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Syntaxonomic revision of chasmophytic vegetation
of the Centaureo cuspidatae-Portenschlagiellion
ramosissimae (Southeastern Adriatic)
Massimo Terzi & Nenad Jasprica
To cite this article: Massimo Terzi & Nenad Jasprica (2018): Syntaxonomic revision of
chasmophytic vegetation of the Centaureo�cuspidatae-Portenschlagiellion�ramosissimae
(Southeastern Adriatic), Plant Biosystems - An International Journal Dealing with all Aspects of
Plant Biology, DOI: 10.1080/11263504.2018.1549602
To link to this article: https://doi.org/10.1080/11263504.2018.1549602
© 2018 The Author(s). Published by Informa
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Syntaxonomic revision of chasmophytic vegetation of the Centaureo
cuspidatae-Portenschlagiellion ramosissimae (Southeastern Adriatic)
Massimo Terzia
and Nenad Jaspricab
a
Institute of Bioscience and Bioresources, Italian National Council of Research, Bari, Italy; b
Institute for Marine and Coastal Research
University of Dubrovnik, Dubrovnik, Croatia
ABSTRACT
The Centaureo-Portenschlagiellion describes the cliff vegetation of the Southeastern Adriatic. This habitat
type harbours many endemic taxa and has a high value for biodiversity conservation.
Notwithstanding its importance, knowledge of the syntaxonomy of this alliance is still poor. This article
aims at revising the synchorological, coenological and floristic relationships of the associations of the
Centaureo-Portenschlagiellion. The revision is based on a data set of 103 releves of Mediterranean xerothermic
cliffs from Croatia, Bosnia i Herzegovina and Montenegro. The releves were clustered by using
the Flexible beta method. An indicator species analysis was used to identify the diagnostic taxa of the
main clusters of releves and non-metric multidimensional scaling ordination was undertaken to visualize
the floristic relationships among them. Results revealed that only seven associations belong to the
Centaureo-Portenschlagiellion, whereas the others already described in phytosociological literature were
invalidly described or should be treated as syntaxonomic synonyms of the previous ones. Two associations
(Inulo-Centaureetum cuspidatae and Portenschlagiello-Campanuletum portenschlagianae) were
reduced to the rank of subassociations of the Moltkio-Inuletum verbascifoliae. Two other subassociations
were described for the first time. Data provided with this revision may be considered as essential
base-line information that should aid in evaluating the state of this vegetation type in the future.
ARTICLE HISTORY
Received 21 March 2018
Accepted 12 November 2018
KEYWORDS
Asplenietea trichomanis;
Centaureo-Campanuletalia;
cliff vegetation;
phytosociology; rupicolous
communities; syntaxonomy;
the Balkans
Introduction
Limestone cliffs harbour a high number of endemic and rare
plant and animal species. The European Council Directive 92/
43/EEC, the so-called “Habitat Directive”, lists “Calcareous
rocky slopes with chasmophytic vegetation” (habitat code:
8210) among the habitat types constituting the European
protected area within the Natura 2000 network. Although
the Habitat Directive refers to the member states of the
European Union, many neighbouring countries are going to
align their protected area systems with the Natura 2000 criteria
(e.g. Petrovic et al. 2012 for Montenegro; Milanovic
et al. 2015 for Bosnia and Herzegovina).
The Interpretation Manual of the Habitat Directive
(European Commission 2013) provides a general description
for each habitat type, including also syntaxonomical references.
Thus, syntaxonomy is an important source of information
for identifying habitats and coordinating conservation
measures within and beween countries.
The syntaxonomy of xerothermic limestone chasmophytic
vegetation of the Adriatic area was reviewed by Trinajstic
(1980) who described the order “Centaureo-Campanuletalia”,
including two alliances, “Centaureo-Campanulion” and
“Centaureo-Portenschlagiellion”. The latter alliance covers an
elongated area in the central and southern part of the eastern
Adriatic pertaining to Croatia, Bosnia and Herzegovina;
Montenegro and Albania (Jovanovic et al. 1986; Petrovic
et al. 2012; Terzi et al. 2018). Trinajstic (1980, 2008) classified
five associations in the Centaureo-Portenschlagiellion:
Portenschlagiello ramosissimae-Campanuletum portenschlagianae,
Campanulo pyramidalis-Moltkietum petraeae, Inulo verbascifoliae-Centaureetum
cuspidatae, Centaureetum ragusinae
and Seslerio robustae-Putorietum calabricae. The distribution
areas of this alliance include some protected and narrowly
endemic taxa, such as Centaurea cuspidata, C. ragusina, C.
radichii, Fibigia triquetra and Campanula portenschlagiana.
Before this syntaxonomic scheme, Lakusic (1968, 1970)
had classified the limestone cliff vegetation of the eastern
Adriatic in another order, Moltkietalia petraeae, consisting of
two alliances, the Centaureo dalmaticae-Campanulion and the
Edraianthion tenuifolii, including coastal and inland vegetation,
respectively (see also Terzi and Di Pietro 2016). Thus,
several other chasmophytic associations (Fibigio triquetraeCerinthetum
tristis, Micromerio kerneri-Onosmetum dalmaticae,
Moltkio petraeae-Campanuletum lepidae, Moltkio petraeaeCentaureetum
voraginicolae nom. inval. (Art. 3 l of ICPN),
CONTACT Nenad Jasprica nenad.jasprica@unidu.hr Institute for Marine and Coastal Research, University of Dubrovnik, Kneza Damjana Jude 12, P. O. Box
83, Dubrovnik, 20000 Croatia
Supplemental data for this article can be accessed here.
