Cladistics
Cladistics 24 (2008) 708–722
10.1111/j.1096-0031.2008.00202.x

Family ties: molecular phylogeny of crab spiders
(Araneae: Thomisidae)
Suresh P. Benjamina,b* , Dimitar Dimitrova, Rosemary G. Gillespieb and Gustavo
Hormigaa
a

The George Washington University, Department of Biological Sciences, 2023 G Street NW, Washington DC, 20052, USA; bUniversity of California,
Berkeley, Insect Biology Division—ESPM, 201 Wellman Hall 3112, Berkeley, CA 94720-3112, USA
Accepted 2 November 2007

Abstract
The first quantitative phylogenetic analysis of three sequenced genes (16S rRNA, cytochrome c oxidase subunit I, histone 3) of 25
genera of crab spiders and 11 outgroups supports the monophyly of Thomisidae. Four lineages within Thomisidae are recovered.
They are informally named here as the Borboropactus clade, Epidius clade, Stephanopis clade and the Thomisus clade, pending
detailed morphology based cladistic work. The Thomisus clade is recovered as a strongly supported monophyletic group with a
minimal genetic divergence. Philodromidae previously widely considered a subfamily of Thomisidae do not group within thomisids
and is excluded from Thomisidae. However, Aphantochilinae previously generally considered as a separate family falls within the
Thomisus clade and is included in Thomisidae. The recently proposed new family Borboropactidae is rejected, as it is paraphyletic.
Ó The Willi Hennig Society 2008.

Thomisidae Sundevall, 1833, commonly called crab
spiders are often cryptically colored sit-and-wait predators that do not build capture webs (Fig. 1). Thomisidae
is the sixth largest spider family. It includes 2062
described species in 171 genera (Platnick, 2007) with
many more species remaining to be described. They are
mainly active during the day and ambush insects with
their well-adapted first and second legs (Homann, 1934;

Comstock, 1948). Not surprisingly, they are an important component of terrestrial ecosystems (Riechert,
1974). As predators of agricultural pests, thomisids play
an important part in natural pest control (Riechert and
Lockley, 1984; Nyffeler and Benz, 1987; Uetz et al.,
1999).
Some thomisids (e.g., Misumena, Diaea, Runcinia and
Thomisus) possess the ability to change color and blend
into their habitat, in most cases flowers (Packard, 1905;
*Corresponding author:
E-mail address:
 Present address: Department of Entomology, National Museum of
Natural History NHB 105, PO Box 37012, Smithsonian Institution,
Washington, DC 20013-7012, USA.
Ó The Willi Hennig Society 2008

Gabritschevsky, 1927; Comstock, 1948). Misumena vatia
(Clerck, 1757) has a remarkable ability to change color,
which takes place during migration to flowers of
different color from spring to the early part of summer
(Comstock, 1948). Crab spiders are attracted by fragrance components of flowers (Aldrich and Barros,
1995; Krell and Kraemer, 1998) and use visual and
tactile cues for selecting flowers (Morse, 1988; Greco
and Kevan, 1994). They reach their ambush sites in a
step-by-step process using several draglines and ballooning events (Homann, 1934).
There are social crab spiders with maternal care in the
Eucalyptus forest of Australia (Main, 1988; Evans,
1995). Mother Diaea ergandros Evans (1995), catch
larger prey for their own offspring, but not for the
adopted offspring leading to large size and possibly
better survival rates for the natural offspring (Evans,

1998). Mothers further increase survival of natural
offspring by producing trophic oocytes used in a system
of sacrificial care (Evans, 1998).
Myrmecomorphism is known in a number of thomisids: Strigoplus albostriatus Simon, 1885, Amyciaea
forticeps (O.P.-Cambridge, 1873), A. lineatipes


S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

A

C

709

B

D

Fig. 1. Living thomisids sampled in this study. (A) ‘‘Monaeses’’ sp. A, Sri Lanka, Central Province; (B) Diaea placata, Sri Lanka, Western Province
(not sampled); (C) Oxytate taprobane, Sri Lanka, Central Province; (D) ‘‘Lysiteles’’ sp. B, Sri Lanka, Central Province. All photos by SPB.

