Nuclear DNA Amounts in Angiosperms: Progress, Problems and Prospects
M. D. BENNETT* and I. J. LEITCH
Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK
Received: 23 July 2004 Returned for revision: 12 August 2004 Accepted: 1 September 2004
CONTENTS
INTRODUCTION 45
PROGRESS 46
Improved systematic representation (species and families) 46
(i) First estimates for species 46
(ii) First estimates for families 47
PROBLEMS 48
Geographical representation and distribution 48
Plant life form 48
Obsolescence time bomb 49
Errors and inexactitudes 49
Genome size, `complete' genome sequencing, and, the euchromatic genome 50
The completely sequenced genome 50
Weeding out erroneous data 52
What is the smallest reliable C-value for an angiosperm? 52
What is the minimum C-value for a free-living angiosperm and other free-living organisms? 53
PROSPECTS FOR THE NEXT TEN YEARS 54
Holistic genomics 55
LITERATURE CITED 56
APPENDIX 59
Notes to the Appendix 59
Original references for DNA values 89
 Background The nuclear DNA amount in an unreplicated haploid chromosome complement (1C-value) is a key
diversity character with many uses. Angiosperm C-values have been listed for reference purposes since 1976, and
pooled in an electronic database since 1997 (http://www.kew.org/cval/homepage). Such lists are cited frequently and
provide data for many comparative studies. The last compilation was published in 2000, so a further supplementary
list is timely to monitor progress against targets set at the first plant genome size workshop in 1997 and to facilitate
new goal setting.
 Scope The present work lists DNA C-values for 804 species including first values for 628 species from 88 original
sources, not included in any previous compilation, plus additional values for 176 species included in a previous
compilation.
 Conclusions 1998­2002 saw striking progress in our knowledge of angiosperm C-values. At least 1700 first values
for species were measured (the most in any five-year period) and familial representation rose from 30 % to 50 %.
The loss of many densitometers used to measure DNA C-values proved less serious than feared, owing to the
development of relatively inexpensive flow cytometers and computer-based image analysis systems. New uses of the
term genome (e.g. in `complete' genome sequencing) can cause confusion. The Arabidopsis Genome Initiative
C-value for Arabidopsis thaliana (125 Mb) was a gross underestimate, and an exact C-value based on genome
sequencing alone is unlikely to be obtained soon for any angiosperm. Lack of this expected benchmark poses a
quandary as to what to use as the basal calibration standard for angiosperms. The next decade offers exciting
prospects for angiosperm genome size research. The database (http://www.kew.org/cval/homepage) should become
sufficiently representative of the global flora to answer most questions without needing new estimations.
DNA amount variation will remain a key interest as an integrated strand of holistic genomics.
 2005 Annals of Botany Company
Key words: Angiosperm DNA amounts, DNA C-values, nuclear genome size, plant DNA C-values database.
INTRODUCTION
It has been possible to estimate the amount of DNA in plant
nuclei for over 50 years, and since the key role of DNA in
biology was discovered in 1953, such research has increased
in each successive decade. Work on plants has played a
leading part in research to describe and understand the
origin, extent and effects of variation in the DNA amount
in the unreplicated haploid nuclear chromosome comple-
ment (defined by Swift, 1950, as the 1C-value) of different
taxa. Indeed, angiosperms are probably the most intensively
Annals of Botany 95/1  Annals of Botany Company 2005; all rights reserved
* For correspondence. E-mail m.bennett@kew.org
Annals of Botany 95: 45­90, 2005
doi:10.1093/aob/mci003, available online at www.aob.oupjournals.org
studied major taxonomic `group' of organisms, with
published C-values for over 4100 species.
Early research to address questions such as possible
relationships between DNA C-value and the rate of cell
development (e.g. Van't Hof, 1965) usually required
work to estimate C-values for most of the taxa concerned,
as these were unavailable. Later, as taxa with `known'
C-values increased, it was possible to use such data in
new comparisons (supplemented by further first estimates
made for sample taxa). However, it was often difficult to
know whether a C-value existed for a particular taxon, and if
so, where to find it. Such estimates were widely scattered in
the literature or even unpublished. Small lists of nuclear
DNA amounts were published in reviews and research
papers, but the first large list of DNA amounts for angio-
sperms, compiled primarily as a reference source was
published in 1976. This contained data for over 750 species
from 54 original sources (Bennett and Smith, 1976), and
noted an intention to publish supplementary lists for refer-
ence purposes at intervals. Five such lists, together giving
pooled data for over 2900 species from 323 original sources,
have followed (Bennett et al., 1982, 2000; Bennett and
Smith, 1991; Bennett and Leitch, 1995, 1997). Data from
the first five publications were pooled in an electronic
form ­ the Angiosperm DNA C-values database, which
went live in April 1997. This was updated as release 3.1
and incorporated, with databases for gymnosperms,
pteridophytes and bryophytes, into the Plant DNA C-values
database (release 1.0) in 2001.
These data are clearly much used, as the published lists
have been cited over 1400 times, including over 700 times
since 1997, whilst the electronic database has received over
50 000 hits. Recently they have provided the large samples
of data needed for many diverse comparative studies, such
as testing for possible relationships between nuclear DNA
amount and risk of extinction (Vinogradov, 2003), ecolog-
ical factors in California (Knight and Ackerly, 2002), lead
pollution in Slovenia (B. Vilhar, University of Ljubljana,
Slovenia, pers. comm.); ploidy level (Leitch and Bennett,
2004), and land plant evolution (Leitch et al., 2005).
Given their ongoing use as reference sources, publication
of a sixth supplementary list of angiosperm C-values is
timely, if not overdue. The present work lists DNA C-values
for 804 species from 88 original sources, including first
estimates for 628 species not included in any previous com-
pilation, plus additional estimates for 176 species already
included in one or more previous compilation. Data in the
Appendix table were prepared for analysis at the second Plant
Genome Size Discussion Meeting in September 2003, so it is
fitting that they are included in this special supplement.
Whilst they represent most of the new C-value data published
or estimated in 2000­2002, we are already aware of a further
large sample estimated but unpublished either by late 2002,
or subsequently. Thus, despite its large size, the present list
will soon be followed by a seventh supplement.
PROGRESS
Research on DNA C-values in angiosperms is unique in
having been subject to detailed analyses of its quantity
and quality over a long period (Bennett and Leitch,
1995). The importance of identifying gaps in our knowledge
concerning this key biodiversity character, of recommend-
ing targets for new work to fill them by collaboration of
international partners, and of monitoring progress to ensure
that any shortfall is recognized, was confirmed by the first
plant genome size workshop in 1997 (http://www.kew.org/
cval/conference.html#outline, Bennett et al., 2000) and
reviewed by participants at the second plant genome size
workshop in 2003. Thus, what follows is mainly a summary
of the overall progress for angiosperms against key targets
set in 1997 for the following quinquennium (1998­2002).
However, it also notes meaningful statistics for the data
included in the Appendix table, or known to us from
personal communications made after the Appendix table
was closed.
In 1997 C-values for 2802 species (approximately 1 %)
of angiosperm species had been estimated in the previous
40 years. The 1997 workshop concluded that the ideal of
a C-value for all taxa was unrealistic, but long-term,
estimates for 10­20 % of angiosperms seemed both ulti-
mately achievable and adequate for all conceivable uses
provided they were carefully targeted to be representative
of the various taxonomic groups, geographical regions,
and life forms in the global flora. So the first recom-
mended target was to estimate first C-values for the
next 1 % of angiosperm species (i.e. another 2500 spe-
cies) by 2003. Many saw this goal as aspirational, as
achieving it would mean estimating as many C-values
in five years as in the past 40. Others thought that new
technology (e.g. flow cytometry) would make it easy to
achieve.
Improved systematic representation (species and
families)
(i) First estimates for species. In September 1997 the
Angiosperm DNA C-values database contained data for
2802 species. By September 2003 C-values were listed
for 4119 species, including 689 first values for species listed
in Bennett et al. (2000) and 628 such values for species
in the Appendix table. Progress toward the first target in
the five year period (1998­2002) considerably exceeded the
average of $110 first values for species per annum in the
early 1990s. Clearly, the 1997 workshop stimulated an
increase in the total output of first C-values for species to
its highest level for any five-year period (almost 200 per
annum; Fig. 1A). Moreover, the proportion of newly pub-
lished C-values that were also first estimates for species,
which had previously fallen (Bennett et al., 2000), rose as a
result of recent targeting and averaged 72Á5 % for values
published since 1997 (Fig. 1B). Nevertheless, the total
number of published first C-values for species (1032)
listed since 1997 was only 41 % of the 1997 target of
approx. 2500.
The real total of first C-values for angiosperms esti-
mated after 1996 but unpublished by 2003 was much higher,
but is difficult to determine exactly. For example, several
hundred values were measured by Ben Zonneveld (pers.
comm.) using flow cytometry but not published. Listing
46 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
for the Appendix table closed in August 2002, ready for the
workshop; however, we saw 158 first C-values published by
other authors later in 2002, and 22 such values were esti-
mated at RBG, Kew. Adding these data to those listed in our
compilations suggests that the total number of first C-values
for species estimated in 1997­2002 was probably at least
1700 and hence not less than approx. 66 % of the target set
in 1997. Analysis shows that this was achieved by interna-
tional collaboration involving at least 18 research groups in
ten countries. Whilst a target of 2500 was aspirational, it
seems attainable as a future five-year goal. However, at the
present rate achieving 20 % species representation would
take 100 years, so an ultimate goal of 10 % (approx. 25 000
angiosperm species) is more sensible.
(ii) First estimates for families. The 1997 workshop
noted that a first C-value was available for only 30 % of
angiosperm families recognized at that time. Thus, a second
recommended target was `To obtain at least one C-value
estimate for a species in all angiosperm families'. Monitor-
ing first C-values for species listed in Bennett et al. (2000)
showed that progress towards this goal was initially very
slow. Indeed, `since 1997 first C-values had been listed for
691 angiosperm species, but only 12 (1Á7 %) were also first
estimates for families'. Work to correct this began at RBG,
Kew in 1999. In 2001 two papers reported first C-values for
50 families (Hanson et al., 2001a, b), and 30 more followed,
including five basal angiosperm families (Leitch and
Hanson, 2002; Hanson et al., 2003), all included in the
present Appendix table.
Analysis of listed data for 4119 species shows that a
first published value is available for at least 217 of the
457 angiosperm families currently recognized by the
Angiosperm Phylogeny Group (APG) (APG II, 2003).
Together with first estimates for 11 unlisted families
(Hanson, RBG, Kew, pers. comm.; Koce et al., 2003)
measured or seen after listing for the Appendix table was
closed, the total is 228. Thus, since 1997 (after losses owing
to new familial circumscriptions--APG II, 2003; Hanson
et al., 2003) first values for at least 85 such families have
been measured, so good progress has been made. However,
the proportion of families represented rose only from 30 %
to 49Á9 % (Fig. 2), which is less than one third of the target
(100 %) set in 1997. Major factors limiting progress were
0
50
100
150
200
250
300
1950­1954
1960­1964
1970­1974
1980­1984
1990­1994
2000­2002
Period
Meannumberofestimates
0
20
40
60
80
100
1965 1970 1975 1980 1985 1990 1995 2000
Year
PercentageofC-valueestimatespublished
thatare`new'
A
B
FI G . 1. (A) Mean numberper year of total (open symbols)and `first' (closed
symbols) DNA C-value estimates communicated in ten successive 5-year
periods and the 3-year period 2000­2002, between 1950 and 2002. Based on
analysis of data listed in the present Appendix table, and the Angiosperm
DNA C-values database (release 4.0, January 2003). (B) Percentage of
C-value estimates published or communicated during 1965­2002 that are
first values for species listed in the present Appendix table and the
Angiosperm DNA C-values database (release 4.0, January 2003).
0
10
20
30
40
50
60
1950 1960 1970 1980 1990 2000
Year
Cumulative%ofAPGfamilies
FI G . 2. Cumulative percentage of angiosperm families recognized by the
Angiosperm Phylogeny Group (APG) (APGII, 2003) with a first C-value
represented in the present Appendix table, the Angiosperm DNA C-values
database (release 4.0, January 2003), plus eleven known to the present
authors in September 2003.
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 47
discussed previously (Hanson et al., 2003). Unlike progress
towards the species target, which involved many research
groups, movement towards the goal for families since 1997
has depended mainly on work by one institution, as RBG
Kew estimated 65 of the 74 (87 %) first values for families
listed in the Appendix table.
The Plant Genome Size Workshop in 2003 confirmed that
global capacity for estimating DNA C-values (determined
by available equipment, funding and trained operators)
remains very limited. Consequently, any increased focus
on targets for other plant groups (e.g. bryophytes, pterido-
phytes, gymnosperms--reviewed in Leitch and Bennett,
2002b) inevitably reduces progress to improve representa-
tion for angiosperm targets, a problem discussed below.
However, it should not detract from the highly successful
progress to make C-values more representative of the global
flora described above.
PROBLEMS
Geographical representation and distribution
We first noted the need to improve geographical representa-
tion for angiosperm C-values in Bennett and Leitch
(1995). This was confirmed by the 1997 plant genome
size workshop, although no specific regional targets were
recommended. Perhaps, in consequence, progress in this
area has never been monitored in detail, although we
have been at pains to advertise the problem in a general
way and to provoke action to rectify it in particular
regions, such as southern Africa (Leitch and Bennett,
2002a).
There are two critical concerns regarding the geographical
distribution of angiosperm C-value work. (i) The first
concerns the small number of publications with original
C-values by first authors in many regions (Table 1). This
reflects a serious imbalance between the geographical dis-
tribution of research scientists working on genome size and
of taxa whose C-values are unknown. Bennett et al. (2000)
noted that `Africa remains an unexplored continent' and that
`Whereas six out of 377 original sources have first authors
with addresses in Africa, still none has an angiosperm
C-value estimated in Africa, as all six reported work done
in Europe or the USA.' Analysis of the 88 original sources
in the present work shows no improvement, as the number
of original sources from Africa (2), China (2), and South
America (2) remains low (Table 1). (ii) The second concerns
the small number of first C-values by any authors for
species endemic to several large geographical regions.
With some exceptions, the sample is still dominated by
crops and their wild relatives, model species grown for
experimental use, and other species growing near labora-
tories in temperate regions, mainly in Western Europe and
North America. Analysis of data in the Appendix table
shows that none presented data for other taxa endemic to
China, Japan, Brazil, Mexico or Central Africa. Similarly,
although island floras are known to be rich in endemics, no
original source has reported C-values for any large islands
such as Borneo, New Guinea or Madagascar, where 80 % of
the 12 000 described plant species are endemic (Robinson,
2004).
Plant life form
There is also a need for the overall sample to represent
better the full range of plant types and life forms. We
previously identified several associations and life forms
as being poorly represented in the database (Bennett and
Leitch, 1995), yet taxa from bog, fen, tundra, alpine
and desert environments, and halophytic, insectivorous,
parasitic, saprophytic and epiphytic species and their asso-
ciated taxa are all still under-represented.
Solving this problem needs a proactive approach, as
recent experience with first C-values for angiosperm
families shows. First, a target must be set for each gap.
Second, monitoring newly published data against targets
TA B L E 1. The number and percentage (in brackets) of original references with first authors from various geographical areas
among the total of 465 sources contributing to the present Appendix table and the six lists of angiosperm DNA amounts previously
compiled for reference purposes that were pooled in the Angiosperm DNA C-values database (release 4.0, January 2003)
DNA C-value compilation
Area 19761
19822
19913
19954
19975
20006
Present Appendix Total
Europe 34 (63.0) 38 (71.7) 30 (53.6) 43 (40.6) 18 (48.6) 38 (51.4) 54 (61.4) 255 (54.8)
UK 28 (51.9) 13 (24.5) 22 (39.3) 23 (21.7) 5 (13.5) 8 (10.8) 10 (11.4) 109 (23.4)
North America 14 (25.9) 11 (20.8) 16 (28.6) 19 (17.9) 5 (13.5) 11 (14.9) 13 (14.8) 89 (19.1)
South and Meso America 0 (0.0) 0 (0.0) 3 (5.4) 9 (8.5) 1 (2.7) 6 (8.1) 2 (2.3) 21 (4.5)
Africa 1 (1.9) 0 (0.0) 0 (0.0) 1 (0.9) 2 (5.4) 1 (1.4) 2 (2.3) 7 (1.5)
Asia 1 (1.9) 3 (5.7) 4 (7.1) 30 (28.3) 11 (29.7) 8 (10.8) 17 (19.3) 74 (15.9)
India 1 (1.9) 1 (1.9) 2 (3.6) 28 (26.4) 11 (29.7) 4 (5.4) 11 (12.5) 58 (12.5)
China 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) 0 (0.0) 2 (2.3) 2 (0.4)
Australasia 4 (7.4) 1 (1.9) 3 (5.4) 4 (3.8) 0 (0.0) 7 (9.5) 0 (0.0) 19 (4.1)
Australia 4 (7.4) 1 (1.9) 3 (5.4) 1 (0.9) 0 (0.0) 1 (1.4) 0 (0.0) 10 (2.2)
Total 54 53 56 106 37 71 88 465 (100)
1
Bennett and Smith (1976); 2
Bennett et al. (1982); 3
Bennett and Smith (1991); 4
Bennett and Leitch (1995); 5
Bennett and Leitch (1997); 6
Bennett
et al. (2000).
48 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
must begin. Third, if work on poorly-represented floras or
life forms does not increase, then established research
groups must re-focus on target material available in exist-
ing collections. Unless global capability for estimating
plant DNA C-values is significantly increased by new
technology, funds or skilled operators, then this change
in strategy will reduce progress towards achieving other
targets. However, the prime objective remains: to generate
a sample representative of the global flora that is able
to support most comparative studies. Managers of the
limited global capacity for estimating genome size should
keep this firmly in mind when targeting taxa for new
work.
Obsolescence time bomb
Several methods have been used to measure plant DNA
C-values, but most values have been estimated by Feulgen
microdensitometry (Fe), both overall and since 1997.
In 1997 we identified an imminent problem, likely to
limit future estimations. This was the failure and non-
replacement of densitometers long used by many groups
to estimate DNA C-values. Manufacturers were ending
their support for such equipment, and users faced difficulty
in funding new equipment for this purpose. Moreover, this
problem was likely to be most acute in regions where some
of the greatest gaps in our knowledge lay. Reviewing
the position at the second Plant Genome Size Workshop
confirmed that, as predicted, the `obsolescence time bomb'
had exploded. By 2003 several laboratories that had
long published C-values listed in the Angiosperm DNA
C-values database were now unable to estimate C-values
by this (e.g. in Mexico, the USA), or any method (e.g. in
Argentina). Vickers Instruments no longer supports their
M85 microdensitometer, and spare parts for it are unobtain-
able. A few laboratories, including ours, can still use such
machines, but now without servicing and only until they fail
catastrophically.
As expected, one response to this problem was the
increased use of flow cytometry (FC) to replace Fe. Analysis
of data in the Appendix table shows a higher proportion of
C-values obtained by FC (58Á4 %), mostly since 1997, than
noted previously (48Á6 %) for data listed in Bennett and
Leitch (2000), whilst in Bennett and Leitch (1995, 1997) FC
averaged 26Á7 %. Several groups have undertaken careful
studies to compare DNA estimates made by FC and Fe, to
define best practice for FC, or to show that FC can be
applied widely to most plants across the full range of
known DNA C-values (e.g. see review by Dolezzel and
Bartos, 2005, this volume). Fortunately, the cost of a
basic flow cytometer for such work has fallen, and suitable
models (e.g. Partec PAII) have recently been set up for this
use for approx. 20K (US$30K). If this technology con-
tinues to improve, and its costs continue to fall, FC should
be more easily available. However, FC easily yields poor
data in unskilled hands and by itself does not provide the
cytological view of test material(s) that is essential to count
chromosome number(s). Its use in some less-developed
countries (where the greatest gaps in our knowledge still
remain) will depend on training local operators, but such
capacity-building may be thwarted by a lack of in-country
support by the suppliers of flow cytometers.
A second solution to the problem is a new availability
of relatively inexpensive computer-based image analysis
(CIA) systems, which can estimate DNA amounts using
Feulgen-stained cytological preparations in place of a micro-
densitometer. Although proprietary hard-wired CIA systems
have been available since the 1970s (e.g. Zeiss Quantimet
system), they cost much more than microdensitometers, and
analysis of the literature shows they have not been used to
estimate plant C-values. However, in the 1990s, with
advances in computer technology, less expensive systems
were developed (e.g. CIRES system) primarily for medical
use, and these have also been used to good effect for plant
C-value estimations (e.g. Temsch et al., 1998; Greilhuber
et al., 2000).
Sadly, the CIRES system that adapted well for this pur-
pose is no longer available, as the software is incompatible
with the operating system used on modern PCs. However,
computer-literate groups can assemble the kit needed to
estimate C-values using CIA, and several software packages
written specifically for this purpose are available (Vilhar
et al., 2001; Hardie et al., 2002). Hardie et al. (2002) give
an excellent review of this technique and practical issues
concerned with its use for animal materials, and Vilhar et al.
(2001) have compared CIA, Fe and FC, to help define
best practice for CIA, demonstrating that CIA can be
applied to plants with an approx. 100-fold range of
C-values. Vilhar et al. (2001) concluded that `DNA image
photometry gives accurate and reproducible results, and
may be used as an alternative to photometric cytometry
in plant nuclear DNA measurements'. CIA can use an
existing microscope, costs less than FC to set up, and is
easier to service in countries that lack FC manufacturers'
support. The field would benefit from development of a
standard inexpensive CIA `kit', an agreed best practice
CIA technique, and easy access to leading laboratories
for training and technology transfer. Given this, CIA could
soon become the method of choice for estimating C-values
in angiosperms, replacing Fe as a method of choice along
with FC, but with the advantage that, unlike FC, it uses
microscope slide preparations, allowing users to make cyto-
logical observations.
Errors and inexactitudes
Swift (1950) defined the DNA content of an unreplicated
haploid complement as its 1C-value (C standing for
`constant'). Thus, replicated diplophase nuclei have a
4C DNA amount and produce two unreplicated 2C nuclei
by mitotic division, and four 1C gametic nuclei after meio-
sis, irrespective of the organism's ploidy level. This con-
vention applies well to polyploid taxa with diploidized
meiotic chromosome pairing such as hexaploid breadwheat,
which produce mainly functional, balanced polyhaploid
gametes with 1C DNA amounts at meiosis (Rees and
Walters, 1965). Consequently, for several previous refer-
ence lists, 4C DNA estimates for all taxa were divided
by 2 and 4 to generate 2C- and 1C-values respectively
(e.g. Bennett et al., 2000). However, a problem with this
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 49
practice was identified for the few taxa with odd ploidy
levels in release 3.1 of the Angiosperm DNA C-values
database (namely 45 out of 3493 listed taxa, $1Á3 %), as
the resulting 1C-values are not biologically meaningful. For
example, triploids with a 4C amount in their fully replicated
metaphase nuclei do regularly produce two 2C nuclei at
mitosis, but do not regularly produce four 1C products at
meiosis. The authors are grateful to several colleagues who
noted this problem and suggested solutions.
This problem has several practical consequences.
(i) Regrettably, researchers who use 1C data from the
literature or downloaded from the Angiosperm DNA
C-values database may have included this error in the
samples that they used for comparative analyses. However,
this is unlikely to have influenced their conclusions signific-
antly, since the magnitude of the error is relatively small
(ranging between 0Á25C for a triploid, to +0Á25C for a
pentaploid--which tend to cancel out), and affects only
1Á3 % of all taxa listed. Overall, errors in mean DNA
amounts for samples are probably less than 0Á5 %. Studies
that used data from the 2C or 4C columns for samples of
odd-ploid taxa are unaffected by the error. (ii) To ensure that
researchers are aware of the problem and do not generate 1C
data for taxa with odd ploidy levels in the future, release 4.0
of the Angiosperm DNA C-values database gives 2C- and
4C-values for the 45 out of 3493 entries with odd ploidy
levels, plus a warning note in response to any queries for
1C-values. This approach is also followed in the present
Appendix table (see footnote t). (iii) This problem also high-
lights a general need to re-assess definitions of `C-value'
and `genome size' in light of recent usage and new theore-
tical understanding, a topic explored by Greilhuber et al.
(2005). Indeed, the above problem shows the need for care
when handling data, and the danger of using computer-
generated numbers uncritically. It is clearly perilous to
ignore basic biology or the literature, as the recent history
of genome size, `complete' genome sequencing, and interest
in the smallest angiosperm genome clearly shows.
Genome size, `complete' genome sequencing,
and, the euchromatic genome
A growing semantic problem concerns different uses of
the term `genome' (Greilhuber et al., 2005). As originally
defined by Winkler (1920), genome referred to a monoploid
chromosome complement. Since a monoploid is defined as
`having one chromosome set with the basic (x) number of
chromosomes' (Rieger et al., 1991), it followed by defini-
tion that any polyploid taxon had three or more genomes.
However, an alternative meaning, now in common usage,
uses genome as an interchangeable alternative for the
1C-value to refer to the DNA content of an unreplicated
gametic nuclear complement, irrespective of ploidy level.
Unless the meaning intended is clearly defined on each
occasion, this can be confusing, especially when authors
use both meanings for a polyploid taxon in the same
paper. For example, Devos and Gale (1997) used the
term `genome' to refer to both the entire complement of
nuclear DNA in a hexaploid wheat nucleus and to the
individual A, B and D `genomes'.
Further potential for confusion comes from new uses of
the term `genome' recently spawned by genome sequencers.
These concern the counter-intuitive meaning of a `wholly',
`completely' or `entirely' sequenced genome, or of equating
`genome' with `euchromatic genome'--a confusing con-
cept in which `genome' equals the parts which could be
cloned and sequenced, but not the rest (see below). None of
these qualitative new uses of genome equates to its quanti-
tative use to mean either a 1C-value, or one monoploid
parental genome in a polyploid.
The completely sequenced genome
Since 2000 the scientific and popular press has reported
and celebrated the `complete' sequencing of the first insect
(Drosophila melanogaster) and plant genome (Arabidopsis
thaliana) and the human genome (in 2001). For example,
a title in Nature reported: `The sequencing of an entire plant
genome is now complete.' Readers could be forgiven for
assuming this meant the entire linear sequence of the
nuclear DNA had been sequenced and assembled, so that
the total size of the nuclear genome in these organisms was
now known with certainty, and hence much more accurately
than any previous estimate based on other methods subject
to various experimental errors. The popular and scientific
literature easily gives that impression, and unfortunately
that is what many, incorrectly, understood. The truth is
otherwise, as a `completely sequenced' genome is a very
relative concept. In the same issue of Science where Brenner
(2000) wrote `We have the complete sequence of the
125-megabase genome of the fruit-fly Drosophila', Pennisi
(2000) noted that `the fly sequence still has c. 1000 small
gaps'--referring only to the sequenced euchromatin part.
But what of the rest? Speaking of heterochromatin, Adams
et al. (2000) explained that the `genomes of eukaryotes
generally contain heterochromatic regions surrounding
the centromeres that are intractable to all current sequencing
methods' and that `Because of the unclonable repetitive
DNA surrounding the centromeres it is highly unlikely
that the genomic sequence of chromosomes from eukar-
yotes such as Drosophila or human will ever be `complete'.
Moreover, Adams et al. (2000) stated that the unsequenced
centric heterochromatin regions comprised `one third' of the
approx.180 Mb genome of Drosophila. But how was its size
determined? Careful reading revealed that the Mb size of
these unsequenced centromeric heterochromatic segments
was measured not by any modern molecular method, but
by using a ruler on one cell of a plate in a paper by
Yamamoto et al. (1990). This important detail is not stated
in the main text, but in the legend to fig. 1 in Adams et al.
(2000). As Bork and Copley (2001) clearly explain, `There
are regions, often highly repetitive, that are difficult or
impossible to clone (one of the initial steps in a sequencing
project) or sequence with current technology. . . . The extent
of these regions varies widely in different species. So,
rather than applying a universal gold standard, each sequen-
cing project has made pragmatic decisions as to what
constitutes a sufficient level of coverage for a particular
genome. For example, as much as one-third of the
sequence of the fruitfly Drosophila melanogaster was not
50 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
stable in the cloning systems used, and so was not
sequenced.'
