tergites 7 of males with tergal glands. (A, C, E) dorsal view, (B, 0 , F) ventral view; (A, B) Ph. chladeki;
(C, 0) Ph. transylvanica ; (E, F) Ph. harzi. Scale same (in mm) for (A-F).
61
Dark spot as long as the forewings (Fig. 3E); the hindwings almost entirely pale, transparent
(Fig. 3F); tergal gland at the posterior border of the opening with bent rampart (Figs 5C, D).
Occurrence: Romania - Transylvania Ph. transylvanica sp. nov.
5. Forewings black with narrow whitish lateromarginal bands (Fig. 2A); hindwings dark with
widely rounded apex (Fig. 2B); tergal gland as in Figs 4A, B. Occurrence: middle and
eastern Slovakia, north-eastern Hungary, Poland ..
.............................. Ph. maculata marani CHLADEK& HARZ, 1980
Pale forewings with two clearly separated dark spots (Fig. 2E)(light-coloured individuals
with inconspicuous spots); hindwings only in the apical third dark, apex narrowly rounded
(Fig. 2F); tergal gland as in Figs 4C, D. Occurrence: western Slovakia, Austria, northwestern
Hungary, Bohemia, Germany, Poland, Romania ..
............................................................................... Ph. maculata maculata (SCHREBER, 1781)
Key for the identification of females of the central European
species of the maculata - group
1. Forewings with one large spot (Figs 2C, K, 3E ) 2
- Forewings with 2 (exceptionally one) small spots 4
2. Apex of the hindwings black, wings narrow (Fig. 2L); forewings as in Fig. 2K. Occurrence:
Hungary - BOkkmountains and surroundings Ph. hungarica VIDLICKA, 1993
- Apex of the hindwings without black spot 3
3. Hindwings narrow, pointed (Fig. 3H); forew ings subtruncate, dark spot less extended,
yellow stripe along posterior and apical margin broader (Fig. 3G). Occurrence: Romania Transylvania
Ph. transylvanica sp. nov.
Hindwings broad (Fig. 2D); forewings broadly rounded with a large dark spot leaving only a
narrow yellow stripe along the posterior and apical margin (Fig. 2C). Occurrence: middle
and eastern Slovakia, north-eastern Hungary, Poland ..
................. Ph. maculata marani CHLADEK& HARZ, 1980
4. Apex of the hindwings black (Fig. 3L), forewings with two small spots at the anterior and
posterior end (Fig. 3K). Occurrence: Slovakia - Slovensky Kras mountains ..
......................................................................................................... Ph. harzi CHLADEK, 1977
Apex of hindwings not darker than remaining part 5
5. Forewings obliquely truncated, tricoloured (Fig. 3C); hindwings with pale spots at the base
(Fig. 3D). Occurrence: Slovakia - Muranska planina plateau .......... Ph. chladeki HARZ, 1977
- Forewings obliquely rounded with 2 larger or smaller spots (light-coloured individuals
sometimes wholly without spots) (Fig. 2G); hindwings without pale spots at the base (Fig.
2H). Occurrence: western Slovakia, north-western Hungary, Austria, Bohemia, Germany,
Poland, Romania Ph. maculata maculata (SCHREBER, 1781)
Acknowledgements
I thank Dr. Gyrgy Sziraki from the Zoologic al Department of the Hungarian Natural History Museum in
Budapest (Hungary), Dr. Bela Kis from University "Babes-Bolyai" in Klausenburg (Romania) and Dr. Oto Majzlan
from the Department of Biology and Pathobiology of Pedagogical faculty of Comenius University in Bratislava
(Slovakia) for lending me the cockroach specimens. I am deeply grateful to Dr. Horst Bohn from Zoologisches
Institut der Universitat MOnchen (Germany), for critically reviewing my manuscript.
References
BEY-BIENKO, G. JA., 1950: Nasekomyie Tarakanovyie. Fauna SSSR. N. Ser., 40: 1-342.
CHLADEK, F. & HARZ, K., 1977 : Zwei neue Phyllodromica - Arten aus der Siowake i. Articulata, Bd.I., 4: 21-24.
CHLADEK, F. & HARZ, K., 1980 : Zur Varlabllitat der Oothek von Phyllodromica maculata (Schreb.). (Blattoptera) .
Articulata, Bd.l., 16: 176-178.
VIDLICKA, 1:., 1993: Phyllodromica hungarica sp. nov., a new cockroach species from Hungary (Insecta: Blattodea:
Blattellidae: Ectobiinae). Entomol. Probl., 24(1): 63-68.
Manuscript received: 28. 9. 1994
62
Príloha č. 6
VIDLIČKA, Ľ., MAJZLAN, O. 1997.
Revision of the megerlei - group of the cockroach genus Phyllodromica Fieber
(Blattaria: Blattellidae, Ectobiinae).
Entomologica Scandinavica 28: 163-173.
Revision of the megerlei-group of the cockroach genus
Phyllodromica Fieber (Blattaria: Blattellidae, Ectobiinae)
L'UBOMIR VIDLICKA and OTO MAJZLAN
Ent. scand. Vidlicka, L'. & Majzlan, 0 .: Revision of the megerlei-group of the cockroach genus Phyllodromica
Fieber (Blattaria: Blattellidae, Ectobiinae). Ent. scand. 28: 163-173. Copenhagen,
Denmark. August 1997. ISSN 0013-8711.
The species of the megerlei-group are characterised by shortened wings with typical scattered
dark spots and large simple glands on tergite 7 without inner structures . Two of the species
are widely distributed from south-eastern Germany to Romania and Ukraine (P. megerlei
Fieber) or from Turkey to Syria (P. asiatica Bey-Bienko), the third is endemic for eastern
Bulgaria (P. pulcherrima sp. n.). The characteristics of the megerlei-group are described and
the relationships with the tyrrhenica-group are discussed.
L' .Vidlicka, Institute of Zoology SAS, Diibravska cesta 9,84206 Bratislava, Slovakia.
O. Majzlan, Department of Biology and Pathobiology, PedFCU, Moskovska I, Bratislava,
Slovakia.
Introduction
Phyllodromica megerlei (Fieber, 1853) was originally
described by Charpentier (1825) as Blatta
punctata. Since Eschsholz (1822) applied same
name to another cockroach species (now Diploptera
punctata) Charpentier's name is a homonym
and the name suggested by Fieber is commonly
used at the present time. Princis (1971) discerned
five subgenera within the genus Phyllodromica. P
megerlei is the type species of the genus Phyllodromica
and the subgenus Phyllodromica. BeyBienko
(1950) divided the subgenus Phyllodromica
into 5 groups. The megerlei group included
only one species - P megerlei. The second one
was described by Bey-Bienko (1950) as a subspecies
P megerlei asiatica. Ramme (1951)
described the same species as P megerlei f. eryth-
ronota.
Collections made by the second author in Bulgaria
have revealed one new species (P pulcherrima)
in this group.
Specimens used in this study are deposited in
the following collections:
BN - Collection of Dr Barnabas Nagy, Budapest,
Hungary.
L'V - Collection of first author.
© Entomologica scandinavica (Grp.10)
HNHM - Magyar Termeszettudornanyi Miizcum,
Budapest, Hungary ; Dr Gyorgy Sziraky,
MMG - Matra Muzeum, Gyongyos, Hungary ; Dr
Tibor Kovacs.
MNB - Museum fur Naturkunde, Berlin, Germany ;
Dr Kurt K. GUnther.
NMP - Narodnf muzeum v Praze, Prague, Bohemia;
Dr Ivo Kovar.
NMW - Naturh istorisches Museum zu Wien, Vienna,
Austria; Dr Ulrike Aspock.
OM - Collection of second author.
PFUK - Prfrodovedecka fakulta Univerzity
Komcnskeho, Bratislava, Slovakia; coli. Dr
Jan Gulicka.
SNMB - Slovenske narodne rmizeurn - Prfrodovedne
rmizeum, Bratislava, Slovakia; Dr Ilja oisu.
55MB - Stredoslovenske muzeum, Banska Bystrica,
Slovakia; Dr Tomas Kizek.
Characteristics of the megerlei-group
Bey-Bienko (1950) was the first to attempt a classification
of Phyllodromica species living in the
former Soviet Union and adjacent territories. His
classification is based on differences in the degree
of tegminal development, on different venation
and on the structure of the male tergal gland. The
last character is most important also for the division
of the closely related genus Ectobius into
various groups (Failla & Messina 1978).
164 vuua« L. & Majzlan , O. ENT. SCA ND. VOL. 28:2 (1997 )
I/
)
D
Plate I. Phyllodromica spp., habitu s: (A) P. megerlei , male; (B) P. asiatica, male; (C) P. pulcherrima, male; (D) P.
pulcherrima, female (orig . P. Kuliffay).
Description. - Wings: Forew ings in males reaching
to the middle of the fourth abdominal tergite
or at most to the end of abdomen, in females more
strongly shortened, reaching to the second
abdominal tergite, with typical irregular net venation,
the costo-radial area without oblique veins ;
hindwings rudimentary. Coloration: Whitish to
transparent with different small black or brown
spots, subcostal area sparsely or not at all spotted.
Legs: Front femur Type B2; pulvilli present on the
1-4 proximal tarsomeres of all legs, tarsal claw s
asymmetrical (the posterior claw is longer than
the anterior), arolia well developed.
Males. Tergite structures: Posterior margin of
ENT. SCAND. VOL. 28:2 ( 1997)
sixth abdominal tergite mediall y deeply excavated,
elevated dorso-medially, in its normal position
overlapping part of the glandular pit. The glandular
pit on tergite 7 simple, transversely oval, inner
structures absent. Surface of the pit with sparsely
dispersed long bristles with curved tips, the bristles
in the center are more densely distribut ed.
Genitalia: Visible part of subgenital plate slightly
asymmetrical, broadly triangular, apex rounded,
with a single style located to the left of the midline.
Stylus well developed , relatively large. The
end of the stylus curved and covered with dense
short microtrichae (Figs IG, 31, 4E). Right stylus
only indicated. Retractable hook (L3) slender
(Figs IH, 3K, 4D), endophallic apodeme at the
anterior end broadened, cleft sclerite (R2) present.
Females. Genitalia (Fig. 3L): The valves narrow,
bent; posterior lobes of valvifer conical with
tip on the posterior end; paratergites short. Coloration:
The pronotum with yellowish white to
transparent borders and with black disk. Sternites
black or with narrow whitish lateral borders, tergites
black with narrow light posterior borders.
On the lateral margins of tergites 5-7 bigger whitish
spots. Legs in male brown to black with yellow
regions at the coxa-trochanter jo int and at the
tarsi, tibial spines and sometimes posterior end of
tibia yellow. Legs in female brown to yellow, tibial
spines yellow.
Systematic relationships
The lIlegerlei-group is very closely related to the
tyrrhenica-group (P. tyrrh enica, P. clavata, P.
pavani ). Similar characters are: the simple form
of the glandular pit; a well developed stylus on
the left side with dense short bristles, and the
coloration of the forewings. The distinguishing
character is the venation of the costo-radial area:
in the lIlegerlei-group obliqu e veins are absent.
The floor of the glandul ar pit is covered only with
a small number of bristles and the absence of other
structures in the glandular pit in both megerleiand
tyrrh enica-groups suggest their partial separation
from the remaining groups of the subgenus
Phyllodromica.
Key to males of the megerlei-group
I . Forewings strongly reduced, reachin g at most
to the middle of the fourth abdominal tergite
................................................. P. pulcherrima sp. n.
- Forewin gs reachin g to the end of abdomen 2
Revision of the Phyllodromica megerlei-group 165
2. Disk of the pronotum black; sixth tergite with
small, elongated whitish to tran sparent spots
on the posterior lateral margins .. P. megerlei Fieber
- Disk of the pro notum orange -yellow (ce ntral
part) to yellowish (posterior part and corne rs);
sixth tergite dark, without whitish to transparent
spots on the posterior lateral margins
............................................. P. asiatica Bey-Bien ko
I. Pltyllodromica megerlei (Fieber, 1853)
(Plate I: A; Figs IA-H , 2, 4G, 5)
Blatta punctata Charpentier, 1825: 77 (nee Esc hsc holtz,
(822).
Blatta (Phyllodromica) megerlei Fieber, 1853: 94.
Aphlebia punctata (Charpentier): Brunner v. W. 1865:
71; 1882: 41 ; Jakobson & Biank i 1905: 126.
Hololampra megerlei (Fieber): Bazyluk 1956: 28.
Phyllodromica megerlei Fieber: Rehn 1903: 266; BeyBienko
1938: 26; 1950: 232; Prin cis 1971: 1094;
Harz 1976: 276; Bazyluk 1977 : 103.
Hololampra punctata (Charpenti er) : Kirby 1904 : 69;
Shelford 1907 : II ; Harz 1957: 34; 1960: 17.
Phyllodromica punctata (Ch arpentier) : Us & Matvejev
1967 : 8.
Type material. - Missing.
Material studied. - SLOYAKIA: Bucany, 150 m, I 0',
22.v.1994, L' . Yidlicka (L' Y); Piest' any, 160 m, 2 0' , 2
9, 16.v.1992, L'.Yidlicka (L'Y); Nova Bosaca, 350 m, 1
0' , 19, 30.v.1993, O. Majzlan (L' Y); Jakubov, 140 m, I
0' , 22.iv.1994, L' . Yidlicka (L' Y); Horne Plachtince, I 9 ,
30.v.1990, o. Majzlan (OM) ; Modry Kamen, 1 9 ,
13.vi.1990, Bitusfk (SSMB); Plavecke Podhrad ie, 190
m, 1 9, 15.v.1959, I. LobI (SNMB), 1 0', 23.v.1994, L' .
Vidlicka (L' Y); Devin, Mt. Devfnska Kobyla, 250 m, 2
0' , 24.iv.1994, L' . Yidlicka (L' Y); Brati slava, YICie
hrdl o, 130 m, I 0', 16.v.1992 , o. Majzlan (OM); Bratislava,
1 0' , 14.vi.1992, O. Majzlan (OM) ; Ivanka pri
Dun aji, 130 m,) 0', 18.v.1992, L' . Yidlicka, I 9,
30.vi.1992, K. Sm idakova, 3 9, 19.v.1993, I 9,
2I.v.1993, L' .Yidlicka (L' Y); Kalinkovo, 130 m, I 9,
Il.vi.1 975, I. Okali (SN MB); Oremov Laz - Lest' , I 9,
27.vi.1958, J.Gul icka (PFUK); _Ma lacky, I 9,
15.vi.1958, J.Guli cka (PFUK); Jur - Sur, pan6nsky haj,
I 9, 8.ix.1957, J.Gul icka (PFUK); Studienka, 5 0', 10 9,
10.vi.1996, L' . Yidlicka (L' Y); Vel'ke Levarc, I 0' ,
10.vi.1996, L' .Yidl icka (L' Y); Tisovec, I 9, 15.vi.1996,
V. Jansky (L' Y); Zahorie, Rohoznfk, Obora, I 9,
4.vi.1 969, L.PospiSilova (MMB); Cajkov, 1 9,
5.vi.197I , 1.0kali (SNMB); CZECH REPUBLI C (part
MORAYIA): Pavlovske kopce, 1 9, 2I.vii.1 957 , 10 0' ,
37 9, 25.vi.1962 (NMP); Pouzdrany, I 0', 2 9 ,
26.vi.1962 (NMP); AUSTRIA: MOdling, 2 0' , 2 9, H.
TUrk, 2 9, 1862 (NMW) ; Eichkogl bei MOdling, I 0',
30. v.1909, I 9, vii.1912, I 9, 22.vii.1921, 2 9 ,
7.vi.1950, R. Ebner (NMW); Guntramsdorf, 1 9, R.
Ebner (NMW) ; Gurhofgraben bei Aggsbach, Wachau,
Holdhaus, 1 9, 30.v. 1909 (NMW); Hackelsberg, Burgenland,
1 9, Koisi (NMW) ; Hermann skogel, I 9,
28.vii. 1907, Karny (NMW); Herzogenburg, I 0', before
1930, L. Mader (NMW) ; Ma uer, 20', 6 9, Brunn er v.
w., I 0', 1870, TUrk (NMW) ; Plank am Kamp, I 9,
vii.65, I 9, 12.v.1948 (NMW) ; Purgstall, I 9,
166 vidlicka, L'. & Majzlan. O.
17.vii.1955 , Sandbruch (NMW); Sulzer Berg bei Wien,
I nymph, 25.v.1949 (NMW); surroundings Vorau, I 0',
before 1952, H. Fran z (NMW); St. Veit bei Wien, 20', 4
9 (one with ootheca), Brunner v. W. (NMW); Wien,
Weidling , I 0', I 9, before 1946, L. Mader (NMW);
Wien, Lainzer Tiergarden, 500 m, 2 0', I 9, 26.v.1951, 2
nymphs, 20.v.1950, I nymph, 4.x.1951, R. Ebner
(NMW); Eichkogl, I 9 with ootheca, 18.vi.1908
(MNB); Austria, Kat.-Nr.3724, coli. Schaum, 1 9
(MNB); HUNGARY: Nagyborzsony, Hosszii -volgy, I
9, 27-3I.vii.1975, J. Jablonkay & A. Varga (MMG);
Szokolya, Szenpatak-volgy., 300 m, I 0', 3 9, 27.vi.
1965, B. Nagy (BN) ; Szokolya, Szenpatak-volgy,
300m, I 9 (HNHM); Budapest, Csucs-hcgy, 1 9,
26.vi.1983, B. Nagy (BN) ; Cserepfalu, Also-Csakany, I
9, Il.vi.1984 , Merkl, Korsos (HNHM); Balatonszepezd,
150 m, 10', 24.v.1990, B. Nagy (BN) ; Pilisszentkereszt,
Dobogoko, 680m, 1 9, 16.vii.l961, B. Nagy
(BN) ; Hcgyhatszcntrnarton, I 9, 2.vii.1982, B. Nagy
(BN); Hortobagy, Ujszentmargita, I 0', 9.iv.-8.v.1974,
Kaszab, 60',49, 9.v.-ll.vi.1974, Kaszab (BN) ; Keszthely,
Koponar-teto, I 9, 28.vi.1994, B. Nagy (BN) ;
Korosujfalu, I 9, 10.viii. 1980, B. Nagy (BN); Koszeg,
Also-erdo, I 9, 13.vii.1937 , Visnya (HNHM); Hortobagy,
Ohati-erdo, I 0', 3 9, I nymph, 15.v.1947 , B.
Nagy (BN); Tard, Baba-volgy, I 9, I.vi .1959 , S. T6th
(HNHM); Harornhuta, Istvan-kilt, I 9, 6-12.vi.
1955, Kaszab , Szekcssy (HNHM); Pomaz, I 9, 2.vii.
1978, B.Nagy, (BN); Szentendre, 2 9, 8.vii.1962, B.
Nagy (BN) ; Ujszentmargita, 3 0', I 9, 26.v.1966, B.
Nagy (BN); ROMANIA: Aiud, 1 9, 30.v.1909, 2
nymphs, 4.iii.191O, I. Nagy (HNHM), 29, 25.vi.1961,
B. Kis (HNHM); Arcalia, 10', I 9 with ootheca, 16.vi.
1976, B. Kis (HNHM); Baisoara, I 9, 16.vii.1962, B.
Kis (HNHM); Caraorrnan, 2 9, 27.vii.1967, B. Kis
(HNHM); Cluj, Mt. Szaf, I 9, 22.vi.l943, Kolosvary
(HNHM); Cluj, Bacsie, I 9, 22.vi .1943, Kolosvary
(HNHM); Cluj, I 9, 10.vii.1962 , 29, 28.v.1964, I 9,
10.vii.1969, 2 9, 5.vi.1977 , B. Kis (HNHM); Cluj,
Manastur, 1 0', 12.v.1963 , B. Kis (HNHM); Ciurtuci, I
9, 5.vii.1972 , B. Kis, (HNHM); Giurcuta de Jos, 2 9,
4.vii.1972 , 29, 8.vii.1972 , B. Kis (HNHM); Deva , 3 0',
7 9, 25.v.1959 , B. Kis (HNHM); Dezmir, I 0', I 9,
18.vi.1977, B. Kis (HNHM); Foeni, 19, 9.vii.1983 , B.
Kis (HNHM); Cheile Turzii, 1 0', 24.v.1964, B. Kis
(HNHM); Mociu, 10', I 9, 7.v.1976, B. Kis (HNHM);
Mraconia, 2 0', I 9, 12.vi.l968, B. Kis (HNHM);
Talagiu, I 9; Transilvania, I 0', I 9, Pungar, 2 0', I 9
(NMW); Heltau, I 9, 15.vi.1924, A. MUller (MNB);
Hammersdorf, 19, lO.v.I923, A. MUller (MNB);
Reghin, 19, 23.vii.1917, A. MUller (MNB); UKRAINE:
Cherson, 19, lO.v.1903, Ewert (MNB).
Literature records. - SLOVAKIA: Banska Stiavnica
(Petricsk6 1892); Streda nad Bodrogom (Chladek
1965); Vinne (Gulicka 1967), Plesivecka planina (Chladek
1988); Muranska planina Mts: Tesna skala, Kl' ak
(Cejchan 1992); Silicka planina, Koniarska planina
(Chladek 1994); CZECH REPUBLIC: Chot'ovice, Tfemosnice,
.Pal'fzov (Cejchan, 1959); Havlfckiiv Brod,
Lednice, Zelesice (Chladek 1965); HUNGARY: Budaihegyscg
(Frivaldszky 1867); Sarospatak, Satoraljaujhely,
Budapest (Chyzer 1897, Pungur I~OO); Koszegihegyscg
(Pongracz 1940); Egyek, Ujszentmargita
(Nagy 1983); ROMANIA: Oradea - Jud. Crisana (FriENT.
SCAND. VOL. 28:2 (1997)
valdszky 1867); Cluj , Zsul de Cimcie, Baita, Marpod
(Herman 1871); Baile I Mai (Mocsary 1871); Nusfalau,
Zalau, Fetindia, Fagetul, Girceiu, Jac (Pungur 1891);
Transilvania (Redtenbacher 1900); Babadag (Jaquet
1903) ; Marpod, Cluj , Zsul de Cimpie, Baita, Zalau,
Nusfalau, Hunedoara, Reghin, Ocna Sibiului (MUller
1924) ; Zalau, Reghin, Nu~falau, Sibiu, Clui (Knechtel
& Popovici-Biznosanu 1959) ; AUSTRIA: Pitten (Handlirsch
1889); Kalksburg, Mauer, Modling, Hainbach,
Bruck a.d. Leitha (Redtenbacher 1900); Wien (Brunner
v. W. 1885; TUmPlil 190I); Neusiedler See (Ebner
1951) ; GERMANY : SAXONY, BAVARIA: Regensburg
(Ramwe 1927; Harz 1957; Harz 1960); SWITZERLAND
: Saleve, Siders, Martigny (Fruhstorfer
1921; Harz 1957); MOLDAVIA: Chisinau-Kisinev,
Baurci (Bey-B ienko 1938, 1950); UKRAINE: Kirovogradska
oblast - Nerubajka (Bey-Bienko 1938, 1950);
YUGOSLAVIA: Srem [Szerem] (Pungur 1900);
CROATIA: Vinkovci [Nustar] (Graber 1870); SLOVENIA,
BOSNIA-HERCEGOVINA (Us & Matvejev
1967).
*The findings in Germany and Switzerland are very doubtful.
Description. - Size (mm) : Body length: 0' 5.8-
6.5,96.2-7.4; pronotum length x width, 0' 1.9-2.1
x 3-3.2, 9 2.0-2.2 x 3.3-3.6; forewing length, 0' 4-
4.8,92.5-3.2 .
Male (Plate 1: A). Head with few short setae,
interocular width larger than the ocular length
(IW/OL index =1.6). Pronotum semicircular (Fig.
lC), widest near the hind margin, surface sparsely
setose. Forewings partly shortened (Fig. ID),
reaching at most to the end of abdomen, distal
end narrowly rounded, veins in costo-radial area
lost. Posterior margin of the tergite 6 strongly
concave (Fig. IE). Tergite 7 is divided by two longitudinal
furrows into three parts, middle part
covered by the tergal gland (Fig. IF) . Glandular
pit large, trapezoid-oval, reaching to the posterior
margin of tergite. Hind margin of supraanal plate
rounded, right and left paraprocts are dissimilar,
right paraproct is much broader than the left. Cerci
long, with 8-9 segments, very often at least one
abbreviated. Genitalia: The curved hook portion
long, slender, pointed (Fig. IH); cleft sclerite (R2)
present on the right side. Coloration: Head black
except for pale labrum, lower part of the clypeus
and ocellar spots, proximal region of antennae
yellowish (about 13 segments), distal part brownish
to black. The maxillary palpomeres brown to
black, labial palps brownish. Pronotum black
except for narrow, yellowish, anterior and posterior
borders and broad, yellowish to transparent,
lateral margins. Forewings white to transparent
with a great number of small spots, hindwings
dark. Abdominal tergites black, sixth to eighth
ENT. SCAN D. VOL. 28:2 (1997) Revision of the Phyllodromica megerlei-group 167
1 mm
(A-F)
E
0.1 mm
(G -H)
.' - <0 • \ • •
'-rA\~?
Fig.!. Phyl/odromica megerlei, (A-B) female, (C-H) male: Pronotum (A, C), right tegmen (B, D), tergite 6 (E), tergite
7 with glandular pit (F), part of subgenital plate with stylus (G), hook (H).
168 viaua « L'. & Majzlan, O. ENT. SCAND. VOL. 28:2 (1997)
ex.
Fig. 2. Seasonal
dynamics of Phyllodromica
megerlei.o oothecae
• males EJfemales
60 I..N=219
70 r;=======:::::;--- - --- - -----;=============::;l
50 .. . . - . . . .
40 . -- -- -- 0-
30
20
10
o'--- ----'- 1---
o
August September
months
tergites with small, elongated whitish to transparent
spots on the posterior lateral margins. Abdominal
sternites black. Subgenital plate pale to dark
brown. Cerc i dark brown to black. Femur and tibia
black, tarsal segments (tarsomeres) yellow with
brown posterior ends, at the coxa-troc hanter joint
white regio ns.
Female. Head sparsely setose, interoc ular space
distinctly wider than the ocular length (IW/OL
index = 1.6-1.7 ). Forewings short, transversally
truncated (Fig. I B). Genitalia very similar to P.
pulcherrima. Coloration: Head black, interocular
space brown. Antennae, pronotum and forewings
as in male. Abdominal tergites and sternites black,
tergites and sternites 2-7 with triangula r white lateral
margins and often with narrow pale posterior
margins. Subgenital plate black with pale lateral
margins. Legs brown, tarsal segments yellow with
brown posterior ends , tibial bristles yellow.
Trochanter and adjacent part of coxa and anteroventral
margins of all coxae whitish. Ootheca:
Length 2.8 mm, width 2 mm . Surface brown, with
10-12 strong, longitudinal ridges on each side, the
keel is serra ted, the teeth are small (Fig. 4G).
Geographical distribution. - (Fig. 5) Phyllodrom ica
megerlei is widely distributed: SE Germ any
(Bavaria, Saxony - old, doubtful records only),
Czec h Republic, Slovakia, Austria, Switzerland
(old record s only), Hungary, Romania, nordeastern
Italy, Slovenia, Croatia (Istria, Dalmatia),
Yugoslavia (Serbia - Vojvodina), Bosnia-Hercegovina
(Hercegovina), Moldavia, Ukraine (Kirovogradska
oblast), but everywhere it is rare. On
the territory of Slovak ia and Czec h Republic it
reaches the northern border of its distri bution.
Biology. - Pliyllodromica megerlei occurs at altitudes
between sea level and about 700 m. The
animals live most freq uently in tussocks of grass
on xerotherm and mesohygrophilous meadows
with bushes and trees, banks of rivers, brooks
overgrown with grass, and clearings. Adults occur
from April to August and are most abundant at the
end of May and at the end of June (Fig. 2) .
2. Phyllodromica asiatica Bey-Bienko, 1950
stat. n.
(Plate I: B; Figs 3A-E, 5)
Phyllodromica megerlei asiatica Bey-Bienko, 1950:
233.
Aphlebia pallida (Brunner v. W.): Brunner v. W. 1882:
42, partim; Jakobson & Bianki, 1905: 127. partim.
Misidentifications.
Aphlebia punctata (Charpentier): Bolivar 1899: 585.
Misidentification.
Hololamp ra punctata (Charpentier) var.: Ebner 1919:
152.
Hololampra punctata f. erythronota Ramme, 1951: 324
(syn. Karabag 1958).
ENT. SCAND. VOL. 28:2 (1997) Revision of the Phyllodromica megerlei-group 169
I
II
8
c
0.1 mm
(D-E)
1 mm
(A-C)
Fig. 3. Phyllodromica asiatica, (A-E) male: Pronotum and tegmina (A), tergite 6 (B), tergite 7 with glandular pit
(C), hook (D), part of subgenital plate with stylus (E); ootheca of Phyllodromica pulcherrima (F) and P. megerlei
(G).
170 vidlicka, L'. & Majzlan, O. ENT. SCAND. VOL. 28:2 (1997)
H
1 mm
F
B (A-I)
0.1 mm
(J-K)
G
Fig. 4. Phyllodromica pulcherrima sp. n., (A-D , L) female, (E-K) male : Pronotum (A, E), metanotum (B, F), subgenital
plate (C), right tegmen (D, I), internal ovipositor (L), tergite 6 (G), tergite 7 with glandular pit (H), part of
subgenital plate with stylus (1), hook (K).
ENT. SCAND. VOL. 28:2 (1997)
Material studied. - Holotypect of Hololampra punctata
f. erythronota: Adana, Kleinasien, Rolleleg. (MNB).
Literature records. - TURKEY: Amasya(Brunnerv. W.
1882); Akbes, Kahramanmaras (Bolivar 1899); Bahce,
Amanos Mts (Ebner 1919); Gi.ilek (Bey-Bienko 1950);
Adana (Ramrne 1951 ; Karabag 1958); SYRIA: El
Ladhaqiye - Lattakia(Bey-Bienko, 1950).
Description. - Size (mm). Body length: a 5.47;
pronotum length x width, a 1.5 x 2.88; forewing
length , a 3.97.
Male (Plate I : B). Head with few short setae,
interocular width larger than the ocular length
(IW/OL index = 1.8). Pronotum trapeziform, surface
sparsely setose. Forewings reaching to the
end of abdomen, distal end narrowly rounded,
veins in costo-radial area only indicated. Radial,
medial and cubital veins can by discriminated.
Posterior margin of tergite 6 concave (Fig. 3B).
The middle part of tergite 7 covered with the tergal
gland (Fig. 3C). Glandular pit large, trapezoid-oval,
reaching to the posterior margin of tergite.
Hind margin of supraanal plate rounded.
Cerci long, with 8-9 segments. Coloration: Head
brown , ocellar spots yellowish-white, antennal
region pale . Proximal region of antennae yellowish-brown,
distal part brown. First and second
segments of maxillary palpomeres yellow, thirth
black, labial palps brownish. Disk of the pronotum
orange-yellow (central part) to yellowish
(posterior part and comers). Anterior and posterior
borders of pronotum white to transparent.
Forewings white to transparent with a great number
of small dark spots, basal part with big brownish-black
spot (Fig. 3A). Costal field and outer
margin of forewings without spots . Hindwings
dark to transparent. Abdominal tergites black.
Abdominal stemites black, lateral margins of sternite
2.-7. with yellow border. Subgenital plate
pale-brown, in the middle white. Femur black,
tibia brown-black, tarsal segments (tarsomeres)
yellow with brown posterior ends. Cerci black,
stylus brown with yellow tip.
Female (not examined, description from BeyBienko
1950). Very like as a P. megerlei, but usually
the disk of pronotum yellowish-orange, rarely
blackish-brown. Posterior border of pronotum
pale, equally wide as anterior border. Abdomen
with pale lateral border equally wide as pale posterior
border of pronotum. Ootheca: As in P.
megerlei, but with 18-19 longitudinal ridges on
each side.
Revision of the Phyllodromica megerlei-group 171
Geographical distribution (Fig. 5). - Known only
from Asia Minor - Turkey and Syria.
3. Phyllodromica pulcherrima sp, n.
(Plate 1: C, D; Figs 3F,4A-L, 5)
Type material. - Holotype cr, Bulgaria: Enime plane,
Vias (near Nesebar), 150 m, 30.v.1995, Majzlan
(SNMB). Paratypes: 6 ct , 8 9, same locality as holotype,
30.v.1995 and 26.v.1996, Majzlan (SNMB and
authorscollection).
Etymology. - The species name refers to its very decorative
appearance (from latin pulcherrimus = nicelooking,
magnificent).
Description. - Size (mm) : Body length: a 5.2-
5.7,96.2-6.6; pronotum length x width, a 1.5 x
2.4,9 1.8 x 2.8; forewing length, a 2.3-2.5, 9 2.2-
2.3.
Male (Plate 1: C). Head with few short setae,
interocular space slightly greater than distance
between ocellar spots, interocular width larger
than the ocular length (IW/OL index = 1.7).
Forewings strongly reduced, reaching at most to
the middle of the fourth abdominal tergite, distal
end widely rounded, veins in costo radial area
absent (Fig. 41). Hindwings strongly reduced,
nearly reach to posterior border of the metanotum
(Fig. 4F). Posterior margin of the tergite 6 slightly
concave (Fig. 4G). Seventh tergite with wide
oblique lateral margins reaching to the tergal
gland (Fig. 4H). Glandular pit large, oval. Genitalia
: The curved end of hook is very narrow, the
tip strongly bent (Fig. 4K), cleft sclerite present.
Coloration: Head with clypeus, labrum and ocellar
spots whitish, remainder shiny blackish. First,
second and fifth maxillary palpomeres brown,
third and fourth yellow. Labial palps blackishbrown.
First ten segments of antennae yellow,
remainder segments gradually brown to black.
Pronotum (Fig. 4E) with black disc and yellow to
transparent margins, anterior and posterior margins
narrower than lateral margins. Forewings
white to transparent with a great number of small
spots, hindwings dark. Abdominal tergites black,
tergites 2-5 with narrow, whitish posterior and lateral
borders; tergites 6-8 with large, whitish to
transparent spots in the posterior comers. Abdominal
stemites black. Subgenital plate pale to dark
brown. Femur and tibia dark brown to black, tarsomeres
yellow with brown posterior ends, at the
coxa-trochanter joint white regions.
Female (Plate 1: D). Head sparsely setose,
172 Yidlicka, I..:. & Majzlan, O. ENT. SCAND . VOL. 28:2 (1997)
40·
20·
MEDITERRANEAN
30·
oe Ph. megerfei
O. Ph. asiatica
* Ph. pulcherrima
Fig. 5. Geographical distribution of megerlei-group of cockroach genus Phyllodromica (data from literature - open
marks, material studied - filled marks).
interocular space distinctly wider than the ocular
length (IW/OL index = 1.6). Forewings short,
transversally truncated, their interior margins contacted
(Fig. 4D). Genital ia as in Fig. 4L. Coloration:
Head black, interocular space orange-brown.
Labrum, lower part of clypeus and ocellar spots
whitish. Antennae, pronotum (Fig. 4A) and
forewings as in male. Abdominal tergites and sternites
black, tergites and stemites 2-7 with white
lateral margins and often with narrow pale posterior
margins. Subgenital plate black with pale lateral
margins. Femora yellow-brown, longitudinally
striped Tibiae and tibial bristles yellow. Tarsal
segments yellow with brown posterior ends. All
trochanters, adjacent parts of coxae and their
anteroventral margins whitish, remaining parts of
coxae brownish. Ootheca: Length 2.3±0.2 mm,
width 1.8±0.1 mm. Surface brown, with 11-12
strong, longitudinal ridges on each side. The keel
is serrated, with 12-13 small teeth (Fig. 3F).
Biology. - The Bulgarian species P. pulcherrima
sp. n. lives at an altitude of about 150 m. It occurs
in submediterranean, xero-termophilous oak
woods (the alliance Quercion pubescentipetraeae).
The specimens were found in grass during
May.
Geographical distribution (Fig. 5). - Known only
from the type locality in Bulgaria.
Acknowledgements
For loan of material we are grateful to the following
curators and private persons : Dr Gyorgy Sziraky
(HNHM) , Dr Tibor Kovacs (MMG), Dr Kurt K.
Gunther (MNB), Dr Ivo Kovai' (NMP), Dr Ulrike
Aspock (NMW), Dr Ladislav Jedlicka (PFUK - Coli.
Dr Jan Gulicka), Dr Ilja Okali (SNMB), Dr Tomas Kizek
(SSMB), Dr Barnabas Nagy (Budapest).
We would like to thank Dr Louis M. Roth and Dr
Horst Bohn for valuable comments and linguistic help
with the English text.
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Journal and subscription to reprints of particular groups to be ordered from:
APOLLO BOOKS. Kirkeby Sand 19, DK-5771 Stenstrup. Denmark.
Príloha č. 7
VIDLIČKA, Ľ. 1999.
Caeparia sausai sp.nov. from Laos, and description of the male Caeparia donskoffi
(Blattaria: Blaberidae: Panesthiinae).
Entomological Problems 30(2): 1-5.
Entomological Problems. 30 (2): 1-5, December 1999
Caeparia sausai sp.nov. from Laos, and description of the male Caeparia
donskofji (Blattaria: Blaberidae: Panesthiinae)
Lubomir VIDLICKA
Institute of Zoology Slovak Academy of Sciences, Dubravska cesta 9, 842 06 Bratislava, Slovakia. E-mail: uzaevidl@savba.savba.sk
VIDLICKA, U. 1999. Caeparia sausai sp.nov, from Laos, and description of the male Caeparia donskoffi
(Blattaria: Blaberidae: Panesthiinae). Entomol. Probl. 30(2): 1-5. - The female of a new cockroach species of
Panesthiinae, Caeparia sausai is described from Laos. The male of Caeparia donskoJfi ROTH is described. Its
habitus is similar to that of the female: pronotum bicolored with dense long hairs; hind margin of supraanal plate
entire; genital phallomere L2d strongly toothed . The species is found in North and South Vietnam, Laos, and
Thailand.
Key words: Caeparia sausai,Caeparia donskoJfi, cockroaches, Blattaria, southeast Asia, Laos.
Introduction
Four species of Caeparia STAL, 1877, from
mainland southeast Asia, are known. Both sexes are
known for 2 species, namely C. saussurii (WOODMASON,
1876) and C. crenulata (BRUIJNING, 1948).
The males of C. donskofji ROTH and C. kaltenbachi
ROTH are unknown (ROTH 1979). ROTH (1982)
described an unnamed nymph from Thailand which
may prove to be either of these last 2 species; the
colour pattern of this nymph differs from that of the
adults of C. donskoffi or C. kaltenbachi, but the hind
margin of its supraanal plate is similar to that of C.
donskoffi.
In this paper the female of new species of
Caeparia (C. sausai) from Laos and the male of C.
donskoffi are described.
Caeparia sausai sp.nov,
Holotype !jl (SNM, Bratislava): Laos south, Attapu prov.,
Bolaven Plateau, 18-30.IY.1999, 15 km SE of Ban Houaykong,
NONG LOM (lake) env., N 15°02', E 106°35', alt. 800 m, E.
Jendek & O. Sausa leg.
Description of female (Figs 1 - 3). Head
sparsely punctate, ocell i very obvious. Antennae
short, thick, widest in the centre, consisting of 35
segments.
Pronotum transverse, widest near the hind
margin, the surface of pronotum hairless, sparsely
covered with large pits, on the anterior parts cresFig.
I: Caeparia sausai sp.nov. - habitus.
cent-like groove. Lateral margins broadly rounded.
Mesonotum "between tegmina densely foveolate.
Tegmina reaching the hind margin of the 6th tergite,
basal half enlarged, the apex rounded. Anterior parts
of coxae wrinkled, densely hairy. Femora smooth,
anterior and posterior margins with hairs; tibia
armed. Euplatulae (pulvilli) present on tarsal
segments 1 to 4; tarsal claws smooth, symmetrical,
arolia absent.
Abdominal tergites pitted, the hind margins of
tergites 3-7 with tubercles, which become gradually
longer in posterior tergites. Surface of tergites with
large, round impressions'. On T7 two very large
sublateral tubercles, th e lateral angles of T7
deflexed. Supraanal plate with round punctations,
hind margin with lateral angles, the area between
them subentire. Abdominal sternites narrow, surface
medially punctate, laterally with larger pits. S2 in
the middle with longitudinal keel. Cerci short,
conical, dorsal surface non-setose, ventrally with
long setae. StyIi not present.
Measurements [mm]. Total length 29;
pronotum length x width 5.5 x 9; tegmen length 22.
Coloration. Vertex black, the longitudinal black
band between ocelli continued over the clypeus and
labrum, remaining parts of head yellow (Fig. 2).
Eyes reddish brown , ocelli bright red. Maxillary and
labial y palps yellow, apical segments black. First 15
antennomeres shiny black, the succeeding 15
segments dull black, 4 segments yellow and the last
one dull black.
Pronotum yellow, in the centre. with big Iyrelike
black spot reaching to the hind margin.
Fig. 2: Caeparia sausai sp.nov, - color pattern of head.
Base of left tegmen black, central part with
yellow triangular spot and black band, preapical pali
membranous, yellowish and apex black. Anterior
half of right tegmen near to the left one, posterior
half dark with yellowish spot (Fig. 3). Legs black,
apical parts of coxae and femora, and basal parts of
tibiae yellow, spines black with reddish brown apex,
tarsi and claws black.
Abdominal tergites black, with small irregular
reddish-yellow spots sub laterally. Supraanal plate
2
black, the hind margin brownish red. Abdominal
sternites brownish black. Cerci brownish black.
Fig. 3: Caeparia sausai sp .nov, - tegmina.
Comments. The head colour of Caeparia
sausai is similar to that of C. crenulata, and its
pronotal colour pattern resembles those of C.
donskoffi and C. kaltenbachi. The same colour
combination is found in a male nymph from
Thailand (Caeparia sp. - Ron -I 1982), but probably
it is not the same species.
The species is named in honour of O. Sausa, who
caught this specimen.
Caeparia donskoffi ROTH
Caeparia donskofji R OTH, 1979: 101, Figs. 84A - F, 8SA
- F, 89G.
Male (Figs 4a - c, 5, 6a - c, 7a - c). Head
sparsely punctate, vertex not foveolate, largely
exposed. Clypeus with scattered, slender, long hairs;
eyes large, protruded; ocell i well developed ;
antennae short, consisting about 40 mostly broad,
short segments.
Pronotum transverse, anterior margin slightly
concave, posterior margin slightly excised medially,
lateral margins broadly rounded; anterior half of
pronotum with a pair of oblique grooves; surface of
pronotum densely covered with long hairs, some
black-coloured areas hairless, lateroposterior angles
th inly punctate. Mesonotum region between
tegmina with long hairs. Tegmina constricted,
surpassing the end of the abdomen, their apices
rounded. Thoracic sternites and coxae with long
hairs. Euplantulae (pulvilli) present on tarsal
segments 1 to 4 on all legs; tarsal claws simple,
symmetrical, arolia absent.
Abdominal tergites pitted, the hind margins
with some tubercles laterally, lateral margins of
abdominal segment 7 with 2 bluntly rounded teeth
on the anterior half, the lateral angle large, directed
caudal. Posterior margin of the supraanal plate with
collar-like elevation; medial part of hind margin
weakly con vex, entire. Cerci small, dorsal surface
non-setose, ventrally with long setae. Second
sternite narrow, medially with a longitudinal keel.
Hind margin of second sternite incised medially.
1 mm
I
....
Fig. 5: Caeparia donskoffi - genitalia of male from Viet-
nam.
Measurements [mm]. Total length 26 - 29.5;
pronotum length x width 4 - 5 x 7.5 - 8.5; tegmen
length 21 - 23.
Coloration. Head yellow, labrum black, an
incomplete or narrow complete black band on the
front femur unarmed except for slender hairs.
Genital phallomere L2d well developed, darkly
sclerotized with many large spines alon g the
posterior margin; L2vm long, narrow, stro ngly
sclerotized; LI small, only lightly sclerotized; R2
lightly sclerotized, markedl y reduced, the curved
part of the hook absent (Fig. 5).
Right paraproct with two strongly scle rotized
apical processes, left paraproct smaller, without
apical processes.
a
Fig. 4 Caeparia donskoffi: a) habitu s of male from Vietnam,
b - c) pronotum of males from Laos (a - leg. Kubao ,
b - leg. Jendek & Sausa).
The surfaces of abdominal sternites thinly punctate,
the lateral punctures denser; sternite 7 punctate on
apical half. Subgenital plate small, almost hidden,
non punctate. Styli absent. Antero-ventral margin of
clypeus, a bro ader tran sverse black band between
the antennae (split around ocelli); 4 broad black
vertical bands extend from between the eyes over
the vertex to the hind margin of the head, with a
black spot behind the eye s (Fig. 6a - c). Eyes reddi sh
brown, ocelli bright red. Maxillary palps yellow,
apical segments black. First 10 antennomeres shiny
black, the succeeding 20 seg ments dull blacki sh
grey and about 10 apical segments browni sh to
reddish-yellow.
Pronotum yellow with black anterior and lateral
margins, central part with two symmetrical Lshaped
black spots, hind angles with black spots, the
middle of the hind margin s with a black spot; hairs
on the pronotum yellow. Left tegmen with a large
black basal spot followed by a testaceous area, then
3
a
b
c
d
Fig. 6 Caeparia donskoffi - colour pattern of head: a)
ma le from Laos (leg. Kuban); b) male from Laos (leg.
Jendek & Sausa); c) ma le from Vietnam; d) female from
Thai -land.
a wide blackish to brownish band fo llowed by a
transparent zone which reaches the apex of the
tegmen. Right tegmen with a large black basal spot
followed by a testaceous area, then ,a triangular spot
followed by a wide brownish area (Fig. 7a - c).
Abdominal tergites brownish black with large
irregular reddish yellow spots laterally and medially.
Supraanal plate brown to brownish red . Abdominal
sternites brown, laterally w ith or without large
yellow spots. Cerci dorsally brown ish black,
ventrally yellow ish brown. Legs blackish with
transverse yellowish bands, tarsi blackish brown;
spines dark brown.
Material examined : 1 e: Laos north, 24-30.Y.1997, 20
km NW Louang Namtha, N 21°09.2, E 101°18 .7, al. 900±100 m,
E. Jendek & O. Sausa leg.; 1 e: Laos, Lou ang Namtha pr.,
2 1°09' N, 101°19' E, Namtha-eMuang Sing, 5-3 I.Y. 1997, 900-
1200 m, Vit Kuban leg.; 1 «. Vietnam north , l5 .Y.-16 .VI.1991 ,
Tam Dao nat. park, 75 km NW from Hanoi , E. Jendek leg.
Addition to description of female (Figs 6d, 7d,
8). Ocelli present, flat, less distinct than in ma le.
Antennae 39 segmented; first 15 antennal segments
shiny black, the succeeding 15 segments dull blackish
grey and 9 apical segments brownish to yellow.
. Pronotum nearl y hairless, but the coloration
was typical for C. donskoffi.
Coloration of tegmina is very similar to male
with left-right colour asymmetry (Fig. 7 d).
4
Fig. 7 Caeparia donskoffi - colour pattern of tegmina: a)
male from Vietnam ; b) mal e from Laos (leg. Jendek &
Sausa); c) male from Laos (leg. Kuban); d) female from
Thailand.
Measurements [mm]. Total length 33; pronotum
length x width 6.5 x 10.5; tegmen length 24.
Material exa mined: 1 ~ : Thailand, Chiang Mai prov.,
18°49 ' N, ~8°54 ' E, 1600 m, Doi Pui Mt., 2-6 .Y.1996. Vit
Kuban leg.
Comments. The male genital phallomeres are
very similar to those of known males of Caeparia.
Previously, C. donskoffi was known only from
North and South Vietnam. The present specimens
come from North Vietnam, Laos and Thailand. Thi s
species is widely distributed in southeast Asi a.
Acknowledgements
I am grateful to Dr Ondrej Sausa, Dr Eduard
Jendek and lng. Vit Kuban who gave me the
cockroach material from their own collections from
South East Asia and Dr Jan Kodada (Comenius
University) for photos.
I would like to thank Dr Louis M. Roth
(Harvard University) for valuable comments and
linguistic help with the English text.
References
ROTH, L.M., 1979. A taxonomic revision of the Panesthi inae of
the wo rld II. Th e ge nera Salganea STAL. Microdina KIRBY.
and Cae paria STAL (Dictyo ptera: Blattaria: Blaberidae).
Australian Journal ofZoology, Supplementary Series No .
69 : 1-201.
Fig. 8: Caeparia donskoffi - habitus female
ROTH, L.M., 1982. A taxonomic revision of the Panesthiinae of
the world IV. The genus Ancaudelia SHAW, with additions
to parts I-III, and a general discussion of distribution and
relationships of the compone.nts of the subfamily
(Dictyoptera: Blattaria: Blaberidae). Australian Journal of
Zoology, Supplementary Series No. 82: 1-142.
5
Manuscript received: 14.4. 1999
Príloha č. 8
VIDLIČKA, Ľ. 2002.
The new cockroach species from the genus Chorisoserrata from Laos
(Blattaria: Blattellidae: Pseudophyllodromiinae).
Entomological Problems 32(2): 145-147.
Eutomological Problems. 32(2): 145-147, March 2002
The new cockroach species from the genus Chorisoserrata from Laos
(Blattaria: Blattellidae: Pseudophyllodromiinae)
Lubomfr VIDLICKA
lnstilLilc of Zoology, Slovak Academy of Sciences. 842 06 Bratislava, Slovakia. E-mail: uzaevidl@savba.sk
VmuCKA, L. 2002. The new cockroach species from the genus Chorisoscrrat;) from Laos (Biattaria: Blattellidae:
Pscudophyllodromiinac). Eutomol. Probl. 32(2): 145-147. - The genus Chorisosermta Rol'll. 1998 contained up to now only
two species. Ch. CIJiicalis H ANITSCII from Sumatra and Ch. .mgiuaria H ANITSCII (only o known) from Vietnam and East Borneo.
The male and female of third specie~ Chorisosermtajeudeki from Laos arc described here. Unlike Ch. sagiuaria the male or Ch.
jeudeki have developed tergal gland on abdominal tergum?. The key or the genus Chorisosermta is presented.
Key words: Blauaria. Chorisoserr(lla. Laos
Introduction
The genus C/l()risoserraw descri bed RoTH ( 1998).
Symmetrical serrated tarsal claws, truncate vertex, maxillary
palpomerc fourth longer that the fifth and unspecialiscd
seventh abdominal tergum were used for separation
of this genus from the genus Chorisoueura BRUNNER OE
Wt~Tr~;Nwn. 1865. RoTH ( 1998) added into the genus
Clwrisosermw two species: Ch. sagillaria described by
HANITSCH ( 1927) from Vietnam and Ch. apicalis described
by HANITSCH ( 1929) from Sumatra. PlliNCIS ( 1950) recordeel
the second species from East 13orneo. Both species were
previously inscnccl in Chorisoueura.Species that remained
in the genusCltorisoueura arc distributed only in New Word
(South, Central and North America).
Key to males of Chorisoserrata
Hind margin of supraanal plate w ith a distinct ushapcd
excavation; apex of intcrstylar margin rounded
(Fig. 21) ...................................................................... 2
Hind margin ofsupraanal plate without excavation; apex
of intcrstylar margin more or less even. (Vietnam) ..........
................................................................ Ch. sagillaria
2 Rounded apex or interstylar margin with deep u-shapcd
excavation; pronotum semicircular (Figs. 2C. D. I).
(Laos) .......................................................... Ch.jendeki
Rounded apex or intcrstylar margin without excavation;
pronotum suboval. (Sumatra, East Borneo) ... Ch. apicalis
Chorisoserratajendeki, sp. nov.
(Figs. I . 2A- 1)
Male: Head exposed, markedly triangular. compound
eyes 1·elativcly small, intcrocular space wide. vertex flat,
maxillary palpomcrc 2 short, palpomeres 3 and 4 very long
and slender, palpomcrc 5 bulbous, more as 2 time shorter
that previous. densely hairy (f-igs. 2G, H); antennae longer
as the body, scapus 2 time longer as wide, pedicel short.
145
first segmcnl ol' f'lagcllum about thrice the length as following
segments, the segments arc lengthen in the direct
ol' apex; pronotum semicircular, posterior margin nearly
straight; tegminanarrow, lanceolate. exceeding beyond the
end of the abdomen, venation distinct. between veins and
branches orcostall'icld obvious screens c1·catcd spatial cell
pattern (Fig. 2A): hind wings well developed. costal t'icld
narrow with indisttnct visible veins. radial vein simple. in
the apex branched. median vein indistinct. visible only nca1·
the apex, cubital vein in last fourthY-bifurcatc. on the apex
bent upwards, intercalary apical triangle small but distinct
(
{
Fig. I. C/wri.,·o.<l!t-rawjendeki sp. nov. - habitu~. male.
G
~ · \I( I
I j
\. '
'v )
Figs. 2A-1. C/rorisoserratajendeki sp. nov.: A, pronotum and left tegmen: B. right hind wing: C. D. end of male abdomen. dor;al and
ventral view; E, F, end of female abdomen (dorsal and ventral view): G. H. head. vertex and fromal view: I. male subgenital plate,
dorsal.
146
(Fig. 2B); hind wings jelly out tegmen at rest. Front coxae
narrow and enlarge. Anteroventral margin of the front femur
with 6 small spines near the middle succeeded by severalslendersetaeand
terminating in one large spine; puvilli
developed on 4 proximal tarsomeres, enlarged. especially
fourth ; tarsal claws symmetrical. distinctly serrated, arolia
well developed. Abdominal tergum 7 narrow. with indistinct
but visible tergal gland in the middle; supra-anal plate
symmetrical, hind margin with a distinct u-shapcd excavation
(Figs. 2C, D). Cerci elongated, 11-segmented, ventral
surface with a few setae, segments I0-1 1 very narrow,
elongated und cylindrical. Exposed portion of subgenital
plate symmetrical, lateral corners produced, turned inside;
styles small, slim, placed at the deep indentation. the interstyJar
region produced, apex with shallow u-shapcd excavation.
Hook on the right side. apex narrowed. widely
rounded; median phallomcrc with pointed apex; left
phallomere with finger-like protrusion; long tapering filament
stick up out of body (Fig. 21).
Female: Head triangular. truncated vertex and compound
eyes visible, not overlapped by pronotum, intcrocular
space broad: antennae long; maxillary palpus long and slim.
same as male; pronotum semicircular; tegmina lanceolate.
reached beyond the abdomen: hind wings well developed,
long. Anteroventral margin of front femur with 9 small
spines. several setae a nd terminal large spine, tarsal claws
symmetrical, serrated; posterior margin o f supraanal plate
with deep u-shapcd excavation (Fig. 2E); apex ofsubgcnital
plate truncated, with long setae (Figs. 2E. F); cerci elongated,
siim.
Coloration: Vertex and frons dark brown with white
strip between eyes: first ten segments ofantenna! flagcllum
dark brown. remains flagcllomerc yellow; the lateral margins
of pronotum hyaline, disc yellow to brown with dark
brown margins; tegmina hyaline. subbasaly with shoe-like
dark brown macula (Fig. I ); hind wings smoked, costal
area yellowish; legs and spines yellow. poste rior parts of
tarsal segments dark; dorsal pan of abdomen yellowish
brown with broad longitudinal dark band in the middle;
ventral part of abdomen yellow.
Measurements (mm) ( 2 in parentheses): Length,
12.75 ( 12.75); pronotum length x width, 1.9-2.0 x 2.3-2.5
(2.2 x 2.7); tegmen length, 10.4 (10.3); interocular width,
0.8 (0.9).
Holotypc: o. LAOS: Bolikhamxai prov.: Ban Nok env..
N 18°08.7' E 104°28.1". 220 m. 9.-14.v.I998. Coli. 0. Sausa &
E. Jendek; (Slovak National Museum- Natural History Museum.
Bratislava).
Paratypes: LAOS: Same data as holotype. I 2: Luang
Namtha prov.: Namtha ---7 Muang Sing, 21°09. IOI 0
19.E.
900-1200 m, I o, 5.-31.v.l997. Coli. V. Kubait: (Institute of
Zoology SAS. Bratislava).
Etymology: The species is dedicated to Dr. E. Jendck
(Institute of Zoology), entomologist who collected specimens
of this new species.
Remarks: RoTH ( 1998) mentioned, that seventh
abdomimal tcrgum ofthe males from genus Chorisoserrata
is unspecialised, without tergnl gland. He had available one
male of Ch. sagittaria and destroyed male of Ch. apicalis
(from abdomen only with terminalia on slide). Seventh
abdominal tergum of here described males Ch. jendeki is
specialized although the tcrgal gland is small and indistinct.
Other characters of the genus arc identical. and suggest
close relationship ofall three species ofClwrisosermw.
Acknowledgements
I am grateful to Dr. Eduard Jendek. Dr. Ondrcj Sausa and
Ing. Vit Kub:li\ who provided me the cockroach material from
their own collections from Laos. I would like to thank Dr. Louts
M. Rot11 (1-Iarvard University) for valuable comments and linguistic
revision of the manuscript. This study was partly supported
by Slovak Scientific Grant Agency. Project No. 217I67/
20.
References
147
1-IANnscn. R.. I927. On a collection or Blattidae from southern
Annan. The Journal ofthe Siam Society. Natural History
St~pplemelll 7 ( I): 7--48.
1-IANITSCH. R.. 1929. Dr. E. Mjiiberg·s zoological collections from
Sumatra. I I. Blattidac. Arkit·for Zoologi 21 A (2): 1-20.
PRtNcts, K.. 1950. lndomalaiischc und australische Blattarien aus
dem Entomologischen Museum der Universitat in Lund.
Opuscula Emomolo~ica 15(3): 161-188.
RoTII. L. M., 1998. Thecockroach generaChorisoncura Brunncr.
Sorincuchora Caudcll, Chorisoneurodcs Princi~. andChorisoscrrata.
gen. nov. (Biattaria: Blattcllidae: Psc:.tdophyllodromiinac).
Oriental insects 32: 1-33.
Manuscript received: 10. 5. 2001
Príloha č. 9
VIDLIČKA, Ľ., VRŠANSKÝ, P., SHCHERBAKOV, D.E. 2003.
Two new troglobitic cockroach species of the genus Spelaeoblatta
(Blattaria: Nocticolidae) from North Thailand.
Journal of Natural History 37(1): 107-114.