ß 2018 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://creativecommons.org/licenses/by-nc-nd/4.
0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited, and is not altered, transformed, or built upon in
any way.
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https://doi.org/10.1080/11263504.2018.1549602
Published online 31 Dec 2018
Moltkio petraeae-Inuletum verbascifoliae, PuccinellioCentaureetum
crithmifoliae nom. inval. (Art. 2b) and Teucrio
arduinii-Seselietum globiferi) have been described for xerothermic
limestone cliffs occurring in an area potentially falling
within the distribution range of the CentaureoPortenschlagiellion,
but they were classified in the Moltkietalia
petraeae (Lovric and Rac 1987; Lovric and Bedalov 1987;
Lovric and Rac 1989, Lovric et al. 2002).
The two orders, Centaureo-Campanuletalia and Moltkietalia
petraeae, at least in their original descriptions, show some
floristic, coenological and geographical overlaps, especially
along the more thermophilous fringe of the Moltkietalia petraeae
(see Terzi and Di Pietro 2016). The authors of the
abovementioned associations followed one of these two syntaxonomic
concepts, but did not compare or discuss their
findings in the light of the two different possible interpretations,
leaving unresolved the coenological overlaps between
the syntaxonomic schemes proposed by Lakusic (1968) and
Trinajstic (1980).
More recently, Mucina et al. (2016) solved this issue, differentiating
the two orders and restricting the CentaureoCampanuletalia
to the thermo-mesomediterranean chasmophytic
vegetation of limestone cliffs of the Adriatic coastal
regions and leaving the Moltkietalia petraeae for the montane
and alpine belts of the Central and Southern Dinarides.
Within this overall pattern, the coenological and floristic
relationships among the associations of the CentaureoPortenschlagiellion
and those originally classified in the
Moltkietalia petraeae but clearly belonging to the thermomesomediterranean
belt, remain unclear, as well as the diagnostic
species of each of them. For example, the stenoendemic
Campanula portenschlagiana is considered to be a
character species of the Moltkio petraeae-Inuletum verbascifoliae
(Lovric and Bedalov 1987), Campanulo-Moltkietum petraeae
(Horvatic 1963) and also of the Portenschlagiello
ramosissimae-Campanuletum portenschlagianae (Trinajstic
1980). Moreover, this species was also recorded with high
frequency in the Inulo verbascifoliae-Centaureetum cuspidatae
and in the Campanulo pyramidalis-Moltkietum petraeae in the
Biokovo Mountains (compare Trinajstic 1987).
Notwithstanding the importance of this habitat type,
knowledge about the syntaxonomic differentiation of the
associations of the Centaureo-Portenschlagiellion is still poor.
The nomenclature of the two orders, CentaureoCampanuletalia
and Moltkietalia petraeae, was reviewed in
some recent articles through the validation and/or typification
of some syntaxon names (Terzi and Di Pietro 2016,
Jasprica and Terzi 2017, Terzi et al. 2017). Moreover, in a syntaxonomic
revision of the xerothermic chasmophytic vegetation
of the central part of the Mediterranean Basin, Terzi
et al. (2018) defined the syntaxonomic position of the
Centaureo-Portenschlagiellion compared with the other alliances
occurring in the area. The aim of this article is to revise
the synchorological, coenological and floristic features of the
associations of the Centaureo-Portenschlagiellion.
Materials and methods
This revision is based on releves already published in the
phytosociological literature and classified in the syntaxa
listed in Supplementary Appendix S1 (Figure 1). Three releves
on Kolocep Island, originally published by Horvatic (1971),
and assigned to the Seslerio robustae-Putorietum calabricae
were excluded from the data set. In fact, the occurrences of
Anthyllis barba-jovis and Helichrysum italicum indicate a transition
towards the Anthyllidion barbae-jovis (sensu Mucina
et al. 2016).
Two other associations of the Centaureo-Portenschlagiellion
(“Puccinellio-Aurinietum leucadae” and “Aurinio-Brassicetum
frutescentis”) from Vis Archipelago and Palagruza (Middle
Adriatic, Croatia) were also excluded from this revision
because they were invalidly and briefly described without any
suppporting phytosociological table (Art. 2b, Lovric and
Bedalov 1987, Lovric and Rac 1991).
A data matrix (releves x species) was prepared replacing
the original Braun-Blanquet scores with the ordinal value of
the combined scale proposed by van der Maarel (1979). Taxa
indicated only at the genus level were omitted from the
data set as well as mosses and lichens that were not
always recorded.
