Global Ecology and Biogeography, (Global Ecol. Biogeogr.

) (2005) 14, 455–468

How many branchiopod crustacean

Blackwell Publishing, Ltd.

RESEARCH
PAPER
species are there? Quantifying the
components of underestimation
Sarah J. Adamowicz* and Andy Purvis

Department of Biological Sciences, Imperial ABSTRACT

College London, Silwood Park Campus, Ascot,
UK, SL5 7PY
*Correspondence: Sarah J. Adamowicz,
Division of Biology, Imperial College London,
Silwood Park Campus, Ascot, UK, SL5 7PY.
E-mail: sarah.adamowicz@imperial.ac.uk
Aims For the crustacean class Branchiopoda, we estimated global species richness,
assessed the reliability of relative diversity estimates among higher taxa and made
predictions about the kinds of species remaining to be described.
Location Global.
Methods We calculated correction factors to account for three different sources of
diversity underestimation, analysed taxonomic patterns of species descriptions over
time and tested possible correlates of description date.
Results Our work suggests that there are c. 2.1 times more branchiopod species in
nature than the c. 1180 currently described. The largest correction factor was at the
scale of single biogeographical regions, mainly attributed to poor knowledge of
species diversity within described morphospecies. Relative diversity estimates among
major taxonomic groups are apparently quite reliable, with diversity differences
greater than 1.5-fold robust to the heterogeneity of the taxonomic process. Current
knowledge of branchiopod diversity is biased towards species with large geographical
ranges, whereas body size and higher taxonomic group were only weak predictors of
description date.
Main conclusions A proper understanding of species boundaries in nature,
which is greatly aided by molecular studies, is key in accurately estimating species
richness for invertebrate groups. However, we suggest that while many species
remain to be described, numbers of presently described species are still useful for
macroevolutionary studies of diversification when interpreted with caution.
Keywords
Cladocera, clam shrimp, conchostracan, estimating diversity, extrapolation, fairy
shrimp, freshwater, tadpole shrimp, water fleas.

(e.g. Colwell & Coddington, 1994; Longino et al., 2002). The

INTRODUCTION
The questions of how many kinds of life there are, how they are
distributed and how best to conserve them are becoming ever
more pressing (Pimm et al., 1995; Wilson, 2003; Mace, 2004).
The urgency is especially great for hyper-diverse but poorly
known groups such as arthropods (Hammond, 1994), and those
inhabiting environments that face particularly severe threats,
such as fresh water (IUCN, 2003).
Extrapolating from known information to estimate total
species richness is a difficult problem even with extensive
sampling and at a local scale, where the numerous rare species
and biases in catchability must be analytically accommodated
problem of estimating total richness becomes more contentious
at a global scale because of the necessity of choosing among
major assumptions about scaling. For example, new insights into
host specificity of tropical arthropods on tree species at large geo-
graphical scales and across different tree species necessitated a
substantial downward revision in Erwin’s (1982) famous rough
calculation of 30 million tropical arthropod species worldwide
(Ødegaard, 2000). Similarly, May (1994) argued that Grassle and
Maciolek’s (1992) suggestion of c. 10 million benthic marine
invertebrate species was an overestimate, because of the general
poor knowledge of how heterogeneity and turnover at regional
scales relate to the patterns at global scales. Rather, May (1994)

© 2005 Blackwell Publishing Ltd www.blackwellpublishing.com/geb DOI: 10.1111/j.1466-822x.2005.00164.x 455

S. J. Adamowicz & A. Purvis
suggested that the best indicators of total diversity may be ratios
of unknown to known species in samples. Similarly, Hammond
(1994) advocated avoiding the use of ecological principles,
because of our current inadequate knowledge of the rules of the
distribution and turnover of biodiversity, and basing extrapolations
on well-estimated simple ratios instead, such as ratios in the
diversity of different geographical regions or taxonomic groups.
In this paper, we take a novel intensive approach to the
problem of estimating species diversity in a globally distributed
freshwater arthropod taxon, the crustacean class Branchiopoda.
In light of the criticisms of some earlier studies noted above,
we extrapolate using only information for species in this clade
and within given spatial scales. The modest size of Branchiopoda
(c. 1200 species) permits analyses on all known species. The
group contains important members of communities in most
types of standing inland waters, including temporary pools,
ponds, lakes and saline lakes. Ranging from about 0.18 to
100 mm in length (Dumont & Negrea, 2002), most are filter
feeders as in the well-known taxon Daphnia, but the class also
contains grazers on vegetation or substrates, predators,
omnivores and interstitial benthic dwellers. We estimate global
species richness for the class and assess the magnitude of
three factors likely to have led to the underestimation of true
branchiopod diversity: (1) different levels of taxonomic effort
among continents; (2) incorrect assumptions of multicontinental
distributions of many species; and (3) incomplete knowledge of
faunas on single continents. This last factor involves two com-
ponents: rare species may be missed in surveys and the knowledge
of ‘intraspecific’ variability is often superficial, resulting in
described morphospecies comprising multiple true species.
Secondly, recognizing that described diversity is generally the
only diversity data readily available for studies of macroecology
and macroevolution, we also assess the reliability of relative
diversity estimates among orders and families. Finally, we
address the issue of what is not known (i.e. what kinds of species
probably remain to be described?). We analyse potential cor-
relates of description date, assuming that species yet to be
discovered will be more similar to those described in the recent
than in the distant past. We conclude with some comparisons
and recommendations for speeding the species discovery process.
METHODS
In this work, we address one aspect of biological diversity —
species richness — which we define as the total number of species
in a taxonomic group worldwide. We note that, although diversity
can be characterized in many different ways, we use the terms
‘richness’ and ‘diversity’ interchangeably here.
The taxonomic data set
We compiled a list of 1176 valid branchiopod species (290 anos-
tracans, 187 spinicaudatans, 35 laevicaudatans, 2 cyclestheridans,
15 notostracans, 560 anomopods, 52 ctenopods, 33 onychopods
and 2 haplopods; c. 10 species were not included here because of
literature unavailability) described up until 1 November 2003.
456 Global Ecology and Biogeography, 14, 455–468 © 2005 Blackwell Publishing Ltd
For each species, we recorded the description date and bio-
geographical regions it occupied. References and details regarding
the assembly of the branchiopod taxon list are presented in
Appendix S1 (see Supplementary Material).
Estimation of global species richness
We estimated three diversity correction factors to be applied to
described branchiopod diversity in order to extrapolate to global
diversity. Each of the factors was constructed to be a multiplier,
to facilitate comparison of their magnitudes (so that the order in
which they were calculated would not matter). The nature of the
data available does not permit the use of ‘normal’ statistical
measures (such as confidence intervals) to account for un-
certainty in these values. However, for each factor, we used the
information that was available about uncertainty to propose
a plausible range of values for the multipliers. We have also
demonstrated how this uncertainty influences the estimate of
global species richness.
Differential taxonomic effort among biogeographical regions
If taxonomic effort has varied greatly among biogeographical
regions, then one would predict that understudied regions might
have fewer species than expected for their area. Korovchinsky
(1996) identified the Oriental and Ethiopian regions as having
had the poorest taxonomic work for the cladoceran branchio-
pods. To estimate the possible effect of poor sampling on known
diversity, we plotted the log species–log area relationship
among seven biogeographical regions (Nearctic, Neotropics,
Europe + Palaearctic Africa, Ethiopian + Malagasy regions,
Palaearctic Asia, Oriental region, and Australasia; see Appendix
S2 in Supplementary Material for justification of the use of these
regions). Estimates of the numbers of species presumed
‘missing’ from the Oriental and Ethiopian regions as a result of
undersampling were then obtained by adjusting their diversity
to match the species–area relationship refit using only the five
better-studied regions. The number of projected new species
was corrected based on the overall proportion of continental
endemism in the whole data set. Since branchiopods are not a
tropical group (indeed, the diversity of the planktonic groups
is regarded to be highest in temperate regions; Fernando & Paggi,
1998), we made no additional correction for elevated tropical
diversity, as would be required in many other groups. Further, we
made no corrections for regional heterogeneity in taxonomic
composition because, although present in branchiopods, it is
weak (see Appendix S3 in Supplementary Material).
The correction for differential taxonomic work among regions
was calculated as the new total number of species (including the
adjustment to the Oriental and Ethiopian richness values)
divided by the old total. A range of plausible correction factors
was obtained by taking into consideration the degree of fit of
the species–area relationship among the five best-studied bio-
geographical regions. Thus, the diversity of the two poorly studied
regions was further adjusted by first adding and then subtracting
the average of the absolute value of the residual diversities from