J  N H, 2003, 37, 107–114
Two new troglobitic cockroach species of the genus Spelaeoblatta
(Blattaria: Nocticolidae) from North Thailand
Lˇ UBOMI´R VIDLICˇ KA†, PETER VRSˇANSKY´ ‡§ and
DMITRIJ E. SHCHERBAKOV§
†Institute of Zoology, Slovak Academy of Sciences, Du´bravska´ cesta 9,
842 06 Bratislava, Slovakia; e-mail: uzaevidl@savba.sk
‡Department of Zoology, Comenius University, Mlynska´ dolina B1,
842 15 Bratislava, Slovakia; e-mail: vrsansky@zutom.sk
§Paleontological Institute of the Russian Academy of Sciences,
123 Profsoyuznaya St., 117868 GSP Moscow, Russia;
e-mail: dima@iitp.ru
(Accepted 15 June 2001)
We describe two new troglobitic cockroach species Spelaeoblatta myugei n. sp.
and S. thailandica n. sp. (Blattaria: Nocticolidae) mountain caves of North
Thailand. We divided the genus Spelaeoblatta into two species groups. The Myugei
group includes both newly described species, which contain unique large tubercles
or elevations on abdominal terga. Using scanning electron microscopy we show
details of these special tubercles. The Thamfaranga group includes only
S. thamfaranga Roth is lacking these structures. The entire genus is further
characterized by the location of a tergal gland on abdominal segments three and
four. A key to identify the males is given. We also describe two specimens of
female nymphs, which possibly represent a new genus of cavernicolous cockroaches.
Elongated and narrow pronotum as well as very long front legs of these
nymphs resemble mantids.
K: Blattaria, Nocticolidae, cockroaches, Spelaeoblatta myugei n. sp.,
Spelaeoblatta thailandica n. sp., Thailand.
Introduction
Only two species were included in the genus Spelaeoblatta Bolivar, 1897. First,
S. gestroi was described by Bolivar (1897: 32) based on a single female from Burma
(Myanmar). Second, S. thamfaranga was described by Roth (Roth and McGavin,
1994: 1321) based on several males and females from Thailand. Here we describe
two species, S. myugei and S. thailandica, from caves in the Nam Lang region (Mae
Hong Son province), North Thailand. Specimens of both cockroach species were
captured in caves located near the well-known cave Tham Pha Mon about 15 km
SE and Red Cliff Cave about 20 km SW from Tham Nam Lang (figure 1).
Journal of Natural History
ISSN 0022-2933 print/ISSN 1464-5262 online © 2003 Taylor & Francis Ltd
http://www.tandf.co.uk/journals
DOI: 10.1080/00222930210135640
Lˇ . Vidlicˇka et al.108
F. 1. Distribution of Spelaeoblatta species in Thailand. (A) Thailand (grey—Mae Hong
Son province); (B) Mae Hong Son province (grey—Nam Lang region); (C) Nam Lang
region. (+) S. myugei, ($) S. thailandica, (&) S. thamfaranga.
Spelaeoblatta myugei n. sp.
(figures 2–4)
Type material. H: W, north-west Thailand: cave Tham Pha Mon (Mae
Hong Son Province, Nam Lang region, 19°28∞N, 98°14∞E), 20 March 1997, N. Myuge
(H. Miore) leg., coll. Slovak National Museum–Natural History Museum Bratislava
(SNM-NHMB). P: 1W, 3X, same data as holotype (Vidlicˇka collection).
Etymology. The species is named in honour of N. Myuge, who collected this
species.
Description
Size (mm). Body length: W 10.0–10.75, X 9.9–11.8; pronotum length×width,
W 2.5×3.9, X 2.4–2.75×3.5–4.1; forewing length, W 4.5–5.0, X 1.75–2.0.
Male (figures 2A, C, D, E, F, 3A–H, 4A–H). Head longitudinal, oval, partly
visible from above; faceted eyes small, strongly reduced, facetes indistinct; ocelli
absent; fifth maxillary palpomere shorter than the fourth, apex rounded. Antennae
long, slender, about 55 segments, scapus long, pedicel short, third segment as long
as scapus, other segments shorter. Pronotum parabolic, lateral hind corner slightly
oblong. Tegmina reaching to the base of fourth tergite, obliquely truncate, surface
covered by rather sparse trichia. Trichia are present mostly apically on the reduced
intercalary veins. Sc is highly expanded and supported by series of branches, clavus
is strongly reduced, radial field covers almost the half of the wing’s width, media is
Two new troglobitic cockroach species 109
F. 2. Spelaeoblatta myugei. (A) Male habitus; (B) female habitus; (C) male abdomen
(lateral view); (D) male genitalia (ventral view); (E) male subgenital plate (dorsal
view); (F) male front femur (anteroventral view).
four branched. Hind wings reduced, lobate (figure 3F), slightly shorter than metanotum.
Anteroventral margin of front femur with three large proximal spines succeeded
by a row of piliform spinules terminating in a large distal spine (type B
1
) (figures 2F,
4C); anterior margin of front tibia with several small spines, followed by three large
spines near the centre and terminating in two large spines, posterior margin with
many piliform spinules, two large spines at the distal end; tarsal claws simple,
symmetrical, pulvilli and arolia absent (figure 4D). Ten terga (T1–T10) are visible
on the abdomen (figures 2A, 3A, 4B). Abdominal terga (T2–T8) are strongly
specialized: the posterior margin of T2 is medially strongly concave, T3 contains
medially large round glandular pit with longitudinal elevated part in the middle, T4
has a large deep fossa, a pair of large tubercles at each posterior border of T5–T8
Lˇ . Vidlicˇka et al.110
F. 3. Spelaeoblatta myugei, male. (A) Thorax and abdomen; (B) tergal gland—T3+T4
(dorsal view); (C, D) tergal gland—detail (dorsal view); (E) tergal gland (dorsolateral
view); (F) reduced hind wing; (G, H) apex of left cercus.
F. 4. Spelaeoblatta myugei, male. (A) Abdomen (dorsolateral view); (B) abdomen—
T5–T10 (dorsal view); (C) front femur and tibia (ventral view); (D) tarsal claws;
(E) left tubercle on T5; (F) left tubercle on T6; (G) left tubercle on T7; (H) left
tubercle on T5—detail of structure.
(unique in Blattaria; figure 4A, B, E–H). Hind margin of supraanal plate rounded
with a shallow medial depression. Cerci 9–10 segmented, surface dorsally smooth,
ventrally with long setae, long sensilla on the apex (figure 3G, H). Subgenital plate
is symmetrical with two similar setose styles, interstylar margin convexly rounded
Two new troglobitic cockroach species 111
(figure 2E). Genitalia are shown in figure 2D: genital hook projects on the left side,
the apex strongly narrow, curved; right phallomere strongly sclerotized, curved to
S shape.
Colour: the specimens are pale brownish yellow, nearly translucent. Only maxillae
and tubercles on abdomen are darker.
Female (figure 2B). Head exposed, faceted eyes reduced, but present. Pronotum
parabolic, lateral hind corner slightly oblong. Tegmina strongly reduced to lateral
pads slightly overlapping mesomotum, veins indistinct. Metanotum laterally distinct
elongated. Hind margin of supraanal plate convexly rounded with long setae.
Subgenital plate with distinct valves. Pattern of front femur same as in the male.
Colour: the specimens are pale brownish yellow, nearly translucent.
Remarks
This species differs from S. thamfaranga described by Roth principally by the
presence of large tubercles on the hind margins of abdominal terga 5–8. Roth
claimed that the tergal glands were present on the second and third segments, but
our observations suggest that he may have missed the narrow first segment. His
drawing (figure 2H in Roth and McGavin, 1994: 1322) is very similar to our
figure 2A, except for the apparent absence of segment 1.
Spelaeoblatta thailandica n. sp.
(figure 5A–C)
Type material. H: W, north-west Thailand: Red Cliff Cave (Mae Hong
Son Province, Nam Lang region, 19°36∞N, 98°04∞E), 24 March 1997, N. Myuge
(H. Miore) leg., coll. SNM-NHMB.
Etymology. The specific name refers to the country of origin.
Description
Size (mm). Body length: W 8.75; pronotum length×width, W 2.4×3.4; forewing
length, W 5.0.
Male (figure 5A–C). Head longitudinal, oval, partly visible from above; compound
eyes strongly reduced but present; ocelli absent; apex of fifth maxillary
palpomere rounded. Antennae long, slender, about 60 segments, scapus and third
segment long, other segments markedly shorter. Pronotum unicoloured, parabolic,
the middle of hind margin straight, lateral corners obliquely truncated. Tegmina
reaching to the end of fifth tergite, obliquely truncate, surface covered by rather
sparse trichia, especially on apical part of venation. Hind wings reduced, only lobate,
reaching to a half of the metanotum length. Ten abdominal terga visible. The
posterior margin of T2 medially concave. T3 has a medial, inverted Y-like glandular
fossa, with hind margin medially elongated and elevated. T4 with a small medial
indentation, and T4–T7 with collar-like elevations on the hind margins. Hind margin
of supraanal plate is oval with medial shallow depression. Cerci are composed of
10 segments, surface dorsally smooth, ventrally with long setae, apically with long
sensilla. Subgenital plate symmetrical with two similar setose styles, interstylar
margin plane contains small medial elevation (figure 5C). The genital hook on the
left side, the apex strongly curved.
Colour: the specimen is pale brownish yellow, nearly translucent.
Male nymph. Size (mm): body length: 6.9; pronotum length×width, 1.9×2.5;
Lˇ . Vidlicˇka et al.112
F. 5. Spelaeoblatta thailandica, male. (A) Habitus; (B) abdomen (lateral view);
(C) subgenital plate (dorsal view).
front femur length, 1.9. Head with compound eyes having approximately 100 separated
ommatidia that become smaller towards the antennae. Wing pads on mesoand
metanotum markedly long. Anteroventral margin of front femur with three
large proximal spines followed by a row of piliform spinules (type B
1
); tarsal claws
simple, symmetrical, pulvilli and arolia absent. T3 medially elongated as in the adult
male. Subgenital plate with two small styles.
Remarks
The species S. thailandica and S. gestroi were only described from a single male
and a single female, respectively. So far, their opposite sexes are not known. Until
the male of S. gestroi is found, we cannot exclude a possibility that S. thailandica
is a male of S. gestroi. However, S. gestroi was captured about 170 km SW from
the locality inhabited by S. thailandica. Known cave-dwelling cockroaches have only
very limited areas of distribution (Vandel, 1965; Izquierdo and Oromi, 1994), thus
we believe that both described sexes are very probably different species.
Species groups
(1) Thamfaranga species-group: Hind margin of male abdominal tergites without
elevations or paired tubercles. Species: S. thamfaranga Roth.
Two new troglobitic cockroach species 113
F. 6. Female nymph from Red Cliff Cave. (A) Pronotum; (B) front femur
(anteroventral view).
(2) Myugei species-group: The hind margin of several male abdominal tergites
with elevations or paired tubercles. Species: S. myugei n. sp., S. thailandica
n. sp.
Key to males of Spelaeoblatta from Thailand
(The male of S. gestroi Bolivar from Burma is unknown.)
1 Hind margins of the abdominal tergites without elevations or paired tubercles (see
Roth and McGavin, 1994: figure 2F) . . . . . . . . . S. thamfaranga
– Hind margins of several abdominal tergites with elevations or paired tubercles . . 2
2 Paired lateral tubercles on hind margins of segments 5–8 . . . . . . S. myugei
– Elevations on hind margins of segments 4–7 . . . . . . . . S. thailandica
Note
Together with caged adult males and nymphs of S. thailandica were found two
female nymphs, possibly of a new genus of cavernicolous cockroach. Nymphs are
markedly different from all known species of Nocticolidae. They are completely
blind, very slender, yellowish white nearly translucent. The head is sparsely but
distinctly setose. Pronotum is long and narrow (figure 6A) similar to that of mantids.
Wing pads are indistinct. Front legs are very long, front femur of type C (figure 6B).
Subgenital plate without styles. Size of smaller nymph (mm): body length: 6.9;
pronotum length×width, 2.13×1.75; front femur length, 3.1. Size of larger nymphs
(mm): body length: 8.25; pronotum length×width, 2.25×1.85; front femur length,
3.4.
Lˇ . Vidlicˇka et al.114
Acknowledgement
We thank Nikolai Myuge (Koltzov Institute of Developmental Biology, Russian
Academy of Sciences) who provided specimens from the Thailand caves. We would
like to thank Dr Louis M. Roth (Harvard University) for valuable comments and
critical reading of the manuscript. This work was supported by a grant from the
Slovak Grant Agency, VEGA (2/7167/20).
References
B, I., 1897, Viaggio di Leonardo Fea in Birmania e regioni vicine. LXXVIII. Nouvelle
espe`ce cavernicole de la famille des Blattaires, Annali del Museo Civico di Storia Naturale
di Genova, 38, 32–36.
I, I. and O, P., 1994, Dictyoptera—Blattaria, in Ch. Juberthie and V. Decu
(eds) Encyclopaedia Biospeologica, Vol. 1 (Moulis, Bucarest: Socie´te´ de Biospe´ologie),
pp. 295–300.
R, L. M. and MG, G. C., 1994, Two new species of Nocticolidae (Dictyoptera:
Blattaria) and a rediagnosis of the cavernicolous genus Spelaeoblatta Bolı´var, Journal
of Natural History, 28, 1319–1326.
V, A., 1965, Biospeleology. The Biology of Cavernicolous Animals (Oxford: Pergamon
Press), 524 pp.
Príloha č. 10
VIDLIČKA, Ľ. 2013a.
New species of Macrophyllodromia (Blattaria, Blattellidae) from Ecuador
and a key to males of the genus.
Zootaxa 3635(2): 185-193.
Accepted by E. Bernard: 11 Feb. 2013; published: 26 Mar. 2013
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)Copyright © 2013 Magnolia Press
Zootaxa 3635 (2): 185–193
www.mapress.com/zootaxa/
Article
185
http://dx.doi.org/10.11646/zootaxa.3635.2.6
http://zoobank.org/urn:lsid:zoobank.org:pub:C4AEA110-8A93-4368-8CD5-322D639B6342
New species of Macrophyllodromia (Blattaria, Blattellidae) from Ecuador
and a key to males of the genus
ĽUBOMÍR VIDLIČKA1,2
1
Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 842 06 Bratislava, Slovakia.
E-mail: lubomir.vidlicka@savba.sk
2
Department of Biology, Faculty of Education, Comenius University, Moskovská 3, 813 34 Bratislava, Slovakia.
E-mail: vidlicka@fedu.uniba.sk
Abstract
Representatives of the cockroach genus Macrophyllodromia are widely distributed in Central and South Americas. The
genus involves 10 species, including one new species described here, that are divided into 3 species groups—splendida,
amabile and maximiliani. Macrophyllodromia onorei sp. n. is described as the fourth species of this genus from Ecuador.
A key is provided for males of all known species of Macrophyllodromia and a species distribution map is presented.
Key words: Macrophyllodromia, cockroach, distribution, Central and South America, taxonomy
Introduction
The Neotropical zone is known for very rich biodiversity. The cockroach fauna (Blattaria) of this region comprises
about 2,000 species belonging to approximately 180 genera. The Ecuadorean cockroach fauna includes 115 species
belonging to 45 genera (Vidlička 2013, Vršanský et al. 2012). However, descriptions of many new species from
this area are expected in close future.
The Neotropical cockroach genus Macrophyllodromia is widespread in Central America and the northern half
of South America. The genus was first described by Saussure and Zehnter (1893) and M. maximiliani was defined
as its type species by monotypy. This species was originally described as Pseudophyllodromia maximiliani by
Saussure (1873) from Mexico. It is widely distributed in Central America (Mexico, Guatemala, Honduras, Costa
Rica and Panama). The second species of the genus, M. splendida Hebard, 1920 and M. panamae Rocha e Silva
Albuquerque, 1962, the most recently described Central American species, have been found only in Panama.
Six species are known from South America. Almost all species are poorly represented in insect collections;
only M. nigrigena has been recorded from more than one country—in Guyana, French Guiana (Hebard 1926) and
Suriname (Bruijning 1959). This species also was listed from Bolivia (Rocha e Silva Albuquerque 1962), but the
data need reexamination. Evidence on this species in Bolivia is based only on 1 female, the identification of which
is problematic. Descriptions of most representatives of Macrophyllodromia are based on males and females are
unknown. The record from Bolivia indicates a disjunct distribution area of M. nigrigena.
Until now, only 2 species (M. lanceolata and M. multipunctata) have been described from Brazil, both from
the same locality, but they look markedly different (Lopes & Oliveira 2006). All other known species (including
the species described herein) come from diverse areas of Ecuador. The holotype of M. ecuadorana is probably a
native from Guayaquil, Ecuador, but it was intercepted at San Diego, California, USA, on bananas. All other
examined material from this species derivable from Ecuador was also intercepted in USA. One specimen of this
species came from Honduras but its occurrence there is controversial (Rocha e Silva Albuquerque 1962).
Two other species (M. amabile and M. nobile) were collected in NE Ecuador in 2005 (Anisyutkin 2007). Here
I describe a new species from the highland region in central Ecuador. Interpretation of protrusions on the subgenital
plate follows Anisyutkin (2007).
VIDLIČKA186 · Zootaxa 3635 (2) © 2013 Magnolia Press
Systematic entomology
Order Blattaria Latreille, 1810
Family Blattellidae Karny, 1908
Subfamily Pseudophyllodromiinae Hebard, 1929
Genus Macrophyllodromia Saussure and Zehnter, 1893
Type of genus: Pseudophyllodromia maximiliani Saussure, 1873.
Diagnosis. The genus Macrophyllodromia consisting of relatively large cockroaches (about 20 mm length) with a
broad, dark longitudinal band on the head stretching from interocular field to labrum (absent in M. nigrigena
Hebard and M. nobile Anisyutkin) and a pair of dark longitudinal bands on the pronotum (very modified in M.
multimaculata Lopes & Oliveira). Males without pheromonal tergal gland on abdomen, subgenital plate
asymmetrical with obvious protrusions. Tegmina and wings in both sexes fully developed.
Distribution: Central and South America.
The genus Macrophyllodromia shows close relationship to Latiblattella Hebard, 1917, Antitheton Hebard,
1919, and Eurylestes Hebard, 1940 (after Rocha e Silva Albuquerque 1962).
Macrophyllodromia onorei Vidlička, sp. n.
Figs. 1a–e, 2a
Type material. Holotype male, Ecuador, Otongachi reserve, 00°19'15"S; 78°57'06"W, near the village of La Union
del Toachi, 33 km E of Santo Domingo de los Colorados, elev. ca 850 m, piedmont evergreen woodland, 19–25
September 2009, Ľ. Vidlička & M. Kozánek leg., deposited at the Institute of Zoology SAS, Bratislava, Slovakia.
Description. General colour of body yellowish brown (Fig. 1a).
Head with occiput pale, epicranium and vertex mostly black, but vertex over the ocelli with narrow irregular
transverse yellow strip. Face yellow with broad longitudinal black band continued to labral suture. On clypeus
band brown, but distinct. Genae black. Eyes and ocellar spots (fenestrae) yellowish. Maxillary palps yellowish, last
joint brownish (Fig. 1b). Antennae brownish, tenuous, longer than whole body.
Pronotum widest in caudal third. Two black longitudinal bands (widest caudally) beginning on front margin,
not reaching hind margin of pronotum. Sides of pronotum yellowish translucent, with scattered fine but visible
dark dots (Fig. 2a). Tegmina and wings fully developed, folded and overlapping apex of abdomen. Veins reddish
brown. Anal fields of forewings with eight anal veins, first vein with four rami. Interveinal areas in anal fields of
forewings with narrow yellow stripes. Legs yellow, apical part of tibiae and tarsal segments on all legs black (Fig.
1c). Tarsal joints with large distal pulvilli. Tarsal claws simple, unspecialized, between them well developed arolia
with bilobed distal end. Antero-ventral margin of front femur armed with 18 heavy spines of type A (sensu BeyBienko
1950, Roth 2003).
Abdominal dorsal surface unspecialised. Supra-anal plate short, triangular, bilobed at apex (Fig. 1d). Cerci
elongated, composed of 15 distinctly separated segments (Figs. 1d, e). Subgenital plate strongly asymmetrical and
of complicated design (Fig. 1e). Apical part divided into three protrusions. Protrusion on left side (LP) forming two
large projections, the first extending straight back as a large, sharp horn with finely bifurcate apex (LP1), the
second extending to the right as a very long, narrow rod-like sclerite (LP2), partly enclosed by small central growth
(CG), and ending just right in deep groove of mitten-like right protrusion (RP) (Figs 1e, 3a). Styli completely
absent, protrusions apparently only representing projections of the subgenital plate.
Measurements. Holotype (mm): length of body 20.7, length of pronotum 4.4, width of pronotum 7, length of
tegmen 21.4, width of tegmen 6.4.
Etymology. This new cockroach species named in honour of Professor Giovanni Onore, missionary,
entomologist and President of the Otonga Foundation in Ecuador.
Zootaxa 3635 (2) © 2013 Magnolia Press · 187NEW MACROPHYLLODROMIA FROM ECUADOR
FIGURE 1. Macrophyllodromia onorei sp. n., male: (a) habitus, (b) head with appendages, (c) right front leg (ventral view),
(d) abdomen with cerci (sap: supra-anal plate) (dorsal view), (e) the tip of abdomen (sgp: subgenital plate) (ventral view).
VIDLIČKA188 · Zootaxa 3635 (2) © 2013 Magnolia Press
FIGURE 2. Male heads and pronotal shields of Macrophyllodromia spp. (a) M. onorei sp. n., (b) M. panamae, (c) M.
ecuadorana, (d) M. nigrigena, (e) M. nobile, (f) M. multipunctata, (g) M. maximiliani, (h) M. splendida, (i) M. lanceolata, (j)
M. amabile. Figures 2b–2j are modified from drawings, descriptions and photos in Hebard (1920, 1926), Rocha e Silva
Albuquerque (1962), Lopes & Oliveira (2006) and Anisyutkin (2007).
Zootaxa 3635 (2) © 2013 Magnolia Press · 189NEW MACROPHYLLODROMIA FROM ECUADOR
FIGURE 3. Subgenital plate of Macrophyllodromia spp. (ventral view). (a) M. onorei sp. n., (b) M. panamae, (c) M.
ecuadorana, (d) M. nigrigena, (e) M. nobile, (f) M. multipunctata, (g) M. maximiliani, (h) M. splendida, (i) M. lanceolata, (j)
M. amabile. RP: right protrusion, LP: left protrusion, LP1, LP2: projections of left protrusion, CG: central growth. Figures 3b–j
are modified from descriptions and illustrations in Hebard (1920, 1926), Rocha e Silva Albuquerque (1962), Lopes & Oliveira
(2006) and Anisyutkin (2007).
VIDLIČKA190 · Zootaxa 3635 (2) © 2013 Magnolia Press
Differential diagnosis. Macrophyllodromia onorei sp. n. is similar to M. panamae in the length of the dark
longitudinal bands of the pronotum (bands not reaching posterior margin of pronotum) but not in their form (Figs.
2a, 2b). Structure of the subgenital plate is also similar in both species, but the central growth (CG) is markedly
shorter in M. onorei (Figs. 3a, 3b). The new species is similar to M. ecuadorana, M. nigrigena, M. nobile and M.
amabile in the coloration of the pronotum (Figs. 2c, 2d, 2e, 2j), but it is easily distinguishable from all those species
in the presence and form of the dark longitudinal band on the face (nigrigena and nobile have the whole face dark;
see Figs. 2d, 2e) interrupted with a narrow pale transversal strip over the ocelli. In M. ecuadorana and M. amabile
this band is without interruption (Figs. 2c, 2j).
Survey and distribution of Macrophyllodromia species (Fig. 4)
1. Macrophyllodromia maximiliani (Saussure, 1873)
Mexico, 1 male (Saussure 1873); Guatemala, 1 male, Dept. Petén, near Lake Eckibix (=Laguna Oquevix,
located on the savanna about 40 km south of town Flores) (Rocha e Silva Albuquerque 1962); Honduras, 1 male, 3
females, Guaimas Dist., Tela (Rocha e Silva Albuquerque 1962); Costa Rica, 1 male, intercepted at San Francisco,
USA (Rocha e Silva Albuquerque 1962); Panama, 1 female, Barro Colorado Island (Hebard 1933); 1 male, Barro
Colorado Island; 1 male, 1 female, intercepted at San Francisco, USA (Rocha e Silva Albuquerque 1962); 1 male,
Barro Colorado Island (Roth & Naskrecki 2001).
2. Macrophyllodromia splendida Hebard, 1920
Panama, 2 males, Porto Bello (=Portobelo) (Hebard 1920); 1 female, Barro Colorado Island (Hebard 1933).
3. Macrophyllodromia nigrigena Hebard, 1926
French Guiana, 1 male, 1 female, St Jean du Maroni; 1 female, St. Laurent du Maroni (Hebard 1926); Guyana,
2 males, 1 juv. male, 1 female, Distr. Bartica, Kartabo (Hebard 1926); Suriname, 1 male, Coppename (Bruijning
1959); Bolivia, 1 female, Tumupasa (Rocha e Silva Albuquerque 1962)
4. Macrophyllodromia ecuadorana Rocha e Silva Albuquerque, 1962.
Ecuador, 1 male, Guayaquil, intercepted at San Diego, USA; 2 males, 4 females, 1 nymph, intercepted in USA
(Rocha e Silva Albuquerque 1962); Honduras ?, 1 male, intercepted at New Orleans, USA (Rocha e Silva
Albuquerque 1962).
5. Macrophyllodromia panamae Rocha e Silva Albuquerque, 1962
Panama, 1 male, Barro Colorado Island (Rocha e Silva Albuquerque 1962); 1 male, Barro Colorado Island
(Roth & Naskrecki 2001).
6. Macrophyllodromia lanceolata Lopes & Oliveira, 2006
Brazil, 8 males, Acre, Senador Guiomard, Reserva Catuaba (Lopes & Oliveira 2006).
7. Macrophyllodromia multipunctata Lopes & Oliveira, 2006
Brazil, 1 male, Acre, Senador Guiomard, Reserva Catuaba (Lopes & Oliveira 2006).
8. Macrophyllodromia amabile Anisyutkin, 2007
Ecuador, 1 male, 1 female, env. of waterfall San Rafael on Rio Coca, forest, 95 km E of city Quito (Anisyutkin
2007).
9. Macrophyllodromia nobile Anisyutkin, 2007
Ecuador, 1 male, 80–85 km E of town Lago Agrio, env. of lake Lago Grande, Rio Cuyabeno (Anisyutkin
2007).
Zootaxa 3635 (2) © 2013 Magnolia Press · 191NEW MACROPHYLLODROMIA FROM ECUADOR
10. Macrophyllodromia onorei Vidlička, sp. n.
Ecuador, 1 male, Otongachi reserve, near the village of La Union del Toachi, 33 km E of Santo Domingo de los
Colorados.
FIGURE 4. Macrophyllodromia species distribution map (Central and South America). (1) M. maximiliani, (2) M. splendida,
(3) M. nigrigena, (4) M. ecuadorana, (5) M. panamae, (6) M. lanceolata, (7) M. multipunctata, (8) M. amabile, (9) M. nobile,
(10) M. onorei sp. n.
VIDLIČKA192 · Zootaxa 3635 (2) © 2013 Magnolia Press
Key to males of Macrophyllodromia (species groups and species).
1. Caudal end of subgenital plate with two distinctly separated protrusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Left and right protrusions on the caudal end of subgenital plate slightly asymmetrical, separated only with narrow groove (Fig.
3h); two pronotal bands fused at the caudal margin (Fig. 2h) (splendida group) . . . . . . . . . . . . . . . . . . . . . . . . . . M.splendida
2. Right protrusion outward rotated (Figs. 3i, 3j) (amabile group). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Right protrusion inward rotated (Figs. 3a–3g) (maximiliani group). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Left protrusion comb-like (Fig. 3i), two black longitudinal bands on pronotum parallel, on posterior margin markedly
expanded (Fig. 2i) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..M. lanceolata
- Left protrusion compact (Fig. 3j), black longitudinal bands on pronotum convergent towards anterior margin of pronotum (Fig.
2j) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. amabile
4. Right protrusion straight, on left stylus a small leftward protrusion (Fig. 3g); tegmen with row of dark, obvious spots. . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. maximiliani
- Right protrusion moderately but visibly curved (Figs. 3a–3f); tegmen without spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Interior part of left protrusion (LP2) curved toward right stylus, marginal part (LP1) short and straight (Figs. 3e, 3f). . . . . . . . 6
- Interior part of left protrusion (LP2) narrow, transverse; marginal part (LP1) obviously curved inward or its tip cuspidate and
bent (Figs. 3a–3d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6. Protrusions RP and LP2 equally tapering (Fig. 3e); dark bands on pronotum obvious (Fig. 2e); most of facial part of head dark
(Fig. 2e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. nobile
- Protrusions RP and LP2 thick, tapered only on last third (Fig. 3f); pronotum with intricate dark mosaic not in band-like form
(Fig. 2f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. multipunctata
7. Protrusion LP1 wholly curved inward (Fig. 3d); facial part of head dark (Fig. 2d); two pronotal longitudinal bands briefly separated
at the caudal margin (Fig. 2d). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. nigrigena
- Protrusion LP1 pointed, straight or the tip distinctly curved, facial part of head with longitudinal dark band; pronotal longitudinal
bands widely separated at the caudal end . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Central growth (CG) not at all overlapping protrusion LP2 (Fig. 3c); extended caudal end of pronotal dark bands reaching posterior
margin of pronotom (Fig. 2c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. ecuadorana
- Central growth (CG) overlapping protrusion LP2 at least partially; caudal end of pronotal dark bands not reaching posterior
margin of pronotum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Central growth (CG) very long, completely overlapping central part of LP2 (Fig. 3b); pronotal longitudinal bands relatively
narrow with smooth margins (Fig. 2b); facial part of head with uninterrupted longitudinal dark band (Fig. 2b) . . M. panamae
- Central growth (CG) only slightly protuberant, partially overlapping central part of LP2 (Fig. 3a); pronotal longitudinal bands
wide with indented margins (Fig. 2a); facial part of head with longitudinal dark band on facial part of head with transverse
light stripe over ocellar spots (Fig. 2a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. onorei sp. n.
Notes. In only five of Macrophyllodromia species have females been described. Several unnamed females also have been
reported. Anisyutkin (2007) mentioned 4 females from 3 various localities in Ecuador. They probably represent at least two
different species, but their assignment to the known species was not possible at that time. Roth & Naskrecki (2001) also
mentioned an unknown female (probably a new species) from Costa Rica.
The transverse light stripe over the ocellar spots is characteristic for females of M. amabile and also some unnamed
females from Rio Aguarico, Ecuador (Anisyutkin 2007). Macrophyllodromia onorei is the first species found in which the male
has a strong transverse stripe over the ocellar spots.
Acknowledgements
The study was supported by the Slovak Research and Development Agency (APVV) using financial support no.
APVV-0213-10. I thank Dr. Giovanni Onore for his all-round assistance in Ecuador, Dr. Ladislav Roller and Dr.
Juraj Majtan for valuable comments.
References
Anisyutkin, L.N. (2007) New species of the genus Macrophyllodromia Saussure & Zehntner, 1893 (Dictyoptera: Blattina:
Blattellidae) from Ecuador. Cockroach Studies, 2, 27–41.
Bey-Bienko, G.Y. (1950) Nasekomye tarakanovye [Cockroaches]. Fauna SSSR, N.S. 40. Moscow-Leningrad, 342 pp.
Bruijning, C.F.A. (1959) The Blattidae of Surinam. Studies on the Fauna of Suriname and Other Guyanas, 4, 1–103.
Hebard, M. (1920) The Blattidae of Panama. Memoirs of the American Entomological Society, 4, 1–148.
Hebard, M. (1926) The Blattidae of French Guiana. Proceedings of the Academy of Natural Sciences of Philadelphia, 78,
135–244.
Zootaxa 3635 (2) © 2013 Magnolia Press · 193NEW MACROPHYLLODROMIA FROM ECUADOR
Hebard, M. (1933) Notes on Panamanian Dermaptera and Orthoptera. Transactions of the American Entomological Society,
59(2), 103–144.
Lopes, S.M. & de Oliveira, E.H. (2006) Duas espécies novas de Macrophyllodromyia do Estado do Acre, Brasil (Blattaria,
Blattellidae) coletadas em ninhos de vespas. Iheringia, Série Zoologia, 96(2), 257–260.
http://dx.doi.org/10.1590/S0073-47212006000200017
Rocha e Silva Albuquerque, I. (1962) Synopsis of the neotropical cockroach genus Macrophyllodromia (Orthoptera:
Blattoidea, Epilamprinae). Proceedings of the United States National Museum, Smithsonian Institution, 113, 421–428.
Roth, L.M. (2003) Systematics and phylogeny of cockroaches (Dictyoptera: Blattaria). Oriental Insects, 37, 1–186.
http://dx.doi.org/10.1080/00305316.2003.10417344
Roth, L.M. & Naskrecki, P. (2001) Trophobiosis between a blattellid cockroach (Macrophyllodromia spp.) and fulgorids
(Enchophora and Copidocephala spp.) in Costa Rica. Journal of Orthoptera Research, 10(2), 189–194.
http://dx.doi.org/10.1665/1082-6467(2001)010[0189:TBABCM]2.0.CO;2
Saussure, H. (1873) Mélanges orthoptérologiques. IVme Fascicule, Mantides et Blattides. Mémoires de la Société de Physique
et d’Histoire Naturelle de Geneve, 23(1), 1–164.
Saussure, H. & Zehntner, L. (1893) Biologia Centrali-Americana. Insecta. Orthoptera. Vol I. Blattidae. Suborder Orthoptera
Genuina, 1, 13–123.
Vidlièka, Ľ. (2013) Cockroaches (Blattaria) of Ecuador—checklist and history of research. Zootaxa, 3599 (5), 401–445.
http://dx.doi.org/10.11646/zootaxa.3599.5.1
Vršanský, P., Chorvát, D., Fritzsche, I., Hain, M. & Ševèík, R. (2012) Light-mimicking cockroaches indicate Tertiary origin of
recent terrestrial luminescence. Naturwissenschaften, 99(9), 739–749.
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Príloha č. 11
VIDLIČKA, Ľ. 2013b.
Cockroaches (Blattaria) of Ecuador – checklist and history of research.
Zootaxa 3599(5): 401-445.
Accepted by F.A. Crespo: 22 Nov. 2012; published: 9 Jan. 2013
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)Copyright © 2013 Magnolia Press
Zootaxa 3599 (5): 401–445
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Article
401
http://dx.doi.org/10.11646/zootaxa.3599.5.1
http://zoobank.org/urn:lsid:zoobank.org:pub:8662421E-F753-46B2-ACDE-0A5868DE6716
Cockroaches (Blattaria) of Ecuador—checklist and history of research
ĽUBOMÍR VIDLIČKA
Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 845 06 Bratislava, Slovakia.
E-mail: lubomir.vidlicka@savba.sk
Department of Biology, Faculty of Education, Comenius University, Moskovská 3, 813 34 Bratislava, Slovakia.
E-mail: vidlicka@fedu.uniba.sk
Abstract
Cockroaches are an understudied group and the total number of described taxa increases every year. The last checklist of
Ecuador species was published in 1926. The main aim of this study was to complete a new checklist of cockroach species
recorded in Ecuador supplemented with a research history of cockroaches (Blattaria) on the territory of continental Ecuador.
In addition, the checklist contains comments on Ecuadorian faunistic records, including the Galápagos Islands. A total
of 114 species (105 in continental Ecuador and 18 in Galápagos Islands) belonging to 6 families and 44 genera are listed.
Forty species (38.1 %) occur solely in continental Ecuador and five (27.8 %) are endemic on Galápagos Islands.
The results indicate that further research on the cockroach fauna of Ecuador as well as determination of museum
collections from this territory is needed.
Key words: cockroach distribution, catalogue, South America, faunistic record
Introduction
Cockroaches (Blattaria) are a mainly tropical and subtropical order of insects. The Neotropical zone (South and
Central America, including the Caribbean Islands) have, in comparison with other zoogeographic regions, the
richest fauna of cockroaches. About 2,000 of approximately 5,500 known recent species are native to South
America. Cockroaches are an understudied group and the total number of described taxa increases every year,
especially from the Neotropical region. In the last decade checklists of cockroaches from several South American
countries have been published. The greatest number of species was recorded from Brazil, probably due to its great
size and diverse biotopes. In 2008, 647 species were listed from Brazil (Pellens & Grandcolas 2008), an increase of
almost 70% over the 383 species listed in 1964 (Rocha e Silva Albuquerque 1964a). Recent regional faunal lists in
the Neotropics have been published for Colombia (133 species; Vélez 2008), Argentina (89 species; Crespo et al.
2010), Venezuela (87 species; Bonfils 1987; Perez 1988), Nicaragua (97 species; Maes 1992a,b,c), Puerto Rico (48
species; Gutiérrez 1999) and Cuba (85 species; Gutiérrez 1995).
Cockroaches of Ecuador are insufficiently explored. The last summaries were compiled by Campos (1923,
1926). Additional data on occurrence of cockroaches are scattered in various publications. The Galápagos Islands
are the best studied Ecuadorian region, but a number of species is very low (18 species; Peck & Herrera 2011). An
annotated list of Galápagos Blattaria is given in the paper of Peck & Roth (1992).
This study is based on a critical evaluation of the published data on Ecuadorian cockroaches. The base sources
were the checklists of Campos (1923, 1926) and the catalogues of Princis (1962, 1963, 1964, 1965, 1966, 1967,
1969, 1971). The classification is based on McKittrick (1964) and Roth (1970d, 2003a), and also takes into account
the works of Grandcolas (1996) and Djernaes et al. (2012).
Determination and revision of cockroach material deposited in many museums will be carried out in following
years.
VIDLIČKA402 · Zootaxa 3599 (5) © 2013 Magnolia Press
Cockroaches of continental Ecuador—history of research
The oldest reference about cockroaches in the territory of Ecuador was listed in a book from Italian Jesuit Mario
María Cicala (19 January 1718, Fiume di Nisi–after 1785, Viterbo). Cicala was a missionary in Ecuador and he
stayed there for 23 years (1743–1767). He described his experiences in two books, but only the manuscript of the
second part from the year 1771 (Cicala 1994, 2004) has been preserved. He divided cockroaches into three groups,
but he did not mention any particular species. At the end he wrote, “All houses and apartments are full of
cockroaches at all times.” (Cicala 2004:189).
The first cockroach species known and described from Ecuador was Paratropes aequatorialis (Saussure,
1864). Swiss orthopterologist Henri Louis Frédéric de Saussure (27 September 1829, Geneva–20 February 1905,
Geneva) only indicated that this species was from the locality “Respublica Equator” and did not give the name of
the collector or more accurate locality data. Saussure visited the West Indies, Mexico and the United States in
period of time from 1854 to 1856, where he collected insects (Bedot 1906a, b). He traveled with another Swiss
naturalist, Adrien Jean Louis François de Sumichrast (15 October 1828, Yvonne, Canton de Vaud,
Switzerland–26 September 1882, Tonalá, Chiapas, Mexico). Their valuable collections were taken to Geneva by
Saussure in 1856, while Sumichrast decided to stay in Mexico (Papavero & Ibáñez-Bernal 2001). Orthopteroid
specimens (now in the Natural History Museum of Geneva) were described by Saussure in a long series of papers
(Saussure 1859, 1861, 1862, 1864. 1868, 1869, 1893–1899). These works contained descriptions of orthopterans
not only from Central America but also from northern South America countries (Brazil, Guyana, French Guiana,
Suriname, Peru, Bolivia, Chile, Venezuela and Ecuador). These specimens may have been obtained during his
travel. The aforementioned specimen of P. aequatorialis is deposited in the Geneva Museum and has slightly more
detailed labeling than the published record: “Quito, Andes, M. H. de Saussure” (J. Hollier, personal
communication). One grasshopper species (Chromactis ictera (Pictet & Saussure, 1887)) is labelled as “Quito, M.
H. de Saussure” (Hollier 2011). These two cases suggest that Saussure was the collector of this cockroach species.
Subsequent data on Ecuadorian cockroaches were published by English entomologist Francis Walker (31
July 1809, Southgate–5 October 1874, Wanstead). Between 1844 and 1873 Walker was employed as a curator in
the British Museum Natural History where he worked on collections of orthopterous insects. His catalogue of
cockroaches (Walker 1868) includes five species from the territory of Ecuador. All five species were described as
new to science, but two of them were later synonymized. Two of these species (Blaberus parabolicus and
Paratropes lanceolatus (syn. of P. aequatorialis)) originated from the Fraser’s collection. Louis Fraser (1819,
England–1884, USA) was an excellent British naturalist, zoologist and collector. He was curator at the Museum of
the Zoological Society in Brewer Street, Golden Square (London). Fraser participated in a number of expeditions
and collected vertebrates and insects in Ecuador in 1857–1859. His first collecting place was Cuenca, the capital
city of the Azuay Province (formerly Cuenca) (October–November 1857, March–May 1858). In this locality both
species (each of one specimen) were collected. From December 1857 through February 1858 he was at Gualaquiza
(Morona Santiago Province) and Zamora (Zamora Province). Afterwards, from June to August 1858 he was at
Riobamba (Chimborazo Province) and its sourroundings (Guana, Pinipi, Matos, Titiacun, Punin). He spent four
months (August–November 1858) at Pallatanga, a pueblo southeast of Riobamba. In the middle of January 1859 he
returned to Riobamba and then made an excursion to Panza (southern slope of Chimborazo), returned to Riobamba,
and then moved to Quito. During period of February–April 1859 Fraser visited various places in the surroundings
of Quito. From July to September 1859 he visited the environs of Guayaquil, then left for Esmeraldas. During
October–December 1859 Fraser paused at Esmeraldas, on the coast of Ecuador and on his route from Guayaquil to
Panama by sea (Sclater 1858a, b, 1859, 1860a–e; Sharpe 1906; Moore 2004).
Two other roaches described by Walker (Megaloblatta longipennis and Polyzosteria crassipes (syn. of
Pycnoscelus surinamensis)) came from Stevens’ collection. Samuel Stevens (11 March 1817, St Martin-in-theFields,
Middlesex–29 September 1899, Beulah Hill, Croydon, Surrey) was a Fellow of the Linnean and
Entomological Societies of London and Entomological Club and a familiar figure at meetings of naturalists. In
1848 he established the Natural History Agency in Bloomsbury Street (London), where bought and sold insects
from all over the world. He supported various explorers and collectors, the best known of whom were Alfred
Russel Wallace and Henry Walter Bates, who collected natural history specimens in South America, too. Stevens
acted as their agent in the distribution of the animals and insects that they obtained. Many specimens were bought
from Stevens by the British Museum of Natural History. After Stevens’ death his collections of insects were sold at
the auction (Anonymous 1899a, 1899b; Sharpe 1906; Baker 2001).
Zootaxa 3599 (5) © 2013 Magnolia Press · 403COCKROACHES (BLATTARIA) OF ECUADOR
The last of Walker’s cockroach species (Panchlora nigricornis) was donated to the British Museum by French
naturalist and collector Jules Bourcier (1797, Cuisery, Saône-et-Loire–9 March 1873, Batignolles, Paris), who
acted as French Consul in Ecuador from 1849 to 1850. All of Walker’s specimens are deposited in the Natural
History Museum, London.
Further cockroach species from Ecuador were described by the American orthopterologist Samuel Scudder
(13 April 1837, Boston–17 May 1911, Cambridge, Massachusetts) in 1869. Four of nine species mentioned in his
study (Scudder 1869) were newly described, but three of them were later synonymized: Blabera armigera (syn. of
Blaberus parabolicus), Blabera cubensis (syn. of Blaberus discoidalis), Blabera femorata (syn. of Eublaberus
posticus). Only Ischnoptera pallipes is a valid species. The other five species (Panchlora exoleta, Lanxoblatta
rudis (?), Periplaneta americana, Periplaneta australasiae and Ischnoptera melasa [as I. melana]) were new
records for Ecuador.
Cockroaches determined by Scudder came from the collection of the first expedition of the well-known
American theologian, naturalist and collector James H. Orton (21 April 1830, Seneca Falls, New York–25
September 1877, shore of Lake Titicaca, Peru) in South America. His expedition team was composed mostly of
students from Williams College, and the expedition was made under the auspices of the Smithsonian Institution.
The expedition sailed from New York on 1 July 1867, crossed the Isthmus of Panama and reached Paita (NW coast
of Peru). Exploration started on 19 July 1867, in the coastal tropical forest of Guayaquil (Ecuador). The first “huge
cockroach” (Blaberus discoidalis) was caught in the city of Guayaquil (Orton 1870). The general route was from
Guayaquil up the Rio Guayas, over the western Cordillera near Chimborazo into the valley of Quito; thence over
the eastern Cordillera to the Rio Napo and down that river to the Marañón. Orton’s second expedition into South
America was in 1873 and the third in 1876. Orton died during the third expedition (Treadwell 1934; Miller 1982).
Additional faunistic records of Ecuadorian cockroaches were made by English entomologist and professor at
Oxford University John Obadiah Westwood (22 December 1805, Sheffield, Yorkshire–2 January 1893, Oxford)
in 1876. In an orchid bulb from Ecuador, he found nine living specimens of cockroaches comprising six different
species (Rhyparobia maderae, Nauphoeta cinerea, Blatta orientalis, Periplaneta americana and two unknown
species). The bulb was purchased by British naturalist William Chapman Hewitson (9 January 1806, Newcastleupon-Tyne-28
May 1878 Oatlands Park) with a collection of roots from Ecuador. Hewitson was wealthy collector
who purchased insects (especially Lepidoptera and Coleoptera), bird’s eggs and other products of nature from
travellers throughout the world (Evans 1908).
The excellent Spanish orthopterologist Ignacio Bolívar y Urrutia (9 November 1850, Madrid–20 November
1944, Mexico City) described four Ecuadorian cockroach species: Blabera aequatoriana (syn. of Blaberus
parabolicus), Zetobora martinezi (now Lanxoblatta martinezi), Anaplecta pallida, Blatta nigrita (incertae sedis)
and provided new Ecuadorian records of six species: Panchlora pulchella, Anaplecta lateralis, Ischnoptera
consobrina (syn. of Ischnoptera rufa), Chorisoneura pellucida (syn. of Chorisoneura diaphana), Chorisoneura
minuta, and Blatta adspersicolis (now Neoblattella adspersicollis) (Bolivar 1881). Two species new for Ecudorian
fauna (Panchlora viridis [probably Panchlora nivea] and Blattella germanica) were published by Bolivar (1884).
Quote specimens comes from two collectors:
The first group of specimens was collected during an important Spanish scientific expedition, “The Pacific
Scientific Commision”, by the Spanish entomologist Francisco de Paula Martínez y Sáez (30 March 1835,
Madrid–26 February 1908, Madrid). The members of this expedition began its survey of South America in San
Salvador (Brazil) on 9 September 1862. On 31 August 1864 they arrived at Guayaquil (Ecuador). They proceeded
to Quito (8 October 1864), then travelled across the Andes to River Napo. They also visited Cotopaxi Volcano
and on 24 August 1865 reached River Marañón near the Brazilian boundary. Collections were placed in Madrid on
18 January 1866, of which the insect collection contained 19,522 specimens, 10,427 of them caught by Martínez y
Sáez. The cockroach species Lanxoblatta martinezi was named by Ignacio Bolívar in his honour (Almagro 1866;
Martínez y Sáez 1994; López-Ocón 2003).
The second collection was assembled by Polish zoologist Jean Stanislaus Stolzmann (or Jan Sztolcman; 19
November 1854, Warsaw–28 April 1928, Warsaw), Director of the Zoological Museum of Branicki at Warsaw
from 1887 and Vice Director of the Polish Museum of Natural History from 1919. Stolzmann’s zoological
expeditions in Peru (1875–1881) and Ecuador (1882–1884) were financially covered by the Branicki brothers.
Stolzmann himself focused mainly on birds and mammals, but also collected insects. All cockroaches collected by
Stolzmann were caught at the locality “Palmal”. Identification of this locality is difficult. Palmal is a humid
VIDLIČKA404 · Zootaxa 3599 (5) © 2013 Magnolia Press
rainforest in the southern area of the western coast of Ecuador, near the city of Tumbes and the Tumbes River
(Taczanowski 1877; Taczanowski & Berlepsch 1885), and probably is identical with “Palmales”, situated 10 km
from the Peruvian border near the city of Arenillas. Tumbes is located on the northwest coast of Peru, 40 km from
Palmales.
The English mountaineer, explorer and illustrator Edward Whymper (27 April 1840, London–16 September
1911, Chamonix, France) organized a climbing expedition in Ecuador from 9 December 1879 to 27 July 1880.
During this expedition he also collected insects. Regarding his unique orthopteroid collection from high elevations
of Chimborazo Volcano, he wrote: “The whole of this collection remains undescribed. Leaping Orthoptera were
found very numerous up to the height of 12–13,000 feet, and upon Chimborazo were obtained in the Vallon de
Carrel at 16,000 feet” (4,877 m; Whymper 1892). Cockroaches specifically were mentioned from environs of Altar
Volcano near Riobamba, upon river Collane: “Blatta–Not identified–Valley of Collanes, Altar 12,500 feet” (3,810
m), where Whymper camped on 17–19 June 1880. At the end of expedition, when he was lodged in Guayaquil
(16–27 July 1880), he “collected fifty species of vermin in a single room”. One of this “vermin” was a big
cockroach of the genus Blaberus, displayed very faithfully in a table in Whymper (1892).
Swiss entomologist Henri Louis Frederic de Saussure and his collaborator Leo Zehntner (19 December
1864, Reigoldswil—3 April 1961 Liestal), assistant at the Natural History Museum in Geneva, described in 1895
one new cockroach species, Hormetica strumosa, from Ecuador but without detailed faunistic data.
The first comprehensive work on orthopteroid insects from Ecuador was published by the Italian entomologist
Ermanno Giglio-Tos (25 March 1865, Chiaverano, Turin–18 August 1926, Turin) (Giglio-Tos 1898). Among 30
listed cockroach species were 14 new records for Ecuador and 10 species new to science (Epilampra stigmosa,
Epilampra josephi, Panchlora festae, Zetobora aberrans, Ischnoptera josephina, Xestoblatta hamata, Xestoblatta
sancta, Neoblattella festae, Pelmatosilpha cothurnata, Lamproblatta zamorensis). Specimens studied by GiglioTos
are deposited in the Turin Museum of Natural History. This extensive collection was gathered by the Italian
naturalist Enrico Luigi Festa (11 August 1868, da Corrado e da Teodora Vola–30 September 1939, Moncalieri)
during an expedition in South America in May 1895–April 1898. He journeyed in Ecuador during 1896 and1897,
beginning in Guayaquil. He then travelled to Cuenca and afterwards to the headwaters of the Rio Santiago of the
Marañón system, and northwards by Quito to Julian, near the Colombian border (Festa 1909). During this
expedition Mr. Festa collected about 150,000 invertebrates.
English orthopterologist Robert Walter Campbell Shelford (3 August 1872, Singapore–22 June 1912,
Margate) started work in the Department of Zoology, Oxford University Museum as Assistant Curator in 1905,
studying Orthoptera and particularly Blattaria. During 1906 and 1907 he described eight new cockroach species
from Ecuador (Anaplecta fusca, Anaplecta pavida, Anaplecta varipennis, Pseudomops albostriatus, Pseudomops
bicolor, Pseudomops burri, Pseudomops magnus and Chorisoneura morosa) which originated from the collection
of William Frederick Henry Rosenberg (1868, Paddington, Hampstead-1957, Hendon, Middlesex). Rosenberg
was professional collector and dealer of exotic animals (including insects). His first expedition in 1894 explored
Colombia, where he remained about a year; and the second was conducted in western Ecuador from 2 November
1896 to September 1897. Collections of insects (chiefly butterflies, moths and beetles) from this expedition were
acquired by the Oxford Museum in 1899 and 1903 (Smith 1986). Rosenberg’s cockroach collection originated
from two localities: Cabaché – a small village near the river of that name (NW Coast, Prov. Esmeraldas), and
Paramba – a farm on the west bank of the River Mira (Prov. Imbabura) (Boulenger 1898).
The type specimen of the cockroach species from Ecuador described by Shelford in 1912 (Melyroidea
magnifica) came from the Saunders collection deposited in the Oxford Museum. William Wilson Saunders (4
June 1809, Little London near Wendover, Buckinghamshire–13 September 1879 Worthing, West Sussex) was a
British insurance broker, entomologist and botanist. He was an original member of the Entomological Society and
its President from 1841 to 1842 and again from 1856 to 1857. He was also Treasurer of the Linnean Society of
London (1861–1873) and a Fellow of several societies including the Royal Society and the Zoological Society.
Saunders himself probably never collected insects in Ecuador or in South America, but his collection contained
insects collected by other collectors. Orthopteroid insects (including Blattaria) were collected by the English
naturalist and explorer Henry Walter Bates (8 February 1825, Leicester–16 February 1892, London) during his
legendary 11-year expedition to the Amazon (1848–1859). Saunders sold his collection of cockroaches (467
specimens, containing Bates types) to the University Museum, Oxford, in 1873 (Smith 1986). The collector of
these Ecuadorian specimens is not clear.
Zootaxa 3599 (5) © 2013 Magnolia Press · 405COCKROACHES (BLATTARIA) OF ECUADOR
Shelford (1913) described three new species of cockroaches from Ecuador (Ceratinoptera alticola,
Eurycotis riveti and Zetha rufescens (syn. of Zetha vestita)) and four species recorded for the first time in
Ecuador. The collector of these specimens was a French ethnologist, Paul Adolph Rivet (7 May 1876, Wasigny,
Ardennes–25 March 1958, Paris). In the role of physician, he took part in the Second French Geodesic Mission
and scientific expedition (1899–1906), which arrived in Ecuador in 1901. He had spent six years in South
America and had worked on an ethnographic study of the Huaorani (Jívaro) people of the Ecuadorian Amazon.
In addition, he collected insects throughout Ecuador. When Rivet returned to France (1906), he donated his
collections to the Muséum National d’Histoire Naturelle in Paris. One new cockroach species (Eurycotis riveti)
was dedicated to the collector by Shelford.
The first checklist of Ecuadorian cockroaches was published by Ecuadorian scientist (entomologist),
researcher and educator Francisco Campos Ribadeneira (11 January 1879, Guayaquil–5 May 1962,
Guayaquil) in 1923. On the base of literature data and own collection he included 54 species in the checklist.
Several cockroaches collected by Campos were sent to Morgan Hebard for identification. Campos, as professor
of medical zoology at the University of Guayaquil, created the first entomological collection in Ecuador
(Barragán et al. 2009).
American orthopterologist Morgan Hebard (23 February 1887 Cleveland, Ohio–28 December 1946,
Philadelphia, Pennsylvania) determined not only material from the Campos collection but also cockroaches
deposited in the American Museum of Natural History (AMNH). The museum’s collection of South American
cockroaches consists mostly of specimens acquired by American mammalogist George Henry Hamilton Tate
(30 April 1894, Strout Green, London–24 December 1953, Morristown, New Jersey, USA), assistant and later
curator of the Department of Mammals in AMNH. During 1921–1929 he participated in several collecting
expeditions to various South American countries, including Ecuador (1922), Bolivia, Brazil, Venezuela, and
British Guyana. Tate assembled a large collection, especially mammals (Dorst 1954). Hebard (1924) published
an extensive work on Ecuadorian cockroaches, describing two species new to science (Panchlora irrorata and
Anaplecta falcifer) and providing eight new records for the country. Campos (1926) added these and other
species to the checklist to give a total of 67 cockroach species in Ecuador.
Another Ecuadorian cockroach was described about 15 years later. American entomologist Ashley Buell
Gurney (16 May 1911, Cummington, Massachusetts–10 March 1988, Montgomery, Maryland) began his career
at the United States Department of Agriculture in 1935, focusing on cockroaches. He started by processing
museum collections of cockroaches from North, South and Central America, including the West Indies. In a
1939 paper he analysed the genus Xestoblata and described Xestoblatta ecuadorana from Ecuador. All 3
specimens of this species (1 male and 2 females) were collected by a young American collector, Edward
Jackson Brundage Jr. (20 October 1915, Chicago, Illinois). These cockroach specimens were part of his
large collection of South American insects, which was sold to the United States National Museum in
Washington, D.C., in 1938 (Wetmore & Abbot 1939). Mr. Brundage previously (January 1938) donated almost
500 arthropods from United States, Panama and Colombia to the Field Museum of Natural History in Chicago
(Anonymous 1938). Gurney also donated 400 Ecuadorian cockroaches to the United States National Museum in
Washington, D.C., in 1938 (Wetmore & Abbot 1939).
Since 1960 a number of new species have been described from Ecuador: Macrophyllodromia ecuadorana,
Nahublattella ecuadorana, Xestoblatta bananae (Rocha e Silva Albuquerque 1962a, b, c); Xestoblatta
amedegnatae (Grandcolas 1992); Litopeltis compleptera (Roth & Gutiérrez 1998); Macrophyllodromia amabile,
Macrophyllodromia nobile (Anisyutkin 2007); Nahublattella alexandri, Nahublattella cuyabeno, Nahublattella
incurvata, Nahublattella reticulata, Nahublattella ultima (Anisyutkin 2009); Euphyllodromia propinqua,
Euphyllodromia rasnitsyni (Anisyutkin 2011).
The current checklist comes nearly 90 years after that of Campos (1926) and includes 105 species from
continental Ecuador and 18 species from Galápagos Islands.
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Checklist of cockroaches (Blattaria) from Ecuador
Structure of checklist
An italicized name of country means “terra typica”; i.e, the country from which the type specimens originate. A
question mark before the species name indicates the occurrence of the species in Ecuador is uncertain. A question
mark after the species name indicates that determination of the species is questionable. A question mark after the
name of a country suggests uncertain or problematic occurrence.
Family Blaberidae Brunner de Wattenwyl, 1865
Blaberus atropos (Stoll, 1813)
Synonymy:
Blatta atropos Stoll, 1813
Blabera fusca Brunner de Wattenwyl, 1865
Blabera laticollis Walker, 1868
Distribution: Trinidad, Guyana, Ecuador, Chile (terra typica was not defined).
Comments to faunistic records in Ecuador:
Bonsall (1995:33): Misahualli, Jatun Sacha (eventual confusion with Blaberus discoidalis).
Blaberus boliviensis Princis, 1946
Distribution: Bolivia, Ecuador.
Comments to faunistic records in Ecuador:
Princis (1952:6): 1 male, 3.IX.1948, Ecuador, Guayaquil (in a shipment of bananas from near Puna).
Roth & Willis (1960:147): the same data as mentioned by Princis (1952).
Roth (1969:220, 221, 241, 248): 1 male, Guayaquil (det. Princis).
Roth (1970b:310): the same data as mentioned by Roth (1969).
Blaberus craniifer (Burmeister, 1838)
Synonymy:
Blabera craniifer Burmeister, 1838
Blabera limbata Burmeister, 1838
Blabera trapezoidea Burmeister, 1838
Blabera varians Serville, 1838
Blabera luctuosa Stål, 1855
Blatta (Blabera) atropos Guérin-Méneville, 1857 non Stoll, 1813
Blabera ferruginea Saussure, 1864
Libisoca aequalis Walker, 1868
Blabera quadrifera Walker, 1868
Distribution: Mexico, Costa Rica, Guatemala, Honduras, Panama, Venezuela, Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:15) (Blabera trapezoidea): San José, Valle del Santiago, Gualaquiza.