The plot size is indicated for only 68% of the releves. The
average plot size of releves was 43 m2
with minimum and
maximum values of 1 m2
and 200 m2
, respectively. Since
excessively small or excessively large plot sizes may affect
the outcomes of statistical analyses (Otypkova and Chytry
2006), releves recorded on plots outside the range 10 m2
to
100 m2
were removed. Eventually the data matrix consisted
of 93 releves and 188 taxa. To reduce noise in the dataset
(compare McCune and Grace 2002), rare taxa occurring in
only 1 or 2 releves were removed before analyses.
The releves were clustered by using the Flexible beta
method, with the Bray-Curtis coefficient. The b value was set
at –0.25 because with this value the Flexible beta is a spaceconserving
sorting strategy (McCune and Grace 2002). The
resulting dendrogram was pruned at the level yielding the
Figure 1. The study area with distribution of releves. Abbreviations: SLO:
Slovenia; HR: Croatia; BIH: Bosnia and Herzegovina; MNE: Montenegro;
AL: Albania.
2 M. TERZI AND N. JASPRICA
highest number of indicator species (IndSp, compare
McCune and Grace 2002). Indicator species analysis (ISA,
Dufr^ene and Legendre 1997) was run for the first 20 partitioning
levels of the dendrogram, further partitions dealing
with minor variations. A taxon was considered as IndSp of a
cluster if its indicator value (IndVal) turned out to be higher
for that cluster than for the others of the same partitioning
level, and if it turned out to be significant (p < .05) in a
Monte Carlo test using 10,000 permutations.
IndSp were also used to characterize floristically the clusters
of releves. To increase interpretability of results, each
IndSp was assigned to only one cluster along the descending
hierarchical typology of the dendrogram. In accord with
Dufr^ene and Legendre (1997), the best clustering level for a
species was considered as the one for which the IndVal first
reached its maximum value.
Results were interpreted from a syntaxonomic standpoint
on the basis of the occurrences of the association type-releves
within the clusters and the set of IndSp associated to
each cluster. Diagnostic taxa of the higher syntaxonomic
ranks (alliance, order and class) were deduced from the scientific
literature (compare Terzi et al. 2018).
A non-metric multidimensional scaling (NMDS) ordination
was undertaken to visualize the floristic relationships among
the main clusters of releves. The NMDS was run in the “slow
and thorough” autopilot mode of the PC-ORD (McCune and
Mefford 2011), selecting the Bray-Curtis coefficient as dissimilarity
measure.
All the statistical analyses described above were performed
using the software PC-ORD, version 6.22 (McCune
and Mefford 2011).
For each association, a brief description of its syntaxonomic
and nomenclatural vicissitudes, the holotypus, locus
classicus, distribution range, diagnostic, frequent and dominant
taxa were reported. The locus classicus of a syntaxon
is here defined as the locality from which the original diagnosis
of that syntaxon (or at least its nomenclatural type)
was described. To identify the “dominant taxa,” (i) first we
selected those taxa occurring in at least one releve of the
phytosociological table with a score (on the ordinal scale)
of 5 or more and then (ii) we calculated for each of these
taxa the sum of the scores for all the releves included in
the phytosociological table. Those taxa yielding the highest
values were considered as “dominant taxa.” The diagnostic
taxa were subjectively selected from the IndSp list
(Supplementary Appendix S2).
Results
The dendrogram was pruned at the 10th
partitioning level
that being the partition yielding the highest number of
IndSp. The occurrence of the type-releves of the associations
in the clusters allowed us to assign each of the 10 main clusters
of the dendrogram (Figure 2) to one or more
association.
The first dendrogram division separates the releves of the
Centaureetum ragusinae (Figure 2: cluster 2b) which were further
arranged into two clusters. The first cluster (Crl1) represents
the Centaureetum ragusinae subass. limonietosum
anfracti. The second one (Crt) includes the releves of the
Centaureetum ragusinae typicum and another sub-group,
namely Crl2, which includes releves from Bisevo Island (HR)
representing a third coastal subassociation, C.r. subass. limonietosum
cancellati. Several IndSp are associated with these
clusters, some of them having high IndVal, such as Centaurea
ragusina, Crithmum maritimum, Lotus cytisoides, Limonium dictyophorum
and L. cancellatum. Most of these taxa are ingressive
from the Crithmo-Staticetea class and indicate that most
of the sampled sites were salt-sprayed coastal cliffs.
Four other clusters of releves were characterized by many
IndSp, with high IndVal, and they are floristically well distinguishable
from the rest of the releves (Supplementary
Appendix S2). Three of them (cluster 4a, 4 b and 5b) represent
the following associations: Campanulo austroadriaticaeMoltkietum
petraeae (CaM), Seslerio robustae-Putorietum calabricae
(SrP) and Teucrio arduinii-Seselietum globiferi (TaS). The
fourth one (MkO) includes the type releves of two associations,
namely, Micromerio kerneri-Onosmetum dalmaticae and
Moltkio petraeae-Campanuletum lepidae, together with
another releve of the Moltkio petraeae-Centaureetum voraginicolae.
The latter association was invalidly described since one
of the name-giving taxa (Centaurea voraginicola) had not
been validly published before (or simultaneously with) the
description of the new association (Art. 3l, Weber et al.