the five-point relationship. Note that the resulting interval is
much narrower than a 95% confidence interval would be.
Evaluation of cosmopolitanism
Incorrect assumptions of cosmopolitan (i.e. global) or multi-
continental distributions result in an underestimation of diversity.
Thus, we searched for genetic studies that shed light on the status
of supposedly conspecific populations on different continents
and determined the proportion of these morphospecies that
probably actually contain two or more species. We estimated an
overall proportion for the branchiopods by pooling our results
with a survey of detailed morphological studies on the anomopod
family Chydoridae (Frey, 1995), as our surveys are based on
different taxonomic samples.
The new estimate of branchiopod diversity was calculated by
tallying the number of species currently thought to have multi-
continental distributions and then revising diversity upwards
according to the rate of non-cosmopolitanism indicated by the
surveys. The correction multiplier was calculated as the new
diversity value divided by the previous total diversity. A range of
plausible values for this multiplier was obtained by using the
results of the two data sources separately.
Diversity within biogeographical regions: morphology and
genetics
We quantified the missing diversity at a continental scale by
comparing species richness based on traditional morphological
taxonomy with assessments of diversity based on regional genetic
investigations. We would expect these estimates to converge if the
morphology, including intra- and interpopulation variation,
were completely known. Subtle but diagnostic morphological
differences are generally found between species identified by
genetic analyses (e.g. De Melo & Hebert, 1994; Korí˜ek &
Hebert, 1996), although at least one case of a virtually cryptic
branchiopod species is known (King & Hanner, 1998). Our main
criterion for the inclusion of a genetic study was that it must
include more than one sample per named species. For genera
that have received a lot of attention (e.g. Daphnia), we selected
the most comprehensive studies and regions.
We found that most recent authors, in practice, employ a
concordance species concept (Avise & Ball, 1990), or a proxy
thereof (see below), in counting the number of branchiopod
species among their samples. For example, most use traditional
morphospecies as a starting point and then consider other
types of evidence to identify distinct species with their own
evolutionary trajectories. Here, we also adopted a concordance
species concept, as we expected that this approach would provide
the best indication of true species boundaries (i.e. where natural
breaks lie in genetic, morphological and ecological variation).
Therefore, after we had evaluated the levels of genetic variation
observed among named species, lineages that were similarly
divergent from others became candidates for species status. Then
we considered other factors, such as new morphological or
ecological characters. Thus, we outline a set of pragmatic quanti-
Global Ecology and Biogeography, 14, 455– 468 © 2005 Blackwell Publishing Ltd
Estimating true species diversity
tative criteria by which we counted species, which nearly always
agreed with the original assessments of the authors. Our pre-
ferred species criteria involved using multiple lines of con-
cordant evidence, requiring that at least two of the following
conditions be met: (1) a fixed allozyme difference; (2) a fixed
allozyme substitution at a different locus; (3) sequence diver-
gence (> 5% in mtDNA); (4) clear morphological difference; and
(5) clear habitat difference or other ecological segregation. In
cases where such data were not available, we used a proxy for the
concordance definition, based on analogy with other studies. We
treated lineages showing substantial sequence divergence (> 10%
mtDNA) as species, because in cases where there was more infor-
mation, lineages differing by this amount always were different
species. We recognized one species not conforming to the
above criteria, Daphnia arenata Hebert, 1995, because Pfrender
et al. (2000) provide evidence of a habitat shift and genetic differ-
entiation and suggest that this may represent an example of
incipient speciation.
For all studies, we calculated the ratios of the total to old
number of species. We used linear regression to evaluate the
influence of four factors on the resulting ratio: (1) (log10) sample
size of populations, with the expectation that more species will
be discovered with increased sampling; (2) geographical scale of
the study, with the expectation that more species should be
found at larger scales [three levels: (i) local, such as northern
Italy; (ii) subcontinental, such as western USA; or (iii) conti-
nental]; (3) year of the most recent taxonomic monograph for
the group, with the expectation that more recently revised groups
should have smaller ratios; and (4) higher taxonomic group
(ordinal level), with the expectation that larger-bodied groups
should be more completely described. For this analysis, the total
number of new species was the dependent ( y) variable, whereas
the number of old species was controlled for by including it in
the regression as one of the independent (x) variables.
Upon finding sample size to be the only significant predictor
of the ratio, we estimated diversity correction factors for the
branchiopods as a whole in two ways. First, we averaged the
ratios for the five best studies, judged by the number of samples
per species and the inclusion of a large number of morphospecies,
with the added criterion that they should be in different families
to avoid having biased the result towards a particular taxon.
Secondly, we used information from all studies, tallying the
total number of genetic species encountered among studies and
dividing them by the total number of morphospecies included.
We obtained a plausible range of correction factors by using the
lowest and highest ratios observed among the five best-studied
groups.
Evaluation of reliability of relative diversity estimates
We analysed the pattern of taxonomic descriptions over time
to assess how well described diversity performs as an indicator of
relative diversity among orders and families. Visual inspection of
the description date curves for branchiopods (Fig. 1) suggests
that most have a similar shape, with curves for those taxa with
quite different current diversity crossing few times during the
457
S. J. Adamowicz & A. Purvis
Figure 1 Species description curve for the Branchiopoda for (a) all species, (b) species found in Europe, (c) the four most diverse orders
separately, (d) the five most diverse families of Anostraca, and (e) the five most diverse families of Anomopoda.
history of discovery. We evaluated three decreasingly stringent
hypotheses about relative diversity in branchiopods:
(1) Is there evidence of taxonomic heterogeneity in description
over time? If not, current estimates of relative diversity could be
considered a reliable surrogate for true relative diversity in
nature. Thus, after sorting the list of description dates into
chronological order, we divided them into 12 bins of 100 descrip-
tions (the 12th bin contained only 75 species) and tested for
heterogeneity in ordinal composition among bins using a chi-
square (χ 2) test. (We repeated this analysis using temporal bins
with uneven sample sizes: bins were divided mainly into 20-year
intervals, but with the first interval containing 100 years. Alter-
native binnings did not change the results in any of the analyses
and are not reported further.) Some orders were pooled to
avoid small number problems with the χ 2 test, the groups being:
(1) Anostraca, (2) Spinicaudata, (3) other large-bodied branchi-
opods (Laevicaudata, Cyclestherida, Notostraca), (4) Anomopoda,
and (5) other Cladocera (Ctenopoda Onychopoda, Haplopoda).
We analysed familial composition within the two largest orders.
The groups of Anomopoda were: (1) Bosminidae, (2) Chydoridae
(including the former subfamilies Eurycercidae and Sayciidae),
(3) Daphniidae, (4) Ilyocryptidae and (5) Macrothricidae
(including the new species-poor families it formerly included,
Acantholeberidae, Neothricidae, and Oxyuridae). Also, we used
five bins of 110 descriptions, the 5th bin with 119 (and we
also tried 11 bins of 50 descriptions, the last bin with 59; results
were similar and are not shown). The groups of Anostraca were:
(1) Branchinectidae, (2) Branchipodidae, (3) Chirocephalidae,
(4) Streptocephalidae, (5) Thamnocephalidae, and (6) other
families (Artemiidae, Artemiopsidae, Galaziellidae, Linderi-
ellidae and Polyartemiidae); and we used six bins of 50 species
(the last with only 39 species).
458 Global Ecology and Biogeography, 14, 455–468 © 2005 Blackwell Publishing Ltd
(2) Upon finding heterogeneity and noting that this hetero-
geneity does not appear to be declining over time (see Appendix S4
in Supplementary Material), we asked: Is heterogeneity in new
descriptions essentially random over time? Or, do some groups
appear to become relatively more or less diverse over the course of
taxonomic discovery? We tested each taxonomic grouping (same
groups as in part 1) separately for trends using binomial regression
in  version 1.7.1 (Ihaka & Gentlemen, 1996). The raw data were the
proportions of descriptions in each bin belonging to that taxon.
Slopes significantly different from zero would be interpreted as
evidence that described relative diversity has changed over time,
and that current relative diversity estimates may not be stable.
(3) Finally, are diversity rankings reliable given that there is
heterogeneity in the description process? We treated our sets of
bins as samples, and resampled them 1000 times with replace-
ment. We then calculated the total diversity of each taxonomic
group in each of these replicates and determined the number of
cases where taxonomic groups exchanged places in terms of their
ranking in species richness, as well as the diversity differentials of
the taxa involved.
Undescribed diversity: what is out there?
We analysed the correlations between various branchiopod traits
and description date in order to make predictions about what
kinds of species remain to be described. We tested for correlations
with:
Body size
We hypothesize that smaller-bodied species will tend to have
been described later, as found, for example, in birds (Gaston