Thus, workers interested in C-values should clearly
understand that a `completely', `entirely' or `wholly'
sequenced genome is not what those words might imply
if taken at face value, and the size given for such a genome
may indicate either the amount of DNA sequenced, or the
size of that euchromatic genome sequenced plus a best-
guess estimate of a lot of unsequenced heterochromatin.
Further, it can mean that every type of sequence in an
organism has been sequenced, but it need not mean that
all copies of all types have been sequenced, or that their
copy numbers are known. Without this information total
genome size (the DNA C-value) cannot be determined
based on genome sequencing (Bennett et al., 2003).
Swift (1953) stated that, `in general estimates of the
nucleic acids in cells are at present accurate to 10 or
20 %'. Later, Bennett and Smith (1976) concluded that
`While a few estimates are not accurate even to within
20 %, careful measurements of 4C DNA amounts in species
with 0Á5­2Á0 times that of a standard species are probably
accurate to within 5­10 %'. Greilhuber (1998) noted `much
suspect or demonstrably wrong data have accumulated and
continue to be accumulated in the literature'. Sadly, the
`complete' genome sequencing of Arabidopsis (Arabidopsis
Genome Initiative, 2000), which was expected to provide
a new baseline, only added to this phenomenon.
Plant genome size researchers have long recognized the
need for an exact calibration standard, whose C-value is not
subject to technical errors. Thus, the publication of a precise
C-value for the first plant to have its genome completely
sequenced was eagerly awaited, as it was expected to
provide a baseline, gold-standard reference point, against
which all other plants could be compared and expressed.
Arabidopsis thaliana ecotype `Columbia' was chosen for
complete genome sequencing, partly because its tiny
genome should be less costly to sequence than larger
genomes in other species.
In 2000 the Arabidopsis Genome Initiative (AGI) pub-
lished the genome size of Arabidopsis thaliana as 125 Mb,
comprising 115Á4 Mb in the sequenced regions plus a rough
estimate of 10 Mb in unsequenced centromere and ribo-
somal DNA regions. The accuracy of this estimate was
set not by the precision of sequencing and assembling con-
tigs, but by the total inaccuracy in the sizes assumed for the
unsequenced gaps (Bennett et al., 2003) and hence was no
more accurate than many estimates in the range 150­180 Mb
made by other methods. Further analysis showed that the
AGI's rough estimate of 10 Mb in the unsequenced gaps
was highly inaccurate. Thus, new comparisons using flow
cytometry, which co-ran A. thaliana ecotype `Columbia'
with three animal species including Caenorhabditis elegans
Bristol N2 (whose genome size is accurately established by
genome sequencing as just over 100 Mb), gave C-value
estimates for A. thaliana in the range 154­162 Mb (with
157 Mb when C. elegans was used as the standard) (Bennett
et al., 2003). This value is about 25 % larger than the AGI
estimate of 125 Mb which was clearly a gross under-
estimate, and hence is not the long-awaited first benchmark
C-value for a completely sequenced plant genome--giving
those words their natural meaning. Other molecular work
has confirmed this conclusion (e.g. Hosouchi et al., 2002).
More recently, the draft DNA sequence of the rice (Oryza
sativa) genome was published (O. sativa ssp. japonica,
Goff et al., 2002; O. sativa ssp. indica, Yu et al., 2002).
However, while the estimated genome sizes based on DNA
sequencing did not suffer from the serious shortcomings of
the Arabidopsis estimate, neither did they fulfil the criteria
essential for a new benchmark calibration standard. Yu et al.
(2002) gave a new C-value of 466 Mb for O. sativa ssp.
indica calculated by adding up the DNA sequencing data for
362 Mb of sequenced scaffolds and 104 Mb of `unas-
sembled data'. In contrast Goff et al. (2002) reported the
sequencing of DNA which covered a total of 389 809 244 bp
of the O. sativa ssp. japonica genome. They stated that this
represented 93 % of the 420 Mb rice genome but did not
give a reference to the source of 420 Mb. It is therefore
unclear whether the C-value of 420 Mb given by Goff et al.
(2002) represents a new C-value based on genome sequen-
cing alone. The 1C-value for rice may yet prove to be
slightly higher than the values assumed by Goff et al.
(2002) and Yu et al. (2002), and approach 490 Mb, equival-
ent to the 0Á5 pg estimated by Bennett and Smith (1991).
Exact C-values based on complete genome sequences
would be invaluable (Bennett et al., 2003). The need to
complete sequencing gaps in Arabidopsis remains tech-
nically difficult, and it is unclear how, when, or if it will
be achieved. Genome sequencing becomes more difficult as
genome size increases, and experience with Arabidopsis
implies that exact C-values are unlikely to be obtained in
this way soon for any larger plant genomes, including the
established plant C-value standard Oryza sativa.
The current situation poses a quandary for the plant
genome size community, who have long paid serious
attention to trying to maximize the accuracy and compar-
ability of plant DNA C-values by using agreed calibration
standards (both materials and assumed values; e.g. see
http://www.rbgkew.org.uk/cval/conference.html#outline,
Bennett et al., 2000), while eagerly awaiting the first absol-
ute measurement for a plant obtained by really complete
DNA sequencing. Current options include: (i) continue to
use the existing small group of plant calibration standards
until a plant C-value which meets the required criteria
becomes available; (ii) adopt an animal C-value which
meets these criteria as the baseline reference for expressing
all other plant species values, e.g. Caenorhabditis elegans
Bristol N2, whose C-value is known with confidence to
within 1 % from genome sequencing to be just above
100 Mb (or roughly 0Á1 pg); (iii) adopt a plant value
based on direct comparisons with C. elegans, as the base
calibration standard for plants, and create a ladder of
secondary calibration standards all measured against it in
a study replicated between several groups able to use best
practice. The C-value for Arabidopsis thaliana (1C = 157 Mb
or 0Á16 pg), recently measured against C. elegans (Bennett
et al., 2003), could be adopted as the basal plant calibration
standard. Seed is readily available from stock centres and
gives small, easily grown plants. Moreover, the ladder of
values for its many endopolyploid nuclei would also provide
convenient calibration reference points for higher values up
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 51
to approx. 2500 Mb or 2Á5 pg (i.e. 0Á64  4C, 1Á28  8C,
and 2Á56  16C).
Weeding out erroneous data
The value of the database is determined by the accuracy of
the data it contains. Ideally, values should be exact, but in
reality they are all subject to various technical and other
errors, as noted above. This raises questions as to how accur-
ate data are, and what level of error is acceptable in practice,
or makes a datum valueless for a particular use or study.
The existence of a database itself is a valuable means of
identifying real or potential errors, and hence of improving
the accuracy and quality of the whole body of data. For
example, where estimates for the same taxon (with the same
chromosome number) disagree greatly this suggests an
error. Further, where a body of data for a taxon shows
close agreement except for one major departure, this iden-
tifies the outlier as almost certainly incorrect. For example,
in the Appendix table, the 2C-value for diploid Acacia
dealbata (1Á7 pg) reported by Blakesley et al. (2002) is
similar to that reported by Bukhari (1997) of 2C = 1Á6 pg
(listed in Bennett et al., 2000), but both values differ con-
siderably from the 2C-value of 2Á9 pg reported by Mukerjee
and Sharma (1993b) (see Notes to the Appendix bb).
Another example concerns Brachypodium distachyon.
In 1991, the PhD thesis of Shi reported a 1C-value of
0Á15 pg, but later Shi et al. (1993) gave its 1C-value as
0Á3 pg. To resolve this discrepancy, RBG, Kew obtained
some original material studied by Shi and estimated its
1C-value to be 0Á36 pg, confirming the larger C-value for
this species (see also footnote br). Thus, real errors can be
identified with certainty, and potential errors flagged up for
users in cautionary footnotes following Appendix tables.
DNA C-values in angiosperms vary approx. 1000-fold
(over three orders of magnitude) from approx. 0Á1 pg to
over 100 pg. It is, therefore, often useful to know whether
a species' 1C DNA amount has approximately 0Á1, 1, 10 or
100 pg, even if there is still uncertainty regarding whether a
species with approximately 1 pg is really closer to 0Á8 pg
than to 1Á2 pg (an error 620 %). In terms of its predictive
value in nucleotypic correlations, such an error still permits
useful conclusions to be drawn. The Arabidopsis com-
munity laboured long under the misapprehension that its
1C DNA amount was approx. 70 Mb (Leutwiler et al.,
1984), and later approx. 100 Mb (Meyerowitz, 1994),
when in reality it is much higher (about 157 Mb, Bennett
et al., 2003). The level of inaccuracy involved (approx.
50­100 %) was considerable, yet it did not prevent the
selection of Arabidopsis as the model plant for first
complete genome sequencing, in no small part on the
basis of its `small genome size' (NSF, 1990; Somerville
and Somerville, 1999). The Convention on Biological
Diversity (United Nations Environment Programme,
1992) noted the need to make biodiversity data available,
despite imperfections; a view which merits support
(Bennett, 1998). Thus, it is better to list available C-value
data subject to errors, until improved data with fewer errors
become available. The body of data is needed by the
scientific community and can clearly already be used to
draw important conclusions, to make valuable predictions,
and as a basis for necessary planning.
What is the smallest reliable C-value for an angiosperm?
The above examples show how seeing data in the
comparative context of the database can help to identify
real or potential errors in particular species. It can also
facilitate broader enquiries such as `what is the smallest
reliable C-value for an angiosperm?'. Again, the comparat-
ive approach has enabled researchers to be active in iden-
tifying potential errors in species with the smallest reported
C-values, and to be transparent in correcting mistakes.
Because of error variation, a population of 1C-value
estimates for one taxon should vary according to a normal
curve, so those in the lower tail are all too low (Fig. 3A).
Too low Too high
Numberofestimates
1C DNA amount
0
20
40
60
0­010 011­020 021­030
1C DNA amount range (pg)
Numberofspecies
A
B
AboutAbout
rightright
About
right
FI G . 3. (A) Expected error variation in a large population of DNA C-value
estimates for one genotype as underestimates (in the lower tail) and
overestimates (in the upper tail) surround more accurate, intermediate,
genome size estimates. (B) Histogram showing frequency of C-values for
the 85 smallest species in the database or Appendix.
52 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
Such values are lost in the frequency histogram for all
angiosperm C-value estimates except at its lowest tail
where some of the lowest C-values claimed are expected
to be too low. This expectation is strongly supported in
practice, as shown below. There are 53 1C estimates in
the Angiosperm DNA C-values database or the present
Appendix with 0Á21­0Á30 pg, 29 with 0Á11­0Á20 pg,
but only three with 0Á10 pg or below (Fig. 3B). Table 2
lists the 24 lowest estimates listed with 0Á175 pg or less, but
how robust are they?
Thirteen of the 24 estimates in Table 2 are for
Arabidopsis thaliana. A comparative approach suggests
that some, which featured among the lowest C-values
reported for angiosperms, are too low. Thus several
C-value estimates made by molecular means in the range
0Á05­0Á125 pg (Leutwiler et al., 1984; Francis et al., 1990;
Arabidopsis Genome Initiative, 2000) are now seen as gross
underestimates, while many others in the range 0Á15­0Á18 pg
are shown to span the true value of about 0Á16 pg (Bennett
et al., 2003).
After discounting the 1C estimate for Arabidopsis
thaliana of 0Á051 pg by Francis et al. (1990), the next
smallest estimate listed is 0Á055 pg for Cardamine amara
(communicated from S. R. Band in 1984). With only a third
the DNA amount of its related crucifer A. thaliana (0Á16 pg),
it seemed suspiciously low. Cardamine amara seed cannot
survive drying, so it is unavailable from seed banks.
However, we recently used flow cytometry to compare
diploid C. amara collected near Sheffield with several cali-
bration standards including A. thaliana ecotype `Columbia'.
The 1C-value we obtained was around 0Á24 pg (almost five-
fold the earlier report). This is in close agreement with
independent estimates made elsewhere (e.g. see Bennett
and Leitch, 2005; Johnston et al., 2005).
Once the underestimates for Arabidopsis thaliana
and Cardamine amara are discounted, few 1C-values of
0Á125 pg or below remain for other species. One is the
1C estimate of 0Á125 pg for Rosa wichuriana, estimated
using callus material from Dr Andy Roberts (Bennett and
Smith 1991). This value seemed questionably low in the
context of the database, especially as it became clear that
culturing may induce stain inhibitors. This concern led to
a new collaboration with RBG, Kew using non-callous
material, and our doubts were confirmed when it was
re-estimated as 1C = 0Á575 pg (Yokoya et al., 2000).
Perhaps all estimates below 0Á125 pg should be
doubted until confirmed. Another candidate was Aesculus
hippocastanum whose 1C-value was listed as 0Á125 pg
(Bennett et al., 1982). This material is rich in tannins
and a likely candidate for underestimating its DNA amount
(Noirot et al., 2000, 2005). Recent work using flow cytome-
try at RBG, Kew, showed that the 1C-value of 0Á125 pg was
clearly an underestimate, as the true value is approx. 0Á60 pg
(L. Hanson, RBG, Kew, pers. comm.). With 0Á125 pg for A.
hippocastanum rejected, only one estimate below 0Á14 pg
remains, namely 0Á11 pg for the Green strawberry, Fragaria
viridis. Since there is considerable interest in knowing the
smallest possible angiosperm genome, checks to establish
whether this estimate is robust are now urgently required.
What is the minimum C-value for a free-living angiosperm
and other free-living organisms?
Such comparative approaches can also facilitate broader
questions such as: `what is the minimum genome size in
angiosperms and other free-living organisms?'. There is a
minimum compendium of nuclear genes essential for the
life of any organism. This concept was behind Craig
Venter's declared intension to synthesize from scratch a
minimal bacterial genome (Check, 2002), and a project
for a minimal eukaryote genome may eventually follow.
Meanwhile we can only speculate on how small the
minimum genome is for an angiosperm, and how closely
extant species approach the minimum. It is, of course, below
the lowest robust C-value for the group, i.e. less than 0Á16 pg
established for Arabidopsis thaliana. The presence of six
other species with C-values of 0Á15­0Á169 pg in Table 2
strongly supports this conclusion. The estimate(s) of approx.
0Á108 pg for Fragaria viridis may indicate a minimum
C-value for extant angiosperms of about 100 Mb, but if
so, is it a diploid, or a polyploid with three or more even
smaller ancestral genomes?
Whilst the robust 1C-value for A. thaliana is 0Á16 pg, this
includes >25 % of repeated DNA (Bennett et al., 2003) and
analysis of sequenced regions shows that >70 % of coding
genes are duplicated (Bowers et al., 2003). Thus, in theory,
a minimal genome without duplicated coding genes or rep-
etitive DNA should not exceed approx. 50 Mb. Currently,
there is no robust 1C estimate below 0Á1 pg for an angio-
sperm, but if any of the seven species with C-values
TA B L E 2. The 24 lowest angiosperm 1C DNA estimates among
data listed in the present Appendix table and the Angiosperm
DNA C-values database (release 4.0, January 2003)
Taxon 1C (pg) Original reference
Arabidopsis thaliana 0.051 Francis et al. (1990)
Cardamine amara 0.055 Band SR (pers. comm. 1984)
Arabidopsis thaliana 0.073 Leutwiler et al. (1984)
Fragaria viridis 0.108 Antonius and Ahokas (1996)
Rosa wichuriana 0.125 Bennett and Smith (1991)
Aesculus hippocastanum 0.125 Bennett et al. (1982)
Arabidopsis thaliana 0.128 Arabidopsis Genome Initiative (2000)
Sedum album 0.145 Hart (1991)
Arabidopsis thaliana 0.150 Arumaganathan and Earle (1991)
Carex nubigera 0.150 Nishikawa et al. (1984)
Carex paxii 0.150 Nishikawa et al. (1984)
Epilobium palustre 0.150 Band SR (pers. comm. 1984)
Hypericum hirsutum 0.150 Hanson, Leitch and Bennett
(pers. comm. 2002)
Thlaspi alpestre 0.150 Band SR (pers. comm. 1984)
Arabidopsis thaliana 0.153 Bennett et al. (2003)
Arabidopsis thaliana 0.160 Bennett et al. (2003)
Arabidopsis thaliana 0.160 Galbraith et al. (1991)
Arabidopsis thaliana 0.165 Galbraith et al. (1991)
Arabidopsis thaliana 0.167 Krisai and Greilhuber (1997)
Arabidopsis thaliana 0.167 Bennett et al. (2003)
Amoreuxia wrightii 0.168 Hanson et al. (2001a)
Arabidopsis thaliana 0.170 Galbraith et al. (1991)
Arabidopsis thaliana 0.175 Bennett and Smith (1991)
Arabidopsis thaliana 0.175 Marie and Brown (1993)
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 53
between 0Á14­0Á15 pg is a tetraploid, this would indicate a
minimum genome size in extant taxa of approx. 75 Mb, or
approx. 50 Mb if it is a hexaploid.
The comparative approach is usefully extended to include
other groups of organisms. Table 3 shows minimum C-value
estimates for multicellular organisms in several widely
differing groups obtained by genome sequencing or other
methods. Such minima for groups as diverse as nematodes,
insects, algae and angiosperms range from 59 to 160 Mb.
Thus, the minimum C-value known in extant free-living
multicellular higher organisms is around 60 Mb. All may
be diploidized paleopolyploids (Wendel, 2000), but except
for one early and unconfirmed report of a 1C-value of
approx. 39 Mb in a most simple multicellular placozoan
animal (Ruthmann and Wenderoth, 1975) there is no evi-
dence for extant diploid multicellular eukaryotic life forms
with only 40­50 Mb. This tantalizing possibility will be an
interesting driver for new work to find a first angiosperm
whose 1C-value is <100 Mb, or a first free-living multi-
cellular plant or animal with a robust 1C-value <50 Mb.
PROSPECTS FOR THE NEXT TEN YEARS
Apart from better defining the limits of genome size
variation, what key developments are targeted, or likely,
to occur in angiosperm genome size research in the next
decade?
The first concerns the expected progress to increase the
total number and representation of angiosperms in the
C-values database. As noted above, estimating first values
for species reached a historic high during recent years
(Fig. 1). At least 1700 such values were added in 1997­
2002, and the total number of species' C-value estimates
probably reached around 4300. In 2003 the second Plant
Genome Size Workshop set a goal of estimating a further
1 % (i.e. approx. 2500 species) in the next five years, and a
similar target is likely for the following quinquennium.
If so, there is a reasonable prospect that the number of
species with a C-value estimate will reach, or significantly
exceed 7500 by 2014.
More important than the expected increase in total
numbers is the predicted improvement in the spread of
new values across taxa, geographical regions and life
forms, making the sample more representative of the global
angiosperm flora, based on careful targeting to identify and
fill knowledge gaps. The next decade should see almost
complete representation for families, and a greatly increased
representation for genera (especially in monocots), as work
focuses increasingly at this taxonomic level. Representation
at the generic level is currently approx. 1042 out of an
estimated 14 000 genera (7Á4 %) and is targeted to rise to
10 % by 2009, and might reasonably be expected to
approach 15 % within a decade. Moreover, this may
approach 100 % for monocots, as they are targeted for
holistic genomic studies (including C-values) for the global
Monocot Checklist Project (Govaerts, 2004).
Recent experience shows that identifying a gap and
setting a target may still not provoke the work needed to
fill it. Positive monitoring of trends in published C-value
data may also be required to achieve a significant change in
research activity (e.g. as with the level of family representa-
tion in angiosperms; Hanson et al., 2003). Thus it will be
important to monitor by 2009 whether the gaping chasms in
the representation of African, South American and Chinese
floras noted previously have yet resulted in a significant rise
in first estimates for taxa from those regions. If not, then a
major effort will be needed to correct this. The same applies
to other groups of plants that have been identified as poorly
represented in the Angiosperm DNA C-values database
(e.g. halophytes, parasitic species and their hosts and tundra
species). Whilst less certain, there is a good prospect that
this vital process will occur in the next decade, driven by
the Genome Size Initiative (GESI: see Bennett and Leitch,
2005).
TA B L E 3. Robust minimum 1C-value estimates for several widely different groups of free-living, multicellular, higher
organisms obtained by genome sequencing (*), other best practice techniques, or static cytometry using the fluorochrome
DAPI for algae1
Group Species Mb Original reference
ANIMALS
Nematode Caenorhabditis elegans 100* C. elegans sequencing Consortium (1998)
Platyhelminthes (flatworms) Stenostomum brevipharyngium 59 Gregory et al. (2000)
Crustacea Scapholeberis kingii (water flea) 157 Beaton (1988)
Annelid Dinophilus gyrociliatus (polychaete worm) 59 Soldi et al. (1994)
Tardigrades (water bears) Isohypsibius lunulatus 78 Redi and Garagna (1987)
Insect Peristenus stygicus 98 TR Gregory (pers. comm.)
Arachnid Tetranychus urticae (spider mite) 78 TR Gregory (pers. comm.)
Urochordates (tunicates) Oikopleura dioica 72 Seo et al. (2001)
PLANTS
Chlorophyta (green alga) Caulerpa paspaloides 88 Kapraun (2005)
Rhodophyta (red algae) Heydrichia wolkerlingii 69 Kapraun (2005)
Phaeophyta (brown algae) Stilophora rhizodes 98 Kapraun (2005)
Bryophyte Holomitrium arboreum 167 Voglmayr (2000)
Lycophyte Selaginella kraussiana 157 Obermayer et al. (2002)
Angiosperm Arabidopsis thaliana 157 Bennett et al. (2003)
1
Theuseofthebase-specificfluorochromeDAPIforestimatingDNAamountsmaybelessreliablethanusingintercalatingfluorochromessuchaspropidium
iodide (e.g. Dolezzel et al., 1992).
54 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
Hitherto, when a question regarding C-value was framed
(e.g. is genome size related to weediness?), it was often
necessary to estimate C-values for many species before it
could be addressed (Bennett et al., 1998). Clearly, the prime
aim is to create a sample of C-values that is sufficiently
representative for systematic, regional and life form varia-
tion as to allow most questions to be answered with
confidence using the available dataset, without recourse
to further C-value estimations. This goal is likely to be
achieved in the next ten years. Thus, the next decade
may be the last to see major efforts devoted to estimating
first DNA C-values for taxa. Thereafter, new C-value
research will probably concentrate on using and under-
standing such data, rather than acquiring them.
What important questions regarding genome size in
angiosperms are likely to be answered in the next decade?
Three closely interrelated issues concerning the possible
significance of genome size for extinction, conservation
and pollution are worth mentioning here.
The possibility that a large C-value might correlate with
an increased risk and rate of extinction was suggested by
Rejmanek (1996) and by Bennett et al. (2000). To test this,
Vinogradov (2003) identified 3036 diploid species from
the Plant DNA C-values database and compared each
one against the United Nations Environmental Programme
World Conservation Monitoring Centre (UNEP-WCMC)
species database to determine its conservation status (i.e.
global concern, local concern or no concern). He noted a
striking relationship between genome size and conservation
status; species with large genomes appeared to be at greater
risk of extinction that those with smaller genomes.
Clearly, this was an important finding that now requires
independent confirmation, drawn from further independent
samples of species in different local regions and environ-
ments. Obtaining data for meaningfully large samples of
species for such studies will probably be one main driver
determining which taxa are targeted for C-value estimates in
the future. If so, the next decade offers the prospect of a
more definite and detailed understanding of any relation-
ships between C-value and/or genome size and the risk of
extinction. This, in turn, may have important practical
and theoretical implications for conservation models and
strategies. A key question is whether a large nuclear
DNA amount gives an increased risk of extinction equally
in diploid or polyploid taxa? Vinogradov (2003) tested
whether ploidy played a role in increasing a species' risk
of extinction, concluding that C-value per se was most
important. Polyploidy is supposed to confer many advant-
ages based on increased gene dosage and diversity, but do
such advantages overcome the possible risks of a high
C-value? If so, the proportion of polyploids should be higher
for species with very high C-values than for those with
lower C-values. In a test different from that of Vinogradov
(2003), we compared the percentage of polyploids in 3400
extant species with known DNA amount and ploidy level in
the Angiosperm DNA C-values database, ranked in order of
increasing DNA amount and divided into five groups each
containing 680 species. We found the percentage of poly-
ploids for species in group 5 with the highest C-values
(29Á9 %) was actually lower than for species in group 4
(32Á1 %) (Fig. 4). This confirms Vinogradov's finding
that the prime factor determining increased risk of extinc-
tion is high C-value, and that polyploidy does not reduce
this risk.
Other enquiries should test whether the risk of extinction
in relation to high C-value or genome size varies for
different threats and environments. This should compare
variation in internal factors affecting the structure and eco-
logy of the genome (e.g. increased ploidy level, and hetero-
chromatin distribution), and in external factors (e.g.
pollution and increased competition for space, minerals,
light, and pollinators). Vilhar (pers. comm., and Vidic
et al., 2003) investigated the effect of genome size on
plant survival in lead-polluted soils. With increasing lead
concentration in the soil the percentage of species with
large genomes decreased significantly, suggesting that spe-
cies with large genomes were at a selective disadvantage.
Similar work on local floras in different areas with various
threats is now needed to test whether their results are typical
for other pollutants and environments. Understanding which
species survive locally is always important, but especially
as local loss equals global extinction if a species range
is restricted to just that one locality. Such work will
increasingly inform local environmental action plans and
conservation strategies.
Holistic genomics
Early interest in plant genome size variation (c. 1950s
and 1960s) ranged broadly across many fields including
its genetic, developmental, ecological and evolutionary
implications. However, after the molecular revolution the
field fragmented somewhat as interest in DNA sequences
was largely separated from more macro interests in C-
values. However, given `complete sequences' for genomes
and homoeologous segments, and greater computing
299321
232256
132
0
20
40
60
80
100
1 2 3 4 5
Group
%ofspecies
FI G . 4. The percentage of diploids (open bars) and polyploids (closed bars)
among 3400 species of known DNA amount and ploidy level ranked in order
of increasing DNA amount and divided into five groups with 680 species per
group. Data taken from the Angiosperm DNA C-values database (release
4.0, January 2003) and the present Appendix.
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 55
power, this post-genomic age is seeing a strong converg-
ence of these interests. Thus, leading scientists who work at
comparative sequence levels can also work on questions of
genome size and evolution (e.g. Zhang and Wessler, 2004;
Bennetzen et al., 2005). This is the age of holistic genomics
in which knowledge of variation in genome size and
C-value can be seamlessly joined up with information at
all other levels to embrace information from sequences to
ecology and from evolution to the environment. This power-
ful approach should permit or provoke quantum leaps in
understanding the significance of extant variation in C-value
and genome size, the processes that produce it, the rate at
which it occurs, the factors that limit its extent and the
advantages and disadvantages that it confers. Together,
such understanding will link across biological fields to
explain patterns of genome size variation in development,
floras, ecological niches and evolution. The next ten years
offer many exciting prospects for angiosperm genome size
research. Work on DNA amount will remain a key core
interest in biological research, but will increasingly become
one integrated strand in holistic genomic studies and under-
standing, covering its origin(s), mechanisms of change,
phenotypic and phenological effects, and its significance
for ecological, developmental and environmental issues.
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APPENDIX
Notes to the Appendix
The Appendix appears on pp. 59­88.
Named references in the following notes are given above
in `Literature Cited', while numbered references are given
in `Original references for DNA values' below.
(a) The original references for species DNA amounts
in the Appendix are given in a numbered list following
the Appendix table. Reference numbers follow on sequen-
tially from those given in `Notes to Table 8' by Bennett and
Smith (1976, references 1­54) `Notes to Table 1' by Bennett
et al. (1982, references 55­107), Bennett and Smith (1991,
references 108­163), `Notes to the Appendix' by Bennett
and Leitch (1995, references 164­269), `Notes to the
Appendix' by Bennett and Leitch (1997, references 270­
306), and `Notes to the Appendix' by Bennett et al. (2000,
references 307­377).