2000). These three associations are represented in the data
set by very few releves so that a definitive interpretation of
their floristic composition cannot be expressed. Despite
being grouped in the same cluster, some phytosociologically
important species distinguish the Micromerio kerneriOnosmetum
dalmaticae (e.g. Micromeria kerneri, Seseli tomentosum)
from the Moltkio petraeae-Campanuletum lepidae (e.g.
Figure 2. Clustering of releves. Partitioning levels are indicated by numbers;
the two clusters originated at each partition are labelled as “a” and “b”. The
main clusters represent the following associations: CaM: Campanulo austroadriaticae-Moltkietum
petraeae; MkO: Micromerio kerneri-Onosmetum dalmaticae
and Moltkio petraeae-Campanuletum lepidae; TaS: Teucrio arduinii-Seselietum
globiferi; CpM1/2: Campanulo pyramidalis-Moltkietum petraeae; IvC: Inulo verbascifoliae-Centaureetum
cuspidatae; SrP: Seslerio robustae-Putorietum calabricae;
Crl1: Centaureetum ragusinae limonietosum anfracti; Crt: Centaureetum ragusinae
typicum; Crl2: Centaureetum ragusinae limonietosum cancellati; FtC: Fibigio triquetrae-Cerinthetum
tristis; MpI: Moltkio petraeae-Inuletum verbascifoliae.
PLANT BIOSYSTEMS - AN INTERNATIONAL JOURNAL DEALING WITH ALL ASPECTS OF PLANT BIOLOGY 3
Campanula fenestrellata subsp. fenestrellata, Arenaria orbicularis,
Iris adriatica). Therefore, these associations are provisionally
maintained as independent associations and further
phytosociological data are needed to clarify their syntaxonomic
relationship.
The cluster 5a is differentiated by only one IndSp
(Portenschlagiella ramosissima) and its further subdivisions
highlight two main clusters representing the Campanulo pyramidalis-Moltkietum
petraeae from one side (cluster 6a, CpM1-
2) and four associations (cluster 6b, IvC and PrC) scarcely differentiated
on the other: Inulo verbascifoliae-Centaureetum
cuspidatae, Portenschlagiello ramosissimae-Campanuletum portenschlagianae,
Fibigio triquetrae-Cerinthetum tristis and
Moltkio petraeae-Inuletum verbascifoliae. The last two associations
are represented in the data-set by only one releve, i.e.
their holotypes.
The NMDS ordination resulted in a two axis solution with
a final stress of 19.4, a fairly high value (Figure 3). The ordination
explained only 64.5% of the total proportion of variance,
axes 1 and 2 accounting for 52.0% and 12.5% of
variance, respectively.
Axis 1 roughly represents an altitudinal gradient, with the
Centaureetum ragusinae and the Seslerio robustae-Putorietum
calabricae developing near the coastline at one end, and the
Campanulo austroadriaticae-Moltkietum petraeae, of the
Lovcen Mt. in Montenegro, at the other end. The other associations
lie in the middle part of the diagram with those
recorded along the eastern Adriatic canyons by Lovric and
Rac (1987, 1989) and Lovric et al. (2002) in the lower part of
the diagram. The cluster 6a, representing the Campanulo pyramidalis-Moltkietum
petraeae, shows some overlaps with the
cluster 6b, that is, Inulo verbascifoliae-Centaureetum cuspidatae
(IvC) and Portenschlagiello ramosissimae-Campanuletum portenschlagianae
(PrC). The results of the NMDS ordination confirm
the general relationships among the associations as they
are revealed by the cluster analysis. Online Supplementary
Table SI shows the percentage frequencies of the diagnostic
species for their corresponding vegetation units.
Discussion
Based on the results, the following syntaxonomic scheme is
here proposed:
Class Asplenietea trichomanis (Braun-Blanquet in Meier et
Braun-Blanquet 1934) Oberdorfer 1977
Order Centaureo dalmaticae-Campanuletalia pyramidalis
Trinajstic ex Terzi et Di Pietro 2016
 Alliance Centaureo cuspidatae-Portenschlagiellion ramosissimae
Trinajstic ex Terzi et Di Pietro 2016
– Campanulo pyramidalis-Moltkietum petraeae
Horvatic ex Trinajstic 1964
– Centaureetum ragusinae Horvat ex Terzi, Jasprica et
Cakovic 2017
C.r. typicum Terzi, Jasprica et Cakovic 2017
C.r. limonietosum anfracti Hecimovic ex Terzi, Jasprica et
Cakovic 2017
C.r. limonietosum cancellati subass. nov. hoc loco
– Micromerio kerneri-Onosmetum dalmaticae Lovric ex
Terzi, Jasprica et Cakovic 2017 [syn. Moltkio petraeaeCentaureetum
voraginicolae Lovric nom. inval. (3l)]
– Moltkio petraeae-Campanuletum lepidae Lovric ex
Terzi, Jasprica et Cakovic 2017
– Moltkio petraeae-Inuletum verbascifoliae
Wendelberger in Lovric et Rac 1987 [syn. Fibigio triquetrae-Cerinthetum
tristis Lovric in Lovric et Rac 1987,
Inulo verbascifoliae-Centaureetum cuspidatae Trinajstic
ex Terzi et Di Pietro 2016, Portenschlagiello ramosissimae-Campanuletum
portenschlagianae Jasprica et
Terzi 2017]
M.p.-In.v. typicum subass. nov. hoc loco [holotypus rel.