& Blackburn, 1994) and mammalian carnivores (Collen et al.,
2004); however, body size is a poor predictor of description date
in primates (Collen et al., 2004), North American amphibians
and reptiles (Reed & Boback, 2002), Australian scarab beetles
(Allsopp, 1997) and South American passerines (Blackburn &
Gaston, 1995).
Geographic range
We predict that widely distributed species will have been dis-
covered earlier, as this is generally the best predictor of descrip-
tion date among variables examined in past studies (e.g. Blackburn
& Gaston, 1995; Allsopp, 1997; Collen et al., 2004).
Latitudinal range
We predict that species with a wide latitudinal range will have
been discovered earlier, as they are more likely to be present in
the temperate zone. Bias towards description of temperate
species is known in e.g. insects (Gaston, 1994) and carnivores
(Collen et al., 2004).
Latitudinal centroid
Again, we predict that species whose ranges are centred in
temperate regions will have been described earlier.
Biogeographical region of occupancy
Our prediction is that species occurring in Europe will have been
described first, because of the origin of western-style taxonomy
there.
Data and analyses
Body size was measured as the (log10) maximum length (in milli-
metres) per species. Because of the lack of detailed information
on the distributions of many species, geographical range was
coded as a pseudo-continuous trait on a scale from one to seven:
(1) known from type locality only; (2) narrow endemism of a
few habitats within a local area; (3) small provincial distribution
(e.g. ‘Cape region’ of South Africa); (4) large provincial distribu-
tion (up to one-half the area of a continent); (5) continental
distribution (more than half the area of a continent); (6) inter-
continental distribution (more than one continent, with Eurasia
considered as just one continent); and (7) global distribution
(found on 5 or more continents). Latitude was also coded as
pseudo-continuous: (1) 0 –20° N or S latitude; (2) 20 –30° N or S;
(3) 30 – 40° N or S; (4) 40–50° N or S; (5) 50–60° N or S; (6) 60 –
75° N or S; and (7) 75 –90° N or S. Latitudinal range was taken as
the number of zones inhabited by each species. (Those species
with a geographical range of a ‘3’ or less were considered to occur
in a single latitudinal zone even if their true range crossed two of
the above latitudinal categories, as their latitudinal range was
restricted by their narrow total distribution.) Latitudinal
centroid was the mean of all latitudinal zones occupied and was
Global Ecology and Biogeography, 14, 455– 468 © 2005 Blackwell Publishing Ltd
Estimating true species diversity
only considered for species endemic to a single biogeographical
region. The seven biogeographical regions were coded as binary
traits, with each species being present or absent in each region.
The trait data were analysed in two ways. First, all species were
considered as independent data points. Year of description was
regressed against each (pseudo-) continuous trait individually
using . The mean description dates for biogeogeraphical regions
were compared using Tukey’s honestly significant difference
test, with only those species endemic to a single region included,
in order to compare the different regions as independent
samples. Differences among higher taxa (orders) were also com-
pared using Tukey’s test. The significance of results obtained
during single-parameter analyses was evaluated by including the
species–level traits, biogeographical regions, and higher taxa (i.e.
with order treated as a single factor with nine levels) in multiple
regression models. Secondly, because species’ traits are not statis-
tically independent (Felsenstein, 1985; Freckleton et al., 2002), a
phylogenetic comparative analysis was also performed based on
a composite high-level phylogeny. (Findings were very similar to
the taxonomic analysis, and thus the details and results are
included in Appendix S5 in Supplementary Material.)
RESULTS
Estimation of total diversity
Comparison of biogeographical regions
There were 280 branchiopod species recorded in the Nearctic,
287 in the Neotropics, 260 in Europe + North Africa, 306 in
Palaearctic Asia, 260 in the Ethiopian + Malagasy region, 185 in
the Orient and 262 in Australasia (eight occur in Antarctica and
Antarctic islands, but these were not included in the global
analysis). In a log species–log area plot for all branchiopods
(Fig. 2), the Oriental and Ethiopian regions fell below the fitted
Figure 2 The relationship between total branchiopod species
richness and land area of biogeographical regions. With seven
biogeographical regions included, the slope was 0.229 (dotted line;
P = 0.068; r 2 = 0.518). With two understudied regions excluded
(open circles), the slope was 0.119 (solid line; P = 0.069;
r 2 = 0.722).
459
S. J. Adamowicz & A. Purvis