(b1) Bennett and Smith (1991) gave absolute 4C DNA
values for 11 angiosperm species recommended for use as
calibration standards to estimate DNA amounts in other
species. These species and their 4C DNA amounts are
given in Table 4. If a species was calibrated in direct com-
parison with any one or more of the 11 standard species then
the standard species used is identified in column 15 of the
Appendix by the appropriate Key letter given above (e.g. F is
Hordeum vulgare, etc.). If a species was first calibrated
using a standard species listed above, then the original
standard species is identified first and the intermediate stand-
ard species used to calibrate those species listed with it is
also denoted by its number in column 1 of the Appendix.
For instance, standard G (P. sativum) was used to calibrate
Capsicum annuum `Doux Long des Landes' (species 212 h
in the Appendix), which was then used as an intermediate
standard to estimate other Capsicum species given by
Belletti et al. (1998, Ref. 434). The calibration standard
for such Capsicum species is therefore given as G-212h.
(b2) In Refs 444 (Wendel et al., 2002) and 447 (Lin et al.,
2001) Pisum sativum `Minerva Maple' was used as the
calibration standard but they assumed a 4C DNA value
of 19Á12 pg (Johnston et al., 1999) instead of 19Á46 pg,
which is the value given in Bennett and Smith (1976)
and listed in Table 4. The 4C-value of P. sativum `Minerva
Maple' used in Refs 444 and 447 was estimated using
Hordeum vulgare `Sultan' as the calibration standard
with an assumed 4C DNA content of 22Á24 pg (Johnston
et al., 1999).
(c) In several references listed in `Original references for
DNA values' the authors used a cultivar of a standard
species different from that listed in Table 4, these are listed
in Table 5. In some cases the C-value of the cultivar used
was assumed to be the same as that of the cultivar given in
Table 4. Evidence of intraspecific variation in a number of
species suggests that such assumptions may sometimes be
incorrect. In other cases the C-value of the cultivar was
determined by the authors and was different from that of
the standard species listed in Table 4. For example Refs 386,
TA B L E 4. The eleven angiosperm species recommended for
use as calibration standards (see Notes to the Appendix, b1)
Key Standard species 4C DNA amount (pg)
A Triticum aestivum `Chinese Spring' 69.27
B Allium cepa `Ailsa Craig' 67.00
C Vicia faba PBI, inbred line 6 53.31
D Anemone virginiana line AV 200 35.67
E Secale cereale `Petkus Spring' 33.14
F Hordeum vulgare `Sultan' 22.24
G Pisum sativum `Minerva Maple' 19.46
H Zea mays `W64A' 10.93
I Senecio vulgaris (PBI population) 6.33
J Vigna radiata `Berken' 2.12
K Oryza sativa `IR36' 2.02
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 59
397, and 453 used the cultivar `Express Long' of Pisum
sativum with a 4C DNA value of 16Á74 pg. This value is
lower than the 4C DNA amount of the cultivar `Minerva
Maple' of 19Á46 pg given in Table 4.
(d) In References 407, 408 and 457 the cultivar of the
calibration standard was not given. Refs 407 and 408 used
Vicia faba as a calibration standard, whereas Ref. 457 used
Allium cepa. In Ref. 408 Cremonini et al. (1992) assumed
the same 4C-value for Vicia faba as for PBI line 6 (i.e.
53Á3 pg) given in Table 4. If this species exhibits intra-
specific variation then such assumptions may be incorrect.
(e) In a number of original references the authors used a
plant species not listed in Table 4 as a calibration standard.
These are listed in Table 6.
(f ) Several papers listed in `Original references for DNA
values' used animal cells as the calibration standards. Thus
Refs 387, 417, 426, 442, 456, 463 used chicken erythrocytes
with an assumed 4C DNA value of 4Á66 pg (Galbraith et al.,
1983). The calibration standard is abbreviated to Gallus in
column 15 of the Appendix. In Ref. 438 blood cells from the
catfish Ictalurus punctatus were used as a standard with an
assumed 4C-value of 4Á00 pg (Tiersch et al., 1989), this is
abbreviated to Ictal. in the Appendix. In Ref. 438 domestic
swine (Sus scrofa) erythrocytes were used as a standard with
an assumed 4C-value of 11Á34 pg (Taliaferro et al., 1997),
and is abbreviated to Sus in the Appendix. Human cells with
an assumed 4C DNA amount of 14Á00 pg (Tiersch et al.,
1989) were used as calibration standards in Refs 384, 428
and 419 (leucocytes, Ref. 384, 428; lymphocytes, Ref. 419)
and the abbreviation of Homo is used in the Appendix.
Finally, Drosophila melanogaster with an assumed 1C DNA
amount of 180 Mb (Adams et al., 2000) and Caenorhabditis
elegans with a 1C DNA amount of 100Á25 Mb, based on
complete genome sequencing (see C. elegans Sequencing
Consortium, 1998 and http://wormbase.org), were used as
calibration standards for Ref. 461, and the abbreviation of
Dros. and Caeno. respectively are used.
If a plant species was calibrated using an animal species
and then subsequently used as the calibration species for
other plants, then the animal species is identified first,
and the intermediate plant species is identified by its
entry number given in column 1 of the Appendix. Thus
Mishiba et al. (2000, Ref. 387) used Gallus with an assumed
4C DNA amount of 4Á66 pg (Galbraith et al., 1983) to
calibrate Hordeum vulgare `New Golden' (species 398p
in Appendix), this was then used as the calibration standard
to estimate DNA C-values of Petunia and Calibrachoa
species given by Mishiba et al. (2000). The calibration
standard for these Petunia and Calibrachoa species is
given as Gallus-398p.
(g) When a new estimate (or estimates) is given for a
species or subspecies already listed by Bennett and Smith
(1976, 1991), Bennett et al. (1982, 2000) or Bennett
and Leitch (1995, 1997), the estimate is given a number
and a lower case letter in column 1 of the Appendix. An `a'
implies that the value is preferred to any estimate for
that species listed previously by the first author. Where
several estimates are available for the same species, the
`a' value would automatically be chosen in any arithmetical
or statistical calculations. In this context, single estimates
for species and `a' values are referred to as `prime
entries'.
(h) Intraspecific variation in nuclear DNA amount is
claimed to occur in this species. Consequently the values
given in the Appendix should not be assumed to be correct
for all accessions of the species. Where several C-values
are listed for a single species with the same ploidy level
or chromosome number within a taxon, then only the
minimum and maximum values reported from a single
reference are listed in the Appendix.
(i) A range of DNA amounts was reported for this species
in the reference cited in column 13 of the Appendix. Intra-
specific variation was not claimed to occur, so the nature of
this variation is unclear. Where estimates differed by more
than 10 % the minimum and maximum values are given
for the same ploidy level or chromosome number in the
Appendix, otherwise only the highest value is given.
(j) According to the International Code of Botanical
Nomenclature (Greuter et al., 1994), the names of plant
families must end in -aceae. However, eight plant families
are exceptions in that each has two alternative names, both of
TA B L E 5. Cultivars of standard species used that differ from
those listed in Table 4
Original
reference number
Plant calibration
standard used
Assumed 4C DNA
amount and reference
(pg)
Allium cepa
413 `Fruuhstamm' 67.0 (reference not given)
409 `Kantar topu' 67.0 (Van't Hof, 1965;
Bennett and Smith, 1976)
435, 443, 449,
454, 455
`Nasik Red' 67.0 (Van't Hof, 1965)
427 `Stuttgarter
Reisen'
67.0 (Bennett and Smith, 1976)
388, 411,
420, 422
var. aggregata ­ (amount and reference not given)
Hordeum vulgare
417 `Stark' 21.36 (reference not given)
423 strain NE 86954 20.48 (Lee et al., 1997)
Oryza sativa
424 type japonica 2.20 (Bennett and Smith, 1991)
Pisum sativum
386, 397, 453, `Express Long' 16.74 (Marie and Brown, 1993)
393, 394, 395,
396, 429, 458
`Kleine
Rheinla¨nderin'
17.68 (Greilhuber and Ebert, 1994)
434 `Lincoln' 18.14 (Dolezzel et al., 1992)
418 ssp. sativum
convar. sativum
var. ponderosum
`Viktoria'
18.18 (Dolezzel et al., 1998)
Secale cereale
418 ssp. cereale 32.38 (Dolezzel et al., 1998)
394 `Dankovske' 31.16 (Dolezzel et al., 1998)
Vicia faba
418 ssp. minor var.
minor subvar.
rigida `Tinova'
54.00 (Dolezzel et al., 1998)
405 `Aquadulce' 53.31 (pers. comm. 2002)
430 `Superguadulce' 53.30 (Bennett and Smith, 1976)
Zea mays
391 `CE-777' 10.86 (Lysák and Dolezzel, 1998)
437 `C-777' 10.86 (Lysák and Dolezzel, 1998)
382 Va35 10.93 (Bennett and Leitch, 1995)
60 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
which are correct under the Botanical Code. One is a standard
name, ending in -aceae, the other is an exception, sanctioned
by long usage. These and their alternatives are the follow-
ing: Palmae (Arecaceae), Gramineae (Poaceae), Cruciferae
(Brassicaceae), Leguminosae (Fabaceae), Guttiferae
(Clusiaceae), Umbelliferae (Apiaceae), Labiatae
(Lamiaceae) and Compositae (Asteraceae). To be con-
sistent with previous DNA lists (Bennett and Smith,
1976, 1991; Bennett et al., 1982, 2000; Bennett and
Leitch, 1995, 1997) the `non-standard' plant names are
retained in the present work.
(k) Recent cladistic analysis using both molecular and
non-molecular phylogenetic data has resulted in a revised
classification of families by the Angiosperm Phylogeny
Group (APG) (APG II, 2003). Familial names used in
the APG classification are followed in the Appendix.
Thus, although Zonneveld (2002, Ref. 440) placed Aloe
in Aloeaceae, recent molecular and non-molecular phylo-
genetic data recognizes that this family is embedded within
the newly circumscribed Xanthorrhoeaceae (APG II, 2003)
so Xanthorrhoeaceae is given in the Appendix. Similarly,
the APG II (2003) now recognizes that Hostaceae is
embedded within the Asparagaceae, so Hosta, which was
placed in Hostaceae in Ref. 384 (Zonneveld and Van Iren,
2000), is listed under Asparagaceae in the Appendix.
(l) The authority for this species is either unknown or
unclear to the present authors.
(m) Whether or not voucher specimens exist for this
species is unknown to the present authors.
(n) The chromosome number of this species is either
unknown or unclear to the present authors.
(o) The chromosome count for this species was taken
from the literature and not determined by the authors of
the reference cited.
(p) The ploidy level of this species is either uncertain or
unclear to the present authors.
(q) The life cycle type of this species is either unknown or
unclear to the present authors.
(r) The method used to measure the DNA amount is
unclear.
(s) The factor of 1 pg = 980 Mbp was used to convert
picograms to Mbp (Cavalier-Smith, 1985; Bennett et al.,
2000).
(t) As a rule, replicated diplophase nuclei contain a 4C
DNA amount producing two unreplicated 2C nuclei by
mitotic division and four 1C gametic nuclei after meiosis
(irrespective of ploidy level). This convention applies well
to polyploid taxa with diploidized meiotic chromosome
pairing which produce functional balanced polyhaploid
gametes with 1C DNA amounts at meiosis. Thus 4C esti-
mates were automatically divided by 4 to generate 1C-
values given for all taxa of even ploidy level listed in the
Appendix. However the resulting `1C' data are not biologi-
cally meaningful for taxa with odd ploidies. Consequently
the Appendix gives only 2C- and 4C-values for such taxa.
(u) There is no obvious basic number for the genus
Luzula due to the presence of holocentric chromosomes.
It is therefore impossible to allocate Luzula species
TA B L E 6. Plant species used as calibration standards but not listed in Table 4
Original reference
number
Plant calibration
standard used
Assumed 4C DNA amount and reference
(pg)
Abbreviation used in
column 15 of Appendix
383 Agave americana 31.80 (Zonneveld and Van Iren, 2001) Agave
Arabidopsis thaliana
389 `Columbia' ­ (amount and reference not given) Arab.
414 `Columbia' 0.53 (Kaneko et al., 1998) ''
427 Cerastium eriophorum 5.20 (Boscaiu et al., 1999) Cerastium
Glycine max
393 `Ceresia' 4.54 (Greilhuber and Obermayer, 1997) Glycine
429 `Ceresia' 4.51 (Obermayer and Greilhuber, 1999) ''
421 var. Palmetto 5.00 (Dolezzel et al., 1994) ''
432 `Burlison' 5.56 (Graham et al., 1994) ''
402 `Polanka' 5.00 (Dolezzel et al., 1994) ''
Lycopersicon esculentum
380, 382, 465 `Gardener's Delight' 4.00 (Obermayer et al., 2002) Lycopers.
437 `Stukické' 3.92 (Dolezzel et al., 1992) ''
385 `Montfavet' 4.02 (Marie and Brown, 1993) ''
Nicotiana tabacum
427 `Petit Havana SR1' 18.00 (Bennett and Leitch, 1995) Nicot.
417 `Samsun' 18.15 (reference not given) ''
Petunia hybrida
390, 410, 433, 452 `PxPC6' 5.70 (Godelle et al., 1993; Marie and Brown, 1993) Petunia
390 `Hit Parade Blau' R 5.70 (Marie and Brown, 1993) ''
401 No cultivar given 5.70 (Marie and Brown, 1993) ''
Rhaphanus stativus
404 `Saxa' 2.20 (Dolezzel et al., 1992) Rhaphanus
Sorghum bicolor
439 Line TX623 3.52 Price and Levin (pers. comm.) Sorghum
462 No cultivar given 3.20 (Bennett and Smith, 1991) ''
406 Vicia narbonensis 29.10 (Frediani et al., 1992) Vicia narb.
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 61
with high chromosome numbers to any ploidy level with
certainty.
(v) Unal and Callow (1995, Ref. 412) obtained a
regression of the nuclear fluorescence of Allium cepa
(4C = 67Á0 pg), Crepis capillaris (4C = 9Á6 pg), Hordeum
vulgare (4C = 22Á2 pg), Pisum sativum (4C = 20Á2 pg),
Secale cereale (4C = 33Á2 pg) and Vicia faba (4C =
47Á9 pg) versus nuclear DNA content, and used this to
estimate the DNA C-values of 13 Lathryus species.
However, it is noted that the 4C-values for P. sativum,
and Vicia faba are non-standard values compared with
those given for these species in footnote (b1) above.
(w) The standard species used to convert arbitrary units
into absolute DNA amounts is unclear to the present
authors.
(x) The DNA value given for this species in the original
reference differs considerably (i.e. >100 %) from that given
in other original references cited in previous compiled lists
of DNA amounts (i.e. Bennett and Smith, 1976, 1991;
Bennett et al., 1982, 2000; Bennett and Leitch, 1995,
1997). The reason(s) for this is unknown. This C-value
should therefore be used with caution until the question is
resolved.
(y) The specific status of the material available for study
is unclear. The data are included since information on DNA
amounts for this genus is relatively sparse, so an indication
of genome size in the genus may be useful.
(z) Zonneveld (2001, Ref. 383) gave C-values for
16 hybrid cultivars which fall within the range that he
reported for Helleborus species listed in the Appendix.
Our compiled lists have usually been restricted to C-values
for species. Following this practice, C-values for Helleborus
hybrids were not included in the present Appendix.
(aa) Zonneveld and Van Iren (2000, Ref. 384) gave DNA
amounts for 94 accessions of Hosta which were recognised
as 23 different species. Their table 1 gives a DNA amount
for each accession together with a mean value for each
recognised species. Only the later value is given in
the present Appendix. They also included C-values for
16 Hosta cultivars (in their table 3). These were once
recognized as species, but following pollen viability tests
Zonneveld and Van Iren (2000) concluded they were
hybrids. Our compiled lists have usually been restricted
to C-values for species. Accordingly, C-values for Hosta
hybrids were not included in the present Appendix.
(ab) Zonneveld and Van Iren (2000, Ref. 384) and
Zonneveld (2002, Ref. 440) used male human leucocytes
(2C = 7Á0 pg; Tiersch et al., 1989) as their primary standard
to estimate the DNA amount of three Agave species,
namely: (i) Agave stricta (ii) A. americana and (iii) A.
sisalana. A. americana was then used as internal calibration
standard for most Hosta (Ref. 384) and Aloe (Ref. 440) taxa.
However, in a few cases where the DNA content of Hosta or
Aloe coincided with that of A. americana, one of the other
two Agave species was used as the internal standard.
As neither reference identified which Agave species was
used, the calibration standard in column 15 of the present
Appendix is given as Agave sp.
(ac) Thibault (1998, Ref. 385) claimed intraspecific
variation ranging from 6 to 11 % in the Salix species he
studied, but only a mean DNA C-value for each species was
given in his table 3. It is these values that are listed in the
present Appendix. Thibault (loc. cit.) also included
C-values for five hybrids. Our compiled lists have usually
been restricted to C-values for species, thus C-values
for Salix hybrids are not included in the Appendix. In
addition, Thibault (1998, Ref. 385) listed a C-value for
`S. triandra?' but concluded its identity was `hard to
specify'. Consequently, this taxon was not included in
the Appendix.
(ad) Thibault (1998, Ref. 385) used DNA C-values to
predict the ploidy level of each Salix species given in his
table 3, assuming direct proportionality. Moreover, their
chromosome numbers were not counted by him, but derived
by him assuming a constant basic chromosome number
of n = 19 for the genus. These predictions are entered in
columns 6 and 7 of the Appendix.
(ae) Some taxa once included in Petunia are now
included in Calibrachoa. The taxonomy for most Petunia
species listed in Mishiba et al. (2000, Ref. 387) follows that
of Wijsman (1990) who split the genus Petunia sensu
Jussieu (1803) into two; Petunia sensu Wijsman and
Calibrachoa. However, five species listed in Mishiba
et al. (2000) were not reclassified by Wijsman and so
they were listed under the generic name of Petunia sensu
Jussieu in Mishiba et al. (2000) although they `were
regarded as Calibrachoa' (see their table 3). By following
the taxonomy of Petunia sensu Wijsman, several species
originally listed under the genus Petunia, now belong to
Calibrachoa (e.g. parviflora was listed in the genus Petunia
by White and Rees, 1985, 1987) and this generic name was
used in the list of Bennett and Leitch (1995). Yet Mishiba
et al. (2000) assigned this species to Calibrachoa. To avoid
confusion readers looking under Petunia are referred to
Calibrachoa in the Appendix.
(af ) Joachimiak et al. (2001, Ref. 391) reported chromo-
some numbers and C-values for six Bromus species.
Chromosome numbers varied considerably in roots of
three species, but variation in C-values was `virtually
absent within leaf mesophyll cells'. The C-values given
by Joachimiak et al. (2001) were obtained using leaf
mesophyll cells and are listed in the Appendix.
(ag) The study by Rosato et al. (1998, Ref. 392) was
primarily concerned with polymorphism in Zea mays ssp.
mays races with B-chromosomes, but gave C-values only
for plants lacking B-chromosomes. Thus, they listed DNA
amounts for 17 populations which differed by 36 % (2C =
5Á008­6Á757 pg) in plants with 2n = 20. Similar intraspecific
variation in this species was reported previously (Laurie and
Bennett, 1985; Rayburn et al., 1985). Mean DNA amounts
for only the populations with the largest and smallest
C-values for A-chromosomes, are listed in the Appendix.
(ah) Dimitrova and Greilhuber (2000, Ref. 394) reported
significant intraspecific variation in Crepis biennis (P < 0Á05)
and C. sancta (P < 0Á01), some of which had variable
numbers of B-chromosomes. As only means were given
for material with 0­2 B-chromosomes, it was impossible
to give values (presumably the largest) for the 2B comple-
ment. Consequently, the Appendix just lists the smallest and
largest C-values for accessions without B-chromosomes.
62 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
(ai) Dimitrova and Greilhuber (2000, Ref. 394)
reported significant (P < 0Á001) intraspecific variation of
11 % for Crepis pulchra. They suggested that the two
accessions with the higher C-values may belong to sub-
species turkestanica. This is not recognized in the
Bulgarian flora (where these accessions were collected),
but was described by Babcock (1947). In the Appendix
the higher C-values listed for this species (entry
numbers 305c and e) may thus correspond to C. pulchra
ssp. turkestanica.
(aj) Temsch and Greilhuber (2000, Ref. 395) estimated
C-values in 11 accessions of Arachis hypogaea using both
Feulgen microdensitometry and flow cytometry. C-values
for different accessions showed great stability, so they
calculated a mean C-value for each method in the `Results
and Discussion' of their paper. Only these mean values are
listed in the Appendix.
(ak) Previous estimates for Vicia melanops (2n = 10)
(e.g. Chooi, 1971; Raina and Rees, 1983; Raina and
Bisht, 1988) all report a 4C-value of approx. 40 pg,
which is much higher than the value of 27Á6 pg given in
Cremonini et al. (1992, Ref. 408, entry number 780d). Thus,
this estimate should be viewed with caution until confirmed
independently.
(al) Akpinar and Bilaloglu (1997, Ref. 409) gave
a 2C-value of 13Á1 pg for Vicia cracca ssp. cracca
(with 2n = 2x = 14; their original count). However, six
previous reports for V. cracca listed in the database (Bennett
and Leitch, 2003) gave similar 2C-values (from 10 to 13 pg),
but for 2n = 4x = 28. The cause of this discrepancy is
unknown, thus the estimate by Akpinar and Bilaloglu
(loc. cit.) should be viewed with caution until confirmed
independently.
(am) Sakamoto et al. (1998, Ref. 414) estimated the
C-value of Cannabis sativa using Arabidopsis thaliana
`Columbia' (1C = 130 Mb, Kaneko et al., 1998) as the
calibration standard. However, the 1C-value assumed for
A. thaliana was low compared with its recently confirmed
estimate of 157 Mb (Bennett et al., 2003). If 157 Mb is
assumed for A. thaliana, then the 1C-value for C. sativa
would be 988 Mb = 1Á01 pg (female) and 1016 Mb = 1Á04 pg
(male).
(an) Gammar et al. (1999, Ref. 416) gave DNA amounts
for eight Lupinus species in arbitrary units (a.u.) listed as
Mn(x) values in their Figs. 1­4. Bennett and Smith (1976)
gave the 4C DNA amount of L. luteus as 4Á0 pg (allowing
for recalibration of Senecio vulgaris from 5Á88 pg to 6Á33 pg,
see Bennett and Smith, 1991). Gammar et al. (loc. cit.) gave
Mn(x) values for three L. luteus populations as 60Á4, 58Á4,
and 63Á8 in figure 1A, noting they were not statistically
different. The mean of these three values was calculated
to be 60Á86 a.u. To convert the Mn(x) values for each
Lupinus species into absolute DNA amounts, they were
multiplied by a conversion factor of 0Á07 (i.e. 4Á0 pg
, 60Á86 a.u.).
In some Lupinus species more than one population was
studied, and several Mn(x) values were listed. If these did
not differ significantly, the average Mn(x) value was calcul-
ated and converted into absolute DNA amounts. However,
chromosome counts of 2n = 38, 42 and 44 were reported in
L. angustifolius, so variation in Mn(x) may correspond to
different cytotypes.
Some absolute DNA amounts calculated for Gammar
et al. (loc. cit) do differ greatly from those previously
reported for the same species (e.g. L. pilosus, 4C = 4Á9 pg,
is almost double the value of 2Á5 pg given by Obermayer
et al., 1999). Similarly, the 4C-value of 3Á1 pg calculated for
L. angustifolius with 2n = 38, 42, or 44 is similar to the
estimate (4C = 3Á7 pg) by Barlow (pers. comm., listed in
Bennett et al., 1982), yet the latter was for material with
2n = 26. Data from Gammer et al. give a useful approxi-
mation of C-values in the five species not previously listed,
but should be treated with caution unless confirmed
independently.
(ao) Brandizzi and Grilli Caiola (1996, Ref. 419) gave
2n = 18 for Crocus biflorus in their table 1, but 2n = 8 in
the first paragraph of their text. They also stated in their
final paragraph: `However, C. biflorus and C. etruscus,
having half the chromosome number with respect to
C. thomasii and C. cartwrightianus . . . ..' As C. thomasii
and C. cartwrightianus were both recorded with 2n = 16
by Brandizzi and Grilli Ciola (1996), we conclude that
C. biflorus had 2n = 8, and so this number is entered in
the Appendix.
(ap) The 4C DNA amounts reported by Mukerjee and
Sharma (1993a, Ref. 420) for Luzula nivea and L. luzuloides
are over 50 % larger than those reported by Barlow (pers.
comm. 1976; reference 36 in Bennett and Smith, 1976).
The chromosome numbers for each species were the
same, so the cause of the discrepancy is unknown. How-
ever, Mukerjee and Sharma (1993a) used a single wave-
length method, which may suffer from distributional error
(Greilhuber, 2005, this volume). Thus, estimates for
Luzula in Mukerjee and Sharma (1993a) should be viewed
with caution until confirmed independently for these
species.
(aq) Asif et al. (2001, Ref. 421) estimated DNA amounts
in 14 genotypes of Musa acuminata. Genotype BC3
(belonging to the separate subspecies truncata) had the
highest DNA amount and its C-value was shown to be
significantly different (P < 0Á01) from the other thirteen
genotypes. Only the DNA amount of genotype BC3, corre-
sponding to M. acuminata ssp. truncata, and the highest
DNA amount out of the 13 other genotypes of M. acuminata
are entered in the Appendix.
(ar) Chaudhuri and Sen (2001, Ref. 422) examined two
Scilla indica cytotypes (entry numbers 710b and c) which
differed considerably in both DNA amount and karyotype
structure, although both had 2n = 30. The differences may
reflect problems with taxonomy. Studies by Greilhuber and
colleagues (Greilhuber, 1979; Greilhuber and Speta, 1985)
have shown that large intraspecific differences in C-values
in other Scilla species (e.g. S. bifolia) reduce to a level
hardly more than methodological error following taxonomic
splitting.
(as) Chung et al. (1998, Ref. 423) estimated C-values
in 12 soybean (Glycine max) strains varying in seed size.
They reported statistically significant differences of 4Á6 %
in the 2C-values between strains. Only the smallest and
largest C-values are entered in the Appendix.
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 63
(at) Hartman et al. (2000, Ref. 425) estimated C-values in
22 Leucaena species using flow cytometry. Three species
(Pisum sativum 4C = 17Á6 pg, Oryza sativa 4C = 1Á8 pg and
Vicia faba 4C = 53Á0 pg) were used as calibration standards
at various times, but unfortunately the authors did not state
which standard(s) was compared with which Leucaena
species.
(au) Boscaiu et al. (1999, Ref. 427) referred to plants
of Cerastium with 2n = 36 as diploids in contrast with
various other authors who consider them as tetraploids.
The assumption was based on Boscaiu et al.'s observa-
tions that, while the base chromosome number in
Cerastium may be x = 9, no Cerastium species is known
with 2n = 18.
(av) The C-values reported for Hedera helix by
Obermayer and Greilhuber (1999, Ref. 429) agree well
with previous reports by Ko¨nig et al. (1987) of 2C = 3Á0 pg,
but are only about one third the value reported by Marie and
Brown (1993) of 2C = 8Á2 pg, which is unsupported.
(aw) Blanco et al. (1996, Ref. 431) gave DNA amounts
for Dasypyrum hordaceum and D. villosum in arbitrary units
(a.u.), listed as mean values in their fig. 3. The value for
D. hordaceum was converted into an absolute DNA amount
by multiplying the mean value of 381Á7 a.u. by a conversion
factor of 0Á11. This conversion factor was obtained as the
ratio of the 4C estimate for Dasypyrum villosum (listed as
the synonym Haynaldia villosa) reported by Bennett (1972)
as 21Á4 pg, and the estimate of 193Á7 a.u. reported by Blanco
et al. (1996).
(ax) Rayburn et al. (1997, Ref. 432) estimated C-values
in 90 accessions of Glycine max. Accessions showed a 12 %
variation in DNA amount and these differences were
statistically significant. Only the smallest and largest
C-values are listed in the Appendix.
(ay) Comparing C-values given by Belletti et al. (1998,
Ref. 434) with those previously published showed DNA
amounts for Capsicum baccatum, C. chinese, C. eximium,
C. frutescens and C. pubescens were around one third
greater than those of Owens (pers. comm.) listed in Bennett
and Smith (1976). Belletti et al. (1998) suggested that the
cause of the discrepancy could be that Owens used Allium
cepa as the calibration standard, whose 2C-value of 33Á5 pg
differs considerably from those reported in Capsicum
species studied.