O.P.-Cambridge, 1901 and Aphantochilus rogersi
O.P.-Cambridge, 1870, are known to be ant mimics.
A. forticeps is of the same color as the ant Oecophylla
smaragdina and bears on the posterior part of the
abdomen a pair of eye-like spots that correspond to eyes
of the ant (Shelford, 1902). The similarity of A. rogersii
to some spiny South American ants is striking, and
forms a further instance of myrmecomorphism (Oliveira

and Sazima, 1984).
Given their ecological significance and appealing
adaptations one would expect to see a plethora of
phylogenetic studies. However, no such studies of
thomisids exist. Moreover, understanding the exact
taxonomic limits of this large family has always been
problematic. Thomisidae was proposed to accommodate spiders with legs generally extended sideways
(laterigrade), instead of being oriented towards the front
or back as in most other spiders. Originally all spiders
with laterigrade legs such as Sparassidae and Philodromidae were included. Simon (1892) was the first to
propose a hypothesis of generic groups for all thomisid
genera recognized during his time. Although his
‘‘groups’’ were neither evolutionary nor phylogenetic
in the modern sense, he provided some information on
Thomisidae morphology and arguments supporting his
ideas. His Stephanopsinae contained spiders with cheliceral teeth; Aphantochilinae and Strophiinae contained species with modifications such as elongated
maxillae related to their ant mimicking habits; Stiphropodinae included species with an enlarged tarsus; spiders

that did not fit into the above categories were included
in Misumeninae and Philodrominae. Species groups
were then formed within these subfamilies based on eye
pattern and shape of prosoma.
Since Simon (1892) the understanding of generic
relationships has not changed greatly. Holm (1940), on
the grounds of embryological studies, and Homann
(1975), on the grounds of eye morphology, excluded
Philodromids from Thomisidae. Philodromids were
later given family status (Ono, 1988), which has been
accepted since (but see Roberts, 1995). Separately,
family status was proposed for some thomisids, Stephanopidae (Pickard-Cambridge, 1871) and Aphantochilidae (Thorell, 1873). Levi (1982) placed the Thomisidae

in a superfamily along with Aphantochilidae, which was
not accepted by Ono (1988). In the most current study
Thomisidae was separated into seven subfamilies (Ono,
1988): Stephanopinae, Thomisinae, Bominae, Stiphropodinae, Dietinae, Strophiinae and Aphantochilinae
using characters proposed by Simon (1892). Recently
(Wunderlich, 2004b) proposed a new family ‘‘Borboropactidae’’ to include parts of Thomisidae. Thus, to date,
the monophyly of Thomisidae remains untested.
A great part of our knowledge of evolutionary history
is derived from phylogenies, reconstructed by sampling
and grouping characters. Harvey and Pagel (1991)
illustrated the richness of evolutionary questions that
can be approached with phylogenies. Our current
understanding of relationships within Thomisidae is
largely based on Simon (1892) and modifications to his


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S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

ideas by different workers (Schick, 1965; Homann, 1975;
Ono, 1988). However, they do not provide much
information, partly because the family has never been
subject to quantitative phylogenetic analysis. Recent
taxonomic work on tropical Asian crab spiders (Tikader,
1980; Barrion and Litsinger, 1995) added more confusion, illustrating the fragile systematic state of the
family.
Here we present the first cladistic analysis of Thomisidae. This analysis of molecular data aims to test the
monophyly of Thomisidae, including the validity of
‘‘Borboropactidae’’ and the placement of some enigmatic taxa such as Epidius and Cebrenninus, which share

characters such as the presence of a conductor and
median apophysis with ‘‘Borboropactidae’’. Epidius was
provisionally placed in Thomisidae owing to its unusual
male palp (Benjamin, 2000). Owing to the considerable
volatility in thomisid systematics and paucity of morphological information for a large number of taxa, we
do not formally name any higher-level taxa here.
However, we provide putative morphological synapomorphies for most of the largest clades. The first author
is presently undertaking a detailed morphological revision of Thomisidae.