Table 1 Evaluation of alleged cosmopolitan distributions on the basis of genetic evidence. Of the 21 morphospecies evaluated below, only 10
were confirmed to occur on more than one continent, whereas the remaining 11 should each be split into at least two species. It should be borne
in mind that the majority of these examples are from a single genus (Daphnia), whereas the intercontinental phylogeography of species in other
groups is poorly studied. Full references are listed in Appendix S6 in Supplementary Material. Abbreviations: AFR (Africa),
AUS (Australia), EUR (Europe), NA (North America) and SA (South America)

Single
Species Alleged distribution species? Notes Reference code

Order Anostraca
Artemia franciscana NA and SA Yes 1
Artemia parthenogenetica EUR, ASIA, AFR Yes A single species at least in Europe, south-western 2
Asia, and India
Order Anomopoda
Daphnia pulex Holarctic, SA No A panarctic species (in NA and northern Europe) and a 3,4,5,6
different temperate European species
Daphnia pulicaria Holarctic, SA No At least two species in the Holarctic 3,4,5,6
Daphnia middendorffiana arctic NA and EUR No One species restricted to arctic NA 3,4,5
Daphnia tenebrosa arctic EUR, ASIA, NA Yes One species with broad circumarctic distribution 3,4,5
Daphnia magna Holarctic Yes Single species at least in EUR and NA 7
Daphnia lumholtzi AUS, AFR, ASIA, Yes Single species everywhere 7
introduced to NA
Daphnia ambigua NA, SA, introduced to EUR Yes Single species everywhere 8
Daphnia similis NA, SA, EUR, AFR, ASIA No One species in NA and SA; 9,10
Old World species are different
Daphnia obtusa Holarctic, AFR, SA No Different species complexes 6,9,11
present in NA, SA and EUR
Daphnia rosea NA and EUR No Different species in EUR and NA 12
Daphnia galeata NA and EUR No Different species in EUR and NA 12
Daphnia parvula NA, SA, introduced to EUR No Different species in NA and SA 9
Daphnia curvirostris NA (arctic) and EUR Yes 6,13
(broader distribution)
Daphnia longiremis Cold NA and EUR Yes 13
Daphnia umbra Cold NA and EUR Yes 13
Daphnia laevis NA, SA, AFR No Different species in NA and SA; AFR unknown 9

Order Ctenopoda
Sida crystallina NA and EUR No Former subspecies on the two continents shown to 14
be different species
Holopedium amazonicum NA and SA No 15
Holopedium gibberum Holarctic Yes One species in Eurasia and arctic NA (an additional 15
cryptic species in NA)

line as expected, although they could not be classified as statistical
outliers. After fitting a new regression without these two regions
(log species = 0.1189*log area + 1.5808), and adjusting their
diversity to fit this line, we estimated the richness of the Orient to
be 256 species and Ethiopia to be 290 species. Given that the
overall degree of continental endemism in our entire data set was
73%, we assumed that there were an additional 52 undiscovered
species in the Orient and 22 new species in the Ethiopian region.
This would bring the global estimate for Branchiopoda to 1251
species, up from 1176. The resulting ratio, a correction factor of
1.06-fold, is an estimator of the magnitude of differential
sampling among biogeographical regions as a source of diversity
underestimation.
The mean residual magnitude in the five-point log species–
log area relationship was 0.0127. Adding and subtracting
the antilog of this value from the diversity of the Oriental and
Ethiopian regions give lower probable diversity values of 248
and 282 and upper values of 263 and 299 for the two regions,
respectively. Again, adjusting for overall continental endemism,
these values would result in 62 to 85 additional species likely
undiscovered in those two regions (i.e. because of differential
sampling among regions). These figures produce a range of
diversity correction ratios of only 1.05 to 1.07.
Evaluation of cosmopolitanism
We found 21 morphospecies with supposedly intercontinental
distributions that have been genetically studied on two or more
continents (Table 1). Only ~half (10) of these are actually
comprised of conspecific populations on different continents,