(az) Širokyy et al. (2001, Ref. 437) investigated C-values
in four Silene species, including S. latifolia, which has
previously been listed by Bennett and Leitch (1995)
under its synonym Melandrium album.
(ba) Taliaferro et al. (1997, Ref. 438) gave DNA C-values
for 18 accessions of Cynodon corresponding to two species:
C. transvaalensis (2n = 2x = 18), and C. dactylon var.
dactylon (2n = 4x = 36 and 2n = 6x = 54). Only small
differences in DNA amounts were noted between five
diploid and five tetraploid accessions, and a mean
2C-value for each ploidy level was also given in table 2
of their paper. This mean value is listed in the Appendix.
However, the three hexaploid accessions examined
comprised one accession of C. dactylon var. dactylon and
two hybrids. Thus, only the C-value estimate for hexaploid
C. dactylon var. dactylon is entered in the Appendix, rather
than the mean for the three hexaploid accessions given in
table 2 of Taliaferro et al. (1997).
(bb) Blakesley et al. (2002, Ref. 441) examined seven
populations of Acacia dealbata and four of A. mangium
to determine ploidy and DNA amount. In A. dealbata,
they identified naturally occurring diploid, triploid and
tetraploid genotypes. Chromosome numbers were counted
in only one diploid and one tetraploid genotype, and
C-values for only these populations are given in the
Appendix. In naturally occurring A. mangium only
diploid populations were found, and the C-value for the
only population whose chromosome number was deter-
mined is given in the Appendix. C-values for colchicine-
induced tetraploid genotypes of A. mangium are not
included.
The 2C-value for diploid A. dealbata (1Á7 pg) is similar to
that reported by Bukhari (1997) as 2C = 1Á6 pg. In contrast
the 2C-value (2Á9 pg) reported by Mukherjee and Sharma
(1993b) is nearly twice that of Blakesley et al. (2002).
Perhaps this discrepancy reflects the use of Allium cepa
(2C = 33Á5 pg), whose genome size is over an order of
magnitude greater than that of Acacia, as a calibration
standard by Mukherjee and Sharma (1993b). Similar
discrepancies were noted between DNA estimates for
A. mangium reported by Blakesley et al. (loc. cit.) of
2C = 1Á3 pg, and those by Mukherjee and Sharma (1995)
of 2C = 2Á3 pg.
(bc) Ohri and Singh (2002, Ref. 443) listed C-values for
20 wild relatives of cultivated pigeon pea (Cajanus cajan).
However, C-values for 14 of these species had already been
communicated to MD Bennett in 1996 and listed in the
Appendix of Bennett and Leitch (1997) under Original
reference number 303. To avoid duplication of data in the
database, only C-values for six species not listed previously
are included in the Appendix.
(bd) Wendel et al. (2002, Ref. 444) listed DNA amounts
for 13 species in the tribe Gossypieae. However, C-values
for three of these species had already been communicated to
MD Bennett in 1999 and listed in the Appendix of Bennett
et al. (2000) under Original reference number 349. To
avoid duplication of data in the database, only C-values
for ten species not listed previously are included in
the Appendix.
(be) The C-value of Arabidopsis thaliana given by the
Arabidopsis Genome Initiative (2000, Ref. 448) was based
on DNA sequencing data for 115Á4 Mb of the genome, plus
a guestimate of 10 Mb for several unsequenced gaps in the
genome. Recent work places its 1C-value around 157 Mb
(Bennett et al., 2003).
(bf ) Ohri (2002, Ref. 449) listed DNA amounts for 36
tropical hardwood species belonging to 13 families.
However, C-values of 35 of these had already been
communicated to MD Bennett in 1996 and listed in the
Appendix of Bennett and Leitch (1997) under Original
reference number 301. To avoid duplication of data in the
database, a C-value for the only species not included in
a previous compilation (Drypetes roxburghii) is listed in
the present Appendix. A new C-value for Melaleuca
leucadendra, double that given in Bennett and Leitch
(1997), is also listed in the Appendix, to correct
64 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
an error in communication which confused the 2C- and 4C-
values for this species.
(bg) The C-value of 466 Mb for Oryza sativa ssp. indica
given in Yu et al. (2002, Ref. 450) was based on DNA
sequencing data for 362 Mb of sequenced scaffolds, and
104 Mb of `unassembled data' subject to numerous assump-
tions (see Yu et al., loc. cit. ­ page 80).
(bh) The C-value of 420 Mb for Oryza sativa ssp.
japonica given in Goff et al. (2002, Ref. 451) was derived
from DNA sequencing data for 389Á9 Mb, and their assump-
tion that this equals 93 % of the genome, perhaps using
some previously published 1C-value. The source of this
assumption, as of any such DNA estimate, and the method
by which it was obtained, was not clearly cited by Goff
et al. (2002).
(bi) Redondo et al. (1996, Ref. 453) estimated C-values in
four populations of Saxifraga granulata. In one population
chromosome numbers ranged from 2n = 44 to 56, but
2n = 44 was predominant. They noted that the DNA amount
was also variable but gave only one C-value, which is listed
in the Appendix. However, intraspecific variation in DNA
amount may occur in this species, so the C-value listed may
not apply to all members of the population.
(bj) Redondo et al. (1996, Ref. 453) estimated DNA
amounts in four populations of Saxifraga granulata. They
reported DNA amounts for a population in which they could
not obtain a chromosome count (entry number 706), but
based on the DNA amount, they suggested this population
may have 2n = 30.
(bk) Emshwiller (2002, Ref. 456) estimated C-values
in 10 accessions of cultivated oca (2n = 8x = 64; Oxalis
tuberosa), two tetraploid wild species, and 78 diploid
accessions which were provisionally identified as 35 species.
As variation in 2C DNA amounts was usually no more than
0Á1 pg, and considered to be technical in nature, only the
highest C-value was reported in table 3 of Emshwiller
(2002) for most species and is entered in the Appendix.
Variation in DNA amounts greater than 0Á1 pg was con-
sidered real for O. spiralis (2C = 1Á062­1Á339 pg) and O.
peduncularis (2C = 0Á927­1Á163 pg), so both the lowest and
highest values are entered in the Appendix for these species.
Emshwiller (2002) noted that this variation may reflect
problems of taxonomy and species boundaries.
(bl) Emshwiller (2002, Ref. 456) estimated the DNA
amounts in ten accessions of cultivated oca (2n = 8x = 64;
Oxalis tuberosa). Variation was noted, even in measure-
ments made for all accessions estimated on one day (see
table 3 of Emshwiller, 2002), but she did not consider it to
represent intraspecific variation and a mean 2C estimate
calculated from all measurements made was given in her
`Results' section as 2C = 3Á514 pg. It is this value that is
entered in the Appendix.
(bm) Nagl et al. (1983, Ref. 457) included DNA amounts
for 49 species. However, C-values for 20 of these had
been published elsewhere, and already included in previous
compilations by Bennett and colleagues (listed under
original reference numbers 34, 36, 60, 61, 81, 82, 84,
85, 86). To avoid duplication only C-values for 29 species
that had not been listed previously are included in the
Appendix.
(bn) Values for Phaseolus coccineus and P. vulgaris
given in Nagl et al. (1983, Ref. 457) are around twice
those given in another paper by Nagl and Treviranus
(1995, Ref. 390), listed in the present Appendix. 2C-values
for both species given in Nagl et al. (loc. cit.) agree with
those reported by Ayonoadu (1974), but are around twice
that reported in Ingle et al. (1975) and Arumuganathan and
Earle (1991). The basis of this discrepancy is unclear,
so C-values for these Phaseolus taxa should be viewed
with caution until confirmed independently.
(bo) The value for Sambucus nigra (2C = 30Á5 pg ) given
by Nagl et al. (1983, Ref. 457) is similar to the value of
2C = 21Á8 pg for a related species, S. racemosa, reported by
Nagl et al. (1979) and listed under Ref. 86 in Bennett et al.
(1982). However, it is very different from 2C = 3Á1 pg
reported for S. nigra by Mowforth (1986) and listed under
Ref. 158 in Bennett and Smith (1991). The cause for the
discrepancy remains unclear, so C-values for S. nigra
should be used with caution until confirmed independently.
(bp) Baranyi et al. (1996, Ref. 458) investigated C-values
in 75 accessions of four wild Pisum species. Results
were given as percentages relative to P. sativum `Kleine
Rheinla¨nderin' ( = 100 %) which was used as the calibration
standard. To convert these into absolute DNA amounts the
4C-values were multiplied by the value of Pisum sativum
`Kleine Rheinla¨nderin' of 17Á68 pg (Greilhuber and Ebert,
1994) and then divided by 100.
Pisum fulvum was homogeneous in DNA amount size
(4C = approx. 19Á3 pg), but wide variation was seen between
accessions of the other species studied (P. abyssinicum,
P. humile and P. elatius). This variation was interpreted
to show that these taxa with variable genome sizes were
genetically heterogeneous, suggesting that the current spe-
cies delimitations did not reflect the true biological species
groups adequately. Only the smallest and largest C-values
for each of these species are listed in the Appendix.
(bq) Punina and Alexandrova (1992, Ref. 459) estimated
DNA amounts in 11 Paeonia species but gave the results
as percentage values relative to P. caucasica. Since
Mulry and Hanson (pers. comm. 1999) had estimated the
4C DNA value of this species as 65Á2 pg (see entry number
602 in Bennett et al., 2000), the relative percentage values
given in Punina and Alexandrova (loc. cit.) were converted
into absolute DNA amounts by multiplying by 0Á652.
(br) A PhD thesis by Shi (1991) gave a 1C-value of
0Á15 pg for two accessions of diploid Brachypodium dis-
tachyon (2n = 10), plus values for four other Brachypodium
species. Later, Shi et al. (1993, Ref. 460) gave the 1C-value
for diploid B. distachyon as 1C = 0Á3 pg, but cited the PhD
thesis (Shi, 1991) as the source for this figure. In order to
confirm which was correct, Clive Stace kindly supplied seed
of one original accession (B306) and RBG, Kew estimated
its DNA amount as 0Á36 pg using Oryza sativa `IR36'
(4C = 2Á02 pg) as a calibration standard (see entry number
161a in the present Appendix). This was much closer to
the value in Shi et al. (1993). As C-values for four other
Brachypodium species in Shi (1991) may also be under-
estimates, they are therefore not included in the present
Appendix, and should be viewed with caution until
confirmed independently.
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 65
APPENDIX.Chromosomenumber,ploidylevel,life-cycletype,andnuclearDNAcontentin804angiospermspecies(thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
1aAcaciadealbataLink.NoLeguminosaeE262P8530.91.73.5441bb
OJFC:PI
2AcaciadealbataLink.NoLeguminosaeE39
3P--t
--t
2.55.1441bb
OJFC:PI
3AcaciadealbataLink.NoLeguminosaeE524P1,6711.73.46.8441bb
OJFC:PI
4aAcaciamangiumWilld.NoLeguminosaeE262P6370.71.32.6441bb
OJFC:PI
5AcridocarpusnatalitiusA.Juss.NoMalpighiaceaeEc.21624P1,4901.53.06.1379OJFe
6AdenantheramicrospermaTeijsm&Binn.NoLeguminosaeE--n
--p
P6810.71.42.8454OBc
Fe
7AdenantherapavoninaL.NoLeguminosaeE26
--p
P6660.71.42.7454OBc
Fe
8Adinacordifolia(Roxb.)Hook.f.NoRubiaceaeE22
2P8160.81.73.3454OBc
Fe
9AeoniumhaworthiiWebb&Berth.NoCrassulaceaeE72
4or8P7600.81.63.1378OJFe
10aAesculushippocastanumL.NoSapindaceaeE402P5880.61.22.4465OLycopers.c
FC:PI
11bAgaveamericanaL.--m
AsparagaceaeM1204P7,7918.015.931.8384aa
OHomof
FC:PI
12dAgavesisalanaPerr.--m
AsparagaceaeM1505P--t
--t
20.040.0384aa
OHomof
FC:PI
13AgavestrictaSalm.--m
AsparagaceaeM602P3,8223.97.815.6384aa
OHomof
FC:PI
14AgrostispalustrisHuds.NoGramineaeM284P2,7692.85.711.3417OGallusf
FC:PI
15AilanthusgrandisPrainNoSimaroubaceaeE64
--p
P2,1342.24.48.7454OBc
Fe
16aAlbucapendulaB.MathewNoAsparagaceaeM162P2,9673.06.112.1465OGFe
16bAlbucapendulaB.MathewNoAsparagaceaeM142P3,0333.16.212.4465OGFe
17kAlliumcepaL.NoAlliaceaek
M16
2P16,41516.833.567.0457bm
OBd
Fe
18Allocasuarinaverticillata(Lam.)
L.Johnson
NoCasuarinaceaeE20-28
2P9311.01.93.8452OPetuniae
FC:PI
19Alocasiacucullata(Lour)SchottNoAraceaeM98--p
AP8,2008.416.733.5411OBc
Fe
20AlocasiahilobeautyHost.NoAraceaeM32--p
A3,6803.87.515.0411OBc
Fe
21AloealbifloraGuillauminNoXanthorrhoeaceaek
M14
2P15,33715.731.362.6440OAgavesp.ab
FC:PI
22Aloealooides(Bolus)DrutenNoXanthorrhoeaceaek
M14
2P13,08313.426.753.4440OAgavesp.ab
FC:PI
23Aloeantandroi(Decary)H.PerrierNoXanthorrhoeaceaek
M14
2P17,19917.635.170.2440OAgavesp.ab
FC:PI
24Aloearborescens(yellowflowers)Mill.i
NoXanthorrhoeaceaek
M14
2P13,67114.027.955.8440OAgavesp.ab
FC:PI
25aAloearistataHaw.NoXanthorrhoeaceaek
M14
2P15,72916.132.164.2440OAgavesp.ab
FC:PI
25bAloearistatavar.parvifoliaBakerHaw.NoXanthorrhoeaceaek
M14
2P16,02316.432.765.4440OAgavesp.ab
FC:PI
26AloebakeriScott-ElliotNoXanthorrhoeaceaek
M14
2P15,92516.332.565.0440OAgavesp.ab
FC:PI
27AloebarberaeDyerNoXanthorrhoeaceaek
M14
2P15,04315.430.761.4440OAgavesp.ab
FC:PI
28AloebellatulaReynoldsNoXanthorrhoeaceaek
M14
2P16,26816.633.266.4440OAgavesp.ab
FC:PI
29AloeboiteauiGuillauminNoXanthorrhoeaceaek
M14
2P16,02316.432.765.4440OAgavesp.ab
FC:PI
30AloebowieaSchult.&Schult.f.NoXanthorrhoeaceaek
M14
2P16,26816.633.266.4440OAgavesp.ab
FC:PI
31AloebrevifoliaMill.NoXanthorrhoeaceaek
M14
2P14,55314.929.759.4440OAgavesp.ab
FC:PI
32bAloecameroniiHemsl.NoXanthorrhoeaceaek
M14
2P17,05217.434.869.6440OAgavesp.ab
FC:PI
33AloecapitataBakerNoXanthorrhoeaceaek
M14
2P15,38615.731.462.8440OAgavesp.ab
FC:PI
34AloechabaudiiSchonlandNoXanthorrhoeaceaek
M14
2P17,93418.336.673.2440OAgavesp.ab
FC:PI
35Aloeciliarisvar.tidmarshiiSchonlandHaw.NoXanthorrhoeaceaek
M14
2P10,53510.821.543.0440OAgavesp.ab
FC:PI
36AloeciliarisHaw.NoXanthorrhoeaceaek
M35
5P--t
--t
53.3106.6440OAgavesp.ab
FC:PI
37AloeciliarisHaw.NoXanthorrhoeaceaek
M42
6P30,72331.462.7125.4440OAgavesp.ab
FC:PI
38AloecomptoniiReynoldsNoXanthorrhoeaceaek
M14
2P13,42613.727.454.8440OAgavesp.ab
FC:PI
39aAloecryptopodaBakerNoXanthorrhoeaceaek
M14
2P14,16114.528.957.8440OAgavesp.ab
FC:PI
39bAloecryptopoda``Wickensii''BakerNoXanthorrhoeaceaek
M14
2P14,35714.729.358.6440OAgavesp.ab
FC:PI
40AloedaweiBergerNoXanthorrhoeaceaek
M28
4P35,23136.071.9143.8440OAgavesp.ab
FC:PI
41aAloedescoingsiiReynoldsNoXanthorrhoeaceaek
M14
2P15,97416.332.665.2440OAgavesp.ab
FC:PI
66 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
41bAloedescoingsiiReynoldsssp.augustina
Lavranos
NoXanthorrhoeaceaek
M14
2P16,21916.633.166.2440OAgavesp.ab
FC:PI
42aAloedichotomaMassonvar.ramosissima
(Pillans)Glen&D.S.Hardy
NoXanthorrhoeaceaek
M14
2P12,00512.324.549.0440OAgavesp.ab
FC:PI
42bAloedichotomaMassonNoXanthorrhoeaceaek
M14
2P12,10312.424.749.4440OAgavesp.ab
FC:PI
43AloedinteriA.BergerNoXanthorrhoeaceaek
M14
2P16,36616.733.466.8440OAgavesp.ab
FC:PI
44bAloedistansHaw.NoXanthorrhoeaceaek
M14
2P13,62213.927.855.6440OAgavesp.ab
FC:PI
45AloedorotheaeA.BergerNoXanthorrhoeaceaek
M14
2P15,28815.631.262.4440OAgavesp.ab
FC:PI
46AloeelegansTod.NoXanthorrhoeaceaek
M14
2P17,34617.735.470.8440OAgavesp.ab
FC:PI
47AloeerinaceaD.S.HardyNoXanthorrhoeaceaek
M14
2P12,10312.424.749.4440OAgavesp.ab
FC:PI
48AloeferoxMill.NoXanthorrhoeaceaek
M14
2P14,89615.230.460.8440OAgavesp.ab
FC:PI
49AloefleurentiniorumLavranos&
L.E.Newton
NoXanthorrhoeaceaek
M14
2P18,17918.637.174.2440OAgavesp.ab
FC:PI
50Aloegariepensis(?)PillansNoXanthorrhoeaceaek
M14
2P15,72916.132.164.2440OAgavesp.ab
FC:PI
51AloeglaucaMill.NoXanthorrhoeaceaek
M14
2P15,68016.032.064.0440OAgavesp.ab
FC:PI
52AloeglobuligemmaPole-EvansNoXanthorrhoeaceaek
M14
2P16,61117.033.967.8440OAgavesp.ab
FC:PI
53AloehaemanthifoliaA.Berger&MarlothNoXanthorrhoeaceaek
M14
2P7,9388.116.232.4440OAgavesp.ab
FC:PI
z
Chromosomenumber.
x
E,ephemeral;A,annual;B,biennial;P,perennial.
y
O,originalvalue;C,calibratedvalue
*
Thestandardspeciesusedtocalibratethepresentamount.
yy
Fe,Feulgenmicrodensitometry;FC,flowcytometryusingoneofthefollowingfluorochromes:PI,propidiumiodide;DAPI,40
,6-diamidinophenylindole;EB,ethidiumbromide;MI,mithramycin;HO,
Hoechst33258;GS,genomesequencing;CIA,computerimageanalysis;RK,reassociationkinetics.
#
E,eudicot;M,monocot;BA,basalangiosperm.
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 67
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
54AloehaworthioidesBakerNoXanthorrhoeaceaek
M14
2P14,74915.130.160.2440OAgavesp.ab
FC:PI
55AloehereroensisEngl.NoXanthorrhoeaceaek
M14
2P18,13018.537.074.0440OAgavesp.ab
FC:PI
56Aloehumilis(smallform)(L.)Mill.i
NoXanthorrhoeaceaek
M14
2P16,56216.933.867.6440OAgavesp.ab
FC:PI
57AloejacksoniiReynoldsNoXanthorrhoeaceaek
M28
4P32,48733.266.3132.6440OAgavesp.ab
FC:PI
58AloejucundaReynoldsNoXanthorrhoeaceaek
M14
2P17,59118.035.971.8440OAgavesp.ab
FC:PI
59bAloejuvennaBrandham&CarterNoXanthorrhoeaceaek
M28
4P34,79035.571.0142.0440OAgavesp.ab
FC:PI
60Aloekrapohlianavar.dumoulinii
Lavranos
NoXanthorrhoeaceaek
M14
2P17,34617.735.470.8440OAgavesp.ab
FC:PI
61AloelinearifoliaA.BergerNoXanthorrhoeaceaek
M14
2P12,93613.226.452.8440OAgavesp.ab
FC:PI
62AloelomatophylloidesBalf.f.NoXanthorrhoeaceaek
M14
2P17,24817.635.270.4440OAgavesp.ab
FC:PI
63AloelongistylaBakerNoXanthorrhoeaceaek
M14
2P15,58215.931.863.6440OAgavesp.ab
FC:PI
64AloemacrosiphonBak.NoXanthorrhoeaceaek
M14
2P17,93418.336.673.2440OAgavesp.ab
FC:PI
65AloemaculataAllioniiNoXanthorrhoeaceaek
M14
2P18,62019.038.076.0440OAgavesp.ab
FC:PI
66aAloemarlothiiA.Berger``Spectabilis''NoXanthorrhoeaceaek
M14
2P15,43515.831.563.0440OAgavesp.ab
FC:PI
66bAloemarlothiiA.Bergervar.bicolor
Reynolds
NoXanthorrhoeaceaek
M14
2P15,63116.031.963.8440OAgavesp.ab
FC:PI
67bAloemcloughliniiChristianNoXanthorrhoeaceaek
M14
2P16,21916.633.166.2440OAgavesp.ab
FC:PI
68AloemelanacanthaA.BergerNoXanthorrhoeaceaek
M14
2P12,29912.625.150.2440OAgavesp.ab
FC:PI
69AloemicrostigmaSalm-DyckNoXanthorrhoeaceaek
M14
2P15,09215.430.861.6440OAgavesp.ab
FC:PI
70AloemitriformisMill.NoXanthorrhoeaceaek
M14
2P13,47513.827.555.0440OAgavesp.ab
FC:PI
71bAloengobitensisReynoldsNoXanthorrhoeaceaek
M28
4P28,42029.058.0116.0440OAgavesp.ab
FC:PI
72Aloeoccidentalis(H.Perrier)L.E.Newton&
G.D.Rowley
NoXanthorrhoeaceaek
M14
2P20,28620.741.482.8440OAgavesp.ab
FC:PI
73AloeparvulaA.BergerNoXanthorrhoeaceaek
M14
2P16,56216.933.867.6440OAgavesp.ab
FC:PI
74AloepearsoniiSchonlandNoXanthorrhoeaceaek
M14
2P12,34812.625.250.4440OAgavesp.ab
FC:PI
75bAloepeckiiBally&VerdoornNoXanthorrhoeaceaek
M14
2P17,44417.835.671.2440OAgavesp.ab
FC:PI
76AloepegleraeSchonlandNoXanthorrhoeaceaek
M14
2P15,72916.132.164.2440OAgavesp.ab
FC:PI
77AloepetricolaPole-EvansNoXanthorrhoeaceaek
M14
2P15,09215.430.861.6440OAgavesp.ab
FC:PI
78AloepillansiiL.GuthrieNoXanthorrhoeaceaek
M14
2P12,59312.925.751.4440OAgavesp.ab
FC:PI
79Aloeplicatilis(L.)Mill.NoXanthorrhoeaceaek
M14
2P8,6248.817.635.2440OAgavesp.ab
FC:PI
80AloepluridensHaworthNoXanthorrhoeaceaek
M14
2P14,16114.528.957.8440OAgavesp.ab
FC:PI
81AloepolyphyllaSchonlandNoXanthorrhoeaceaek
M14
2P13,37713.727.354.6440OAgavesp.ab
FC:PI
82AloeprinslooiVerdoorn&HardyNoXanthorrhoeaceaek
M14
2P17,44417.835.671.2440OAgavesp.ab
FC:PI
83Aloeprostrata(H.Perrier)L.E.Newton&
G.D.Rowley
NoXanthorrhoeaceaek
M14
2P20,13920.641.182.2440OAgavesp.ab
FC:PI
84AloerauhiiReynoldsNoXanthorrhoeaceaek
M14
2P15,33715.731.362.6440OAgavesp.ab
FC:PI
85AloerichardsiaeReynoldsNoXanthorrhoeaceaek
M14
2P21,75622.244.488.8440OAgavesp.ab
FC:PI
86AloeriviereiLavranos&
L.E.Newton
NoXanthorrhoeaceaek
M14
2P16,56216.933.867.6440OAgavesp.ab
FC:PI
87AloesecundifloraEngl.NoXanthorrhoeaceaek
M14
2P17,59118.035.971.8440OAgavesp.ab
FC:PI
88AloesinkatanaReynolds(redflowers)i
NoXanthorrhoeaceaek
M14
2P17,54217.935.871.6440OAgavesp.ab
FC:PI
89AloesladenianaPole-EvansNoXanthorrhoeaceaek
M14
2P15,97416.332.665.2440OAgavesp.ab
FC:PI
90AloespeciosaBakerNoXanthorrhoeaceaek
M14
2P14,11214.428.857.6440OAgavesp.ab
FC:PI
91AloespicataL.f.NoXanthorrhoeaceaek
M14
2P14,25914.629.158.2440OAgavesp.ab
FC:PI
92AloestriataHaw.NoXanthorrhoeaceaek
M14
2P18,91419.338.677.2440OAgavesp.ab
FC:PI
93AloesuprafoliataPole-EvansNoXanthorrhoeaceaek
M14
2P14,01414.328.657.2440OAgavesp.ab
FC:PI
68 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
94AloesuzannaeDecaryNoXanthorrhoeaceaek
M14
2P16,31716.733.366.6440OAgavesp.ab
FC:PI
95bAloetenuiorHaw.NoXanthorrhoeaceaek
M14
2P10,63310.921.743.4440OAgavesp.ab
FC:PI
96AloetrichosanthaBergerNoXanthorrhoeaceaek
M14
2P18,17918.637.174.2440OAgavesp.ab
FC:PI
97AloevanbaleniiPillansNoXanthorrhoeaceaek
M14
2P13,96514.328.557.0440OAgavesp.ab
FC:PI
98aAloevariegataL.``Ausana''NoXanthorrhoeaceaek
M14
2P16,26816.633.266.4440OAgavesp.ab
FC:PI
98bAloevariegataL.NoXanthorrhoeaceaek
M14
2P16,56216.933.867.6440OAgavesp.ab
FC:PI
99Aloevera(L.)Burm.f.NoXanthorrhoeaceaek
M14
2P16,07216.432.865.6440OAgavesp.ab
FC:PI
100AlstoniamacrophyllaWall.ex
G.Don.