Materials and methods
Ingroup taxa
We sampled 41 ingroup and 13 outgroup taxa for
DNA sequencing (Table 1). Representatives of all major
thomisid subfamilies except for Stiphropodinae and
Strophiinae are included. They are: Stephanopinae,
Thomisinae, Bominae, Dietinae and Aphantochilinae.
Additionally, several sequences of thomisids and outgroups available in GenBank were added to the analyses. Accession codes for all sequences are given in
Table 1.

(P. Lehtinen, pers. comm.). We have included six species
of Salticidae, one species each of Miturgidae and
Corinnidae in our study. As Philodromidae have been
in the past included as a subfamily of Thomisidae, we
include three philodromid taxa in our study.
DNA sequencing and alignment
Genomic DNA was extracted from either fresh or
ETOH-preserved leg tissue using the Qiagen DNeasy
Tissue Kits (Qiagen, Valencia, CA, USA). Otherwise
intact spiders, preserved in alcohol have been deposited
as voucher specimens (Table 1). Partial fragments of the

mitochondrial genes cytochrome c oxidase subunit I
(COI) and 16S rRNA (16S) and the nuclear gene histone
H3 (H3) were amplified using the following primer pairs:
(COI) C1-J-1751 and C1-N-2191 (Simon et al., 1994)
(16S) LR-N-13398 (Simon et al., 1994) and LR-J-12864
(Arnedo et al., 2004) and (H3) H3aF and H3aR (Colgan
et al., 1998). PCR products were purified using the
QIAquick PCR Purification Kit (Qiagen) and sequenced
directly in both directions using an ABI 3730 automated
sequencer (Applied Biosystems, Foster City, CA, USA),
in combination with the ABI PRISM Big Dye Terminator Cycle Sequencing Ready Reaction Kit. The
chromatograms produced for each DNA sequence were
edited in Sequencher 3.1 (Gene Codes, Ann Arbor, MI,
USA).
The protein-coding COI and H3 sequences were easily
aligned unambiguously. For non-protein-coding sequences there are two basic approaches to multiple
sequences alignment. The first are the static alignment
methods with fixed homology statements, in which the
alignment is independent of the phylogenetic analysis.
The second is the optimization alignment method where
alignment is an integral step of phylogenetic analysis
(Wheeler, 1996). We explored our data using both
analytical philosophies. In all analyses gaps were treated
either as missing data (parsimony and Bayesian inference) or as a fifth state (POY).

Outgroup taxa
Direct optimization
Thomisids fall within the large clade Dionycha
(Coddington and Levi, 1991), which are characterized
by the loss of the unpaired tarsal claw. The monophyly

of Dionycha has, however, been elusive. Furthermore,
the phylogenetic structure within Dionycha has not been
fully explored [however, see Davila (2003) who included
a small sample of Dionycha in her study of Ctenidae
interrelationships]. Thus, the only character-based outgroup hypothesis for thomisids is by Loerbroks (1984);
in a functional study of the male palp of M. vatia, he
found characters to relate Salticidae to Thomisidae.
Morphological comparisons of representatives of families within Dionycha have suggested that Creugas
(Corinnidae) might be the sister to thomisids

Direct optimization (Wheeler, 1996; Wheeler et al.,
2006) was implemented with the software package POY
Version 3.0.11. Unlike analysis of statically aligned data,
in POY alignments are tree dependent and as a result the
homology statements are dynamic. In all the analyses
the protein-coding genes H3 and COI were treated as
prealigned. This avoids the use of in ⁄ del transformations
for these fragments and saves computing time without
alteration of results (Giribet, 2001). To further reduce
computational time and to minimize the probability of
unwanted artifacts in the alignment the 16S was
divided into two homologous fragments (Giribet,
2001). Homolog partitions in 16S were defined based


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S. P. Benjamin et al. / Cladistics 24 (2008) 708–722
Table 1
Specimens sequenced for molecular analyses. Fragments successfully sequenced and GenBank accession number are given

Family

Genus

Salticidae

Corinnidae
Miturgidae
Thomisidae

Species

Country

Locality

COI

16S

EU168142 ⁄
EU168143
Proernus
stigmaticus USA
Hawaii, HI
EU168156 EU168144
Philodromus
sp.
USA
Berkeley, CA