J1, tab. 6, p. 126, Lovric and Rac (1987)]
M.p.-In.v. centaureetosum cuspidatae (Terzi et Di Pietro
2016) stat. nov. hoc loco [¼Inulo verbascifoliaeCentaureetum
cuspidatae Trinajstic ex Terzi et Di Pietro
2016: 1367]
M.p.-In.v. fibigietosum triquetrae (Jasprica et Terzi 2017)
stat. nov. hoc loco [¼Portenschlagiello ramosissimaeCampanuletum
portenschlagianae Trinajstic ex Jasprica et
Terzi 2017: 332]
– Seslerio robustae-Putorietum calabricae Horvatic ex
Birac 1973
– Teucrio arduinii-Seselietum globiferi Lovric, Rac et
Solic ex Terzi, Jasprica et Cakovic 2017
Order Moltkietalia petraeae Lakusic 1968
 Alliance Edraianthion tenuifolii Lakusic 1968
– Campanulo austroadriaticae-Moltkietum petraeae
(Tomic-Stankovic ex Terzi, Jasprica et Cakovic 2017)
Terzi, Jasprica, Cakovic et Di Pietro 2018
Figure 3. NMDS ordination of releves. Abbreviations are the same used in the
dendrogram (Figure 2).
4 M. TERZI AND N. JASPRICA
Campanulo pyramidalis-Moltkietum petraeae
This association was invalidly described by Horvatic (1963)
for the Mediterranean mountain belt (see also Trinajstic
1998) of Central and Southern Dalmatia, Croatia. It was validated
by Trinajstic (1964) with an original diagnosis consisting
of only one releve recorded on Korcula Island at nearly
300 m a.s.l. in the eu-Mediterranean belt. Our results showed
that the Campanulo pyramidalis-Moltkietum petraeae is widely
distributed, from the Southern Dalmatian islands, such as
Korcula and Mljet, where the association turned out to be
distinguished by some xerothermic species (i.e. Erica manipuliflora,
Brachypodium retusum), up to Mt. Biokovo where a
Mediterranean mountain variant is identified. No character
taxa were found for this association which is distinguished
only by three transgressive taxa from alliance and order
(Campanula pyramidalis, Moltkia petraea and Portenschlagiella
ramosissima).
Holotypus: releve p. 158, in Trinajstic (1964); Locus classicus:
Korcula Island; Distribution area: Central and Southern
Dalmatia; Differential taxa of the eu-mediterranean variant:
Erica manipuliflora, Brachypodium retusum; Frequent and
dominant taxa: Inula verbascifolia, Moltkia petraea,
Brachypodium retusum, Campanula pyramidalis and Asplenium
ceterach subsp. ceterach (the latter is frequent but
not dominant).
Centaureetum ragusinae
This association was invalidly described by Horvat (1942:
464) and subsequently renamed as Phagnalo-Centaureetum
ragusinae by Horvatic (1963). Hecimovic (1984) described the
halophytic subassociations (C.r. limonietosum anfracti) distinguished
by the steno-endemic Limonium dictyophorum. Our
results show a floristic differentiation between the releves
from this subassociation and those carried out on Bisevo
Island and containing Limonium cancellatum (cluster Crl2).
These two groups of releves represent the halophytic aspects
of the Centaureetum ragusinae association, with some ingressive
taxa from the Crithmo-Staticetea class. The releves from
Bisevo Island are here included in the new subassociation
C.r. limonietosum cancellati (holotypus, rel. 7, tab. 5 in Pavletic
1973: Centaurea ragusina 3, Crithmum maritimum 1, Inula verbascifolia
1, Limonium cancellatum þ, Capparis orientalis þ,
Phagnalon graecum þ, Juniperus phoenicea þ, Allium ampeloprasum
þ, Reichardia picroides þ, Convolvulus cneorum þ,
Daucus carota subsp. hispanicus þ).
The Centaureetum ragusinae is usually classified within the
Centaureo cuspidatae-Portenschlagiellion (e.g. Trinajstic 1980,
2008). However, Terzi et al. (2018) showed floristic similarities
with the associations of the Centaureo-Campanulion alliance,
in the Northern Adriatic. These similarities are due to the
occurrence of the halophytic taxa recorded in several other
cliff associations of the Centaureo-Campanulion. On the other
hand, the Centaureetum ragusinae lacks almost completely
the character species of both of the abovementioned alliances.
Therefore, we provisionally classify the Centaureetum
ragusinae within the Centaureo-Portenschlagiellion for
geographical reasons, the distribution area of the main character
taxon of the association, Centaurea ragusina, extending
over the central and southern parts of Dalmatia.
Holotypus: rel. 2, tab. of “Phagnalo-Centaureetum
ragusinae,” p. 258, in Trinajstic (1995); Locus classicus: cliffs in
the northwestern part of Mljet Island, Southern Dalmatia;
Distribution area: Dalmatia; Diagnostic taxa: Centaurea ragusina,
Convolvulus cneorum, Phagnalon rupestre subsp. graecum
(locally). Frequent taxa: Centaurea ragusina, Inula
verbascifolia, Crithmum maritimum, Allium ampeloprasum,
Silene vulgaris, Convolvulus cneorum; Dominant taxon:
Centaurea ragusina.