NoApocynaceaeE--n
--p
P7180.71.52.9454OBc
Fe
101cAlstroemeriaaureaGrahamh
NoAlstroemeriaceaeM162P24,84325.450.7101.4436OBFC:PI
101dAlstroemeriaaureaGrahamh
NoAlstroemeriaceaeM162P27,09727.755.3110.6436OBFC:PI
102AlstroemeriaaureaGrahamh
NoAlstroemeriaceaeM243P--t
--t
80.9161.8436OBFC:PI
103dAlstroemerialigtuL.ssp.
incarnataL.h
NoAlstroemeriaceaeM162P34,30035.070.0140.0436OBFC:PI
103eAlstroemerialigtuL.ssp.simsiih
NoAlstroemeriaceaeM162P31,94832.665.2130.4436OBFC:PI
103fAlstroemerialigtuL.ssp.simsiih
NoAlstroemeriaceaeM162P38,66139.578.9157.8436OBFC:PI
103gAlstroemerialigtuL.ssp.ligtuh
NoAlstroemeriaceaeM162P34,00634.769.4138.8436OBFC:PI
103hAlstroemerialigtuL.ssp.ligtuh
NoAlstroemeriaceaeM162P33,36934.168.1136.2436OBFC:PI
104cAlstroemeriamagnificaHerb.
ssp.magnificah
NoAlstroemeriaceaeM162P17,88518.336.573.0436OBFC:PI
104dAlstroemeriamagnificaHerb.
ssp.magnificah
NoAlstroemeriaceaeM162P20,53121.041.983.8436OBFC:PI
105AlstroemeriamagnificaHerb.
ssp.magnificah
NoAlstroemeriaceaeM243P--t
--t
61.6123.2436OBFC:PI
106AmborellatrichopodaBaill.NoAmborellaceaeBA26
--p
P8700.91.83.6381OKFC:PI
107AmoreuxiawrightiiA.GrayNoCochlospermaceaeEc.12-142P1640.20.30.7378OJFe
108Anthemisaltissimal
NoCompositaej
E--n
--p
--q
7,7427.915.831.6457bm
OBd
Fe
109Anthemismontanal
NoCompositaej
E--n
--p
--q
8,2818.516.933.8457bm
OBd
Fe
110AnthuriumgrandeHost.NoAraceaeM28--p
P13,25213.527.054.1411OBc
Fe
111AnthuriumtetragonumSchottNoAraceaeM30--p
P7,4857.615.330.6411OBc
Fe
112AphyllanthesmonspeliensisL.NoAsparagaceaeMc.32--p
P6350.61.32.6380OJFC:PI
113Arabidopsiskorshynskyil
NoCruciferaej
E--n
--p
--q
2450.30.51.0457bm
OBd
Fe
114aArabidopsisthaliana(L.)Heynh.
ecotypeColumbia
NoCruciferaeE102A1570.20.30.6461OCaeno.f
FC:PI
114gArabidopsisthaliana(L.)Heynh.
ecotypeColumbia
NoCruciferaeE102A1640.20.30.7461OGallusf
FC:PI
114hArabidopsisthaliana(L.)Heynh.
ecotypeColumbia
NoCruciferaeE102A1500.20.30.6461ODros.f
FC:PI
114iArabidopsisthaliana(L.)Heynh.--m
CruciferaeE102A125be
0.1be
0.3be
0.5be
448be
O--GS
114jArabidopsisthaliana(L.)Heynh.
ecotypeColumbia
NoCruciferaeE102A1670.20.30.7463OGallusf
FC:DAPI
114kArabidopsisthaliana(L.)Heynh.
ecotypeColumbia
NoCruciferaeE102A1620.20.30.7463OGallusf
FC:HO
114lArabidopsisthaliana(L.)Heynh.
ecotypeColumbia
NoCruciferaeE102A1570.20.30.6463OGallusf
FC:MI
114mArabidopsisthaliana(L.)Heynh.
lineLandsbergerecta
NoCruciferaeE102A510.050.100.21464O--RK
115aArachisduranensisKrapov.&
W.C.Gregoryh
NoLeguminosaej
E202A1,2431.32.55.1396OGc
FC:PI
115bArachisduranensisKrapov.&
W.C.Gregoryh
NoLeguminosaej
E202A1,3241.42.75.4396OGc
FC:PI
115cArachisduranensisKrapov.&
W.C.Gregoryh
NoLeguminosaej
E202A1,3331.42.75.4396OGc
Fe
116aArachishypogaeaL.NoLeguminosaeE404A2,8132.95.711.5395aj
OGc
Fe
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 69
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
116bArachishypogaeaL.NoLeguminosaeE404A2,8983.05.911.8395aj
OGc
FC:PI
116rArachishypogaeaL.NoLeguminosaej
E40
4A1,5681.63.26.4457bm
OBd
Fe
117aArachismonticolaKrapov.&
Rigoni
NoLeguminosaeE404A2,8913.05.911.8395aj
OGc
Fe
117bArachismonticolaKrapov.&
Rigoni
NoLeguminosaeE404A2,9303.06.012.0395aj
OGc
FC:PI
118Archidendronmonadelphum(Roxb.)
I.C.Neilsen
NoLeguminosaeE--n
--p
P1,4701.53.06.0454OBc
Fe
119cArtemisiaabsinthiumL.NoCompositaej
E182P4,1754.38.517.0386OG-120dFC:PI
120dArtemisiaannuaL.NoCompositaej
E182A1,7151.83.57.0386OGc
FC:PI
121ArtemisiabarrelieriBesserNoCompositaej
E364P6,3506.513.025.9386OGc
FC:PI
122ArtemisiacaerulescensL.ssp.gallica
(Willd.)K.Persson
NoCompositaej
E182P3,2633.36.713.3386OGc
FC:PI
123ArtemisiacampestrisL.NoCompositaej
E182P2,8762.95.911.7386OGc
FC:PI
124ArtemisiacampestrisL.NoCompositaej
E364P5,3905.511.022.0386OGc
FC:PI
125ArtemisiacanaPursh.NoCompositaej
E728P12,56912.825.751.3386OGc
FC:PI
126ArtemisiachamaemelifoliaVill.NoCompositaej
E182P2,9603.06.012.1386OGc
FC:PI
127ArtemisiacrithmifoliaL.NoCompositaej
E546P7,6447.815.631.2386OGc
FC:PI
128ArtemisiadracunculusL.NoCompositaej
E9010P11,37811.623.246.4386OGc
FC:PI
129ArtemisiafragransWilld.NoCompositaej
E182P2,6222.75.410.7386OGc
FC:PI
130Artemisiaherba-albaAssossp.
valentina(Lam.)Mascl.
NoCompositaej
E182P3,2193.36.613.1386OGc
FC:PI
131Artemisiaherba-albaAssossp.
herba-alba
NoCompositaej
E364P6,1156.212.525.0386OGc
FC:PI
132cArtemisiajudaicaL.NoCompositaej
E162P5,6455.811.523.0386OGc
FC:PI
133ArtemisialucenticaO.Bolos,
Valles&Vigo
inO.Bolos&Vigo
NoCompositaej
E162P3,7633.87.715.4386OG-120dFC:PI
134ArtemisiamolinieriQuezel,Barbero&
R.Loisel
NoCompositaej
E182P2,9203.06.011.9386OGc
FC:PI
135ArtemisiamonospermaDelileNoCompositaej
E364P5,4005.511.022.0386OGc
FC:PI
136ArtemisiasplendensWilld.NoCompositaej
E324P6,6596.813.627.2386OGc
FC:PI
137ArtemisiathusculaCav.NoCompositaej
E182P5,1555.310.521.0386OGc
FC:PI
138ArtemisiatournefortianaReichenb.NoCompositaej
E182AB3,2783.36.713.4386OGc
FC:PI
139ArtemisiatridentataNutt.ssp.spiciformis
Kartesz&Gandhi
NoCompositaej
E182P4,0084.18.216.4386OG-120dFC:PI
140ArtemisiaumbelliformisLam.
ssp.umbelliformis
NoCompositaej
E344P6,0816.212.424.8386OGc
FC:PI
141bArtemisiavulgarisL.NoCompositaej
E162P2,9793.06.112.2386OGc
FC:PI
142ArtemisiavulgarisL.NoCompositaej
E344P4,7734.99.719.5386OGc
FC:PI
143bArummaculatumL.NoAraceaeM56
8P10,68210.921.843.6457bm
OBd
Fe
144AsarumeuropaeumL.NoAristolochiaceaeBA--n
--p
P4,7534.99.719.4457bm
OBd
Fe
145AsteliafragransColensoNoAsteliaceaeMc.608P1,2401.32.55.1380OKFC:PI
146Atalantiaceylanica(Arn.)Oliv.)NoRutaceaeE18
2P5150.51.12.1426OGallusf
FC:PI
70 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
147AustrobaileyascandensC.T.WhiteNoAustrobaileyaceaeBA44
--p
P9,3279.519.038.1381OGFC:PI
148AverrhoacarambolaL.NoOxalidaceaeE--n
--p
P2350.20.51.0454OBc
Fe
149AzadirachtaindicaA.JussNoMeliaceaeE28
--p
P3850.40.81.6454OBc
Fe
150Bauhiniahookeri(F.Muell.)PedleyNoLeguminosaeE26
--p
P6200.61.32.5454OBc
Fe
151bBauhiniapurpureaL.NoLeguminosaeE28
2P5730.61.22.3454OBc
Fe
152BauhiniatomentosaL.NoLeguminosaeE28
2P6130.61.32.5454OBc
Fe
153BellevaliarixiiP.WendalboNoAsparagaceaeM82P9,1029.318.637.2465OBFe
154BerberidopsiscorallinaHook.f.NoBerberidopsidaceaeEc.426P2520.30.51.0379OJFe
155Berryacordifolia(Willd.)BurretNoMalvaceaeE--n
--p
P5490.61.12.2454OBc
Fe
156aBixaorellanaL.NoBixaceaeE142P1910.20.40.8379OJFe
156bBixaorellanaL.NoBixaceaeE14
2P2030.20.40.8454OBc
Fe
157BlandfordiapuniceaSweet.NoBlandfordiaceaeM684P7,9708.116.332.5380OGFe
158BombaxceibaL.NoMalvaceaeE92
--p
P1,5901.63.26.5454OBc
Fe
159BoswelliaserrataRoxb.NoBurseraceaeE22
--p
P6840.71.42.8454OBc
Fe
160BrachychitondiscolorF.MuellNoMalvaceaeE40
--p
P1,1321.22.34.6454OBc
Fe
161aBrachypodiumdistachyon(L.)P.Beauv.NoGramineaeM102A3550.40.71.5465OKFC:PI
161bBrachypodiumdistachyon(L.)P.Beauv.--m
GramineaeM102A2940.30.61.2460bh
OJFe
162eBrassicanapusL.NoCruciferaej
E38
4AB1,5681.63.26.4457bm
OBd
Fe
163bBromusarvensisL.NoGramineaeM142A5,6995.811.623.3391af
OH-164bFC:PI
164bBromuscarinatusHooker&Arnottcv.BromaNoGramineaeM568P11,24111.522.945.9391af
OHc
FC:PI
165cBromuserectusHudsonNoGramineaeM568P12,07912.324.749.3391af
OHc
FC:PI
166cBromushordeaceusL.NoGramineaeM284A11,28511.523.046.1391af
OHc
FC:PI
167bBromusinermisLeysserNoGramineaeM568P12,02512.324.549.1391af
OHc
FC:PI
168BromuswilldenowiiKnuthNoGramineaeM426P6,3656.513.026.0391af
OHc
FC:PI
169Buchloedactyloides(Nutt.)Engelm.NoGramineaeM404P7790.81.63.2417OGallusf
FC:PI
170BuddlejaglobosaHopeNoBuddlejaceaeE382P8400.91.73.4378OJFe
171BulbinealooidesWilld.NoXanthorrhoeaceaeM142P10,60110.821.643.3465OBFe
172BulbinefallaxPoelln.NoXanthorrhoeaceaeM142P11,20111.422.945.7465OBFe
173Bulbinelagopus(Thunb.)N.E.BrownNoXanthorrhoeaceaeM--n
--p
P7,9388.116.232.4465OBFe
174BulbinepraemorsaSpreng.NoXanthorrhoeaceaeM142P12,21312.524.949.9465OBFe
175aBuniaserucagoL.NoCruciferaeE142A2,0292.14.18.3393OGc
Fe
175bBuniaserucagoL.NoCruciferaeE142A2,1362.24.48.7393OGlycinee
FC:PI
176aBuniasorientalisL.NoCruciferaeE142P2,5382.65.210.4393OGc
Fe
176bBuniasorientalisL.NoCruciferaeE142P2,6362.75.410.8393OGlycinee
FC:PI
177BuxuspapillosaC.K.Schneid.NoBuxaceaeE--n
--p
P1,3891.42.85.7454OBc
Fe
178Buxussempervirensl
NoBuxaceaeE282or4P7940.81.63.2380OKFC:PI
179ByblislinifloraSalisb.NoByblidaceaeE322A8700.91.83.6378OJFe
180Cajanusalbicans(Wight.&Am.)MaesenNoLeguminosaeE222A1,2591.32.65.1443bc
OBc
Fe
181Cajanusmollis(Benth.)MaesenNoLeguminosaeE222B8040.81.63.3443bc
OBc
Fe
182Cajanussericeus(Benth.exBak.)MaesenNoLeguminosaeE222P1,4141.42.95.8443bc
OBc
Fe
183CaladiumbicolorVent.var.redpolkaNoAraceaeM32--p
A5,4075.511.022.1411OBc
Fe
184CaladiumbicolorVent.var.redpolkalargeNoAraceaeM66--p
A9,92710.120.340.5411OBc
Fe
185CalceolariaacutifoliaWitasekNoScrophulariaceaeE--n
--p
--q
1,3481.42.85.5465OJFe
186Calceolariagracilisl
NoScrophulariaceaeE--n
--p
--q
1,3351.42.75.5465OJFe
187Calibrachoacalycina(Sendtn.)WijsmanNoSolanaceaeE182P1,5041.53.16.1387ae
OGallus-398pFC:PI
188Calibrachoadusenii(R.E.Fr.)Stehmann&
Semir.
NoSolanaceaeE182P1,4011.42.95.7387ae
OGallus-398pFC:PI
189CalibrachoaeglandulataStehmann&Semir.NoSolanaceaeE18
2P1,4111.42.95.8387ae
OGallus-398pFC:PI
190Calibrachoaelegans(Miers)Stehmann&
Semir.
NoSolanaceaeE182P1,5631.63.26.4387ae
OGallus-398pFC:PI
191Calibrachoaericaefolia(R.E.Fr.)WijsmanNoSolanaceaeE182P1,4361.52.95.9387ae
OGallus-398pFC:PI
192Calibrachoaheterophylla(Sendtn.)WijsmanNoSolanaceaeE18
2P1,4551.53.05.9387ae
OGallus-398pFC:PI
193Calibrachoalinearis(Hook.)WijsmanNoSolanaceaeE182P1,4851.53.06.1387ae
OGallus-398pFC:PI
194Calibrachoalinoides(Sendtn.)WijsmanNoSolanaceaeE182P1,3971.42.95.7387ae
OGallus-398pFC:PI
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 71
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
195Calibrachoamacrodactylon(L.B.Sm.&Downs)
Wijsman
NoSolanaceaeE182P1,4801.53.06.0387ae
OGallus-398pFC:PI
196Calibrachoamicrantha(R.E.Fr.)
Stehmann&Semir.
NoSolanaceaeE182P1,4111.42.95.8387ae
OGallus-398pFC:PI
197Calibrachoaparviflora(Juss.)WijsmanNoSolanaceaeE182A9361.01.93.8387ae
OGallus-398pFC:PI
198Calibrachoapygmaea(R.E.Fr.)WijsmanNoSolanaceaeE182A7640.81.63.1387ae
OGallus-398pFC:PI
199Calibrachoarupestris(Dusen)WijsmanNoSolanaceaeE18
2P1,5971.63.36.5387ae
OGallus-398pFC:PI
200Calibrachoaselloviana(Sendtn.)WijsmanNoSolanaceaeE182P1,4551.53.05.9387ae
OGallus-398pFC:PI
201Calibrachoasendtneriana(R.E.Fr.)
Stehmann&Semir.
NoSolanaceaeE18
2P1,4501.53.05.9387ae
OGallus-398pFC:PI
202Calibrachoaserrulata(L.B.Sm.&Downs)
Stehmann&Semir.
NoSolanaceaeE182P1,4461.53.05.9387ae
OGallus-398pFC:PI
203Calibrachoaspathulata(L.B.Sm.&Downs)
Stehmann&Semir.
NoSolanaceaeE182P1,4161.42.95.8387ae
OGallus-398pFC:PI
204Calibrachoathymifolia(A.St.-Hil.)
Stehmann&Semir.
NoSolanaceaeE182P1,4851.53.06.1387ae
OGallus-398pFC:PI
205Callistemoncitrinus(Curtis)SkeelsNoMyrtaceaeE22
--p
P1,0141.02.14.1454OBc
Fe
206CallistemonrigidusR.Br.NoMyrtaceaeE--n
--p
P1,5261.63.16.2454OBc
Fe
207CamelliasinensisKuntzeNoTheaceaeE302P3,8243.97.815.6379OGFe
208Cannaindical
NoCannaceaeM182P7060.71.42.9379OJFe
209aCannabissativaL.(female)NoCannabaceaeE202A818.am
0.8am
1.7am
3.3am
414OArab.e
FC:DAPI
209bCannabissativaL.(male)NoCannabaceaeE202A843.am
0.9am
1.7am
3.4am
414OArab.e
FC:DAPI
210CanotiaholacanthaTorr.NoCelastraceaeE302P1810.20.40.7378OJFe
211bCapsellabursa-pastoris(L.)Medic.NoCruciferaej
E32
4A6860.71.42.8457bm
OBd
Fe
212hCapsicumannuumL.cv.DouxLongdes
Landes
NoSolanaceaeE242--q
3,7343.87.615.2434OGc
FC:PI
213cCapsicumbaccatumL.ssp.pendulumNoSolanaceaeE242--q
4,1114.28.416.8434ay
OG-212hFC:PI
213dCapsicumbaccatumL.ssp.baccatumNoSolanaceaeE242--q
4,1314.28.416.9434ay
OG-212hFC:PI
214CapsicumcardenasiiHeiser&SmithNoSolanaceaeE24
2--q
4,3954.59.017.9434OG-212hFC:PI
215CapsicumchacoenseA.T.Hunz.NoSolanaceaeE24
2--q
3,7533.87.715.3434OG-212hFC:PI
216bCapsicumchinenseJacq.NoSolanaceaeE242--q
3,9404.08.016.1434ay
OG-212hFC:PI
217bCapsicumeximiumA.T.Hunz.NoSolanaceaeE24
2--q
4,2634.48.717.4434ay
OG-212hFC:PI
218bCapsicumfrutescensL.NoSolanaceaeE24
2--q
3,8914.07.915.9434ay
OG-212hFC:PI
219CapsicumpraetermissumHeiser&SmithNoSolanaceaeE24
2--q
4,4744.69.118.3434OG-212hFC:PI
220bCapsicumpubescensR.&P.NoSolanaceaeE24
2--q
4,7634.99.719.4434ay
OG-212hFC:PI
221CapsicumtovariiEshbaugh,Smith&NickrentNoSolanaceaeE24
2--q
3,8864.07.915.9434OG-212hFC:PI
222aCardamineamaraL.NoCruciferaeE162P2380.20.51.0465OLycopers.c
FC:PI
223CastanospermumaustraleA.Cunn.&C.FraserNoLeguminosaeE--n
--p
P5540.61.12.3454OBc
Fe
224CasuarinaglaucaSieb.exSpring.NoCasuarinaceaeE18
2P3430.40.71.4452OPetuniae
FC:PI
225Catunaregamspinosa(Thunb.)TrivengadumNoRubiaceaeE22
--p
P3430.40.71.4454OBc
Fe
226bCentaureascabiosal
NoCompositaeE--n
--p
P1,2541.32.65.1465OJFe
227CephalotusfollicularisLabill.NoCephalotaceaeE20
2P6250.61.32.6378OJFe
228aCerastiumalpinumL.NoCaryophyllaceaeE724P1,8131.93.77.4427OCerastiume
FC:DAPI
228bCerastiumalpinumL.NoCaryophyllaceaeE724P1,9702.04.08.0427OBc
Fe
229CerastiumarcticumLanges.str.NoCaryophyllaceaeE1086P3,1263.26.412.8427OCerastiume
FC:DAPI
72 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
230CerastiumarvenseL.ssp.glandulosum
(Kit.)Soo
NoCaryophyllaceaeE362au
P6660.71.42.7427OCerastiume
FC:DAPI
231CerastiumarvenseL.ssp.arvenseNoCaryophyllaceaeE724P1,2741.32.65.2427OCerastiume
FC:DAPI
232aCerastiumbanaticum(Rochel)Heuff.NoCaryophyllaceaeE362au
P1,4701.53.06.0427OBc
Fe
232bCerastiumbanaticum(Rochel)Heuff.NoCaryophyllaceaeE362au
P1,5291.63.16.2427OCerastiume
FC:DAPI
233CerastiumcarinthiacumVestNoCaryophyllaceaeE362au
P1,4801.53.06.0427OCerastiume
FC:DAPI
234aCerastiumeriophorumKit.inSchult.NoCaryophyllaceaeE362au
P1,2641.32.65.2427OBc
Fe
234bCerastiumeriophorumKit.inSchult.NoCaryophyllaceaeE362au
P1,2741.32.65.2427ONicot.e
FC:DAPI
235bCerastiumfontanumBaumg.NoCaryophyllaceaeE1448P3,4693.57.114.2427OCerastiume
FC:DAPI
236aCerastiumlatifoliumL.NoCaryophyllaceaeE362au
P1,4211.52.95.8427OBc
Fe
236bCerastiumlatifoliumL.NoCaryophyllaceaeE362au
P1,4701.53.06.0427OCerastiume
FC:DAPI
237aCerastiumtranssylvanicumSchurexGriseb.&
Schenk
NoCaryophyllaceaeE1086P3,0383.16.212.4427OCerastiume
FC:DAPI
237bCerastiumtranssylvanicumSchurexGriseb.&
Schenk
NoCaryophyllaceaeE1086P3,0483.16.212.4427OBc
Fe
238CeratophyllumdemersumL.NoCeratophyllaceaeBAc.706P6740.71.42.8381OJFe
239ChenopodiumalbumL.h
NoAmaranthaceaeE18
2A7500.81.53.1455OBc
Fe
240ChenopodiumalbumL.h
NoAmaranthaceaeE36
4A1,5971.63.36.5455OBc
Fe
241bChenopodiumalbumL.h
NoAmaranthaceaeE54
6A2,4232.54.99.9455OBc
Fe
242Chenopodiumberlandieri(Saff.)Wilson&
Heiserssp.nuttalliaeh
NoAmaranthaceaeE36
4A1,4461.53.05.9455OBc
Fe
243ChenopodiumbushianumAellenNoAmaranthaceaeE36
4A1,5581.63.26.4455OBc
Fe
244ChenopodiumficifoliumSm.NoAmaranthaceaeE18
2A6490.71.32.7455OBc
Fe
245ChenopodiumgiganteumD.DonNoAmaranthaceaeE54
6A2,1512.24.48.8455OBc
Fe
246ChenopodiummuraleL.NoAmaranthaceaeE18
2A6100.61.22.5455OBc
Fe
247ChenopodiumopulifoliumSchrad.exKoch&
Ziz
NoAmaranthaceaeE36
4A1,3031.32.75.3455OBc
Fe
248bChenopodiumpallidicauleAellenNoAmaranthaceaeE18
2A6170.61.32.5455OBc
Fe
249bChenopodiumquinoaWilld.h
NoAmaranthaceaeE36
4A1,5851.63.26.5455OBc
Fe
250Chenopodiumugandae(Aell.)Aell.NoAmaranthaceaeE32
--p
A1,4011.42.95.7455OBc
Fe
251ChenopodiumvulvariaL.NoAmaranthaceaeE18
2A6220.61.32.5455OBc
Fe
252ChloranthusspicatusMak.NoChloranthaceaeBA30--p
P3,5263.67.214.4381OGFC:PI
253ChorisiaspeciosaSt.HillNoMalvaceaeE86
--p
P8450.91.73.5454OBc
Fe
254Ciccaacida(L.)Merr.NoEuphorbiaceaeE--n
--p
P9381.01.93.8454OBc
Fe
255CicersongaricumSteph.exDC.NoLeguminosaeE162P1,3281.42.75.4435OBc
Fe
256CienfuegosiatripartitaH.B.K.Gurke--m
MalvaceaeE202--q
9311.01.93.8444bd
OGb2
FC:PI
257CienfuegosiayucatanensisMillspaugh--m
MalvaceaeE222--q
9801.02.04.0444bd
OGb2
FC:PI
258CistusalbanicusE.F.WarburgexHeywoodNoCistaceaeE18
2P2,6172.75.310.7404ORaphanuse
FC:PI
259CistusalbidusL.NoCistaceaeE18
2P2,3422.44.89.6404ORaphanuse
FC:PI
260CistusclusiiDunalNoCistaceaeE18
2P2,5872.65.310.6404ORaphanuse
FC:PI
261CistuscreticusL.NoCistaceaeE18
2P2,1272.24.38.7404ORaphanuse
FC:PI
262CistuscrispusL.NoCistaceaeE18
2P1,9212.03.97.8404ORaphanuse
FC:PI
263CistusheterophyllusDesf.ssp.carthaginensis
(Pau)Crespo&Mateo
NoCistaceaeE18
2P2,3622.44.89.6404ORaphanuse
FC:PI
264CistusladaniferL.NoCistaceaeE18
2P2,1812.24.58.9404ORaphanuse
FC:PI
265CistuslaurifoliusL.NoCistaceaeE18
2P2,1852.24.58.9404ORaphanuse
FC:PI
266CistuslibanotisL.NoCistaceaeE18
2P2,8272.95.811.5404ORaphanuse
FC:PI
267CistusmonspeliensisL.NoCistaceaeE18
2P2,8812.95.911.8404ORaphanuse
FC:PI
268CistusosbeckiaefoliusWebbexPitard&ProustNoCistaceaeE18
2P2,0242.14.18.3404ORaphanuse
FC:PI
269CistusparviflorusLam.NoCistaceaeE18
2P2,4302.55.09.9404ORaphanuse
FC:PI
270CistuspopulifoliusL.NoCistaceaeE18
2P2,1022.14.38.6404ORaphanuse
FC:PI
271CistuspsilosepalusSweetNoCistaceaeE18
2P2,5582.65.210.4404ORaphanuse
FC:PI
272CistussalviifoliusL.NoCistaceaeE18
2P2,3322.44.89.5404ORaphanuse
FC:PI
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 73
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
273CistussymphytifoliusLam.NoCistaceaeE18
2P2,4062.54.99.8404ORaphanuse
FC:PI
274bCitrusaurantiumL.NoRutaceaeE18
2P4310.40.91.8426OGallusf
FC:PI
275bCitrusgrandis(L.)OsbeckNoRutaceaeE18
2P3770.40.81.5426OGallusf
FC:PI
276bCitruslimon(L.)Burm.f.i
NoRutaceaeE18
2P3920.40.81.6426OGallusf
FC:PI
277CitruslimoniaOsbeckcv.BromeRangpurNoRutaceaeE18
2P4020.40.81.6426OGallusf
FC:PI
278bCitrusparadisiMacfad.NoRutaceaeE18
2P3920.40.81.6426OGallusf
FC:PI
279CitrusreshniHort.exTanakaNoRutaceaeE18
2P4020.40.81.6426OGallusf
FC:PI
280eCitrussinensis(L.)Osbeckcv.Sargoins
GrosseRondeh
NoRutaceaeE18
2P3720.40.81.5426OGallusf
FC:PI
280fCitrussinensis(L.)Osbeckcv.Pineappleh
NoRutaceaeE18
2P4170.40.91.7426OGallusf
FC:PI
280gCitrussinensis(L.)OsbeckNoRutaceaeE18
2P5880.61.22.4457bm
OBd
Fe
281CitrusvolkamerianaTen.&Pasq.NoRutaceaeE18
2P3870.40.81.6426OGallusf
FC:PI
282CoccolobadiversifoliaJacq.NoPolygonaceaeE--n
--p
P1,1271.22.34.6454OBc
Fe
283hCoffeaarabicaL.--m
RubiaceaeE444P1,2791.32.65.2424OKc
FC:PI
283iCoffeaarabicaL.NoRubiaceaeE44
4P1,1221.12.34.6454OBc
Fe
284cCoffeabrevipesHiern.--m
RubiaceaeE222P7600.81.63.1424OKc
FC:PI
285dCoffeacanephoraPierre.exFroehn.--m
RubiaceaeE222P7550.81.53.1424OKc
FC:PI
286cCoffeacongensisFroehn.--m
RubiaceaeE222P7940.81.63.2424OKc
FC:PI
287dCoffeaeugenioidesS.Moore--m
RubiaceaeE222P6810.71.42.8424OKc
FC:PI
288cCoffeahumilisA.Cheval.--m
RubiaceaeE222P8720.91.83.6424OKc
FC:PI
289dCoffealibericaL.ssp.dewevreiWild&
Dur.Hiern
NoRubiaceaeE222P7030.71.42.9401OPetuniae
FC:PI
289eCoffealibericaL.--m
RubiaceaeE222P8230.81.73.4424OKc
FC:PI
290dCoffeapseudozanguebariaeD.M.Bridson--m
RubiaceaeE222P5340.51.12.2424OKc
FC:PI
291cCoffearacemosal
--m
RubiaceaeE222P4660.51.01.9424OKc
FC:PI
292cCoffeasessilifloraD.M.Bridson--m
RubiaceaeE222P5100.51.02.1424OKc
FC:PI
293Coffeasp.F.Bridsony
--m
RubiaceaeE222P6520.71.32.7424OKc
FC:PI
294Coffeasp.Moloundouy
--m
RubiaceaeE222P8280.81.73.4424OKc
FC:PI
295cCoffeastenophyllaG.Don.--m
RubiaceaeE222P6620.71.42.7424OKc
FC:PI
296bColocasiaantiquorumSchottvar.1i
NoAraceaeM32--p
AP4,9515.110.120.2411OBc
Fe