Campos (1923:10 and 1926:55) (Blabera trapezoidea): the same locality as mentioned by Giglio-Tos (1898).
Maes (1992a:23): Ecuador.
Note: According to Roth (1969) Blaberus trapezoideus Burmeister, 1838 is synonym of Blaberus craniifer
Burmeister, 1838.
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Blaberus discoidalis (Serville, 1839)
Synonymy:
Blabera discoidalis Serville, 1839
Blabera varians Serville, 1839
Blabera cubensis Saussure, 1864
Blabera subspurgata Walker, 1868
Blabera rufescens Saussure & Zehntner, 1894
Distribution: Cuba, Haiti, Jamaica, Costa Rica, Dominican Republic, Puerto Rico, Vieques Island, Panama,
Colombia, Venezuela, Trinidad, Nicaragua, Ecuador.
Comments to faunistic records in Ecuador:
Scudder (1869:342) (Blabera cubensis): 2 specimens, Guayaquil.
Giglio-Tos (1898:15) (Blabera rufescens): 1 specimen, La Concepción, Valle del Mira.
Campos (1923:11 and 1926:55) (Blabera rufescens): the same locality as mentioned by Giglio-Tos (1898);
(Blabera discoidalis): Guayaquil, Durán, Naranjito, Barraganetal, Posorja, El Morro, Daule, Bucay, Vinces
(leg. F. Campos R.) and the same locality as mentioned by Hebard (1924).
Hebard (1924:130): 2 males, 1 female, Guayaquil, Guayas (leg. F. Campos R.), 1 male, Machala, El Oro (leg. Capt.
Moirel) and 1 female, Riobamba, Chimborazo (leg. R. Bourgeois).
Roth & Willis (1960:75): Ecuador.
Princis (1963:127): Ecuador ?.
Roth (1969:240, 248): 1 specimen, Ecuador.
Maes (1992a:23): Ecuador.
Note: Princis (1963:128) marked data of Giglio-Tos and Campos with “?” but Scudder’s and Hebard’s data (with
same localities) were not questioned.
Blaberus parabolicus (Walker, 1868)
Synonymy:
Blabera parabolica Walker, 1868
Blabera armigera Scudder, 1869
Blabera aequatoriana Bolívar, 1881
Distribution: Ecuador (+Galápagos Islands), Bolivia, Brazil, Colombia, Peru, Suriname.
Comments to faunistic records in Ecuador:
Walker (1868:8) (Blabera parabolica): 1 male, Cuenca (from Mr. Fraser’s collection).
Scudder (1869:343) (Blabera armigera): 1 male, Napo or Marañón (Ecuador or Peru).
Finot (1897:210) (Blabera parabolica): the same locality as mentioned by Walker (1868).
Bolivar (1881:479) (Blabera aequatoriana): Archidona and Coca, Ecuador (leg. Martínez y Saez).
Bolivar (1884:24) (Blabera aequatoriana): the same data as mentioned by Bolivar (1881).
Kirby (1904:164) (Blabera parabolica): Ecuador; (Blabera armigera): Napo?.
Campos (1923:10 and 1926:55, 56) (Blabera aequatoriana): the same localities as mentioned by Bolivar (1881);
(Blabera parabolica): the same locality as mentioned Walker (1868).
Rehn (1933a:58): Cuenca, Gualaquiza and valley of Santiago, Ecuador.
Princis (1958:74): the same locality as mentioned by Walker (1868); the study identifies as a female the holotype
described by Walker (1868).
Bruijning (1959:8): the same localities as mentioned by Rehn (1932).
Princis (1963:129): Ecuador.
Roth (1969:220, 221, 236, 248): Ecuador and the same locality as mentioned by Scudder (1869).
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2214): Isla San Cristóbal.
Peck (1996:1502): the same locality as mentioned by Peck & Roth (1992).
Peck et al. (1998:223): Isla San Cristóbal, Isla Santa Cruz.
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Eublaberus immaculus (Saussure & Zehntner, 1894)
Distribution: Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:106) (Blabera immacula): 1 male, Vinces.
Campos (1923:11 and 1926:55) (Blabera immacula): the same locality as mentioned by Giglio-Tos (1898).
Princis (1963:135): Ecuador.
Eublaberus posticus (Erichson, 1848)
Synonymy:
Blabera postica Erichson, 1848
Blatta ferruginea Stoll, 1813 [primary homonym]
Blabera femorata Scudder, 1869
Blabera thoracica Saussure & Zehntner, 1894
Blabera lindmani Shelford, 1911
Distribution: Guyana, Suriname, French Guiana, Brazil, Peru, Ecuador, Costa Rica, Nicaragua, Panama,
Colombia, Trinidad.
Comments to faunistic records in Ecuador:
Scudder (1869:342) (Blabera femorata): 1 male, Napo or Marañón.
Giglio-Tos (1898:15) (Blabera thoracica): Valle del Santiago.
Kirby (1904:165) (Blabera femorata): Napo.
Campos (1923:11 and 1926:55) (Blabera thoracica): the same locality as mentioned by Giglio-Tos (1898).
Princis (1963:134): Ecuador.
Hormetica strumosa Saussure & Zehntner, 1895
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Saussure & Zehntner (1895:56): 1 male, Ecuador (Equator).
Note: The syntype of Hormetica strumosa is deposited in Muséum d’Histoire Naturelle de la Ville de Genève,
Genève, Switzerland with the data label “[male] Ecuador” (J. Hollier, personal communication 2012).
Kirby (1904:197): Ecuador.
Hebard (1924:130): 1 juv. female, Cuenca, Azuay (leg. F. Campos R.).
Campos (1926:56): the same locality as mentioned by Hebard (1924).
Princis (1963:116): Ecuador.
Phoetalia circumvagans (Burmeister, 1838)
Synonymy:
Nauphoeta circumvagans Burmeister, 1838
Blatta marginicollis Stål, 1860
Distribution: Brazil, Ecuador, Colombia (Circumtropical).
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:13) (Nauphoeta circumvagans): La Concepción, Valle del Mira, Ibara.
Campos (1923:10 and 1926:54) (Nauphoeta circumvagans): the same locality as mentioned by Giglio-Tos (1898).
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Phoetalia pallida (Brunner de Wattenwyl, 1865)
Synonymy:
Nauphoeta pallida Brunner de Wattenwyl, 1865
Blatta laevigata Serville, 1839 non Palisot de Beauvois, 1805
Nauphoeta marginalis Walker, 1868
Distribution: Cuba, Ecuador (+Galápagos Islands), Colombia, Brazil (Circumtropical).
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:13) (Nauphoeta laevigata Pal. de Beauv.): Gualaquiza.
Campos (1923:6, 10 and 1926:50, 54) (Leurolestes pallidus): Guayaquil, Durán (leg. F. Campos R.), (Nauphoeta
laevigata Pal. de Beauv.): the same locality as mentioned by Giglio-Tos (1898).
Hebard (1924:128) (Leurolestes pallidus): 1 female, Guayaquil, Guayas (leg. F. Campos R.).
Bonsall (1995:33): Misahualli, Puerto Lopez.
Comments to faunistic records in Galápagos:
Howard (1889:194) (Nauphoeta laevigata ? (Pal.)): Galápagos Islands, 3 immature specimens (det. L. Bruner),
(according Hebard (1920b:317) these juveniles belong to Nauphoeta cinerea; Princis (1967:654) stated it as
Phoetalia circumvagans).
Scudder (1893:7) (Nauphoeta circumvagans): Galápagos Islands: Charles Island, 3 immature females (the same
specimens as studied by Howard (1889); according to Hebard (1920b:317) these juveniles belong to
Nauphoeta cinerea; Princis (1967:653) stated it as Phoetalia circumvagans).
McNeill (1901:489) (Nauphoeta circumvagans): Galápagos Islands: the same locality as mentioned by Scudder
(1893).
Hebard (1920b:316) (Leurolestes pallidus): Galápagos Islands: Chatham Island, 1 female, January 1906 (leg. F.X.
Williams).
Linsley & Usinger (1966:125) (Leurolestes pallidus): Galápagos Islands: San Cristóbal.
Peck & Roth (1992:2214): Galápagos Islands: Isla San Cristóbal (=Chatham), January (the same specimens as
studied by Howard (1889) and by Scudder (1893)), but they stated a different locality than Scudder (1893).
Peck (1996:1502) & Peck et al. (1998:223): Galápagos Islands: the same locality as mentioned by Peck & Roth
(1992).
Epilampra azteca Saussure, 1868
Distribution: Mexico, Cuba, Panama, Colombia, Ecuador, Venezuela, Trinidad, French Guiana, Costa Rica,
Suriname, Brazil.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:7): males and female, Valle dello Zamora.
Shelford (1910a:16): Ecuador.
Campos (1923:7 and 1926:50): the same locality as mentioned by Giglio-Tos (1898).
Princis (1967:694): Ecuador.
Roth (1970c:470): 1 male, Napo, Santa Cecilia, Rio Aguarico, Ecuador (det. Roth).
Epilampra conferta Walker, 1868
Synonymy:
Epilampra stigmosa Giglio-Tos, 1898
Distribution: Brazil, Bolivia, Colombia, Ecuador, Peru, Costa Rica, Panama.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:8) (Epilampra stigmosa sp.n.): 4 males, Valle del Santiago.
Kirby (1904:127) (Epilampra stigmosa): Ecuador.
Shelford (1910a:16) (Epilampra stigmosa): Ecuador.
Campos (1923:7 and 1926:50) (Epilampra stigmosa): the same locality as mentioned by Giglio-Tos (1898).
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Hebard (1924:128) (Epilampra stigmosa): 1 female, 2 juveniles, Allpa-yuca, Rio Pastaza, Tunguragua (from
Rosenberg).
Rehn (1933a:54) (Epilampra stigmosa): the same locality as mentioned by Giglio-Tos (1898) and Hebard (1924).
Princis (1948:10) (Epilampra stigmosa): the same locality as mentioned by Giglio-Tos (1898) and Hebard (1924).
Princis (1967:689): Ecuador.
Roth (1970c:441): 1 male, Napo, Santa Cecilia, Rio Aguarico (det. Roth), and the same data as mentioned by
Giglio-Tos (1898).
Note: Hebard (1926:201) states: “This species (E. opaca) may prove to be a synonym of the Brazilian E.
maculicollis (Serville), and the Ecuadorian E. stigmosa Giglio-Tos may fall in the same synonymy.“ Princis
(1958:63) synonymized E. stigmosa with E. conferta. According to Roth (1970c) Epilampra stigmosa is
probably a different species from E. conferta.
Epilampra josephi Giglio-Tos, 1898
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:9): 2 males, San José.
Shelford (1910a:16): Ecuador.
Campos (1923:7 and 1926:50): the same locality as mentioned by Giglio-Tos (1898).
Princis (1967:688): Ecuador.
Roth (1970c): the same data as mentioned by Giglio-Tos (1898).
Epilampra mexicana Saussure, 1862
Distribution: Mexico, Panama, Guatemala, Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:7): 1 female, Valle del Santiago.
Shelford (1910a:16): Ecuador.
Campos (1923:7 and 1926:50): the same locality as mentioned by Giglio-Tos (1898).
Princis (1946:153): Ecuador.
Princis (1967:689): Ecuador.
Roth (1970c:442) (Epilampra sp.): 1 male, Rio Blanco or Ecuador; this specimen was originally determined by
Rocha e Silva Albuquerque as Epilampra mexicana. Roth (1970c) stated that this male has genitalia very
different from E. mexicana.
Epilampra opaca Walker, 1868
Distribution: Guyana, Brazil, French Guiana, Suriname, Ecuador, Bolivia, Venezuela.
Comments to faunistic records in Ecuador:
Roth (1970c:460): 3 males, Napo, Santa Cecilia, Rio Aguarico, Ecuador.
Note: According to Princis (1958:62) E. opaca Walker is a synomym of E. substrigata Walker. Roth (1970c:465,
471) indicated, that E. opaca and E. substrigata are distinct species. Both species belong to subgroup B of the
Burmeisteri Group.
Epilampra substrigata Walker, 1868
Distribution: Brazil, Venezuela, Ecuador, Colombia (terra typica was not defined).
Comments to faunistic records in Ecuador:
Roth (1970c:461): 2 males, Napo, Santa Cecilia, Rio Aguarico, Ecuador (det. Roth and det. Princis).
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Homalopteryx laminata Brunner von Wattenwyl, 1892
Distribution: St. Vincent, Grenada, Trinidad, Venezuela, Colombia, Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:10): Valle del Santiago.
Campos (1923:8 and 1926:50): the same locality as mentioned by Giglio-Tos (1898).
Princis (1967:658): Ecuador ?.
Note: Princis (1967:658) mentioned the data of Giglio-Tos with “?”.
Litopeltis compleptera Roth & Gutiérrez, 1998
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Roth & Gutiérrez (1998:199): 4 females, Gualea, Ecuador (leg. F. Campos).
Note: It is the first record of the genus Litopeltis in South America. All other species are known from Central
America.
Rhyparobia maderae (Fabricius, 1781)
Synonymy:
Blatta maderae Fabricius, 1781
Blatta major Palisot de Beauvois, 1805
Blatta tuberculata Thunberg, 1810
Blatta maderensia Jones, 1859
Proscratea illepida Walker, 1868
Nauphoeta kukenthali Shelford, 1910
Distribution: Island Madeira, Ecuador (+Galápagos Islands), Colombia (Circumtropical).
Comments to faunistic records in Ecuador:
Westwood (1876:xxxii) (Blatta Maderae): in bulb of orchid from Ecuador.
Giglio-Tos (1898:11): La Concepción, Valle del Mira.
Shelford (1913:59): Santo Domingo de los Colorados.
Campos (1923:9 and 1926:52): Guayaquil, Durán, Naranjito, Bucay, Posorja, El Morro, Daule, Quito (leg. F.
Campos R.), Tambillo (leg. R.P. Luis Mille, S. J.) and the same locality as mentioned by Giglio-Tos (1898) and
Shelford (1913); vertical distribution in Ecuador: 0–2850 m.
Hebard (1924:128) (Leucophaea maderae): 2 males, 1 female, Guayaquil, Guayas (leg. F. Campos R.); 1 juv.
female, Tambillo, Pichincha (leg. F. Campos R.).
Gurney (1953:49) (Leucophaea maderae): Ecuador.
Roth & Willis (1960:151, 162): the same data as mentioned by Westwood (1876).
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2214): Isla San Cristóbal, Baquerizo Morena, 11–23.II.1989, 1 male (leg. S.B. Peck).
Peck (1996:1502) & Peck et al. (1998:223): the same locality as mentioned by Peck & Roth (1992).
Nauphoeta cinerea (Olivier, 1789)
Synonymy:
Blatta cinerea Olivier, 1789
Blatta gallica Fabricius, 1793
Blatta elegans Eschscholtz, 1822
Nauphoeta bivittata Burmeister, 1838
Nauphoeta grisea Burmeister, 1838
Distribution: Mauritius, Ecuador (+Galápagos Islands), Brazil (Circumtropical).
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Comments to faunistic records in Ecuador:
Westwood (1876:xxxii) (Blatta cinerea): in bulb of orchid from Ecuador.
Giglio-Tos (1898:106): 1 female, Vinces, 1 female, Guayaquil.
Campos (1923:10 and 1926:54): Guayaquil, Durán, Naranjito, Daule (leg. F. Campos R.) and the same locality as
mentioned by Giglio-Tos (1898).
Hebard (1924:130): 1 male, Guayaquil, Guayas (leg. F. Campos R.).
Roth & Willis (1960:151, 162): the same data as mentioned by Westwood (1876).
Comments to faunistic records in Galápagos:
Howard (1889:194) (Nauphoeta bivittata): 3 specimens, Chatham Island (det. L. Bruner).
Scudder (1893:6): the same exemplars as mentioned by Howard (1889).
Hebard (1920b:317): Chatham Island, 3 males, 4 females, 3 juv., October 1905; Charles Island, 6 males, 8 females,
3 juv., October 1905 (leg. F.X. Williams).
Linsley & Usinger (1966:125): San Cristóbal, Floreana.
Peck & Roth (1992:2214); Peck (1996:1502); Peck et al. (1998:223): the same locality as mentioned by Linsley &
Usinger (1966).
Achroblatta luteola (Blanchard, 1843)
Synonymy:
Blatta luteola Blanchard, 1843
Paratropes histrio Saussure, 1862
Pseudomops lituriceps Walker, 1868
Zetobora sigillata Walker, 1868
Panchlora tripartita Walker, 1868
Distribution: Bolivia, Mexico, Honduras, Costa Rica, Colombia, Brazil, French Guiana, Suriname, Ecuador,
Venezuela.
Comments to faunistic records in Ecuador:
Bruijning (1959:15): distribution from Mexico to Bolivia and French Guiana.
Gurney & Roth (1972:525): Ecuador, Guayaquil, intercepted at San Diego, Calif., with bananas, 1 female, 30
March 1953.
Panchlora exoleta Burmeister, 1838
Synonymy:
Panchlora punctum Saussure & Zehntner, 1893
Distribution: Brazil, Costa Rica, Panama, Colombia, Ecuador, Peru, Paraguay, Argentina, Venezuela, Suriname,
Jamaica, Mexico.
Comments to faunistic records in Ecuador:
Scudder (1869:342): 1 specimen, Napo or Marañón.
Hebard (1920a:106): 1 male, Alpayacu, eastern Ecuador.
Princis (1964:177): Ecuador.
Panchlora festae Giglio-Tos, 1898
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:12): 1 female, Valle del Santiago.
Kirby (1904: 154): Ecuador.
Campos (1923:9) and (1926:53): the same locality as mentioned by Giglio-Tos (1898).
Princis (1964:187): Ecuador.
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Panchlora irrorata Hebard, 1924
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Hebard (1924:129): 1 female, Bucay, Guayas, 1922 (leg. G.H.H. Tate).
Campos (1926:53): the same locality as mentioned by Hebard (1924).
Princis (1964:188): Ecuador.
Panchlora nigricornis Walker, 1868
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Walker (1868:28): Quito (M. Bourcier).
Finot (1897:203): Quito.
Hebard (1924:129): Ecuador.
Campos (1926:53): the same data as mentioned by Hebard (1924).
Princis (1964:188): Ecuador.
Panchlora nivea (Linnaeus, 1758)
Synonymy:
Blatta nivea Linnaeus, 1758
Blatta chlorotica Pallas, 1772
Blatta alba Strøm, 1783
Blatta hyalina Stoll, 1813
Blatta virescens Thunberg, 1826
Panchlora cubensis Saussure, 1862
Panchlora poeyi Saussure, 1862
Panchlora luteola Saussure, 1864
Ischnoptera lucida Walker, 1868
Pycnosceloides aporus Hebard, 1919
Distribution: Central and South America; Ecuador.
Comments to faunistic records in Ecuador:
Bolivar (1884:17) (Panchlora viridis Burmeister [sic]): Baeza, Ecuador (leg. Martínez y Saez)
Giglio-Tos (1898:12) (Panchlora virescens): Gualaquiza.
Campos (1923:9) and (1926:53) (Panchlora virescens): the same locality as mentioned by Giglio-Tos (1898);
(Panchlora cubensis): the same locality as mentioned by Hebard (1924).
Hebard (1924:128) (Panchlora cubensis): 1 male, 1 female, Ventura, Chimborazo, 12 April 1922 (leg. G.H.H.
Tate).
Panchlora peruana Saussure, 1864
Distribution: Peru, Ecuador, Brazil.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:12): S. José, Gualaquiza, Valle del Santiago.
Campos (1923:9) and (1926:53): the same locality as mentioned by Giglio-Tos (1898).
Princis (1964:186): Ecuador.
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Panchlora pulchella Burmeister, 1838
Distribution: Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Bolivar (1881:469): Palmal (leg. Stolzmann).
Campos (1923:10) and (1926:53): the same locality as mentioned by Bolivar (1881).
Pycnoscelus surinamensis (Linnaeus, 1758)
Synonymy:
Blatta surinamensis Linnaeus, 1758
Blatta indica Fabricius, 1775
Blatta melanocephala Stoll, 1813
Blatta corticum Serville, 1839
Perispherus laevis Le Guillou, 1841
Pycnoscelus obscurus Scudder, 1862
Panchlora celebesa Walker, 1868
Polyzosteria latipes Walker, 1868
Polyzosteria crassipes Walker, 1868
Panchlora occipitalis Walker, 1871
Epilampra tatei Tepper, 1894
Epilampra dimorpha Shiraki, 1906
Pycnoscelus surinamensis major Roeser, 1940
Pycnoscelus surinamensis minor Roeser, 1940
Distribution: Suriname; Ecuador (+Galápagos Islands) (Circumtropical).
Comments to faunistic records in Ecuador:
Walker (1868:166) (Polyzosteria crassipes): Napo River, South America (Mr. Stevens’ collection).
Finot (1897:197) (Polyzosteria crassipes): Napo River.
Giglio-Tos (1898:12) (Leucophaea surinamensis): La Concepcion, Valle del Mira.
Kirby (1904:196) (Parasphaeria ? crassipes): the same locality as mentioned by Walker (1868).
Campos (1923:10) and (1926:54): the same locality as mentioned by Giglio-Tos (1898).
Bonsall (1995:33): Misahualli.
Comments to faunistic records in Galápagos:
Butler (1877:87) (Panchlora surinamensis): Charles Island, 2 adult, 2 immature, June 1875 (leg. W. E. Cookson).
Howard (1889:194) (Leucophaea surinamensis): 8 specimens (adults and juv.), 1887–88 (det. L. Bruner).
Scudder (1893:7) (Leucophaea surinamensis): Chatham Island, 2 immature specimens obtained by Dr. Baur, and
specimens determined by Bruner (Howard 1889) from Charles and Chatham Islands.
McNeill (1901:494) (Leucophaea surinamensis): Chatham, 2 juv. females, and older data from 3 previous
expeditions: Charles Island and Chatham.
Hebard (1920b:317): Albemarle Island, Villamil, 1 female; Chatham Island, 3 males, 3 juv. females, October 1905;
Charles Island, 1 juv. female, October 1905 (leg. F.X. Williams).
Hebard (1934:280): Chatham Island, 1 female, 2 juveniles, 28 December 1925.
Linsley & Usinger (1966:125): Isabela, San Cristóbal, Floreana.
Parkin et al. (1972:102): 1 specimen, Floreana, 31 August 1968; 2 juveniles, Santa Cruz, 8 September 1968 (det.
D.R. Ragge).
Linsley (1977:4): Floreana, Isabela, Santa Cruz.
Peck & Roth (1992:2214): Isla Floreana, San Cristóbal, Santa Cruz, Islas Campéon, Fernandina, Gardner (at
Floreana), Isabela, Santiago.
Peck (1996:1502) & Peck (1998:223): the same locality as mentioned by Peck & Roth (1992).
Peck et al. (2001:152): Campéon, Española, Ferdinandina, Gardner at Floreana, Isabela, San Cristóbal, Santa Cruz,
Santiago (only older data).
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Lanxoblatta emarginata (Burmeister, 1838)
Synonymy:
Zetobora emarginata Burmeister, 1838
Zetobora cicatricosa Burmeister, 1838
Zetobora perspicua Walker, 1868
Distribution: Brazil, Guyana, French Guiana, Suriname, Ecuador, Colombia.
Comments to faunistic records in Ecuador:
Roth (1970a:228): 1 male, Ecuador.
Lanxoblatta martinezi (Bolivar, 1881)
Synonymy:
Zetobora martnezii Bolivar, 1881
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Bolivar (1881:478) (Zetobora martinezii): 1 female, Archidona, Ecuador (leg. Martínez y Saez).
Bolivar (1884:20) (Zetobora martinezii): the same data as mentioned by Bolivar (1881).
Giglio-Tos (1898:13) (Zetobora martinezii): 1 female, Valle del Santiago.
Kirby (1904:158) (Zetobora martinezi): Ecuador.
Campos (1923:10 and 1926:54) (Zetobora martinezii): the same locality as mentioned by Bolivar (1881) and by
Giglio-Tos (1898).
Princis (1964:259): Ecuador.
Lanxoblatta rudis (Walker, 1868)
Synonymy:
Zetobora rudis Walker, 1868
Distribution: Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Scudder (1869:342) (Zetobora rudis): 1 specimen, unlabelled, on either side of the Andes of equatorial South
America (James Orton coll.).
? Parasphaeria ovata (Blanchard, 1851)
Synonymy:
Blatta ovata Blanchard, 1851
Hormetica chilensis Saussure, 1862
Polyzosteria geissei Philippi, 1863
Polyzosteria valdiviana Philippi, 1863
Distribution: Chile, Ecuador ?.
Comments to faunistic records in Ecuador:
Shelford (1913:58): 1 male, Santo Domingo de los Colorados, 510 m.
Campos (1923:6 and 1926:56): the same locality as mentioned by Shelford (1913).
Princis (1964:240): Ecuador.
Note: According to Rehn (1933b:189) Parasphaeria ovata is limited in distribution to the southern half of Chile.
The unique male specimen examined by Shelford probably belongs to one of the other superficially similar
genera of Perisphaerinae.
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Zetobora aberrans Giglio-Tos, 1898
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:13): 1 male, San José.
Kirby (1904:158) (Tribonidium aberrans): Ecuador.
Campos (1923:10 and 1926:54): the same locality as mentioned by Giglio-Tos (1898).
Princis (1964:256): Ecuador.
Family: Anaplectidae Walker, 1868
Anaplecta alaris Saussure & Zehntner, 1893
Distribution: Peru, Ecuador, Brazil.
Comments to faunistic records in Ecuador:
Hebard (1924:121): 1 male, Ventura, Chimborazo, 12 April 1922 (leg. G.H.H. Tate).
Campos (1926:45): the same locality as mentioned by Hebard (1924).
? Anaplecta brunneri Shelford, 1906
Distribution: Brazil, Ecuador ?
Note: Princis (1965:369) listed this species as recorded in Ecuador, but detailed distribution data are available only
from Brazil.
Anaplecta cabimae Hebard, 1920
Distribution: Panama, Ecuador.
Comments to faunistic records in Ecuador:
Hebard (1924:122): 1 male, Ventura, Chimborazo, 12 April 1922 (leg. G.H.H. Tate).
Campos (1926:45): the same locality as mentioned by Hebard (1924).
Princis (1965:373): Ecuador.
Anaplecta falcifer Hebard, 1924
Distribution: Ecuador, Brazil.
Comments to faunistic records in Ecuador:
Hebard (1924:119): 11 males, 2 females, Ventura, Chimborazo, 1400 feet, 12 April 1922 (leg. G.H.H. Tate).
Campos (1926:45): the same locality as mentioned by Hebard (1924).
Rehn (1933a:11): western Ecuador.
Princis (1965:371): Ecuador.
Anaplecta fusca Shelford, 1906
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1906:246): Cachabi, December 1896, 1 female (W.F.H. Rosenberg coll., Oxford Museum).
Shelford (1907a:10): Ecuador.
Hebard (1921:222): the same locality as mentioned by Shelford (1906).
Hebard (1924:122): Ecuador.
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Campos (1926:45): the same locality as mentioned by Hebard (1924).
Princis (1965:368): Ecuador.
Anaplecta lateralis Burmeister, 1838
Distribution: Colombia, Panama, Venezuela, Brazil, Ecuador (+Galápagos Islands), Argentina.
Comments to faunistic records in Ecuador:
Bolivar (1881:463): Palmal, Ecuador (leg. Stolzmann).
Campos (1923:5 and 1926:44): the same locality as mentioned by Bolivar (1881).
Princis (1965:372): Ecuador.
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2206): Isla Santa Cruz, Puerto Ayora, Charles Darwin Research Station, 3 males, 3–7 March
1992; 5 km N Puerto Ayora, 1 male, 1–30 May 1991; 1.7 km N Santa Rosa, 3 males, 1 female, 1–30 May 1991.
Peck (1996:1502) & Peck et al. (1998:223): the same locality as mentioned by Peck & Roth (1992).
Anaplecta mexicana Saussure, 1868
Synonymy:
Anaplecta gemma Hebard, 1920
Distribution: Mexico, Costa Rica, Panama, Ecuador.
Comments to faunistic records in Ecuador:
Hebard (1924:119): 1 female, Rio Pescado, Guayas, 1600 feet, 15 May 1922 (leg. G.H.H. Tate).
Campos (1926:45): the same locality as mentioned by Hebard (1924).
Princis (1965:367): Ecuador.
Anaplecta pallida Bolivar, 1881
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Bolivar (1881:463): Palmal (leg. Stolzmann).
Kirby (1904:67): Ecuador.
Shelford (1907a:10): Ecuador.
Campos (1923:5 and 1926:45): the same locality as mentioned by Bolivar (1881).
Princis (1965:369): Ecuador.
Anaplecta pavida Shelford, 1906
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1906:245): Cachabi, December 1896, 1 female (W.F.H. Rosenberg coll., Oxford Museum).
Shelford (1907a:10): Ecuador.
Princis (1965:368): Ecuador.
Anaplecta varipennis Shelford, 1906
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1906:246): 5 females, Paramba, 3500 feet, May 1897 (W.F.H. Rosenberg coll., Oxford Museum).
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Shelford (1907a:10): Ecuador.
Princis (1965:368): Ecuador.
Maraca fossata Hebard, 1926
Distribution: French Guiana, Venezuela, Brazil, Ecuador, Suriname.
Comments to faunistic records in Ecuador:
Rocha e Silva Albuquerque (1965:2): Ecuador.
Family: Blattellidae Karny, 1908
Blattella germanica (Linnaeus, 1767)
Synonymy:
Blatta germanica Linnaeus, 1767
Blatta transfuga Brünnich, 1763
Blatta daurica Laxmann, 1769
Blatta asiatica Pallas, 1773
Blatta obliquata Daldorf, 1794
Blatta bivittata Serville, 1839
Ischnoptera paralella Tepper, 1893
Phyllodromia magna Tepper, 1895
Blattella germanica shuguroffi Karny, 1908
Phyllodromia niitakana Shiraki, 1931
Phyllodromia cunei-vittata Hanitsch, 1925
Blattella stylifera Chopard, 1938 (in part)
Distribution: Danmark, Ecuador (+Galápagos Islands) (Cosmopolitan).
Comments to faunistic records in Ecuador:
Bolivar (1884:12) (Blatta germanica): Baeza, Ecuador (leg. Martínez y Saez).
Giglio-Tos (1898:3) (Blatta germanica): Sig-sig (nec Sis-sig), Cuenca, La Concepción, Valle del Mira.
Shelford (1913:57) (Phyllodromia germanica): males and juv. females, Tulcan, Balsabamba, Riobamba.
Campos (1923:5 and 1926:46): Quito (leg. F. Campos R.); Tambillo, Cotocollao (leg. R.P. Luis Mille, S. J.);
Azógues, Biblián (Dr. Leoncio A. Cordero); Loja (Clodoveo Carrión); vertical distribution in Ecuador: 0–2980
m.
Hebard (1924:123): 1 female, Durán, Guayas, 25 June 1914 (leg. Herbert S. Parish); 1 male, Azogues, Cañar (leg.
F. Campos R.).
Bonsall (1995:33): Misahualli, Puerto Lopez, Jatun Sacha, Tena.
Comments to faunistic records in Galápagos:
Hebard (1920b:315): Hood Island (=Isla Española), 1 female, 1 June 1906 (leg. F.X. Williams).
Hebard (1929:358): Charles Island, 1 male, August 1924 (leg. Miss Cheesman).
Linsley & Usinger (1966:124): Hood (=Isla Española).
Peck & Roth (1992:2205): Isla Española (old data) and new records (1990–1992) only from boats based on San
Cristóbal and Santa Cruz.
Peck (1996:1502): Española, Santa Cruz.
Peck et al. (1998:223): the same locality as mentioned by Peck (1996).
Ischnoptera josephina Giglio-Tos, 1898
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:6): 4 specimens, San José.
Kirby (1904: 83): Ecuador.
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Shelford (1908a:8): Ecuador.
Campos (1923:6 and 1926:47): the same data as mentioned by Giglio-Tos (1898).
Princis (1969:746): Ecuador.
? Ischnoptera melasa Walker, 1868
Distribution: Brazil, Ecuador ?.
Comments to faunistic records in Ecuador:
Scudder (1869:342) (Ischnoptera melana): 1 specimen, Napo River.
Note: Species is presented by Scudder with “?”, because the specimen from Napo does not entirely agree with
Walker’s description. Scudder used an incorrect name, Ischnoptera melana, but his species description clearly
indicates Ischnoptera melasa and not Pseudomops melana (=melanus) as presented by Princis (1969:949).
Ischnoptera pallipes (Scudder, 1869)
Synonymy:
Phyllodromia pallipes Scudder, 1869
Distribution: Ecuador, Brazil, Peru.
Comments to faunistic records in Ecuador:
Scudder (1869:342) (Phyllodromia pallipes): 1 male, Napo or Marañón.
Kirby (1904:96) (Phyllodromia pallipes): the same locality as mentioned by Scudder (1869).
Ischnoptera peckorum Roth, 1988
Distribution: Ecuador–Galápagos Islands (endemic).
Comments to faunistic records in Galápagos:
Roth (1988:307): St. Cruz, 1 km east of Bellavista, 210 m, Bellavista Cave No. 2, 14 July 1985, S. & J. Peck leg.
Peck (1990:371); Peck & Roth (1992:2210); Peck (1996:1502) & Peck (2001:151): the same locality as mentioned
by Roth (1988).
Ischnoptera rufa (De Geer, 1773)
Synonymy:
Blatta rufa De Geer, 1773
Blatta rufescens Palisot de Beauvois, 1805
Ischnoptera fumata Burmeister, 1838
Ischnoptera consobrina Saussure, 1862
Ischnoptera rubiginosa Walker, 1868
Ischnoptera terminalis Walker, 1868
Ischnoptera conformis Saussure & Zehntner, 1893
Distribution: Suriname, Caribbean, Costa Rica, Jamaica, Nicaragua, Panama, Colombia, Guyana, Trinidad,
Ecuador, Brazil.
Comments to faunistic records in Ecuador:
Bolivar (1881:463) (Ischnoptera consobrina): Palmal (leg. Stolzmann).
Bolivar (1884:14) (Ischnoptera consobrina): Pichincha (leg. Martínez y Saez).
Campos (1923:6 and 1926:48) (Ischnoptera consobrina): the same data as mentioned by Bolivar (1881).
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Ischnoptera santacruzensis Roth, 1992
Distribution: Ecuador–Galápagos Islands (endemic).
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2210): Isla Santa Cruz: Puntudo, Cerro Crocker, Los Gemelos, many males and females.
Peck (1996:1502) & Peck (2001:151): the same locality as mentioned by Peck & Roth (1992).
Ischnoptera snodgrassi (McNeill, 1901)
Synonymy:
Temnopteryx snodgrassii McNeill, 1901
Distribution: Ecuador—Galápagos Islands (endemic).
Comments to faunistic records in Galápagos:
McNeill (1901:493) (Temnopteryx snodgrassii): Albemarle Island (=Isabela), Iguana Cove Mountain, 2000 feet, 1
male, 3 nymphs.
Hebard (1920b:315) (Anisopygia snodgrassii): Albemarle Island (=Isabela), 1 male, 1 female, 3 nymphs, 9 June
1899 (the same exemplars as mentioned by McNeill (1901)).
Linsley & Usinger (1966:124) (Anisopygia snodgrasii): The same locality as mentioned by McNeill (1901).
Princis (1969:774) (Anisopygia snodgrasii): Galápagos Islands.
Peck & Roth (1992:2006–2210): Isla Isabela (Cerro Azul volcano, Sierra Negra, Volcano Chico on Sierra Negra,
Volcano Alcedo).
Peck (1996:1502): the same locality as mentioned by Peck & Roth (1992).
Peck (2001:151): Isabela (Cerro Azul, Sierra Negra, Volcano Alcedo, Volcano Wolf).
Ischnoptera taczanowskii Bolivar, 1881
Distribution: Peru, Ecuador.
Comments to faunistic records in Ecuador:
Hebard (1924:123): 1 male, Bucay, Guayas, 1922 (leg. G.H.H. Tate); 1 male, 1 female, Milagro, Guayas, December
1922 (leg. G.H.H. Tate); 1 male, Ventura, Chimborazo, 12 April 1922 (leg. G.H.H. Tate); 1 female,
Chaguarpata, Chimborazo, 5 April 1922 (leg. G.H.H. Tate)
Campos (1926:48): the same locality as mentioned by Hebard (1924).
Princis (1969:745): Ecuador.
Pseudomops albostriatus Shelford, 1906
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1906:259) (Pseudomops albostriata): 1 female, Cachabi, November 1896 (W.F.H. Rosenberg coll.,
Oxford Museum).
Shelford (1908a:4) (Pseudomops albostriata): Ecuador.
Princis (1969:947): Ecuador.
Pseudomops bicolor Shelford, 1906
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1906:260): 1 male, Paramba, March 1897 (W.F.H. Rosenberg coll., Oxford Museum).
Shelford (1908a:4): Ecuador.
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Rehn (1933a:50): the same locality as mentioned by Shelford (1906).
Princis (1969:947): Ecuador.
Pseudomops burri Shelford, 1906
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1906:257): 3 females, Cachabi, December 1896 (W.F.H. Rosenberg coll., Oxford Museum).
Shelford (1908a:4): Ecuador.
Princis (1969:952): Ecuador.
Pseudomops magnus Shelford, 1906
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1906:259) (Pseudomops magna): Paramba, 1 male, 2 females, February and May 1897 (W.F.H.
Rosenberg coll., Oxford Museum).
Shelford (1908a:4) (Pseudomops magna): Ecuador.
Princis (1969:947): Ecuador.
Symploce pallens (Stephens, 1835)
Synonymy:
Ectobius pallens Stephens, 1835
Blatta capitata Saussure, 1862
Blatta deprivata Walker, 1868
Phyllodromia hospes Perkins, 1899
Symploce lita Hebard, 1916
Ischnoptera platystyla Chopard, 1938
Symploce benzoni Princis, 1951
Symploce kevani Chopard, 1954
Symploce vicentina Princis, 1959
Cahita gracilis Rocha e Silva, 1965
Distribution: Cuba, Ecuador—Galápagos Islands (Circumtropical).
Comments to faunistic records in Galápagos:
Hebard (1920b:316) (Symploce lita): Chatham Island (= Floreana), 1 male, October 1905 (leg. F.X. Williams).
Hebard (1934:279) (Symploce hospes): Charles Island (= Floreana), Postoffice Bay, 16 males, 2 females, 12
juveniles, 2 oothecae, 20 August –15 September 1925.
Linsley & Usinger (1966:125) (Symploce lita): San Cristóbal.
Roth (1984:51): Charles Island, 1 male, 1 September 1925, 1 male, 10–14 September 1925, 1 male, 20–30 August
1925, 1 female, 1–15 September 1925, Wollebaeks (all determined by Hebard as S. hospes); Santa Cruz I.,
0–100 m, 2 males, September 1970 (J. & M. Sedlacek)
Peck & Roth (1992:2206): Islas Floreana, San Cristóbal, Santa Cruz, Bartolomé, Campeón, Fernandina, Genovesa,
Isabela, Marchena, Pinta, Santa Fe, Santiago.
Peck (1996:1502) & Peck et al. (1998:223): the same locality as mentioned by Peck & Roth (1992).
Xestoblatta amedegnatae Grandcolas, 1992
Distribution: Ecuador.
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Comments to faunistic records in Ecuador:
Grandcolas (1992b:163): Napo, Tena, Misahualli, Jatun Sacha, 25 March 1990 and 30 March 1990 (leg. S.
Poulain).
Xestoblatta bananae Rocha e Silva Albuquerque, 1962
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Rocha e Silva Albuquerque (1962c:244): 1 male, intercepted in plant quarantine from Ecuador, at San Diego,
Calif., 14 April 1953, with bananas, through R. F. Wilkey; 1 female, intercepted in plant quarantine from
Ecuador, in California, 30 March 1953.
Princis (1969:896): Ecuador ?.
Xestoblatta ecuadorana Gurney, 1939
Distribution: Ecuador, Bolivia.
Comments to faunistic records in Ecuador:
Gurney (1939:106): 1 male, 1 female, Baños–Mera Trail, Rio Pastaza watershed, Ecuador, 22 September 1937, at
an altitude of 1200 m (leg. E.J. Brundage Jr.); 1 female, Puyo, Oriente, Ecuador, 900 m, 14 October 1937 (leg.
E.J. Brundage, Jr.).
Princis (1948:14): the same locality as mentioned by Gurney (1939).
Princis (1969:896): Ecuador.
Xestoblatta hamata (Giglio-Tos, 1898)
Synonymy:
Ischnoptera hamata Giglio-Tos, 1898
Distribution: Ecuador, Colombia, Panama, Brazil.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:4) (Ischnoptera hamata): 2 females, Valle del Santiago, Ecuador; 1 male, Gualaquiza, Ecuador.
Shelford (1908a:8) (Ischnoptera hamata): Ecuador.
Hebard (1916:371): the same data as mentioned by Giglio-Tos (1898).
Campos (1923:6 and 1926:47) (Ischnoptera hamata): the same data as mentioned by Giglio-Tos (1898).
Rehn (1933a:31): eastern Ecuador.
Princis (1969:899): Ecuador.
Xestoblatta sancta (Giglio-Tos, 1898)
Synonymy:
Ischnoptera sancta Giglio-Tos, 1898
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:5) (Ischnoptera sancta): 5 females, San José, Ecuador.
Kirby (1904: 83) (Ischnoptera sancta): Ecuador.
Shelford (1908a:8) (Ischnoptera sancta): Ecuador.
Hebard (1916:371): the same data as mentioned by Giglio-Tos (1898).
Campos (1923:6 and 1926:47) (Ischnoptera sancta): the same data as mentioned by Giglio-Tos (1898) and Hebard
(1924).
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Hebard (1924:124): 1 female, Pasaje, Del Oro (leg. F. Campos R.); 1 female, Chaguarpata, Chimborazo, 5 April
1922 (leg. G.H.H. Tate).
Gurney (1939:117): the same data as mentioned by Hebard (1924).
Rehn (1933a:31): eastern Ecuador.
Princis (1969:899): Ecuador.
Eushelfordia pica (Walker, 1868)
Synonymy:
Paratropes pica Walker, 1868
Distribution: Brazil, Ecuador, Peru.
Comments to faunistic records in Ecuador:
Campos (1923:8 and 1926:49) (Paratropa/Paratropes pica): Baños (leg. F. Campos R.).
Hebard (1924:126): 1 female, Baños, Tunguragua (leg. F. Campos R.).
Princis (1967:633): Ecuador.
Megaloblatta blaberoides (Walker, 1871)
Synonymy:
Epilampra blaberoides Walker, 1871
Megaloblatta rufipes Dohrn, 1887
Distribution: Nicaragua, Costa Rica, Panama, Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:6) (Megaloblatta rufipes): 1 male, 1 female, Valle del Santiago; 1 male, San José; 1 nymph,
Gualaquiza.
Campos (1923:7 and 1926:49) (Megaloblatta rufipes): Bucay, Chimbo, Baños (leg. F. Campos R.) and the same
data as mentioned by Giglio-Tos (1898).
Princis (1967:632): Ecuador ?.
Megaloblatta longipennis (Walker, 1868)
Synonymy:
Blabera longipennis Walker, 1868
Megaloblatta peruviana Dohrn, 1887
Distribution: Ecuador, Panama, Peru.
Comments to faunistic records in Ecuador:
Walker (1868:8) (Blabera longipennis): 1 male, Guayaquil (Mr. Stevens’ collection).
Finot (1897:210) (Blabera longipennis): the same locality as mentioned by Walker (1868).
Kirby (1904:109): the same locality as mentioned by Walker (1868).
Shelford (1908b:4): Ecuador.
Hebard (1920a:92): 1 female, Ecuador.
Campos (1923:41): Bucay (leg. F. Campos R.).
Hebard (1924:127): 1 female, San Domingo de los Colorados, Pichincha (leg. F. Campos R.).
Campos (1926:49): the same data as mentioned by Hebard (1924).
Princis (1967:631): Ecuador.
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Megaloblatta regina (Saussure, 1870)
Synonymy:
Blabera regina Saussure, 1870
Distribution: Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Hebard (1924:127): 1 female, 1 juv. male, ootheca, Allpa-yacu, Rio Pastaza, Tunguragua, 3600 feet (Rosenberg).
Campos (1926:49): the same locality as mentioned by Hebard (1924).
Gurney (1959:133): 1 female, Banos, Ecuador.
Princis (1967:631): Ecuador.
Bell et al. (2007:6): 1 ootheca, Ecuador.
Nyctibora glabra Giglio-Tos, 1897
Distribution: Bolivia, Peru, Brazil, Ecuador, Argentina.
Comments to faunistic records in Ecuador:
Shelford (1913:58): 1 female, 1 larve, Santo Domingo de los Colorados.
Campos (1923:7 and 1926:48): the same data as mentioned by Shelford (1913).
Princis (1967:626): Ecuador.
Note: Giglio-Tos (1897:9) listed only localities from Peru (La Masa) and Bolivia. Hebard (1921:271) mentioned
the same localities as Giglio-Tos (1897): San Francisco and Caiza (San Francisco Solano in Gran Chaco,
Bolivia and Caiza, Yacuíba, Bolivia), but both were incorrectly placed in Ecuador.
? Nyctibora humeralis Dohrn, 1888
Distribution: Brazil, Ecuador ?, Bolivia ?.
Note: Princis (1967:624) reported the occurrence of this species in Brazil, Ecuador and Bolivia, but published data
are available only from Brazil: Fonte Boa (Brazil, Alto Amazonas). Literature data: Dohrn (1888:129): Alto
Amazonas, Fonteboa; Kirby (1904:107): Upper Amazonas; Shelford (1908b:2): Upper Amazonas; Rocha e
Silva Albuquerque (1964a:18): Amazonas.
Nyctibora noctivaga Rehn, 1902
Distribution: Nicaragua, Panama, Jamaica, Virgin-Islands, Colombia, Ecuador.
Comments to faunistic records in Ecuador:
Campos (1923:41): San Remo (leg. F. Campos R.).
Hebard (1924:126): 1 male, San Xavier [A. N. S. P.]; 1 female, San Remo, Los Rios (leg. F. Campos R.).
Campos (1926: 48): Bucay and the same data as mentioned by Hebard (1924).
Princis (1967:628): Ecuador ?.
Note: Princis (1967:628) used “?” in the case of Ecuador and in the Campos papers, but not in the case of Hebard
paper, athough the same Ecuadorian locality is mentioned.
Nyctibora obscura Saussure, 1864
Distribution: Brazil, Colombia, Trinidad, Ecuador, Peru.
Comments to faunistic records in Ecuador:
Hebard (1924:125): 1 female, Chaguarpata, Chimborazo, 5 April 1922 (leg. G.H.H. Tate).
Campos (1926:48): the same data as mentioned by Hebard (1924).
Princis (1967:626): Ecuador.
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Paratropes aequatorialis Saussure, 1864
Synonymy:
Paratropa lanceolatus Walker, 1868
Distribution: Ecuador, Bolivia, Brazil, Peru.
Comments to faunistic records in Ecuador:
Saussure (1864:309): 1 female, Ecuador (Respublica Equator).
Note: The syntype of Paratropes aequatorialis is deposited in Muséum d’Histoire Naturelle, Genève, Switzerland
with the data label “Quito, Andes, [female] M. H. de Saussure” (J. Hollier, personal communication 2012).
Walker (1868:150) (Paratropes lanceolatus): 1 female, Cuenca (Mr. Fraser’s collection).
Finot (1897:190) (Paratropa aequatorialis): Ecuador (République de l’Équateur).
Kirby (1904:109): Ecuador.
Shelford (1908b:3): Ecuador.
Princis (1958:72): the same locality as mentioned by Walker (1868).
Princis (1967:619): Ecuador.
Family: Pseudophyllodromiidae Hebard, 1929
? Amazonina conspersa (Brunner de Wattenwyl, 1865)
Synonymy:
Phyllodromia conspersa Brunner de Wanttenwyl, 1865
Neoblattella sooretamensis Rocha e Silva Albuquerque, 1958
Distribution: Brazil, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Brazil, Peru, Paraguay, Argentina,
Ecuador ?.
Comments to faunistic records in Ecuador:
Rocha e Silva Albuquerque (1974:57): Ecuador, Rio Branco (2,000 m alt.).
Note: In other Rocha papers, the locality Rio Branco is mentioned in Amazonas (Brazil) – Rocha e Silva
Albuquerque (1964a, 1972).
Ceratinoptera alticola Shelford, 1913
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1913:57): 2 males and 5 larves, Borma (3,100 m), Loja.
Campos (1923:7 and 1926:48): Loja (Clodoveo Carrión) and same data as Shelford (1913).
Princis (1969:762): Ecuador.
Ceratinoptera picta Brunner de Wattenwyl, 1865
Synonymy:
Phyllodromia conspersa Brunner de Wanttenwyl, 1865
Phyllodromia binotata Bruner, 1906
Distribution: Brazil, Guatemala, Panama, Colombia, Trinidad, Suriname, French Guiana, Ecuador, Peru.
Comments to faunistic records in Ecuador:
Hebard (1924:122): 1 female, Rio Pescado, Azuay, 1600 feet, 15 May 1922 (leg. G.H.H. Tate).
Campos (1926:48): the same data as mentioned by Hebard (1924).
Princis (1969:760): Ecuador.
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Chorisoneura carpenteri Roth, 1992
Distribution: Ecuador–Galápagos Islands (endemic).
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2210): Isla Isabela: Santo Tomas, Siera Negro, Volcan Siera Negro, Volcan Alcedo, Volcan
Cerro Azul; Isla Santa Cruz: Academy Bay, Los Gemelos, Media Luna, Puntudo, Santa Rosa; Isla Floreana:
Black Beach; Isla Santiago: Aguacate Camp.
Peck (1996:1502): the same locality as mentioned by Peck & Roth (1992).
Chorisoneura cristobalensis Roth, 1992
Distribution: Ecuador–Galápagos Islands (endemic).
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2213): Isla San Cristóbal – El Junco.
Peck (1996:1502): the same locality as mentioned by Peck & Roth (1992).
Chorisoneura diaphana Princis, 1965
Synonymy:
Blatta pellucida Saussure, 1864
Distribution: Mexico, Ecuador, Colombia ?.
Comments to faunistic records in Ecuador:
Bolivar (1881:468) (Chorisoneura pellucida): Palmal (leg. Stolzmann).
Campos (1923:9 and 1926:52) (Chorisoneura pellucida): the same locality as mentioned by Bolivar (1881).
Chorisoneura minuta Saussure, 1869
Distribution: Argentina, Ecuador.
Comments to faunistic records in Ecuador:
Bolivar (1881:469): Palmal (leg. Stolzmann).
Campos (1923:9 and 1926:52): the same locality as mentioned by Bolivar (1881).
Princis (1965:339): Ecuador.
Chorisoneura morosa Shelford, 1907
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1907b:44): 1 specimen, Cachabi, December 1896 (W.F.H. Rosenberg coll., Oxford Museum).
Princis (1965:340): Ecuador.
Chorisoneura translucida (Saussure, 1864)
Synonymy:
Blatta translucida Saussure, 1864
Distribution: Mexico, Guatemala, Costa Rica, Panama, Colombia, Ecuador.
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Comments to faunistic records in Ecuador:
Giglio-Tos (1898:11): 1 specimen, Valle del Santiago.
Campos (1923:9 and 1926:52): the same locality as mentioned by Giglio-Tos (1898).
Princis (1965:340): Ecuador ?.
Euphyllodromia erythromelas Rehn, 1932
Distribution: Peru, Colombia, Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2011:383): 7 males, 5 females, prov. Sucumbíos, E of town Lago Agrio, November 2005 (coll. A.
Gorochov, A. Ovtshinnikov).
Euphyllodromia hystrix (Saussure, 1869)
Synonymy:
Pseudophyllodromia hystrix Saussure, 1869
Pseudophyllodromia histrio Saussure, 1870
Distribution: Venezuela, Colombia, Ecuador, Peru, Costa Rica.
Comments to faunistic records in Ecuador:
Hebard (1924:124): 10 females, Rio Pescado, Azuay, 1600 feet, 15 May 1922 (leg. G.H.H. Tate).
Hebard (1929:360) (Euphyllodromia histrio): 1 female, Ecuador.
Campos (1926:47): the same locatity as mentioned by Hebard (1924).
Princis (1969:937): Ecuador.
Anisyutkin (2011:383): 1 male, prov. Sucumbíos, E of town Lago Agrio, November 2005 (coll. A. Gorochov, A.
Ovtshinnikov).
Euphyllodromia nigrochlamys Rehn, 1928
Distribution: Peru, Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2011:394): 9 males, 3 females, prov. Sucumbíos, SE of town Lago Agrio, November 2005 (coll. A.
Gorochov, A. Ovtshinnikov); 1 male, prov. Morona Santiago, bank of Rio Morona, January 2010 (coll. A.
Gorochov).
Euphyllodromia peruana (Saussure, 1864)
Synonymy:
Blatta peruana Saussure, 1864
Distribution: Peru, Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2011:378): 2 males, 1 female, prov. Sucumbíos, E of town Lago Agrio, November 2005 (coll. A.
Gorochov, A. Ovtshinnikov); 1 male, 1 female, prov. Morona Santiago, bank of Rio Morona, January 2010
(coll. A. Gorochov).
Euphyllodromia propinqua Anisyutkin, 2011
Distribution: Ecuador.
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Comments to faunistic records in Ecuador:
Anisyutkin (2011:390): 5 males, 5 females, prov. Sucumbíos, SE (or E) of town Lago Agrio, November 2005 (coll.
A. Gorochov, A. Ovtshinnikov).
Euphyllodromia rasnitsyni Anisyutkin, 2011
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2011:391): 1 male, 2 females, 95 km E of city Quito, November 2005 (coll. A. Gorochov, A.
Ovtshinnikov).
Euphyllodromia venezuelica Princis, 1951
Distribution: Venezuela, Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2011:380): 1 male, prov. Sucumbíos, SE of town Lago Agrio, November 2005 (coll. A. Gorochov, A.
Ovtshinnikov).
? Imblattella albida (Saussure, 1869)
Synonymy:
Blatta albida Saussure, 1869
Distribution: Colombia, Ecuador ?.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:2) (Blatta albida): 1 female, San José (determination is questionable).
Campos (1923:5 and 1926:45) (Blatta albida): the same locality as mentioned by Giglio-Tos (1898).
Princis (1969:797): Ecuador.
? Latiblattella vitrea (Brunner de Wattenwyl, 1865)
Synonymy:
Phyllodromia vitrea Brunner de Wattenwyl, 1865
Blatta latimargo Walker, 1868
Blatta alaris Saussure & Zehntner, 1893
Blatta maya Saussure & Zehntner, 1893
Distribution: Mexico, Honduras, Costa Rica, Ecuador ?.
Comments to faunistic records in Ecuador:
Campos (1923:6) (Phyllodromia vitrea ?) (determination is questionable) and (1926:47) (Phyllodromia vitrea ?):
Bucay (leg. F. Campos R.).
Lophoblatta brevis Rehn, 1937
Distribution: Guyana, Venezuela, Suriname, Ecuador, Brazil.
Comments to faunistic records in Ecuador:
Rocha e Silva Albuquerque (1964b:4): Ecuador, Puyo, 900m, 16 males, 9 females, 14 October 1937 (Clarke,
Macintyre, Brundage coll.).
Princis (1969:785): Ecuador.
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Lopes & Oliveira (2003:346): Ecuador, Pastaza: Puyo, 900m, 2 males, 1 female, 14 October 1937 (Clarke,
Macintyre, Brundage col., Museu Nacional, Universidade Federal do Rio de Janeiro).
Lopes (2004:345): Ecuador (on the map of South America).
Lophoblatta speerae Roch e Silva Albuquerque & Gurney, 1963
Distribution: Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Rocha e Silva Albuquerque (1964b:6): Ecuador, Puyo, 900m, 1 male, 14 October 1937 (Clarke, Macintyre,
Brundage coll.).
Princis (1969:786): Ecuador.
Lopes & Oliveira (2003b:352): Ecuador.
Lopes (2004:345): Ecuador (on the map of South America).
Macrophyllodromia amabile Anisyutkin, 2007
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2007:29): 1 male, 1 female, E. Ecuador, env. of waterfall San Rafael on Rio Coca, 95 km E of city
Quito, forest, 1300 m (leg. A. Gorochov, A. Ovitshinnikov).
Macrophyllodromia ecuadorana Rocha e Silva Albuquerque, 1962
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Rocha e Silva Albuquerque (1962a:425): 1 male, Guayaquil, Ecuador, intercepted at San Diego, California, in
bananas, 14 April 1953 (leg. R. Wilkey); 1 female, Ecuador, intercepted in California, 16 April 1953, in
bananas (leg. R. Wilkey).
Princis (1969:773): Ecuador.
Macrophyllodromia nobile Anisyutkin, 2007
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2007:32): 1 male, E. Ecuador, 80–85 km E of town Lago Agrio, env. of lake Lago Grande, Rio
Cuyabeno (leg. A. Gorochov, A. Ovitshinnikov).
Melyroidea magnifica Shelford, 1912
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1912:375): 1 female, Ecuador (Saunders collection, Oxford museum).
Princis (1965:357): Ecuador.
Nahublattella alexandri Anisyutkin, 2009
Distribution: Ecuador.
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Comments to faunistic records in Ecuador:
Anisyutkin (2009:611): 7 males, E. Ecuador, 80–85 km E of Lago Agrio, near lake Lago Grande (Rio Cuyabeno),
2–9 November 2005 (leg. A. Gorochov, A. Ovchinnikov).
Nahublattella cuyabeno Anisyutkin, 2009
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2009:610): 1 male, E. Ecuador, 80–85 km E of Lago Agrio, near lake Lago Grande (Rio Cuyabeno),
2–9 November 2005 (leg. A. Gorochov, A. Ovchinnikov).
Nahublattella ecuadorana Rocha e Silva Albuquerque, 1962
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Rocha e Silva Albaquerque (1962b:114): 1 male, Ecuador, on bananas, in cargo at San Pedro, CA, USA, 22 August
1959; 1 female, Ecuador, on bananas at New Orleans, LA, USA, 20 February 1953.
Princis (1969:795): Ecuador.
Nahublattella incurvata Anisyutkin, 2009
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2009:611): 1female, W Ecuador, 10 km E of Aqua Blanca, env. of Puerto Lopez, 26–29 October 2005
(leg. A. Gorochov, A. Ovchinnikov).