460 Global Ecology and Biogeography, 14, 455–468 © 2005 Blackwell Publishing Ltd

whereas 11 consist of at least two species each. Since Frey (1995)
found that 16 of the 21 widespread chydorids studied were prob-
ably different species on different continents, our pooled results
suggest that c. 64.3% (27 of 42) of branchiopod species thought
to have multicontinental distributions do not. Beginning with
our revised estimate of 1251 species of branchiopod, this would
include about 338 species with purported intercontinental distri-
butions (based on a 73% endemism rate for the described species).
If 64.3% of these are not actually very widespread (and assuming,
conservatively, that each should be split into only two species), an
additional 217 species need to be added to the total, bringing our
running estimate of global diversity to 1468 species. Thus, a factor
of c. 1.17-fold is necessary to account for incorrect assumptions
of intercontinental distributions. Using each set of results
separately suggests a plausible range of ratios of 1.14 to 1.21.
Diversity within continents: morphology vs. genetics
We found 16 examples of continental and regional genetic
surveys that provide information on species diversity in the
Branchiopoda (Table 2). We found that ratios of genetic:
morphological species ranged from 1.0 to 2.5, whereas the ratio
for the most comprehensively sampled genus and region (Daphnia
in North America) was 2.33-fold. Regression analysis indicated
that the number of genetic species was best explained by the
combined effect of the number of morphospecies included and
the number of populations sampled (Table 3; r 2 = 0.974;
P << 0.0001). Comparison of this model with more complex
models by  tests indicated that no additional variation was
explained by geographical scale (F5,13 = 25.24; P = 0.954), year of
the most recent taxonomic monograph (F9,13 = 0.700;
P = 0.611), or higher taxonomic group (F4,13 = 0.229; P = 0.969).
The average ratio among the five comprehensive studies
selected (see Table 2) was 1.78 genetic to original species. Pooling
across studies, the ratio was 1.58 (as 91 morphospecies were
included among studies and 144 species were recovered by
genetic surveys). Taking a mid-point ratio, we estimate that a
correction factor of c. 1.68-fold is necessary to account for the
missing diversity as a result of insufficient knowledge of vari-
ability within morphospecies. Applying this correction factor
to our running estimate results in a global diversity estimate of
c. 2452 species, more than doubling the current described diversity
of the class. Applying the variety of continental ratios among the
best studies provides a range of plausible ratios of about 1.17 to
2.50, which would produce global diversity estimates of about
1710 to 3650 species (using the middle ratio for each of the other
correction factors).
Uncertainty in the global estimate of branchiopod diversity
The middle estimates for each of the three ratios indicate that there
are c. 2.1 times more species of branchiopod in nature than
currently described. However, there is considerable uncertainty
in the estimate of c. 2450 species (Table 4). Because the ratio for
correcting for unknown diversity within continents is by far the
largest of our three correction factors, the estimate of total rich-
Global Ecology and Biogeography, 14, 455– 468 © 2005 Blackwell Publishing Ltd
Estimating true species diversity
ness is most sensitive to estimation errors in this factor. Applying
the lower and upper ratio for all factors gives a broad range of
global diversity estimates of about 1650 to 3810 species.
Analysis of relative diversity
Our analyses of the species discovery process revealed that:
(1) There was significant heterogeneity over time in the taxo-
nomic composition of new descriptions, both at the ordinal level
(12 bins; χ 2 = 197.9, d.f. = 44; P < 0.001) and familial level in
anomopods (5 bins, χ2 = 39.5, d.f. = 16; P < 0.001) and anostra-
cans (6 bins, χ2 = 59.1, d.f. = 25; P < 0.001).
(2) There was little evidence for trends over time in the relative
diversity of species described within bins. The results of quasibi-
nomial regression are presented in Table 5, as the residual errors
were overdispersed relative to the expectations for the binomial
model. While ‘other large branchiopod’ species (Notostraca,
Laevicaudata and Cyclestherida) are described less often towards
the present, and Ilyocryptidae and Thamnocephalidae appear to
be increasing in relative diversity, none of these results remained
significant following the sequential Bonferroni correction (Rice,
1989).
(3) Diversity rankings were relatively stable despite heterogeneity
in the description process. During the bin-resampling analysis at
the ordinal level, ‘other large branchiopods’ (with 51 described
species) became more diverse than ‘other Cladocera,’ which has
87 species, in only 34 of 1000 trials, whereas Spinicaudata (189
species) overtook Anostraca (289 species) in just nine trials. In all
other trials and taxa, the rankings in relative ordinal diversity
remained the same as for actual described diversity. For anomo-
pod families, Bosminidae (23 species) overtook Ilyocryptidae
(24 species) in 392 of 1000 trials, and Daphniidae (with 150
species) overtook Chydoridae (with 229 species) in just two
replicates. There was more swapping of diversity ranks among
anostracan families; all families (except ‘other families’) exchanged
places with another, with all but two pairs swapping places in
> 100 times in 1000 replicates.
Among the three data sets (orders and families of anostracans
and anomopods), swapping of ranks occurred in > 50 of 1000
trials only between pairs of taxa displaying a maximum ratio of
up to 1.525 : 1 in described diversity. Clear separation of relative
diversity occurred among all pairs of taxa having higher diversity
differentials.
Correlates of description date
Geographical and latitudinal ranges were the largest single
predictors of description date, explaining about 24 and 27% of
the variation in the taxonomic analysis, respectively (Table 5).
Body size and latitudinal centroid were also negatively related
to description date, but explained little variation in description
date (c. 2% each). All four of these factors remained significant
when attempting to simplify a multiple regression model
including endemic species (P < 0.0001), although the explana-
tory power of the model was lower than the single-predictor
models (r 2 = 0.19) because of the variation in geographical range
461
S. J. Adamowicz & A. Purvis
Table 2 Comparison of species diversity estimates within continents or smaller regions from morphological vs. genetic studies. Asterisks
designate those studies selected for calculating the first diversity correction ratio to account for incomplete knowledge of faunas within
continents (see text). Full references are listed in Appendix S7 in Supplementary Material
Taxonomic group and region
Order Anostraca
*Branchinella in Australia
Parartemia in Australia
Artemia from China
Artemia from Italy
Artemia in South America
Chirocephalus in Italy
Order Notostraca
*Lepidurus in temperate
North America
Triops in North America
Order Spinicaudata
Three genera in northern Italy
Order Anomopoda
*Bosmina in North America
*Daphnia in North America
Daphnia in Argentina
Daphnia in Australia
(including Daphniopsis species,
which are now known to be
part of Daphnia)
Daphnia in the former
Czechoslovakia
Simocephalus in North America 5
Order Ctenopoda
*Holopedium in North
and South America
Sida in North America
462
Species richness (based on Reference Species richness Reference
morphological study) (code) (after genetic study) Extent of genetic sampling (code)
12 (12 species were 1–4 14 33 samples from across 5
included in the study, much of Australia, including
but 27 are known 12 of the known species
in Australia)
8 6,7 10 17 samples from across 8
Australia, focusing on the west
including all known species
1 9 1 Nine samples 10
2 9 2 Eight samples 11
2 9 2 Nine populations from Chile, 12
Brazil, Peru, and Argentina
6 13 6 using mtDNA 22 populations (all six 14
(5 using allozymes) morphospecies)
4 15 5 18 habitats (mostly from 15
western USA)
1 16,17 2 31 populations of the 18
morphospecies Triops
longicaudatus in western USA
3 (included in study) 3 Seven samples 19
6 20 10 72 habitats (98 single-species 21,22
samples) from across much
of North America
15 23 35 More than 2000 samples 24–39
ranging from Mexico to arctic
Canada collected by Hebert
and colleagues
11 40–42 16 154 water bodies from 43 for 15 species;
across most of the country for Daphnia pulex
(Adamowicz and
Marinone,
unpublished data)
12 44–48 24 About 130 samples 49–52
(most intensively sampled in
the south-eastern and
south-western regions)
6 53 6 35 54
55 8 24 samples from across 56
North America
2 57 5 180 habitats 58 – 60
1 57 1 68 habitats 61
Global Ecology and Biogeography, 14, 455–468 © 2005 Blackwell Publishing Ltd
 Estimating true species diversity