EU168157 EU168145
Onomastus
sp. A
Sri Lanka
Central Province
EU168158 EU168146
Onomastus
sp. B
Sri Lanka
Central Province
EU168159 EU168148
Portia
labiata
NA
NA
AY297361 AY296653
Lyssomanes
virides
NA
NA
AY297360 AY296652
Neon
nelli
NA
NA
AF327988 AF327959
Helvetia
cf. zonata
NA
NA

AY297394 AY296685
Castianeira
sp.
NA
NA
AY297419 AY296710
Cheiracanthium sp.
NA
NA
AY297421 NA
Borboropactus
cinerascens Singapore
Bukit Timah NR
NA
EU168138
Borboropactus
sp.
South Africa Limpopo
EU168187 EU168151
Epidius
parvati
Sri Lanka
Western Province
EU168163 EU168141
Cebrenninus
rugosus
Thailand
Chumphon Province
EU168175 EU168139
Cebrenninus

rugosus
Malaysia
Kelantan
EU168177 EU168140
‘‘Epicadinus’’
sp.
Argentina
Misiones
NA
EU168153
‘‘Stephanopis’’
sp. A
Chile
Region IX
EU168167 EU168137
‘‘Stephanopis’’
sp. B
Australia
Tasmania
EU168185 NA
‘‘Stephanopis’’
sp. C
Australia
Tasmania
NA
NA
‘‘Stephanopis’’
sp. D
Australia
Western Australia

NA
NA
Pseudoporrhopis granum
Madagascar Fianarantsoa
EU168170 EU168131
Oxytate
taprobane
Sri Lanka
Central Province
EU168161 EU168133
Amyciaea
forticeps
Thailand
Montha Tarn
NA
NA
‘‘Lysiteles’’
sp. A
Sri Lanka
Central Province
EU168184 EU168128
‘‘Lysiteles’’
sp. B
Sri Lanka
Central Province
EU168183 EU168129
Coriarachne
versicolor
USA
VA

NA
NA
Xysticus
californicus USA
CA
EU168181 EU168115
Xysticus
sp.
USA
CA
EU168182 EU168116
‘‘Monaeses’’
sp. A
Sri Lanka
Central Province
EU168172 EU168117
‘‘Monaeses’’
sp. B
Australia
Western Australia
EU168186 EU168150
Tmarus
angulatus
USA
CA
EU168179 EU168120
Tmarus
angulatus
USA
CA

EU168180 EU168121
Talaus
sp.
Vietnam
Kien Gian Province
NA
EU168135
Runcinia
albostriata Sri Lanka
Central Province
EU168178 EU168125
Runcinia
albostriata Myanmar
Bago Division
EU168165 EU168124
Runcinia
acuminata
Myanmar
Bago Division
EU168166 EU168126
Diaea
subdola
Sri Lanka
Central Province
EU168174 EU168118
Aphantochilus
sp.
Argentina
Misiones
NA

EU168152
Diaea
nr subdola
Myanmar
Yangon Division
EU168169 EU168127
Cyriogonus
sp.
Madagascar Antananarivo
EU168168 EU168130
Western Province
EU168162 EU168122
Thomisus
granulifrons Sri Lanka
Thomisus
sp. A
Sri Lanka
Sabaragamuwa Province EU168164 NA
Thomisus
sp. B
Thailand
Phuket Province
EU168176 EU168123
Thomisops
piger
Sri Lanka
Western Province
EU168171 EU168134
Haplotamarus
sp.

Sri Lanka
Western Province
EU168173 EU168132
Camaricus
sp.
Sri Lanka
Central Province
NA
EU168119
Boliscus
sp.
Malaysia
Kelantan
NA
EU168136
Mecaphesa a
semispinos
NA
NA
DQ174382 DQ174339
Mecaphesa
naevigera
NA
NA
DQ174344 DQ174387
Xysticus
sp. B
NA
NA
AY297423 AY296714


Philodromidea Pagiopalus

nigriventris

USA

Maui, HI

on a preliminary static multiple alignment with
ClustalW (Thompson et al., 1994) using the default
parameters.
Sensitivity of the results to various parameters was
explored using five different values of gap opening
costs. As an objective criterion to select among
different parameter combinations, maximum congru-