297CommiphoramossambicensisEngl.NoBurseraceaeE262P6130.61.32.5379OJFe
298CoriariamyrtifoliaL.NoCoriariaceaeEc.728P3260.30.71.3378OJFe
299Cosmosatrosanguineusl
NoCompositaeE48--p
P7,1917.314.729.4465OFFe
300cCrepisbiennisL.h
NoCompositaeEc.4010B7,4487.615.230.4394ah
OGc
CIA
300dCrepisbiennisL.h
NoCompositaeEc.4010B7,9288.116.232.4394ah
OGc
CIA
300eCrepisbiennisL.h
NoCompositaeEc.4010B8,1738.316.733.4394ah
OGc
Fe
300fCrepisbiennisL.h
NoCompositaeEc.4010B8,5558.717.534.9394ah
OGc
Fe
301aCrepisbithynicavar.pirinicaAcht.i
NoCompositaeE102P3,1563.26.412.9394OGc
Fe
301bCrepisbithynicavar.bithynicaBoiss.i
NoCompositaeE102P3,2443.36.613.2394OGc
Fe
302hCrepiscapillaris(L.)Wallr.NoCompositaej
E6
2A2,5972.75.310.6457bm
OBd
Fe
303aCrepisconyzaefolia(Gouan)A.Kerneri
NoCompositaeE82P5,4005.511.022.0394OGc
Fe
303bCrepisconyzaefolia(Gouan)A.Kerneri
NoCompositaeE82P5,5765.711.422.8394OEc
FC:PI
304aCrepispaludosa(L.)Moenchi
NoCompositaeE122P4,0774.28.316.6394OGc
Fe
304bCrepispaludosa(L.)Moenchi
NoCompositaeE122P4,3614.58.917.8394OEc
FC:PI
305bCrepispulchraL.h
NoCompositaeE82A4,4594.69.118.2394OGc
CIA
305cCrepispulchraL.h
NoCompositaeE82A5,4495.611.122.2394ai
OGc
CIA
74 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
305dCrepispulchraL.h
NoCompositaeE82A4,8515.09.919.8394OGc
Fe
305eCrepispulchraL.h
NoCompositaeE82A5,4005.511.022.0394ai
OGc
Fe
305fCrepispulchraL.h
NoCompositaeE82A5,3805.511.022.0394OEc
FC:PI
305gCrepispulchraL.h
NoCompositaeE82A5,9886.112.224.4394OEc
FC:PI
306aCrepissancta(L.)Babc.h
NoCompositaeE102A2,0482.14.28.4394ah
OGc
Fe
306bCrepissancta(L.)Babc.h
NoCompositaeE102A2,1762.24.48.9394ah
OGc
CIA
307CrepisschachtiiBabc.i
NoCompositaeE102P2,7642.85.611.3394OGc
Fe
308bCrepissetosaHallerf.i
NoCompositaeE82A1,6951.73.56.9394OGc
CIA
309aCrepisviscidulaFroel.i
NoCompositaeE122P4,2834.48.717.5394OGc
CIA
309bCrepisviscidulaFroel.i
NoCompositaeE122P4,7824.99.819.5394OEc
FC:PI
310bCrepiszacintha(L.)Babc.i
NoCompositaeE62A1,0581.12.24.3394OGc
CIA
311CrocusbiflorusMill.NoIridaceaeM8ao
2P4,2534.38.717.4419OHomof
FC:PI
312CrocuscartwrightianusHerb.i
NoIridaceaeM162P3,8864.07.915.9419OHomof
FC:PI
313CrocusetruscusParl.NoIridaceaeM82P3,5533.67.314.5419OHomof
FC:PI
314CrocussativusL.i
NoIridaceaeM243P--t
--t
11.823.6419OHomof
FC:PI
315CrocusthomasiiTen.NoIridaceaeM162P4,2584.38.717.4419OHomof
FC:PI
316Cyclamenhederifoliuml
NoPrimulaceaeE--n
--p
P2,9233.06.011.9465OGFC:PI
317aCyclamentrochopteranthumO.Schwarzh
NoPrimulaceaeE30--p
P10,37310.621.242.3465OGFe
317bCyclamentrochopteranthumO.Schwarzh
NoPrimulaceaeE30--p
P13,45813.727.554.9465OGFe
318bCynodondactylon(L.)Pers.NoGramineaeM364P9561.02.03.9417OGallusf
FC:PI
318cCynodondactylon(L.)Pers.var.dactylonNoGramineaeM364P1,1031.12.34.5438ba
OSusf
FC:PI
319Cynodondactylon(L.)Pers.var.dactylonNoGramineaeM546P1,4361.52.95.9438ba
OIctal.f
FC:PI
320aCynodontransvaalensisBurtt-DavyNoGramineaeM182P5050.51.02.1417OGallusf
FC:PI
320bCynodontransvaalensisBurtt-DavyNoGramineaeM182P5440.61.12.2438ba
OIctal.f
FC:PI
321DalbergiahorridaDennst.NoLeguminosaeE202P1,9282.03.97.9445OGFe
322aDalbergialanceolariaLin.f.NoLeguminosaeE202P1,4311.52.95.8445OGFe
323aDalbergialatifoliaRoxb.NoLeguminosaeE202P1,6811.73.46.9445OGFe
324DalbergiamalabaricaPrain.NoLeguminosaeE202P1,8471.93.87.5445OGFe
325DalbergiamelanoxylonGuill&Perr.NoLeguminosaeE202P1,8061.83.77.4445OGFe
326DalbergiapaniculataRoxb.NoLeguminosaeE202P1,5511.63.26.3445OGFe
327DalbergiarubiginosaRoxb.NoLeguminosaeE202P1,8031.83.77.4445OGFe
328DalbergiasissoidesGrah.NoLeguminosaeE202P1,7641.83.67.2445OGFe
329aDalbergiasissooRoxb.exDC.NoLeguminosaeE202P1,5851.63.26.5445OGFe
329bDalbergiasissooRoxb.exDC.NoLeguminosaeE20
2P6910.71.42.8454OBc
Fe
330DalbergiavolubilisRoxb.NoLeguminosaeE202P1,9011.93.97.8445OGFe
331DamasoniumalismaMill.NoAlismataceaeM--n
--p
A23,14323.647.294.5465OBFC:PI
332DasypogonhookeriDrumm.NoDasypogonaceaeM142P4260.40.91.7380OJFC:PI
333Dasypyrumhordeaceum(Cosson&Durieu)
Candargy
NoGramineaeM284P10,28810.521.042.0431C--aw
Fe
334dDasypyrumvillosum(=Haynaldiavillosa)
(L.)P.Candargyh
NoGramineaeM142A10,40810.621.242.5430OCc
Fe
334eDasypyrumvillosum(=Haynaldiavillosa)
(L.)P.Candargyh
NoGramineaeM142A6,2626.412.825.6430OCc
Fe
335bDecaisneafargesiiFranch.NoLardizabalaceaek
E40
2P2,4502.55.010.0457bm
OBd
Fe
336DeutziaprunifoliaRehderNoHydrangeaceaeE524P1,8351.93.77.5378OJFe
337DictamnusalbusL.NoRutaceaeE36
--p
P3,3813.56.913.8457bm
OBd
Fe
338DieffenbachiapictaSchottNoAraceaeM36--p
P12,08312.324.749.3411OBc
Fe
339DiospyrosdiscolorWilld.NoEbenaceaeE30
--p
P1,1741.22.44.8454OBc
Fe
340DiospyrosmalabaricaKost.NoEbenaceaeE30
--p
P1,4361.52.95.9454OBc
Fe
341DissotiscanescensHook.f.NoMelastomataceaeEc.28-32--p
P1810.20.40.7378OJFe
342aDoritispulcherrimaLindl.NoOrchidaceaeM382P6,6106.713.527.0447OGb2
FC:PI
343DoryanthespalmeriW.HillexBenth.NoDoryanthaceaeM48--p
P3,2393.36.613.2380OGFC:PI
344DrimysvickerianaA.C.SmithNoWinteraceaeBA--n
--p
P1,1051.12.34.5381OKFC:PI
345DrypetesroxburghiiWall.NoPutranjavaceaek
E42
--p
P1,0021.02.04.1449bf
OBc
Fe
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 75
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
346Ehretialaevis(RottlerexG.Don)Roxb.NoBoraginaceaeE--n
--p
P3,5333.67.214.4454OBc
Fe
347Eremochloaophiuroides(Munro)Hack.NoGramineaeM182P8130.81.73.3417OGallusf
FC:PI
348Eriocaulonaquaticuml
NoEriocaulaceaeM324P4,1014.28.416.7380OGFC:PI
349Escalloniarubral
NoEscalloniaceaeE242P4140.40.81.7380OJFe
350EucnidegrandifloraRoseNoLoasaceaeEc.38-402or6P5880.61.22.4378OJFe
351EucommiaulmoidesOliverNoEucommiaceaeE342P7250.71.53.0379OGFe
352aFagussylvaticaL.var.tortuosaPepinWillk.NoFagaceaeE24
2P5440.61.12.2433OPetuniae
FC:PI
352bFagussylvaticaL.NoFagaceaeE24
2P5440.61.12.2433OPetuniae
FC:PI
352cFagussylvaticaL.var.purpureaAit.NoFagaceaeE24
2P5490.61.12.2433OPetuniae
FC:PI
352dFagussylvaticaL.var.pendulaLodd.NoFagaceaeE24
2P5540.61.12.3433OPetuniae
FC:PI
353dFestucaarundinaceaSchreb.NoGramineaeM426P7,6397.815.631.2417ONicot.e
FC:PI
354FestucalongifoliaThuill.NoGramineaeM426P6,2236.412.725.4417OFc
FC:PI
355Firmianacolorata(Roxb.)R.Br.NoMalvaceaeE40
--p
P1,6151.63.36.6454OBc
Fe
356FlagellariaguineensisSchum.NoFlagellariaceaeM38
2P8800.91.83.6380OKFC:PI
357FlemingiabracteataWightNoLeguminosaeE222P1,5701.63.26.4443bc
OBc
Fe
358FortunellahindsiiSwing.NoRutaceaeE36
4P6220.61.32.5426OGallusf
FC:PI
359FouquieriasplendensEngelm.NoFouquieriaceaeE244P5190.51.12.1378OJFe
360Fragariaxananassacv.RedcoatDuch.--m
RosaceaeE568P5980.61.22.4442OGallusf
FC:M
361aGagealutea(L.)KerGawl.NoLiliaceaeM726P19,35519.839.579.0413OBc
Fe
361bGagealutea(L.)KerGawl.NoLiliaceaeM726P19,82520.240.580.9413OBc
FC:EB
362GardeniaresinifluaHiernNoRubiaceaeE--n
--p
P1,2691.32.65.2454OBc
Fe
363GarryafremontiiTorr.NoGarryaceaeEc.202P1,4901.53.06.1380OLycopers.c
FC:PI
364jGlycinemax(L.)Merr.strainT215h
NoLeguminosaeE40
2A1,1611.22.44.7423as
OFc
FC:PI
364kGlycinemax(L.)Merr.strainPI423.894h
NoLeguminosaeE40
2A1,2151.22.55.0423as
OFc
FC:PI
364lGlycinemax(L.)Merr.h
--m
LeguminosaeE40
2A1,2501.32.65.1432ax
CGlycinee
FC:PI
364mGlycinemax(L.)Merr.h
--m
LeguminosaeE40
2A1,4011.42.95.7432ax
CGlycinee
FC:PI
365GoodeniamimuloidesS.MooreNoGoodeniaceaeE162A5070.51.02.1379OGFe
366GossypioidesherbaceumL.--m
MalvaceaeE26
2--q
1,8131.93.77.4444bd
OGb2
FC:PI
367GossypioidesraimondiiUlbrich--m
MalvaceaeE26
2--q
9801.02.04.0444bd
OGb2
FC:PI
368GunneramanicataLindenNoGunneraceaeE342P7,2867.414.929.7379OFFe
369Gymnostomadeplancheana(Miq.)L.JohnsonNoMoraceaek
E16
2P3680.40.81.5452OPetuniae
FC:PI
370Haldinacordifolia(Roxb.)RidsdaleNoRubiaceaeE44
4P1,2961.32.65.3454OBc
Fe
371Hampeaappendiculata(J.Donnell-Smith)
Standley
--m
MalvaceaeE26
2--q
2,8913.05.911.8444bd
OGb2
FC:PI
372HanguanamalayanaMerrillNoHanguanaceaeMc.170--p
P1,6121.63.36.6380OFFe
373aHederacanariensisWilld.NoAraliaceaeE48
2P1,3721.42.85.6429OGlycinee
FC:PI
373bHederacanariensisWilld.NoAraliaceaeE48
2P1,5091.53.16.2429OGc
CIA
374aHederacolchicaC.Koch.NoAraliaceaeE1928P5,3415.510.921.8429OGlycinee
FC:PI
374bHederacolchicaC.Koch.NoAraliaceaeE1928P5,5865.711.422.8429OGc
CIA
375eHederahelixL.NoAraliaceaeE482P1,4601.53.06.0429av
OGc
CIA
375fHederahelixL.NoAraliaceaeE482P1,3721.42.85.6429av
OGlycinee
FC:PI
375gHederahelixL.f.arborescensC.K.SchneiderNoAraliaceaeE482P1,5091.53.16.2429OGc
CIA
375hHederahelixL.f.arborescensC.K.SchneiderNoAraliaceaeE482P1,3821.42.85.6429OGlycinee
FC:PI
376rHelianthusannuusL.NoCompositaeE342A3,5773.77.314.6403OGFC:PI
377HeliconiarostrataRuiz&Pav.NoHeliconiaceaeM242P4410.50.91.8379OJFe
76 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
378HelleborusargutifoliusViv.NoRanunculaceaeE322P9,2619.518.937.8383z
OAgavee
FC:PI
379aHelleborusatrorubensWaldst.&Kit.
`Cupreus'i
NoRanunculaceaeE322P14,50414.829.659.2383z
OAgavee
FC:PI
379bHelleborusatrorubensWaldst.&Kit.i
NoRanunculaceaeE322P15,09215.430.861.6383z
OAgavee
FC:PI
380Helleboruscyclophyllus(A.Br.)Boiss.NoRanunculaceaeE322P14,65115.029.959.8383z
OAgavee
FC:PI
381HelleborusdumetorumWaldst.&Kit.NoRanunculaceaeE322P15,87616.232.464.8383z
OAgavee
FC:PI
382aHelleborusfoetidusL.`WesterFlisk'NoRanunculaceaeE322P11,41711.723.346.6383z
OAgavee
FC:PI
382bHelleborusfoetidusL.i
NoRanunculaceaeE322P11,46611.723.446.8383z
OAgavee
FC:PI
383HelleboruslividusAitonNoRanunculaceaeE322P9,3109.519.038.0383z
OAgavee
FC:PI
384aHelleborusmultifidusVis.ssp.hercegovinus
(Martinis)B.Mathew
NoRanunculaceaeE322P14,50414.829.659.2383z
OAgavee
FC:PI
384bHelleborusmultifidusVis.ssp.istriacus
(Schiffner)Merxm&Podl.
NoRanunculaceaeE322P14,74915.130.160.2383z
OAgavee
FC:PI
384cHelleborusmultifidusVis.ssp.multifidusNoRanunculaceaeE322P14,79815.130.260.4383z
OAgavee
FC:PI
384dHelleborusmultifidusVis.ssp.bocconeisiculusNoRanunculaceaeE322P15,04315.430.761.4383z
OAgavee
FC:PI
384eHelleborusmultifidusVis.ssp.bocconei
(Tenore)B.Mathew
NoRanunculaceaeE322P15,09215.430.861.6383z
OAgavee
FC:PI
385aHelleborusnigerL.(doubleflower)i
NoRanunculaceaeE322P13,72014.028.056.0383z
OAgavee
FC:PI
385bHelleborusnigerL.i
NoRanunculaceaeE322P13,86714.228.356.6383z
OAgavee
FC:PI
385cHelleborusnigerL.ssp.macranthus
(Freyn)Schiffneri
NoRanunculaceaeE322P14,40614.729.458.8383z
OAgavee
FC:PI
386HelleborusodorusWaldst.&Kit.i
NoRanunculaceaeE322P15,04315.430.761.4383z
OAgavee
FC:PI
387aHelleborusorientalisLamarckssp.orientalisi
NoRanunculaceaeE322P14,55314.929.759.4383z
OAgavee
FC:PI
387bHelleborusorientalisLamarcki
NoRanunculaceaeE322P14,72515.030.160.1383z
OAgavee
FC:PI
387cHelleborusorientalisLamarckssp.guttatus
(A.Br.&Sauer)B.Mathewi
NoRanunculaceaeE322P14,74915.130.160.2383z
OAgavee
FC:PI
387dHelleborusorientalisLamarckssp.
abchasicus(A.Br.)B.Mathewi
NoRanunculaceaeE322P14,79815.130.260.4383z
OAgavee
FC:PI
387eHelleborusorientalisLamarck`Kochii'i
NoRanunculaceaeE322P14,99415.330.661.2383z
OAgavee
FC:PI
388HelleboruspurpurascensWaldst.&Kit.NoRanunculaceaeE322P14,94515.330.561.0383z
OAgavee
FC:PI
389HelleborusthibetanusFranchetNoRanunculaceaeE322P17,49317.935.771.4383z
OAgavee
FC:PI
390aHelleborustorquatusArcherHind`Dido'
(doubleflowers)i
NoRanunculaceaeE322P14,60214.929.859.6383z
OAgavee
FC:PI
390bHelleborustorquatusArcherHindi
NoRanunculaceaeE322P14,74915.130.160.2383z
OAgavee
FC:PI
390cHelleborustorquatusArcherHind`Croaticus'i
NoRanunculaceaeE322P14,70015.030.060.0383z
OAgavee
FC:PI
391HelleborusvesicariusAucherNoRanunculaceaeE322P13,86714.228.356.6383z
OAgavee
FC:PI
392aHelleborusviridisL.ssp.viridisNoRanunculaceaeE322P14,89615.230.460.8383z
OAgavee
FC:PI
392bHelleborusviridisL.ssp.occidentalis(Reut.)
Schiffner
NoRanunculaceaeE322P15,09215.430.861.6383z
OAgavee
FC:PI
393Hernandianymphaeifolia(C.Presl.)KubitzkiNoHernandiaceaeBA--n
--p
P2,3402.44.89.6454OBc
Fe
394HerniariaglabraLinn.NoCaryophyllaceaeE182AP5150.51.12.1465OJFe
395bHieraciumaurantiacumL.NoCompositaej
E36
4P3,6263.77.414.8457bm
OBd
Fe
396HolopteleaintegrifoliaPlanch.NoUlmaceaeE28
--p
P6660.71.42.7454OBc
Fe
397HomalomenarubescensKunthNoAraceaeM34--p
AP8,9559.118.336.6411OBc
Fe
398pHordeumvulgareL.cv.NewGoldenNoGramineaeM14
2A5,0965.210.420.8387ae
OGallusf
FC:PI
399Hostacapitata(Koidzumi)NakaiNoAsparagaceaek
M602P9,4579.719.338.6384aa
OAgavesp.ab
FC:PI
400Hostaclausavar.normalisF.MaekawaNoAsparagaceaek
M602P9,4089.619.238.4384aa
OAgavesp.ab
FC:PI
401HostaclausaNakaivar.clausaNoAsparagaceaek
M903P--t
--t
28.557.0384aa
OAgavesp.ab
FC:PI
402HostagracillimaF.MaekawaNoAsparagaceaek
M602P10,82911.122.144.2384aa
OAgavesp.ab
FC:PI
403HostahypoleucaMurataNoAsparagaceaek
M602P12,49512.825.551.0384aa
OAgavesp.ab
FC:PI
404HostajonesiiM.ChungNoAsparagaceaek
M602P8,5758.817.535.0384aa
OAgavesp.ab
FC:PI
405HostakikutiiF.MaekawaNoAsparagaceaek
M602P11,17211.422.845.6384aa
OAgavesp.ab
FC:PI
406HostakiyosumiensisF.MaekawaNoAsparagaceaek
M602P11,90712.224.348.6384aa
OAgavesp.ab
FC:PI
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 77
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
407Hostalongipesvar.longipesMatsumuraNoAsparagaceaek
M602P12,74013.026.052.0384aa
OAgavesp.ab
FC:PI
408HostalongissimaHondaNoAsparagaceaek
M602P9,6049.819.639.2384aa
OAgavesp.ab
FC:PI
409HostaminorNakai`Gosan'NoAsparagaceaek
M602P8,4288.617.234.4384aa
OAgavesp.ab
FC:PI
410Hostaplantaginea(Lamarck)AschersonNoAsparagaceaek
M602P12,10312.424.749.4384aa
OAgavesp.ab
FC:PI
411HostapulchellaN.FujitaNoAsparagaceaek
M602P10,63310.921.743.4384aa
OAgavesp.ab
FC:PI
412HostapycnophyllaF.MaekawaNoAsparagaceaek
M602P10,87811.122.244.4384aa
OAgavesp.ab
FC:PI
413bHostarectifoliaNakaiNoAsparagaceaek
M602P10,43710.721.342.6384aa
OAgavesp.ab
FC:PI
414HostarupifragaNakaiNoAsparagaceaek
M602P12,98513.326.553.0384aa
OAgavesp.ab
FC:PI
415HostashikokianaN.FujitaNoAsparagaceaek
M602P11,22111.522.945.8384aa
OAgavesp.ab
FC:PI
416Hostasieboldianavar.sieboldiana
(Hooker)Engler
NoAsparagaceaek
M602P11,56411.823.647.2384aa
OAgavesp.ab
FC:PI
417HostasieboldiiP.O.(Paxton)IngramNoAsparagaceaek
M602P11,02511.322.545.0384aa
OAgavesp.ab
FC:PI
418HostatibaeF.MaekawaNoAsparagaceaek
M602P8,6248.817.635.2384aa
OAgavesp.ab
FC:PI
419HostatsushimensisN.FujitaNoAsparagaceaek
M602P8,4778.717.334.6384aa
OAgavesp.ab
FC:PI
420HostaventricosaStearnNoAsparagaceaek
M1204P19,20819.639.278.4384aa
OAgavesp.ab
FC:PI
421HostavenustaF.MaekawaNoAsparagaceaek
M602P8,4778.717.334.6384aa
OAgavesp.ab
FC:PI
422HostayingeriS.B.JonesNoAsparagaceaek
M602P9,3599.619.138.2384aa
OAgavesp.ab
FC:PI
423aHydrangeaanomalaD.Donssp.petiolaris
Sieb.&Zucc.
NoHydrangeaceaeE362P1,3281.42.75.4397OGc
FC:EB
423bHydrangeaanomalaD.Donssp.anomala
McClint.
NoHydrangeaceaeE362P1,5341.63.16.3397OGc
FC:EB
424HydrangeaarborescensL.NoHydrangeaceaeE362P1,1321.22.34.6397OGc
FC:EB
425aHydrangeaasperaDon.ssp.robustaMcClint.
(=H.longipesFranch.)
NoHydrangeaceaeE342P1,4801.53.06.0397OGc
FC:EB
425bHydrangeaasperaDon.ssp.sargentiana
(Rehder)McClint.
NoHydrangeaceaeE342P1,5291.63.16.2397OGc
FC:EB
425cHydrangeaasperaDon.ssp.strigosaMcClint.NoHydrangeaceaeE342P1,7001.73.56.9397OGc
FC:EB
425dHydrangeaasperaDon.ssp.asperaMcClint.NoHydrangeaceaeE362P2,3232.44.79.5397OGc
FC:EB
426HydrangeaheteromallaD.DonNoHydrangeaceaeE362P1,4461.53.05.9397OGc
FC:EB
427HydrangeainvolucrataSieb.NoHydrangeaceaeE302P2,4502.55.010.0397OGc
FC:EB
428aHydrangeamacrophylla(Thunb.)Ser.ssp.
serrata(Thunb.)Makino
NoHydrangeaceaeE362P1,8871.93.97.7397OGc
FC:EB
428bHydrangeamacrophylla(Thunb.)Ser.ssp.
macrophyllaMcClint.
NoHydrangeaceaeE362P2,1072.24.38.6397OGc
FC:EB
429HydrangeapaniculataSieb.NoHydrangeaceaeE362P1,8471.93.87.5397OGc
FC:EB
430HydrangeaquercifoliaBartr.NoHydrangeaceaeE362P9561.02.03.9397OGc
FC:EB
431aHydrangeascandens(L.f.)ssp.scandens
McClint.
NoHydrangeaceaeE362P1,8031.83.77.4397OGc
FC:EB
431bHydrangeascandens(L.f.)ssp.luikinensis
(Nakai)McClint.
NoHydrangeaceaeE362P1,8721.93.87.6397OGc
FC:EB
432HydrangeaseemanniiRileyNoHydrangeaceaeE362P1,0241.02.14.2397OGc
FC:EB
433Hypericumhirsutuml
NoHypericaceaeE--n
--p
P1470.20.30.6465OJFe
434Ingadulcis(Roxb.)Willd.NoLeguminosaeE--n
--p
P4020.40.81.6454OBc
Fe
435aIrisstenophyllaHausskn.exBakerh
NoIridaceaeM242P8,3598.517.134.1465OBFe
435bIrisstenophyllaHausskn.exBakerh
NoIridaceaeM262P10,74311.021.943.9465OBFe
435cIrisstenophyllaHausskn.exBakerh
NoIridaceaeM262P11,43911.723.346.7465OBFe
78 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
436IxiolirionledebouriiFisch.&Mey.NoIxioliriaceaeMc.242P9951.02.04.1380OJFe
437IxoraarboreaRoxb.exSm.NoRubiaceaeE--n
--p
P1,3651.42.85.6454OBc
Fe
438JacquiniaaristataJacq.NoTheophrastaceaeE382P5930.61.22.4378OJFe
439KentranthusruberDruceNoValerianaceaeE324P4070.40.81.7379OGFe
440Khayasenegalensis(Desr.)A.Juss.NoMeliaceaeE--n
--p
P8530.91.73.5454OBc
Fe
441Kigeliaafricana(Lam.)Benth.NoBignoniaceaeE--n
--p
P1,7001.73.56.9454OBc
Fe
442KirkiaacuminataOliverNoKirkiaceaeEc.302P3190.30.71.3378OJFe
443LagerstroemiatomentosaC.Presl.Presl.NoLythraceaeE--n
--p
P9651.02.03.9454OBc
Fe
444LantanacamaraL.NoVerbenaceaeE22
--p
P2,6972.85.511.0454OBc
Fe
445LapageriaroseaRuiz&Pav.NoPhilesiaceaeM30+1B2P6,6446.813.627.1380OGFe
446bLathyrusamphicarposL.--m
LeguminosaeE142A5,1235.210.520.9412O--v
Fe
447eLathyrusannuusL.--m
LeguminosaeE142A6,4296.613.126.2412O--v
Fe
447fLathyrusannuusL.--m
LeguminosaeE142A6,2826.412.825.6418OGc
FC:PI
448jLathyrusaphacaL.--m
LeguminosaeE142A4,5284.69.218.5418OC&Ec
FC:PI
449bLathyruschloranthusBoiss.--m
LeguminosaeE142A5,9446.112.124.3412O--v
Fe
450fLathyrusciceraL.--m
LeguminosaeE142A5,1945.310.621.2418OGc
FC:PI
451eLathyrusclymenumL.--m
LeguminosaeE142A4,2974.48.817.5418OC&Ec
FC:PI
451fLathyrusclymenumL.--m
LeguminosaeE142A6,3996.513.126.1412O--v
Fe
452LathyrusgmeliniiFritsch--m
LeguminosaeE142P8,7919.017.935.9412O--v
Fe
453bLathyrusgrandiflorusSibth.&Sm.--m
LeguminosaeE142P8,7248.917.835.6412O--v
Fe
454bLathyrusheterophyllusL.--m
LeguminosaeE142P8,6398.817.635.3412O--v
Fe
455Lathyruslaevigatus(Waldst.&Kit.)--m
LeguminosaeE142P11,00811.222.544.9412O--v
Fe
456cLathyrusmaritimusBigelow--m
LeguminosaeE142P6,7776.913.827.7412O--v
Fe
457eLathyrusnissoliaL.--m
LeguminosaeE142A4,8535.09.919.8412O--v
Fe
458eLathyrusochrus(L.)DC--m
LeguminosaeE142A4,5424.69.318.5418OC&Ec
FC:PI
459iLathyrusodoratusL.NoLeguminosaej
E14
2A8,1348.316.633.2457bm
OBd
Fe
460fLathyrussativusL.--m
LeguminosaeE142A6,5516.713.426.7418OGc
FC:PI
460gLathyrussativusL.--m
LeguminosaeE142A6,8997.014.128.2412O--v
Fe
461fLathyrussylvestrisL.--m
LeguminosaeE142P10,10410.320.641.2412O--v
Fe
462gLathyrustingitanusL.--m
LeguminosaeE142A7,6937.915.731.4418OGc
FC:PI
462hLathyrustingitanusL.--m
LeguminosaeE142A7,6917.815.731.4412O--v
Fe
465LawsoniainermisL.NoLythraceaeEc.30-344P3330.30.71.4378OJFe
464LebronneciakokioidesFosberg--m
MalvaceaeE262--q
1,7641.83.67.2444bd
OGb2
FC:PI
465LemnaminorL.NoAraceaeM1266P1,4261.52.95.8400OGFe
466LeucaenacollinsiiBritton&RoseNoLeguminosaeE52,?56
2P5290.51.12.2425O--at
FC:EB
467bLeucaenaconfertifloraS.ZarateNoLeguminosaeE112
4P8280.81.73.4425O--at
FC:EB
468LeucaenacuspidataStandleyNoLeguminosaeE--n
--p
P6860.71.42.8425O--at
FC:EB
469Leucaenadiversifolia(Schltdl.Benth.NoLeguminosaeE1044P1,3281.42.75.4425O--at
FC:EB
470cLeucaenaesculenta(Sesse&Moc.exDC)
Benth.