Table 3 Regression analysis of factors influencing the total number of species in an area, as revealed by genetic studies

Factor Coefficient Standard error t-test value P-value

Intercept −0.245 2.225 −0.110 0.914

No. of morphospecies included 0.163 0.232 0.701 0.495
(log)number of populations samples 1.096 1.581 0.693 0.500
Morphospecies* (log)number of populations 0.599 0.141 4.239 0.00097***

Overall model: r 2 = 0.975; P << 0.00001.

*** highly significant (P < 0.001).

Table 4 Estimates of diversity correction factors to account for three likely sources of diversity underestimation in branchiopods. The effects of
uncertainty in different factors on our estimate of global branchiopod diversity are explored

Estimates of multipliers (and global diversity)

Source of diversity underestimation Lower Mid Upper

1. Differential taxonomic effort among major biogeographical regions 1.05 1.06 1.07
(Estimate of global diversity using mid ratio for other factors) (2429) (2452) (2475)
2. Incorrect assumption of multicontinental distributions 1.14 1.17 1.21
(Estimate of global diversity using mid ratio for other factors) (2389) (2452) (2536)
3. Incomplete knowledge of continental faunas 1.17 1.68 2.50
(Estimate of global diversity using mid ratio for other factors) (1707) (2452) (3649)
Range of overall correction (product of the three difference sources) 1.39 2.08 3.24
using lower and upper values for all three ratios
(Overall range of diversity values using lower and upper values for all ratios) (1648) (2452) (3810)

Table 5 Results of a test for trends in the taxonomic composition of new species descriptions over time. The relative diversity of higher taxa
across description bins was regressed against ordered bin number using quasi-binomial regression, and the significance of the slope was tested
using F-tests (with 1 d.f.) comparing the two-parameter with the one-parameter (intercept) model. The three P-values marked with asterisks are
significant if interpreted as individual tests, but none of these remained significant (at a total α = 0.05) after sequential Bonferroni correction
(Rice, 1989) within each of the three data sets

F-statistic Slope of
Taxon (comparing models) regression P-value

Orders
Anomopoda 0.0487 — 0.831
Other Cladocera (Ctenopoda, Onychopoda and Haplopoda) 1.375 — 0.268
Anostraca 2.546 — 0.142
Spinicaudata 1.111 — 0.317
Other large-bodied orders (Cyclestherida, Laevicaudata and Notostraca) 6.926 −0.13 0.025*
Anomopod families
Bosminidae 0.103 — 0.756
Chydoridae 0.727 — 0.416
Daphniidae 3.789 — 0.0834
Ilyocryptidae 9.128 0.147 0.0145*
Macrothricidae 0.794 — 0.396
Anostracan families
Branchinectidae 2.302 — 0.204
Branchipodidae 0.0003 — 0.986
Chirocephalidae 4.670 — 0.0968
Streptocephalidae 1.175 — 0.339
Thamnocephalidae 16.006 0.286 0.0161*
Other families 0.9793 — 0.3784

Global Ecology and Biogeography, 14, 455– 468 © 2005 Blackwell Publishing Ltd 463
S. J. Adamowicz & A. Purvis

Table 6 Relationships between several types of branchiopod traits and year of species description. Only the endemics were included in Tukey’s
test when comparing biogeographical regions so that these could be compared independently without the pull of cosmopolitan species (which
tend to be described early). These taxonomic results are based on all species being treated as independent data points; see Appendix S5 in
Supplementary Material for phylogenetic results

Trait Analysis Trait Analysis

Species trait Linear regression of single predictor variable Higher taxon Tukey’s test comparing multiple means
n Slope r2 P value Mean description date (total n = 1176)
(Log) body size 925 −15.24 0.016 0.0001*** Anomopoda (n = 560) 1930.7 55 Eight
Geographic range 1175 −14.13 0.244 < 2.2 × 10−16*** Anostraca (n = 290) 1939.6 55 5 44 significant
Latitudinal range 1172 −20.02 0.268 < 2.2 × 10−16*** Ctenopoda (n = 52) 1934.5 5 5 44 4 44 differences
Latitudinal centroid 852 −5.16 0.020 2.9 × 10−5*** Cyclestherida (n = 2) 1923.0 3 4 64 4 66 at a family-
(endemics) Haplopoda (n = 2) 1863.0 16 6 4 wise
4 6
Laevicaudata (n = 35) 1915.7 7 4 4 44 α = 5%
5 74 4 74
Notostraca (n = 15) 1879.8 7
Onychopoda (n = 33) 1896.5 6 74 7
Spinicaudata (n = 187) 1922.7 7 7

Biogeographical region Tukey’s test comparing multiple means Traits in models Multiple regression (taxonomic analyses)
Mean description date F-statistic r2 P value
(total n = 852 endemic species)
1. Species traits* 161.73,921 0.345 << 0.0001***
Nearctic 1945.7 2. Biogeographical 64.097,1168 0.278 << 0.0001***
Neotropics 1953.0 #55 regions†
Europe 1936.6 $6 4 Three significantly 3. Higher taxa (order) 6.1098,1167 0.0402 << 0.0001***
4
Palaearctic Asia 1936.3 7 different pairs at a 4. Species traits + 64.0910,914 0.412 << 0.0001***
6
Ethiopian 1932.7 family-wise α = 5% regions‡
4
Oriental 1952.2 4 5. Species + regions + 43.2418,906 0.462 << 0.0001***
Australia 1933.8 7 higher taxon§