EU168155

H3

Depository

EU157106 USNM
EU157107
EU157108
NA
EU157109
NA
NA

NA
NA
NA
NA
EU157126
NA
EU157114
EU157134
NA
NA
EU157117
EU157138
EU157137
EU157139
EU157120
EU157112
EU157135
EU157133
NA
EU157136
EU157131
EU157132
EU157122
NA
EU157110
EU157111
EU157127
EU157130
EU157116
NA

EU157124
EU157140
EU157119
EU157118
EU157113
EU157115
EU157129
EU157121
EU157123
EU157125
EU157128
NA
NA
NA

USNM
USNM
USNM
USNM
NA
NA
NA
NA
NA
NA
MHNG
MHNG
MHNG
MHNG
MHNG

MACN-Ar
CAS
USNM
USNM
USNM
CAS
MHNG
MHNG
MHNG
MHNG
USNM
USNM
USNM
MHNG
CAS
USNM
USNM
MHNG
MHNG
CAS
CAS
MHNG
MACN-Ar
CAS
CAS
MHNG
MHNG
MHNG
MHNG
MHNG

MHNG
MHNG
NA
NA
NA

ence between the 16S and the COI + H3 partitions
was used (Wheeler and Hayashi, 1998). Congruence
was measured using the incongruence length difference
(ILD) test (Mickevich and Farris, 1981). Tables 2 and
3 contain the information about the parameter combinations investigated together with statistics from the
analyses. All POY searches were run on a


712

S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

Table 2
Statistics resulting from the exploration of the effect of a range of gap
opening ⁄ gap extension cost on the data set used to generate the tree
shown in Fig. 1
Gap
cost

16S
length

COI + H3
length


Combined
length

ILD

1
2
3
4
5

1362
1489
1569
1642
1709

1834
1834
1834
1834
1834

3279
3414
3501
3578
3649


0.025312595
0.02665495
0.027992002
0.028507546
0.029049055

Table 3
Statistics resulting from the exploration of the effect of a range of gap
opening ⁄ gap extension cost on the data set used to generate the tree
shown in Fig. 2

matrices (elision matrix; Wheeler et al., 1995; Hedin and
Maddison, 2001). In principle, an elision matrix should
generally recover relationships that are robust to alignment differences. This was done as suggested by Hedin
and Maddison (2001).
Topological congruence between the results from the
analysis of each particular cost scheme and the elision
matrix was measured by the number of nodes in
common between the consensus tree, which they produced and the average symmetric-difference distances
between all trees from the individual and the elision
matrices (Hedin and Maddison, 2001; Swofford, 2002).
Based on these criteria the alignment built with gap
opening ⁄ gap extension cost of 20 ⁄ 2 was selected and
used in all further analyses.
Phylogenetic analysis of static alignments

Gap
cost

16S

length

COI + H3
length

Combined
length

ILD

1
2
3
4
5

1423
1539
1591
1631
1671

1772
1772
1772
1772
1772

1774
1798

1807
1809
1828

0.053893989
0.019543974
0.027191206
0.035430839
0.041481069

computer cluster at The George Washington
University.
Initial trees were built using an approximation shortcut (command –approxbuild). Thirty replicates of
random addition of sequences were performed (command –replicates 30) holding up to 300 trees
(command –holdmaxtrees 300). Tree fusing and
tree drifting (TBR) strategies were used for tree searching (commands –tbr –treefuse –drifttbr). In
order to increase the search efficiency and guarantee that
POY will save only disparate trees the command
-fitchtrees was used. To reduce error from heuristic
operations suboptimal trees with 1% length distance
from the shortest trees were submitted to a further
round of TBR (command –checkslop 10). Indices
and topology specific implied alignments were saved
using the commands –indices and –implied
alignment.
Static alignments
Static alignments were built using ClustalW (Thompson et al., 1994) with the following gap opening ⁄ gap
extension costs: 8 ⁄ 2, 8 ⁄ 4, 15 ⁄ 6.66 (default), 20 ⁄ 2, 24 ⁄ 4
and 24 ⁄ 6 and transition weight fixed to 0.5. Owing to the
lack of an a priori basis for deciding on one alignment

over the other and to avoid having to use all six
alignments, we adopted phylogenetic congruence to
select an ‘‘optimal’’ alignment. Phylogenetic congruence
works by comparing phylogenetic results from each
alignment matrix with results of a concatenation of all