Micromerio kerneri-Onosmetum dalmaticae
The Micromerio kerneri-Onosmetum dalmaticae was invalidly
described by Lovric and Rac (1987) for the canyons of
Zrmanja and Cikola (Northern Dalmatia) and classified in the
Edraianthion tenuifolii and Moltkietalia petraeae. This association
has been recently moved to the CentaureoPortenschlagiellion
(Terzi et al. 2018).
The releves of the Micromerio kerneri-Onosmetum dalmaticae
were grouped in the same dendrogram cluster together
with those of the Moltkio petraeae-Campanuletum lepidae
(Figure 2). Despite the floristic similarities between these two
associations, they cannot be reunited in only one association
because their character taxa are mutually exclusive. However,
their syntaxonomic relationships need to be reassessed when
more releves are available.
On the other hand, the Moltkio petraeae-Centaureetum
voraginicolae, invalidly described by Lovric (in Lovric and Rac
1987), is here considered as a synonym of the Micromerio
kerneri-Onosmetum dalmaticae because of the high frequency
of two character taxa, Micromeria kerneri and Asplenium ceterach
subsp. bivalens (tab. 1).
Holotypus: rel. 5, tab. 2, p. 113, in Lovric and Rac (1989);
Locus classicus: Krupa canyon, Northern Dalmatia;
Distribution area: Northeastern Adriatic, Northern Dalmatia;
Diagnostic taxa: Onosma echioides subsp. dalmatica (d),
Micromeria kerneri, Seseli tomentosum, Asplenium ceterach
subsp. bivalens; Frequent taxa: Onosma echioides subsp. dalmatica,
Inula verbascifolia, Seseli tomentosum, Galium corrudifolium,
Iris illyrica, Sesleria tenuifolia subsp. tenuifolia, Sedum
hispanicum, Micromeria kerneri, Asplenium trichomanes, As.
ceterach subsp. bivalens, Campanula pyramidalis, Sedum dasyphyllum;
Dominant taxa: Onosma echioides subsp. dalmatica,
Inula verbascifolia, Seseli tomentosum.
Moltkio petraeae-Campanuletum lepidae
This association is here classified in the CentaureoPortenschlagiellion
due to the presence of Moltkia petraea,
Portenschlagiella ramosissima and Tanacetum cinerariifolium.
However, lying in the northern part of the alliance distribution
area, the Moltkio petraeae-Campanuletum lepidae
includes also a character taxon of the CentaureoCampanulion
(Campanula fenestrellata subsp. fenestrellata)
among its most frequent taxa.
PLANT BIOSYSTEMS - AN INTERNATIONAL JOURNAL DEALING WITH ALL ASPECTS OF PLANT BIOLOGY 5
Holotypus: rel. 2, tab. 2, p. 113, in Lovric and Rac (1989);
Locus classicus: Krupa Canyon, Northern Dalmatia;
Distribution area: Northern Dalmatian inland; Diagnostic taxa:
Campanula fenestrellata subsp. fenestrellata, Arenaria orbicularis,
Iris adriatica (d); Frequent taxa: Campanula fenestrellata
subsp. fenestrellata, C. pyramidalis, Moltkia petraea, Galium
corrudifolium, Asperula aristata, Asplenium trichomanes;
Dominant taxa: Campanula fenestrellata subsp. fenestrellata,
Ficus carica.
Moltkio petraeae-Inuletum verbascifoliae
The “Inula candida-Moltkea petraea-Ass. (Horv. 41) Wendelb
63” was invalidly described by Wendelberger (1963) for the
Biokovo mountain ridge, and classified in the Micromerion
croaticae (Wendelberger 1963, 1965). This association was
accidentally validated by Lovric and Rac (1987) through the
publication of only one releve, and classified in the
Edraianthion tenuifolii. Subsequently, Lovric et al. (2002)
moved this association to the Centaureo-Campanulion.
According to Lovric and Rac (1987), the Moltkio petraeaeInuletum
verbascifoliae is characterized by Campanula portenschlagiana
and corresponds pro parte to the CampanuloMoltkietum
petraeae sensu Horvatic 1963, replacing at higher
altitude the Fibigio triquetrae-Cerinthetum tristis along the Mt.
Biokovo coastal cliffs. However, they did not mention or discuss
the syntaxonomic proposals of Trinajstic (1980, 1987)
who described two other associations with Campanula portenschlagiana,
namely Portenschlagiello ramosissimaeCampanuletum
portenschlagianae and Inulo verbascifoliaeCentaureetum
cuspidatae, characterized by Fibigia triquetra
and Centaurea cuspidata, respectively. In the cluster analysis,
the four associations mentioned above turned out to be
grouped in the same cluster. Moreover, this cluster showed
some floristic overlaps with the Campanulo pyramidalisMoltkietum
petraeae.