*Body size, geographical and latitudinal ranges were included; all terms remained significant upon attempting model simplification.
†All regions were retained in the model.
‡All species traits remained in the model, but only Europe, Asia and Ethiopia remained of the biogeographical regions, upon model simplification.
§All species traits, higher taxon, Europe, Asia and Ethiopia remained in the model after model simplification.

being truncated. After omitting latitudinal centroid so that all

species could be included, the multiple model explained c. 34.5%
of the variation in description date. Although latitudinal and
geographical ranges were moderately correlated (r = 0.48), they
both remained significant predictors of description date.
Species from the Neotropics tended to have been described
more recently than those in several other regions, suggesting
that a higher proportion of new species are expected to be
described from there. Multiple regression analysis on
biogeographical regions for all species indicated that differences
among biogeographical regions explained c. 28% of the variation
in description date.
Comparison of description date among orders revealed that
anostracans and anomopods tend to be described more recently
than many other orders, whereas notostracans and onychopo-
dans tend to be described earlier than others (Table 6). Variation
in description date attributable to differences among orders was
low (c. 4%) compared to species traits or biogeographical region.
Together, species traits, regions and orders accounted for about
46% of the variation in description date, with all three sets of
predictors being significant (Table 6).
DISCUSSION
Most of what is missing is probably known
This study is the first to quantify the degree to which species
diversity is underdescribed in a group of crustaceans and to iden-
tify the geographical scales at which diversity underestimation
occurs. We estimate that there are about 2.1-fold more species in
nature than currently described and suggest that three correction
factors are required to account for the likely missing diversity in
the class Branchiopoda. A factor of only c. 1.1-fold corrects for
differential knowledge of diversity among biogeographical
regions. Similarly, a multiplier of c. 1.2-fold is sufficient to
account for incorrect assumptions about global or interconti-
nental distributions. These rather low values were unexpected
given the attention accorded to variation in taxonomic quality
among regions (Korovchinsky, 1996) and to the question of
cosmopolitanism (e.g. Frey, 1982, 1986, 1995) in branchiopods.
By contrast, the most important source of missing species, at a
factor of c. 1.7-fold, is the underestimation of diversity within
biogeographical regions. While this result is initially surprising, it