Both parsimony and likelihood methods were used as
optimality criteria for the phylogenetic analyses. Bayesian inference was preferred over maximum likelihood
because performing Bayesian analysis is computationally
less demanding than traditional likelihood methods.
Manipulations of data matrices and trees were performed
with MacClade (Maddison and Maddison, 2001),
PAUP* (Swofford, 2002) and WinClada (Nixon, 2002).
Parsimony
Parsimony searches were conducted using TNT versus
1.0 (Goloboff et al, 2003a). All multistate characters
were treated as non-additive (Fitch, 1971). In all
parsimony analyses, heuristic searches under the ‘‘traditional search’’ option were performed using 1000
replicates, holding 10 trees per replicate to a maximum
of 10 000 trees. Clade support was assessed by means of
non-parametric bootstrap analysis (Felsenstein, 1985) as
implemented in TNT with 1000 pseudoreplicates of
heuristic searches with 100 interactions of random
addition of taxa and holding 10 trees per interaction.
The same parameters were used to perform a jackknife
analysis (Farris et al., 1996). Additionally, Poisson
weighting bootstrap and symmetric re-sampling support
values (Goloboff et al., 2003b) were calculated using the
same search parameters. These methods try to avoid
errors where high support values are found for groups

not supported by the original data (Goloboff et al.,
2003b). While bootstrap and Poisson weighting bootstrap re-sample characters by building a matrix of the
same size as the original, jackknife and symmetric resampling removes part of the characters, constructing a
smaller matrix where character replacement does not
occur (Goloboff et al., 2003b). In both cases the
analyses were performed using the default setting in
TNT (probability of character removal in the pseudoreplicates was set to 0.36).


S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

In addition to the parsimony analysis with equal
character weights, analyses using implied weighting were
conducted in TNT. Implied weighting was used to build
trees using differential character weighting (Goloboff,
1993a). It is superior to the successive weighting as it
provides an optimality criterion (maximum fit) to build
trees and weight characters (Goloboff, 1993b). This
helps to avoid iterative searches from preliminary trees,
which were one of the most criticized aspects of the
successive weighting method [Swofford and Olsen
(1991), but see Steel (1994) and Farris (2001)]. Fit is
calculated as a function in which concavity depends on
the constant K. With the increase of K the penalization
against homoplasy decreases (Goloboff, 1993b). Implied
weighting analyses, for K-values from 1 to 100 were
done in TNT for 500 replicates.
Bayesian inference (likelihood approach)
MrBayes 3.4 (Ronquist and Huelsenbeck, 2003) was
used to estimate topologies and to calculate posterior

probabilities of inferred clades. The best-fit model of
sequence evolution was selected using Modeltest 3.6
using the Akaike information criterion (Posada and
Crandall, 1998). Four replicate analyses were run
simultaneously from random starting trees using default
priors. Metropolis-coupled MCMC (one cold and three
heated) were run for 10 million generations. Topologies
were sampled at intervals of 1000 generations within
each chain. Average likelihood scores (–ln) were examined to assess convergence. Sampled trees were plotted
against generations in order to determine point of
stability. Trees under the stability value were discarded
as ‘‘burn-in’’. There is some disagreement on the
reliability of the posterior probability, which has been
criticized by Suzuki et al. (2002) for overestimating
statistical confidence when concatenated gene sequences
are used. However, others have countered: Wilcox et al.
(2002) based on simulations have shown that Bayesian
support values represent much better estimates of
phylogenetic accuracy than do non-parametric bootstrap support values. However, when model misspecifications are present (which is absent in simulation-based
studies) inflation of posterior probabilities may occur.

Results
The lengths of the sequenced fragments excluding the
primers were as follows: 16S, 430 bp; H3, 328 bp; and
COI, 557 bp. All sequences have been deposited in
GenBank and their acquisition numbers are listed
in Table 1. The ILD test (Farris et al., 1994), done in
NONA, did not find significant incongruence between
the three gene fragments. Thus, all three gene fragments
were combined into a single matrix.