Nahublattella reticulata Anisyutkin, 2009
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2009:621): 1 male, 2 females, E. Ecuador, 80–85 km E of Lago Agrio, near lake Lago Grande (Rio
Cuyabeno), 2–9 November 2005 (leg. A. Gorochov, A. Ovchinnikov).
Nahublattella ultima Anisyutkin, 2009
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Anisyutkin (2009:616): 2 males, E. Ecuador, 70 km NE of Lago Agrio, near S. Pablo de Kantesiya Vill., on Rio
Aguarico, 10–17 November 2005 (leg. A. Gorochov, A. Ovchinnikov); 1 male, 80–85 km E of Lago Agrio,
near lake Lago Grande (Rio Cuyabeno), 2–9 November 2005 (leg. A. Gorochov, A. Ovchinnikov).
Neoblattella adspersicollis (Stål, 1860)
Synonymy:
Blatta adspersicollis Stål, 1860
Distribution: Brazil, Ecuador.
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Comments to faunistic records in Ecuador:
Bolivar (1881:466) (Blatta adspersicolis): Palmal (leg. Stolzmann).
Campos (1923:5 and 1926:46) (Blatta adspersicolis): the same locality as mentioned by Bolivar (1881).
Neoblattella festae (Giglio-Tos, 1898)
Synonymy:
Blatta festae Giglio-Tos, 1898
Distribution: Ecuador, Venezuela ?.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:2) (Blatta Festae): 1 male, 1 female, Gualaquiza, Ecuador.
Kirby (1904: 94) (Phyllodromia festae): Ecuador.
Shelford (1908a:14) (Phyllodromia festae): Ecuador.
Campos (1923:5 and 1926:46) (Blatta Festae): the same data as mentioned by Giglio-Tos (1898).
Rehn (1903:268) (Blattella festae): the same data as mentioned by Giglio-Tos (1898).
Rehn (1949:12) (Blattella festae): the same data as mentioned by Giglio-Tos (1898).
Princis (1969:794): Ecuador.
Note: Based on the Campos papers Princis (1969:898) listed by mistake Ecuador as the country of distribution for
Xestoblatta festae (Griffini, 1896). Campos evidently referred to Neoblattella festae (Giglio-Tos, 1898).
Supella longipalpa (Fabricius, 1798)
Synonymy:
Blatta longipalpa Fabricius, 1798
Blatta (Phyllodromia) supellectilium Serville, 1839
Blatta cubensis Saussure, 1862
Blatta phalerata Saussure, 1863
Blatta extenuata Walker, 1868
Blatta incisa Walker, 1868
Ischnoptera quadriplaga Walker, 1868
Ischnoptera vacillans Walker, 1868
Blatta subfasciata Walker, 1871
Blatta transversalis Walker, 1871
Distribution: Southern America (“India orientali”), Ecuador (Circumtropical).
Comments to faunistic records in Ecuador:
Campos (1923:6 and 1926:47) (Supella supellectilium): Guayaquil, Durán, Yaguachi, Milagro, Naranjito, San
Rafael, Bucay, Posorja, El Morro (leg. F. Campos R.).
Hebard (1924:122) (Supella supellectilium): 2 males, 3 females, Guayaquil, Guayas (leg. F. Campos R.).
Hebard (1929:358) (Supella supellectilium): 1 male, Ecuador (leg. G. Hammond).
Family: Blattidae Stephens, 1835
Blatta orientalis Linnaeus, 1758
Synonymy:
Blatta lucifuga Poda, 1761
La Blatte des cuisines Geoffroy, 1762
Blatta secunda Schaeffer, 1769
Blatta tertia Schaeffer, 1769
Blatta culinaris De Geer, 1773
Blatta ferruginea Thunberg, 1810
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Blatta europaea Bartsch, 1846
Blatta castanea Blanchard, 1851
Blatta hemialata Gistel, 1856
Pulex imperator Westwood, 1858
Blatta badia Saussure, 1863
Kakerlac pallipes Philippi, 1863
Kakerlac platystetho Philippi, 1863
Stylopyga orientalis gracilis Adelung, 1903
Stylopyga orientalis spontanea Semenov-Tian-Shansky, 1909
Distribution: America, Oriente; Ecuador (Cosmopolitan).
Comments to faunistic records in Ecuador:
Westwood (1876:xxxii): in bulb of orchid from Ecuador.
Roth & Willis (1960:151, 162): the same data as mentioned by Westwood (1876).
Pelmatosilpha cothurnata (Giglio-Tos, 1898)
Synonymy:
Eurycotis cothurnata Poda, 1761
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:11) (Eurycotis cothurnata): Guajaquil (sic), Canar, S. José, Cuenca.
Kirby (1904:144) (Eurycotis cothurnata): Ecuador.
Shelford (1910b:12) (Eurycotis cothurnata): Ecuador.
Shelford (1913:58) (Eurycotis cothurnata): 1 female, Cuenca.
Campos (1923:8 and 1926:51) (Eurycotis cothurnata): the same locality as mentioned by Giglio-Tos (1898) and by
Hebard (1924).
Hebard (1924:125) (Eurycotis cothurnata): 1 male, Cuenca, Azuay, (Captain Noirel); 1 male, 1 female, 7800 feet,
12 July 1922 (leg. G.H.H. Tate); 1 female, Mount Chimborazo, 12467 feet (leg. Captain Noirel).
Gurney (1965:7): Ecuador.
Princis (1966:462): Ecuador.
Pelmatosilpha praestans Dohrn, 1887
Synonymy:
Periplaneta aterrima Walker, 1869
Distribution: Peru, Brazil, Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1913:58) (Pelmatosilpha aterrima): 1 male, Santo Domingo de los Colorados.
Campos (1923:9 and 1926:52) (Pelmatosilpha aterrima): the same locality as mentioned by Shelford (1913) and
by Hebard (1924).
Hebard (1924:125): 1 male, Canelos, Rio Bobonaza, Oriente (leg. Rosenberg).
Gurney (1965:7): Ecuador.
Princis (1966:461): Ecuador.
Periplaneta americana (Linnaeus, 1758)
Synonymy:
Blatta americana Linnaeus, 1758
Blatta ferrugineo-fusca Gronovius, 1764
Blatta kakerlac De Geer, 1773
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Blatta orientalis Sulzer, 1776 non Linnaeus, 1758
Blatta aurelianensis Fourcroy, 1785
Blatta siccifolia Stoll, 1813
Blatta heros Eschscholtz, 1822
Periplaneta stolida Walker, 1868
Periplaneta americana colorata Rehn, 1902
Distribution: America, Ecuador (+ Galápagos Islands) (Cosmopolitan).
Comments to faunistic records in Ecuador:
Scudder (1869:342): 1 specimen, Napo or Marañón.
Westwood (1876:xxxii) (Blatta Americana): in a bulb of orchid from Ecuador.
Giglio-Tos (1898:106): 1 male, Vinces
Campos (1923:8 and 1926:51): Guayaquil, Durán, Naranjito, Posorja (leg. F. Campos R.) and same locality as
Giglio-Tos (1898).
Roth & Willis (1960): the same information as mentioned by Westwood (1879).
Bonsall (1995:33): Misahualli, Puerto Lopez, Jatun Sacha, Tena.
Comments to faunistic records in Galápagos:
Butler (1877:87): Charles Island, 2 adult specimens, June 1875 (leg. W.E. Cookson).
Howard (1889:193): Chatham Island, 10 specimens, 1887–88 (det. L. Bruner).
Scudder (1893:6): only the same exemplars as stated by Butler (1877) and by Howard (1889).
Hebard (1934:279): Charles Island, (= Floreana), Postoffice Bay, 1 female, 1 juv., 15–19 November 1925.
Linsley & Usinger (1966:125): San Cristóbal, Floreana.
Peck & Roth (1992:2205): Islas Floreana, San Cristóbal, Genovesa, Isabela, Santa Cruz (Bellavista, El Chato).
Peck (1996:1051); Peck et al. (1998:223) & Peck (2001:149): the same data as mentioned by Peck & Roth (1992).
Periplaneta australasiae (Fabricius, 1775)
Synonymy:
Blatta aurantiaca Stoll, 1813
Periplaneta zonata Haan, 1842
Periplaneta inclusa Walker, 1868
Periplaneta rapenda Walker, 1868
Periplaneta subcincta Walker, 1868
Periplaneta emittens Walker, 1871
Polyzosteria subornata Walker, 1871
Distribution: Ecuador (+ Galápagos Islands) (Circumtropical); (terra typica was not defined).
Comments to faunistic records in Ecuador:
Scudder (1869:342): 2 specimens, Napo or Marañón.
Bolivar (1884:16): Coca, Ecuador (leg. Martínez y Saez).
Giglio-Tos (1898:10): Gualaquiza, Valle dello Zamora, La Concepcion nella valle del Mira – Ecuador.
Shelford (1913:59): Santo Domingo de los Colorados, Riobamba, Balsabamba.
Campos (1923:8): Guayaquil, Durán, Naranjito, San Rafael, Bucay, Posorja, El Morro (leg. F. Campos R.) and the
same locality as mentioned by Giglio-Tos (1898).
Hebard (1924:125): 1 male, Guayaquil, Guayas, (leg. F. Campos R.); 1 female, Rio Zamora, Loja (leg. F. Campos
R.).
Campos (1926:50): Daule (leg. F. Campos R.), Portoviejo (leg. Justino Cornejo V.) and the same locality as
mentioned by Giglio-Tos (1898), Shelford (1913) and Campos (1923).
Roth & Willis (1960:80): Ecuador (according Campos (1926)).
Bonsall (1995:33): Misahualli, Puerto Lopez, Jatun Sacha, Tena.
Comments to faunistic records in Galápagos:
Howard (1889:194): Charles Island, 3 images and 3 nymphs, 1887–88 (det. L. Bruner).
Scudder (1893:6): only the same exemplars as mentioned by Howard (1889).
McNeill (1901:494): Chatham, 1 male, and the same locality as mentioned by Howard (1889).
Hebard (1920b:316): Chatham Island, 1 female, October 1905–January 1906; Charles Island, 3 males, 6 females,
October 1905 (leg. F.X. Williams).
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Hebard (1929:364): Charles Island, 1 juv. female, August 1924 (leg. Miss Cheesman).
Hebard (1934:279): Charles Island, (= Floreana), Postoffice Bay, 2 males, 3 females, 1 juv., 5 August–25
November 1925.
Linsley & Usinger (1966:125): San Cristóbal, Floreana.
Parkin et al. (1972:102): 1 specimen, Floreana, 31 August 1968 (det. D.R. Ragge).
Maes (1992b:3): Galápagos.
Peck & Roth (1992:2205): Islas Floreana, San Cristóbal, Isabela, Santa Cruz.
Peck (1996:1501); Peck (1998:223) & Peck (2001:149): the same data as mentioned by Peck & Roth (1992).
Periplaneta brunnea Burmeister, 1838
Synonymy:
Periplaneta patens Walker, 1868
Periplaneta concolor Walker, 1868
Periplaneta truncata Krauss, 1892
Periplaneta ignota Shaw, 1925
Distribution: Chile and Demerary, Ecuador (+ Galápagos Islands) (Circumtropical).
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:106) (Periplaneta truncata): Guayaquil.
Campos (1923:8 and 1926:51) (Periplaneta truncata): Guayaquil, Durán (leg. F. Campos R.).
Bonsall (1995:33): Misahualli, Puerto Lopez, Jatun Sacha, Tena.
Comments to faunistic records in Galápagos:
Hebard (1920b:316): Chatham Island, 1 female, January 1906 (leg. F.X. Williams).
Linsley & Usinger (1966:125): San Cristóbal.
Peck & Roth (1992:2205): Isla San Cristóbal (according Hebard (1920b)).
Peck (1996:1501); Peck et al. (1998:223) & Peck (2001:149): the same data as metioned by Peck & Roth (1992).
Lamproblatta zamorensis (Giglio-Tos, 1898)
Synonymy:
Stylopyga zamorensis Giglio-Tos, 1898
Distribution: Ecuador, Peru.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:10) (Stylopyga zamorensis): 1 female, Valle del Zamora.
Kirby (1904: 139) (Blatta zamorensis): Ecuador.
Shelford (1910b:15) (Stylopyga zamorensis): Ecuador.
Campos (1923:8 and 1926:51) (Stylopyga zamorensis): the same locality as mentioned by Giglio-Tos (1898).
Princis (1966:546): Ecuador.
Roth (2003a:286): Ecuador.
Eurycotis riveti Shelford, 1913
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Shelford (1913:58) (Eurycoti Riveti): 3 females, 1 nymph, Loja.
Campos (1923:8 and 1926:52): the same locality as mentioned by Shelford (1913).
Princis (1966:555): Ecuador.
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Family: Polyphagidae Walker, 1868
Compsodes mexicanus (Saussure, 1868)
Synonymy:
Latindia mexicana Saussure, 1868
Latindia tolteca Saussure & Zehntner, 1894
Distribution: Mexico, Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:15) (Latindia mexicana): 1 specimen, Gualaquiza.
Campos (1923:10 and 1926:54) (Latindia mexicana): the same locality as mentioned by Giglio-Tos (1898).
Princis (1963:104): Ecuador ?.
Holocompsa nitidula (Fabricius, 1781)
Synonymy:
Blatta nitidula Fabricius, 1781
Holocompsa capsoides Shelford, 1911
Corydia collaris Burmeister, 1838
Distribution: Suriname, Brazil, Ecuador–Galápagos Islands (Circumtropical).
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2205): Isla Isabela, 4 males, 3 females, 9 March 1989.
Peck (1996:1501); Peck (2001:149): the same locality as mentioned by Peck & Roth (1992).
Holocompsa sp.
Distribution: Ecuador—Galápagos Islands.
Comments to faunistic records in Galápagos:
Peck & Roth (1992:2205): Isla Santa Cruz (Academia Bay, 15 March 1970; Mirador, 10 July 1970), 3 males
different from H. nitidula.
Peck (1996:1501): the same locality as mentioned by Peck & Roth (1992).
Zetha simonyi (Krauss, 1892)
Synonymy:
Holocompsa simonyi Krauss, 1892
Blatta vestita Brullé, 1838 [primary homonym]
Holocompsa chavesi Bolívar, 1894
Zetha freyi Chopard, 1942
Zetha rufescens Shelford, 1913
Distribution: Canary Islands, Azores Islands, Kapeverde, Peru, Ecuador, Guatemala, Hawai.
Comments to faunistic records in Ecuador:
Shelford (1913:59) (Zetha rufescens sp.n.): 1 female, Loja.
Campos (1923:10 and 1926:55) (Zetha rufescens): the same locality as mentioned by Shelford (1913).
Princis (1963:89): Ecuador.
VIDLIČKA436 · Zootaxa 3599 (5) © 2013 Magnolia Press
Species incertae sedis
Blatta (?) nigrita Bolivar, 1881
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Bolivar (1881:477(31)) (Blatta nigrita): Baeza, Ecuador (leg. Martínez y Saez).
Bolivar (1884:12) (Phyllodromia nigrita): the same locality as mentioned by Bolivar (1881).
Kirby (1904:96) (Phyllodromia nigrita): Ecuador.
Shelford (1908a:14) (Phyllodromia nigrita): Ecuador.
Campos (1923:5 and 1926:46) (Blatta nigrita): the same locality as mentioned Bolivar (1881).
Princis (1969:1010): Ecuador.
Note: More accurate classification is not possible in this species. The type material is not preserved.
Theganopteryx (?) sp. (Anaplectinae)
Distribution: Ecuador.
Comments to faunistic records in Ecuador:
Giglio-Tos (1898:2): 1 specimen, Valle del Zamora.
Campos (1923:4 and 1926:44): the same locality as mentioned by Giglio-Tos (1898).
Note: All known species from the genus Theganopteryx occur only in Africa and Madagascar. The determination
was evidently erroneous.
In conclusion, records of 114 cockroach species (Blattaria) in the territory of Ecuador 105 in continental
Ecuador and 18 in Galápagos Islands) are summarized. These species belong to 6 families and 42 genera (Table 1).
Occurrence of another 8 species is questionable. Forty species (38.5 %) were recorded exclusively from the
territory of continental Ecuador. Five of 18 species living on the Galápagos Islands (27.8 %) are endemic.
Currently, the continental Ecuadorian cockroach fauna shares the highest number of species with the Brazilian
(43.3%) and Colombian faunas (28.8 %) (Table 2). However, further studies on Ecuadorian as well as other
Neotropical cocroaches are needed.
TABLE 1. Number of genera and species of cockroaches in continental Ecuador and in Galápagos Islands. Species of
questionable occurrence are not included.
Family
Ecuador (continental) Galápagos Island Ecuador + Galápagos
Genera Species Genera Species Genera Species
Blaberidae 14 33 5 5 14 33
Anaplectidae 2 10 1 1 2 10
Blattellidae 8 22 3 5 9 27
Pseudophyllodromiidae 9 28 1 2 9 30
Blattidae 5 8 1 3 5 8
Polyphagidae 2 2 1 2 3 4
Species incertae sedis 2 2 0 0 2 2
Total 42 105 12 18 44 114
Zootaxa 3599 (5) © 2013 Magnolia Press · 437COCKROACHES (BLATTARIA) OF ECUADOR
TABLE 2. Distribution of Ecuadorian cockroach species in surrounding countries.
Species / Country Ec GI Br Co Pe Ve GS Ar Bo Ch
Blaberidae
Blaberus atropos (Stoll, 1813) X X X
Blaberus boliviensis Princis, 1946 X X
Blaberus craniifer (Burmeister, 1838) X X
Blaberus discoidalis (Serville, 1839) X X X
Blaberus parabolicus (Walker, 1868) X X X X X X X
Eublaberus immaculus (Saussure & Zehntner, 1894) X X
Eublaberus posticus (Erichson, 1848) X X X X X X
Hormetica strumosa Saussure & Zehntner, 1895 X
Phoetalia circumvagans (Burmeister, 1838) X X X x x x x
Phoetalia pallida (Brunner von W., 1865) X X X X x x x x
Epilampra azteca Saussure, 1868 X X X X X
Epilampra conferta Walker, 1868 X X X X X
Epilampra josephi Giglio-Tos, 1898 X
Epilampra mexicana Saussure, 1862 X
Epilampra opaca (Walker, 1868) X X X X X
Epilampra substrigata Walker, 1868 X X X X
Homalopteryx laminata Brunner von W., 1892 X X X
Litopeltis compleptera Roth & Gutiérrez, 1998 X
Rhyparobia maderae (Fabricius, 1781) X X x X x X x x
Nauphoeta cinerea (Olivier, 1789) X X X x x x x x
Achroblatta luteola (Blanchard, 1843) X X X X X X X X
Panchlora exoleta Burmeister, 1838 X X X X X X X
Panchlora festae Giglio-Tos, 1898 X
Panchlora irrorata Hebard, 1924 X
Panchlora nigricornis Walker, 1868 X
Panchlora nivea (Linnaeus, 1758) X X X x X x x
Panchlora peruana Saussure, 1864 X X X
Panchlora pulchella Burmeister, 1838 X X
Pycnoscelus surinamensis (Linnaeus, 1758) X X X X x X X X x x
Lanxoblatta emarginata (Burmeister, 1838) X X X X
Lanxoblatta martinezi (Bolivar, 1881) X
Lanxoblatta rudis (Walker, 1868) X X
Parasphaeria ovata (Blanchard, 1851) ? X
Zetobora aberrans Giglio-Tos, 1898 X
Anaplectidae
Anaplecta alaris Saussure & Zehntner, 1893 X X X
Anaplecta brunneri Shelford, 1906 ? X
Anaplecta cabimae Hebard, 1920 X
......to be continued on the next page
VIDLIČKA438 · Zootaxa 3599 (5) © 2013 Magnolia Press
TABLE 2. (continued)
Species / Country Ec GI Br Co Pe Ve GS Ar Bo Ch
Anaplecta falcifer Hebard, 1924 X X
Anaplecta fusca Shelford, 1906 X
Anaplecta lateralis Burmeister, 1838 X X X X X X
Anaplecta mexicana Saussure, 1868 X
Anaplecta pallida Bolivar, 1881 X
Anaplecta pavida Shelford, 1906 X
Anaplecta varipennis Shelford, 1906 X
Maraca fossata Hebard, 1926 X X X X
Blattellidae
Blattella germanica (Linnaeus, 1767) X X X X x X x X x X
Ischnoptera josephina Giglio-Tos, 1898 X
Ischnoptera melasa Walker, 1868 ? X
Ischnoptera pallipes (Scudder, 1869) X X X
Ischnoptera peckorum Roth, 1988 X
Ischnoptera rufa (De Geer, 1773) X X X X
Ischnoptera santacruzensis Roth, 1992 X
Ischnoptera snodgrassi (McNeill, 1901) X
Ischnoptera taczanowskii Bolivar, 1881 X X
Pseudomops albostriatus Shelford, 1906 X
Pseudomops bicolor Shelford, 1906 X
Pseudomops burri Shelford, 1906 X
Pseudomops magnus Shelford, 1906 X
Symploce pallens (Stephens, 1835) x X X x x x x x
Xestoblatta amedegnatae Grandcolas, 1992 X
Xestoblatta bananae Rocha e Silva, 1962 X
Xestoblatta ecuadorana Gurney, 1939 X X
Xestoblatta hamata (Giglio-Tos, 1898) X X X
Xestoblatta sancta (Giglio-Tos, 1898) X
Eushelfordia pica (Walker, 1868) X X X
Megaloblatta blaberoides (Walker, 1871) X
Megaloblatta longipennis (Walker, 1868) X X
Megaloblatta regina (Saussure, 1870) X X
Nyctibora glabra Giglio-Tos, 1897 X X X X X
Nyctibora humeralis Dohrn, 1888 ? X ?
Nyctibora noctivaga Rehn, 1902 X X
Nyctibora obscura Saussure, 1864 X X X X
Paratropes aequatorialis Saussure, 1864 X X X X
Pseudophyllodromiidae
Amazonina conspersa (Brunner v. Wattenwyl, 1865) ? X X X X X
......to be continued on the next page
Zootaxa 3599 (5) © 2013 Magnolia Press · 439COCKROACHES (BLATTARIA) OF ECUADOR
TABLE 2. (continued)
Species / Country Ec GI Br Co Pe Ve GS Ar Bo Ch
Ceratinoptera alticola Shelford, 1913 X
Ceratinoptera picta Brunner von W., 1865 X X X X X
Chorisoneura carpenteri Roth, 1992 X
Chorisoneura cristobalensis Roth, 1992 X
Chorisoneura diaphana Princis, 1965 X ?
Chorisoneura minuta Saussure, 1869 X X
Chorisoneura morosa Shelford, 1907 X
Chorisoneura translucida (Saussure, 1864) X X X
Euphyllodromia erythromelas Rehn, 1932 X X X X
Euphyllodromia hystrix (Saussure, 1869) X X X X
Euphyllodromia nigrochlamys Rehn, 1928 X X
Euphyllodromia peruana (Saussure, 1864) X X X
Euphyllodromia propinqua Anisyutkin, 2011 X
Euphyllodromia rasnitsyni Anisyutkin, 2011 X
Euphyllodromia venezuelica Princis 1951 X X
Imblattella albida (Saussure, 1869) ? X
Latiblattella vitrea (Brunner von W., 1865) ?
Lophoblatta brevis Rehn, 1937 X X X X
Lophoblatta speerae Rocha e Silva & Gurney, 1963 X X
Macrophyllodromia amabile Anisyutkin, 2007 X
Macrophyllodromia ecuadorana Rocha e Silva, 1962 X
Macrophyllodromia nobile Anisyutkin, 2007 X
Melyroidea magnifica Shelford, 1912 X
Nahublattella alexandri Anisyutkin, 2009 X
Nahublattella cuyabeno Anisyutkin, 2009 X
Nahublattella ecuadorana Rocha e Silva, 1962 X
Nahublattella incurvata Anisyutkin, 2009 X
Nahublattella reticulata Anisyutkin, 2009 X
Nahublattella ultima Anisyutkin, 2009 X
Neoblattella adspersicollis (Stål, 1860) X X
Neoblattella festae (Giglio-Tos, 1898) X ?
Supella longipalpa (Fabricius, 1798) X X X x X x X x x
Blattidae
Blatta orientalis Linnaeus, 1758 X X x X X X
Pelmatosilpha cothurnata (Giglio-Tos, 1898) X
Pelmatosilpha praestans Dohrn, 1887 X X X
Periplaneta americana (Linnaeus, 1758) X X x X x X x X x X
Periplaneta australasiae (Fabricius, 1775) X X X X X X X x x x
Periplaneta brunnea Burmeister, 1838 X X X X x X X X x X
......to be continued on the next page
VIDLIČKA440 · Zootaxa 3599 (5) © 2013 Magnolia Press
Abbreviations: Ec—Ecuador, GI—Galápagos Islands, Br—Brazil, Co—Colombia, Pe—Peru, Ve—Venezuela, GS—Guyana,
Suriname and French Guiana, Ar—Argentina, Bo—Bolivia, Ch—Chile; X—occurrence based on published data,
?—questionable occurrence, (?)—questionable status in classification, x—expected occurrence of circumtropical and
synantropic species
Acknowledgements
The study was supported by the Slovak Research and Development Agency (APVV) through financial support no.
APVV-0213-10. I thank Dr. Ladislav Roller and Dr. Juraj Majtan for valuable comments.
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Príloha č. 12
VRŠANSKÝ, P., CIFUENTES-RUIZ, P., VIDLIČKA, Ľ., ČIAMPOR, F., Vega, F.J. 2011.
Afro-Asian cockroach from Chiapas amber
and the lost Tertiary American entomofauna.
Geologica Carpathica 62: 463-475.
www.geologicacarpathica.sk
GEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICA, OCTOBER 2011, 62, 5, 463—475 doi: 10.2478/v10096-011-0033-8
Introduction
Chiapas amber is peculiar for its content of plant remains,
occasional fungi, scarce pulmonate gastropods, arachnids,
crustaceans, relatively abundant insects, amphibians, reptiles,
bird feathers and mammal hair (Poinar 2003; Engel
2004; Solórzano-Kraemer 2007; Vega et al. 2009a,b). Insects
are represented by 235 species all of which belong to
146 living families and the living or closely related (n=3)
genera. The termite Kalotermes nigritus Snyder, 1946
(Isoptera, Kalotermitidae), which still lives in South America
today (Zherikhin 1970) is famous.
It is worth mentioning that the earliest living insect species
are much older, originating from Eocene Baltic amber. These
species are Tetracha carolina (Linnaeus, 1766) (Coleoptera,
Cicindelidae), Colasposoma metallicum Clark, 1865; Palaeomymar
anomalum (Blood & Kryger, 1922) (Hymenoptera,
Mymaridae), Cupes tesselatus (Motschulsky, 1856) (Coleoptera,
Cupedidae) and Setodes picescens Ulmer, 1912 (Trichoptera,
Leptoceridae). Drosophila rubrostriata Becker,
1908 is known from the Oligocene of Fontainebleau.
The specimen reported here was collected from the Los
Pocitos locality (Fig. 1) of the amber-bearing Mazantic Shale,
Afro-Asian cockroach from Chiapas amber and the lost
Tertiary American entomofauna
PETER VRŠANSKÝ1,2
, PAULINA CIFUENTES-RUIZ3
, UBOMÍR VIDLIČKA4,5
, FEDOR ČIAMPOR JR4
and FRANCISCO J. VEGA3
1
Geological Institute, Slovak Academy of Sciences, Dúbravská cesta 9, P.O. BOX 106, 840 05 Bratislava, Slovak Republic;
geolvrsa@savba.sk
2
Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117868 Moscow, Russia; lab@palaeoentomolog.ru
3
Instituto de Geología, UNAM, Ciudad Universitaria, Coyoacán, 04510 México D.F., Mexico; vegver@unam.mx
4
Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 845 06 Bratislava, Slovak Republic;
uzaevidl@savba.sk; f.ciampor@savba.sk
5
Department of Biology, Faculty of Education, Comenius University, Moskovská 3, 813 34 Bratislava, Slovak Republic;
vidlicka@fedu.uniba.sk
(Manuscript received January 28, 2011; accepted in revised form March 17, 2011)
Abstract: Cockroach genera with synanthropic species (Blattella, Ectobius, Supella, Periplaneta, Diploptera and ?Blatta),
as well as other insects such as honeybees, although natively limited to certain continents nowadays, had circumtropic
distribution in the past. The ease of their reintroduction into their former range suggests a post-Early Miocene environmental
stress which led to the extinction of cosmopolitan Tertiary entomofauna in the Americas, whilst in Eurasia,
Africa and Australia this fauna survived. This phenomenon is demonstrated here on a low diversity (10 spp.) living
cockroach genus Supella, which is peculiar for the circumtropical synanthropic brownbanded cockroach S. longipalpa
and also for its exclusively free-living cavicolous species restricted to Africa. S. (Nemosupella) miocenica sp. nov. from
the Miocene amber of Chiapas in Mexico is a sister species to the living S. mirabilis from the Lower Guinea forests and
adjacent savannas. The difference is restricted to the shape of the central macula on the pronotum, and size, which may
indicate the around-Miocene origin of the living, extremely polymorphic Supella species and possibly also the isochronic
invasion into the Americas. The species also has a number of characteristics of the Asian (and possibly also
Australian) uniform genus Allacta (falling within the generic variability of Supella) suggesting Supella is a direct
ancestor of the former. The present species is the first significant evidence for incomplete hiati between well defined
cockroach genera – a result of the extensive fossil record of the group. The reported specimen is covered by a mycelium
of a parasitic fungus Cordyceps or Entomophthora.
Key words: Mexico, synanthropic, fossil insects, parasitic fungi, Allacta, Supella miocenica sp. nov.
overlain by the Balumtum Sandstone (Fig. 1). Both lithostratigraphic
bodies are regarded as informal units. The age of the
Chiapas amber has been a matter of debate. A Late Oligocene
to Early Miocene age has been proposed by Langenheim
(1966), Tomasini-Ortíz & Martínez-Hernández (1984), Santiago-Blay
& Poinar (1993), Bousfield & Poinar (1994), Poinar
& Brown (2002), Poinar (2003), Engel (2004), Castan~edaPosadas
& Cevallos-Ferriz (2007). Ferrusquía-Villafranca
(2006) described an artiodactyl from the Los Pocitos locality,
and considered a Late Oligocene age for these sediments,
based on previous biostratigraphic interpretations of Frost &
Langenheim (1974) and unreferenced paleomagnetic studies.
He suggested that the age of the Los Pocitos strata falls within
the 28—26 Ma (Ferrusquía-Villafranca 2006, p. 993). Other
authors suggest that the amber-bearing stratigraphic units are
of Middle Miocene age, and thus correlatives with the units
that produce amber in the Dominican Republic (MenesesRocha
2001; Solórzano-Kraemer 2007; Solórzano-Kraemer
& Mohrig 2007; Perrilliat et al. 2010). At the Los Pocitos locality
(Fig. 1), dark grey shales of the Mazantic Shale contain
amber, benthic foraminifera, gastropods, bivalves and
crustaceans. Based on 87
Sr/86
Sr measurements taken from a
well-preserved shell of Turbinella maya from Los Pocitos,
464 VRŠANSKÝ, CIFUENTES-RUIZ, VIDLIČKA, ČIAMPOR Jr. and VEGA
GEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICA, 2011, 62, 5, 463—475
Vega et al. (2009a, p. 53) obtained an absolute age of 23 Ma
for the Mazantic Shale, placing it right on the boundary between
the Oligocene and Miocene. The Mexican amber has
been interpreted as the resinous exudates of Hymenaea sp., a
leguminose tree whose communities developed near the ancient
coast, in estuarine environments, very similar to mangroves
(Poinar 1992). It is worth mentioning, that the closest
relative of amber producing H. mexicana is the relictuous H.
verrucosa from East Africa (Poinar & Brown 2002). Another
amber species, H. allendis is a relative to H. courbaril of
Americas (Calvillo-Canadell et al. 2009). The amber was consequently
transported to a shallow marine environment (Langenheim
1995; García-Villafuerte 2008).
The native representatives of the genus Supella are recently
limited to Africa and are considered to have diverged early
in the phylogeny of the family, with the genus being the
second basalmost offshoot (after Nahublattella) of the Blattellidae
(Klass 1997). Nevertheless, their position is obscure
in some other analyses (see Ware et al. 2008), and there is a
single plesiomorphy recorded in respect to most Symploce
Hebard, 1916 (M and CuA longitudinal), which is considered
to be the most primitive blattellid in our previous studies
(Vršanský 1997). Thus Supella is most likely derived
from this genus (Vršanský et al. 2011) or its predecessors.
Thanks to the synanthropic Supella longipalpa Fabricius,
1798 (circumtropic – introduced by commerce), the morphology
of the present genus is well studied at the microstructural
level (Roonwal & Rathore 1983). For example,
olfactory, gustatory, and mechanosensory receptors with a
density of 73.700 sensilla/mm2
were found on the most distal
segments of maxillary as well as labial palps (Prakash et
al. 1995).
The ecology of the genus is much less known. S. longipalpa
reproduces in temperatures between 25—33 °C (Tsai & Chi
2007). It can be a carrier of human intestinal parasites (Kinfu
& Erko 2008), Salmonella (Fathpour et al. 2003), perhaps a
cryptic species related to Gregarina blattarum von Siebold,
1839 (Clopton & Gold 1996) and an allergen source
(Tungtrongchitr et al. 2004). Its ootheca may often be parasitized
(over 19 %) by host-specific hymenopterans Comperia
merceti (Compere, 1938) (Encyrtidae) and Anastatus tenuipes
Bolivar & Pieltain, 1925 (Eupelmidae) (for details see
Narasimham 1992). Individuals infected by archiacanthocephalan
Moniliformis moniliformis (Bremser in Rudolphi
1819) spend more time in the shade (Moore & Gottelli 1992).
S. longipalpa nymphs may be more capable of producing
and utilizing extra metabolic water from food than Blattella
germanica (Linnaeus, 1767) nymphs (Melton 1995) and its
average motility and dispersion ability is also significantly
higher than those of B. germanica (Khrustalyova 1993).
The average life cycle takes 161 days.
The extremely close relation of the species described here
with the exclusively African S. mirabilis (Shelford, 1908) may
also be an indication of the possibility of living (at least morpho-)
species of cockroaches occurring in the Miocene. Living
cockroach species are still unknown in the fossil record, possibly
because of rapid phylogeny at their species level (Vršanský
2008). Living genera of cockroaches (both primitive and
advanced) are known since the Eocene (Vršanský et al. 2011).
What is notable is the identity of the pronotum colouration
with numerous representatives of another unrelated genus
that includes synanthropic species, the genus Periplaneta of
the Blattidae, for example, circumtropical P. australasiae
(Fabricius, 1775). Additionally, Periplaneta such as P. indica
Karny, 1908 from China can be smaller (forewing length
16 mm) (Karny 1908) and superficially could be easily confused
with Nemosupella Rehn, 1947. Periplaneta is known
from Eurasia, starting from the Middle Eocene – P. eocaenica
Meunier, 1921, P. relicta Meunier, 1921 and possibly others
occur in Messel sediments (Schmied 2009). Miocene representatives
are P. hylecoeta Zhang, 1989 and P. lacera
Zhang, 1989 from the Shanwang and P. sphodra Zhang, Sun
& Zhou, 1994 from Shandong in China. P. houlberti Piton,
1940 is reported from the Late Oligocene of Menat in
France. The living genus and even some species have a cosmopolitan
distribution. The origin of the synanthropic P.
americana is currently anticipated to have shifted to South,
Central and south of North America with the slave ships
from tropical Africa (Rehn 1945). Notably, both P. americana
and P. australasiae live both synanthropically and free
(Vidlička 2001). Supella can be easily discriminated from all
Periplaneta by the characteristic pale “band” and simplified
venation. The identical, but homoplastic habitus of cockroaches
significantly differing in size (ca. 10 mm in amber
specimen; up to 30 mm in living Nemosupella and up to
60 mm in living Periplaneta) is obscure.
Another similar and related genus Allacta Saussure &
Zehntner, 1895 is really problematic, because the only significant
difference of species placed in this genus is the geographical
distribution which is limited to Asia, shape and
colouration of the pronotum and shape of subgenital plate.
The vast majority of characters are overlapping between Supella
and Allacta and most likely these two genera represent
Fig. 1. Location and stratigraphic column of the Los Pocitos locality
of the Chiapas amber, Mexico.
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a single holophyletic taxon. Roth (1993) noticed the extreme
similarity of S. longipalpa and A. diluta. Nevertheless, although
we have studied all Allacta and Supella species, it is
not the aim of the present publication to revise both taxa. Because
both, and especially Supella is extremely polymorphic,
we retain the determination of the present species
within Supella and its subgenus Nemosupella, but it is evident
Supella is a direct ancestor of Allacta.
The taxon is evidence of the rich cosmopolitan Tertiary
entomofauna, which went extinct in the Americas, but still
survives on other continents, in this case in Africa. The African
taxa apparently also gave birth to the rich but uniform
genus Allacta, radiated in the whole of Asia, and comparatively
recently also in Australia. Allacta itself was also
present in America during the Eocene.
It follows that the horse was not the only group which
went extinct in the Americas, but could easily be reintroduced
there by humans.
Material and methods
A single completely preserved specimen was collected by
Luis Zú±iga Miganjos in the Los Pocitos locality and is deposited
in Museo Comuntario del Ambar, Simojovel, Chiapas
(MUCAS, supported by Instituto Nacional de Antropología e
Historia – INAH). It was photographed using a Canon EOS
Mark II and drawn with a camera lucida from an Olympus
SZH10. Wing terminology follows Vršanský (1997). The living
undescribed Allacta or Supella sp. provided in Fig. 3 was
collected in Central Laos, Bolikhamsai province, Ban NapeKaew
Nua Pass [N 18°22.3’/ E 105°09.1’], by Ondrej Šauša
and Eduard Jendek (deposited in IZ SAS Bratislava).
Parsimony analysis was performed using PAUP* software
version 4.0b10 (Swofford 2002), with a TBR heuristic search
of 1,000,000 replicates and the option ‘save multiple trees’ activated.
All characters were treated as unordered (0 – plesiomorphic,
1, 2, 3 – apomorphic states). MaxTrees option was
set to 1000. Characters were weighted regarding their evolutionary
relevance (characters 9, 12 – weight = 10, character
13 – weigh = 1, remaining characters – weight = 5). Details
of the procedure of each character are explicitely stated
in the character analysis below. A heuristic search produced
1000 equally parsimonious trees with length 368 (consistency
index CI= 0.29, retention index RI=0.77). Majority consensus
tree was constructed and post-edited (coloured) in
MESQUITE software version 2.6. Only clades with frequency
>50 % were retained.
Results
Systematic entomology
Order: Blattida Latreille, 1810 (= Blattaria Latreille,
1810=Blattodea Brunner von Wattenwyl, 1882)
Family: Blattellidae Karny, 1908
Subfamily: Pseudophyllodromiinae Hebard, 1929
Supella Shelford, 1911
Fig. 2. Supella miocenica sp. nov. from the Miocene Chiapas amber of Mexico. Holotype MUCAS-001. a – Dorsal view; b – Ventral
view; c – Head; d – Fore leg; e – Fore tarsus. Original by PC and FV. Total specimen length (from the head to the end of wings) 10 mm.
466 VRŠANSKÝ, CIFUENTES-RUIZ, VIDLIČKA, ČIAMPOR Jr. and VEGA
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Type: Blatta longipalpa Fabricius, 1798 (circumtropical,
native in Africa). 
= Blatta supellectilium (Serville, 1839)
= Blatta incisa (Walker, 1868)
= Ischnoptera quadriplaga (Walker, 1868)
= Blatta extenuata (Walker, 1868)
and numerous other synonyms of diverse specimens from the West
Indies (see Rehn 1947).
Composition: Besides the type; S. vicina Chopard,
1958 [Comores islands]; S. abbotti Rehn, 1947 [Kenya,
Tanzania, Malawi]; S. dimidiata Gerstaecker, 1869 [Kenya,
Congo, Angola, Rhodesia, Malawi, Mosambique, Botswana,
Natal, Transvaal]; S. orientalis Grandcolas, 1994 [Saudi
Arabia]; S. (Mombutia) chapini Rehn, 1947 [Congo]; S.
(Nemosupella) gemma Rehn, 1947 [Ghana]; S. (Nemosupella)
mirabilis [Cameroon, Gabon, Congo, Uganda, Kenya,
Tanzania]; S. (Nemosupella) occidentalis Princis, 1963
[Guinea]; S. (Nemosupella) tchadiana Roth, 1987 [Chad].
All extant in Africa.
Diagnosis (Rehn 1947, in part): “Pronotum ovate subtrapezoideal
in outline. Tegmina of female varying in length
from covering but half the abdomen to surpassing the abdominal
apex by a distance equal to the pronotal length, in outline
ranging from 1.5 times as long as broad. Apex well-rounded.
Costal veins numerous, straight oblique, several of the more
distal ones usually ramose, reduced in number in the abbreviated
tegmined forms; discoidal sectors oblique, tending toward
sublongitudinal in males of S. abbotti and S. mirabilis.
Anal field pyriform, anal veins five or more, regular; discoidal
sectors similarly developed, anal field always fully indicated.
Cephalic femora with ventro-cephalic margin bearing a
regular series of spines, evenly reducing in length and
strength distad (sometimes replaced by setae (e.g. in one
limb) – e.g. in S. mirabilis), apical spines of same margin
two-three in number, the terminal one much the longer; median
and caudal femora with ventral margins spined; caudal
tarsi with metatarsus in length surpassing the other articles
combined. Arolia well developed; tarsal claw of equal
length, their margins unarmed, simple.”
Subgenus: Nemosupella Rehn, 1947
Type: Phyllodromia mirabilis Shelford, 1908.
Composition: Supella mirabilis, S. gemma, S. tchadiana,
S. occidentalis.
Diagnosis (ex Rehn 1947 in part – only relevant
characters): “Females more robust with tegmina and wings
broader and in length less markedly surpassing the apex of
the abdomen, apex well rounded. Head pyriform in outline,
transverse facial ridge nearly straight transverse; palpi with
penultimate and antepenultimate articles elongate. Caudal
tarsi moderately slender, metatarsus in length somewhat exceeding
that of the remaining tarsal articles combined”.
Supella (Nemosupella) miocenica sp. nov.
(Figs. 2a—c, 3a—c)
Holotype: MUCAS-001. A complete female.
Type locality: Los Pocitos, Simojovel de Allende,
Chiapas amber.
Type horizon: Lower Miocene, Mazantic Shale, Tertiary.
Differential diagnosis: The present species differs
from its consubgeners, S. mirabilis in being smaller (total
body length with wings ca. 10 mm vs. 16—25.5 mm in S.
mirabilis), in having discoidal sectors oblique (oblique to
sublongitudinal in S. mirabilis), and in having the central dark
pronotal macula divided into two parts; from S. gemma in size
(similar as S. mirabilis), in having wings more coloured and
pronotal central macula smaller; from S. tchadiana of a comparable
size (12 mm), in having pronotum without markings;
and from S. occidentalis in colouration and size.
Description: Very small species (overall body length
without wings about 9 mm). Head small (length to width:
1.6/1.3 mm) with very fine antenna covered by a row (basal
segments) or up to four rows with four short (roughly corresponding
to segments’ width) sensilla chaetica in each.
Pronotum ovate subtrapezoidal in outline, significantly
vaulted (1.7/2.9 mm), pale, with dark macula covering the
whole posterior margin and central macula, divided into two
separate parts.
Body slender, sterna (especially the posteriormost ones)
widely carved, cerci with up to 16 segments, very long
(1.8 mm) with dense fine sensilla chaetica of diverse length
(0.2—2 times as long as the width of the median cercal segment).
Legs slender, long (including fore legs), cursorial. Fore
legs terminated with claw and arolium; femora (1.7/0.1 mm)
with dense chaetica, tibia (0.9/0.07 mm) with at least 5 fine
spurs (arrangement of spines along the tibiae in 3 rows), tarsi
5-segmented (0.7, 0.2, 0.1, 0.05, 0.2 mm). Front femur Type
B2, with four proximal stout spines succeeded by a row of
uniform piliform spinules, terminating in two large spines;
pulvilli present only on the fourth tarsomere, tarsal claws symmetrical
and unspecialized (simple), simple arolia present.
Mid femora wide and with numerous sensilla (about 26
spines in two rows) (2.1/0.6 mm), tibia also robust (1.6/
0.2 mm), with long fine spurs (10 or more); tarsi curious, extremely
short (0.7/0.1 mm), with an indistinct claw and arolium.
Hind legs long, femora robust (2.6/0.8 mm), with two
rows with numerous (about 19 2) fine spurs and terminal two
fine spurs; tibia long (2.9/0.2 mm), with numerous (up to 30)
fine spines; tarsi long (1.3, 0.3, 0.1, 0.1, 0.3 mm), densely
haired, with distinct claw and arolium. First third of the first
Fig. 3. a—d – Supella miocenica sp. nov. from the Miocene Chiapas amber of Mexico. Holotype MUCAS-001. Ventral view. a – Complete
specimen with curculionid beetle; b – detail on head with mycelia of parasitic fungus Cordyceps or Entomophthora (white “bubbles”);
c – detail on cercus; d – dorsal view, total specimen length (from the head to the end of wings) 10 mm. e – Undescribed male of
Allacta or Supella sp. from Central Laos. Total length ca. 12 mm. Nemosupella is even more similar in having the nearly identical pronotal
shape and colouration (the central macula is divided into 2 parts only in S. miocenica). Photograph of the sister species, S. (Nemosupella)
mirabilis is available free on the web (FOW).
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Fig. 3.
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tarsal segment sparsely (5) haired, the rest with at least 33 sensilla;
second segment with 9, third with two, fourth with six
and terminal one with at least two sensilla.
Forewing dark, with characteristic pale anterior margins
and central stripe making the illusion of the body being separated
into two halves. Apex rounded. Venation simple, with
minimum deformities. Total number of veins meeting the
margin (without A) ca. 30. Sc simple, short, R nearly
straight, R branches simple except for the few terminal
branches (18 R veins meet margin). RS not clearly differentiated.
Discoidal sectors oblique. M secondarily branched,
with about 10 branches at the margin. CuA is fused with M
and perhaps consists of a single terminally dichotomized
branch. Anal field pyriform, fully indicated, anal veins simple,
regular, apparently five or more. Intercalaries locally
distinct (in basal R and CuA) and probably common.
Remarks: For comparison and details see Discussion.
Derivation of name: miocenica is after the Miocene epoch.
Character analysis: (0 – plesiomorphy; 1—3 – apomorphies
relative to other species within genera Supella and
Allacta, and/or Cariblattoides Rehn & Hebard, 1927 and
Symploce (and fossil Piniblattella Vršanský, 1997), which
were chosen as outgroups based on high similarity, and
based on retension of all original states of characters due to
standard habitus (not derived like in Supella and Allacta):
1. Head with interocular (IO) space roughly identical with
the distance between antennal sockets: apomorphy; plesiomorphic
is partially reduced IO space (as in Symploce and
most regular cockroaches); IO space nearly absent is a strong
apomorphy (2), eyes very small and remote (3).
2. Pronotum large ovate subtrapezoidal in outline, significantly
vaulted: plesiomorphic (as in most cockroaches including
Symploce and Piniblattella); apomorphic states are
1) small and round; 2) subelliptical with margins parallel.
3. Pronotal colouration with basal dark stripe and two central
maculas: plesiomorphic (central maculas are present in
most primitive blattellids such as Piniblattella), derived states
are alternative colourations (dark with cental pale macula).
4. Subgenital plate long: apomorphy (plate is plesiomorphically
of normal length in Symploce, Piniblattella and
most other blattellids).
5. Forewing with apex of radial area reduced to mostly
simple branches (RS indistinct): apomorphy (RS distinct in
primitive Symploce and Piniblattella).
6. Forewing M and CuA branched and curved: plesiomorphy
(as in Mesozoic cockroaches); these branches are apomorphically
longitudinal even in some Symploce; serrate (2).
7. Forewing colouration with characteristic transversal
stripes: apomorphy at level of common ancestor of Supella
and Allacta. Plesiomorphic state is colouration uniform but
not strong as in Cariblattoides, and other primitive blattellids
(Nahublattella, Neoblattella etc.).
8. Colouration of wings soft: plesiomorphy; pronotum
strongly dark, with sophistic pale stripes on forewing is apomorphic
(funebris spp. group).
9. Colouration of wings and pronotum with continuous colouration:
plesiomorphy; derived apomorphic state is colouration
in dark dots, lines and blotches. This character was
found as a global irreversive reorganization of morphology
and colouration and thus has been given higher weight in
the cladistic analysis.
10. Hindwing R1 distinct: plesiomorhy (as in most primitive
blattellids including Symploce and Piniblattella); R1 is
apomorphically reduced to a single vein.
11. Fore tarsi of B-type: apomorphy (tarsi are plesiomorphically
A-type in most primitive blattelids including most
Symploce).
12. Pulvilli exclusively on 4th
tarsomere: apomorphy (pulvilli
are plesiomorphically on 4 tarsomeres in primitive blattellids
including Symploce). In addition to diagnosis of Allacta (Saussure
& Zehntner, 1895), this character was found unique, never
occurring homoplastically in any other group (additionally unrelated
with respect to size changes) and thus has been given
higher weight in the cladistic analysis.
13. 1 or 2 terminal fore femoral spurs: apomorphy (plesiomorphic
state is with 3 spines in both A- and B-types of
Symploce). This character was found polymorphic within
species and even on one specimen (L/R sides – Roth 1991,
1993, 1996 and our observation) and thus has been given
lower weight in the cladistic analysis.
14. Habitus robust: plesiomorphy (as in Piniblattella), derived
apomorphic states are slender (1), extremely fragile (2)
and extremely elongated (3).
Allacta Saussure & Zehntner, 1895
= Abrodiaeta Brunner von Wattenwyl, 1893
= Pseudochorisoblatta Bruijning, 1948
= Arublatta Bruijning, 1947
Type: Abrodiaeta modesta Brunner de Wattenwyl, 1893
from Carin Ghecu in Burma, by selection.
Composition: Funebris species group (sensu Roth
1993): basivittata (Bruijning, 1947) [New Guinea, Aroe and
Aru Islands], bipunctata (Walker, 1869) [Celebes, Aru Islands,
New Guinea], funebris (Walker, 1868) [Borneo] (Roth
1993); grandcolasi Roth, 1995 [Irian-Jaya], megamaculata
Roth, 1995 [Papua New Guinea], straatmani Roth, 1995
[Papua New Guinea] (Roth 1995), diagrammatica (Hanitsch,
1923) [Malacca, Singapore, Mentawai islands, Sumatra, Java].
Hamifera species group (sensu Roth 1993): bimaculata
Bey-Bienko, 1969 [China], diluta (Saussure, 1863) [Ceylon,
India], figurata (Walker, 1871) [Ceylon, India], hamifera
(Walker, 1868) [Malacca, Java, Borneo, Philippinen], interrupta
(Hanitsch, 1925) [Borneo], luteomarginata (Hanitsch,
1923) [Singapore], maculicollis (Hanitsch, 1927) [Vietnam],
parva Shelford, 1906 [Borneo], pantherina (Hanitsch, 1933)
[Borneo] (Roth 1993); svensonorum Roth, 1995 [Malaysia,
Borneo] (Roth 1995).
Polygrapha species group (sensu Roth 1993): fascia Roth,
1993 [Indonesia], immunda (Brunner von Wattenwyl, 1893)
[Burma, Malacca], polygrapha (Walker, 1868) [Thailand,
Malacca, Singapore, Sumatra, Borneo], picturata (Shelford,
1907) [Singapore, Sumatra, Thailand, Malaysia, Borneo],
marmorata Walker, 1869 [Burma, Sumatra, Malaysia],
mcgavini Roth, 1991b [Indonesia], robusta Bey-Bienko, 1969
[China], transversa Bey-Bienko, 1969 [Vietnam]; arborifera
(Walker, 1868) [Malaysia, Java, Borneo, Mentawai Islands],
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Cariblattoides with A-type and pulvilli on 4 tarsomeres, supporting
this state as ancestral.
Phylogenetically obscure is also the distribution of knee
spines at the distal end of the femora – 2 in present species
and some Allacta (see character matrix) and Supella, but more
often 3 (also in figured most closely related Allacta sp.).
The present amber species clearly differs from representatives
of the A. funebris species group, which are very dark
including the pronotum (eventually with a narrow pale margin
only) and with basal narrow pale stripes. A. polygrapha
spp. group differs in having a large symmetrical pronotal
pattern of dark dots, lines and blotches and forewings chequered
with dark cells between veinlets and with larger
blotches (A. confluens placed here is somewhat different
from other representatives of this species group, it has more
coherent colouration and pronotum colouration most similar
to the amber specimen. Nevertheless, the shape of the pronotum
of S. miocenica is different: dissimilar to any described
Allacta spp.). A. puncticollis is also completely different,
with a subparabolic reddish brown pronotum with small yellowish
spots, but has pulvilli limited to the 4th
tarsomere. A.
crassivenosa is categorized as incertae sedis and most likely
belongs to another genus. The most related to the present
amber species within Allacta is the A. hamifera spp. group,
which differs in having smaller pronota with a different colouration
pattern and in the shape of the subgenital plate which is
never as long in Allacta. Even more similar, and hardly recognizable
from the Nemosupella spp. is an undescribed representative
putatively attributed to Allacta on the basis of identical
legs (pulvilli limited to the 4th
tarsomere – Figure 2b). Alternatively
it can mean that this species belongs to Supella and
is its only Asian representative, but more likely it is a transitional
taxon leading to Allacta. The figured undescribed species
(Fig. 3e) cannot be placed into any spp. groups of
Allacta, but is most closely related to the A. hamifera spp.
group in the shape of the head and underived colouration. It
is apparent that A. funebris and A. polygrapha spp. groups
were derived much later, the latter apparently derived via A.
confluens, which has similar pronotum colouration and underived
forewing colouration.
It is clear, that there is no strict hiatus between these two
genera, but this cannot be used as a reason for their synonymization
or for the erection of additional genera. The problem
is that there are known “missing link” taxa in all cockroach
families and we also know a half-cockroach—half termite
(Vršanský 2010) and also half-cockroach—half mantodean. All
studied living cockroach genera with fossil records have these
transitional stages too, and the present taxon is no exception.
This is a half-Supella—half-Allacta, but better Supella than
Allacta. So splitting or erecting does not have a proper place
here just because the group is well studied. Paraphyletic taxa
and incomplete hiati are present in the vast majority of studied
cockroaches, which is a result of the extensive fossil record
with ca. 100,000 specimens. So our specimen has synapomorphies
of Allacta+Supella (colouration, venation), autapomorphies
of Allacta (extremities), but major autapomorphies of
Supella (pronotum, subgenital plate).
The cladogram (Fig. 4) supports all the above-mentioned inferences,
with nearly ideal separation of all spp. groups, but it
australiensis Roth, 1991 [Queensland], confluens (Hanitsch,
1925) [Borneo], labyrinthica (Hanitsch, 1927) [Vietnam],
loconti Roth, 1993 [Indonesia], megaspila (Walker, 1868)
[Malacca, Mentawai Islands, Java, Borneo], ornata Bey-Bienko,
1969 [China], modesta (Brunner von Wattenwyl, 1893)
[Burma; type], karnyi (Hanitsch, 1928) [Mentawai Islands,
Sumatra] (Roth 1993), brossuti Roth, 1995 [Irian-Jaya],
deleportei Roth, 1995 [Papua New Guinea], gautieri Roth,
1995 [Papua New Guinea], nalepae Roth, 1995 [Papua New
Guinea], persoonsi Roth, 1995 [Papua New Guinea], srengi
Roth, 1995 [Papua New Guinea] (Roth 1995).
A. puncticollis (Brunner von Wattenwyl, 1898) [Borneo]
(not placed sensu Roth 1993) and A. crassivenosa Bolívar,
1897 [India] (incertae sedis sensu Roth 1993). Except for
problematic A. australiensis, all are extant in Asia.
Diagnosis: Front femur Type B [B2 or B3 according to
Roth 1993, or C (right and left femur can differ in type (B2-B3
or even B-C) according to Roth (1991, 1996))]; pulvilli
present on fourth tarsomere only, tarsal claws simple, symmetrical,
arolia present (Roth 1995).
Discussion
Supella-Allacta complex
Because there is an immense similarity and relation between
the genera Supella and Allacta, it is necessary to provide
arguments for the categorization of the present fossil
within Supella.
These genera are clearly distinguished among other blattellids
by autapomorphies including the characteristic colouration
with a pale stripe appearing to divide the body into
two (or, in combination with the pronotal colouration into 3)
separate parts; M and CuA descending in an obtuse angle;
mostly simple R branches with indistinct RS (homoplastic
with Pseudomops Serville, 1831 and Ectobius Stephens,
1835); and in other characters unseen in the present fossil (see
Rehn 1947). Supella restricted to Africa is much more diverse
and polymorphic (including forms identical with Allacta)
which suggests its direct ancestral position in respect to
Asian and Australian Allacta (the latter is restricted to
a single species A. australiensis from Queensland, which has
an indicated hindwing R1 as in Supella, but is very different
from both Supella and Allacta in having eyes nearly connected,
and in subelliptical – with anterior and posterior
margins parallel – form of pronotum; thus it can simply
represent Supella or a different genus).
The present amber species has fore legs of B type identical
(!including the number of proximal spines and terminal
spurs) with Allacta, dissimilar to most Supella (A-type), but
these types can be polymorphic (L/R) in a single specimen
of Allacta (B2/B3 or even B/C – see above), and the Supella
subgenus Nemosupella can have this B-type pattern too. The
most primitive living blattellid, Symploce has mostly A3
type, but ocassionally B3, thus it is likely A is the original
type, but the above-mentioned polymorphisms are evidence
for a convergent nature of this character changes. Closely related
and perhaps derived from Supella is also the genus
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was weak in the position of A. interrupta (well nested within
hamifera group by intuition (Roth 1993), very closely related
to A. hamifera) within the polygrapha group. The second problem
is the terminal position of A. confluens which is expected to
be the basalmost polygrapha group stem (it has a very basal
position when all characters were equally weighted). Third, A.
bimaculata+A. pantherina (hamifera spp. group) appear nested
within Supella. Notably, all three problems are absent in the
cladogram with the normal weight of character 13 (weight 1 in
contrast to weight 5 of all other characters except weight 10 of
characters 9 and 12), which is polymorphic (and thus has been
given a lower weight), A. bimaculata+A. pantherina (hamifera
spp. group) are nested (as basalmost offshots) within hamifera
spp. group using the same weights but using different (500)
number of maximal search trees. A. puncticollis (out of any spp.
group) appears to be a sister taxon to A. svensonorum and A. luteomarginata
(trichotomy at cladogram). When all characters
were weighted normally and in many other options tried, S. dimidiata
were often nested within Allacta. S. dimidiata is a good
Supella, and this placement is evidence for numerous homoplasies
within the group. Generally, the homoplasies wihin cockroaches
are enormous.