NoLeguminosaeE52(?56,
?112)
--p
P7060.71.42.9425O--at
FC:EB
471LeucaenagreggiiS.WatsonNoLeguminosaeE56
2P8870.91.83.6425O--at
FC:EB
472LeucaenainvolucrataS.ZarateNoLeguminosaeE--n
--p
P1,1221.12.34.6425O--at
FC:EB
473cLeucaenalanceolataS.WatsonNoLeguminosaeE522P7060.71.42.9425O--at
FC:EB
474LeucaenalempiranaC.E.HughesNoLeguminosaeE--n
--p
P4260.40.91.7425O--at
FC:EB
475bLeucaenaleucocephala(Lam.)DeWit.NoLeguminosaeE1044P1,4551.53.05.9425O--at
FC:EB
476LeucaenamacrophyllaBenth.NoLeguminosaeE?52
--p
P2990.30.61.2425O--at
FC:EB
477Leucaenamagnifica(C.E.Hughes)C.E.HughesNoLeguminosaeE?52
--p
P5000.51.02.0425O--at
FC:EB
478Leucaenamatudae(S.Zarate)C.E.HughesNoLeguminosaeE?52
--p
P5190.51.12.1425O--at
FC:EB
479LeucaenamulticapitulaScheryNoLeguminosaeE?52
--p
P4700.51.01.9425O--at
FC:EB
480LeucaenapallidaBritton&RoseNoLeguminosaeE104
(?110/112)
4P7740.81.63.2425O--at
FC:EB
481LeucaenapueblanaBritton&RoseNoLeguminosaeE--n
--p
P4900.51.02.0425O--at
FC:EB
482Leucaenapulverulenta(Schltdl.)Benth.NoLeguminosaeE562P6860.71.42.8425O--at
FC:EB
483LeucaenaretusaBenth.NoLeguminosaeE56
2P7640.81.63.1425O--at
FC:EB
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 79
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
484LeucaenasalvadorensisStandley
exBritton&Rose
NoLeguminosaeE?56
--p
P8870.91.83.6425O--at
FC:EB
485LeucaenashannoniiJ.D.SmithNoLeguminosaeE52
2P6910.71.42.8425O--at
FC:EB
486Leucaenatrichandra(Zucc.)UrbanNoLeguminosaeE52(?56)
2P7640.81.63.1425O--at
FC:EB
487Leucaenatrichodes(Jacq.)Benth.NoLeguminosaeE52
2P5390.61.12.2425O--at
FC:EB
488LimnanthesdouglasiiR.Br.NoLimnanthaceaeE102A1,3621.42.85.6379OJFe
489Litseaglutinosa(Lour.)C.B.RobinsonNoLauraceaeBA48
--p
P2,7662.85.611.3454OBc
Fe
490cLoliumperenneL.NoGramineaeM142P2,7732.85.711.3417OGallusf
FC:PI
491Loranthuseuropaeusl
NoLoranthaceaek
E--n
--p
--q
8,0858.316.533.0457bm
OBd
Fe
492bLupinusangustifoliusL.NoLeguminosaeE38,42,44--p
A752.an
0.8an
1.5an
3.1an
416C--an
FC:PI
493LupinusatlanticusGladst.NoLeguminosaeE38--p
A1,458.an
1.5an
3.0an
6.0an
416C--an
FC:PI
494LupinuscosentiniiGuss.NoLeguminosaeE32--p
A1,126.an
1.1an
2.3an
4.6an
416C--an
FC:PI
495LupinusdigitatusForssk.NoLeguminosaeE36--p
A1,286.an
1.3an
2.6an
5.3an
416C--an
FC:PI
496LupinusmicranthusGuss.NoLeguminosaeE52--p
A461.an
0.5an
0.9an
1.9an
416C--an
FC:PI
497LupinuspalaestinusBoiss.NoLeguminosaeE42--p
A1,201.an
1.2an
2.5an
4.9an
416C--an
FC:PI
498cLupinuspilosusMurr.NoLeguminosaeE42--p
A1,201.an
1.2an
2.5an
4.9an
416C--an
FC:PI
499cLuzulacampestris(L.)DC.NoJuncaceaeM12--u
P1,4431.52.95.9420OBc
Fe
500cLuzulaelegansGuthnickNoJuncaceaeM6--u
P1,5121.53.16.2420OBc
Fe
501dLuzulaluzuloides(Lam.)Dandy&WilmottNoJuncaceaeM12--u
P1,722.ap
1.8ap
3.5ap
7.0ap
420OBc
Fe
502dLuzulaniveaLam.&DC.NoJuncaceaeM12--u
P1,482.ap
1.5ap
3.0ap
6.1ap
420OBc
Fe
503cLuzulapedemontanaBoiss&Reut.NoJuncaceaeM30--u
P1,7171.83.57.0420OBc
Fe
504cLuzulapediformisDC.NoJuncaceaeM12--u
P1,5831.63.26.5420OBc
Fe
505LuzulaspicataDC.NoJuncaceaeM24--u
P1,9041.93.97.8420OBc
Fe
506LuzulasudeticaDC.NoJuncaceaeM48--u
P1,6861.73.46.9420OBc
Fe
507kLycopersiconesculentumMill.cv.
Gardener'sDelight
NoSolanaceaeE--n
--p
A9801.02.04.0382OHc
FC:PI
508bMalvasylvestrisL.--m
MalvaceaeE42
6P1,4701.53.06.0444bd
OGb2
FC:PI
509bMangiferaindicaL.NoAnacardiaceaeE40
4P8820.91.83.6454OBc
Fe
510Matthiolaincanal
NoCruciferaej
E--n
--p
--q
2,5972.75.310.6457bm
OBd
Fe
511aMelaleucaleucadendraL.NoMyrtaceaeE22
--p
P1,1101.12.34.5449bfOBc
Fe
512MelampyrumarvenseLinn.NoOrobanchaceaeE--n
--p
P8,0738.216.533.0465OGFC:PI
513MeliaazedarachL.NoMeliaceaeE28
--p
P4210.40.91.7454OBc
Fe
514MelianthusmajorL.NoMelianthaceaeE36,38
2or4P6270.61.32.6378OJFe
515Menthalongifolial
NoLabiataeE242P3850.40.81.6465OJFe
516Merrilliodendronmegacarpum(Hemsl.)
Sleum.
NoIcacinaceaeE302P1,0711.12.24.4380OKFC:PI
517MimusopselengiL.NoSapotaceaeE24
--p
P2740.30.61.1454OBc
Fe
518aMonsteradeliciosaLiebm.NoAraceaeM50--p
P9,3849.619.238.3411OBc
Fe
519MonsteraobliquaMiq.NoAraceaeM44--p
P8,8229.018.036.0411OBc
Fe
520MontiniacaryophyllaceaThunb.NoMontiniaceaeE242P5540.61.12.3380OJFe
521dMusaacuminataCollassp.banksii--m
MusaceaeM222P6000.61.22.5402OGlycinee
FC:PI
521eMusaacuminataCollassp.siamea--m
MusaceaeM222P6180.61.32.5402OGlycinee
FC:PI
521fMusaacuminataCollassp.banksiih
NoMusaceaeM22
2P5880.61.22.4410OPetuniae
FC:EB
521gMusaacuminataCollassp.malaccensis
AccessionSelangorh
NoMusaceaeM22
2P5980.61.22.4410OPetuniae
FC:EB
521hMusaacuminataCollassp.banksiih
NoMusaceaeM22
2P6370.71.32.6410OPetuniae
FC:EB
80 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
521iMusaacuminataCollassp.siamea
AccessionSiamh
NoMusaceaeM222P6520.71.32.7410OPetuniae
FC:EB
521jMusaacuminataCollassp.truncata--m
MusaceaeM222P6260.61.32.6421aq
OGlycinee
FC:PI
521kMusaacuminataCollagenotypePPC--m
MusaceaeM222P6100.61.22.5421aq
OGlycinee
FC:PI
522cMusabalbisianaColla--m
MusaceaeM222P5560.61.12.3421OGlycinee
FC:PI
522dMusabalbisianaColla--m
MusaceaeM222P5490.61.12.2402OGlycinee
FC:PI
522eMusabalbisianaCollaAccessionTani(1120)NoMusaceaeM22
2P5680.61.22.3410OPetuniae
FC:EB
523MusaornataRoxb.NoMusaceaeM22
2P6030.61.22.5410OPetuniae
FC:EB
524Musaviolascensl
--m
MusaceaeM202P6910.71.42.8421OGlycinee
FC:PI
525MuscariadiliiM.B.Guner&H.DumanNoAsparagaceaeM182P2,5502.65.210.4465OGFe
526Muscarimcbeathianuml
NoAsparagaceaeM182P2,9893.16.112.2465OBFe
527MyoporummauritianumA.DC.NoMyoporaceaeE728P1,9041.93.97.8379OGFe
528MyricagaleLinn.NoMyricaceaeE16,48,
80,96
--p
P4090.40.81.7379OJFe
529MyrsineafricanaL.NoMyrsinaceaeE46--p
P1,2051.22.54.9380OJFe
530NartheciumossifragumHuds.NoNartheciaceaeM26
2P4040.40.81.7380OJFC:PI
531NavarretiasquarrosaHook.&Arn.NoPolemoniaceaeE182A1,2891.32.65.3378OJFe
532NelumbonuciferaGaertn.NoNelumbonaceaeE162P2380.20.51.0379OJFe
533NemophilamenziesiiHook.&Arn.NoBoraginaceaeE182A1,2201.22.55.0465OJFe
534NepenthespervilleiBlumeNoNepenthaceaeE--n
--p
P2740.30.61.1378OJFe
535kNicotianatabacumL.NoSolanaceaeE48
4A7,5467.715.430.8457bm
OBd
Fe
536Nyctanthesarbor-tristisL.NoOleaceaeE44
--p
P1,2031.22.54.9454OBc
Fe
537Odontitesluteal
NoOrobanchaceaeE--n
--p
P5540.61.12.3465OJFe
538OdontitesvernaDum.NoOrobanchaceaeEc.182P5590.61.12.3465OJFe
539OdontostomumhartwegiiTorr.NoTechophilaeaceaeM202P2,5062.65.110.2380OGFC:PI
540OenotheraammophilaFockeNoOnagraceaeE142BP1,1421.22.34.7439OSorghume
FC:PI
541OenotherabiennisL.NoOnagraceaeE142BP1,1961.22.44.9439OSorghume
FC:PI
542OleaafricanaMill.NoOleaceaeE462P1,5481.63.26.3462OSorghume
Fe
543OleacuspidataWall.NoOleaceaeE462P2,0292.14.18.3462OSorghume
Fe
544aOleaeuropaeaL.cv.DolceAgogiah
NoOleaceaeE462P1,9112.03.97.8462OSorghume
Fe
544bOleaeuropaeaL.cv.Pendolinoh
NoOleaceaeE462P2,2832.34.79.3462OSorghume
Fe
545OleaferrugineaRoyaleNoOleaceaeE462P1,8131.93.77.4462OSorghume
Fe
546OleaindicaKleinNoOleaceaeE462P1,6461.73.46.7462OSorghume
Fe
547rOryzasativaL.ssp.indicaNoGramineaeM242A4660.51.01.9450bg
O--GS
547sOryzasativaL.ssp.indicacv.IR8--m
GramineaeM242A4400.40.91.8389OArab.e
FC:PI
547tOryzasativaL.ssp.japonicacv.Nipponbare--m
GramineaeM242A4010.40.81.6389OArab.e
FC:PI
547uOryzasativaL.ssp.japonicacv.NipponbareNoGramineaeM242A4200.40.91.7451bh
O--GS
548OxalisbolivianaBrittonNoOxalidaceaeE--n
--p
P4690.51.01.9456bk
OGallusf
FC:PI
549OxaliscoralleoidesR.KnuthNoOxalidaceaeE16
2P5210.51.12.1456bk
OGallusf
FC:PI
550OxaliscuzcensisR.KnuthNoOxalidaceaeE16
2AP5750.61.22.3456bk
OGallusf
FC:PI
551OxalisherreraeR.KnuthNoOxalidaceaeE16
2P4540.50.91.9456bk
OGallusf
FC:PI
552OxalishumbertiiR.KnuthNoOxalidaceaeE16
2P5100.51.02.1456bk
OGallusf
FC:PI
553OxalislotoidesKunthNoOxalidaceaeE16
2P4320.40.91.8456bk
OGallusf
FC:PI
554aOxalislucumayensisR.Knuthssp.
lucumayensis
NoOxalidaceaeE16
2P4550.50.91.9456bk
OGallusf
FC:PI
554bOxalislucumayensisR.Knuthssp.subiens
Lourteig
NoOxalidaceaeE16
2P4830.51.02.0456bk
OGallusf
FC:PI
555OxalismarcapatensisR.KnuthNoOxalidaceaeE16
2P4400.40.91.8456bk
OGallusf
FC:PI
556OxalismedicagineaKunthNoOxalidaceaeE16
2P4170.40.91.7456bk
OGallusf
FC:PI
557OxalismegalorrhizaJacquinNoOxalidaceaeE14or18
2P3990.40.81.6456bk
OGallusf
FC:PI
558OxalismollisKunthNoOxalidaceaeE16
2P4370.40.91.8456bk
OGallusf
FC:PI
559OxalisoulophoraLourteigNoOxalidaceaeE16
2P4330.40.91.8456bk
OGallusf
FC:PI
560OxalispaucartambensisR.KnuthNoOxalidaceaeE16
2P4700.51.01.9456bk
OGallusf
FC:PI
561aOxalispeduncularisKunthh
NoOxalidaceaeE16
2P4540.50.91.9456bk
OGallusf
FC:PI
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 81
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
561bOxalispeduncularisKunthvar.pilosah
NoOxalidaceaeE16
2P5700.61.22.3456bk
OGallusf
FC:PI
562OxalispetrophilaR.KnuthNoOxalidaceaeE16
2P4820.51.02.0456bk
OGallusf
FC:PI
563OxalisphaeotrichaDielsNoOxalidaceaeE32
4P8200.81.73.3456bk
OGallusf
FC:PI
564OxalispicchensisR.KnuthNoOxalidaceaeE32
4P8210.81.73.4456bk
OGallusf
FC:PI
565aOxalisptychocladaDielsNoOxalidaceaeE16
2P4340.40.91.8456bk
OGallusf
FC:PI
565bOxalisptychocladaDielsvar.
trichocarpaLourteig
NoOxalidaceaeE16
2P4660.51.01.9456bk
OGallusf
FC:PI
566Oxalissan-migueliiR.KnuthNoOxalidaceaeE16
2P4280.40.91.7456bk
OGallusf
FC:PI
567Oxalissp.cfr.melilotoidesZuccariniy
NoOxalidaceaeE16
2P4560.50.91.9456bk
OGallusf
FC:PI
568Oxalissp.cfr.teneriensisR.Knuthy
NoOxalidaceaeE16
2P4730.51.01.9456bk
OGallusf
FC:PI
569aOxalisspiralisR.&P.exG.Donh
NoOxalidaceaeE16
2AP5200.51.12.1456bk
OGallusf
FC:PI
569bOxalisspiralisR.&P.exG.Donh
NoOxalidaceaeE16
2AP6560.71.32.7456bk
OGallusf
FC:PI
570OxalistabaconasensisR.KnuthNoOxalidaceaeE16
2P5150.51.12.1456bk
OGallusf
FC:PI
571OxalistuberosaMolinaNoOxalidaceaeE64
8P1,7221.83.57.0456bl
OGallusf
FC:PI
572Oxalisunduavensis(Rusby)R.KnuthNoOxalidaceaeE16
2P5050.51.02.1456bk
OGallusf
FC:PI
573OxalisurubambensisR.KnuthNoOxalidaceaeE16
2P4310.40.91.8456bk
OGallusf
FC:PI
574OxalisvulcanicolaDonn.Sm.NoOxalidaceaeE16
2P4340.40.91.8456bk
OGallusf
FC:PI
575aPaeoniacaucasica(Schipcz.)Schipcz.h
--m
PaeoniaceaeE102P15,59215.931.863.6459C--bq
Fe
575bPaeoniacaucasica(Schipcz.)Schipcz.h
--m
PaeoniaceaeE102P16,08716.432.865.7459C--bq
Fe
576aPaeoniadauricaAndr.h
--m
PaeoniaceaeE102P11,80412.024.148.2459C--bq
Fe
576bPaeoniadauricaAndr.h
--m
PaeoniaceaeE102P12,97013.226.552.9459C--bq
Fe
577aPaeonialagodechianaKem.-Nath.h
--m
PaeoniaceaeE102P11,98112.224.548.9459C--bq
Fe
577bPaeonialagodechianaKem.-Nath.h
--m
PaeoniaceaeE102P14,00914.328.657.2459C--bq
Fe
578aPaeoniamacrophylla(Albov)Lomak.h
--m
PaeoniaceaeE204P29,44930.160.1120.2459C--bq
Fe
578bPaeoniamacrophylla(Albov)Lomak.h
--m
PaeoniaceaeE204P30,08630.761.4122.8459C--bq
Fe
579bPaeoniamlokosewitschiLomak.h
--m
PaeoniaceaeE102P16,79017.134.368.5459C--bq
Fe
579cPaeoniamlokosewitschiLomak.h
--m
PaeoniaceaeE102P17,57117.935.971.7459C--bq
Fe
580bPaeoniaofficinalisL.h
--m
PaeoniaceaeE204P25,99526.553.1106.1459C--bq
Fe
581aPaeoniaruprechtianaKem.-Nath.h
--m
PaeoniaceaeE102P15,22415.531.162.1459C--bq
Fe
581bPaeoniaruprechtianaKem.-Nath.h
--m
PaeoniaceaeE102P17,33117.735.470.7459C--bq
Fe
582aPaeoniastevenianaKem.-Nath.h
--m
PaeoniaceaeE204P27,95528.557.1114.1459C--bq
Fe
582bPaeoniastevenianaKem.-Nath.h
--m
PaeoniaceaeE204P29,93930.661.1122.2459C--bq
Fe
583bPaeoniatenuifoliaL.h
--m
PaeoniaceaeE102P7,7007.915.731.4459C--bq
Fe
583cPaeoniatenuifoliaL.h
--m
PaeoniaceaeE102P11,59811.823.747.3459C--bq
Fe
584aPaeoniatomentosa(Lomak.)N.Buschh
--m
PaeoniaceaeE204P25,26025.851.6103.1459C--bq
Fe
584bPaeoniatomentosa(Lomak.)N.Buschh
--m
PaeoniaceaeE204P27,39128.055.9111.8459C--bq
Fe
585aPaeoniawittmannianaHartwissexLindl.h
--m
PaeoniaceaeE204P27,51428.156.2112.3459C--bq
Fe
585bPaeoniawittmannianaHartwissexLindl.h
--m
PaeoniaceaeE204P31,04231.763.4126.7459C--bq
Fe
586dPapaverrhoeasL.NoPapaveraceaeE14
2A2,5482.65.210.4457bm
OBd
Fe
587ParmentieracereiferaSeem.NoBignoniaceaeE--n
--p
P6470.71.32.6454OBc
Fe
588PaspalumnotatumFlugge.NoGramineaeM202P7060.71.42.9417OGallusf
FC:PI
589Peltophorumpterocarpum(DC.)Bakerex
K.Heyne
NoLeguminosaeE26
--p
P7770.81.63.2454OBc
Fe
590aPetroselinumcrispumcv.ChampionMoss
Curledl
NoUmbelliferaeE--n
--p
P2,2052.34.59.0382OLycopers.c
FC:PI
82 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
Petunia­Sometaxaonceincludedin
Petuniaarenowincludedin
Calibrachoa(seefootnoteae)
591PetuniaalpicolaL.B.Sm.&DownsNoSolanaceaeE182P1,4501.53.05.9387ae
OGallus-398pFC:PI
592PetuniaaltiplanaT.Ando&Hashim.NoSolanaceaeE142P1,2741.32.65.2387ae
OGallus-398pFC:PI
593bPetuniaaxillaris(Lam.)Britton,Sterns&
Poggenb.ssp.axillaris
NoSolanaceaeE142P1,4361.52.95.9387ae
OGallus-398pFC:PI
593cPetuniaaxillaris(Lam.)Britton,Sterns&
Poggenb.ssp.subandina
NoSolanaceaeE14
2P1,4651.53.06.0387ae
OGallus-398pFC:PI
593dPetuniaaxillaris(Lam.)Britton,Sterns&
Poggenb.ssp.parodii
NoSolanaceaeE142P1,4701.53.06.0387ae
OGallus-398pFC:PI
594PetuniabajeensisT.Ando&Hashim.NoSolanaceaeE14
2P1,4501.53.05.9387ae
OGallus-398pFC:PI
595PetuniabonjardinensisT.Ando&Hashim.NoSolanaceaeE142P1,4211.52.95.8387ae
OGallus-398pFC:PI
596PetuniaexsertaStehmann.NoSolanaceaeE14
2P1,5391.63.16.3387ae
OGallus-398pFC:PI
597PetuniaguarapuavensisT.Ando&Hashim.NoSolanaceaeE142P1,4991.53.16.1387ae
OGallus-398pFC:PI
598PetuniahelianthemoidesSendtn.NoSolanaceaeE182P1,4361.52.95.9387ae
OGallus-398pFC:PI
599ePetuniahybridaVilm.cv.PearlSkyBlueNoSolanaceaeE14
2P1,4411.52.95.9387ae
OGallus-398pFC:PI
600aPetuniaintegrifolia(Hook.)Schinz&
Thell.ssp.inflata(R.E.Fr.)
NoSolanaceaeE142A1,3331.42.75.4387ae
OGallus-398pFC:PI
600bPetuniaintegrifolia(Hook.)Schinz&
Thell.ssp.integrifoliavar.integrifolia
NoSolanaceaeE142P1,4361.52.95.9387ae
OGallus-398pFC:PI
600cPetuniaintegrifolia(Hook.)Schinz&
Thell.ssp.integrifoliavar.
depauperata(R.E.Fr.)
NoSolanaceaeE142P1,4901.53.06.1387ae
OGallus-398pFC:PI
601PetuniainteriorT.Ando&Hashim.NoSolanaceaeE142P1,4551.53.05.9387ae
OGallus-398pFC:PI
602PetuniakleiniiL.B.Sm.&DownsNoSolanaceaeE182P1,4361.52.95.9387ae
OGallus-398pFC:PI
603PetunialittoralisL.B.Sm.&DownsNoSolanaceaeE142P1,4551.53.05.9387ae
OGallus-398pFC:PI
604PetuniamantiqueirensisT.Ando&Hashim.NoSolanaceaeE14
2P1,5241.63.16.2387ae
OGallus-398pFC:PI
605PetuniaoccidentalisR.E.Fr.NoSolanaceaeE142A1,3621.42.85.6387ae
OGallus-398pFC:PI
606Petuniapubescens(Spreng.)R.E.Fr.NoSolanaceaeE182P1,4461.53.05.9387ae
OGallus-398pFC:PI
607PetuniareitziiL.B.Sm.&Downs.NoSolanaceaeE142P1,4061.42.95.7387ae
OGallus-398pFC:PI
608PetuniariograndensisT.Ando&Hashim.NoSolanaceaeE14
2P1,4601.53.06.0387ae
OGallus-398pFC:PI
609PetuniasaxicolaL.B.Sm.&DownsNoSolanaceaeE142P1,4111.42.95.8387ae
OGallus-398pFC:PI
610PetuniascheideanaL.B.Sm.&DownsNoSolanaceaeE142P1,4361.52.95.9387ae
OGallus-398pFC:PI
611PetuniavariabilisR.E.Fr.NoSolanaceaeE182P1,4411.52.95.9387ae
OGallus-398pFC:PI
612PhalaenopsisamboinensisJ.J.SmithNoOrchidaceaeM382P7,0367.214.428.7447OGb2
FC:PI
613PhalaenopsisaphroditeRchb.f.NoOrchidaceaeM382P1,3721.42.85.6447OGb2
FC:PI
614Phalaenopsisbellina(Rchb.f.)CristensonNoOrchidaceaeM382P7,3657.515.030.1447OGb2
FC:PI
615Phalaenopsiscornu-cervi(Breda)Bl&Rchb.f.NoOrchidaceaeM382P3,1563.26.412.9447OGb2
FC:PI
616aPhalaenopsisequestris(Schauer)Rchb.f.NoOrchidaceaeM382P1,6511.73.46.7447OGb2
FC:PI
617PhalaenopsisfasciataRchb.f.NoOrchidaceaeM382P3,2143.36.613.1447OGb2
FC:PI
618PhalaenopsisgiganteaJ.J.SmithNoOrchidaceaeM382P2,5872.65.310.6447OGb2
FC:PI
619aPhalaenopsislueddemannianaRchb.f.NoOrchidaceaeM382P3,1803.26.513.0447OGb2
FC:PI
620PhalaenopsismanniiRchb.f.NoOrchidaceaeM382P6,6156.813.527.0447OGb2
FC:PI
621PhalaenopsismariaeBurb.exWarn.&Wms.NoOrchidaceaeM382P3,1753.26.513.0447OGb2
FC:PI
622PhalaenopsismicholitziiRolfeNoOrchidaceaeM382P3,1803.26.513.0447OGb2
FC:PI
623PhalaenopsismodestaJ.J.SmithNoOrchidaceaeM382P2,5242.65.210.3447OGb2
FC:PI
624PhalaenopsisparishiiRchb.f.NoOrchidaceaeM382P8,1398.316.633.2447OGb2
FC:PI
625Phalaenopsispulchra(Rchb.f.)SweetNoOrchidaceaeM382P3,1213.26.412.7447OGb2
FC:PI
626PhalaenopsissanderianaRchb.f.i
NoOrchidaceaeM382P1,3721.42.75.6447OGb2
FC:PI
627PhalaenopsisstuartianaRchb.f.NoOrchidaceaeM382P1,5341.63.16.3447OGb2
FC:PI
628PhalaenopsissumatranaKorth.&Rchb.f.NoOrchidaceaeM382P3,2443.36.613.2447OGb2
FC:PI
629PhalaenopsisvenosaShim&Fowl.NoOrchidaceaeM382P4,6654.89.519.0447OGb2
FC:PI
630cPhaseolusacutifoliusvar.latifoliusG.Freeman--m
LeguminosaeE22
2A7940.81.63.2390OPetuniae
FC:DAPI
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 83
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
630dPhaseolusacutifoliusvar.tenuifolius
(Wood&Standl)A.Gray
--m
LeguminosaeE22
2A7990.81.63.3390OPetuniae
FC:DAPI
630ePhaseolusacutifoliusvar.acutifoliusA.Gray--m
LeguminosaeE22
2A8620.91.83.5390OPetuniae
FC:DAPI
631PhaseolusangustissimusA.Gray--m
LeguminosaeE22
2--q
6470.71.32.6390OPetuniae
FC:DAPI
632ePhaseoluscoccineusL.--m
LeguminosaeE22
2P7840.81.63.2390OPetuniae
FC:DAPI
632fPhaseoluscoccineusL.ssp.Purpurascens--m
LeguminosaeE22
2P7940.81.63.2390OPetuniae
FC:DAPI
632gPhaseoluscoccineusL.ssp.coccineus
cv.Hammond'sDwarfScarleth
--m
LeguminosaeE22
2P7940.81.63.2390OPetuniae
FC:DAPI
632hPhaseoluscoccineusL.ssp.coccineuscv.