This result is biased by the low number of releves for the
Moltkio petraeae-Inuletum verbascifoliae and Fibigio triquetraeCerinthetum
tristis, which were represented in our data set by
only their holotypes. Nonetheless, these two associations
were validly published and their names have priority over
Portenschlagiello ramosissimae-Campanuletum portenschlagianae
and Inulo verbascifoliae-Centaureetum cuspidatae (Terzi
and Di Pietro 2016; Jasprica and Terzi 2017). Therefore, we
consider only one association – whose correct name is
Moltkio petraeae-Inuletum verbascifoliae, characterized by
Campanula portenschlagiana.
The Portenschlagiello ramosissimae-Campanuletum portenschlagianae
and Inulo verbascifoliae-Centaureetum cuspidatae
are here lowered to the rank of subassociations of the
Moltkio petraeae-Inuletum verbascifoliae as M.p.-In.v. fibigietosum
triquetrae and M.p.-In.v. centaureetosum cuspidatae,
respectively. Actually, the distribution range of the stenoendemic
Centaurea cuspidata, restricted to the southern part of
the Biokovo ridge, is included within the distribution area of
Fibigia triquetra. However, these two stenoendemics species
seem to be mutually exclusive, at least in the releves
included in the data set. In fact, Jasprica and Terzi (2017)
collected 10 releves of the Portenschlagiello ramosissimaeCampanuletum
portenschlagianae with no record of
Centaurea cuspidata, whereas the releves of the Inulo verbascifoliae-Centaureetum
cuspidatae carried out by Trinajstic
(1980, 1987) include no record of Fibigia triquetra. For this
reason, we have considered three subassociations of the
Moltkio petraeae-Inuletum verbascifoliae: the typical one (M.p.In.v.
typicum), the M.p.-In.v. centaureetosum cuspidatae characterized
by Centaurea cuspidata and the M.p.-In.v. fibigietosum
triquetrae, characterized by Fibigia triquetra.
Holotypus: rel. J1, tab. 6, p. 126, in Lovric and Rac (1987);
Locus classicus: Mt. Biokovo; Distribution area: Central and
Southern Dalmatia; Diagnostic taxa: Campanula portenschlagiana,
Centaurea cuspidata, Fibigia triquetra and locally Cerinthe
minor subsp. auriculata. Frequent and dominant taxa:
Campanula pyramidalis, Inula verbascifolia, Portenschlagiella
ramosissima, C. portenschlagiana.
Seslerio robustae-Putorietum calabricae
This association is represented in our dataset by the releves
of the Birac’s thesis (1971) from which the original diagnosis
of the association was derived and effectively published 2
years later (Birac 1973). The lectotypus hoc loco of the
Seslerio robustae-Putorietum calabricae is releve 6 of tab. II in
Birac (1971), as here reported: Putoria calabrica 3, Inula verbascifolia
2, Parietaria judaica 1, Centaurea glaberrima 1,
Sesleria robusta þ, Campanula pyramidalis þ, Asplenium ceterach
þ, Micromeria juliana þ, Brachypodium retusum þ,
Helichrysum italicum þ, Dittrichia viscosa þ, Aurinia leucadea
þ, Euphorbia dendroides þ, Lavandula latifolia þ, Picris hispidissima
þ, Sonchus asper subsp. glaucescens þ, Smyrnium olusatrum
þ.
Locus classicus: The Ombla Estuary near the city of
Dubrovnik; Distribution area: Southern Dalmatia, Montenegro
and Northwestern Albania; Diagnostic taxa: Putoria calabrica,
Sesleria robusta, Aurinia leucadea, Lavandula latifolia.
Frequent and dominant taxa: Putoria calabrica, Inula verbascifolia,
Campanula pyramidalis s.l., Sesleria robusta.
Teucrio arduinii-Seselietum globiferi
This vegetation exhibits a wide altitudinal range, from sea
level (e.g. in the Kotor Bay, Montenegro) up to 1,000 m a.s.l.,
in Bosnian and Herzegovinian canyons (Lovric et al. 2002,
Milovic 2015). Additionally, stands of this community show
some similarities with the thermophilous chasmophytic vegetation
of walls (Jasprica et al. 2017).
Holotypus: rel. 7, tab. 1, p. 45, in Lovric et al. (2002); Locus
classicus: The Ombla Estuary, near the city of Dubrovnik;
Distribution area: Dalmatia, Southern Bosnia and
Herzegovina, Montenegro; Diagnostic taxa: Seseli globiferum,
Hieracium waldsteinii subsp. plumulosum (d), Teucrium arduinii;
Frequent taxa: Seseli globiferum, Campanula pyramidalis
s.l., Alyssoides utriculata, Asplenium ceterach subsp. ceterach;
Dominant taxa: Seseli globiferum.