Paleogeographically, the basalmost blattellids were preserved
in Asia, but their ancestors within the Mesoblattinidae
were also common in Europe (Vršanský & Ansorge
2007). The basalmost Supella is up to recently clearly nested
within Africa, but the present American species is clear evidence
for the past circumtropic distribution. Allacta was possibly
derived in Africa but radiation was apparently not
limited to Asia (except for the mentioned synanthrope, Allacta
is present in sediments of the Green River in Colorado).
According to this cladogram A. australiensis was
derived quite recently from one of the species in the
polygrapha spp. group.
From the most primitive blattellids, Nahublattella and
Symploce (for position of these genera within Blattellidae
see also Klass & Meyer (2006)), there is a significant reduction
and simplification of venation, which is evidence of the
very early divergence of the whole complex from the main
blattellid stem. Another eventual conclusion considers the
small size of the present species, which may be a plesiomorphic
character, which would explain the significant simplification
of venation even in large living species. Notably, in
the derived genus Allacta, terminal radial veins are simplified
only in the smallest species (A. parva).
Notwithstanding, some living Supella and Nemosupella in
particular have more primitive traits and likely diverged before
the speciation of S. miocenica. Its single insignificant
deformity (not clearly visible, insignificantly changing the
wing geometry) supports the fact that Eocene and Miocene
species have few accumulated wing deformities and their occurrence
in living species represents support for them being
inheritable mutations (see Vršanský 2005).
Extinct American Supella
In spite of the close relation of its 3 known subgenera, the
general habitus of the respective taxa in Supella is very diverse.
S. longipalpa is a slender, fine cockroach, while S.
orientalis has extremely elongated wings (—1:4.5), and the
subgenus Nemosupella is clearly differentiated by robust
habitus with a robust unplain pronotum and more or less normal
longitudinal veins. The present species share all the autapomorhies
of the genus and subgenus Nemosupella and
can be safely categorized within this taxon.
The colouration and general appearance is hardly recognizable
from females of S. mirabilis, its sister species, although
the shape of the pronotum is somewhat transitional
between its males and females. Except for the significantly
smaller size of the present new species, the sole difference
between the two taxa is the divided central pronotal macula.
There are no additional plesiomorphies, which indicate the
present as well as the living species of Nemosupella diverged
near the Mid-Miocene. The other three representatives of the
subgenus are closely related to S. mirabilis, but are dissimilar
to the present fossil due to different colouration.
On the other hand, other representatives of the genus such
as S. longipalpa reveal significant divergence from the main
morphological standard and suggest rapid phylogeny at the
subgeneric level.
The similarity with the undescribed representatives of the
genus Allacta (Fig. 3e) is so striking (and involves size – Nemosupella
are much larger), that it is apparent Allacta is derived
from Nemosupella via the predecessors of this
undescribed taxon and also via predecessors of Supella miocenica.
Its direct ancestry can be excluded based on the derived
pronotum of S. miocenica. While there is a standard
Allacta placed within a living spp. group (polygrapha)
present in much older Eocene Green River sediments, the
Chiapas is apparently another case of the presence of primitive
species in amber, when compared to isochronous sedimentary
record. Relic character of amber cockroach (and all insect to
some extent) assemblages is characteristic also for the only
two studied Mesozoic ambers (Lebanon and Archingeay, but
also in Baltic amber), which might either be caused by different
methods of dating, or by the more humid, dark and
colder source microclimates of amber forests. This is in contrast
to Cretaceous ecosystems, where primitive cockroach
forms of the Jurassic type are restricted to younger, but dry
to semiarid ecosystems (Vršanský et al. 2002).
Very little can be said about the ecology of the present species.
Generally the ecosystems of Chiapas were perhaps diverse,
ranging from lowland tropical dry forest tending toward
open forest and mangroves (Solórzano-Kraemer 2007). The
wide range of ecosystems from the rainforest down to savannas,
of the closely related S. mirabilis, indicate this genus is
highly adaptable to diverse conditions. This ecological plasticity
could have resulted in invasions of early Supella (Nemosupella)
into the Americas before the Mid-Miocene (and later
into Asia as the genus Allacta). Living species of Supella are
cavicolous (Grandcolas 1994) and the genus most likely also
originated in Africa because the most primitive blattellid, Symploce
Hebard, 1916 is circumtropic, but rare in America. On the
other hand, the genus Nahublattella Bruijning, 1959 considered
to be even more primitive by Klass (1997) is native to Central
and South America, which could indicate the opposite.
The parasitic (or predatory) fungus Entomophthora or
Cordyceps is indeterminable, but the mycelium is richest in
471AFRO-ASIAN COCKROACH AND LOST TERTIARY AMERICAN ENTOMOFAUNA
GEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICAGEOLOGICA CARPATHICA, 2011, 62, 5, 463—475
Fig. 4. Parsimony analysis of all the known Supella and Allacta species (excluding obscure A. crassivenosa, S. occidentalis, S. vicina, and
including 3 undescribed Allacta and/or Supella species) with their geographic distribution. S. miocenica and Piniblattella vitimica are extinct.
Position of A. interrupta (top) is illusory due to numerous homoplasies with A. robusta and other polygrapha spp. group species, as
this species apparently belongs to the hamifera spp. group. 50% majority consensus tree from 1000 equally parsimonious trees was gained
with maximum parsimony search (PAUP). Numbers above branches represent clade frequencies in %.
the junction of head and pronotum and could have its epicentre
in the head. This fungus provides a contribution to the
poorly known microorganisms of the Chiapas amber. Only a
?Bacillus-like cell and two types of budding-bacteria-like
microrganisms were reported previously (Veiga-Crespo et
al. 2007).
Comments on synanthropism in cockroaches
The fifty species of synanthropic cockroaches comprise
only an insignificant fraction of the total of about 5000 (Bell
et al. 2007) described species of living cockroaches. Nevertheless,
they are important for their number and ecological significance.
Most of the species had their genera recently limited to
certain continents and only nowadays have become cosmopolitan
(or circumtropical). On the contrary, their history on a
geological scale is much richer than we would expect and
their original distribution was also circumtropical.
The genus Blattella was until very recently limited to Africa
(26 species), Asia and the Pacific islands (23 species) (Roth
1985), and the synanthropic species B. germanica spread to
the whole world from east Asia (Roth 1985). The occurrence
of this genus in the Mesozoic of Europe as a single nymph
(adult could eventually differ) (Vršanský 2008) and in the
Eocene of the USA (Green River, Colorado) is thus surprising.
Free Ectobius is limited to Europe, but in the Tertiary it was
cosmopolitan. The same situation is found with the present
Supella introduced from North Africa to Central America on
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slave ships (Rehn 1947). The Miocene of Mexico is quite distant
from the recent distribution limited to Africa.
According to Princis (1954), Blatta originates from the
Near and Middle East – the native place of the closely related
Shelfordella Adelung, 1910, but the other predicted origin
of this genus is North Africa (Rehn 1945; Cornwell 1968),
and Blatta furcata Bohn, 1985 is known from the Near East
and North Africa (Bohn 1985). Their close relative also occurred
in the Eocene of the USA (Green River).
The last significant synanthrop is Periplaneta, recently
limited to Asia.
The only viviparous cockroach group met the same fate:
the Diplopteridae, now restricted to two genera (one African,
one Asiatic), were common in America during the Eocene
(in Green River). This family is also reported from the
Eocene of Quilchena (Archibald & Methewes 2000), but the
figured specimen (Q-0040) is very different from all known
Diplopteridae (details to be provided elsewhere).
Thus it is apparent that all the synanthropic species belong
to cosmopolitan genera (cosmopolitan genera are otherwise
rare), very likely with a broad environmental tolerance – and
thus pre-adapted for synanthropism.
Lost Tertiary American entomofauna
The occurrence of some cosmopolitan synanthropic species
in the Americas and their absence prior to re-introduction,
triggered the present discussions about the causes of the
extinction of these entomofaunas during the Tertiary.
It was Eocene Ectobius from the Green River – a member
of an extinct genus, but which was extremely easily reintroduced
in North America several times (with 3 species),
which concentrated our efforts on the search for this fauna in
2006. It was very rapidly supported by the discovery of a
honey bee in the Miocene sediments of Nevada (Engel et al.
2009), a genus extinct in the Americas.
Poinar et al. (1999) also noticed the Early Tertiary North
American extinctions of species of living tropical ant genera
Technomyrmex Mayr, 1872, Leptothorax Mayr, 1855 and
Dolichoderus Lund, 1831, recorded in the Eocene of British
Columbia. These records comprise only the species level,
which is insignificant on the present time scale, but Technomyrmex
is now, with the exception of a single Central American
species (and its abundance in the Dominican amber),
limited to the tropics of the old world; Leptothorax is holarctic
today; Dolichoderus is cosmopolitan.
The Eocene of the Okanagan Highlands reveals a representative
of the Myrmeciinae, currently limited to the Australian
region (Archibald et al. 2006). The only genus – determined
hemipteran from Quilchena, Megymenum is today found in
only in the Oriental biotic region and Australia (G. Gross,
personal communication in Archibald & Mathewes 2000).
The post-Miocene cooling was unlikely to be a reason for
this extinction, as both Ectobius and Apis Linnaeus, 1758
occur in Northern Europe today.
The loss of another taxon from North America is now apparent
– Supella, in which case cooling could be the reason
as nowadays this genus is restricted to Africa. On the other
hand it is hardly possible that this taxon went extinct in
warm Central and South America. Their historical absence in
South America is also difficult to anticipate as other cockroach
taxa from the Dominican amber are present in South
America (see below).
Thus, of fourteen studied cockroach genera (all still living)
from the Eocene—Miocene of North America only two (Cariblattoides,
Sigmella) survive nowadays in (South and Central)
America and only Cariblattoides is characteristic for Central
America (although it occurs in Brazil). It is perhaps not incidental
that a representative of Sigmella was dominant during
the Eocene and also in the present Mexican amber. An additional
taxon reported from the Mexican amber is Ischnoptera
sp., currently distributed in Central and South America, but
this determination is obscure (determination may be correct,
but no diagnostic characteristics for the genus are provided,
and the species (Ischnoptera sp. 1 in Solórzano-Kraemer
2007) may well belong to Supella or some other blattellid
taxon). All things being equal, the diversity of cockroaches in
Chiapas amber was certainly high: 7 specimens belong to
7 different genera and species of the family Blattellidae.
Another cockroach genus, exclusively African today,
known from the Tertiary of North America is Namablatta
Rehn, 1937.
The closely related termites are ubiquitous in the present
context. While Kalotermes nigritus still lives in South America,
the whole cosmopolitan family Mastotermitidae (present
as Mastotermes electromexicus Krishna & Emerson, 1983
and Mastotermes electrodominicus Krishna & Grimaldi,
1991 occurring in the Dominican amber according to Solórzano-Kraemer
(2007)) went extinct in the Americas and survives
only in Australia.
Some other insect groups from the Mexican amber (Solórzano-Kraemer
2007) and Green River (our data) reveal a
similar pattern, to be analysed in detail elsewhere.
It is of special consideration that the Dominican amber
shows a very different pattern in respect to the distribution of
cockroach genera. If the determinations of Arillo & Ortun~o
(2005) are correct, then there are no shared taxa (even on the
generic level) between the Mexican and the Dominican amber,
and all Dominican amber cockroach genera are not only
highly advanced, but with the exception of the circumtropical
Anaplecta Burmeister, 1838 all – Euthlastoblatta Hebard,
1917, Pseudosymploce Rehn & Hebard, 1927, Plectoptera
Saussure, 1864, Cariblatta Hebard, 1916, Holocompsa Burmeister,
1838 (a single species (H. debilis (Walker, 1868))
also occurs in Ceylon, Java, Sumatra, Borneo and Phillipines)
– are characteristic of Central and/or South America.
Taxa described by Gorochov (2007), including obscure
Agrabtoblatta Gorochov, 2007 and Erucoblatta Gorochov &
Anisyutkin, 2007, also appear limited to South America.
Taking all this preliminary information together, it is apparent
that sometime after the Mid-Miocene some extensive
environmental change influenced North and probably also
Central and South America, resulting in the loss of cosmopolitan
Early Tertiary entomofaunas. Judging from the modern
composition of the Dominican amber, this may (if the
abovementioned determinations are correct) mean a recovery
occurred during the time between the Early Miocene Mexican
amber (23—7.1 Ma) and the Dominican amber times
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(20.5—16.4 Ma). The dating of both of ambers is still uncertain
(the abovementioned datings are after EDNA database),
more counterbalanced by the Late Barstovian (14.5—14 Ma)
dating of the Nevada (with honeybee) sediments. Recently,
the age of 23 Ma was designated for the basalmost amber
bearing strata of Chiapas (Vega et al. 2009a).
As it is very difficult to imagine some geological or ecological
process which would be able to trigger such an extensive
change (aridization and/or cooling would not influence
some of the cockroaches), the change was probably biological
– either caused by diversification of cockroach parasites
which were consequently reduced (as reintroduction has
been easy), or diversification of more progressive insect
groups. The parasite hypothesis may be valid in the present
case, as Comperia merceti eradicated populations of synanthropic
cockroaches in Europe, even when its function as a
control of Supella is still not validated (Goudey-Perriere
1991) and Encyrtidae and Eupelmidae parasitizing ootheca
have Tertiary origin (A.P. Rasnitsyn, personal communication
2010), known only from Europe starting with the
Eocene Baltic amber (Trjapitzin 1963). (These parasites
could also cause extinctions of external ovipositor bearing
cockroaches which did not lay eggs in ootheca.)
The occurrence of advanced taxa in the Dominican amber
(isochronous with the Mexican amber according to Solórzano-Kraemer
(2007), however see above) would favour the
diversification and radiation of the modern South-American
cockroach taxa hypothesis. Nevertheless, Diploptera is the
most advanced cockroach that has ever lived, and thus its extinction
in the Americas falsifies the latter hypothesis. On
the other hand, it is possible that its viviparity evolved only
in the common ancestor of Asian and African species.
Plants were perhaps not as influenced as fauna, as Eocene
flora of British Columbia in Canada is characteristic of the
modern eastern North American deciduous forest zone, principally
the mixed mesophytic forest, but also including extinct
taxa: taxa known only from eastern Asian mesothermal
forests, and a small number of taxa restricted to the presentday
North American west coast coniferous biome (Greenwood
et al. 2005). Also, according to Solórzano-Kraemer
(2007), all plants from the Chiapas amber are currently
present in Pacific coastal forest.
Conclusions
The genus Supella with S. miocenica sp. nov. was native to
America during the Miocene time of the Chiapas amber. It
represents another case of rich cosmopolitan Early Tertiary
entomofauna, which suddenly went extinct in America somewhere
around the Miocene (but which still survives in other
continents). Supella/Allacta complex (Allacta was derived
from Supella) is another case of the genera which now includes
synanthropic species, which were natively circumtropic,
and can be easily reintroduced in America nowadays. S.
(Nemosupella) miocenica sp. nov. is the earliest known cockroach
which can be categorized within the living subgenus
and also the first published direct evidence of transitional species
(and thus incomplete hiatus) at the level of living genera.
Acknowledgments: We thank the Museo Comunitario del
Ámbar de Simojovel, Chiapas for making this material available
for study; the generous support of Luis Zún~iga and Jorge
Balcázar is greatly appreciated; Sonia Fraga Lopes (National
Musem, Rio de Janeiro), Dong Ren (Capital Normal University,
Beijing), Maria del Carmen Perrilliat (Institute of Geology,
Mexico), Jozef Michalík (Geological Institute, Slovak Republic)
and five anonymous reviewers for revision of the manuscript,
Ladislav Roller (Institute of Zoology SAS, Bratislava)
for literature supply, Ivona Kautmanová (Slovak National
Museum, Bratislava) for fungus determination and Graeme
Butler (Bratislava) for linguistic correction. Supported by the
UNESCO, AMBA/NUUR, VEGA 6002, 2/0125/09, 2/0167/9,
MVTS and the Literary Fund.
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Príloha č. 13
VRŠANSKÝ, P., VIDLIČKA, Ľ., ČIAMPOR, F., MARSH, F. 2012b.
Derived, still living cockroach genus Cariblattoides (Blattida: Blattellidae)
from the Eocene sediments of Green River in Colorado, USA.
Insect Science 19: 143-152.
Insect Science (2012) 19, 143–152, DOI 10.1111/j.1744-7917.2010.01390.x
ORIGINAL ARTICLE
Derived, still living cockroach genus Cariblattoides
(Blattida: Blattellidae) from the Eocene sediments
of Green River in Colorado, USA
Peter Vrˇsansk´y1,2
, L ubom´ır Vidliˇcka3,4
, Fedor ˇCiampor Jr3
and Finnegan Marsh5
1
Geological Institute, Slovak Academy of Sciences, Bratislava, Slovakia, 2
Paleontological Institute, Russian Academy of Sciences, Moscow,
Russia,3
Institute of Zoology, Slovak Academy of Sciences,4
Department of Biology, Faculty of Education, Comenius University, Bratislava,
Slovakia,5
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA
Abstract Cariblattoides labandeirai sp.n. from the Eocene sediments of Green River in
Colorado, USA bear only two plesiomorphies, but also several significant autapomorphies
within the advanced and highly derived living cockroach genus. Thus, Cariblattoides with
extant occurrence in the Caribbean and South America was historically common in the
Nearctic, and represents important evidence for the occurrence of derived living genera
of cockroaches ∼50 Ma ago. Generally, the vast majority of living genera were absent
during the Palaeocene, thus the diversification of most living cockroach lineages near
the Palaeocene/Eocene boundary must have been extremely rapid. Females of living C.
suave, the type species, have identical (sophisticated) coloration of pronotum, but the most
related living taxa are C. piraiensis and C. fontesi from Brazil (supported by phylogenetical
analysis).
Key words Blattida = Blattaria = Blattodea, Cariblattoides, Eocene, fossil insects,
Green River, Tertiary cockroaches
Introduction
Among 11 cockroach genera (17 species) found in the
Green River locality, Colorado, USA, nine represent still
living recognised taxa (genera). Only the genus Blattella,
although advanced in behavior (female bearing
ootheca until nymphs emerge), can be considered historically
primitive, because it is recorded from the Albian
Cretaceous Mesozoic (Vrˇsansk´y, 2008). All other living
genera of cockroaches are recorded only starting from
the Eocene, and as with Ectobius among those found in
Correspondence: Peter Vrˇsansk´y, Geological Institute, Slovak
Academy of Sciences, D´ubravsk´a cesta 9, P.O. Box 106,
840 05 Bratislava, Slovakia. Tel: +421 2 59203620; email:
geolvrsa@savba.sk
This work was generated during the stay of the author in the
NMNH, Washington, DC, USA.
Green River, are modern. Nevertheless, one of two genera
with derived morphology, the Cariblattoides, suggests
that even derived cockroach genera evolved during or before
the Eocene. Taking into consideration absence of
any living genera except Blattella before the Eocene, and
presence of relic Mesozoic taxa in the Eocene, it seems
that the living cockroach fauna evolved extremely rapidly
near the Palaeocene/Eocene boundary. This is supported
by several thousands cockroaches known from the terminal
Mesozoic, with occurrences of exclusively Mesozoic
cockroach families except the primitive Blattellidae related
to Symploce and Blattella. It is very unlikely that
such a rich record would not reveal some other representatives
of this, starting from the earliest Cretaceous,
dominant cockroach family.
The genus Cariblattoides is contemporary in Cuba,
Puerto Rico, Guadaloupe, French Guiana and Brazil, and
in the past was apparently distributed more widely, at least
in North America, and was present in different remote localities
at Green River.
C 2012 The Authors
Journal compilation C Institute of Zoology, Chinese Academy of Sciences
143
144 P. Vrˇsansk´y et al.
sinnamariensis
fontesi
gruneri
guyanensis
minor
unicolor
suave
belenensis
instigator
neoinstigator
albomarginata
matogrossensis
labandeiraisp.n.
piraiensis
100
38
7
23
15
30
23
53
30
15
46
38
46
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
1
0,9
0,8
0,7
0,6
0,5
0,4
0,3
0,2
0,1
Similarity
Fig. 1 Cluster analysis of Cariblattoides species. Jaccard similarity measure, paired group algorithm, with cophenetic correlation
coefficient 0.838 9.
Material and methods
The material was collected by David Kohls and Louis
Pribyl in the Anvil Points (AP) – Labandeira Site (LS);
Denson Site (DS) and Parachute Creek (PC) of the Green
River Locality in Colorado, USA. All the 13 specimens
of the present taxon are deposited in the National Museum
of Natural History (NMNH), Washington, DC. The
numbers represent official NMNH numbers, site number
(40193 – Anvil Points; 41088, 41678 – Anvil Points –
Labandeira Site; 41619 – Denson Site; 41142 – Denson
Site 3, 4, 5; 40190 – Paleoburn; 41139 – Parachute Creek
– Gunderson)/ official USNM catalogue number (these
do not correspond with the numbers on the rocks).
The photographs were made using an Olympus SZX12
stereozoom microscope and the figure represents redrawn
photographs with Corel Draw 13 and Adobe Photoshop
6.0.
Cluster analysis (Fig. 1) was made using PAST 1.43
(Hammer et al., 2001); parsimony analysis was performed
using PAUP∗
software version 4.0b10 (Swofford, 2002),
with a tree bisection reconnection (TBR) heuristic search
of 10 000 replicates and the option ‘save multiple trees’
activated. All characters were treated as ordered (0 – plesiomorphic,
1 – apomorphic state). MaxTrees option was
set to 500. All 13 morphological characters were set as
ordered and equally weighted except character no. 10. As
simple CuA (present in most living Cariblattoides, but
not in C. labandeirai) is a strong apomorphy, homoplasically
present also in some others, basal blattelid genera
(e.g., in the primitive Supella this character is polymorphic),
we set it at a higher (5) weight (branched CuA is a
very strong plesiomorphy within Cariblattoides). Heuristic
search produced 46 equally parsimonious trees with
length 30 (consistency index [CI] = 0.5, retention index
[RI] = 0.643). Majority rule consensus revealed most of
the nodes resolved in more than 50% of the trees produced
by heuristic search (Fig. 2). Terminology of wings follows
Vrˇsansk´y (1997).
Results
Systematic paleoentomology
Blattida Latreille, 1810
Blattellidae Karny, 1908
Cariblattoides Rehn et Hebard, 1927
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Journal compilation C Institute of Zoology, Chinese Academy of Sciences, Insect Science, 19, 143–152
Eocene Cariblattoides from North America 145
Fig. 2 Majority consensus tree of the parsimony analysis (46 equally parsimonious trees with length 30 [for details see Material and
Methods]) of Cariblattoides species with Cariblatta used as outgroup. Numbers above branches show group frequencies (in %).
Type species Cariblattoides suave Rehn et Hebard,
1927. Extant, Puerto Rico.
Composition – Princis (1969) and additions In addition
to the type species, C. albomarginata Rocha e Silva
Albuquerque, 1967 (Brazil), C. belenensis Rocha e Silva
Albuquerque, 1964a (Brazil); C. fontesi Rocha e Silva
Albuquerque, 1954 (Brazil); C. gruneri Bonfils, 1975
(French Guiana); C. guyanensis Bonfils, 1975 (French
Guiana); C. instigator Rehn et Hebard, 1927 (Cuba);
C. matogrossensis Rocha e Silva Albuquerque, 1958
(Brazil); C. minor Rocha e Silva Albuquerque, 1964b
(Brazil); C. neoinstigator Rocha e Silva Albuquerque,
1958 (Brazil); C. piraiensis Rocha e Silva Albuquerque,
1955 (Brazil); C. sinnamariensis Bonfils, 1975 (French
Guiana); C. unicolor Rocha e Silva Albuquerque, 1964b
(Brazil). All extant.
Diagnosis – Rehn and Hebard (1927): “Size small,
form depressed, females slightly broader and heavier than
males, size approximately similar. Head subdepressed,
distinctly and broadly visible cephalad of pronotum; interocular
space wide . . . , . . . maxillary palpi with third
palpomere (joint in the original text) elongate, slender;
fourth palpomere shorter than third palpomere; first
palpomere slightly shorter or slightly longer than fourth
palpomere. . . . Tegmina elongate lanceolate, considerably
surpassing the abdomen in both sexes. Costal margin moderately
arcuate (proximad), sutural margin almost straight:
radial (scapular in the original) field broad: mediocubital
veins (discoidal sectors in the original) longitudinal, six
to seven in number (including the media (median in the
original) and cubital (CuA – ulnar in the original) and
rami of the median veins); anal groove (sulcus in the
original) strongly arcuate proximad, straight oblique in
greater portion of length; anal field elongate pyriform;
diagonal channel of right tegmen well indicated. Wings
elongate, relatively narrow, moderately iridescent: subcosta
(mediastine in the original) and a number of costal
veins clavate; ulnar vein quadriramose; axillary vein with
three rami in distal two-thirds; intercalated triangle small
but distinctly and clearly defined . . . ”
Character analysis (0 – plesiomorphy; 1 – apomorphy
relative to other species within genus and/or Cariblatta,
which was chosen as an outgroup based on high similarity,
but retention of all original states of characters due to
standard habitus (not derived like in Cariblattoides):
1. Head significantly elongated: apomorphy; plesiomorphic
state is elongated, but more or less
of normal cockroach appearance, i.e., less than
1.2 times longer than wide (head more or less standard
in both Cariblatta Hebard, 1916 and Neoblattella
Shelford, 1911)
2. Pronotum with coloration concentrated in cervical
structures: apomorphy; plesiomorphic state is coloration
simple or dark stripes are more primitive in
the Blattellidae
3. Terminal palpomere short: apomorphy; plesiomorphic
state is the terminal palpomere long (also in
Cariblatta and Neoblattella)
4. Terminal palpomere cup-like: apomorphy; plesiomorphic
state is normal shape of the terminal
palpomere such as in Cariblatta and Neoblattella
C 2012 The Authors
Journal compilation C Institute of Zoology, Chinese Academy of Sciences, Insect Science, 19, 143–152
146 P. Vrˇsansk´y et al.
(even when the terminal palpomere is homoplasically
cup-like in several unrelated Blattellidae)
5. Forewing elongated more than 3.5:1: apomorphy
(elongation itself is an autapomorphy of the genus);
plesiomorhic state is the forewing normal, as in majority
of cockroaches including the outgroup, Cari-
blatta
6. Forewing with distinct costal margin: apomorphy
(plesiomorphically absent in Cariblatta, Neoblattela
and most Cariblattoides species)
7. Forewing monochromatically colored: apomorphy
(plesiomorphic coloration of forewing is medial
(with more dark central stripe) in some Cariblatta
and Neoblattella; the coloration in the later two genera
vary, except for the coloration mentioned, the
coloration is different from those appearing in Cari-
blattoides)
8. Forewing colored monochromatically medially, with
more dark central stripe: plesiomorphy (see above),
monochromatic median coloration without the central
stripe is an apomorphy
9. Forewing RS indistinct, terminal R(+RS) branches
not dichotomized: apomorphy (plesiomorphically is
RS distinct and/or terminally branched like in all
studied Cariblatta and Neoblattella – the eventual
reduction is homoplasic)
10. Forewing M+Cu with over 10 branches: plesiomorphy
(also in Neoblattella); apomorphy is reduced
number
11. Forewing CuA branched: strong plesiomorphy in
primitive Blattellidae (homoplasically simple in
Cariblatta)
12. Hindwing with simplified RS: apomorphy (branched
in otgroup Cariblatta and rest Blattellidae)
13. Hindwing monochromatic, pale: plesiomorphy
(state in Neoblattella and Cariblatta); apomorphic
is any other derived coloration pattern.
Cariblattoides labandeirai sp. n.
Holotype. 41619/542284-AB. Part and counterpart of
a complete ?male (Figs. 3a, 4, 5).
Type locality. Denson Site 1998, Green River, Colorado,
USA.
Type horizon. Green River Formation, Eocene, Ter-
tiary.
Paratypes: 40190/542285; 40193/542288 (AP),
542286, 542287, 542289; 41088/542290, 542291 (AP
LS 95); 41139/542292 (PC) (Diptera collection databasis);
41142/542293, 542294 (DS 4); 41678/542295(8)9,
542297 (LS 99). All the same locality and horizon as the
type (for sublocalities see M&M).
Differential diagnosis The present species can be
differentiated from all living Cariblattoides species by
branched forewing median vein M, terminally branched
CuA and strong, distinct and black, overlapping apex
costa, and by differentiated hindwing R1. All other characters
are present in some of the living species, but in
different combinations (see discussion for remarks and
comparison).
Description Head free, often preserved in upright position;
palps long, the last segment short and cup-like.
Antennae very soft and long, with at least 90 segments.
Pronotum coloration as in Fig. 3A. Both wings strongly
elongated, with reduced venation. Forewing extremely
elongated (length/width 9–10 mm/2.5 mm), with sharpened
apex, with extremely short simple Sc. Radial field
very narrow (not reaching half of the wing’s width) with
main stem of R nearly straight; rich simple R (15 in holotype)
ascending parallel, without secondary branches; M
rich (10 in holotype), but short, with parallel branches descending
directly from the main branch in angle comparable
to ascendance of R, a single M is dichotomised, rest
are simple; CuA reduced to a single branch dichotomised
near margin (2 veins meet margin); few simple A present
(4 or 5 in holotype), except for A1 all meet posterior margin.
Dark forewing coloration restricted to longitudinal
stripe along the clavus and central part of the medial and
cubital areas, reaching to the apical part of the radial area.
Hindwing Sc simple, very short – end before the wings
halve; RS differentiated, mostly simple – a single branch
is dichotomised (9 veins meet margin in the holotype). R1
consists of onerichlybranched(6veins at themargininthe
holotype) branch vein, with dense secondarily branched
veins; M simple, slightly curved; CuA reduced to 2 (possibly
3 branches, with 3–4 veins at margin), CuP simple,
copying the posterior-most CuA; A1 widely branched (4).
Vannus pleating veer-like. Body soft and thin. Sterna and
terga colored posteriorly. Legs long with margins colored,
hindleg femur very long.
Remarks and comparison See discussion.
Derivation of name After Conrad C. Labandeira, a
superb teacher, scientist and one of the collectors of the
Green River material.
Character of preservation 13 complete specimens.
Discussion
In addition to characteristic unconfuseable elongated
habitus and size with extremely long legs,
strong autapomorphies such as elongated head, hindwing
with widely branched A1 or A2, maxillary
palps with characteristic ratio of respective
palpomeres (?:1:0.9:1:0.8 – see Figs. 3A, 4D) with
C 2012 The Authors
Journal compilation C Institute of Zoology, Chinese Academy of Sciences, Insect Science, 19, 143–152
Eocene Cariblattoides from North America 147
Fig. 3 Cariblattoides labandeirai sp.n. Holotype. NMNH 41619/542284-A. (A) A complete ?male. Eocene. Green River, Colorado,
USA. Forewing length 10 mm. (B) pronotum of the female allotype (Aibonito, Guayama, Puerto Rico, July 14–17, 1914; HG Barber;
AMNH New York) of the type species, C. suave Rehn et Hebard, 1927, identical to that of the C. labandeirai sp.n. holotype. (C) palp
of C. guayanensis Bonfils, 1975.
terminal segment cup-like (homoplasically in Supella
abotti Rehn, 1947) allow the cathegorization within
the genus Cariblattoides. Large, deplanate pronotum,
slender and narrow body, pale and widely arcuate base of
forewing makes this genus with 13 living species a good
monophylectic group originating from Supella or its precursors
(retaining the original blattellid bauplan of both
wings and the lack of autapomorphies characteristic for
other genera of the Blattellidae, and synapomorphic reduction
of forewing RS and CuA, differentiate this genus
from the rest of Blattellidae). Alternatively, both genera
can had a shared history during the Green River times.
The present new fossil species share all the apomorphies
with living representatives of the genus, except the
simple CuA and branched hindwing R1. It additionally
has some secondary characters limited to Cariblattoides,
namely the sophisticated coloration of pronotum identical
with C. suave, and the coloration of forewings with
indistinct basal-most R, identical with C. fontesi. Thus it
can be safely catherorized within the genus.
On the other hand, there are some differences, which
need clarification. While the general habitus, details of
head and pronotum and wing coloration are characteristic
for Cariblattoides, the wing venation pattern is identical
with related Supella longipalpa (Fabficius, 1789) as figured
by Rehn (1951) (as Supella supellectilium). Thus
it is necessary to analyse the differences in detail. The
forewing radial area is identical with Supella and closely
related to (Supella) Namablatta Rehn, 1937 in having R
simple and in undifferentiated RS. Nevertheless, some
extinct as well as extant species of Supella (whole subgenus
Nemosupella Rehn, 1947) have RS differentiated
and venation expanded (Rehn, 1947) and this character
is polymorphic even within species of Supella and could
not be treated as diagnostic. Moreover, this character is
polymorphic even within Cariblattoides and thus represents
no contradiction of placing the present fossil within
this genus. R branches are homoplastically simple in diverse
other more or less related living cockroaches such
as in Eustegasta, Pseudomops, Ectobius, Chorisoneura,
Euphyllodromia and others, mostly with reduced venation
and/or size.
The more unusual is the character of branching of M
and CuA, descending in an angle opposite to bifurcations
of R. This character is different to that of the living
Cariblattoides and is present exclusively in Supella and
Ectobius. But again, in these genera this character of dichotomisation
is polymorphic (Rehn, 1947), apparently
due to reduction of venation in smaller species (e.g., in
Supella longipalpa) and as such is without phylogenetic
and/or taxonomic relevance.
The most significant difference with phylogenetical
information is the branched forewing CuA, suggesting
that Cariblattoides (based on the present species)
diverged before the divergence of Cariblatta and
Neoblattella spp. Within the lineage, this character
C 2012 The Authors
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148 P. Vrˇsansk´y et al.
is significantly plesiomorphic (branched) only in the
very basal Symploce (Rehn, 1951), but polymorphic
also in derived Supella (Rehn, 1947). Due
to the terminal dichotomisation, and not a fully
expressed branch, this can also eventually be a unique
character reversal or deformity of the holotype.
The clear separation of branched hindwing R1 is also
different. In contrast to other characters, this one is plesiomorphic
and polymorphic (polymorphic also in Supella),
with R1 indicated (although not richly branched)
in some living Cariblattoides species (see character 9), but
reduced in the closest relative of Supella, the Namablatta
(Rehn, 1937). R1 tends to reduce even within Polyphagidae
and Blattidae – see Rehn (1951).
Thus, Supella is similar in venation and palp, but not
in general habitus, even when S. longipalpa, the most
departured from the standard morphotype of the genus
has also a slender habitus, and S. orientalis Grandcolas,
1994 has general habitus identical with the Cariblattoides
(including the form of forewing, but head is not elongated,
and coloration with characteristic central stripe,
see Grandcolas, 1994). Thus it is possible that the precursors
of Supella were direct ancestors of Cariblattoides.
All this taken together does not contradict the placement
within Cariblattoides, although eventually would allow
us to erect a new subgenus, which we would not consider
oblique.
Additionally, Cariblattoides has flattened pronotum
larger than the most related Cariblatta and Neoblattella,
longer narrow tegmina, with subparallel margins (see
Hebard, 1916 for Cariblatta). It can be further distinguished
from Neoblattella by fewer discoidal sectors of
the tegmina and strongly deplanate pronotum (Rehn &
Hebard, 1927). Thus the sister genus is Cariblatta with
Neoblattella a sister taxon to them (Rehn & Hebard,
1927). Both Neoblattella and Cariblatta are known from
the Eocene sediments of Europe (Schmied, 2009, unpublished
observation), with the whole group apparently
derived from precursors of Supella (synapomorphic in
elongated forewing with numerous parallel M branches
descending directly from the main M branch in an angle
comparable to descention of R, weakly separated RS; A
except A1 ending in posterior margin; simple CuA (in
advanced Supella); and in reduced hindwing CuA and
general venation scheme of the hindwing with separation
of basal-most RS, which is plesiomorphy of basal living
Supella – subgenus Nemosupella.
Cariblattoides labandeirai sp.n. resembles C. suave
from Puerto Rico in maxillary palp fourth joint significantly
longer than fifth, unlike in C. instigator from Cuba
where both segments are of subequal length. Nevertheless,
C. labandeirai has apical segment even shorter and
much more oval. Females of C. suave also have identical
coloration of pronotum, similar to some representatives
of the genus Cariblatta, supporting indication about their
relation (Rehn & Hebard, 1927). (Nevertheless, it must
be noted that both Cariblattoides and Cariblatta contain
species with pronota with two dark stripes as well
as species with characteristic cervical coloration.) On
the other hand, C. suave has coloured hindwing, which
was apparently pale in C. labandeirai sp.n., and more
expanded coloration of forewing, which reaches the posterior
margin in C. suave.
It is notable that in C. suave females have shorter and
broader wings than in males, but the holotype of C. labandeirai
has wings significantly elongate and thus is unlikely
to represent a female (thus this specific coloration pattern
appears plesiomorphic for both sexes). In seven collected
individuals of C. suave, the forewing measurements
(9.5–11.5/2.9–3.5 mm) of C. labandeirai were more elongate,
similar to C. instigator (−9.6/−2.7 mm). The number
of veins of C. suave and C. labandeirai in the radial
area is approximately the same (14–19), but C. labandeirai
apparently has less reduced venation in the medial
and cubital area (12 M + CuA) compared to C. suave
(6–7 discoidal veins – M + CuA), C. instigator (7 discoidal
veins in holotype), C. minor (8), C. unicolor (7),
C. guyanensis (6), C. fontesi (7) and C. sinnamariensis (9
discoidal veins in holotype) (for others, see below). The
single extant species with expanded M is C. piraiensis
(11 including one CuA). There are conservatively 5 anal
veins present in all species of Cariblattoides.
C. gruneri Bonfils, 1975 – a larger species (forewing
length/width 11.7/3.2 mm) also has simple hindwing M,
and 4 CuA branches (3 in C. labandeirai). Another larger
species is C. guyanensis (forewing length/width 11.6–
12.5/2.9–3.1 mm), which has even more expanded venation
in the radial area (±20), and reduced venation of
Media (6). The process of reduction of CuA, characteristic
for the genus, is nearly complete in this species (and
also in C. sinnamariensis) – a single CuA fuses with
the radial stem. Palp of C. guyanensis is very similar
to C. labandeirai (perhaps a synapomorphy). Hindwing
of C. guyanensis has apomorphically widened apexes of
radial branches (even more expressed in C. sinnamariensis),
present cross-veins (most likely an autapomorphy),
but branched M (plesiomorphic even in respect to C. labandeirai).
C. sinnamariensis has plesiomorphically (in
respect to all the known species) branched hindwing
CuA (7).
C. minor is another comparatively large species, with
less prolonged tegmina (12/3.5 mm) and pronotum coloration
similar to C. labandeirai, but with finer dark
pattern.
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Eocene Cariblattoides from North America 149
Fig. 4 Cariblattoides labandeirai sp.n. Holotype. NMNH 41619/542284. A complete ?male. Eocene. Green River, Colorado, USA.
(A) general habitus; (B) forewing; (C) hindwing; (D) pronotum and head; (E) terminalia and hindwing. Forewing length 10 mm.
Scales = 1 mm.
C. unicolor differs in having uniform coloration of significantly
elongated forewing (12/3 mm).
C. fontesi has comparatively robust forewing
(10/3 mm), but coloration similar to C. labandeirai. Similar
lengths are present also in the palps, but the apical
palpomere is not oval as in C. labandeirai and C. piraiensis.
Hindwing of C. fontesi has more expanded venation
in the RS area (plesiomorphy).
C. piraiensis has palp entirely identical with C. labandeirai,
with oval cup-like terminal segment and also
identical forewing coloration (in some specimens – this
character varies). It is also a single species with expanded
forewing M (10). Pronotum coloration is also similar.
Nevertheless, this species differs in having less elongated
forewings (under 4:1) – a plesiomorphy and expanded
hindwing RS (plesiomorphy).
C. mattogrossensis and C. neoinstigator have pronotum
with two dark stripes and expanded forewing RS. Their
tegmina are less elongated (under 3.5:1).
C. albomarginata with the forewing length 11 mm is
likely the taxon with the most primitive characters, as it
has shortest head, widest wing, rich R (20), and standard
dichotomisation of M (with both branches descending
at the same angle) resembling Neoblattella (in Cariblattoides
posterior branches tend to descent from the straight
stem). On the other hand, it has simplified M (6) and simple
CuA (both synapomorphies of living representatives
of the genus; expanded M is characteristic also for C.
piraiensis) and uniformly colored center of the pronotum.
C. belenensis with the most simplified M (5), simple
CuA and (as with all known species) 5 anal veins, and
simplified hindwing CuA (3), appears to be the most
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150 P. Vrˇsansk´y et al.
Fig. 5 Cariblattoides labandeirai sp.n. (A) Holotype NMNH 41619/542284-B; (B) NMNH 41139/542292 (PC) (Diptera collection
databasis); (C) NMNH 40193/542286 (AP); (D) NMNH 41142/542293 (DS 4). Eocene. Green River, Colorado, USA. Scales = 10 mm.
Table 1 Cariblattoides character matrix of 13 extant and the present extinct species of Cariblattoides, and Cariblatta spp. as outgroup
(OG) (the same dataset is provided for related Neoblattella and Supella as well as the rest of Blattellidae).
sp/character 1 2 3 4 5 6 7 8 9 10 11 12 13
Cariblatta (OG) 0 0 0 0 0 0 0 0 0 0 ? 0 0
albomarginata 0 0 0 0 0 0 0 0 0 1 1 ? 1
belenensis 1 1 1 0 1 0 0 0 ? 1 1 ? 0
fontesi 1 0 1 0 0 0 0 1 1 1 1 1 0
gruneri 1 ? 0 0 1 0 1 ? ? 1 1 ? 0
guyanensis 1 1 1 0 1 0 1 ? 0 1 1 0 0
instigator 1 0 0 0 1 0 0 0 ? 1 1 ? 1
labandeirai sp.n. 0 1 1 1 1 1 0 1 1 0 0 1 0
matogrossensis 0 0 1 0 0 0 0 0 0 1 1 0 0
minor 1 1 1 0 1 0 1 ? 1 1 1 ? 0
neoinstigator 0 0 ? ? 0 0 0 0 1 1 1 ? 1
piraiensis 0 1 1 1 1 0 0 0 0 0 1 1 0
sinnamariensis 1 0 0 0 0 0 1 ? 0 1 1 1 0
suave 1 1 1 0 1 0 0 0 0 1 1 1 1
unicolor 1 1 1 0 1 0 1 ? 1 1 1 ? 0
0 – plesiomorphy; 1 – apomorphy; ? unknown character. Data were obtained basing on the present study, unpublished observations and
the following references: Hebard (1916), Rehn and Hebard (1927), Rocha e Silva Albuquerque (1954, 1955, 1958, 1964ab, 1967) and
Bonfils (1975).
C 2012 The Authors
Journal compilation C Institute of Zoology, Chinese Academy of Sciences, Insect Science, 19, 143–152
Eocene Cariblattoides from North America 151
derived species (in spite of its large size with forewing
length 12 mm).
To summarise, C. labandeirai has palp identical with
C. piraiensis (synapomorphy) (nearly identical with C.
fontesi), pronotum identical with females of C. suave
(plesiomorphy) and forewing coloration identical with
C. fontesi and similar to C. piraiensis (synapomorphies).
Thus, the most related living taxon appears C. piraiensis
from Brazil.
The cluster analysis (Fig. 1) and the consensus tree of
the parsimony analysis (Fig. 2) reveal results comparable
with the empiric observation, but the relation of C. suave
and C. belenensis appears artificial.
Generally, variation within the genus includes diverse
variations in the combinations of all studied characters
(see Table 1) except for hindwing (with an exception of
coloration, hindwing venation is principally identical in
all living representatives – variable only in the number of
veins in RS area), CuA (simple in all extant species) and
colored costa of C. labandeirai. The conservative pattern
of hindwing (with unmodified radial area [most significantly
involved in flight], and more similar to that of
Supella and primitive Symploce) in the earliest C. labandeirai
infers the strong selection due to active flight (and
elongated habits) in all living Cariblattoides. It is notable
that extinct C. labandeirai has the hindwing radial area
(most significantly involved in flight) unmodified, and is
more similar to that of Supella and primitive Symploce.
The present species can be more easily identified than
other undescribed species – it is very distinct morphologically
and as such is easily determinable (in contrast
to most other species at the locality, which can be confused
according to the preservation of body only, and as
such belong to indetermined specimens). Otherwise the
preservation is standard (with an exception of the holotype
which is the only specimen at the locality with both
wings visible), and even in these completely preserved
specimens, venation is indistinct due to overlap with body
structures.
All specimens have size and coloration within normal
intraspecific variability range as the type species and thus
very probably belong to a single biological species. Nevertheless,
a closely related taxon due to eventually belonging
to slightly different layers cannot be excluded. This is
a general problem of palaeoentomology and authors are
not aware of any larger type series which would not face
this problem. Thus the material is included in the type
series, clearly representing the same morphospecies, but
eventually not the biological species.
The presence of a derived living genus in the Eocene
suggests the radiation of newly evolved living genera
after the Paleocene/Eocene boundary must have been extremely
rapid (it indicates that nearly all living genera
– even such advanced ones as Cariblattoides – evolved,
speciated and radiated within 5 Ma at most). The earliest
representative of such a derived genus, the present C. labandeirai
cannot be considered more primitive than most
of the living species. Radiation but also origination of
modern genera of cockroaches thus could be associated
with the thermal maximum (PETM) and massive invasions
of tropical elements polewards – into unoccupied
habitats.
Conclusions
• Cariblattoides labandeirai sp.n. was a common
species in the Eocene assemblage of the Green River.
Nevertheless, according to taphonomic advantages,
13 of 289 identified cockroaches may be a little overestimated
figure compared to other species.
• It was a rather advanced taxon within the genus,
apomorphic in 8 of 13 characters, with only two
significant plesiomorphies (dichotomized forewing
CuA and branched hindwing R1).
• Most closely related living species is C. piraiensis
from Brazil.
• The presence of this derived blattelid genus during
the Eocene indicates the radiation of most living
cockroach genera must have taken place in a short
time interval near the Palaeocene/Eocene boundary.
Acknowledgments
We thank David Kohls and Louis Pribyl (NMNH Washingtn
DC) for collecting the material, and Conrad Labandeira
for organising the study; Alexandr P. Rasnitsyn
(PIN Moscow), Sonia M. Lop´es (MN/UFRJ), Pavel ˇStys
(CU Prague) and Heiko Schmied (SI Bonn) for fruitful
advice and revision of the manuscript, and Ladislav
Roller (ZI Bratislava) for literature supply. Supported by
the UNESCO, Amba, National Museum of Natural History,
Washington D.C. award, VEGA 6002, 2/0125/09,
2/0167/09, MVTS and the Literary Fund.
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Príloha č. 14
VRŠANSKÝ, P., VAN DE KAMP, T., AZAR, D., PROKIN, A., VIDLIČKA, Ľ., VAGOVIČ, P. 2013a.
Cockroaches Probably Cleaned Up after Dinosaurs.
PLoS ONE 8(12): e80560.
Cockroaches Probably Cleaned Up after Dinosaurs
Peter Vrsˇansky´1,2
*, Thomas van de Kamp3
, Dany Azar4
, Alexander Prokin5,6
, L’ubomı´r Vidlicˇka7,8
,
Patrik Vagovicˇ3,9
1 Geological Institute, Slovak Academy of Sciences, Bratislava, Slovakia, 2 Arthropoda Laboratory, Paleontological Institute, Russian Academy of Sciences, Moscow, Russia,
3 ANKA/Institute for Photon Science and Synchrotron Radiation (IPS), Karlsruhe Institute of Technology (KIT), Eggenstein-Leopoldshafen, Germany, 4 Faculty of Science II,
Natural Sciences Department, Lebanese University, Fanar, Fanar-Matn, Lebanon, 5 I.D. Papanin Institute for biology of inland waters Russian Academy of Sciences, Borok,
Russia, 6 Voronezh State University, Voronezh, Russia, 7 Institute of Zoology, Slovak Academy of Sciences, Bratislava, Slovakia, 8 Department of Biology, Faculty of
Education, Comenius University, Bratislava, Slovakia, 9 Institute of Multidisciplinary Research for Advanced Materials, Tohoku University, Japan
Abstract
Dinosaurs undoubtedly produced huge quantities of excrements. But who cleaned up after them? Dung beetles and flies
with rapid development were rare during most of the Mesozoic. Candidates for these duties are extinct cockroaches
(Blattulidae), whose temporal range is associated with herbivorous dinosaurs. An opportunity to test this hypothesis arises
from coprolites to some extent extruded from an immature cockroach preserved in the amber of Lebanon, studied using
synchrotron X-ray microtomography. 1.06% of their volume is filled by particles of wood with smooth edges, in which size
distribution directly supports their external pre-digestion. Because fungal pre-processing can be excluded based on the
presence of large particles (combined with small total amount of wood) and absence of damages on wood, the likely source
of wood are herbivore feces. Smaller particles were broken down biochemically in the cockroach hind gut, which indicates
that the recent lignin-decomposing termite and cockroach endosymbionts might have been transferred to the cockroach
gut upon feeding on dinosaur feces.
Citation: Vrsˇansky´ P, van de Kamp T, Azar D, Prokin A, Vidlicˇka L, et al. (2013) Cockroaches Probably Cleaned Up after Dinosaurs. PLoS ONE 8(12): e80560.
doi:10.1371/journal.pone.0080560
Editor: Ulrich Joger, State Natural History Museum, Germany
Received April 23, 2013; Accepted October 4, 2013; Published December 4, 2013
Copyright: ß 2013 Vrsˇansky´ et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: Supported by UNESCO-Amba (MVTS), VEGA 6002, 02/0152, 2/0186/13, Literary fund, Schwarz stipend. This work was supported by the Slovak Research
and Development Agency under the contract No. APVV-0436-12. This paper is a contribution to the team project (ER023: Biodiversity: Origin, Structure, Evolution
and Geology) awarded to DA by the Lebanese University. Most funds covered the home institutions. The funders had no role in study design, data collection and
analysis, decision to publish, or preparation of the manuscript.
Competing Interests: The authors have declared that no competing interests exist.
* E-mail: geolvrsa@savba.sk
Introduction
The Triassic, Jurassic and Early Cretaceous terrestrial ecosystems
differed from extant ecosystems for various reasons, one of
them being the presence of gigantic reptiles. The energy flow was
principally less efficient (more rapid) and also the general
appearance of the landscape was dissimilar [1,2]. Grasses, flowers
with their fruits, large butterflies, and before the latest Jurassic, all
eusocial insects (cockroaches, termites, ants, bees) were absent
[3,4]. Discerning between dinosaur feces decomposers (which were
not identified until now) is also essential as it changes the general
appearance of our assemblage reconstructions. Moreover, the
problem is of a principal, systemic importance. If nothing fulfilled
this role, a large amount of dung would prevent soil regeneration
just as it suffocated the pasture systems and prevented grass
regeneration in present-day Australia [5]. Grasses were absent
before the Early Cretaceous, but such influence will definitely alter
extinct cenoses similar to some extent to the variety of living fern
groups or perhaps taxa such as Gnetum and Ephedra. On the other
hand, bird droppings are known to significantly (often positively)
influence vegetation composition of ombrotrophic bogs [6]. Late
Cretaceous biomes actually contain grasses and silicified plant
tissues (phytoliths) preserved in the Maastrichtian coprolites
(presumably from titanosaurid dinosaurs) from the Lameta
Formation in India show that at least five taxa from extant grass
(Poaceae) subclades were present during the latest Cretaceous [7].
Was the Mesozoic world full of sterile dinosaur dung, clean as a
modern forest, or transitional between these two extremes?
Circumstantial evidence of dinosaur (probably hadrosaur) coprolites
[8,9] suggests that feces were used. The absence of dungbeetles
during the Triassic and near-absence during most of the
Jurassic [10] (roughly half of the age of dinosaurs) and their
radiation associated only with the spread of modern grasslands [1]
is still under discussion [2].
Feces have a greater capacity to retain moisture than the parent
plant tissue [11] and coprophages exploit the microbial consortia
concentrated on these recycled cellulose-based foodstuffs; the
microorganisms serve not only as a source of nutrients and gut
mutualists, but they also pre-digest recalcitrant substrates [12].
Microbial dominance is so pronounced that fecal pellets may be
considered as living organisms [12]. They consist largely of living
cells, they consume and release nutrients and organic matter, and
they serve as food for animals higher on the food chain [13].
Any excrement is a valuable source of nitrogen, and its amount
must have been huge [14] at least seasonally [15], during the age
of dinosaurs. Each single separate dung might have had a volume
of 7 liters [8]. Probably an important feature of dinosaur and
pterosaur excrements (as in birds and reptiles when compared with
mammals) was the large proportion of nitrogen compared with
phosphorus [16]. The association with urine and thus with a high
concentration of phosphoric acid, oxalic and carbonic acids and
salts, primarily sodium chloride, leads to the recent conclusion
PLOS ONE | www.plosone.org 1 December 2013 | Volume 8 | Issue 12 | e80560
about the association of dung-beetles and coprophagy with
mammals (not with dinosaurs) since the very beginning [17]. On
the other hand, some common (11 of the 15 deposits) fossilised
dinosaur coprolites contain 13–85% of rotting conifer wood with
only 0.20–0.30% of nitrogen (conifers are utilized by the living
cockroach Cryptocercus – the most important wood-decomposing
cockroach) with its attendant microbial and detritivore fauna and
thus augmented the resource options of Cretaceous ecosystems
that lacked fodder provided by grasses and other derived
angiosperms [8,18]. The consistency of the coprolites during the
deposition varied from fairly cohesive to viscous liquid and fluid to
some extent – those containing a significant amount of wood are
most easily recognizable as their high wood content prevented
degradation [8].
In addition to dung, it has recently been proposed that the
density of sauropods was high enough to produce the amounts of
methane necessary for sustaining the warm climate during the
Mesozoic [19].
The cockroach family Blattulidae, described by Vishniakova
[20] originated in the Late Triassic and constitutes a (co-)dominant
group of insects (,1%) throughout the whole Jurassic and
Cretaceous [21]. They are often completely preserved [22–24]
and contributed to knowledge of some general patterns such as the
decreasing variability of species over time, and mass mutations
[25,26]. The Blattulidae constitute the sole cockroach fossils
preserved in several Cretaceous localities such as Shin Khudukh
and some others in Mongolia and Verchnebureinskaja Vpadina in
Russia, and are the dominant insect fossils in diverse Mesozoic
ambers [27,28]. The hypothesis tested and supported in the course
of the present research was the heterogeneous character of the diet
of these Mesozoic cockroaches (in contrast to homogeneous one of
all the studied Cenozoic and present ones). There are numerous
Tertiary (Cenozoic) cockroaches preserved with the gut-content,
but all of them have a homogeneous diet. The same holds for the
studied living cockroaches. The occurrence of any wood (digested
twice, a second time by cockroaches, after it was previously
digested by herbivores; Figs. 1E, S1) was entirely unexpected.
Protozoan cysts and helminth eggs preserved in the Early
Cretaceous Iguanodon coprolite represent the only reported case of
dinosaur parasites [29], but the discovered trophic relation of
dinosaur-age vertebrate herbivore and insects might appear
important also due to the structuring of the extinct ecosystems
via parasites (and pathogens) transferred. Trophic association of
Mesozoic vertebrates and insects suggest endoparasite transfer as
well.
A similar transfer is known from numerous living species, e.g.,
from Blatta orientalis and Periplaneta americana feeding on human
excrement that contained cysts of Chilomastix mesnili and rats eating
food that had been contaminated with feces from these
cockroaches became infected with this protozoan [30].
Materials and Methods
The material studied herein is from Mdeirij-Hammana, Baabda
District Governorate Mount Lebanon, Central Lebanon - detailed
coordinates for the localities of completely studied specimens
(mostly immatures: (59, 76A, 623i-m, 778AB, 799, 800, 810CD,
845AB, 934AB, 1062, 1274B,D, FAL -3C (Falougha), 133.C,
JEZ.F-14 (Wadi Jezzine, Jezzine District, Governorate Southern
Lebanon), 1669-B, RIH-33 (Rihane outcrop, Jezzine District,
Governorate Southern Lebanon), (deposited at the Lebanese
University); AMNH Lebaneese amber 22, 77, 84, 91 (Bcharreh
District, Governorate North of Lebanon; Jouar Ess-Souss,
Bkassine, Jezzine District, Governorate Southern Lebanon, all
deposited in the American Museum of Natural History), J. lebani
holotype (Jouar Ess-Souss, Bkassine, Jezzine District, Governorate
Southern Lebanon, Acra collection) can not be revealed due to site
protection [31], in a Lower Cretaceous (ca. 120 Ma) amberbearing
deposit. An enicocephalid assassin bug, three ceratopogonid
biting midges, and two male coccids occur as syninclusions.
Examined specimen (1094A-I) was not embedded in epoxy resin
due to ST examination, but for photography a drop of maple sirup
and a coverslip glass was attached to see inside. It is deposited at
the Lebanese University, Faculty of Sciences II, Lebanon. We
performed a microtomographic scan of the amber piece (0.185 g,
well transparent dark yellow-red sample) at the full-field X-ray
imaging station TopoTomo beamline of the ANKA light source.
The scan covered 180 angular degrees with 2,800 radiographic
projections measured. We used a filtered white beam radiation
with a spectrum peak at ,20 keV. A sample-to-detector distance
of 35 cm resulted in both absorption contrast and edge enhancing
phase contrast in the projection images. These were recorded by
an indirect detector system based on a scintillator coupled to an
optical microscope and a CCD detector [32]. The magnification
factor of the optical microscope was 22.4 which led to an effective
pixel size of 0.4 mm with attached CCD camera pco.4000 with
400862672 pixels. We processed each radiographic projection
using a single distance phase retrieval algorithm [33] integrated in
ANKA phase plugin [34] for ImageJ and reconstructed the
volume by PyHST reconstruction software [35]. The triangle
algorithm is unknown, but the original surfaces contain so many
polygons that the details lost to a reduction to 10% are negligible.
For segmentation of the coprolites we used software Amira 5.4.
After loading the volume data as an image stack of virtual slices,
we labelled the whole coprolites and the dense particles with the
segmentation editor of the program. We exported and reassembled
the surface models from the labels with the software Cinema
4D R12. Volumes were calculated from the polygon meshes using
the GeoTools2010 plug-in.
Before creating the interactive 3D graphics, we reduced the
surface polygons once more to 10%. The objects were saved as
Collada files and opened with the software Right HemisphereH
Deep Exploration 6. After creating the object hierarchy, we saved
the data as Universal 3D files, opened with AdobeH AcrobatH 9
Pro Extended, and integrated into PDF files.