Preisgewinnerh
--m
LeguminosaeE22
2P8090.81.73.3390OPetuniae
FC:DAPI
632iPhaseoluscoccineusL.NoLeguminosaej
E22
2P1,7151.83.57.0457bm
OBd
Fe
633bPhaseolusfiliformisBenth.--m
LeguminosaeE22
2P6910.71.42.8390OPetuniae
FC:DAPI
634bPhaseolusglabellusPiper--m
LeguminosaeE22
2P1,0241.02.14.2390OPetuniae
FC:DAPI
635PhaseolusgrayanusWood.&Standl--m
LeguminosaeE22
2--q
9311.01.93.8390OPetuniae
FC:DAPI
636bPhaseolushintoniiDelgado--m
LeguminosaeE22
2P7150.71.52.9390OPetuniae
FC:DAPI
637Phaseolusleptostachysvar.leptostachysBenth.--m
LeguminosaeE22
2--q
6130.61.32.5390OPetuniae
FC:DAPI
638dPhaseoluslunatusL.var.lunatuscv.
EarlyThorogreenh
--m
LeguminosaeE22
2P6910.71.42.8390OPetuniae
FC:DAPI
638ePhaseoluslunatusL.var.silvesterBaudet--m
LeguminosaeE22
2P6960.71.42.8390OPetuniae
FC:DAPI
638fPhaseoluslunatusL.var.lunatuscv.
HendersonBushh
--m
LeguminosaeE22
2P7010.71.42.9390OPetuniae
FC:DAPI
639bPhaseolusmarechalliDelgado--m
LeguminosaeE22
2P7840.81.63.2390OPetuniae
FC:DAPI
640PhaseolusmicranthusHook.&Arn.--m
LeguminosaeE22
2--q
5880.61.22.4390OPetuniae
FC:DAPI
641PhaseolusmicrocarpusMart.--m
LeguminosaeE22
2--q
5050.51.02.1390OPetuniae
FC:DAPI
642bPhaseolusneglectusHerm.--m
LeguminosaeE22
2--q
9411.01.93.8390OPetuniae
FC:DAPI
643PhaseolusparviflorusG.Freytag--m
LeguminosaeE22
2--q
6370.71.32.6390OPetuniae
FC:DAPI
644bPhaseoluspluriflorusMarechal--m
LeguminosaeE22
2--q
1,0681.12.24.4390OPetuniae
FC:DAPI
645bPhaseoluspolyanthusGreenm.--m
LeguminosaeE22
2P7990.81.63.3390OPetuniae
FC:DAPI
646gPhaseolusvulgarisL.cv.KentuckyWonderh
--m
LeguminosaeE22
2A6860.71.42.8390OPetuniae
FC:DAPI
646hPhaseolusvulgarisL.var.aborigineus
(Burk.)Baudet
--m
LeguminosaeE22
2A7200.71.52.9390OPetuniae
FC:DAPI
646iPhaseolusvulgarisL.var.mexicanus--m
LeguminosaeE22
2A7350.81.53.0390OPetuniae
FC:DAPI
646jPhaseolusvulgarisL.cv.Sanilac--m
LeguminosaeE22
2A7500.81.53.1390OPetuniae
FC:DAPI
646kPhaseolusvulgarisL.NoLeguminosaej
E22
2A1,666.bn
1.7bn
3.4bn
6.8bn
457bm
OBd
Fe
647bPhaseolusxanthotrichusPiper
var.xanthotrichus
--m
LeguminosaeE22
2P6620.71.42.7390OPetuniae
FC:DAPI
647cPhaseolusxanthotrichusPiper--m
LeguminosaeE22
2P8480.91.73.5390OPetuniae
FC:DAPI
648PhilodendronerubescensC.Koch&BouscheNoAraceaeM42--p
P5,1745.310.621.1411OBc
Fe
649PhilodendronselloumC.KochNoAraceaeM36--p
P4,8955.010.020.0411OBc
Fe
650PhilodendronsquamiferumPoepp.&Endl.NoAraceaeM30--p
P4,5574.79.318.6411OBc
Fe
651Phormiumtenaxl
NoHemerocallidaceaeM322P7400.81.53.0379OJFe
652PinguiculaprimulifloraC.E.Wood&GodfreyNoLentibulariaceaeE222P6690.71.42.7378OJFe
653PiptocalyxmooreiOliverNoTrimeniaceaeBA16
2P4,0014.18.216.3381OGFC:PI
654bPistiastratiotesL.NoAraceaek
M282P2500.30.51.0400OGFe
655cPisumabyssinicumA.Braunh
NoLeguminosaeE142A4,3714.58.917.8458bp
Cbp
Gc
FC:EB
655dPisumabyssinicumA.Braunh
NoLeguminosaeE142A4,7514.89.719.4458bp
Cbp
Gc
FC:EB
84 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
656cPisumelatiusStevenexM.Bieb.h
NoLeguminosaeE142A4,2314.38.617.3458bp
Cbp
Gc
FC:EB
656dPisumelatiusStevenexM.Bieb.h
NoLeguminosaeE142A4,9785.110.220.3458bp
Cbp
Gc
FC:EB
657bPisumfulvumSibth.&SmithNoLeguminosaeE142A4,7164.89.619.3458bp
Cbp
Gc
FC:EB
658cPisumhumileBoiss&Noe¨h
NoLeguminosaeE142A4,2584.38.717.4458bp
Cbp
Gc
FC:EB
658dPisumhumileBoiss&Noe¨h
NoLeguminosaeE142A4,8094.99.819.6458bp
Cbp
Gc
FC:EB
659PittosporumtenuifoliumGaertn.NoPittosporaceaeE242P4530.50.91.9379OGFe
660Planchonellaeerwah(F.M.Bailey)vanRoyenNoSapotaceaeEc.242P5270.51.12.2380OJFe
661PlantagoafraL.NoPlantaginaceaeE122A1,1291.22.34.6388OBc
Fe
662PlantagoarenariaW.&K.NoPlantaginaceaeE122A1,1151.12.34.6388OBc
Fe
663PlantagocoronopusL.NoPlantaginaceaeE102AP8450.91.73.5388OBc
Fe
664PlantagoindicaL.NoPlantaginaceaeE122--q
1,0881.12.24.4388OBc
Fe
665bPlantagolagopusL.NoPlantaginaceaeE122--q
1,0461.12.14.3388OBc
Fe
666cPlantagolanceolataL.NoPlantaginaceaeE122P1,2991.32.75.3388OBc
Fe
667cPlantagomajorL.NoPlantaginaceaeE122P8670.91.83.5388OBc
Fe
668PlantagopsylliumL.NoPlantaginaceaeE122--q
1,1421.22.34.7388OBc
Fe
669PlantagoserrariaL.NoPlantaginaceaeE102--q
8820.91.83.6388OBc
Fe
670PlantagostepposaK.NoPlantaginaceaeE244--q
1,5931.63.36.5388OBc
Fe
671PlatanusorientalisL.NoPlatanaceaeE422P1,2741.32.65.2379OJFe
672PoapratensisL.NoGramineaeM58-62--p
P4,1554.28.517.0417OGallusf
FC:PI
673Poncirustrifoliata(L.)Raf.NoRutaceaeE18
2P3770.40.81.5426OGallusf
FC:PI
674Prosopiscineraria(L.)DruceNoLeguminosaeE52
--p
P1,2521.32.65.1454OBc
Fe
675Protiumserratum(Wall.exColebr.)NoBurseraceaeE--n
--p
P9240.91.93.8454OBc
Fe
676PterospermumlanceifoliumRoxb.NoMalvaceaeE38
--p
P7860.81.63.2454OBc
Fe
677PterostyraxpsilophyllaDielsexPerkinsNoStyracaceaeE242P8670.91.83.5380OJFe
678PunicagranatumL.NoSonneratiaceaeE16
2P7060.71.42.9454OBc
Fe
679ResedaluteolaL.NoResedaceaeE262B5000.51.02.0378OJFe
680RhapidophoramontanaSchottNoAraceaeM30--p
P9,82910.020.140.1411OBc
Fe
681RhapidophorapeeplaSchottNoAraceaeM18--p
P8,9849.218.336.7411OBc
Fe
682RhipogonumpapuanumC.T.WhiteNoRhipogonaceaeM302P10,92211.122.344.6380OGFe
683Rhodocomagigantea(Kunth)H.P.LinderNoRestionaceaeM--n
--p
P7280.71.53.0380OJFC:PI
684Rhodohypoxismilloides(Baker)Hilliard&
B.L.Burtt
NoHypoxidaceaeM24+1-2B4P1,3941.42.85.7379OJFe
685cRhoeodiscolorHanceNoCommelinaceaeM12
2P7,9878.216.332.6457bm
OBd
Fe
686RhoiacarposcapensisA.DC.NoSantalaceaeE--n
--p
P3040.30.61.2379OJFe
687RhynchosiacyanospermaBenth.ExBakerNoLeguminosaeE222B2,7272.85.611.1443bc
OBc
Fe
688Rhynchosiaminima(L.)DC.NoLeguminosaeE222P1,2271.32.55.0443bc
OBc
Fe
689Ribesglutinosuml
NoGrossulariaceaeE162P5340.51.12.2379OJFe
690RoridulagorgoniasPlanch.NoRoridulaceaeE122P1860.20.40.8379OGFe
691cRutagraveolensL.NoRutaceaeE--n
--p
P7350.81.53.0457bm
OBd
Fe
692SalixalbaL.h
--m
SalicaceaeE76ad
4ad
P8090.81.73.3385ac
OLycopers.c
FC:PI
693SalixatrocinereaBrot.h
--m
SalicaceaeE76ad
4ad
P8040.81.63.3385ac
OLycopers.c
FC:PI
694bSalixcapreaL.h
--m
SalicaceaeE38ad
2ad
P4700.51.01.9385ac
OLycopers.c
FC:PI
695SalixcinereaL.h
--m
SalicaceaeE76ad
4ad
P8280.81.73.4385ac
OLycopers.c
FC:PI
696SalixelaeagnosScop.h
--m
SalicaceaeE38ad
2ad
P4170.40.91.7385ac
OLycopers.c
FC:PI
697SalixfragilisL.h
--m
SalicaceaeE76ad
4ad
P8430.91.73.4385ac
OLycopers.c
FC:PI
698SalixpurpureaL.h
--m
SalicaceaeE38ad
2ad
P4610.50.91.9385ac
OLycopers.c
FC:PI
699SalixpyrenaicaGouanh
--m
SalicaceaeE38ad
2ad
P4700.51.01.9385ac
OLycopers.c
FC:PI
700SalixtriandraL.h
--m
SalicaceaeE38ad
2ad
P3870.40.81.6385ac
OLycopers.c
FC:PI
701SalixviminalisL.h
--m
SalicaceaeE38ad
2ad
P4020.40.81.6385ac
OLycopers.c
FC:PI
702SalixviminalisL.h
--m
SalicaceaeE76ad
4ad
P7940.81.63.2385ac
OLycopers.c
FC:PI
703aSambucusnigraL.NoAdoxaceaek
E36
2P14,945.bo
15.3bo
30.5bo
61.0bo
457bm
OBd
Fe
704SantalumalbumL.NoSantalaceaeE20
--p
P2820.30.61.2454OBc
Fe
705bSaxifragagranulataL.ssp.granulataNoSaxifragaceaeE22--p
P6620.71.42.7453OGc
FC:EB
706SaxifragagranulataL.NoSaxifragaceaeE--bj
--p
P1,1221.12.34.6453OGc
FC:EB
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 85
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
707aSaxifragagranulataL.NoSaxifragaceaeE52--p
P2,3322.44.89.5453OGc
FC:EB
707bSaxifragagranulataL.ssp.fernandesii
Redondo&Horjalesi
NoSaxifragaceaeE44-56--p
P1,7351.83.57.1453bi
OGc
FC:EB
708Schisandrarubrifloral
NoSchisandraceaeBA--n
--p
P8,9389.118.236.5381OGFC:PI
709Schleicheraoleosa(Lour.)OkenNoSapindaceaeE32
--p
P1,1421.22.34.7454OBc
Fe
710bScillaindica(Roxb.)Baker(cytotypeII)NoAsparagaceaeM30--p
P3,5043.67.214.3422ar
OBc
Fe
710cScillaindica(Roxb.)Baker(cytotypeI)h
NoAsparagaceaeM30--p
P5,7015.811.623.3422ar
OBc
Fe
711Scillanervosa(Burch.)J.P.JessopNoAsparagaceaeM38--p
P3,9644.08.116.2422OBc
Fe
712fScillasibericaHaw.inAndr.NoAsparagaceaeM122P30,13530.861.5123.0422OBc
Fe
713ScillatalosiiD.Tzanoudakis&KypriotakisNoAsparagaceaeMc.150--p
P45,84046.893.6187.1465OBFe
714cScillavindobonensisSpetaNoAsparagaceaeM183P--t
--t
17.935.7422OBc
Fe
715ScindapsuspictusHasskNoAraceaeM60--p
P11,51711.823.547.0411OBc
Fe
716SedumacreL.NoCrassulaceaeE--n
--p
P1,2251.32.55.0457bm
OBd
Fe
717SedumalbumL.--m
CrassulaceaeE342P1420.10.30.6398O--w
--r
718SedumforsterianumSm.--m
CrassulaceaeE242P4510.50.91.8398O--w
--r
719SedummontanumSong.&Perrier--m
CrassulaceaeE342P5150.51.12.1398O--w
--r
720aSedumobtusifoliumC.A.Meyer--m
CrassulaceaeE122P2060.20.40.8398O--w
--r
720bSedumobtusifoliumC.A.Meyer--m
CrassulaceaeE122P2060.20.40.8399OB-723bFe
721SedumobtusifoliumC.A.Meyer--m
CrassulaceaeE305P--t
--t
1.73.4399OB-723bFe
722SedumochroleucumChaix--m
CrassulaceaeE342P4460.50.91.8398O--w
--r
723aSedumrupestreL.ssp.erectum--m
CrassulaceaeE644P1,0141.02.14.1398O--w
--r
723bSedumrupestreL.ssp.rupestre--m
CrassulaceaeE--n
--p
P2,2442.34.69.2399OBFe
724Sedumsediforme(Jacq.)Pau--m
CrassulaceaeE322P5680.61.22.3398O--w
--r
725SedumspuriumBieb.--m
CrassulaceaeE284P1,7351.83.57.1399OB-723bFe
726SedumspuriumBieb.--m
CrassulaceaeE426P2,7642.85.611.3399OB-723bFe
727aSedumstellatumL.--m
CrassulaceaeE102P2890.30.61.2399OB-723bFe
727bSedumstellatumL.--m
CrassulaceaeE102P2890.30.61.2398O--w
--r
728aSedumstoloniferumS.G.Gmelin--m
CrassulaceaeE142P3090.30.61.3399OB-723bFe
728bSedumstoloniferumS.G.Gmelin--m
CrassulaceaeE142P3090.30.61.3398O--w
--r
729SenecioviscosusL.NoCompositaej
E--n
--p
A1,5191.63.16.2457bm
OBd
Fe
730SesamumalatumThonn.NoPedaliaceaeE262A1,6511.73.46.7446OGFe
731SesamumcapenseBurm.NoPedaliaceaeE262A1,1881.22.44.9446OGFe
732SesamumindicumL.NoPedaliaceaeE262A9511.01.93.9446OGFe
733SesamumlaciniatumKlein.NoPedaliaceaeE324A1,1541.22.44.7446OGFe
734SesamumlatifoliumGillett.NoPedaliaceaeE324A9331.01.93.8446OGFe
735SesamummulayanumNair.NoPedaliaceaeE262A8700.91.83.6446OGFe
736SesamumoccidentaleRegel.NoPedaliaceaeE648A1,5511.63.26.3446OGFe
737SesamumradiatumSchumach.NoPedaliaceaeE648A1,3061.32.75.3446OGFe
738SesamumschinzianumAschers.NoPedaliaceaeE648A1,3431.42.75.5446OGFe
739SesamumtriphyllumWelw.exAschers.NoPedaliaceaeEc.262A5240.51.12.1378OJFe
740bSesleriaalbicansKit.exSchult.h
NoGramineaej
M28
4P4,7484.89.719.4428OHomof
FC:PI
740cSesleriaalbicansKit.exSchult.h
NoGramineaej
M28
4P4,8274.99.919.7428OHomof
FC:PI
741Severiniabuxifolia(Poir.)Ten.NoRutaceaeE18
2P3280.30.71.3426OGallusf
FC:PI
742SilenechalcedonicaL.NoCaryophyllaceaeE242P3,2293.36.613.2437OLycopers.c
FC:PI
743cSilenelatifoliaPoiret(female)NoCaryophyllaceaeE242AP2,8082.95.711.5437az
OLycopers.c
FC:PI
743dSilenelatifoliaPoiret(male)NoCaryophyllaceaeE242AP2,8672.95.911.7437az
OLycopers.c
FC:PI
86 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
744SilenependulaL.NoCaryophyllaceaeE242A1,1521.22.44.7437OHc
FC:PI
745Silenevulgaris(Moench)GarckeNoCaryophyllaceaeE242P1,1031.12.34.5437OHc
FC:PI
746aSimmondsiachinensis(Link)C.K.Schneid.NoSimmondsiaceaeE48-50--p
P7230.71.53.0465OJFe
747Spirodelapolyrrhiza(L.)Schleid.NoAraceaek
M804P2920.30.61.2400OGFe
748Spirodelapunctata(G.F.W.Meyer)ThompsonNoAraceaeM462P3630.40.71.5400OGFe
749StemonatuberosaLour.NoStemonaceaeM--n
--p
P7150.71.52.9379OJFe
750Stenotaphrumsecundatum(Walt.)Kuntze.NoGramineaeM182P5290.51.12.2417OGallusf
FC:PI
751StrelitzianicolaiRegel&C.KochNoStrelitziaceaeM14,22
2P5660.61.22.3379OJFe
752StreptocarpuscyaneusS.MooreNoGesneriaceaeEc.302or4P6620.71.42.7378OJFe
753StylidiumadnatumR.Br.NoStylidiaceaeE302P1,4951.53.16.1378OJFe
754StylobasiumspathulatumDesf.NoSurianaceaeE302P1,2841.32.65.2378OJFe
755Syngoniumalbo-lineatumBullNoAraceaeM22--p
AP4,6284.79.418.9411OBc
Fe
756SyngoniumpodophyllumSchottNoAraceaeM24--p
AP4,7554.99.719.4411OBc
Fe
757Tabebuiaargentea(Bureau&K.Schum.)
Britton
NoBignoniaceaeE--n
--p
P7820.81.63.2454OBc
Fe
758TamarindusindicaL.NoLeguminosaeE26
--p
P8210.81.73.4454OBc
Fe
759Tecomastans(L.)Juss.exKunthNoBignoniaceaeE36
--p
P5930.61.22.4454OBc
Fe
760Thespesialampas(Cavanilles)Dalzellex
Dalzell&Gibson
--m
MalvaceaeE26
2--q
1,5681.63.26.4444bd
OGb2
FC:PI
761aThespesiapopulnea(L.)SolanderexCorrea--m
MalvaceaeE262P4,0184.18.216.4444bd
OGb2
FC:PI
761bThespesiapopulnea(L.)SolanderexCorreaNoMalvaceaeE26
2P3,0113.16.113.3454OBc
Fe
762Thespesiathespesioides(R.BrownexBentham)
Fryxell
--m
MalvaceaeE262--q
1,5681.63.26.4444bd
OGb2
FC:PI
763TriteleialaxaBenth.NoAsparagaceaeM28
4P10,43510.621.342.6380OBFC:PI
764TrochodendronaralioidesSiebold&Zucc.NoTrochodendraceaeE382P1,8721.93.87.6380OLycopers.c
FC:PI
765TyphoniumcuspidatumDecne.NoAraceaeM16--p
A5,0645.210.320.7411OBc
Fe
766TyphoniumtrilobatumSchottNoAraceaeM40--p
A6,4536.613.226.3411OBc
Fe
767ViciacanescensLab.NoLeguminosaeE102P3,0583.16.212.5409OBc
Fe
768aViciacraccaL.ssp.tenuifoliaNoLeguminosaeE142P5,7975.911.823.7409OBc
Fe
768bViciacraccaL.ssp.craccaNoLeguminosaeE142P6,409.al
6.5al
13.1al
26.2al
409OBc
Fe
769aViciaepetiolarisBurk.h
NoLeguminosaej
E142A4,0674.28.316.6415OBFe
769bViciaepetiolarisBurk.h
NoLeguminosaej
E142A4,5334.69.318.5415OBFe
770bViciaeristalioidesMaxtedNoLeguminosaeE142A9,4529.619.338.6406OVicianarb.e
Fe
771rViciafabaL.NoLeguminosaej
E12
2A12,74013.026.052.0457bm
OBd
Fe
771sViciafabaL.var.equinaNoLeguminosaeE122A14,37714.729.358.7408OCd
Fe
771tViciafabaL.`FuturaRZ'--m
LeguminosaeE--n
--p
A12,98513.326.553.0384aa
OHomof
FC:PI
772aViciagalilaeaPlitm.&Zoh.NoLeguminosaeE142A6,3926.513.026.1408OCd
Fe
772bViciagalilaeaPlitm.&Zoh.NoLeguminosaeE142A7,9048.116.132.3409OBc
Fe
773dViciagramineaSm.i
NoLeguminosaej
E142AB4,9695.110.120.3415OBFe
774bViciahyaeniscyamusMout.NoLeguminosaeE142A7,6547.815.631.2408OCd
Fe
775eViciahybridaL.NoLeguminosaeE122A8,3068.517.033.9409OBc
Fe
776eViciahyrcanicaFisch.&Mey.NoLeguminosaeE122A7,6347.815.631.2409OBc
Fe
777cViciajohannisTamamsch.NoLeguminosaeE142A6,1456.312.525.1408OCd
Fe
778bViciakalakhensisKhattab,Maxted&BisbyNoLeguminosaeE142A10,34410.621.142.2406OVicianarb.e
Fe
779ViciamacrogramineaBurk.NoLeguminosaej
E142BP5,8215.911.923.8415OBFe
780dViciamelanopsSibth.&Sm.NoLeguminosaeE102A6,764.ak
6.9ak
13.8ak
27.6ak
408OCd
Fe
781ViciananaVog.NoLeguminosaej
E142A4,3764.58.917.9415OBFe
782aViciapampicolaBurk.h
NoLeguminosaej
E142A4,6404.79.518.9415OBFe
782bViciapampicolaBurk.h
NoLeguminosaej
E142A5,2235.310.721.3415OBFe
783dViciaperegrinaL.NoLeguminosaeE142A9,5409.719.538.9409OBc
Fe
784dViciapisiformisL.h
NoLeguminosaeE122P6,2236.412.725.4407OCd
Fe
784eViciapisiformisL.h
NoLeguminosaeE122P8,0388.216.432.8407OCd
Fe
785vViciasativaL.line20.1i
NoLeguminosaeE122A1,8821.93.87.7409OBc
Fe
785wViciasativaL.line31i
NoLeguminosaeE122A2,4452.55.010.0409OBc
Fe
Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms 87
APPENDIX.(continued,thesuperscriptlettersrefertonotesprecedingthistable)
Ploidy
level
(x)
Life
cycle
typex
DNAamount
Entry
numberg
SpeciesVoucherFamily
Higher
group#
2nz
1C
(Mbps
)
1C
(pg)
2C
(pg)
4C
(pg)
Original
ref.a
Present
amounty
Standard
species*b1
Methodyy
785xViciasativaL.ssp.amphicarpaNoLeguminosaeE122A2,1022.14.38.6409OBc
Fe
785yViciasativaL.ssp.nigravar.nigraNoLeguminosaeE122A2,3372.44.89.5409OBc
Fe
786eViciasepiumL.NoLeguminosaeE142P4,7194.89.619.3409OBc
Fe
787cViciaserratifoliaJacq.NoLeguminosaeE142A9,7009.919.839.6405OCc
Fe
788dViscumalbumL.NoLoranthaceaek
E20
2P52,43053.5107.0214.0457bm
OBd
Fe
789VitexnegundoL.NoLamiaceaeE34
--p
P1,5901.63.26.5454OBc
Fe
790VitexpinnataL.NoLamiaceaeE--n
--p
P1,4111.42.95.8454OBc
Fe
791Voacangagrandifolia(Miq.)RolfeNoApocynaceaeE--n
--p
P3580.40.71.5454OBc
Fe
792Wolffiaarrhiza(L.)HorkelexWimmerNoAraceaeM422P1,6001.63.36.5400OGFe
793Wolffiellaoblonga(Phil.)Hegelm.NoAraceaeM422P7420.81.53.0400OGFe
794XanthorrhoeapreisiiEndl.NoXanthorrhoeaceaeM222P1,0141.02.14.1380OJFe
795Xanthosomasagittifolium(L.)SchottNoAraceaeM38--p
P8,6098.817.635.1411OBc
Fe
796XeronemacallistemonW.R.B.Oliv.NoXeronemataceaeM342or4P3,2103.36.613.1380OGFC:PI
797XerophytahumilisTh.Dur.&Schinz.NoVelloziaceaeM48
4or8P5320.51.12.2378OJFe
798XimeniaamericanaLinn.NoOlacaceaeE262P1,5951.63.36.5379OJFe
799XiphidiumcaeruleumAubl.var.caeruleumNoHaemodoraceaeM38
2P7670.81.63.1378OJFe
800Xyrisgracilisssp.gracilisl
NoXyridaceaeM26
2P6,8677.014.028.0380OBFC:PI
801bxZeamaysssp.maysL.lineopaque2h
NoGramineaej
M204A3,2623.36.713.3392OBFe
801byZeamaysssp.maysL.raceAltiplanoh
NoGramineaej
M204A2,4542.55.010.0392ag
OB-801bxFe
801bzZeamaysssp.maysL.raceBlancoyocho
rayash
NoGramineaej
M204A3,3113.46.813.5392ag
OB-801bxFe
801caZeamaysL.NoGramineaej
M20
4A3,2833.46.713.4457bm
OBd
Fe
802ZizyphusglabrataHeyneNoRhamnaceaeE24
--p
P1,5171.53.16.2454OBc
Fe
803Zosteramarinal
NoZosteraceaeM12
2P3090.30.61.3380OJFe
804ZoysiajaponicaSteud.NoGramineaeM404P4210.40.91.7417OGallusf
FC:PI
88 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms
Original references for DNA values
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90 Bennett and Leitch -- Nuclear DNA Amounts in Angiosperms