713

Direct optimization
The results of the analysis with the combination of
gap opening ⁄ gap extension cost of 2 ⁄ 1 produced implied
alignments with maximum congruence among partitions
(Tables 2 and 3). One of the two most parsimonious
trees (L ¼ 3377) shown in Fig. 2, recovers a monophyletic Thomisidae. The strict consensus of the two most
parsimonious trees is given in Fig. 3. Thomisids are
rendered possibly paraphyletic in the strict consensus, as
it unites Cesonia sp. (Gnaphosidae) and Hibana sp.
(Anyphaenidae) in a basal tricotomy with ingroup taxa
(Fig. 3). However, this arrangement of Cesonia sp. and
Hibana sp. within thomisids is not supported and is very
sensitive to taxon sampling. Adding sequence fragments
of four taxa (one Misumenoides, two Mecaphesa and one
Xysticus species; see Table 1) and ⁄ or excluding the
outgroup taxon Portia labiata (Thorell, 1887) recovers
a monophyletic Thomisidae, which excludes Cesonia
and Hibana (Fig. 4). The analysis of this modified
matrix found six most parsimonious trees (L ¼ 3279).
The combination of gap opening ⁄ gap extension cost of
1 ⁄ 1 resulted in higher overall congruence for this matrix.
The strict consensus of this analysis is shown in Fig. 4.
In all analyses under dynamic optimization Anyphaenidae (represented by Hibana) plus Gnaphosidae (represented by Cesonia) is recovered as the closest relative of
Thomisidae.
Parsimony analyses with static homology
Analyses of the static alignment under equal weights
found a single most parsimonious tree (L ¼ 3234, CI ¼

0.32, RI ¼ 0.477) shown in Fig. 5. Thomisids form a
monophyletic family. The main lineages within Thomisidae and their relationships are the same as in the trees
resulting from dynamic optimization. However, only the
Epidius clade and to a lesser extent the Thomisus clade
are well supported. Hibana sp., but not Cesonia sp.
appears as closest relative to Thomisidae.
Parsimony analyses under the criterion of implied
weights constantly recover the monophyly of Thomisidae. Relationships within Thomisidae are as in uweighted parsimony and dynamic optimization analyses
supporting the existence of the same four main lineages.
The only difference refers to the composition of the
Stephanopis clade, which was always recovered as
monophyletic, but for the majority of the concavities it
also included Epicadinus sp. (Fig. 6).
Bayesian inference
The best fitting model for the fragment of 16S was
GTR + I + C; for H3 it was HKY + I + C and for
COI it was GTR + I + C. The results of the first
150 000 generations were discarded as burn-in. The


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S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

Fig. 2. One of the two most parsimonious trees (L ¼ 3377) found under direct optimization. recovers a monophyletic Thomisidae. Gap opening ⁄ gap
extension cost of 2 ⁄ 1. Jackknife values greater than 60 are shown above the branches.

resulting tree is shown in Fig. 7. Results practically
mirror those from the implied weight parsimony analyses and are highly congruent with the tree topologies
found by the unweighted parsimony and direct optimization analyses. Thomisidae is again monophyletic and

includes the same four main lineages as in all previous
analyses. The Borboropactus clade is sister to all other
thomisids. Support for thomisid monophyly is higher
than in other analyses. Most of the main groups within
Thomisidae and their relationships are also well supported. Further, the Bayesian inference results suggest
that Anyphaenidae (represented by Hibana sp.) is the
sister lineage of the family Thomisidae.

Discussion
Monophyly of Thomisidae
The taxonomy of Thomisidae is challenging. Almost
all genera await revision. Despite past attempts (Simon,
1892; Petrunkevitch, 1928; Roewer, 1954; Ono, 1988;
Lehtinen, 2005), the monophyly of Thomisidae was
never established. Our study, based on molecular data
provides strong support for the monophyly of Thomisidae for the first time. Monophyly of Thomisidae is
supported by three molecular synapomorphies in this
study, all from the 16S gene fragment. They are three


S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

715

Fig. 3. The strict consensus of the two most parsimonious trees found under direct optimization. Gap opening ⁄ gap extension cost of 2 ⁄ 1. Jackknife
values greater than 60 are shown above the branches.

changes to T optimized as synapomorphies at the
aligned positions 181, 183, 230, based on the preferred
topology from POY (Fig. 2).