Results
Distribution of the Blattulidae is associated with the abundance
of dinosaurs (fig. 2F). In the Lebanese amber, the Blattulidae
constitute 8 of the 15 identified (21 studied) cockroach samples
including Ocelloblattula ponomarenkoi Anisyutkin et Gorochov, 2007
[36], in addition to the Umenocoleidae (n = 1), Caloblattinidae
(n = 2), Raphidiomimidae (n = 1), Liberiblattinidae (n = 1), Blattellidae
(n = 2), and Mesoblattinidae (n = 2; Nymphoblatta azari) [37].
The present fossil (Fig. 1) can be categorized as belonging to
Blattulidae on the basis of small size, chaetotaxy and a significant
comparative specimens of amber which include both immatures
and adults [28,38]. Its characteristics are a small size, large head,
antennae with corrugated surfaces, and with 2–3 rows of long
sensilla (Fig. 1BC), pronotum and abdomen with two longitudinal
stripes, cerci with long spurs and extremely long sensilla, legs short.
Especially notable are round elevated pronotal structures of the
present nymph (see Fig. 2B), somewhat resembling lanterns (A
lantern is a specialised light-producing organ of cockroaches.) of
the luminescent cockroaches of the genus Lucihormetica [39,40].
The diet of the Blattulidae is revealed for the first time. Five
coprolites (the last one still protruding from the abdomen) that are
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elliptical in shape and circular in cross section (volumes
847,381 mm3
, 2080,512 mm3
, 2401,192 mm3
, 3435,904 mm3
,
4597807 mm3
) (Fig. 1E, S1) amounting to a total volume of
13362,796 mm3
, and about 0.35 mm long contain heterogeneous
material. They are preserved in a single piece of amber, adjacent
to a fossil of the Early Cretaceous cockroach, and represent a
new type of trace fossil (coprolite adjacent to a preserved dead
organism) that will be designated elsewhere. 1.06% (141,081 mm3
)
is filled by partially digested particles of wood. The structure of the
wood is revealed on the largest particles and the lignin bilayer (part
of the numerous parenchymatous tangential ray cells) is apparent
on Fig. 2a and S1. The distance among parenchymatous
tangential cells is roughly 10 mm.
The surfaces are smooth and the edges of the particles are
rounded even in the largest particles (and also inside of cavities).
The size of them (ca. 30,000 mm3
) is still very small when
compared to the mouthpart and mouthful size (e.g., particles of the
cockroaches of this size often reach 0.4 mm at the widest point).
Wood within the present coprolites has a characteristic, possibly
power law distribution of particles larger than 100 mm3
(distribution
curve at Fig. 2F can be characterised with the equation
y = 21.964x +10.695; y = log (size); x = log (number of debris)), but
the frequency of smaller particles decreases (Fig. 2D) at 100 mm3
,
which is far enough to be recorded by the present technique
(effective pixel sizes below 0.5 mm are common for the present
synchrotron (ST)). The wood particles are not distributed
concentrically and/or in an otherwise ordered way.
Additionally, this wood is apparently decayed in the hind gut
(intestine and/or rectum - as in termites - not in mid gut or
stomach) as the last incompletely formed coprolite (caused by
stress-defecation and still extruding from the body) contains
numerous larger wood particles (S1). This enhanced gut activity is
documented by the amorphous structure of the coprolite apparent
in the sections (Fig. 2C).
The distribution curve of the wood particles is ambiguous. The
gut-processed particles are diminished below 100 mm3
, which is
the rough limit for the smooth edges caused by the cockroach gutprocessing.
On the other hand, the linear (in log scale) distribution
of particles, combined with rounded edges in larger scale (up to
10,000 mm3
) and the absence of small particles and isolated
tracheae (only 3 linear particles are present, and they probably do
Figure 1. Dinosaur-age cockroach of the extinct family Blattulidae. (A – head to leg end length: 3.8 mm) with antennal sensory system (B, C)
and five preserved coprolites (D – optical, E – surface rendering of numbered coprolites and dense particles based on the image stack from
synchrotron X-ray microtomography; F – ST orthoslice with labelled boundaries and fragments). Lebanon amber 1094A-I. Scales 0,5 mm.
doi:10.1371/journal.pone.0080560.g001
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not represent tracheae) in the present coprolite suggests external
pre-digestion.
Dinosaurs apparently had consumed leaves along with the
twigs, but the soft parts of leafs are unrecognizable in the ST
signal. Only the hard and dense wood particles are distinct.
Discussion
The most effective exploiters of nitrogen in animals are
cockroaches, often capable of nitrogen extraction and symbiont
transfer even from their own feces or from feces of vertebrates
including the popular guano of diverse vertebrates. Its storage and
transfer to conspecifics is thought to be used as currency in mating
and parental investment strategies [12]. Cockroaches feed on the
droppings of frugivorous, insectivorous, and haematophagous bats,
but not carnivorous bats [41]. Insect communities on the dung of
crocodiles, varanid lizards and big turtles are virtually unstudied,
and bird dung is generally too small to be utilized by a specialized
dung cohort [17]. Nevertheless, several living cockroaches are
associated with bird nests and presumed to feed on bird dung
[12,42–46]. The only large volume bird dung of the oil bird
Steatornis caripensis or guacharo (see Tab. 1) is processed by
cockroaches [47], which is another (indirect) support for the
present inferences as birds are direct descendants of dinosaurs
(often systematically cathegorized directly inside them). Numerous
authors [48] note explicitly but without specification direct
utilisation of reptile dung. Christoffersen & De Assis [49]summarise
pentastomid parasites transferred to cockroaches via feeding
on reptile and amphibian feces (see Tab. 1). Although appearing
trivial, cockroaches, one of the dominant insect orders during the
Mesozoic were never examined as representing top candidates for
partial processors of dinosaur dung.
The present specimen represents a derived secondary trace
within a trace (traces of microorganisms on wood preserved in a
coprolite–a trace of a cockroach within amber–a trace of a tree).
Figure 2. Dinosaur-age wood decomposing cockroach with coprolite and its ecological context. A) wood fragment no. 123 (coprolite
no. 3), volume 23077 mm3
(TRC- parenchymatous tangential ray cells); B) Lebanese amber (Blattulidae 1094A-I), length (head to leg end): 3.8 mm; C)
a virtual synchrotron section (,1.2 mm) through coprolite no. 3, wood particles are pale; D) percentual representation of volume of the respective
wood particles; E) distribution analysis of simple particle count of 280 wood fragments present in all five coprolites plotted over the fragment size; F)
Ratios of the Blattulidae and ‘‘Voltziablatta’’- group – families that replaced each other during the Triassic (interrupted arrow) – to all cockroaches,
plotted over the timescale (in Ma). The origin and extinction of dinosaurs are pointed with arrows. ‘‘N in %’’ means percentual representation of
number of specimens, ‘‘spp in %’’ is a percentual representation of species. Original data.
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Table 1. Distribuition of living dung-feeding cockroaches supporting their common and cosmopolitan distribution [41],
exclusively in dark (nocturnal, cave or under dung) environments.
Species Family Locality Country Dung Host Habitat Continent Reference
Arenivaga grata Corydiidae Tucson
Mountains,
USA, Arizona guano Bat Bat cave North America [83]
Blabverus discoidalis Blaberidae Bogor, Java Indonesia feces Flat-tailed
gecko
Outdoors Asia [84]
Blatta orientalis Blattidae Johannesburg
Hospital
South Africa dung Human Hospital Africa [30]
Blattella germanica Ectobiidae ? Egypt feces Human Villages Africa [85,86]
Ergaula
scarabaeoides
Corydiidae Selangor Malaysia guano Bat Bat cave Asia [87,88]
Eublaberus distanti Blaberidae Guanapo Cave Trinidad and
Tobago
dry guano Fruit bat Bat cave South America [4]
Eublaberus posticus Blaberidae Trinidad island Trinidad and
Tobago
feces Bat Indoors South America [89]
Eublaberus posticus Blaberidae Tamana cave Trinidad and
Tobago
guano Oilbird Bird cave South America [52]
Euthyrrhapha nigra Corydiidae Antsinomy
grotto
Madagascar guano Bat Bat cave Africa [90]
Gyna kazungulana Blaberidae ? East Africa guano Bat Bat cave Africa [91]
Gyna maculipennis Blaberidae Lualaba Dem Rep Congo guano Bat Bat cave Africa [92]
Opisthoplatia
maculata
Blaberidae Formosa Formosa
( = Taiwan)
dung Human Outside Asia Shikano in [93]
Paratemnopteryx
kookabinnensis
Ectobiidae Kookabinna
George
Western Australia guano Bat Cave Australia [94]
Paratemnopteryx
rufa
Ectobiidae Nullarbor Plain Australia guano Bird Cave Australia [95]
Paratemnopteryx
weinsteini
Ectobiidae Rope Ladder
Cave
Queensland guano Bat Cave Australia [94]
Parcoblatta bolliana Ectobiidae Texas USA dry dung Cow Pine woods North America [96]
Parcoblatta
fulvescens
Ectobiidae Florida USA dry dung Cow Pine woods North America [97]
Periplaneta
australasiae
Blattidae Sarawak Mt.
Jibong
Malaysia guano Bird Cave Asia [98]
Periplaneta
australasiae
Blattidae Malaysia feces Small reptiles Outdoors Asia [99]
Periplaneta
australasiae
Blattidae Punta Gorda,
Florida
South Africa dung Goat Outside; vacant
house
North America [100]
Periplaneta
americana
Blattidae Formosa Formosa
( = Taiwan)
feces Macaca
cyclopis
Indoors Asia [101]
Periplaneta
americana
Blattidae Vengurla India guano Bat Bat cave Asia [102]
Periplaneta
americana
Blattidae Sumatra Sawah
Lunto
Indonesia feces Human Coal mine Asia [103]
Periplaneta
americana
Blattidae western Bengal India feces Human Coal mine Asia [104,105]
Periplaneta
americana
Blattidae Johannesburg
Hospital
South Africa dung Human Hospital Africa [30]
Periplaneta
americana
Blattidae ? Egypt feces Human Villages Africa [85,86]
Periplaneta
americana
Blattidae Accra –
laboratory
Ghana (Gold
Coast)
feces Erythrocebus
patas
Indoor (glass
jars)
Africa [106]
Periplaneta
americana
Blattidae Araripe Brazil feces Worm lizard Outdoors South America [107]
Perisphaerus sp. Blaberidae Jalor caves Malaysia guano Bat Cave Asia [108]
Pycnoscelus
surinamensis
Blaberidae St. Croix USA, Virgin
Islands
feces Chicken Chicken roosts CentralAmerica [109]
Pycnoscelus
surinamensis
Blaberidae Puerto Rico
Mona Island
USA dry dung Cow Pine woods CentralAmerica [43]
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Although it represents a unique find in respect to both quality of
preservation in amber as well as the incidental character of the
preserved ‘‘act’’, coprolite feedings of Mesozoic cockroaches from
other families can be excluded based on the positive evidence in
the form of preserved gut contents. Several dozen species from the
sedimentary record of diverse families (Mesoblattinidae, Caloblattinidae,
Ectobiidae, Liberiblattindiae, Umenocoleidae) were
found with the gut content. All of them contain unprocessed
heterogenous organic debris, but no wood (unpublished observation),
which is irreconcilable with coprophagy. Thus the only
family adept for such duties is the family Blattulidae–the last
ecologically significant family with unstudied gut content. The
generic diversity of this family was significantly low, namely only
12 genera are present in their 80 million years of ecological
dominance. This low diversity is also represented in the fossil
inventory of the Lagersta¨tten and is direct evidence for very
uniform, constant niches and probably also for a more or less
uniform diet. This phenomenon is also visible in the unusually
minor differences between genera of the sedimentary and amber
records. This minimal diversity is highlighted to a greater extent
by the sparse disparity. With the exception of two rare species, all
Blattulidae are very similar. Uniformity is especially shown by the
transversally striated extremities. This coloration dominates in the
whole Mesozoic, but was lost at the K/Pg boundary along with the
extinction of dinosaurs, although this colouration occurs in extant,
nocturnal and arboreal Allacta australiensis under different body
colors.
Just a lack of diversity could mean it had a limited niche, one
that could be seen in modern roaches, but combined with the
longest lasting ecological dominance within cockroaches and
unique morphology (such as corrugated surface of antennae–
Fig. 2B,C), indicating the niche of the Blattulidae was different
from that of living cockroaches.
Generally, during the Mesozoic representatives of the family
Blattulidae usually comprise ,1% of all insects and over 30% of
cockroaches (Fig. 2F), and thus were probably associated with a
dominant group of vertebrates–probably sauropod dinosaurs.
Special features of the present specimen such as extremely short
and wide body with very long cerci suggest it is closely related to
Grandocularis kurnubinsi from Jordanian amber (described based on a
nymph [50] of a similar stage and size). It apparently represents a
closely related species, but differs in the form of the pronotum, eye
size, coloration and chaetotaxy. In adults, bioluminescent ‘‘lanterns’’
were apparently absent–adults of at least several species of
the Blattulidae were documented as crepuscular or diurnal, not
nocturnal–on the basis of the eye morphology and common
occurrence together with diurnal species within a single pterosaur
and/or dinosaur coprolites and/or regurgites [51]. Cockroach
nymphs occurring in dung would signal to adult ovipositing
females by a lantern system. But the detection of luminescence of
lanterns embedded in amber would be difficult. Unfortunately, the
ST signal in a large piece of amber is too weak even to reveal
morphological details and thus the presence of these morphofunctional
units cannot be validated.
One can imagine the distinct contrast coloration characterized
by distinct alternating light-and-dark stripes would be advantageous
(for communication) in an open and confined habitat of
dung surfaces. On the other hand, neither cockroach guano
dwellers nor recent ‘‘external’’ coprophages have any conspicuous
coloration. Additionally, all living coprophagous cockroaches live
concealed within and/or under dung. In nocturnal conditions of
caves, nymphs also burrow in the surface of loose guano. They
may be completely concealed, or may rest with their heads on the
surface with their antennae extended up into the air; if the guano is
compacted, the cockroaches remain on its surface and are
attracted to irregularities such as the edge of a wall, a rock, or
even a footprint [52]. In these dark conditions, guano cockroaches
Table 1. Cont.
Species Family Locality Country Dung Host Habitat Continent Reference
Pycnoscelus
striatus
Blaberidae Selangor Malaysia guano Bat Cave Asia [87,88]
Simandoa
conserfariam
Blaberidae Simandou
Mts.
Guinea guano Fruit bat Cave Africa [110]
Symploce cavernicola Ectobiidae Sarawak Mt.
Jibong
Malaysia guano Bird Cave Asia [98]
Tivia
macracantha
Corydiidae Katanga Province Dem Rep Congo guano ? Cave Africa [92]
Tivia sp. Corydiidae Antsinomy
grotto
Madagascar guano ? Cave Africa [90]
Trogloblattella
nullarborensis
Ectobiidae Nullarbor
Plain
Australia guano Bird Cave Australia [95]
Xestoblatta
hamata
Ectobiidae La Selva Costa Rica dung Bird ? Cental America [4]
Xestoblatta
immaculata
Ectobiidae Chilibrillo Panama guano Bat Cave Cental America [111]
unidentified ? ? ? dung Horse, Cow Desert ? [112]
unidentified Corydiidae ? Ecuador dung Bird Outdoors South America [12]
unidentified ? ? Malaysia feces House gecko Indoors Asia [113]
unidentified ? Hawai USA feces Giant toad Outdoors North America [114]
Feeding of diverse cockroaches on bird excrements and also facultative feeding on reptile and amphibian dungs is apparent. Based on Bell et al. [12], Christoffersen &
De Assis [49] and Roth & Willis [115].
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are also present on dung and mostly are absent from cave zones of
dry soil, stones, or pebbles [53,54].
The low diversity may be a consequence of a heterogenous diet
and/or low specialization of herbivorous animals of which
dinosaurs were the most abundant (suggesting there was relatively
little nutritional variability in their excrement and thus less need
for specialized roaches). Low specialization of at least some
dinosaurs is confirmed by phytoliths extracted from the Upper
Cretaceous coprolites (from dicotyledons, conifers, and palms)
from India, suggesting that the suspected dung producers
(titanosaur sauropods) fed indiscriminately on a wide range of
plants, including grasses [7]. With the diversification of mammals
[55], diverse specialized dung-beetles co-evolved [2] and these
cockroaches, possible with low specialization in their feeding
behaviors became extinct.
Generally, before the massive radiation of the Blattulidae at the
beginning of the Jurassic, their niche was occupied by the
superficially similar ‘‘Voltziablatta’’ group of cockroaches, which
became extremely rare along with the radiation of the Blattulidae.
In all Mesozoic sites, ‘‘Voltziablatta’’ and the Blattulidae occur in
congeneric species pairs, discretely differing in size, but not in
general appearance, thus doubtfully representing nocturnal and
diurnal cohorts (occurrence of both sexes in both groups was
validated earlier [51]). This enigmatic observation is unexplained
and needs further investigation. The Voltziablatta group phylogenetically
connects its descendants, the herein studied Blattulidae
and living cockroaches which bear endosymbionts; namely
termites, Sociala and Cryptocercus all descended from Liberiblattinidae.
If this mutualism had a single origin, it must have been in the
Voltziablatta group (fixed to flora and wood of Voltzia plants), where
the lignin consumption must have originally evolved. In the
opposite case, we would need to consider three independent
origins of endosymbionts, which molecular data do not support
[56].
Coprolite and Dung Decomposition
Presence of related endosymbionts in termites and cockroaches
of the family Cryptocercidae was postulated to be an evidence for
their direct relation. Nevertheless, the probable presence of
endosymbionts in the Mesozoic clade which diverged from stem
of higher cockroaches explains the monophyletic origin of these
symbionts in both groups also in the phylogenetic reconstructions
where they are not directly related [3]. The question is why was
this capability lost in most regular cockroaches?
The hypothesis that lignin-decomposing insect and their
endosymbionts originated via the consumption of wood predigested
by herbivore animal needs explanation. Feeding on
lignified wood and also foliage-eating became more widespread in
both dinosaurs and insects only with the radiation of angiosperms
at the Early Cretaceous/Late Cretaceous boundary [1]. Dung
consumption by Mesozoic termites, assisting in decomposition of
processed plant matter was already proposed [14].
Even the wood decay is preserved in a single sample, it is clear
that these cockroaches might have employed at least a semisocial
way of life to provide the horizontal endosymbiont transfer (thus
supporting the view that it evolved just once, as confirmed by the
phylogenetical scheme). In recent tropics, where food is available
for bats throughout the year, guano deposition is predictable and
also supports very large, persistent groups of cockroaches–
guanobies [57].
To summarize the arguments supporting dung processing, this
single sample is decisive in showing a coprolite still extruding from
the body (and thus belonging to the body fossil as a producer,
excluding incidental preservation) and containing modified wood
fibres with typical parenchymatous tangential ray cells. Lignin can
not be processed this way without endosymbionts and even in the
case it has been modified to some extent by some fungi, it must
have been pre-processed externally. The wood was apparently
processed before it entered the cockroach digestive tract as
indicated by the large extent of digestion apparent in cavities
(which definitely exclude the mechanical processing) and the
fragment preservation plotting fragment volume over the fragment
number–Fig. 2e; additional indirect support comes from dungprocessing
of living cockroaches, Tab. 1. It must be stressed, that
the extent of smoothing of large particles including large cavities
excludes the exclusively within insect processing and is evidence
for external pre-digestion. In this respect, a source of the wood
directly from the environment can be excluded. There are only
three possibilities for the pre-digestion, namely the fungal
(excluded below based on selective disadvantage of preference of
large indigestible particles and absence of wood damages before
the Late Cretaceous contrasting with plethora of coprolites
containing wood) and vertebrate pre-processing or their combination.
Large particles are numerous indicating that they were not
selectively avoided during consumption. Underrepresentation of
smaller particles was apparently due to biochemical digestion of
wood lignin as do their eusocial (extinct cockroaches of the family
Socialidae and termites) and semisocial (Cryptcercidae) descendants.
Although it is very probable that dinosaurs preferred wood
processed by fungi, fungi-only pre-digestion and feeding of these
cockroaches can be excluded based on the presence of large
fragments combined with low partition of wood. Such a small
amount would suggest selective feeding on fungi-modified wood,
in which circumstances large particles are contradictive; on the
contrary, unselective feeding on coprolites would contain the
expected spectrum of particles of diverse size. The only possible
explanation is that these were caused by herbivorous vertebrates.
Due to the dominance of these cockroaches for the same 200
million years as dinosaurs, no other vertebrate group is as
promising for this candidature. It can not be excluded that
cockroaches also cleaned up after some small, unknown vertebrate
herbivores, but these can be excluded from the present study as
small vertebrates can not digest wood.
Certainly, in such a case, in any solitary taxa the capability of
symbiont transfer and thus utilizing lignin was necessarily lost.
Termites did not exist before the Middle Jurassic, but their
precursors under study were apparently pre-adapted for wood
decomposition – and thus possessed one of the necessary
conditions for the origin of a eusocial way of life. Nevertheless,
termites were diversified in the very beginning of the Cretaceous as
evidence from the presently studied locality in Lebanon also
indicates [3,58,59].
Transfer of microflora within dinosaurs was proposed via
juvenile coprophagy [60], which facilitates microflora but also
endoparasite transfer with cockroaches. It is actually the intestinal
bacteria and metabolic by-products [61,62] of the herbivore gut
(perhaps dinosaurs), which likely allowed for lignin digestion in
Blattulidae (by protozoans). The small proportion of wood content
(,1% is of only partially processed wood remnants and up to 5%
of completely processed wood, not recognized in the ST) in the
cockroach coprolite indicates that wood was not the primary
constituent of the diet of the present individual, and rather
supports the derived source. This is also indicated by the Late
Cretaceous dung of herbivorous reptiles [63], probably dinosaurs
(entirely of comminuted plant tissue with the predominance of
secondary conifer xylem tissues of Cupressaceae). The unmodified
state of the cells and the absence of gymnospermous wood in dung
[64] is still problematic, but the small size of the plant fragments
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infilling the fossil burrows suggests comminution or sorting by
invertebrates [63]. Also several gymnosperms remains (Cheirolepidiacae
and Araucariacae) were found in the unstudied coprolite
(larger than the present ones) from the same deposit in Lebanese
amber.
The distance among parenchymatous tangential cells of the
wood in the present coprolite is roughly 10 mm, which is
comparable to the structure of wood of fossil Taxodioxylon
vanderburghii or Metasequoia glyptostroboides (20–30 mm [65]). Even
more similar parenchymatous tangential cells (10–20 mm) are
found in unidentified conifer wood from dinosaur coprolites (as
indicated in Fig. 5B, upper part of [8]). Interestingly, this wood
originates from trees growing in warm and semiarid Late
Cretaceous environments preserved in the sediments of the Two
Medicine Formation [8], which is in contrast to the warm and
humid amber-producing Early Cretaceous forest of Lebanon.
Anyway the specific determination of fossil conifer woods is very
difficult and requires comparisons of many features that do not
seem to be present in the small particles of wood in the fecal
pellets.
The wood (the length of the largest fragment was 13 cm)
preserved in dinosaur coprolites is characterized by absence of
cylindrical wood stems (no terminal twigs were digested); damage
to lignin such as the presence of pliant tracheids, uneven cell walls
and deformed and missing cells is also characteristic [8]. This,
along with the fact that the vertebrate gut cannot hold complex
lignolytic organisms, because these protists are anaerobic suggests
fungal decay prior to consumption [8]. On the contrary, the small
amount of small wood particles in these coprolites indicates they
were processed within dinosaurs and support decomposition of the
smallest particles both in dinosaur and cockroach coprolites.
Coprolite and Dung Decomposition-defecation
In spite of the diversity of behaviors reported from amber, a
review by Arillo [66] contains a single defecation, reported from a
Dominican amber termite [67]. Nevertheless, there is a rich
Cretaceous termite record of distinctive fecal pellets with
diagnostic hexagonal cross-sections that commence during the
Hauterivian or Barremian [68] and continue to occur in various
woods to the end of the Cretaceous. Some of these pellets may
have originated from individuals belonging to taxa such as the
eusocial cockroach Sociala that occurs in Mesozoic amber [3].
Fecal pellets from wood are known [69], and most amber
coprolites contain wood remains and are assigned to wood borers
among termites, beetles or some other insects [70,71]. Additional
pellets are known from the Dominican amber [72] and frass
containing fungi are known from Archingeay amber [73].
Defecation was probably often associated with escaping behaviour,
because more than 60 samples of Lebanese amber (coprolites are
often separated) contain coprolite of diverse size and shape (large
elongate, oval). Lots of them were preserved with wood fibers. In
the same piece there are insects like ceratopogonids, chironomids,
archizelmerids (extinct flies) and wasps, but these coprolites are not
associated with insects and are mentioned here to demonstrate the
common defecation behaviour, not the wood processing. No trace
fossils documenting specialized dung provisioning are known
before the Late Cretaceous [2].
General Ecology of Dung Provisioning
Detritic food chains strongly predominated in the Mesozoic [2]
and the dominance of the Blattulidae among cockroaches seems to
be associated with dung being the most valuable source of
nitrogen. It is improbable that there were specialized guilds of
dung feeders in the Mesozoic comparable with modern regarding
structural complexity and ecological efficiency: Sciaridae and
Scatopsidae (flies) with rapid larval development were remarkably
rare [74], as well as dung beetles, although both are present in the
Lebanese amber [75] along with decomposer flies of the families
Psychodidae and Sciaridae. However, they were absent before the
Jurassic and extremely rare during the entire Jurassic [10,64,75].
Alternative opportunistic exploiters of dinosaur dung were
snails. Multiple associations of 132 (with 0–66 specimens each)
fossils (Megomphix, Polygyrella, Hendersonia, Prograngerella, and three
aquatic taxa) have also been observed on or within 6 of the 15
herbivorous dinosaur coprolite deposits [15].
Despite the great diversity and quantity of scarabeid beetles in
the Mesozoic ([10] especially in the Middle Jurassic locality
Daohugou in Inner Mongolia, China), only a few species can be
considered as possibly coprophagous. Only 3 dung ball-makers
from the subfamily Scarabaeinae are known: Prionocephale deplanate
(Upper Cretaceous Lanxi formation, Zhejiang, China [76]),
Cretonitis copripes (Early Cretaceous Zaza Formation, Baissa, Russia)
and an undescribed species [72,77] of the living coprophagous
genus Trypocopris. Representatives of the Geotrupidae were
probably coprophagous: Parageotrupes incanus from the Yixian
Formation [78], and Cretogeotrupes convexus and Aphodius (s.l.)
(Aphodiinae) from Baissa [77,79].
An alternative hypothesis claiming mainly aquatic plant diet of
dinosaurs [80] and thus water defecation does not explain at least
some damage to terrestrial plants.
The dung of known Mesozoic herbivores is composed mainly of
undigested fern and gymnosperm tissues and was utilized by
opportunistic detritivores together with other plant litter [2].
The specialized coprophagy by beetles is recorded as late as the
Late Cretaceous when the diet of grazing dinosaurs apparently
contained angiosperms other than grasses and ecosystems were
based on biomes similar to grasslands [1]. Based on our
investigations, pollen and angiosperms in the Lebanon amber
are indicated by at least 5–6 different taxa.
The decay of wood pre-digested in dinosaur gastrointestinal
tracts explains and predicts the single origin of lignin consumption
in the common ancestor of termites, eusocial cockroaches (Sociala),
and semisocial cockroaches of the family Cryptocercidae. It would
also explain a huge number of termite-like fecal pellets (containing
wood) in Mesozoic ambers with parallel absence of any termite
damage to wood [68]. The fact that termites were a major lineage
responsible for the degradation of plant tissues (when compared
with cockroaches) is irrelevant in this respect as they originated not
earlier than in the Middle Jurassic when their ancestors, certain
Liberiblattinidae appear in the fossil record [4] and thus can not
play any role in the decomposition of early sauropod dung. In
contrast, blattulid cockroaches and their ecological equivalents
originated as early as the Permian–Triassic boundary.
The contemporary robust appearance of Cryptocercidae does
not require a major morphological shift from anticipated dungbeetle-habits.
It is likely that dung processors will also lose wings
like Cryptocercidae, but in caves, wing loss and associated
morphological changes occur more frequently in organisms that
rely on plant debris than those that rely on guano [81]. Under all
circumstances it is apparent, that termite and cryptocercid
ancestors were pre-adapted for lignin decay and, likely, provided
a limited sanitation to herbivorous reptiles. Based on the
correlation of distribution of reptiles and the dominance of the
blattulid cockroaches in Mesozoic ecosystems, and their coeval
occurrence in the present amber-bearing strata [82], these
herbivorous reptiles were most likely the dominant sauropod
dinosaurs.
Dinosaur Cleaner-Ups
PLOS ONE | www.plosone.org 8 December 2013 | Volume 8 | Issue 12 | e80560
Supporting Information
Figure S1 Synchrotron imaging of 5 coprolites of
dinosaur-age immature cockroach from the Lebanese
amber (Blattulidae 1094A-I). Select transparent mode for 3D
visualization and rotation.
(PDF)
Acknowledgments
We thank Georgy V. Nikolajev (Al-Farabi Kazakh National University,
Almaty, Kazakhstan) for the literature supply and fruitful discussion;
Vladimı´r Sˇimo and Adam Toma´sˇovy´ch (both GlU SAV, Bratislava,
Slovakia), Toma´sˇ Holu´bek and Marthe Kaufholz (both ANKA/IPS,
Eggenstein-Leopoldshafen, Germany) for technical help, Russell Garwood
(IC, London, UK), Ralf Kosma (SNHM, Niedersachsen, Germany) and
five anonymous reviewers for revision as well as Graeme Butler and Martin
Styan for linguistic revisions. The ANKA Synchrotron Radiation Facility is
acknowledged for providing beamtime.
Author Contributions
Conceived and designed the experiments: P. Vrsˇansky´ TV P. Vagovicˇ.
Performed the experiments: P. Vrsˇansky´ TV P. Vagovicˇ. Analyzed the
data: P. Vrsˇansky´ TV LV P. Vagovicˇ. Contributed reagents/materials/
analysis tools: P. Vrsˇansky´ TV DA AP P. Vagovicˇ. Wrote the paper: P.
Vrsˇansky´ TV DA AP LV P. Vagovicˇ. Initiated the research, identified and
drew the immature cockroach with coprolite, and erected and validated the
present hypotheses: P. Vrsˇansky´. Provided the material and dinosaur
information from the locality: DA. Provided and designed the mST
experiment, and provided particle measurements and volume segmentation
of reconstructed mST data and edited the text: TV P. Vagovicˇ.
Provided information on dung beetles and ecology: AP.
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Evidence from Amur River region of Russia.
Zootaxa 3635(2): 117-126.
Accepted by F. Crespo: 11 Feb. 2013; published: 26 Mar. 2013
ZOOTAXA
ISSN 1175-5326 (print edition)
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Zootaxa 3635 (2): 117–126
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Article
117
http://dx.doi.org/10.11646/zootaxa.3635.2.2
http://zoobank.org/urn:lsid:zoobank.org:pub:B74B0B78-911E-41FE-879F-7D41C8A77DBF
Paleocene origin of the cockroach families Blaberidae and Corydiidae:
Evidence from Amur River region of Russia
PETER VRŠANSKÝ1,2
, ĽUBOMÍR VIDLIČKA3,4
, PETER BARNA2
,
EUGENIA BUGDAEVA5
& VALENTINA MARKEVICH5
1
Arthropoda Laboratory of the Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117868 Moscow, Russia
2
Geological Institute, Slovak Academy of Sciences, Dúbravska cesta 9, P.O.BOX 106, 840 05 Bratislava, Slovakia.
E-mail: geolvrsa@savba.sk
3
Institute of Zoology, Slovak Academy of Sciences, Dúbravska cesta 9, 845 06 Bratislava, Slovakia. E-mail: uzaevidl@savba.sk;
4
Department of Biology, Faculty of Education, Comenius University, Moskovská 3, Bratislava, 813 34, Slovakia.
E-mail: vidlicka@fedu.uniba.sk
5
Institute of Biology and Soil Science, Far East Branch, Prosp. 100-letiya 159, 690022 Vladivostok, Russia.
E-mail: bugdaeva@ibss.dvo.ru
Abstract
Morphna paleo sp. n., the earliest winged representative of any living cockroach genus and the earliest representative of
the family Blaberidae, is described from the Danian Arkhara-Boguchan coal mine in the Amur River region (Russian Far
East). The branched Sc and A suggest Ectobiidae (=Blattellidae) probably is not the ancestral family because Blaberidae
were derived directly from the extinct family Mesoblattindae. The associated Danian locality Belaya Gora yielded
Ergaula stonebut sp. n., the earliest record of the family Corydiidae. Both species belong to genera codominant in the
Messel locality, thus validating their dominance in early Cenozoic assemblages.
Key words: fossil insects, fossil cockroaches, Tertiary, Blaberidae, Corydiidae, Morphna, Ergaula
Introduction
The Paleocene epoch, with 177 known extinct insect species: 44 coleopterans, 28 dipterans, 28 hemipterans, 27
hymenopterans, 15 odonates, 10 orthopterans, 8 neuropterans, 6 trichopterans, 5 mecopterans, 2 dermapterans, and
1 lepidopteran (EDNA fossil insect database; http://edna.palass-hosting.org; active 2.5. 2012) is the least known
Tertiary period in terms of insect diversity. No cockroaches, only two related termite species and a single
mantodean Arvernineura insignis have been described from Menat (Piton 1940). In contrast, 6124 Eocene, 2663
Miocene and 2550 Oligocene species have been recorded. Pliocene and Pleistocene species are also numerous, but
in EDNA underrepresented due to the presence of living species in these Epochs (EDNA catalogise only original
designations of species).
Cockroaches originated in the Bashkirian Carboniferous, with the oldest record originating from the
Quilianshan in China (Zhang et al. 2012, Guo et al. 2012). Typical Mesozoic families were derived from the
Phyloblattidae near the P/T boundary and the stem of the living families (but also the stem for all mantodeans and
termites) can be traced from the Mesozoic family Liberiblattinidae (Vršanský 2002, 2010, 2012). The earliest
record of any living family is the ectobiid (blattellid) Piniblattella vitimica (Vishniakova, 1964) from the earliest
Cretaceous (Vršanský 1997). Before this study, living cockroach genera, including highly advanced forms, were
known starting from the early Eocene (Archibald & Mathewes 2000) and the modern fauna is considered to
originate around the Paleocene-Eocene Thermal Maximum (PETM—Vršanský et al. 2011, 2012b). (The amber
fossil ?Blattella lengleti, is a nymph and may represent a separate genus.) The present study provides evidence for
the occurrence of at least some extant genera before the Palaeocene side of the PETM, and in parallel provides
earlier evidence for the two living families Corydiidae and Blaberidae.
VRŠANSKÝ et al.118 · Zootaxa 3635 (2) © 2013 Magnolia Press
Material and methods
Two joined forewings were collected by Yu.L. Bolotsky in the stratotype section of the Tsagayan Formation
(Belaya Gora locality). This specimen is deposited in the Amur Natural History Museum of the Institute of
Geology and Nature Management, Far Eastern Branch Russian Academy of Sciences (ANHM IGNM FEB RAS),
Blagoveshchensk, Russia. One isolated wing was collected by E.V. Bugdaeva in the upper plant-bearing bed in the
Arkhara-Boguchan coal mine. This specimen is deposited in Paleontological Institute of Russian Academy of
Sciences, Moscow, Russia. Comparative living material provided in the photograph originates from the collection
of Vít Kubáň (Thailand, Mae Hong Son Province, Soppong, 7–12.V.1996) deposited in collection of the second
author (ĽV) in the Institute of Zoology, SAS, Bratislava.
Geological background
The Zeya-Bureya Basin is located in the middle course of the Amur (Heilongjiang) River (Fig. 1B–D).
Development of its sedimentary cover began in the Late Cretaceous with accumulation of the Kundur Formation
(Santonian-Campanian). This stratigraphic unit is represented by sandstones, siltstones, and mudstones containing
abundant freshwater fauna. These sediments in some structures are oil-and-gas bearing. Over them is the Tsagayan
Formation, which is divided into three subformations. The lower Tsagayan Formation consists of conglomerates,
mainly clays with sandstone interbeds; its geological age is the early-middle Maastrichtian (Bugdaeva 2001;
Markevich 1994, 1995; Markevich et al. 2004, 2010, 2011). The late Maastrichtian part of the middle Tsagayan
Formation includes conglomerates, sandstones siltstone, and lenses with plant remains. The Danian upper part of
the Tsagayan Formation is represented by conglomerates, sandstones, clays and coal seams. The plant-bearing beds
occur here. The fossil plants have been studied since the 19th century (Heer 1878; Kryshtofovich & Baikovskaya
1966; Krassilov 1976). This flora was named Tsagayan flora.
The stratotype section of Tsagayan Fm is outcropped in the mouth of the Darmakan River, along the northern
and north-eastern slopes of Belaya Gora Mount. It is represented by conglomerates, sandstones, siltstones, and
mudstones. We obtained from each bed abundant fossil spores and pollen that allowed us to define the
Maastrichtian and Danian age of deposits and the position of the K-T boundary (Bugdaeva 2001). The clay with
bedded plant remains lies 37 m above that boundary (Fig. 1C); the thickness of this bed is 3 m. The following fossil
plants were collected: Podocarpus tsagajanicus Krassil., Taxodium olrikii (Heer) Brown, Metasequoia disticha
(Heer) Miki, Androvettia catenulata Bell, Potamageton cf. nordenskioldii Heer, Hydrocharis sp., Limnobiophyllum
scutatum (Dawson) Krassil., Trochodendroides arctica (Heer) Berry, Carinalaspermum bureicum Krassil. and
Nyssa bureica Krassil. The burial is dominated by leaves of Limnobiophyllum and shoots and leaves of Taxodium.
Other remains occur rarely. Remains of fossil insects have been found in this locality, including Buprestidae,
Chironomidae, as well as caddisworm cases of Folindusia cf. communita Cockerell, 1925 and Terrindusia minuta
Vialov et Sukacheva, 1976. The upper part of the upper Tsagayan Fm contains productive coal seams and several
coal mines. One of the mines (Arkhara-Boguchan coal mine) is located near Arkhara settlement. It has three plantbearing
beds with abundant fossil plants; the cockroach wing was found in the upper plant-bearing bed (Figs.
1B–C).
The family name Ectobiidae is used instead of Blattellidae and Corydiidae is substituted for Polyphagidae,
based on ICZN Ruling (see Beccaloni & Eggleton 2011; but with reservations of inclusion of termites within order
of cockroches).
Systematic palaeoentomology
Blattaria Latreille, 1810 (= Blattodea Brunner von Wattenwyl, 1882)
Blaberidae Brunner von Wattenwyl, 1865
Epilamprinae Princis, 1960
Zootaxa 3635 (2) © 2013 Magnolia Press · 119PALEOCENE COCKROACHES
Epilamprini Handlirsch, 1925
Morphna Shelford, 1910
= Morphnina Princis, 1958
Diagnosis (after Shelford 1910): Form rather dorsoventrally flattened. Vertex of head covered or almost covered
by pronotum, which is trapezoidal, sub-cucullate and posteriorly produced obtusely. Tegmina and wings fully
developed, exceeding the apex of the abdomen. Supra-anal lamina of typical Epilamprine shape. Cerci moderately
long. Femora moderately armed with spines beneath. Posterior metatarsus equal in length to succeeding joints; all
the joints entirely unarmed beneath, their pulvilli large, pulvillus of metatarsus apical but produced towards the
base of the joint.
Type species. Morphna maculata (Brunner von Wattenwyl, 1865).
Composition (updated from Princis 1967, 1971).
Morphna amplipennis (Walker, 1868) (India)
= Epilampra amplipennis Walker, 1868
Morphna auriculata (Brunner von Wattenwyl, 1865) (India)
= Epilampra auriculata Brunner von Wattenwyl, 1865
Morphna badia (Brunner von Wattenwyl, 1865) (Thailand, Malaysia, Sumatra, Java, Borneo)
= Epilampra badia Brunner von Wattenwyl, 1865
= Epilampra ramifera Walker, 1869
Morphna clypeata Anisyutkin & Gorochov, 2001 (Vietnam)
Morphna decolyi (Bolívar, 1897) (India)
= Molytria decolyi Bolívar, 1897
Morphna dotata (Walker, 1869) (Thailand, Malaysia, Borneo)
= Epilampra dotata Walker, 1869
Morphna humeralis Bruijning, 1948 (Sumatra)
Morphna imperatoria (Stål, 1877) (Philippines)
= Epilampra imperatoria Stål, 1877
Morphna maculata (Brunner von Wattenwyl, 1865) (Malaysia, Sumatra, Java, Borneo)
= Epilampra maculata Brunner von Wattenwyl, 1865
= Epilampra polyspila Walker, 1868
= Molytria shelfordi Kirby, 1903
Morphna moloch (Rehn, 1904) (Thailand)
= Epilampra moloch Rehn, 1904
Morphna plana (Brunner von Wattenwyl, 1865) (India, Sri Lanka)
= Epilampra plana Brunner von Wattenwyl, 1865
= Homalopteryx biplagiata Bolívar, 1897
= Epilampra punctifera Walker, 1868
= Homalopteryx templetoni Kirby, 1903
Morphna pustulata Hanitsch, 1930 (Sumatra)
Morphna sp. (Germany) extinct, Eocene (MES 10188)
Morphna paleo sp. n.
(Figs. 1A, 2C)
Holotype. PIN 5142/12. Right forewing fragment; type locality, Archara-Boguchan, Far East, Russia;
type horizon, Tsagayan Formation, Danian Paleocene.
Diagnosis. Forewing with length about 23 mm, width 9 mm. Numerous cross-veins present in M and CuA.
Anal intercalaries punctuated.
VRŠANSKÝ et al.120 · Zootaxa 3635 (2) © 2013 Magnolia Press
FIGURE 1. A) Morphna palaeo sp. n., holotype PIN 5142/12; Danian sediments (or Paleocene sediments) of ArcharaBogučan
in the Far East of Russia. Forewing length 23 mm; B) sections of Danian localities Belye Gory: Belaya Gora (1) and
Arkhara-Boguchan (2) (1—conglomerate; 2—sandstone; 3—siltstone; 4—claystone; 5—coal; 6—acid tuff; 7—strata with
cross-bedding; 8—locality of fossil flora; 10—locality of fossil insects); C) Profiles of Belaya Gora locality, the stratotype
section of Tsagayan Formation (1—conglomerate; 2—sandstone; 3—siltstone; 4—claystone; 5—coal; 6—acid tuff;
7—deposits of the Maastrichtian middle Tsagayan Formation; 8—deposits of the Danian upper Tsagayan Formation;
9—locality of fossil flora; 10—locality of fossil insects; D) Localization (Japan to the Right).
Zootaxa 3635 (2) © 2013 Magnolia Press · 121PALEOCENE COCKROACHES
FIGURE 2. A) Ergaula stonebut sp. n. Holotype ANHM 4/7; Danian sediments (or Paleocene sediments) of Belaya Gora, Far
East, Russia. Left forewing 31mm long. B) Males of Ergaula capucina, Thailand (Mae Hong Son prov., Soppong,
7.–12.V.1996, Vít Kubáň leg., coll. Ľ. Vidlička, ZIN SAS). Note significant folding line on right forewingand strong fold along
Sc, apparent also in fossil. C) Morphna palaeo sp. n. Holotype PIN 5142/12; Danian sediments (or Paleocene sediments) of
Archara-Bogučan in the Far East of Russia. Forewing length 23 mm.
VRŠANSKÝ et al.122 · Zootaxa 3635 (2) © 2013 Magnolia Press
Description. Forewing without coloration. Venation distinct with apparent intercalaries and rich cross-veins in
M and CuA. Subcostal area wide, with Sc richly branched (secondarily). R regular, parallel; M (5) slightly curved,
running close to R (apomorphy), fusing to CuA. CuA rich (8). Anal veins simple with punctuated intercalaries.
Remarks. The combination of parallel forewing margins, wide and branched Sc, fusion of M with CuA
running close to R, basalmost branches of CuA running parallel to CuP and simple A place this taxon in Morphna.
Morphna has been considered to be a comparatively terminal taxon of Epilamprinae (Rehn 1951).
Nevertheless, the new species points to a very initial stage of the evolution of Blaberidae, since compared with
Ectobiidae (=Blattellidae) fusion of M with CuA running close to R and wide, branched Sc are apomorphies. In the
living fauna, Morphna is restricted to southeast Asia (India, Sri Lanka, Malaysia, Sumatra, Java, Borneo,
Philippines and Thailand). The genus is quite diverse in species and some seem to have little in common (e.g., M.
pustulata is elongated, with curved forewing posterior margin). On the other hand, the most closely related living
species, M. plana (Brunner von Wattenwyll, 1865) from Sri Lanka, differs only in possessing numerous cross-veins
(plesiomorphy) and in size. All the living representatives of the genus are considerably larger than M. paleo sp. n.
(apomorphy), with forewing lengths of 41–50 mm.
Two basal branches of R have teratological fusion of veins (see Vršanský 2005: this particular parallel fusion
of two ascending R branches is unknown in fossils), but this character is without systematic value. Irregularity
between R and M is interpreted as an apomorphy based on the absence of this character in Cretaceous cockroaches.
Etymology. From Greek palaios: ancient or primitive.
Corydiidae Saussure, 1864 (= Polyphagidae Walker, 1868)
Corydiinae Saussure, 1864 (= Polyphaginae Walker, 1868)
Corydiini sensu Rehn, 1951
Diagnosis (after Rehn 1951). Both sexes with at least tegmina present, wings usually present, but sometimes
considerably reduced. Tegmina varying from normal to somewhat reduced, obovate and densely coriaceous
(mostly in females). Humeral area more developed than in Polyphagini, if coriaceous then broadly expanded. Sc
rami regular, not crowded. R without posterior branches, most branches terminating anteriorly, some apically,
instead of curving posteriorly. M with free base, its branching regular and direct. Cu not curving distinctly away
from plical furrow, CuP not joining cubitus.
Diagnosis (after Walker 1868). Female: Body short-elliptical, convex, dull, very thickly and minutely
punctured. Head shining, impressed between the eyes, with a transverse furrow near the mouth. Eyes not far apart.
Second joint of the palpi subclavate; third slightly securiform, very much longer than the second. Antennae
setaceous, submoniliform, not more than half the length of the body; first, second and third joints short; following
joints very short. Prothorax extending somewhat beyond the head and over the basal part of the fore wings when
they are expanded, rounded in front and on each side, slightly furrowed along each side; its breadth along the hind
border more than twice its length; hind border hardly rounded; hind angles slightly falcate; a lyre-shaped mark in
the disk. Mesothorax, metathorax, pectus and abdomen shining, mostly smooth. Abdomen with the segments above
and beneath near the tip retracted in the middle towards the disk; sides fringed, with bristles; subanal lamina small,
bilobed. Cerci lanceolate, submoniliform, setose. Legs stout; tibiae armed with some strong spines; first joint of the
tarsi twice the length of the fifth, which is very much longer than the second. Fore wings coriaceous, membranous
towards the border; costa much rounded; tips conical; principal veins distinct in the coriaceous part; transverse
sectors numerous, irregular. Hind wings membranous, strongly and thickly reticulated; transverse sectors
numerous, irregular.
Type species. Ergaula carunculigera (Gerstaecker, 1861)
Composition (updated from Princis 1963).
Ergaula Walker, 1868
= Dyscologamia Saussure, 1893 (type is cesticulata = pilosa)
= Parapolyphaga Chopard, 1929 (type is erectipilis = pilosa)
?= Netherea Vršanský et Anisyutkin, 2004 (type is haatica)
Zootaxa 3635 (2) © 2013 Magnolia Press · 123PALEOCENE COCKROACHES
Ergaula capensis (Saussure, 1893) (Nigeria, Cameroon, Democratic Republic of the Congo, Congo, Uganda,
Kenya, Tanzania, Zambia, Zimbabwe, Angola)
=Dyscologamia capensis Saussure, 1893
=Dyscologamia wollastoni Kirby, 1909
Ergaula capucina (Brunner von Wattenwyl, 1893) (Myanmar)
=Homoeogamia capucina Brunner von Wattenwyl, 1893
Ergaula carunculigera (Gerstaecker, 1861) (Philippines (Luzon))
=Corydia carunculigera Gerstaecker, 1861
=Ergaula scaraboides Walker, 1868
Ergaula funebris (Hanitsch, 1933) (Borneo)
=Dyscologamia funebris Hanitsch, 1933
Ergaula nepalensis (Saussure, 1893) (Nepal, Myanmar)
=Dyscologamia nepalensis Saussure, 1893
Ergaula pilosa (Walker, 1868) (Sumatra, Malaysia, Java, Borneo)
=Zetobora pilosa Walker, 1868
=Dyscologamia cesticulata Saussure, 1893
=Dyscologamia chopardi Hanitsch, 1923
=Parapolyphaga erectipilis Chopard, 1929
=Polyphaga sumatrensis Shelford, 1908
Ergaula silphoides (Walker, 1868) (Cambodia)
=Polyphaga silphoides Walker, 1868
Ergaula atica Vršanský et Anisyutkin, 2008 (Israel) extinct, ?Eocene (based on male)
?= Netherea haatica Vršanský et Anisyutkin, 2008 (Israel) extinct, ?Eocene (based on female)
Ergaula spp. (Germany) extinct, Eocene (common in Messel, based on both sexes)
Ergaula stonebut sp. n.
Holotype. IGNM FEB RAS ANHM 4/7. Both forewings; type locality, Archara-Boguchan, Belaya Gora locality,
stratotype of the Tsagayan Formation, Far East, Russia; type horizon, Tsagayan Formation, Danian.
Diagnosis. Forewing narrow, length/width: 31/11 mm, its venation reduced to approximately 50 veins at
margin. Sc branched broadly. Intercalaries distinct, coloration indistinct.
Description. Forewing fore margin slightly arcuate. Sc with both anterior (3 on left forewing, 1 on right
forewing) and posterior (4, 4) branches. R more or less regularly branched, with venation more dense towards
apex; veins secondarily branched (19, 16). M with secondary branches, curved posteriorly (11, 12). CuA largely
simplified, reduced to 3 branches at most. Anal veins sparse (6, 7).
Remarks. E. stonebut sp. n. differs from Therea Bilberg, 1820 (India) (the same tribe) in having costal space
comparatively narrow and Sc less expanded and with branches running more longitudinally, M and R reduced to
some extent and fused. Eucorydia Hebard, 1929 (SE Asia) and Miroblatta Shelford, 1906 (Borneo) have
exclusively straight stem of R (without any posterior branches), the latter comprises deviant forms with extremely
wide forewings, sometimes reduced to some extent. Homoeogamia Burmeiser, 1838 is limited to America (Mexico
and South America) today.
Ergaula stonebut sp. n. can be placed within Ergaula by simple exclusion and differs from its congeners only
in minor characters. It is generally very similar to E. atica from the sediments of Israel (presumably Eocene in age),
including the narrowness of forewings with distinct intercalaries and wide space between respective Sc branches (3
symplesiomorphies). E. atica also is very large, (forewing length 35 mm). The single preserved individual is
distinctly coloured and posses numerous deformations. Undescribed specimens from the Messel, Germany are also
very similar (H. Schmied, in preparation).
The E. stonebut sp. n. forewing is without deformities; it is narrower than in any living species. The type
species E. carunculigera differs in having a considerably smaller forewing (21–27 / 13.5 mm) (Gerstaecker 1861).
Ergaula. capucina differs in having all Sc venation dense; E. pilosa has dense Sc venation in the anterior region
only (Rehn 1951). Males of E. capensis are much larger (55–57 mm in total body length) (Hanitsch 1938).
VRŠANSKÝ et al.124 · Zootaxa 3635 (2) © 2013 Magnolia Press
The much smaller forewing of E. funebris (forewing length 22 mm: (Hanitsch 1933)) is monochromatic a
character likely shared by E. stonebut sp. n. However, the wing of E. funebris is much wider. Ergaula nepalensis is
unique in having discoidal veins straight and longitudinal (Saussure 1893), and E. silphoides, like most living
species, has a rounded fore-margin of the forewing (Walker 1868).
Some distinct characters revealed in the course of study of living E. capucina are seen in forewing of the
present fossil. The most distinct among them are asymmetrical sclerotisation (due to folding of wings over each
other) and invagination in the base of R, which represents the huge ventral ridge serving for folding of the hind
wing. Visible are also reticulations caused by sclerotisation in the costal area.
Etymology. stonebut is derived from some Slavic languages (means something).
Discussion
Based on study of terminal Mesozoic as well as Eocene cockroaches, it follows that most living cockroach genera
originated directly at or around PETM (Vršanský et al. 2011, 2012b). Warming not only expanded the geographical
range and the thermic optimum in more northern latitudes, but also produced conditions different from those
present in the original source area. Changes on land resulted in a higher evolutionary tempo as evidenced by
cockroaches (Vršanský 2011, 2012ab). Nevertheless, the present observations are direct evidence for the prePETM
origin of some of cockroach genera, which was unexpected. It is notable that both of the species described
herein belong to genera present (as codominants) in the Eocene Messel (47Ma) assemblage of Germany (Schmied
2009, unpublished observation), suggesting the characteristic Eurasian assemblage was already formed before the
Paleocene side of the PETM.
Ergaula occurs also in the presumably Eocene or Oligocene mangals of Israel (Anisyutkin et al. 2008) and a
leathery wing described as Netherea haatica Vršanský & Anisyutkin, 2008 seems to represent the smaller female
of living Ergaula—a common sexual dimorphism of this genus. This associations are likely very similar unless
identical in respect to generic content and support the Eocene stage for obscure (originally presumed to be
Mesozoic) locality in Israel.
Different were some Eocene North American localities, where predominantly smaller species were preserved
(Vršanský et al. 2011a, 2012).
Very little can be learned from the geography of the two specimens. Ergaula currently is widely distributed in
Africa and Asia, but apparently was also present also in Europe in PETM, but absent in the Americas during the
Eocene. Morphna has a similar wide pattern in Asia (absent in Africa), with occurrence in Europe during PETM.
The important aspect of deformed wings is ambiguous in this respect. While no wing deformation (developmental
change modifying wing geometry, most often fusion of veins or irregularity), is reported in hundreds of Eocene
individuals from the Green River in Colorado, very few are present in Early Miocene localities. Deformations are
common and abundant in more recent fossils and in living cockroaches. The single specimen of Morphna from the
Paleocene possesses at least one such deformity, which may be stochastic.
Morphna is peculiar also in another respect in that it is not only the earliest occurrence of any living genus, but
also the first occurrence of the family Blaberidae. It is possible that the original blaberid genera, representing the
most advanced cockroaches of the time, survived with minor modifications to the present. In any case, the traces of
plesiomorphies are valuable: branched A, branched Sc and punctuated intercalaries are all characteristics of
Mesoblattinidae, and were lost in the initial stage of the evolution of the family Ectobiidae (=Blattellidae).
Therefore, it seems likely that Blaberidae originated directly from the extinct Mesoblattinidae, and not from
Blattellidae as has been generally accepted (see Djernaes et al. 2012). These results do not contradict with living
material-based analyses, as Blattellidae are direct descendants of the Mesoblattinidae.
Acknowledgements
We thank Yurij Bolotskij for collecting the material, Prof. Alexandr P. Rasnitsyn (PIN, Moscow) for initiating this
study, Prof. Ernest Bernard (UTK) for thorough editing, Prof. Sonia M. Fraga Lopes (Museu Nacional, UFRJ) and
an anonymous reviewer for revision of the manuscript. Research supported by UNESCO/AMBA (MVTS), VEGA
125, Literary fund, Presidium of Russian Academy of Sciences (grant 12–I–P28–01) and Russian Foundation for
Zootaxa 3635 (2) © 2013 Magnolia Press · 125PALEOCENE COCKROACHES
Basic Research (grant 12–04–01335), Slovak Research and Development Agency through financial support no.
APVV-0213-10.
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Príloha č. 16
VRŠANSKÝ, P., ORUŽINSKÝ, R., BARNA, P., VIDLIČKA, Ľ. & LABANDEIRA, C.C. 2014.
Native Ectobius (Blattaria: Ectobiidae) from the Early Eocene Green River Formation
of Colorado and Its Reintroduction to North America 49 Million Years Later.
Annals of the Entomological Society of America 107(1): 28-36.
SYSTEMATICS
Native Ectobius (Blattaria: Ectobiidae) From the Early Eocene Green
River Formation of Colorado and Its Reintroduction to North America
49 Million Years Later
P. VRSˇANSKY´ ,1,2
R. ORUZˇ INSKY´ ,3
P. BARNA,1
L’. VIDLICˇ KA,4
AND C. C. LABANDEIRA5,6,7
Ann. Entomol. Soc. Am. 107(1): 28Ð36 (2014); DOI: http://dx.doi.org/10.1603/AN13042
ABSTRACT Ectobius kohlsi sp. n. and three undetermined species of the common Eurasian cockroach
genus Ectobius Stephens, 1835 are reported from the lower middle Eocene of North America.
This species indicates a cosmopolitan distribution of the genus during the mid Paleogene, and supports
its current relict distribution in modern north-temperate and African ecosystems. When compared
with the living species, E. kohlsi was either neutral or plesiomorphic in all characters, but exhibited
a close relationship to the extant Ectobius kraussianus Ramme, 1923 Species Group in the identical
structure of the pronotum. E. kohlsi also was similar to extant Ectobius ticinus Bohn, 2004, in the
character of its wing venation (see Bohn 2004), in particular the forewing vein M, and to extant
Ectobius vittiventris (Costa 1847) in details of forewing coloration. These latter two species are
members of the Ectobius sylvestris Species Group (Bohn 1989). Ectobius balticus Germar et Berendt,
1856Ña conspicuously dominant cockroach from mid-Eocene Baltic amberÑalso appears plesiomorphic
in all characters despite being a few million years younger than E. kohlsi. One reason for the
complete disappearance of this dominant genus from North America is the peculiar consequence that,
after 49 million years, a cool-adapted Ectobius lapponicus (L.) was capable of being reintroduced to
a signiÞcantly cooler North America than that its antecedents which inhabited North America during
a warmer European Eocene. Modern E. lapponicus is synanthropic in North America, even though no
synanthropism is recorded for this species in its native habitat throughout Europe.
KEY WORDS cockroach, Europe, fossil insect, paleoclimate, relict distribution
The global cockroach fauna can be differentiated historically
into three distinct phases that collectively
form deep and varied evolutionary trajectories. These
phases are the late Paleozoic, the Mesozoic, and the
Cenozoic; the latter, or modern fauna, is overwhelmingly
composed of extant taxa. The late Paleozoic and
Mesozoic faunas are characterized by oviposition of
isolated eggs through use of an external ovipositor,
which was long and prominent in Paleozoic forms and
comparatively short to blunt in Mesozoic taxa, with
eggsdepositedinaconglomeraticfashion.Bycontrast,
the Ectobiidae (ϭBlattellidae), originating near JurassicϪCretaceous
boundary, and possibly some of
their predecessors, the Mesoblattinidae (Vrsˇansky´
1997, Wei and Ren 2013), in addition to the entire
modern cockroach fauna, are characterized by taxa
that lay their eggs within a hardened egg-case, the
oo¨theca.MostMesozoicfamilies,suchasthedominant
Blattulidae and Caloblattinidae, disappeared before
the KϪPg boundary at the end of the Cretaceous,
which responded to a major ecological crisis, at least
in North America (Labandeira et al. 2002).