6 M. TERZI AND N. JASPRICA
Campanulo austroadriaticae-Moltkietum petraeae
This association was originally described as a subassociation
(campanuletosum pyramidalis) of the “Campanulo-Moltkietum
petraeae Horvatic 1963“ and classified in the CentaureoCampanulion
(Tomic-Stankovic 1970). Subsequently, the subassociation
name was corrected (by using Campanula austroadriatica
instead of C. pyramidalis) and the syntaxon was
raised to the rank of association (Terzi et al. 2017, 2018). The
Campanulo austroadriaticae-Moltkietum petraeae, is in an
intermediate syntaxonomic position between the Centaureo
cuspidatae-Portenschlagiellion and the Edraianthion tenuifolii
(Terzi et al. 2017, 2018). The association describes the thermophilous
communities of the Mt. Lovcen towards Kotor Bay
and is substituted by the Saxifrago crustatae-Moltkietum petraeae
(Tomic-Stankovic ex Terzi et al. 2017) Terzi et al. 2018
in colder habitats. The latter association contains some typical
taxa of less xerothermic conditions, and it clearly belongs
to the Edraianthion tenuifolii (compare Terzi et al. 2018). For
this reason, the Saxifrago crustatae-Moltkietum petraeae has
not been considered in this revision.
Holotypus: rel. 11, tab. II, in Tomic-Stankovic (1970; see:
Terzi et al. 2018); Locus classicus: Tisove ploce, Mt. Lovcen;
Distribution area: Southeastern Adriatic coast (Croatia,
Montenegro); Diagnostic taxa: Campanula austroadriatica,
Silene tommasinii, Peucedanum longifolium (d), Achnatherum
calamagrostis (d); Frequent taxa: Moltkia petraea, Campanula
austroadriatica, Asplenium trichomanes, Cephalaria leucantha,
Tanacetum cinerariifolium; Dominant taxa: Moltkia petraea,
Campanula austroadriatica, Achnatherum calamagrostis.
Conclusion
The phytocoenological diversity of the limestone cliffs of
Central and Southern Dalmatia has been hidden behind different
syntaxonomic interpretations, involving two overlapping
orders, Moltkietalia petraeae and CentaureoCampanuletalia.
The proposal to restrict the distribution
range of the Moltkietalia petraeae to the montane and alpine
belts of the Central and Southern Dinarides, as provided
with the EVC, has resolved the problem of the ecological
overlapping of this order with the Centaureo-Campanuletalia,
which was restricted to the coastal and subcoastal
Adriatic area.
However, the syntaxonomic and synchorological relationships
between the subcoastal associations already classified
in the Moltkietalia petraeae and in the CentaureoPortenschlagiellion
(Centaureo-Campanuletalia) have
remained unclear.
This revision has shown that four associations (Moltkio
petraeae-Inuletum verbascifoliae, Fibigio triquetrae-Cerinthetum
tristis, Inulo verbascifoliae-Centaureetum cuspidatae, and
Portenschlagiello ramosissimae-Campanuletum portenschlagianae)
are not sufficiently distinguishable to be treated as
autonomous associations. As a consequence they have been
reunited as a sole association whose name is the earliest validly
published one, Moltkio petraeae-Inuletum verbascifoliae.
The Inulo verbascifoliae-Centaureetum cuspidatae (i.e. the
nomenclatural type of the Centaureo-Portenschlagiellion) and
Portenschlagiello ramosissimae-Campanuletum portenschlagianae
have been here treated as subassociations of the Moltkio
petraeae-Inuletum verbascifoliae.
Moreover, the Campanulo austroadriaticae-Moltkietum petraeae
– which is intermediate between the CentaureoPortenschlagiellion
and the Edraianthion tenuifolii (Terzi et al.
2018) – has to be classified in the latter alliance for the
occurrences of several taxa typical of cool conditions.
This revision has clarified the syntaxonomic diversity of
the Centaureo-Portenschlagiellion, which includes only seven
associations, whereas the others already described in phytosociological
literature are invalidly described (e.g. Moltkio petraeae-Centaureetum
voraginicolae, Puccinellio-Aurinietum
leucadae, Aurinio-Brassicetum frutescentis) or are here treated
as syntaxonomic synonyms of the previous ones.
It is well known that the Centaureo-Campanuletalia order
is characterized by numerous endemic and rare plant taxa
and communities on both sides of the Adriatic Sea (e.g. Di
Pietro and Wagensommer 2008; Nikolic et al. 2015) and this
revision represents a contribution to the knowledge of coenological
diversity of the xerothermic cliffs vegetation in the
eastern part of the Adriatic Basin.
Nonetheless, this study also highlights the need for further
investigations in the less known areas, such as the Vis
Archipelago (Middle Adriatic, Croatia) – an area rich in many
stenoendemic taxa of exceptional phytogeographical importance,
such as Puccinellia teyberi, Asperula visianii, Centaurea
friderici, C. crithmifolia – or the coastal and subcoastal areas
of Montenegro and Albania which are still poorly known
from a phytosociological point of view and of great importance
to identify the southern boundary of the distribution
area of the Centaureo-Portenschlagiellion.
Nomenclature
Taxonomic nomenclature follows Flora Croatica Database
(https://hirc.botanic.hr/fcd/, accessed on March 2018), and
The Plant List (http://www.theplantlist.org/, accessed on
March 2018) for taxa not included there. Syntaxonomic
nomenclature follows Terzi et al. (2017), and Skvorc et al.
(2017) for syntaxa not included in Terzi et al. (2017).
Acknowledgements
The authors thank Dr Niksa Glavic for preparation of the map of the
study area and Steve Latham (UK) for improving the English and two
anonymous referees for their useful suggestions.
Disclosure statement
No potential conflict of interest was reported by the authors.
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