464 Global Ecology and Biogeography, 14, 455–468 © 2005 Blackwell Publishing Ltd

has been recognized that species diversity is incompletely known
even within Europe (Frey, 1989), an observation reinforced by the
shape of the species description curve for species occurring in
Europe (Fig. 1b). This high correction factor is attributed partially
to intensive surveys revealing rare species, but the most important
component of this factor was the splitting of described morpho-
species. Most species recovered by the genetic analyses had previ-
ously been pigeon-holed into known species, often ‘sensu lato’, and
thus it is clear that a proper understanding of species boundaries is
necessary to complete the quantification of branchiopod diversity.
Since only 20 branchiopod species have been formally
described so far after being identified as new by genetic analyses,
most morphospecies that require ‘splitting’ still await taxonomic
attention. While splitting will account for most new species,
there is reason to believe that there remain completely new
organisms to be discovered from the wild. Within the past five
years there have been five new genera described that were not
the result of taxonomic revision (Cammaerts & Mertens, 1999,
Chiambeng & Dumont, 1999 and Nagawana, 1999 described the
same genus; Kotov, 2000; Nagawana & Orgiljanova, 2000; Van
Damme et al., 2003), with three of these from the Lake Baikal
region and one from tropical Africa, and researchers have recently
recognized a new family of anomopod (Santos-Flores & Dodson,
2003). Our analyses of correlates of description date suggest that
species yet to be described will have small geographical and
latitudinal ranges, will tend to occur at low latitudes and will
have a higher chance of being found in the Neotropics. There is
not expected to be a trend with regard to body size, but there may
be a weak tendency for new species to belong to the most diverse
orders Anomopoda and Anostraca. Thus, further sampling from
all habitat types and from the (sub)tropics is expected to yield
many new species. For branchiopods, as for life at large, field
sampling will remain for the foreseeable future a vital component
of completing the inventory of earth’s biodiversity (May, 2004).
Characterizing the species discovery process
It is not surprising that we found evidence of heterogeneity in the
higher taxonomic composition of species descriptions over time,
given that taxonomy is conducted by specialists who publish
descriptions or monographic revisions periodically. However,
the fact that we found little evidence for trends in the composi-
tion of descriptions suggests that this heterogeneity is largely
haphazard and appears to even out over time. Moreover, we
suggest that diversity rankings based on species–richness differ-
entials of greater than 1.5-fold are robust to the erratic nature of
the description process. As diversity estimates can have large
effects on the conclusions of macroevolutionary studies (Isaac &
Purvis, 2004; Isaac et al., 2004), such a ‘rule of thumb’ may be relev-
ant for interpreting the reliability of diversity differences among
clades. Thus, while caution is required when using described
diversity data, we suggest that there is a substantial signal in such
information that can be drawn on for evolutionary research.
An unfortunate limitation of our study is that for analysing
relative diversity over time, it would have been desirable to use
the ‘then valid’ species, rather than the ‘now valid’ species, as
Global Ecology and Biogeography, 14, 455– 468 © 2005 Blackwell Publishing Ltd
Estimating true species diversity
taxonomists’ understanding of diversity in the past would have
included many species that have since been synonymized. A
proper analysis of the impact of synonymy on relative diversity
trends would have required very detailed information about all
the species names there have ever been and when all synonymies
were made (see Alroy, 2002), data that are not readily available for
the Branchiopoda. Also, our rule of thumb may be appropriate
only for taxonomic groups of intermediate ranking that share
a general habitat and many characteristics (such as size and dis-
persability). Distantly related clades in vastly different adaptive
zones, especially remote ones, may be at very different points in
the taxonomic discovery process. For example, among deep-sea
benthic crustaceans (Grassle & Maciolek, 1992), about 70% were
found to be new, suggesting that true diversity may be more
than 300% higher than described diversity. Thus, wider diver-
sity differentials are probably required in order to be confident
about diversity differences among higher taxa.
The impact of synonymy on total diversity estimates
A source of error in estimating diversity not considered here is
synonymy. The species diversity encompassed by currently valid
names can be overestimated because of the fact that some species
names are destined to become synonymized (Alroy, 2002).
Within the branchiopods, this problem is particularly severe in
the conchostracans, whose taxonomy is well-known to be
especially poor (as noted by Straßkraba, 1965a,b, 1966; Timms &
Richter, 2002). For example, of the 59 species of spinicaudatan
described by Daday in the early 1900s, 22 have been syn-
onymized since, eight with Cyzicus tetracerus alone (from Brtek,
1997). However, many other species have never been studied
again and are only known from their type locality. One has to
wonder how many more are destined to be lost once a thorough
revision including new collections is complete. Conversely, many
names that have been synonymized are now being resurrected on
the basis of new, especially genetic, evidence (e.g. Triops newberryi,
see King & Hanner, 1998 and Daphnia magniceps, see Taylor
et al., 1998). Moreover, species that morphologists have noted to
be unduly variable, and which tend to have many varieties and a
complex history of synonymy, are found actually to be species
complexes (e.g. Taylor et al., 1996; Sassaman et al., 1997; King &
Hanner, 1998).
Overall, for the branchiopods, it seems that the problem of
overestimating diversity because of synonymy is overshadowed
by the underestimation of diversity due to incomplete sam-
pling of variability within species. Most genetic studies increase
the number of species (see Table 2) and do not favour new
synonymies. The only exceptions we have encountered are
the proposed mergers of Artemia salina + Artemia tunisiana
(Triantaphyllidis et al., 1997) and Daphnia spinulata + Daphnia
notacantha (Adamowicz et al. 2004). However, while some
synonymizing of described species will be necessary during
taxonomic revision of the Branchiopoda, we suspect that this
factor will have relatively minor impact on global diversity
compared to the proliferation of species required by the results of
detailed genetic and morphological surveys.
465
S. J. Adamowicz & A. Purvis
How many species are there in the world?
Our estimate for branchiopods of 2.1-fold more species in nature
than currently described is comparable to other recent estimates
for arthropods, with most authors predicting about a 2- to 5-fold
increase over the described diversity of approximately one million
insect species (e.g. Hodkinson & Casson, 1991; Hammond,
1992; Ødegaard, 2000; Dolphin & Quicke, 2001). Although we
attempted to estimate ‘true’ species diversity while most other
studies only estimate morphospecies richness, our increase
factor was on the low end compared to the range for insects. This
probably reflects the observation that branchiopods are most
diverse in temperate regions, which have been relatively well-
studied, whereas the majority of terrestrial taxa achieve their
highest diversity in the tropics (Gaston, 2000). Moreover, the
narrower distributions and difficulty of collecting many
terrestrial insects would suggest that they are at an earlier stage
in the discovery process. Another consideration in comparing
diversity–increase factors is the methodology used to obtain
them. Our value is perhaps among the most realistic estimates
for global diversity of a particular taxonomic group, as we
examined several sources of underestimation and based conclu-
sions on data from this group only.
This is one of the first studies of global diversity in an inverteb-
rate group to attempt to incorporate information about real
species boundaries in nature. Estimates for other groups based
on statistical analyses on sampled morphospecies may have to be
revised somewhat upwards to account for this source of under-
estimation, the magnitude of the adjustment depending on the
degree of concordance between genetic vs. current taxonomic
species in different groups of organisms. In some cases, the
differences among groups may be large, e.g. a ratio of only about
1.5 : 1 in planktonic foraminiferans (Kucera & Darling, 2002),
perhaps a several-fold to one ratio in nematodes (from Floyd et al.,
2002; Blaxter, 2004), and who-knows-what: whatever-one-is in
bacteria (at least in the range of 100 –1000 : 1, maybe more; see
Oren, 2004 and references therein). In others groups, such as
some lepidopterans, the agreement may be nearly 1 : 1 (Hebert
et al., 2003). Our value of 1.7 : 1, based on a sample of c. 8% of
described branchiopod species, may be a typical sort of figure for
smallish crustaceans. A value of 1.55 : 1 was recently found in
a group of marine copepods, based on a study of 20 morpho-
species and assuming here a sequence divergence cut-off point
similar to that among morphospecies (Goetze, 2003).
Such large differences among taxonomic groups reflect the
degree to which variation among species is accessible to humans.
While differences in chemical cues, mating behaviour, habitat
segregation or subtle morphological features may be very ‘real’
delimiters of species to the individuals themselves (Knowlton,
1993), such traits are not easily amenable to traditional
taxonomic analysis. Combined molecular and morphological
approaches can be instrumental in identifying the kinds of varia-
tion and traits that do separate species in nature. While some
taxonomists view approaches such as ‘DNA taxonomy’ (Tautz
et al., 2003; Blaxter, 2004) or ‘DNA barcoding’ (Hebert et al.,
2003) to be of limited merit (e.g. Lipscomb et al., 2003; Wheeler,
466 Global Ecology and Biogeography, 14, 455–468 © 2005 Blackwell Publishing Ltd
2004), genetic investigations enable us to ‘see’ beyond what we
can perceive and allow us the possibility of viewing the limits of
biological species as experienced by the organisms themselves.
Characterizing the meaningful variation in terms of functional
morphology, ecology, reproduction and evolutionary history
will take much longer, of course, but delimiting the species is a
vital first step. Due to the large variation in the ratio of perceived
to actual species among higher taxa, we suggest that getting this
ratio approximately right for all major groups will be a key step
towards accurately estimating global species diversity.
ACKNOWLEDGEMENTS
We thank Tim Barraclough, Ryan Gregory, Armand Leroi,
Donald Quicke, and two reviewers for helpful comments on the
manuscript. We are grateful to the Natural History Museum in
London for access to their library. Funding for this project
was provided by a Natural Sciences and Engineering Research
Council of Canada postgraduate scholarship, an overseas research
scholarship (UK) and a Beit Fellowship to SJA.
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SUPPLEMENTARY MATERIAL
The following material is available online at
www.blackwell-synergy.com/loi/geb
Appendix S1 Details regarding the assembly of the taxonomic
database for Branchiopoda
Appendix S2 Explanation for biogeographical regions employed
Appendix S3 Assessment of heterogeneity of taxonomic com-
position among biogeographical regions
Appendix S4 Assessment of the question: is heterogeneity in the
taxonomic composition of descriptions declining towards the
present?
Appendix S5 Correlates of description date, based on analysis of
phylogenetically independent contrasts
Appendix S6 References for Table 1
Appendix S7 References for Table 2
BIOSKETCHES
Sarah Adamowicz is interested in the evolution of
biodiversity. Currently a PhD student at the Imperial
College London, she is investigating evolutionary trends
and correlates of species richness in the Crustacea.
Andy Purvis is Professor in Biodiversity in the Division
of Biology at the Imperial College London. He takes a
phylogenetic approach to a range of questions in
macroevolution and conservation biology. Among his
current projects is coediting a symposium volume entitled
Phylogeny and Conservation and writing a book on
macroevolution.