Extant genera of the modern cockroach fauna
evolved during the beginning of the Paleogene Period
(Vrsˇansky´ 2002; Vrsˇansky´ et al. 2002, 2011, 2012, 2013),
and after the KϪPg event. The early Paleogene fauna
consisted of transitional taxa, predating the origin of
most exclusively modern families, but also consisting
of uniquely ancestral, extinct, and primitive genera.
During the mid-Eocene, rare Mesozoic relicts such as
the Mesoblattinidae and Skokidae persisted, although
at that time most other taxa were afÞliated modern
families and genera. It was during the latest early
Eocene that the genus Ectobius appears in the Palearctic
fossil record of Baltic amber (Statz 1939) as a
dominant species. As it now turns out, this also is the
time interval of the earliest record of the genus in the
Nearctic.
1 Geological Institute, Slovak Academy of Sciences, Du´bravska c. 9,
P.O. Box 106, 840 05 Bratislava, Slovakia.
2 Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya
123, 117868, Moscow, Russia.
3 Geological Institute, Slovak Academy of Sciences, Dˇ umbierska 1,
974 01 Banska´ Bystrica, Slovakia.
4 Institute of Zoology, Slovak Academy of Sciences, Du´bravska c. 9,
845 06 Bratislava, Slovakia.
5 Department of Paleobiology, MRC-121, P.O. Box 37012, National
Museum of Natural History, Smithsonian Institution, Washington, DC
20013-7012.
6 Department of Entomology and BEES Program, University of
Maryland, College Park, MD 20742.
7 Corresponding author, e-mail: labandec@si.edu.
Materials and Methods
Our study accessed specimens from the Parachute
Creek Member of the Green River Formation, in the
Piceance Creek Basin, in GarÞeld County, northwestern
Colorado, Ϸ20 km west of Riße, and in cliffs
overlooking the Colorado River. The material originates
from the upper portion of the Parachute Creek
Member, a regionally ponded lacustrine deposit characterized
by a distinctive organic-rich “mahogany
zone” that is 20Ð60 m thick, which extends throughout
theBasinandservesasastratigraphicreferencedatum
at Ϸ48.8 million years old (Ma), to which insectbearing
subjacent localities can be compared (Smith
et al. 2010). The Parachute Creek Member consists of
evaporates and especially laminated, organic-rich, often
kerogenous, shales, and Þne-grained siltstones
(Hail 1992). A recent study (Smith et al. 2010) analyzing
the “Curly tuff” immediately below the mahogany
zone yielded a 40
Ar/39
Ar age date with a weighted
meanvalueof49.02Ϯ0.30MawithaϮ2␴errormargin
(Smith et al. 2010). These results suggest a date of 49 Ma
for the collective insect deposits mentioned herein,
which would vary minimally, attributable to their subtle
variation in stratigraphic position from the 49 Ma data.
This date is equivalent to the latest Ypresian Stage of the
early Eocene Epoch (Gradstein et al. 2012).
Although insect fossils were frequently revealed
along splits in the bedding planes, additional preparation
of occluding matrix by thin needle-like picks
was necessary in most instances for further preparation.
Prepared specimens were photographed with an
Olympus SZH stereomicroscope attached to an Olympus
5060 camedia camera with 5.1 megapixel storage
capability. Lighting was provided by a Þber-optic illuminator
with plane polarized light and a ßuorescentstyle
ring light to occasionally capture cuticular details.
Images of specimens were taken with and
without ethanol immersion, the latter to enhance
specimen contrast with the surrounding rock matrix.
Characters were polarized based on the genus Symploce
Hebard, 1916, which is one of the basalmost and
primitive living representatives of the family Ectobiidae
(Vrsˇansky´ 1997, Klass 1997, Vrsˇansky´ et al. 2011).
Venational nomenclature follows Comstock and
Needham (1898); Vrsˇansky´ (1997); Rasnitsyn (2002);
and Vrsˇansky´ et al. (2012, 2013).
Results
Blattaria Latreille, 1810 (‫؍‬Blattida Latreille,
1810 ‫؍‬ Blattodea Brunner von Wattenwyl, 1882)
Ectobiidae Brunner von Wattenwyl, 1865
Ectobius Stephens, 1835
Type Species. Blatta lapponica L., 1758 (ϭBlatta
lapponicus lapponica).
Ectobius kohlsi Vrsˇansky´, Vidlicˇka et Labandeira,
sp. nov.
(Figs. 1Ð3)
HOLOTYPE. 41679/53274; a complete female. Deposited
in the Department of Paleobiology, National
Museum of Natural History (NMNH), Smithsonian
Institution, Washington, DC.
Type Horizon. Green River Formation, lower middle
Eocene, equivalent to the latest early Eocene
(Gradstein et al. 2012).
Type Locality. Piceance Creek Basin, Ϸ20 km west
of Riße, northwestern Colorado.
Additional Material. 41075/27701; 25971ϩ-, 31499;
41228; 41087/105061ϩ-; 41088/25457; 41093/75726;
41142/55491;57239;57545;58123;41222/27865;41225/
30376; 41227/137007ϩ-; 41236/139337; 41237/147735;
41679/53476, 52833; 41822/53843; SAV97/147911. The
general locality for the additional material is the same
as the type, although specimens were taken from several
stratigraphically adjacent horizons.
Differential Diagnosis. A new species most closely
resembling Ectobius bruneri Seoane, 1879, superÞcially
with partially punctate pronotum, identical coloration,
and three, characteristically large maculae (a plesiomorphy).
It differs in M fused with the R (an apomorphy).
Same venation in E. kohlsi occurs in Ectobius ticinus
Bohn, 2004, possessing a separate and distinct M vein (a
plesiomorphy).
Description. Head coloration as in Figs. 1 and 2. Antennae
delicate; long, at least 6.5 mm; and 0.7 mm wide.
Pronotum transverse; width 1.3 mm, length 1.9 mm,
nearly ovoid; distinctively colored (Fig. 2). Forewing
(length5.9Ð6.0infemales,and6.5inmales),withRvein
expanded, reaching the proximal part of apex. M vein
separatedfromRapically;reducedtoasimplevein.CuA
vein expanded and fused directly to main R branch.
Coloration with characteristic dark dots and four, large,
dark maculae; apex veins pale, with dark margins. Hindwing
angulate, tip somewhat rounded and colored in
posterior aspect. Body width 2.5Ð3.0 mm.
Comparisons. The present species is assigned to the
extant, exclusively Eurasian and African genus, Ectobius,
based on characteristic forewing venation, including
coloration (Figs. 1Ð3). The specimen signiÞcantly
differs from the extant subgenus Ectobiola, such
as Ectobius duskei Adelung, 1904, which has numerous
dark spots on the forewing, the absence of punctae on
the pronotum, and possession of a characteristic
horseshoe-like pattern of coloration on the forewing.
Females bear reduced wings. Owing to a zone of distinct
coloration, Ectobiola is linked to the Ectobius lapponicus
(L.) Species Group by an autapomorphy consisting of a
sharp, pronotal posterior margin; dark maculae investing
the R vein; and slender punctation expanded to the
entire surface of the wing. We suggest that Ectobiola is
distantly related to the extant species, Ectobius tuscus
Galvagni, 1978, which has a pronotum with sparse punctation,
short forewings, and an odd, reduced venation
with large, numerous maculae, perhaps indicating that
this latter taxon should be referred to a separate derived
species group not closely related to the present species.
January 2014 VRSˇANSKY´ ET AL.: EOCENE Ectobius AND ITS REINTRODUCTION 29
Fig. 1. E. kohlsi Vrsˇansky´, Vidlicˇka et Labandeira sp. n. (a) HOLOTYPE. 41679/53274; negative relief; complete female
specimen. (b) 41142/57545; positive relief; complete male specimen. (c1Ð2) 41142/57239; positive relief, complete female
specimen and detail of terminalia. (d) 41075/27701 (negative, sex unknown). (e) 41679/53274; ?positive relief, probable
female. (f) 41142/58123; detail of female terminalia. All specimens are deposited at the National Museum of Natural
History (NMNH), Washington, DC. C2, d, and f photographed under alcohol immersion. The material is from the Green
River Formation, of latest early Eocene age; Piceance Creek Basin, Northwestern Colorado. Scale bars represent
1 mm.
30 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 1
Another extant apomorphic species is Ectobius semenovi
Bei-Benko, 1935, bearing an extremely enlarged
pronotum without punctuation and monochromatic
forewings. Another aberrant species is Ectobius
tadzhicus Bei-Bienko, 1935, with pronotal coloration
and extremely shortened wings similar to that of EcFig.
2. aÐg, E. kohlsi Vrsˇansky´, Vidlicˇka et Labandeira sp. nov. (a) 41075/27701; positive relief; detail of head and pronotum.
(b) 41142/57545; positive relief; male specimen with pronotum. (c) HOLOTYPE. 41679/53274; negative relief, female specimen
with pronotum. (d1Ð2) 41093/75726; positive relief; female specimen. (e1Ð2) 41087/105061; positive relief; male specimen. (f)
41075/25971; positive relief; female specimen. (g) 41088/25457; negative relief, female specimen, with detail of terminalia. All
specimens deposited in the NMNH, Washington, DC, Green River Formation, latest early Eocene. All specimens photographed
while immersed under alcohol, and originate from Green River Formation, northwestern Colorado. Scale bars represent 1 mm.
January 2014 VRSˇANSKY´ ET AL.: EOCENE Ectobius AND ITS REINTRODUCTION 31
tobiola duskei. The Subgenus Capraiellus of the Ectobius
panzeri Stephens 1835 Species Group is similar to
the present species, possessing punctae, pronotal
darkening, a horseshoe-like pattern of coloration and
short female forewings, similar to Ectobiola. The Ectobius
corsorum Ramme, 1923 Species Group, is a more
derived lineage that has a punctate pronotum with a
sharply curved posterior margin, and spots on fore-
wingsthatareplacedmorebasallythanothertaxa.The
Ectobius minutus Failla et Messina, 1977, Species
Group is related to Ectobius montanus (Costa 1866),
the Ectobius sylvestris Species Group (Bohn 1989) and
Ectobius friesanus Princis, 1963, all of which have sim-
ilarglands,butdifferinsensillardistribution,theshape
of the cerci and the presence of a boss at the center
of glandular pits. E. friesanus may deserve placement
in a separate species group but is not directly related
to the present species.
Instead, E. kohlsi possesses a combination of characters
that occur in two lineages. The Þrst lineage is
the Ectobius kraussianus Ramme 1923 Species Group,
such as Ectobius lagrecai Failla et Messina, 1981, and
Ectobius aetneus Ramme, 1927, whose venation differs
in having a punctate forewing radial area. The pronotum
in this species group has characteristically
dense punctae, and two, symmetrical, arcuate ridges
identical in some individuals to those found in the
present species. The second lineage is the E. sylvestris
(Poda 1761) Species Group that lacks pronotal punctae,
although the forewing venation (Fig. 4) is nearly
identical with the new species (Fig. 1). E. sylvestris
females differ in having shortened wings, and in the
form and coloration of the pronotum. The most similar
character to the present species is venation, also occurring
in E. ticinus, with a separate and distinct M
vein (a plesiomorphy). E. sylvestris also is similar in
pronotal ridge features, but strongly differs by having
brachypterous females (an apomorphy; Fig. 4). E. bruneri
has a partially punctate pronotum, but the M is
fused with the R (an apomorphy). Coloration is identical
to the present species, bearing three, characteristically
large maculae (a plesiomorphy), but with an
unseparated M (an apomorphy). In E. vittiventris
(Costa 1847, Baur et al. 2004), the females have distinct,
long wings (a plesiomorphy; Figs. 4 and 5). By
contrast, extinct E. glabelus Statz, 1939, from the Oligocene
of Rott-am-Siebengebirge, differs in having
rather angulate forewings. Ectobius balticus Germar et
Berendt, 1856, from middle Eocene Baltic amber, possesses
all characters, including pronotal morphology
and coloration, which resembles the most basal Ectobiidae
(Fig. 6) (Germar and Berendt 1856).
Undescribed species, presumably assignable to Ectobius,
also are present in middle Eocene strata of
Messel, in central Germany (Schmied 2009), of age
slightly younger than Ectobius mentioned in this re-
port.
Remarks. An investigation of a single population of
living males of E. sylvestris in Slovakia revealed female
brachyptery (Fig. 4), a broad variability of forewing
area (CoefÞcient of Variation ϭ 13.36%, n ϭ 41), a
distinctive venational pattern (Fig. 4), and a distincFig.
3. E. kohlsi Vrsˇansky´, Vidlicˇka et Labandeira n. sp. HOLOTYPE. 41679/53274; A complete female. The repository
is the NMNH, Washington, DC, Green River Formation, latest early Eocene, Piceance Creek Basin, northwestern Colorado.
Dots correspond to dorsal sensilla. Negative compression; the right forewing is outstretched; length 5.9 mm.
32 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 1
tive pattern of sensillar distribution on both surfaces
of fore- and hind wings (Fig. 4) (Quercetum Mtns. of
Jur, Slovakia, collected in 2008 by O. Majzlan and LÕ.
Vidlicˇka). Another analysis revealed the presence of
dorsal mechanoreceptors on minute dark maculae,
which correspond to dark dots preserved on the forewing
surface of the new species (Fig. 3). Surprisingly,
except for microscopic openings (punctae) on the
ventral side, there are no macroscopic sensilla on posterior
three-quarters of the wings.
Etymology. The species designation honors David
Kohls, a collector of considerable Green River fossil
insect material deposited at the National Museum of
Natural History, in Washington, DC.
Character of Preservation. Twenty-one complete
specimens; Þve with wings, and one laterally pre-
served.
Discussion
LivingspeciesofthegenusEctobiusappearasrelicts
of a richer, ancient fauna that are closely related to
European Oligocene species. Currently, only E. sylvestris
and E. lapponicus are widely distributed
throughout the forested Palearctic Region (Zherikhin
1970). However, during the Eocene, the genus Ectobius
apparently was widely distributed in the Palearctic
as well as in the Nearctic. In North America, it
became extinct and typically was not replaced by
othercockroachtaxa.WithintheUnitedStates,similar
cool-temperate habitats were colonized within the
past several decades by Parcoblata cockroaches (12
species), preferring forests under bark. Other Ectobius
species have been found in association with herbaceous
vegetation, such as tansy, Tanacetum vulgare L.
(Asteraceae), in coastal New Hampshire, with males
predominating in a sex ratio of Ϸ2:1. Adults of this
species were seen at or near the crowns of various
plants and on ßowers or leaves, but when disturbed,
quickly dropped to the ground (Chandler 1992).
Fig. 4. (aÐk) Modern forewings of one population of males of E. sylvestris males, occurring at Jur, near Bratislava, Slovakia.
Collection of the Zoological Institute SAS, Bratislava. Specimens Es01/1Ð26/2.
Fig. 5. Left and right forewing of the sensillar system of
a modern male of E. sylvestris, from Jur, near Bratislava,
Slovakia. The collection is from the Zoological Institute SAS,
Bratislava. Specimen Es1Ð2. Black dots represent ventral sensillae;
red dots are dorsal structures providing for the presence
of dorsal sensilla exclusively on dark dots. The right
wing is missing dorsal sensilla in area overlapped by the left
wing.
January 2014 VRSˇANSKY´ ET AL.: EOCENE Ectobius AND ITS REINTRODUCTION 33
Nielsen (1987) found individuals on ßowers of wild
raspberries, Rubus idaeus L. (Rosaceae). By contrast,
in Europe, nymphs and adult males of the “dusky
cockroach” are found on low lying vegetation, with
adult females commonly found on the ground in leaf
litter (Roth and Willis 1960).
The reintroduction of Ectobius into the United
Statesafter49millionyearsispeculiar.E.sylvestriswas
established in the northeast United States (Hoebeke
and Nickle 1981); E. pallidus also was established in
the Northeast and, in addition, the Midwest; and E.
lapponicus as well became a denizen of the Northeast
(Atkinson et al. 1991, Chandler 1985, 1992). In particular,
E. pallidus appears to have been established in
Massachusetts sometime in 1951, and it occurs in
houses and other domestic effects surrounding human
habitation (Helfer 1987). In 1984 the “dusky cockroach,”
E. lapponicus, was found in southeastern New
Hampshire by Chandler (1985), the Þrst North American
record of this European immigrant. A subsequent
collection in eastern Vermont was noted by Nielsen
(1987), and this species has now been collected repeatedly
in coastal and central New Hampshire
(Chandler 1992). Ectobius lucidus was reported recently
from the eastern United States (Hoebeke and
Carter 2010). Interestingly, these reintroduced species
in the United States are associated with human
habitation and thus possess synanthropic behaviors.
By contrast, no synanthropism was recorded for the
same species throughout their native European habitats,
and only rare cases of building infestation are
known from Europe (Mielke 2000).
During the latest middle Eocene, at least four distinct
species that signiÞcantly differed in size were
present at the various Green River localities of Colorado.
Of these species, three were poorly preserved
and larger than E. kohlsi, but remain undescribed.
(These specimens are represented by preliminary
specimen numbers 41088/26584, 41221(2)/27865, and
41221/87663.) However, it is impossible to conclude
whether these additional species occurred contemporaneously
with E. kohlsi or otherwise were allopat-
ric.ThereasonsforthedisappearanceofEctobiusfrom
theNorthAmericancontinentremainobscure.Atemperature
change is known to be responsible for a
similar pattern of extinction and after human-based
reintroduction in some aquatic organisms (Strasser
1998). Cooling could be responsible for the extinctions
of diverse Ectobius taxa in North and Central
America, as the thermophilic representatives of the
genus have specialist associations and have a high
degree of endemism in contrast to more northerly,
widely distributed European generalists (Vidlicˇka
and Szira´ki 1997, Bohn 2004, Scholczova´ 2013). Nevertheless,
the current distribution of Ectobius from
northernmostEuropeextendssouthwardtothesouthernmost
Africa, with a latitudinal biogeographical discontinuity
along an equatorial belt (Bei-Benko 1950),
suggesting that a warming temperature change would
enable survival of the genus in the southern North and
Central America in the near future.
In the late early Eocene Green River ecosystem, E.
kohlsi had a similar wing size for both sexes (a plesiomorphy),
with females of slightly smaller size than
males, the difference being Ͻ1 mm. Representation of
both sexes is roughly equal in the fossil population,
indicating similar ßight activity. This condition contrasts
to living representatives of the genus, wherein
Fig. 6. E. balticus Germar et Berendt, 1856, from the early middle Eocene Baltic amber. Gusakov collection no. VI-008.
Original image courtesy of D. S. Shcherbakov.
34 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 1
the sex ratio is variable but different from equality.
Such a nonequilibrial sex ratio perhaps is associated
with conspeciÞc males involved in active ßight.
Acknowledgments
We thank David Kohls for collecting the specimens described
herein. We are grateful to the staff of the Paleontological
Institute in Moscow, Russia, particularly A. P. Rasnitsyn,
who gave fruitful advice; A. Gusakov loaned amber
material; and D. S. Shcherbakov provided technical help and
a revision of the manuscript. Finnegan Marsh assisted in
tracking down locality information and Þnal sharpening of
images. Heiko Schmied and Adam Toma´sˇovy´ch provided
advice and an entry into the literature. This research was
supported by Amba Projects of the United Nations Educational,
ScientiÞc and Cultural Organization and its Medzina´rodna´
Vedecko-Technicka´ Spolupra´ca (MTVS) supporting
program in Slovakia. A Short-Term Visitor Award from
the National Museum of Natural History, Washington, DC; a
Vedecka´ Grantova´ Agentu´ra (VEGA) grant from 6002,
2/0125/09, and 2/0186/13 funds; an Agentu´ra pre podporu
Vedy a Vy´skumu (APVV) grant from the 0436-12 fund, and
support from the Literary Fund provided resources for completion
of this project. This is contribution 268 from the
Evolution of Terrestrial Ecosystems Consortium at the National
Museum of Natural History, in Washington, DC.
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36 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 107, no. 1
Príloha č. 17
GREENWALT, D.E., VIDLIČKA, Ľ. 2015.
Latiblattella avita sp. nov. (Blattaria: Ectobiidae)
from the Eocene Kishenehn Formation, Montana, USA.
Palaeontologia Electronica 18.1.16A: 1-9.
Palaeontologia Electronica
palaeo-electronica.org
http://zoobank.org/11628551-F184-435E-8B57-748EB630B6C7
PE Article Number: 18.1.16A
Copyright: Palaeontological Association April 2015
Submission: 24 September 2014. Acceptance: 5 March 2015
Greenwalt, Dale E. and Vidlička, Ľubomír. 2015. Latiblattella avita sp. nov. (Blattaria: Ectobiidae) from the Eocene Kishenehn
Formation, Montana, USA. Palaeontologia Electronica 18.1.16A: 1-9.
palaeo-electronica.org/content/2015/1118-new-eocene-cockroach
Latiblattella avita sp. nov. (Blattaria: Ectobiidae) from
the Eocene Kishenehn Formation, Montana, USA
Dale E. Greenwalt and Ľubomír Vidlička
ABSTRACT
Latiblattella avita Greenwalt and Vidlička, 2015, sp. nov., and the first fossil of the
genus, is described. The discovery of a fossil representative of this genus suggests
that Latiblattella was more widely distributed in the Eocene. The Eocene American
cockroach fauna is mostly comprised of what are today, cosmopolitan genera while the
extant genus Latiblattella Hebard, 1917 is restricted in its geographical distribution to
Central America, Mexico, Florida and Arizona. The discovery of Latiblattella avita, in
combination with the recent description of Cariblattoides labandeirai Vršansky et al.,
2012, also documents the presence of rather derived representatives of the family
Ectobiidae as early as the Middle Eocene.
Dale E. Greenwalt. Department of Paleobiology, NMNH, Smithsonian Institution, Washington, District of
Columbia, U.S.A. 20013-7012, GreenwaltD@si.edu.
Ľubomír Vidlička. Institute of Zoology, Slovak Academy of Sciences, Dúbravská cesta 9, 845 06 Bratislava,
Slovakia, lubomir.vidlicka@savba.sk
and Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská dolina, Bratislava,
811 04, Slovakia
Keywords: Fossil insect; Cenozoic cockroaches; Pseudophyllodromiinae; new species
INTRODUCTION
Cockroaches form one of the taxonomically
smaller insect orders with only about 5,000 living
species, the majority of which are found in tropical
forests (Vidlička, 2001; Vršanský et al., 2002; Roth,
2003; Grimaldi and Engel, 2005; Beccaloni and
Eggleton, 2013). This contrasts with the high diversity
of ecological and behavioral niches occupied
by this clade, as exemplified by the recent discoveries
of eusocial, jumping, aquatic, extinct predatory,
pollinating, troglobitic and luminescent
species (Vishniakova, 1973; Zompro and
Fritzsche, 1999; Bohn et al., 2010; Vršanský, 2007;
2010; Vršanský and Chorvat, 2013 – but see
Greven and Zwanzig, 2013). Extinct species number
over 1,000 but nearly 80% of these are Paleozoic
“roachoids” (Mitchell, 2013). The modern
cockroach fauna is thought to have evolved immediately
after the Cretaceous-Paleogene boundary
(Vršanský et al., 2002, 2011, 2012, 2013) although
only 53 fossil species have been described from
the Cenozoic era (Mitchell, 2013; Arillo and Ortuño,
GREENWALT & VIDLIČKA: NEW EOCENE COCKROACH
2
2005; Solórzano Kraemer, 2007; Gorochov, 2007).
Undescribed material includes that presently studied
from the Green River (e.g., the genera Blattella
Caudell, 1903, Namablatta Rehn, 1937, Diploptera
Saussure, 1864, Sigmella Hebard, 1940 and Symploce
Hebard, 1916) in Colorado (Vršanský et al.,
2011, 2012).
The most speciose family within Blattaria is
Ectobiidae (= Blattellidae) with approximately
2,400 species in about 220 genera (Beccaloni and
Eggleton, 2013). Ectobius Stephens, 1835 and
Phyllodromica Fieber, 1853, two genera in the subfamily
Ectobiinae, provide most of the extant cockroach
diversity in the Palaearctic (Bohn et al.,
2013). Ectobiidae, which originated from the family
Mesoblattindae, first appeared in the Early Cretaceous
and became dominant during the Cenozoic
(Vršanský, 1997; 1999; 2002; Anisyutkin et al.,
2008; Wei and Ren, 2013). Fifteen different genera
and 13 described and a number of undescribed
species representing all five subfamilies of Ectobiidae
have been reported in the Cenozoic record
(Table 1). Of these, 10 genera have been reported
from the New World with the majority, seven, in
TABLE 1. The Cenozoic fossils of Ectobiidae. 
Subfamily* Genus Species Epoch Location Reference
Anaplectinae Anaplecta sp. Miocene Dominican amber Gutiérrez and PérezGelabert,
2000
Blattellinae Ischnoptera sp. Miocene Chiapas amber
(Mexico)
Solórzano Kraemer,
2007
Blattellinae ?Symploce rete Pleistocene African copal Gorochov, 2007
Ectobiinae Agrabtoblatta symmetrica Pleistocene African copal Gorochov, 2007
Ectobiinae Ectobius arverniensis Paleocene Menat (France) Piton, 1940
Ectobiinae Ectobius balticus Eocene Baltic amber Germar and Berendt,
1856
Ectobiinae Ectobius menatensis Paleocene Menat (France) Piton, 1940
Ectobiinae Ectobius glabellus Late Oligocene Rott (Germany) Statz, 1939
Ectobiinae Ectobius kohlsi Early Eocene Green River (USA) Vršanský et al., 2014
Ectobiinae Ectobius spp. (3) Early Eocene Green River (USA) Vršanský et al., 2011
Ectobiinae Isoplates longipennis Middle Eocene Gieseltal
(Germany)
Haupt, 1956
Ectobiinae Telmablatta impar Middle Eocene Gieseltal
(Germany)
Haupt, 1956
Nyctoborinae Nyctibora elongata Late Oligocene Rott (Germany) Statz, 1939
Pseudophyllodromiinae Cariblatta spp. (2) Miocene Dominican amber Gutiérrez and PérezGelabert,
2000
Pseudophyllodromiina Cariblattoides labandeirae Early Eocene Green River (USA) Vršanský et al., 2012
Pseudophyllodromiina Euthlastoblatta sp. Miocene Dominican amber Gutiérrez and PérezGelabert,
2000
Pseudophyllodromiinae Latiblattella Avita sp. nov. Middle Eocene Kishenehn
Formation (USA)
Vidlička and Greenwalt
(This study)
Pseudophyllodromiinae Plectoptera sp. Miocene Dominican amber Gutiérrez and PérezGelabert,
2000
Pseudophyllodromiinae Plectoptera electrina Miocene Haitian amber Gorochov, 2007
Pseudophyllodromiinae Pseudosymploce sp. Miocene Dominican amber Gutiérrez and PérezGelabert,
2000
Pseudophyllodromiinae Supella miocenica Miocene Chiapas amber
(Mexico)
Vršanský et al., 2011
*Subfamily assignments based on Beccaloni (2014)
PALAEO-ELECTRONICA.ORG
3
Miocene amber from either Mexico or Hispaniola.
Only Cariblattoides labandeirae Vršanský et al.,
2012 and Ectobius kohlsi Vršanský et al., 2014
from the Green River Formation in Colorado, and
Latiblattella avita Greenwalt and Vidlicka, 2015 sp.
nov. from the Kishenehn Formation in Montana,
are from North American Eocene deposits.
Dates of the Coal Creek Member of the Kishenehn
Formation of northwestern Montana have
been estimated to be 46.2 ± 0.4 Ma (middle
Eocene) by 40Ar/39Ar analysis and 43.5 ± 4.9 Ma
by fission-track analysis (Constenius, 1996). Deposition
of the fossiliferous deposits of the middle
sequence of the Coal Creek Member occurred in a
shallow near-shore setting that exhibited little or no
water flow in a seasonal subtropical/tropical environment
(reviewed in Greenwalt et al., 2015, in
press). The Kishenehn fossil insect fauna is quite
diverse with 15 different orders identified to date
although only a single specimen (USNM 595139)
out of 6,558 is from the order Blattaria (Greenwalt
et al., 2015, in press).
MATERIALS AND METHODS
Specimen USNM 595139 was collected at the
Dakin site on the Middle Fork of the Flathead River
near Pinnacle, Montana in 2013 in accordance with
USFS Authorization HUN281. The piece of oil
shale that contained the fossil also contained a fossil
notonectid (Heteroptera) and a leg of a tipulid
(Diptera). The specimen was photographed with an
Olympus SZX12 microscope equipped with a QColor5
Olympus camera. Image-Pro Plus 7.0 software
(Media Cybernetics, Inc., Bethesda, MD) was
used to capture and record the images. The specimen
was immersed in 95% ethanol for examination
and photography. Measurements were made with
the Image-Pro Plus 7.0 software. All measurements
are in millimeters (mm). Venational terminology
is from Vršanský (1997) as originally
developed by Comstock and Needham (1898).
SYSTEMATIC PALEONTOLOGY
Order BLATTARIA Burmeister, 1829 (= Blattariae
Latreille, 1810; = Blattodea Brunner von
Wattenwyl, 1882)
Family ECTOBIIDAE Brunner von Wattenwyl, 1865
Genus LATIBLATTELLA Hebard, 1917
Type Species. Latiblattella rehni Hebard, 1917
Diagnosis of the genus (after Hebard, 1917), relevant
material only. Size moderately large to
medium, form moderately broad to very broad, for
the group. Tegmina (in fully developed condition,
found in numerous species only in the male) delicate,
moderately broad, with costal and sutural
margins straight and subparallel in greater part,
scapular field very broad; discoidal (radial) sectors
numerous (usually, including their branches, eight
to ten), moderately oblique. Ventral margins of
median and caudal femora supplied with elongate,
moderately stout spines. First three tarsal joints
supplied distad with small pulvilli, brief ventral surface
of fourth joint occupied by a pulvillus. Moderately
large arolia present.
Latiblattella avita sp. nov.
(Figures 1-3)
zoobank.org/7225896D-EC71-4088-8B05-008C85F3A7B6
Etymology. The new species name is derived
from the latin avitus meaning ancient or ancestral.
Holotype. USNM 595139; a fragment containing
an intact tegmen attached (?) to an intact middle
leg. Deposited in the Department of Paleobiology,
National Museum of Natural History (NMNH),
Smithsonian Institution, Washington, District of
Columbia.
Type Horizon. Kishenehn Formation, middle
Eocene (Lutetian).
Type Locality. Dakin site, Middle Fork of the Flathead
River, near Pinnacle, Montana.
Differential Diagnosis. The significantly less
oblique radial sectors of the tegmen of Latiblattella
avita sp. nov. distinguish it from species in the
closely related genera Neoblattella, Shelford, 1911
and Lupparia Walker, 1868. Species of the genus
Balta Tepper, 1893 differ from Latiblattella in having
a protruded clavus. In addition, the marginal and
scapular fields of the tegmina are narrower in Latiblattella
than those in Eoblatta Shelford, 1911
(=Balta Tepper, 1893; synonymized by Roth,
1990). L. avita differs from most living representatives
of the genus Latiblattella in having a basally
forked, wide and darkly pigmented Sc vein as well
as a more pronounced coloration.
Description. Tegmen (forewing) elongated, 15.1
mm long and 4.4 mm wide (width measured at the
distal terminus of the anal field) with a length/width
ratio = 3.40. Sc wide and heavily pigmented, arcuate
basally with a slight inflection near the anterior
margin of the tegmen (Figure 1). Basal of Sc, the
tegmen is mottled with black pigmentation (Figures
2, 3.1). Sc itself is 4.75 mm long and 0.45 mm in
width at the point where it diverges from R, and
extends to a point about 66% of the length of the
anal field. Sc has no anterior branches. The
humeral field is 4.79 mm long, arcuate basally and
without anterior branches. Sc has a single posterior
branch which diverges from Sc at a point
GREENWALT & VIDLIČKA: NEW EOCENE COCKROACH
4
approximately 1/2 the distance from its origin; this
vein is heavily pigmented and is 2.18 mm long and
0.12 mm wide. Both Sc and its single branch reach
the wing’s margin. The basal portion of the radius
is distinctly curved and is pigmented to a point 4.5
mm from the wing’s base. There are 10-12 apical
branches of the radius – the origins of these
branches are not preserved. These branches are
oblique, evenly spaced and approximately half of
them are branched. The apical radial branches are
complexly branched and, with the anterior
branches of M, parallel to subparallel to the anterior
margin of the tegmen. Overall number of R
branches meeting margin is 26. Posterior branches
of M and those of Cu subparallel to the tegmen’s
longitudinal margin. The radial field is 8.5 mm long.
The apex of the tegmen is evenly curved. Intercalated
veins and crossveins are invisible throughout
the membrane. The anal field is 6.58 mm long and
2.51 mm wide at its widest point and contains at
least 10 near parallel simple longitudinal veins. The
plical furrow is subangulate and pigmented basally.
The plical notch is distinct although the posterior
margin of the tegmen is poorly preserved. The tegmen
is brown in coloration, more darkly brown
within a wide longitudinal stripe along the central
part of the wing with the margins of the wing lighter
in color. The anal field is dark brown except for the
postero-apical third which is essentially the same
color as the shale matrix. Given the poor preservation
of the posterior tegminal margin, the distribution
of color may have been affected by
taphonomic processes. Given the length of the tegmen
of this specimen, the insect may have been a
male (see Discussion).
In cockroaches, the dorsal aspect of the base
of the coxa is very closely apposed to the base of
the forewing and, given its size and association
with the forewing, the leg of this fossil may be a
mesothoracic appendage (Figure 1). It is 13 mm in
total length. The coxa, which is attached to the preserved
thoracal-coxal joint, is 4.05 mm long, 2.14
mm wide and black/dark brown in color. The basal
portion of the coxa and the trochanter are light
brown. The trochanter is triangular in shape, 1.13
mm long, 0.80 mm wide and overlaps the basal
femur by about 0.5 mm. Its shape resembles that
of Ectobiinae [vs. species in Blattellinae (Bazyluk,
1977)]. The femur is brown in color, slightly fusiform
in shape, 3.76 mm long and 1.0 mm wide. Its
posterior margin contains seven or eight relatively
short spines approximately 0.4 mm long and 0.04
mm wide, mostly on the apical half of the femur.
The tibia, also brown in color, is 3.3 mm long and
slightly wider apically (0.49 mm vs. 0.56 mm). The
tibia contains 14 visible spines, evenly distributed
FIGURE 1. Latiblattella avita sp. nov. (USNM 595139). Tegmen attached (?) to an intact middle leg. Scale bar equals
5 mm.
PALAEO-ELECTRONICA.ORG
5
over its length, 0.8 mm long and 0.08 mm wide.
Four of the tibial spines originate at the terminus of
the tibia and lie parallel to the first tarsal segment
(Figure 3.2). The dimensions of the five tarsal segments
are 1.3 mm x 0.31 mm, 0.5 mm x 0.2mm,
0.35 mm x 0.22 mm, 0.2 mm x 0.2 mm and 0.46 x
0.17 mm increasing distally to 0.24 mm wide. T1,
T2 and T3 have triangular distal extensions, which
may contain remnants of tarsal pads (pulvilli). T4 is
bilobed basally, as in Ectobiinae [vs. species in
Blattellinae in which the apical and basal margins
are parallel (Bazyluk, 1977)]. The single asymmetrical
claw that is preserved/visible is approximately
0.36 mm long. The arolium is about 0.2 mm in
length and black/dark brown.
DISCUSSION
Assignment of the New Species to the Genus
Latiblattella
The present specimen is categorized within
the genus Latiblattella based on the generally
unspecialized appearance of the comparatively
robust leg, including an asymmetrical claw, nearly
identical venation of tegmen with branched Sc,
short, pigmented and distinctly curved R, more or
less straight M and CuA, very distinct and arcuate
boundary between clavus and rest of the tegmen.
It differs from the related genera Eoblatta and
Supella Shelford, 1911 in the decidedly less
strongly oblique radial sectors of the tegmen. In
addition, the marginal and scapular fields are narrower
in Latiblattella than those in Eoblatta. It differs
from the related genera Neoblattella and
Lupparia in the degree of the obliqueness of the
radial sectors of the tegmen; those of Latiblattella
are decidedly less oblique. The closely related
genus Balta is characterized by a protruded clavus,
a morphological character absent in Latiblattella.
Nevertheless it is necessary to note that the present
specimen has coloration and general appearance
somewhat similar to Lupparia adimonialis
Walker, 1868 (see plate 1, figure 12 in Shelford,
1908) suggesting a close relationship to the colored
representatives of Latiblattella such as the
present species. The present specimen is, in contrast
to living representatives of the genus, somewhat
coloured, but coloration varies greatly within
the related cockroach genera. For example, Balta
varies from completely colorless to strongly
coloured species. The number of veins falls within
the variation of the genus (see Rehn, 1951; Brunner
von Wattenwyl, 1865). Branching of Sc occurs
within the genus, although the basal branching is
considered to be a plesiomorphy (i.e., present in
ancestral Mesoblattinidae) (Vrsansky et al., 2002).
FIGURE 2. Latiblattella avita sp. nov. (USNM 595139). 2.1. A photograph of the tegmen. 2.2. A line drawing of the
forewing venation. The arrow denotes the boundary between the radial and medial fields. M, medial veins; R, radial
veins; Sc, subcostal vein. Scale bar equals 3 mm.
GREENWALT & VIDLIČKA: NEW EOCENE COCKROACH
6
L. avita sp.n. has venation almost identical with L.
rehni. Unfortunately, the data do not reveal enough
information for a phylogenetic analysis. While the
Sc vein is often simple in extant species of Latiblattella,
it exhibits a single posterior branch in both L.
avita and L. vitrea Brunner von Wattenwyl, 1865.
The posterior branch originates closer to the forewing
margin than the origin of Sc in both species.
Similarly, in both species, both Sc and the distinctly
curved basal portion of the radius are heavily pigmented.
Dark pigmentation of the single posterior
branch of Sc is however unique to L. avita.
Extant species of Latiblattella, in addition to
exhibiting sexual dimorphism relative to body and
wing length such that females often have significantly
reduced wings, vary significantly in size
(Hebard, 1917, 1921, 1922, 1932). The holotype of
Latiblattella vitrea (♂) was reported to have a tegmina
length of 10 mm while that of L. mexicana
Saussure, 1864 is 16 mm (Brunner von Wattenwyl,
1865; Saussure, 1864). L avita sp. n. is 15.1 mm in
length and therefore amongst the largest of the
species of this genus. The tegmina length/width
ratio of extant species ranges from 3.0 (L. pavida
Rehn, 1903) to 4.53 (L. azteca Saussure and
Zehntner, 1893) although this latter species is
unusual in that most species exhibit a ratio
between 3.0 and 3.5 (Rehn, 1903); the tegmina of
L. avita sp. n. fits comfortably within this range. The
length of the humeral field, defined as the distance,
on a line parallel to the wing’s anterior margin, from
the tegmina’s base to the fusion of the primary Sc
vein with the wing’s margin, relative to that of the
anal field, is another potentially valuable morphometric
measurement. Unfortunately, most holotypes
and paratypes have not been figured in the
literature and, as a result, the data is limited. This
ratio is equal to 0.92-0.94 (L. rehni), 0.92 (L. lucifrons
Hebard, 1917), 0.79 (L. vitrea) and 0.66 (L.
avita sp. n.) (Hebard, 1917; Rehn, 1951; Brunner
von Wattenwyl, 1865). Given this limited available
data and the single fossil specimen, the significance
of L. avita’s smaller ratio is unknown.
Latiblattella avita sp. nov. and the Fossil
Record
Different extant species of Latiblattella have
extraordinarily diverse habitats given their small
number. L. rehni is found under cracks in the bark
of Pinus caribea and within strands of Dendropogon
usneoides (Spanish moss), L. chichimeca
Saussure and Zehntner, 1893 is found on bromelias,
L. lucifrons feeds on pollen and detritus on the
flowers of Yucca elata and L. zapoteca Saussure,
1862 is found under stones along the margins of
rivers (Rehn, 1906; Hebard, 1917; Ball et al., 1942;
Blatchley, 1920; Picado, 1913). The conserved
fragment of L. avita appears to be a remnant that
could have been dropped from a predator or
washed into the margins of the lake via a small
stream and it is impossible to know the niche occupied
by the insect. Although pine leaves - yet to be
identified - are found in the shales and siltstones of
the Kishenehn Formation, none are known from
the Dakin site although a single pine seed has
been collected there.
Most Cretaceous cockroach clades (e.g., Blattulidae,
Caloblattinidae and nearly all of the Mesoblattinidae
and Skokidae) are not present in the
fossil record of the Cenozoic; only the single extant
genus Blattella has a fossil record in the Cretaceous
(Vršanský, 2008). On the other hand, nearly
all (9/11) cockroach genera from the Green River
FIGURE 3. Latiblattella avita sp. nov. (USNM 595139).
3.1. The subcostal field of tegmen showing the wide
and darkly pigmented Sc vein. 3.2. The five segmented
tarsus. The arrows denote tarsal segments 1 – 5 and
the end of the single visible claw. Scale bars equal 1
mm.
PALAEO-ELECTRONICA.ORG
7
Formation and all genera from Dominican Republic
and Mexican amber are extant. The marked differences
in the composition of the Cenozoic and Cretaceous
entomofauna suggest a very rapid
evolution and radiation of the cockroach biota in
the Paleocene and early Eocene (Vršanský et al.,
2011; 2012; 2013; Gorochov, 2007). Interestingly,
Namablatta, Diploptera, Ectobius, Allacta Saussure
and Zehntner, 1895, Blattella and Supella are
all extant genera that were present in the North
and Central American Paleogene that are now,
except for recent reintroduction, extinct in those
areas (Vršanský et al., 2011). Their absence, in
combination with the highly advanced nature and
restricted geographical distribution of the Dominican
fauna (Central and South America), led
Vršanský et al. (2011) to suggest that an environmentally/biologically-mediated
extinction event
precipitated extinction of Paleogene fauna and set
the stage for the evolution of a distinctly American
fauna. On the other hand, Cariblattoides labandeira,
present in the lacustrine sediments of the
early Eocene Green River Formation, is an extant
species with a present-day distribution in South
America and the Caribbean (Vršanský et al., 2012).
Latiblattella avita sp.n. is a species from yet
another genus that was able to persist from the
middle Eocene to present-day in Central America
(An alternative explanation would be the migration
of L. avita from its Eocene territory to its present
day location sometime over the last 46 million
years due to the cooling temperatures of the latter
half of the Cenozoic.). Extant species of Latiblattella,
restricted to Central America, Cuba, Mexico,
the Bahamas, Florida and Arizona (Princis, 1969),
may be restricted in distribution relative to that of
the Eocene. So too, Cariblattoides Rehn and
Hebard, 1927, which is presently restricted to
Cuba, Puerto Rico and Brazil (Bonfils, 1975).
Given their preference for the tropics and the near
universal subtropical/ tropical environments of the
Eocene (Wolfe, 1995; Zachos et al., 2001), the
presence of these genera in Eocene North America
is not unexpected. Although recent molecular
phylogenetic data has indicated a close relationship
between Latiblattella and the oriental genus
Balta (Inward et al., 2007), there is no fossil record
for the latter genus and a paleobiogeographical link
between the two has yet to be established.
ACKNOWLEDGEMENTS
We thank C. Labandeira and F. Marsh for
administrative support and P. Vršanský (GlU SAV,
Bratislava) for review of the manuscript. We are
also indebted to two anonymous reviewers whose
efforts greatly improved the manuscript. This is
contribution number 326 of the Evolution of Terrestrial
Ecosystems Consortium of the USNM. This
research was supported by VEGA 2/0186/13.
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Society of America, 107:28-36.
Walker, F. 1868. Catalogue of the specimens of Blattariae
in the collection of the British Museum. British
Museum (Natural History), London.
Wei, D. and Ren, D. 2013. Completely preserved cockroaches
of the family Mesoblattinidae from the Upper
Jurassic-Lower Cretaceous Yixian Formation (Liaoning
Province, NE China). Geologica Carpathica,
64:291-304.
Wolfe, J.A. 1995. Paleoclimatic estimates from Tertiary
leaf assemblages. Annual Review of Earth and Planetary
Sciences, 23:119-142.
Zachos, J., Pagani, M., Sloan, L., Thomas, E., and Billups,
K. 2001. Trends, rhythms and aberrations in
global climate 65 Ma to present. Science, 292:686-
693.
Zompro, O. and Fritzsche, I. 1999. Lucihormetica fenestrata
n.gen., n.sp., the first record of luminescence in
an orthopteroid insect (Dictyoptera: Blaberidae:
Blaberinae: Brachycolini). Amazoniana, 15:211-219.
Príloha č. 18
VIDLIČKA, Ľ. 2001.
Blattaria – šváby; Mantodea – modlivky: (Insecta: Orthopteroidea).
1. vyd., Veda, Bratislava, 169 pp.
(Fauna Slovenska) ISBN 80-224-0640-6
Blattaria - šváby
Mantodea - modlivky
(lnsecta: Orthopteroidea)
LUBOMÍR VIDLIČKA
VEDA
vydavateľstvo
Slovenskej
akadémie
vied
SLOVENSKÁ AKADÉMIA VIED
Ústav zoológie
Vedecký redaktor
RNDr. Milan Kozánek, CSc.
Recenzenti
Prof. RNDr. Ivan Országh, DrSc.
RNDr. Ilia Okáli, CSc.
© Ľubomír Vidlička 2001
ISBN 80-224-0640-6
FAUNA SLOVENSKA
Blattaria - šváby
Mantodea - modlivky
(lnsecta: Orthopteroidea)
lubomír Vidlička
VEDA, vydavatel'stvo Slovenskej akadémie vied
Bratislava 2001
Obsah
Rad: BLATTARIA Burmeister, 1829- šváby ............................................................................ 7
Úvod ......................................................................................................................................... 7
A. Všeobecná časť .......................................................................................................................... 8
l. Morfologická charakteristika imág ................................................................................ 8
2. Anatomická charakteristika imág ................................................................................ 20
3. Charakteristika nedospelých štádií .............................................................................. 23
4. Etológia a ekológia ......................................................................................................... 25
Obydlia- biotopy ............................................................................................................. 25
Potrava .............................................................................................................................. 26
Sociálne správanie ............................................................................................................ 28
Rozmnožovanie ............................................................................................................ ... 29
Embryonálny vývoj ................................ .......................................................................... 32
Postembryonálny vývoj .................................................................................................... 32
Starostlivosť o potomstvo ................................................................................................. 33
Prirodzení nepriatelia ........................................................................................... ............ 34
Obrana .............................................................................................................................. 35
S. Geografické rozšírenie švábov........................................................................... 37
Pôvod synantropných druhov a ich rozšírenie ................................................................. 38
6. Hospodársky význam ..................................................................................................... 39
Ochrana pred synantropnými švábmi .............................................................................. 40
Chov švábov ..................................................................................................................... 41
Šváby a povery .............................................................................................................. .. 42
Šváby v medicíne ................................... .......................................................................... 43
7. Prehľad histórie poznania švábov do roku 1758 ............................................................ 43
Vznik národných mien švábov ......................................................................................... 45
8. Vývoj názorov na systematické usporiadanie švábov po roku 1758 ................ ............ 46
9. Klasifikácia švábov a modliviek ....................................................................................... 49
10. Fylogenéza švábov .......................................................................................................... 52
Fylogenetické vzťahy švábov a modliviek k iným radom ................................................ 54
Spoločný predkovia švábov a termitov ............................................................................ 55
ll. História výskumu švábov na Slovensku ....................................................................... 56
B. Systematická časť .......................................... .......................................................................... 60
Rod Blatta Linnaeus, 1758- šváb ................................................................................ 60
Rod Periplaneta Burmeister, 1838 -šváb .................................................................... 62
Rod Blattella Caudell, 1903 - rus ................................................................................. 66
Rod Supella Shelford, 1911 -šváb ...... ......................................................................... 68
Rod: Ectobius Stephens, 1835 - švábik ........................................................................ 70
Rod: Phyllodromica Fieber, 1853 - švábik ...................................................... ............ 80
Rad: MANTODEA Burmeister, 1838- modlivky ................................................................... 94
Úvod ................................................................................................................................ ...... 94
A. Všeobecná časť ........................................................................................................................ 95
l. Morfologická charakteristika imág ............................................................................. 98
2. Anatomická charakteristika imág .............................................................................. l02
3. Charakteristika nedospelých štádií ............................................................................ l03
4. Etológia a ekológia ....................................................................................................... l04
Obydlia - biotopy ..................................................................................... ..................... 105
Potrava ............................................................................................................................ l05
Rozmnožovanie ................................................................................................... ........... 107
Postembryonálny vývoj .................................................................................................. II O
Prirodzení nepriatelia ..................................................................................................... III
Obrana a mimikry .......................................................................................................... 11 2
5. Geografické rozšírenie ........................ ......................................................................... 11 5
6. Hospodársky význam ................................................................................................... l 15
Chov modliviek .............................................................................................................. 11 6
7. Prehľad histórie poznania modliviek do roku 1758 .................................................. l 16
Etymológia vedeckého a národného mena modliviek ................................................... 118
Ludové názvy modlivky a rôzne povery ...................................................................... 11 9
8. Vývoj názorov na systematické usporiadanie modliviek po roku 1758 .................. 11 9
9. História výskumu modliviek na Slovensku ................................................. ............... 12 1
10. Fylogenéza modliviek ................................................................................................... 122
B. Systematická časť ................................................................................................................. 124
Rod Mantis Linnaeus, 1758- modlivka................................................................. 124
Systematický prehľad švábov a modliviek ............................................................. ........... 126
Smn1nary ............................................................................................................................... 127
Key to the families and genera of cockroaches .................................................................... 131
Použitá literatúra ................................................................................................................ 133
Príloha .................................................................................................................................. 157
Register švábov (Biattaria) ................................................................................................. 161
Register modliviek (Mantodea) .......................................................... .................... ........... 164
Register húb, rastlín a živočíchov (okrem švábov a modliviek) ..................................... 166
Rad: BLATTARIA BuRMEISTER, 1829 - šváby
Úvod
Šváb, rus, tarakán- kto by nepoznal aspoň jeden z týchto názvov označujúcich malé zvieratko,
ktoré sa cíti dobre práve v ľudských príbytkoch a sprevádza človeka už niekoľko storoč í (č i
možno tisícročí) na jeho cestách. Tieto synantropné druhy majú často kozmopolitné rozšírenie, ale
ich bližšiemu poznaniu bráni skrytý nočný spôsob života a veľká rýchlosť, akou unikajú pred svetlom.
Iba nepatrné percento ľudí, zväčša iba tí, ktorí sa zaoberajú biológiou, vedia, že prevažná
väčšina druhov švábov žije voľne v prírode a viacero druhov dokonca aj u nás. Napriek tomu, že sú
blízkymi príbuznými už spomínaných synantropných druhov, ich pôsobenie v prírode môžeme
hodnotiť kladne. Všežravosť švábov je v prirodzených podmienkach, najmä v trópoch a subtrópoch,
kde má prevažná väčšina druhov svoj domov, ideálnou vlastnosťou z hľadi ska rozkladu
organických látok.
Šváby sú jednou z najstarších hmyzích skupín. Ich prví prapredkovia sa objavili na Zemi pred
viac ako 300 miliónmi rokov v karbóne (prvohory) a zanedlho sa stali jedným z n ajväčšíc h hmyzích
radov. V súčasnosti žijúce šváby (4 500- 5 000 druhov z asi 460 rodov) sú v porovnaní
s inými radmi hmyzu iba malou skupinou, ale ich zástupcovia sú rozšírení takmer po celej Zemi.
Šváby sú z hospodárskeho a medicínskeho hľadiska významnou skupinou. Aj preto sa v 20.
storočí stali jedným z najčastej ších hmyzích objektov biologického vedeckého výskumu. Značná
pozornosť, ktorá im je venovaná, sa prejavuje vo veľkom počte publikovaných prác a samozrejme
i v množstve poznatkov o tejto skupine. Celosvetový katalóg švábov zahrnujúci približne 3 800
dovtedy opísaných druhov zostavil švédsky blattológ Karlis PRINCIS ( 1962, 1963, 1964, 1965,
1966, 1967, 1969, 1971 ). Komplexnému spracovaniu švábov východného palearktu sa venoval
BEY-BIENKO ( 1950). Jeho práca sa stala spolu so staršou Rammeho revíziou rodu Ectobius (RAMME
1923) základom pri štúdiu stredoeurópskych druhov švábov. Prvý kľúč stredoeurópskych druhov
švábov vypracoval HARZ ( 1957). Následne boli spracované šváby Európy (PRINCIS 1965 a HARZ
1976). Od vydania spomínaných prác bolo z Európy opísaných viacero druhov švábov z rodov
Phyllodromica a Ectobius. Revízii niektorých skupín rodu Phyllodromica sa v posledných rokoch
venovali BoHN ( 1992, 1993, 1999) a VIDLIČKA a MAJZLAN (1997). Mnohé z publikovaných prác
majú podobu obsiahlych monografií zaoberajúcich sa predovšetkým biológiou švábov a ochranou
pred nimi - ROTH & WILLIS (1960), BEIER( 1961 , 1974), CORNWELL (1968, 1976), BELL & ADIYODI
(1981 ), RusT et al. (1995) a ďalšie .
Najstaršie údaje o výskyte švábov z územia dnešného Slovenska pochádzajú spred takmer 200
rokov (BARHIOLOMAEIDES 1806- 1808). Do súčasnosti sa nahromadilo množstvo najmä faunistických
údajov, ktoré bolo potrebné v prvom rade sumarizovať a kriticky zhodnotiť (VIDLi č KA a MAJZLAN
1992; VIDLIČKA a SZIRÁKY 1997). Až nás ledne mohlo vzniknúť monografické spracovanie
švábov.
Predložená práca vzhľadom na svoj rozsah a svoje poslanie zďaleka nie je a ani nemôže byť
úplným súhrnom poznatkov o tejto zaujímavej skupine hmyzu. Snahou bolo podať ucelenú a prehľadnú
charakteristiku celého radu Blattaria a v systematickej časti zhrnúť a obohatiť doterajšie
poznatky o šváboch žijúcich na území Slovenska. Dúfam, že sa to aspoň čiastočne podarilo.
7
A. Všeobecná časť
l. Morfologická charakteristika imág
Veľkosť. Šváby dosahujú väčšinou stredne veľké až veľmi veľké rozmery, malé druhy sú zriedkavejšie.
Najmenšie sú v termitiskách žijúce druhy z rodu Nocticola (asi 3 mm) a v mraveniskách
žijúce druhy z rodu Attaphilla (asi 4 mm). Najväčší známy šváb je Megaloblatta longipennis
s rozpätím krídiel viac ako 18 cm a šváby z rodu Blaherus dlhé okolo 120 mm. Najťažším švábom
je bezkrídly austrálsky druh Macropanesthia rhinoceros, ktorý dosahuje dÍžku 65 mm a hmotnosť
okolo 20 gramov (DAY 1950).
lntegument (integumentum). Šváby, ako aj ostatné článkonožce, majú vonkajšiu kostru
(exosceletum). Exoskelet je tvorený na povrchu nebunkovou kutikulou, ktorá je vylučovaná vonkajšou
(ektodermálnou) vrstvou telových buniek -epidermou. Pod epidermou je tenká bazálna
membrána. Epiderma s kutikulou a bazálnou membránou tvoria integument. Aby bola zabezpečená
pohyblivosť, kutikula netvorí homogénny celok, ale jednotlivé čl ánky sú kryté malými pl atni čkami
(skleritmi) navzájom pospájanými mäkkými a pružnými blankami - intersegmentálnymi
membránami. [ntegumentje u švábov obyčajne mäkký, ale u niektorých skupín, najmä u bezkrídlych
foriem môže dosahovať značnú tvrdosť (napr. Polyphaginae, Panesthinae). Kutikula švábov,
tak ako u ostatného hmyzu, sa skladá z troch hlavných vrstiev. Periplaneta americana má kutikulu
hrubú 40 1-1m. Jej najvnútornejšia vrstva- endokutikula- je hrubá 20- 30 1-1m a skladá sa z vonkajšej
a vnútornej vrstvy. Nad ňou je lO- 20 1-1m hrubá exokutikula, ktorá obsahuje č i erne pigmenty-
melaníny. Extrémne tenká vonkajšia vrstva- epikutikula má 2 flm. Skladá sa z dvoch chemicky
odlišných vrstiev. Tenká vonkajšia vrstva (0,02- 0,03f.lm) je priehľadná, hygrofobická a odolná
voči kyseline. Vnútorná vrstva je jantárovo sfarbená a nie je odolná voči kyseline. Na povrchu
epikutikuly je vosková vrstva, ktorá má dôležitú funkciu pri priepustnosti vody. Vosková vrstva
u švábov sa obnovuje počas celého ich života. Predpokladá sa, že vosk je u švábov vylučovaný na
povrch kutikuly v roztoku, ktorý sa vyparuje len veľmi pomaly. Vosková vrstva u švábov je zložená
z nenasýtených uhľohydrátov, mastných kyselín a z rôznych aldehydov. U posledného nymfálneho
instaru Periplaneta americana je táto vrstva hrubá 0,4 1-1m a u dospelých l - 2 1-1m. Podobne je
to aj u Blatta orientalis, kde vosková vrstva dosahuje na brušku hrúbku 0,6 1-1m (BEAMENT 1945 ;
DENNELL & MALEK 1955a, 1955b, 1956).
Povrch tela švábov je väčšinou hladký a lysý. Hustejšie ochlpenie sa vyskytuje iba zriedkavo,
častejš ie je iba u púštnych a stepných druhov. Rôzne výrastky a tŕne na kutikule sú veľmi vzácne.
Ojedinelá je i zrnitá štruktúra.
Sfarbenie. Šváby majú takmer výlučne pigmentózne sfarbenie. Najbežnejšie je slamovožlté,
žltohnedé a tmavohnedé, zriedkavejšie čierne. Väčšina švábov je sfarbená veľmi nenápadne. Kovové
sfarbenie je veľmi zriedkavé (napr. Eustegasta buprestoides z Kamerunu a Konga alebo
niektoré Plectopterinae). Občas sa vyskytuje i mimetické sfarbenie (bližšie pozri časť Obrana).
Sfarbenie švábov nie je dobrým diagnostickým znakom, vykazuje vysoký stupei'1 variability.
Bežné je odlišné sfarbenie jedincov toho istého druhu z rôznych lokalít.
Hlava (caput) (obr. 3) je voľná, väčšinou hypognátna (caput hypognathale), prieč ne stl ačená
s typickými švami a s dobre vymedzenými skleritmi . Epikraniálny šev (sutura epicranialis) nie je
u dospelých vždy prítomný. Čelo (frons) a čelový štítok (clypeus) sú veľké, dobre vyvinuté. Tentórium
(vnútorná kostra hlavy) je charakteristické otvorom v stredovej časti (v tele tentória). Zhoraje
hlava buď úplne zakrytá štítom alebo je predná časť hlavy čiastoč ne vid iteľná. Zložené oči (oculi
compositi) sú skoro vždy vyvinuté, veľké sú najmä u krídlatých foriem. Bezkrídle formy majú oči
výrazne menšie a medziočný priestor veľmi široký. Zriedkavo môžu byť oč i redukované alebo
8
Obr. l. Ectobius sylvestris o - dorzálny pohl'ad. a - pronótum, b - disk pronóta, c - kostálne pole, d - análne pole,
e - análna brázda, f- subkostálna žilka, g- radiálna+ mediálna žilka, h- zadná holeň , i - l'avé predné krídlo (tegmina),
j - pravé predné krídlo (tegmina), k- zadné chodidlo, l - vankúšik, m - tykadlo. Orig.
Fig. l. Ectobius sylvestris o - dorsal view. a- pronotum, b- disc of pronotum, c- costa] area, d - ana] area, e - ana!
furrow, f- subcostal vein, g- radia]+ media] veins, h- hi nd tibia, i - left fore wing (tegmen), j - right fore wing (tegmen),
k - hind tarsus, l - arolium, m- antenna. Orig.
9
Obr. 2. Ectohius sylvestris o - ventrálny pohrad. a - hlava, b - čerusťové hmatadlo, c - predné stehno, d - predný
trochanter, e - zadná panvi čka, f- zadné stehno, g - zadný trochanter, h - vejárovito zložené zadné krídlo, i - siedme
sternum, j -siedme tergum, k - subgenitálna pl atnička, l - cerkus. Orig.
Fig. 2. Ecrobius sylvestris ô - ventral view. a - head, b - maxillary palp, c- front femur, d - front trochanter, e - hind
coxa, f- hi nd femur, g - hind Lrochanter, h- folding hind wing, i- sternum 7, j- tergum 7, k- subgenital plate, 1- cercus.
Orig.
lO
chýbajú, najmä u myrmekofilných a kavernikolných foriem (napr. u zástupcov rodu Spelaeoblatta
z Barmy a Thajska alebo Trogloblattella zo Sarawaku). Zložené oči sú vytvorené z množstva malých
omatídií, každé omatídium má bikonvexnú šošovku vytvorenú z priehl'adnej kutikuly.
U niektorých druhov švábov sú vyvinuté dve jednoduché očká (ocelii), ale častejš ie sú na ich
mieste iba 2 bledé, na svetlo citlivé, ocelliformné škvrny (fenestrae) pri vnútornom okraji očí.
Fenestrae sú nervami spojené s mozgom a ich histologická štruktúra pripomína degenerované
očká redukované na pigmentovanú kutikulu. U foriem s plne vyvinutými krídlami sú očká obyčajne
dobre vyvinuté, u bezkrídlych (apterných) foriem hrabajúcich v zemi zvyčajne chýbajú.
Tykadlá (antennae) sú mnohočlánkové, vel'mi dlhé (často dlhšie ako telo), nitkovité, smerom
ku koncu sa zužujú. Zriedkavejšie sú v strede zhrubnuté, vretenovité (napr. u tropického rodu
Pseudothyrsocera). Ku hlave sa pripájajú približne v strede vnútorného okraja očí. Vyrastaj ú
z okrúhlej tykadlovej jamky ohraničenej tykadlovým švom (sutura antennalis), ktorý má uprostred
Obr. 3. Ecto!Jius sylveslris ó - hlava. a- epikraniálny šev, b - zložené oko, c - skapus, d - pedicel, e - bičík tykadla,
f- líce, g - čelový štítok, h- horná pera, i- spodnoperové hmatadlo, j- če ľusiové hmatadlo, k- epikránium, l - ocelliformná
škvrna (očko), m- tykadlová jamka, n- čelo, o- čeľusťový kmeň , p- hryzadlo, r- galea, s - paraglosa. Orig.
Fig. 3. Ecto!Jius sylvestris ó - head. a- epicranial suture, b - compound eye, c - scape, d - pedicel, e - antennal
flagellum, f- gena, g- clypeus, h- Iabrum, i - Iabial palp, j - maxillary palp, k- epicranium, I - fenestra, m - antennal
pit, n- frons, o- stipes, p- mandible, r- galea, s - paraglossa. Orig.
ll
vnútorného okraja apofýzu. Prvý tykadlový článok, skapus (scapus) je najdlhší, druhý článok,
pedicel (pedicellus) má tvar guličky, ostatné články sú obdÍžnikovité, navzájom podobné. Všetky
články sú zvyčajne pokryté jemnými krátkymi chÍpkami. Sfarbené sú spravidla jednofarebne tmavo,
ale časté je i biele sfarbenie niekol'kých článkov v strednej časti tykadla. Tykadlá sú orgánom
čuchu a hmatu. Sú veľmi pohyblivé.
Ústne orgány (trophi) (obr. 4) smerujú pri odpočinku dozadu a dolu. Sú hryzavého typu (trophi
masticatorii). Tvorí ich 5 dobre oddelených častí- horná pera (labrum), hryzadlá (mandibulae),
čeľuste (maxillae),jazýčok (hypopharynx) a spodná pera (labium). Hryzadlá sú veľké, nepravidelne
štvoruholníkovité, silne sklerotizované, na vnútornej strane ozubené, na báze s dobre vyvinutými
žuvacími plochami. Hryzadlá švábov nesú prívesky (prostheca). Na zadnom povrchu každého
hryzadla sú pozdÍžne rady štetín. Čeľuste sa skladajú z viacerých častí. Čap (cardo) je krátky,
rozdelený. Na predÍženom kmeni (stipes) je sformovaný osobitný sklerit nazývaný subgalea. Na
kmeni sú na malých palpiferoch umiestnené S-článkové čeľusťové hmatadlá (palpi maxillares).
Galeaje pomerne mäkká, cylindrického tvaru. Lacínia má na vnútornej strane dva dobre vyvinuté
do stredu smerujúce zúbky, v strede je na jej povrchu množstvo dlhých štetín. Jazýčok je vel'ký
s bočnými sklerotizovanými výbežkami (suspenzóriami) a stredovou priehlbinou (sitophora).
Spodná pera tvorí najväčší diel zadnej časti hlavovej kapsuly. Skladá sa z vel'kého, širokého
a oválneho podbradku (submentum), malej brady (mentum) a prementa nesúceho na palpigeroch
pár 3-článkových spodnoperových hmatadiel (palpi labiales). Na distálnom konci spodnej pery sú
dobre vyvinuté párové glosy (glossae) a paraglosy (paraglossae). Na ich vrchole sú rady jemných,
k ústnemu otvoru smerujúcich štetín.
B
f
g
Obr. 4. Ectobius sylvestris ô - ústne orgány. A - čeľusť ; B - hryzadlo; C- spodná pera. a- čap, b - krner'í, c- lacínia,
d - galea, e- čel\rsťové hrnatadlo, f- ostrie, g- rezáky, h- podbradok, i -brada, j- palpiger, k- prernenturn, l -glosa,
rn - paraglosa, n - spodnoperové hrnatadlo. Orig.
Fig. 4. Ectobius sylvestri.1· ô - rnouth par1S. A - rnaxilla; B - rnandible; C - labiurn. a- cardo, b - stipes, c - lacinia,
d- galea, e - rnaxillary palp, f- mola, g- denticles, h- submentum, i- menturn, j- palpiger, k- prementum, l - glossa,
m - paraglossa, n - labial palp. Orig.
Kŕčok (cervix). Hlava je na predohruď pripojená dlhým kŕčkom . Dorzálna strana kŕčka je
zrete!'ne kratšia ako ventrálna strana. To umožňuje ohnutie hlavy pod hruď i jej horizontálne vysunutie.
Kŕčok je pokrytý arthrodiálnou membránou. Na spevnenie kŕčka slúžia dva páry bočných
cervikálnych skleritov (cervicalia) ležiace medzi záhlavnými kondylami a latero-anteriornými
rohmi predohrudného sterna. Keď je hlava zatiahnutá pod hruď, cervikálna membrána na spodnej
12
strane kŕč ka sa prehne pod predný a zadný okraj ventrálnych cervikálnych skleritov. Tým sa dosahuje
skrátenie funkčnej dÍžky ventrálnej strany kŕčka (POPHAM 196 1).
Hruď (thorax). Predohruď (prothorax) nesie dorzálne vel'ký, silne sklerotizovaný predohrudný
štít (pronotum), ktorý väčšinou prekrýva hlavu. Povrch pronóta je slabo klenutý, buď hladký
alebo s nevel'kými hrbo l čekmi v zadnej časti nad základmi prvého páru krídiel. Inokedy môže byť
povrch zrnitý alebo jemne ochlpený. B oč né okraje a prípadne i predný okraj pronóta bývajú často
jasnejšie sfarbené ako jeho stred (disk) a obyčajn e sú pri ezrač n é (napr. Ectobius). Mezoa
metanótum sú viac alebo menej pravouhlé a navzájom podobné, u krídlatých foriem slabo sklerotizované
a úplne zakryté tegminami a krídlami. Obe sú rozdelené na akrotergit (acrotergit), predštít
(prescutum), štít (scutum) a obyčaj ne vypuklý štítok (scutellum). Postnóta nie sú vyvinuté. Na
bokoch sú závesné kÍby (kondyly) na uchytenie tegmín a krídiel. U brachypterných a apterných
foriem je mezo- a metanótum silne sklerotizované, bez akýchkol'vek štruktúr. Vzhl'adom sú podobné
na abdominálne tergá za nimi.
Pleury sú vďaka silne sploštenému telu a mohutnému rozvoju hrudných tergov malé. Episternity
sú omnoho väčšie ako epimeróny. Stredo- a zadohrudné spirákulá sú prítomné.
Sterná sú slabo sklerotizované, trochu stl ačené, väčšin ou zakryté silne vyvinutými panv i čkami.
Prosternit je tvorený drobnou, úzkou, pretiahnutou pl atni č kou rozdelenou priečnym švom na
dve časti - predný eusternit (basisternit) a zadné sternellum (furcasternit). Vzadu je malé prosternellum
(spinasternit). U mezo- a metasternitu je bazisternit redukovaný a po dÍžke rozdelený na
dve malé pl atni čky, furkasternit (sternellum)je silne redukovaný, vo vnútri má furku v tvare písmena
Y. Mezospinasternit je vyvinutý, metaspinasternit chýba.
Nohy (pedes) sú väčšinou behavé (pedes cursorii), nemodifikované, všetky tri páry stavané
podobne (homonómne). Zriedkavejšie sú prispôsobené hrabaniu v zemi (pedes fossorii), najmä
predný pár (napr. u Arenivaga, Panchlora, Nymphytria). Holene sú v tom prípade vpredu zhrubnuté
a vyzbrojené silnými tŕň mi . Panvičky (coxae) sú dlhé, uložené vel'mi blízko pri sebe, často sú
obrúbené. Trochanter je veľmi malý, pripojený k zadnej časti stehien. Stehná (femora) sú dlhé, po
celej dÍžke približne rovnako široké, spodná (ventrálna) strana často s dvomi kýlovitými okrajmi,
vyzbrojenými mnohými dlhými tŕň mi. Otfnenie je charakteristické najmä na spodnom prednom
(anteroventrálnom) okraji predného páru stehien. Typ otŕneni a je dôležitý aj taxonomicky. Vzhľadom
na charakter otŕneni a sú známe 3 typy predných stehien - pôvodnejší typ A a odvodené typy
B a C. Typ A je charakterizovaný radom silných, dlhých tŕňov, ktoré sa postupne mierne skracuj ú
od bázy stehna k jeho vrcholu (napr. Blatta, Periplaneta, Supe/la). Typ B je charakteristický zmenou
väčšiny tŕňov na jemné štetinky, zachovávajú sa iba bazálne tŕne (napr. Phyllodromica, Ectobius).
Pri type C sú všetky tŕne zmenené na štetinky (napr. Nocticola). V rámci typov môžeme
ďal ej roz li šovať rôzne obmeny základného typu (ozn ačované A l, A2, A3, B l , B2 atď.) . Pri
všetkých typoch je distálny vrchol stehna vyzbrojený 2-3 dlhými vrcholovými tŕň mi . Zadný spodný
okraj stehna sa nevyzn ačuj e žiadnymi osobitosťami , otŕneni e je rovnakého typu a rovnakej
vel"kosti. Podobné otŕneni e ako na predných stehnách sa nachádza i na stredných a zadných stehnách.
Na distálnom konci stredných a zadných stehien v mieste pripojenia holene sa nachádza
vel'ký kolenný tŕň (chýba len pri niektorých špecializovaných skupinách, napr. podče ľaď Panesthiinae).
Kolenný tŕň nie je nikdy na predných stehnách. Holene (tibiae) sú vždy vyzbrojené mnohými
tŕň mi. V äčš inou sú dlhé, ale u druhov hrabajúcich v zemi sú krátke a široké. Chodidlá (tarsi) sú
vždy S-čl ánkové. Menší počet chodidlových č l ánkov je výsledkom regenerácie, ak bola noha poškodená
alebo prišlo k jej strate počas nymfálneho vývoja. Prvý chodidlový č lánok býva väčš í ako
ostatné a označuj e sa ako basitarzus alebo metatarzus. Koncový piaty chodidlový č l ánok nesie pár
viac alebo menej zahnutých pazúri kov, medzi ktorými je alebo nie je prítomné arólium. Pazúriky
(unguiculi) sú buď symetrické - rovnako dlhé alebo asymetrické- jeden je skrátený. Len vel'mi
zriedkavo pazúriky chýbajú (rod Nymphytria) alebo je vyvinutý iba jeden (rod Mononychoblatta).
Vnútorná strana pazúrikov môže byť hladká alebo so zúbkami . Chodidlové články (tarsomerae)
13
1 - 4 majú obyčajne na ventrálnej posteriornej strane vankúšikovité orgány- plantulae (u švábov
niekedy nesprávne označované aj pulvili). Plantulae a aróliá pomáhajú pri pohybe na hladkých
alebo strmých povrchoch. U kavernikolných foriem sú často redukované alebo chýbajú, ale môžu
chýbať aj u epigeických foriem.
Krídla (alae) (obr. 5) sú na stredo- a zadohrudi rôzne utvárané (heteronómne). U druhov
s plne vyvinutými letovými orgánmi sú krídla prvého páru (tegminy, mezotorakálne krídla, alae
anticae) obyčajne viac alebo menej sklerotizované, kožovité. Ich úlohou je predovšetkým chrániť
blanité zadné krídla a bruško. Iba u niekoľkých druhov majú oba páry krídiel rovnakú štruktúru
(napr. Cardacus willeyi). Tvar a rozmery tegmín môžu byť často u rôznych pohlaví odlišné, pričom
je badateľná tendencia k ich redukcii. Skrátenie alebo úplnú stratu tegmín (ale aj krídiel) môžeme
R+M Cu A ab Se c
vt Ju CuP+A1
CuA M R Se c
Obr. 5. Ectobius sylvestris ô - tegmina a zadné krídlo. C - kostálna žilka, Se - subkostálna žilka, R- radiálna žilka,
M - mediálna žilka, Cu - kubitálna žilka, CuA -predná kubitálna žilka, CuP- zadná kubitálna žilka, A -análne žilky,
A1 - prvá análna žilka, A2 - druhá análna žilka, Ju- jugálne žilky, ab- análna brázda, vt- vsunutý troj uholník. Orig.
Fig. 5. Ectobius sylvestris ô - tegmen and hind wing. C -costa, Se - subcosta, R - radius, M - media, Cu - cubitus,
CuA- cubitus anterior, CuP- cubitus posterior, A- ana! veins, A1- first ana! vein, A2 - second anal vein, Ju- jugal veins,
ab - ana] furrow, vt- intercalated triangle. Orig.
14
pozorovať najmä u sami či ek, vďaka čomu je u švábov bežný veľký pohl avný dimorfizmus (napr.
tegminy sami č iek rodov Escala a Robshelfordia (Biattellidae) a Blatta (Biattidae) sú redukované
na okrajové l a l ôč iky a zadné krídla chýbajú; u Laxta granicollis (Biaberidae) a Arenivaga bolliana
(Polyphagidae) sú sami čky bezkrídle; samčekovi a týchto druhov majú úplne vyvinuté tegminy aj
krídl a). Zriedkavejšie dochádza u oboch pohlaví ku skráteniu (napr. Loboptera) alebo úplnému
vymi znutiu (napr. Cryptocercus, Gromphadorhina) tegmín i krídiel. Nie je známy žiadny prípad,
že by redukcia tegmín nekorelovala s redukciou krídiel. Tegminy nie sú použitel'né na aktívn y let,
môžu slúžiť nanajvýš na plachtenie. Žilnatina je zvyčajne vel'mi hustá, dobre vyvinutá. Ak sú
tegminy kožovité, jednotlivé žilky sú dobre viditel'né (napr. Ectobius), ak sú silne sklerotizované,
žilnatina sa stáva nezreteľnou alebo sa úplne stráca (napr. Phyllodromica). Časť jednej z tegmín,
ktorá je pri odpoč inku zakrytá, je obyčajne viac blanitá alebo prinajmenšom menej kožovitá ako
zvyšok tegminy. V prevažnej väčš in e je to pravá tegmina. V niektorých prípadoch môže byť bazálna
časť tegmín nepriehl'adná, ochlpená, bez viditel'ných žiliek a vrcholová časť je priehJ'adn á so
zretel'nými žilkami (napr. Holocompsa nitidula z tropickej Ameriky).
Kostálna žilka (C) tvorí okraj tegmín, subkosta (Se) je krátka, radi álna žilka (R) o byčajn e
s viacerými prednými hrebeiíovitými vetvami. M edi álna (M ) a kubitálna (Cu) žilka zaberajú vel'kú
čas ť tegmín. Krátka, zahnutá postkubi tálna žilka (CuP; PCu) odde ľuj e zretel'ne utvárané análne
pole - clavus. An álna brázda (sulcus analis; línia prechodu žiliek C uP+ A 1
) je na rozdi el od rovnokrídlovcov
oblúkovitá. V análnom poli je výrazne vyvinutá iba druhá análna žilka (A2) a jej vetvy.
Jugálne žilky (Ju) sú vel'mi slabé a nejasné. (V zátvorkách sú uvedené skratky názvov žiliek. )
Zadné (metatorakálne) krídl a (alae posticae) sú väčš in ou blanité, prispôsobené lietaniu. Zriedkavo
môžu byť zhrubnuté. Podl'a ich tvaru rozoznávame 2 hlavné typy. Prvý, polyfagoidný typ má
veľké preaxilárne (remígiové) pole a malé análno-jugálne pole. Druhý typ sa vyskytuje u všetkých
ostatných švábov a je charakteri stický relatívne vel'kým preaxilárnym poJ'om a veJ'mi vel"kým análno-jugálnym
poJ'om. Počas odpoč inku je análne pole zložené v mieste medzi CuP a A1
• A1
(prvá
análna žilka) je atrofovaná a zliata s CuP. U niektorých druhov švábov je na konci tejto žilky vyvinutý
menší alebo väčš í tzv. vsunutý trojuholníK: (tri angulum intercalare). Análno-jugálne pole je od
zvyšku krídla oddelené análnym záhybom (plica ana]is). Z análnych žil iek je dobre vyvinutá iba
A2, aj to nanajvýš len s krátkymi vetvi č kami. Jugálne žilky sú uspori adané radiálne. V pokoji je
análno-jugálne pole obyčajne naskl adané do tvaru vejára a zložené pod predanálnu obl asť krídla.
Poskl adané zadné krídl a sú obyčajn e kratšie ako tegminy a sú úplne zakryté. Zri edkavo sú krídla
výrazne dlhšie ako tegminy. V tom prípade sú v pokoji bud"v strede zložené a celé prekryté tegminami
(napr. Diploptera), alebo nie sú zložené a vzadu vytŕčajú spod tegmín (napr. Euthyrrhapha).
Žilnatin a zadných hídi el je taxonomicky dôležitá. Kubitálna žilka je, alebo nie je vetvená. Ak
je vetvená, môže mať rôzny počet vetiev, ktoré sú buď úplné (t. j. dosahuj ú vrchol krídl a), alebo
neúplné (nedosahuj ú vrchol). Druh môže m ať aj brachypterné aj makropterné fo rmy. Niek torí
taxonómovi a kl adú vel'ký dôraz na redukciu letových orgánov a považujú ju za zákl ad ný rodový
alebo druhový znak.
Bruško (abdomen) (obr. 6, 7) je silne dorzoventrálne sploštené. Pôvodne sa skl adalo z ll
č l ánkov. U súčas n ých druhov je zretel'ných desať tergov (T l - T l0), jedenásty (epiprokt) je spl ynutý
so supraanálnou pl atni č kou (Jamina supraanalis = T l0). Preto sa lalokovitý výbežok supraanálnej
pl atni č ky vyskytujúci sa u niektorých švábov niekedy ozn ač uj e ako epiprokt. Tergá T8 a T9 sú
u oboch pohlaví malé a sú skryté pod T7. Sami čky mn ohých druhov majú vyvinuté rôzne abdominálne
žľazy používané pri obrane alebo na prilákanie samčeka ( R OTH 1969; R OTH & ALSOP 1978;
BRossuT & R OTH 1977). Tergálne ž ľazy používané pri sexuálnom správaní sa vys kytuj ú len
u sam čekov. Vyluč ujú feromón atraktívny pre sami č ky. Ich príto mn osť alebo neprítom nost a ich
umiestne nie a tvar sú taxonomicky dôležité. Tergálne žl'azy vytvárajú často preli ačeninu v strede
terga. U podčel'ade Blattellinae sú často na 5. alebo 7. tergu, ale môžu byť na ktoromko J'vek tergu.
U sam čekov rodu Ectobius a Phyllodromica je tergálna žJ'aza vždy na 7. tergu.
15
A
Obr. 6. Ectobius lapponicus o - bruško. A - dorzálny pohl"ad; B - ventrálny pohl"ad. a - vyústenie tergálnej žl"azy,
b - cerkus, c - stilus. Orig.
Fig. 6. Eclobius lapponicus o - terminal segments of abdomen. A - dorsal view; B - ventral view. a - tergal gland,
b- cercus, c- stylus. Orig.
Obr. 7. Eclobius lapponicus S' -bruško. A- dorzálny pohl"ad; B - ventrálny pohl"ad. a- subgenitálna pl atni čka (S7),
b - cerkus. Orig.
Fig. 7. Eclobius lapponicus S' - terminal segments of abdomen. A - dorsal view; B- ventral view. a - subgenital plate
(S7), b - cercus. Orig.
Prvé sternumje malé alebo chýba, siedme sternum u samičiek a deviate u samčekov je predÍžené
a vytvára subgenitálnu platničku (Jamina subanalis). Jedenáste sternum je rozdelené na
paraprokty, ktoré sú ventrálne kryté subgenitálnou pl atni čkou. Z tohto č l ánku vyrastajú i cerkusy.
Cerkusy (cerci) sú väčšinou krátke, niekoľkočlánkové, u rodu Panesthia sú j ednoč l ánkové.
16
Výnimočne môžu byť úzke a dlhé - u Periplaneta americana až viac ako 20-článkové . Zo
spodnej strany sú zvyčajne pokryté množstvom rôznych hmatových , čuchových a sluchových
senzíl.
Spirákulá 2.-7. článku sú otvorené na pleurálnej membráne, na l. a 8. článku sú pripojené
k bočnému okraju tergov.
Na samčej subgenitálnej platničke (S9 = hypandrium) sú obyčajne dva stilusy (styli), ktoré
môžu byť špecificky rozdielne. Samčekovia rodov Ectoneura, Stenectoneura a druhu Richanitschia
luteomarginata (Biattellidae) majú iba jeden stilus. U všetkých zástupcov podčeľade Panesthiinae
(Biaberidae) a u viacerých rodov čeľade Blattellidae (Arawakina , Neoloboptera, Loboptera,
Astyloblatta, Astylella, Caffroblatta, Jacobsonina, Parascalida, Phymatosilpha a Pseudoceratinoptera)
nemajú samčekovia stilusy vôbec vyvinuté (ROTH 1977, 1989, 1993). Niektoré
druhy (napr. Shelfordina orchidae a niektoré ďalšie druhy tohto rodu) majú výbežok vznikajúci pri
báze každého stilusu, takže sa zdá, akoby mali 4 stilusy (ROTH 1990). U dospelých samičiek nie sú
stilusy nikdy vyvinuté.
Vonkajšie genitálie (organa genitalia externa). Samčie genitálie (falické orgány) (obr. 8) sa
skladajú zo skupiny asymetrických sklerotizovaných falomér, medzi ktorými leží genitálny otvor
(gonoporus). Faloméry (phallomerae) sú skryté v blanitom genitálnom vaku ležiacom nad deviatym
sternom a pred paraproktami. Často poskytujú vynikajúce druhové znaky. U čeľade Blattidae
a Polyphagidae sú faloméry zložité, u čeľade Blattellidae a Blaberidae sú jednoduchšie (McKITTRICK
1964). Homológiu samčích genitálnych skleritov študovali MIZUKUBO & HIRASHJMA
( 1987) a BoHN (1987). Falické orgány druhotne vyrastajú z prednej blanitej steny genitálneho
vaku. Vyskytujú sa v niekoľkých odlišných typoch. Primárne sa skladajú z 3 lalokov alebo falomér,
ktoré sú rôzne obkľúčené skleritmi, z ktorých niektoré majú tvar tŕňov, hákov a gombíkov.
Pravá a ľavá faloméra je prítomná u všetkých švábov. Tretia, ventrálna faloméra, ktorá leží pod
otvorom semenometu (ductus ejaculatorius), čiže pod falotrémou (phallotreme), je prítomná len
u Blattidae a Polyphagidae. U niektorých Blattellidae je posteriorným pokračovaním semenometu
kónický blanitý penis (phallus) medzi pravou a ľavou falomérou. Sklerity sa pre lepšiu orientáciu
číslujú v smere od dorzálnych k ventrálnym, na pravej falomére ako Rl, R2, R3 a mediálne až
laterálne, na ľavej falomére ako L l, L2, L3 . Výnimočne u Polyphagidae existuje i prídavný sklerit
L4 (SNODGRASS 1937; McKnTRICK 1964). Funkciu gonopod prevzali stilusy.
Samičie vonkajšie genitálie (obr. 9) sú usporiadané symetricky. Subgenitálna platnička
(7. sternum- S7) je zväčšená, ohnutá a spolu s redukovanými a čiastočne blanitými S8, S9 a S lO
je pohltená do stien veľkého genitálneho átria. Genitálny vak (atrium) je rozdelený do dvoch široko
spojených komôrok - veľké posteriorné vestibul um, kde je vytváraná ootéka a menšia anteriorná
kopulačná komôrka (bursa copulatrix) s otvorom spoločného vajíčkovodu a otvorom spermatéky.
Strecha vestibulaje zložená z pomerne pevného rámu skleritov podopierajúcich tri páry ovipozičných
valvúl. Podlaha, hoci sa skladá čiastočne zo stredového vestibulárneho skleritu, je väčšinou
blanitá. Podlaha kopulačnej komôrky skladajúca sa zo sklerotizovanej laterosternálnej plošiny je
trocha vyvýšená nad podlahu vestibula. Strecha kopulačnej komôrky je primárne stavaná z troch
zliatych skleritov, ktoré sú anteriorne pripojené k laterosternálnej plošine. Skrytý, vnútorný ovipozitor
je zložený z troch párov malých, prstovitých valvúl vo vnútri komôrky. Prvý pár valviferov
leží vždy bočne od báz prvého páru valvúl a je s nimi zliaty alebo spojený. Druhý pár valviferov
leží hneď za prvým párom valviferov. Je zliaty s kruhovitou štruktúrou, ktorá je zložená z predného
oblúka, podporujúceho druhý, vnútorný pár valvúl a z páru zadných lalokov, ktoré podporujú
bázy tretieho páru valvúl. Prvý (ventrálny) pár valvúl má ontogenetický pôvod v 8. sterne a druhý
a tretí pár (bočné a vnútorné valvuly ovipozitora) sa vytvárajú z deviateho sterna (NEL 1929).
Malá vyčnievajúca plošina (centrálna apodéma) vzniká medzi bázami druhého a tretieho páru valvúl.
Medzi bázami druhého páru valvúl ústia dve kolateriálne žľazy a malý vestibulárny orgán.
Valvuly a valvifery sú držané na mieste pomocou páru dlhých, úzkych apodém, ktoré sú natiahnuté
17
A
ht'li'i:'+- f
~~,--g
k:.;:\·&.-h
B
11-----lt----.----- - b
11--- - - - c
Obr. 8. Samčie vonkajšie genitálie. A, B - subgenitálna pl atni čka s falomérami , hák je č iastočn e odsunut ý;
C, D - posteriorný koniec háku. A - Phyllodromica maculalll maculata; B - Ectobius lapponicus; C - Phyllodromica
hungarica; D - Phyllodromica megerlei. a - endofalická apodéma ravej faloméry (L2vm - virga), b - ravá anteriorná
apodéma, c- pravá anteriorná apodéma, d- hák (L3), e- pravá faloméra R2, f- helmet sklerit, g- pravá faloméra R3,
h - subgenitálna platnička, i - stilus. Orig.
Fig. 8. Male external genitalia. A, B - subgenital plate with phallomeres, hook is partly displace; C, D - posterior end of
hook. A - Phyllodromica maculata maculata; B - Ectobius lapponicu.ľ; C - Phyllodromica lumgarica; D- Phyllodromica
megerlei. a - endophalic apodeme of left phallomere (L2vm - virga), b - left anterior apodeme, c - right anterior
apodeme, d - hook (L3), e - right phallomere R2, f- helmet sclerite, g - right phallomere R3, h - subgenital plate,
i - stylus. Orig.
18
Obr. 9. Samičie vonkajšie genitálie u švába Phyllodromica macu/ata mantlata. a- rameno prvého valviferu, b - ventrálna
ča~ť k.ladielka, c - bazivalvula, d- zadný lalok druhého valviferu, e - laterosternit, f- paratergit, g- val.vula, h - siedme
sternum (subgenitálna platnička), i - paraprokt, j - cerkus. Orig.
Fig. 9. Female external genitalia of Phyllodromica macu/ata macu/ata. a- first valvifer arm, b- ventral pan of ovipositor,
c- basivalvula, d- hind lobus of valvifere II, e- laterosternite, f- paratergite, g- valve, h- sternit 7 (subgenital plate),
i - paraproct, j - cereus. Orig.
19
anteroventrálne z bočných okrajov ôsmeho a deviateho terga. Tieto hrebeňovité apodémy sú zložené
zo zliatych predÍžených paratergitov článkov VIII a IX. Paratergity IX sú často dlhšie ako
paratergity VIII. Stredné konce paratergitov VIII sú spojené s prvým párom valviferov. Samičie
genitálie sú u takmer všetkých švábov trochu asymetrické, stupeií asymetrie varíruje od druhu
k druhu (MCK!TTR!CK 1964).
2. Anatomická charakteristika imág
Dýchacia sústava (systema respiratorium). Vzdušnicová (tracheálna) sústava švábov je holopneustická.
S vonkajším prostredím je spojená s desiatimi pármi spirákul- 2 hrudnými a 8 abdominálnymi.
Cez spirákulá vstupuje vzduch do vzdušnicovej sústavy švábov. Spirákulá sú pripojené
k trom párom veľkých paralelných tracheálnych kmeňov spojených pomocou prekrížených komisúr.
Vzdušnice (tracheae) a jemnejšie vzdušničky (tracheolae) respiračného systému švábov tvoria
rozvetvenú sieť rúrok ležiacu vo vnútri hemocelu. Rozvetvujú sa po celom tukovom telese, prenikajú
tkanivá telových stien a vnútorností a privádzaj ú kyslík do všetkých častí tela švábov. Keďže
na hlave nie sú spirákulá, vzdušnica zásobujúca prednú časť tela švábov je tvorená vetvami pochádzajúcimi
z hrude.
Obehová sústava (systema circulatorium). Primárnou funkciou cievnej sústavy je zásobovať
vnútorné orgány a tkanivá hemolymfou. Hemolymfa privádza produkty trávenia a odoberá odpadový
materiál z metabolizmu do exkrečných orgánov. Hemolymfa tvorí tiež médium pre cirkuláciu
hormónov produkovaných neuroendokrinnými orgánmi. U hmyzu je tzv. otvorený systém cirkulácie
hemolymfy. Cirkulujúca tekutina sa pohybuje voľne v telovej dutine (haemocoel). Pohyb
hemolymfy zabezpečuje pulzujúca chrbtová cieva (vas dorsale), ktorej zadný koniec je uzavretý
a predný koniec je otvorený. Chrbtová cieva leží pod dorzálnym povrchom integumentu, jej predná
časť (srdcovnica- aorta) je umiestnená v hrudi a zadná časť (srdce- cor) v prvých deviatych abdominálnych
článkoch. Upevnená je (u Periplaneta) pomocou 12 párov krídlových svalov (musculi
alares). Má tvar priamej rúrky, rozdelenej na srdcové komôrky. Hemolymfa z peri kardiálneho sinusu
vstupuje pri diastole do srdca cez 12 párov otvorov (ústie- ostium). Tri párové laterálne
otvory sú v hrudnej časti a deväť v abdominálnej časti. Zo srdca vychádza šesť párov ciev - 2
hrudné (mezo- a metatorakálne) a 4 abdominálne (v článkoch 3. - 6.). Tie vedú hemo lymťu zo
srdca do okrajových častí tela.
Okrem chrbtovej cievy existujú rôzne prídavné cirkulačné orgány, ktoré zásobujú hemolymťou
dlhšie telové výbežky. U švábaPeriplanetaamericanaje známe tzv. tykadlové srdce umiestnené v hlave
pred mozgom. Skladá sa z dvoch pri báze tykadiel ležiacich baniek (ampullae) spojených s rytmicky
sa sťahujúcim priečnym svalom. Z baniek vychádzajú tykadlové cievy do tykadiel. Počas relaxácie
priečneho svalu tlačia elastické banky hemolymfu do tykadiel (HERTEL & PENZL!N 1992).
Hemolymfa švábov je číra tekutina s vysokým obsahom aminokyselín a dioxidov. Nemá
funkciu nosiča kyslíka, a preto neobsahuje krvné farbivá - pigmenty. Hemolymfu tvorí tekutá
plazma a množstvo rôznych typov buniek - hemocytov.
Tráviaca sústava (systema digestorium). Tráviaca rúra (tractus intestinalis) zač ína ústnou
dutinou (cavum oris) a končí konečníkom (rectum). Šváby sú všežravé (omnivorné), a preto nie sú
ich ústne ústroje špecializované (podrobnejší opis pozri Morfologická charakteristika). Samotná
tráviaca rúra je točitá a asi dvakrát dlhšia ako telo. Zvyčajne rozlišujeme tri dobre odlíšiteľné
časti - predné, stredné a zadné črevo. Predné a zadné črevo majú ektodermálny pôvod, stredné
črevo má endodermálny pôvod. Črevo je dobre zásobované vzdušnicami, ktoré ho držia v telovej
dutine medzi lalokmi tukového telesa. Svalové steny predného čreva sú inervované sympatickým
nervovým systémom a zadné črevo nervami z terminálneho abdominálneho ganglia (CORNWELL
1968).
20
Predné črevo (stomodeum) sa začína ústnou dutinou (cavum oris). V jej hornej časti je hypofarynx
a epifarynx. Odtiaľ potrava prechádza do dolnej časti - do slinovníka (salivarium) a po
premiešaní slinami prechádza do hltanu. Hltan (pharynx) je vybavený rozťahovacími svalmi umožňujúcimi
jeho značné zväčšenie (veľká žravosť je pre šváby typická). Ďalej prechádza potrava cez
pažerák (oesophagus) a dobre vyvinutý hrvoľ (ing!uvies) do puchorčeka (proventriculus). Puchorček
sa skladá vpredu z armária a vzadu zo stomodeálnych valvúl. Armárium je charakteristické 12
veľmi komplikovanými, špecializovanými kutikulárnymi záhybmi. Každý záhyb má vpredu sklerotizované
zúbkovanie, v strede je vankúšik pokrytý chÍpkami a vzadu chlopnička. Zúbkovanie
a vankúšiky sú u švábov druhovo špecifické. S výnimkou druhov čeľade Blaberidae je zúbkovanie
veľké alebo stredne veľké, u čeľade Blaberidae je veľmi malé. Prechod medzi puchorčekom
a stredným črevom tvorí chlopnička predného čreva (valvula cardiaca). Táto chlopničkaje vsunutá
do stredného čreva a tvorí prechod medzi ektodermálnou a endodermálnou časťou čreva .
Stredné črevo (mesenteron) tvorí žalúdok (ventriculus). Vpredu má 8 slepých výbežkov (coeca).
Na rozdiel od predného a zadného čreva je hemocélový povrch stredného čreva zásobovaný
viscerálnymi vzdušnicami idúcimi od všetkých párov abdominálnych spirákul. Stredné črevo
nemá kutikulárny lem. Jeho epitelový lem je zložený zo sekrečných a absorbčných buniek. Zadná
časť stredného čreva je lemovaná Malpighiho rúrkami (vasa Malpighii; tvoria vylučovaciu sústavu).
Je ich okolo 60-100 a sú usporiadané v skupinách po 6 (Biattidae), po 4 (Blattellidae) alebo
po 3 (Biaberidae) (LECONTE et al. 1967). Zúčastňujú sa na regulácii iónovej rovnováhy a obsahu
vody v hemolymfe. U švábov z rodu Periplaneta rúrky obsahujú intracelulárne enzýmy .
Zadné črevo (proctodeum) má za úlohu odčerpávať vodu zo strávenej potravy. Výsledkom sú
suché granule trusu. Zadné črevo je rozdelené na tri časti- tenké črevo (ileum), hrubé črevo (co!on)
a konečník (rectum). Tenké črevo je najkratšou časťou čreva. Hrubé črevo sa od ostatných častí
ľahko rozozná podra tmavej farby jeho obsahu. Na konečníku je 6 konečníkových papíl (papillae
recti) (CoRNWELL 1968; OscHMAN & WALE 1969).
Slinné žľazy sú dobre vyvinuté, ležia v hrudi po oboch stranách pažeráka (oesophagus). Majú
zvláštne vývody spojené do spoločného vývodu vyúsťujúceho v záhybe medzi hypofaringom
a spodnou perou. Na každej strane je jeden veľký rezervoár.
Vylučovacie orgány (organa uropoetica). Vylučovanie je regulačný mechanizmus, pomocou
ktorého sa udržuje množstvo dusíkatých látok, anorganických solí a vody v hemolymfe
v uspokojivej rovnováhe a takto zabezpečuje stabilnú skladbu iónov a osmotický tlak. Konečným
produktom metabolizmu je dusík. Jeho odstránenie v podobe kyseliny močovej je primárnou funkciou
hmyzieho vylučovacieho systému. U švábov existujú štyri miesta zabezpečujúce reguláciu
vylučovania: l. Malpighiho rúrky, ktoré v spojení so zadným črevom sú zodpovedné za elimináciu
odpadov cez konečník (anus), 2. určité bunky tukového telesa, ktoré sú schopné zadržiavať dusík
v procese označovanom ako "vylučovanie skladovaním", 3. močové žľazy, špeciálne rúrky prídavných
žliaz samčekov určitých druhov švábov, ktoré odstraňujú uráty (močovinu) v spojení so
spermatoťórom počas kopulácie a 4. kutikula, do ktorej môže byť odkladaný odpadový materiál ,
ktorý je následne počas zvliekania eliminovaný. Nie je možné povedať, ktoré z týchto miest vylučovania
je u švábov primárne zodpovedné za reguláciu odpadov. Ak uvážime, že v Malpighiho
rúrkach švábov nebola zistená kyselina močová ako základný konečný produkt exkrécie hmyzu,
tak potom by veľkú úlohu pri regulácii odpadov mohlo hrať tukové teleso.
Tukové teleso (corpus adiposum) je u švábov silne vyvinuté, uložené je predovšetkým
v abdominálnej oblasti. Vonkajšiu vrstvu tukového telesa tvoria dva typy buniek- troťocyty (tukové
bunky) a urocyty (urátové bunky). Troťocyty sú zapojené do syntézy, skladovania a mobilizácie
tukov a bielkovín, zatiaľ čo funkciou urocytov je zhromažďovanie urátov (kyselina močová a jej
soli). Samičky švábov inkorporujú počas oogenézy kyselinu močovú do svojich ooték, kde je za
pomoci bakteriocytov využitá počas embryogenézy. Bakteriocyty (mycetocyty alebo všeobecne
symbiocyty) tvoria vnútornú časť tukového telesa. Obsahujú špeciálne intracelulárne bakteriálne
21
symbionty nazývané bakteroidy (Blattabacterium cuenoti), ktoré sú obalené vakuolárnou membránou
produkovanou eukaryotickými bunkami hostitel'a (BIGLIARDI et al. 1989). Bakteroidy sú
vel'mi dôležitou skupinou mikroorganizmov pravdepodobne všeobecne rozšírenou medzi švábmi.
Doteraz boli zistené v desiatkach druhov švábov z mnohých rodov. Švábom poskytujú rôzne výživné
látky a tak sa spolupodiel'ajú na ich normálnej výžive- recyklujú dusíkaté odpadové látky
uložené v urátových bunkách. Odstránenie bakteroidov pomocou antibiotík spôsobuje zmeny
v správaní, predÍženie vývoja, zvýšenie mortality a neschopnosť reprodukcie. Bakteroidy prenikajú
i do vajíčkových buniek a transovariálne sa dostávajú i do embryí švábov.
Nervová sústava (systema nervosum). Nervová sústava švábov zodpovedá typicky hmyziemu
vzoru nervovej sústavy. Skladá sa z troch sústav. Najznámejšia je centrálna nervová sústava
(somatická sústava), ktorú tvorí mozg, ventrálna nervová páska a jej gangliá (nervové uzly). Ďalšie
sú periférna (obvodová) a sympatická (viscerálna) nervová sústava.
Mozog (nadhltanový nervový uzol - ganglion supraoesophagale) leží nad pažerákom. Vznikol
splynutím 3 hlavových ganglií a je rozdelený na tri časti. Predný mozog (protocerebrum), reprezentujúci
splynutý pár ganglií optického článku (prozocefalónu), je rozdelený do dvoch hemisfér
so zrakovými lal ôčikmi (lobi optici) po stranách. Stredný mozog (deutocerebrum), reprezentujúci
splynuté gangliá tykadlového č lánku (deutocefalónu), sa skladá z č uchovýc h l a l ôč ikov (lobi olfactori),
z ktorých vybiehajú tykadlové nervy. Najmenší je zadný mozog (tritocerebrum). Vznikol
z ganglií tretieho hlavového č l ánku (tritocefalónu) a leží boč ne od čreva.
Ventrálna nervová páska je u švábov zložená z lOganglií pospájaných párovými konektívami .
Prvé ganglium (podhltanový nervový uzol- ganglion suboesophagale) vzniklo splynutím ganglií
zvyšných troch hlavových článkov a leží pod pažerákom. S mozgom je spojené pomocou párových
konektív (cirkumezofageálnych konektív) prechádzajúcich cez otvor v tentóriu. Tri gangliá
sú v hrudi (ganglion thoracicum primum, secundum et tertium) a inervujú hrudné svaly (aj svaly
nôh a krídiel). Jedno (prípadne až tri ) z predných abdominálnych ganglií splýva so zadohrudným
gangliom, čo má za následok prítomnosť iba 6 samostatných abdominálnych ganglií (bruškové
nervové uzly - ganglia abdominales). Prvých 5 inervuje telové svalstvo. Šieste, terminálne abdominálne
ganglium je väčšie ako predchádzajúcich 5 ganglií, lebo vzniklo splynutím ganglií viacerých
terminálnych článkov.
Periférna nervová sústava sa skladá z nervov, ktoré vychádzajú z ganglií a inervujú všetky
časti tela.
Sympatická nervová sústava vykazuje tak nervovú, ako aj endokrinnú (hormonálnu) aktivitu
a skladá sa zo senzorických, motorických a asociačných nervov. Tvoria ju tri čas ti: a) stomatogastrická
sústava obsahujúca gangliá a nervy inervujúce prednú časť čreva, b) ventrálny viscerálny
nervový systém- nervy, ktoré vznikli z ventrálnej nervovej pásky, inervujú a spájajú spirákulá a c)
kaudálna (proktodeálna) sústava, ktorá sa skladá z nervov vznikajúcich z terminálneho abdominálneho
ganglia a inervujúca zadnú časť čreva (CoRNWELL 1968). Stomatogastrická nervová sústava je
u švábov dobre vyvinutá. Skladá sa zo 4 ganglií - čel ového ganglia (ganglion frontale), záhlavného
ganglia (ganglion hypocerebrale), ingluviálneho ganglia a páru ganglií ležiacich na povrchu puchorčeka,
z návratného nervu (nervus recurrens) a pažerákového nervu (nervus oesophagealis).
Vnútorné pohlavné orgány (organa genitalia interna). U samčekov švábov sa párové semenníky
(testes) skladajú zo 4 alebo viacerých semenníkových folikulov (folliculi testiculares)
obyčajne uzavretých v peritoneálnom obale. Semenovody (vasa deferentia) smerujú dozadu
a ústia do semenometu (ductus ejaculatorius). Na prednom konci semenovodu je jeden alebo viac
párov semenných váčkov a väčš í počet tubulárnych prídavných žliaz (glandulae accesoriae) rôznej
dÍžky. Tieto žl'azy vylučujú materiál, z ktorého je vytvorený spermatofór. Sú očividne mezodermálneho
pôvodu, u nýmf sa vyvíjajú z ampulky na konci každého semenovodu. U samčekov sa
nachádza aj nepárová "chumáčová žl'aza" rôzneho tvaru, ktorá leží pod prídavnými žl'azami a ústi
oddelene medzi falomérami .
22
Príloha č. 19
KOČÁREK, P., HOLUŠA, J., VIDLIČKA, Ľ. 2005.
Blattaria, Mantodea, Orthoptera & Dermaptera of the Czech and Slovak Republics.
Illustrated key 3.
Blattaria, Mantodea, Orthoptera & Dermaptera České a Slovenské republiky.
Ilustrovaný klíč 3.
Kabourek, Zlín, 349 pp.
ISBN 80-86447-05-7
Petr Kocarek
Jaroslav Holusa
I:ubomir Vidlicka
Blattaria, Mantodea
Orthoptera & Dermaptera
ofthe Czech and Slovak Republics
Ceske a Slovenske republiky
~~KaBOUREK
Zlfn, 2005
Peer reviewers Pavel Pecina, Praha, Czech Republic
t llja Okali, Bratislava, Slovak Republic
Barnabas Nagy, Budapest, Hungary
Gtinter Kohler, Jena, Germany
Sigfried lngrisch. Bad Karlshafen. Germany
Blattaria. Mantodea. Orthoptera & Dermaptera ofthe Czech and Slovak Republics. Illustrated key 3.
60 I figs. incl. 2 1 colour plates. 349 pp.
Blattaria. Mantodea. Orthoptera & Dennaptera Ceske a Slovenske republiky. llustrovany klic 3. 60 I
obr. vc. 2 1 barev. tabulf. 349 str.
© Petr Kocarek, Ostrava, 2005
© Jaroslav Holusa, Frydek-Mistek, 2005
© L',ubomir Yidlicka, Bratislava, 2005
Translation © David Boukal, Ceske Budejovice, 2005
Photographs © Jifi Tronecek, Bysti'ice pod Host)'nem, 2005
Cover design © Vaclav Zalesak, Zlin, 2005
Editor © Vit Kabourek, Zlin, 2005
Publisher © Nakladatelstvi KABOUREK, s.r.o., Sokolska 3923, 760 0 I Zlin
www.kabourek.cz
ISBN 80-86447-05-7
18 Blattaria
The order Blattaria
I..:ubomir Vidlicka
Introduction
Extant cockroaches (Blattaria) are a relatively small insect group encompassing
about 4,000-4,500 species described and classified in approximately 460 genera.
They belong to the exopterygote Neoptera and, together with the orders Grylloblattodea,
Mantodea, Isoptera. Phasmatodea, Orthoptera, Dermaptera, Embioptera
and Zoraptera, to the series Polyneoptera.
Most extant cockroaches inhabit warm forests of the tropics and subtropics. Only
about 2-3% of the known species inhabit the temperate zone. In Europe, cockroaches
only received more attention when the synanthropic species Blatta orienta/is and
Blattel/a germanica entered major European ports with Greek merchant vessels
and unstoppably invaded the entire continent. Various forms of the word "kakerlac",
apparently derived from the Spanish "cucaracha", have been used as vernacular
names for cockroaches. Besides the English name, its equivalents are still used to
denote a cockroach in other countries, such as kakerlac in the Netherlands and
Denmark, kackerlacka in Sweden, kackerlackor in Finland and cackerlac and
cancrelat in France. The Czech and Slovak common name "svab" comes from the
German "Schwab(e)" and "Schabe" . The generic name Blatta, which is also the
stem of the order' s name. is derived from the Greek PJ.mrrco (to harm, be harmful,
injure, damage). Any harmful and cryptic domiciliary insects used to be called by
that name in old Latin literature.
The systematics of cockroaches is rather disparate, although two approaches
are most frequently used at present. The first one developed by Princis ( 1962-197 1)
divides cockroaches into 28 fami lies. The other one recognises 6 families: Polyphagidae,
Blattellidae, Blaberidae, Blattidae, Cryptocercidae and Nocticolidae
(McKittrick, 1964, Roth, 1988). In the Czech and Slovak Republics, one may
find only the native representati ves of the fami ly Ectobi idae (sometimes included
in the family Blattellidae) freely in nature; our domiciliary species belong
to the families Blattidae and Blattellidae. Members of the family Polyphagidae
inhabit more southern and arid parts of the Palaearctic and Nearctic regions. The
family Cryptocercidae has only 5 species distributed in the Palaearctic and earctic
regions; Grandcolas ( 1994) included this family in Polyphagidae. The speciose
family Blaberidae is distributed mainly in the Neotropical, Indomalayan and Ethiopian
regions. Species of a small family Nocticolidae are similarly distributed.
Bey-Bijenko ( 1950) treated cockroaches of the eastern part of the Palaearctic
region, including most central European species in his revision. A key to central
European species was published by Harz ( 1957). Princis (1965) and Harz & Kalten-
Blattaria 19
Rad Blattaria
I..:ubomir Yidlicka
Uvod
Svabi (Blattaria) tvoi'i v soucasnosti relativne malou skupinu hmyzu zahrnuj ici
okolo 4 000-4 500 popsanych druhu v pfiblime 460 rodech. Patri mezi exopterygotni
Neoptera. Spolu s fady Grylloblattodea, Mantodea, Isoptera, Phasmatodea, Orthoptera,
Dermaptera, Embioptera a Zoraptera nalezi do kohorty Polyneoptera.
Pro vetsinu recentnich druhu svabu jsou domovem vlhke a teple lesy tropicke
a subtropicke zony. Mirne pasmo obyvaji jen asi 2-3% znamych druhtr. Y Evrope
se svabi dostali do centra pozornosti az tehdy, kdyz synantropni druhy Blatta
orienta/is a Blattella germanica pronikly i'eck)'mi obchodnimi lodemi do velk)'ch
evropskych pi'istavu a odtud se nezadrzitelne sifily po cele Evrope. Y minulosti
byli svabi oznacovani ruznymi variantami vyrazu ,kakerlac", odvozeneho zrejme
od spane lskeho , cucaracha" . I v soucasnosti se pouzivaji varianty tohoto jmena
na omacovani svabu v rtrmych zemich - kakerlac (Holandsko a Dansko), kackerlacka
(Svedsko), kackerlackor (Finsko), cackerlac, cancrelat (Francie), cockroach
(Anglie). Nemecke vyrazy Schwab(e) a Schabe se staly zakladem pro slovenske i
ceske pojmenovani , svab". Rodove jmeno Blatta, ktere tvori i slovni zaklad nazvu
celeho radu, je odvozeno z reckeho slova ~A<Xmro (skodit, rusit, prekazet). Ve starsich
latinsk)1Ch pisemnostech se timto jmenem oznacoval kazdy skodlivy hmyz zijici
skryte v domacnostech.
Systematika svabCrje znacne nejednotna. V soucasnosti jsou nejpouzivanejsi 2
pi'istupy. Prvni, vytvoi'eny Princisem (1962-1 971), deli svaby na 28 celedi. Podle
druheho se svabi deli na 6 celedi: Polyphagidae, Blattellidae, Blaberidae, Blattidae,
Cryptocercidae a Nocticolidae (McKittrick, I964, Roth, 1988). V pfirode se u nas
vyskytuji pouze zastupci celedi Ectobiidae (nekdy i'azene do celedi Blattellidae).
Nase synantropni druhy jsou z celedi Blattidae a Blattellidae. Zastupci celedi
Polyhagidae ziji v j iznejsich a aridnejsich castech palearktickeho a nearktickeho
regionu. Celed' Cryptocercidae ma pouze 5 druhtl rozsii'enych v palearkticke a
nearkticke oblasti; Grandcolas ( 1994) i'adi tuto celed' k Polyphagidae. Druhove
bohata celed' Blaberidae je rozsii'ena predevsim v neotropicke, indomalajske a
etiopske oblasti. Podobne rozsifeni maji take druhy male celedi Nocticolidae.
Svaby vychodni casti palearkticke oblasti zpracoval Bey-Bijenko ( 1950) a jeho
prace zahrnuje take vetsinu sti'edoevropskych druhu. Klic na urcovani stredoevropskych
druhtt vypracoval Harz ( 1957). Princis ( 1965) a Harz & Kaltenbach
( 1976) pozdeji zpracovali svaby cele Evropy. Podrobnou revizi rodu Ectobius
publikoval Ramme ( 1923, 195 1). Noveji se revizemi nekterych skupin rodu
Phyllodromica zab)'vali Bohn ( I992, I993) a Vidlicka & Maj zlan ( 1997). Celo-
20 Blattaria
bach ( 1976) subsequently treated cockroaches of the entire Europe. A thorough
revision of the genus Ectobius was pubIished by Ramme ( 1923, 195 1). Recent
revisions of some species-groups of the genus Phyllodromica have been carried
out by Bohn (1992. 1993) and Vidlicka & Majzlan (1997). A world catalogue of
cockroaches was published by Princis (1962, 1963, 1964, 1965a, 1966, 1967, 1969.
1971). The biology of the cockroaches was studied by Roth & Willis (1960), Beier
(1961 , 1974), Cornwell ( 1968) and Rust et at. ( 1995).
The first faunistic data from the territory of the Czech Republic and Slovakia
were reported by Seidl ( 1836) and Frivaldszky ( 1868), respectively. Faunistic
data from Slovakia, taken from literature and museum collections, have recently
been summarised by Vidlicka & Majzlan (1992) and those from the Carpathian
Basin by Vidlicka & Sziniky ( 1997). The first key to all species of the former
Czechoslovakia including the Subcarpathian Ukraine was published by Obenberger
(1926a, 1926b). Later on, cockroaches were also treated within the series Key to
the fauna ofCzechoslovakia (Dobsik, 1959b). A key of the genus Phyllodromica
including the recently described species was published by Vidlicka (1994).
Adult morphology
Cockroaches are usually medium-sized to large insects. The smallest species
belong to the genus Nocticola (about 3 mm), and the largest are in the genus
Blaberus (up to 120 mm).
The body is dorsoventrally flattened and often oval. The body surface is usually
smooth and bare, only seldom (especially in desert and steppe species) with a comparatively
dense pubescence. Cuticular projections, spines and a rough sutface are rare.
The head is usually hypognathous, partly visible from above or entirely covered
by the pronotum; it is transversely flattened with typical sutures. The epicranial suture
is not always developed in adults. The frons and the clypeus are large, well developed.
An opening in the central part is characteristic for the tentorium. The composed
eyes are nearly always developed, and composed of a large number of small
ommatidia. They are large, especially in winged species; rarely, they are reduced or
entirely absent, especially in myrmecophilous and cavemicolous species. The ocelli
are developed in some cockroaches; however, they are more often replaced by two
pale, ocelliform spots near the inner eye margins. The antennae are multi-segmented,
often longer than the body. They are filiform and tapering towards the apex, less
frequently thickened in the median part and spindle-shaped. They are articulated to
the head capsule in the middle of the inner eye margin. All antennomeres are usually
covered by short fine hairs. The biting mouthpa1ts consist offive well-separated parts
- the labrum, mandibles, maxillae, hypopharynx and labium. The mandibles are
robust, inegularly quadrangular, strongly sclerotised, and possess a dentate inner part.
Each mandible bears an appendage (prostheca). The maxilla is composed of several
parts. The cardo is short; the stipes is elongate and its inner face often bears a sclerite
276 List of taxa I Seznam taxonu
BLATIARIA
Blattidae
Periplaneta Burmeister, 1838
P americana (Linne. 1758)
P australasiae (Fabricius. 1775)
P brunnea Burmeister. 1838
Blatta Linne. 1758 . ......... . . .
B. orienralis Linne. 1758
Blattellidae
Blattella Caudel, 1903
B. germanica (Linne. 1767)
Supella Shelford. 1911
S. longipalpa (Fabricius, 1798)
= supellectilium Serville. 1839
324 0'
page/str. Figs.fobr.
~ 42/43
-42/43 319
~ 42/43 320-321
-42/43
-42/43
-42/43 322-323
-44/45
~ 44/45 324-325
~ 44/45
-44/45 326-327
Plates I Tabule 1-2
page/str. Figs.fobr.
Ectobiidae
Ectobius Stephens. 1835 . ~ 46/47
£. lapponicus (Linne. 1758)
£. sylves1ris (Poda. 1761)
- 48/49 328-329
~ 46/47 330-332
[£. eryThrOIIO/us eryThronows
(Burr. 1913)] -48/49
[£. ery1hrono1us ater Bazyluk. 1961] -48/49
E. erythronotus nigricans Ramme. 1923 - 48/49
Phyllodromica Fieber. 1853 . -50/51
=Hololampra Saussure, 1864
P dobsiki Chladek. 1996 -54/55
P harzi Chladek. 1977 ~52/53
P hungarica Vidlicka. 1993 ~54/55
P chladeki Harz. 1977 ~50/51
P maculaw macttlata (Schreber, 1781) -52/53
P maculaw schaefferi (Gmelin, 1789) ~52/53
P megerlei Fieber, 1853 ~50/51
= puncw1a Charpentier. 1825
333-334
335-336
337-338
339-340
341-342
343-344
345-346
330 0' 333 0'
335 0' 336 c;>
343 o 344 c;> 346 c;>