Of the 11 outgroup taxa of the families Philodromidae, Salticidae, Miturgidae, Corinnidae, Gnaphosidae
and Anyphaenidae included in our study, Hibana sp.
(Anyphaenidae) or Hibana sp. plus Cesonia sp. (Gnaphosidae) were placed as sister to Thomisidae. This was
unexpected for two reasons: first, Salticidae was previously considered sister to Thomisidae (Loerbroks, 1984)

and second, Philodromidae was considered a subfamily
of Thomisidae (Simon, 1892). Philodromids are still
considered by some contemporary arachnologist as
derived thomisids (Tikader, 1980; Roberts, 1995). In
this study they do not group within Thomisidae, and
may fall at the root of Dionycha. In contrast, Cheiracanthium sp. is placed closer to the root of Thomisidae.
Cheiracanthium was previously a member of Clubionidae, a family also proposed as sister to Thomisidae
(Ono, 1988).


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S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

Fig. 4. The strict consensus with gap opening ⁄ gap extension cost of 1 ⁄ 1 under direct optimization: with the addition of the ingroups Misumenoides
sp., two Mecaphsesa and one Xysticus species and excluding the outgroup Portia labiata. See text for details. Jackknife values greater than 60 are
shown above the branches.

All phylogenetic analyses independent of gap opening ⁄ gap extension cost or taxon sampling, recover four
well-supported lineages within Thomisidae. They are
informally named here as the Borboropactus clade,
Epidius clade, Stephanopis clade and the Thomisus
clade, pending detailed morphology-based cladistic
work. The Borboropactus clade is sister to all remaining
thomisids. All ‘‘derived’’ thomisids are grouped in

the Thomisus clade. Epidius and Stephanopis clades
are more closely related to each other than to the
previous two groups. None of these clades recovered
in our study correspond strictly to any currently
accepted subfamily groupings. In the most current
study Thomisidae was separated into seven sub-

families: Stephanopinae, Thomisinae, Bominae,
Stiphropodinae, Dietinae, Strophiinae and Aphantochilinae (Ono, 1988). The current study shows these
groupings to be paraphyletic and should thus be
treated with caution.
The following morphological synapomorphies are
proposed to define Thomisidae:
1 Legs 1 and 2 longer and stronger then legs 3 and 4
(Ono, 1988; Wunderlich, 2004b; Jocque´ and DippenaarSchoeman, 2006). However, this might not be obvious
for some African genera such as Thomisops Karsch,
1879 (Dippenaar-Schoeman, 1989).
2 Lateral eyes on tubercles, larger and much more
developed than the median eyes (Ono, 1988).


S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

717

Fig. 5. The single most parsimonious tree (L ¼ 3234, CI ¼ 0.32, RI ¼ 0.477): static alignment, analyzed under unweighted parsimony. Support
values are show as follows: above branches Bootstrap ⁄ Poisson Bootstrap; below branches Jackknife ⁄ Symmetric resampling. Support values bellow
50 are omitted.

3 Presence of a group of setae instead of a colulus

(Homann, 1975).
Borboropactus clade
Borboropactus Simon, 1884 is one of the few thomisid
genera represented in amber. It was dubbed a ‘‘relict’’
and elevated to family rank (Wunderlich, 2004a,b). This
study shows that there is no phylogenetic justification
for such a family (unless of course Thomisidae is split to
four families). Borboropactidae would be paraphyletic
as it includes the Epidius clade. As Wunderlich (2004b)
himself mentions Borboropactus shares characters such
as the presence of a conductor and median apophysis
with other thomisids such as Epidius and Cebrenninus
(Benjamin, 2000; Benjamin unpublished data). Further,

the Malagasy endemic Geraesta Simon, 1889 should be
included in this clade based on the three characters given
below (Benjamin unpublished data).
The following morphological synapomorphies might
define the Borboropactus clade: (1) flexibly attached cupshaped median apophysis (Fig. 8A,B); (2) hyaline conductor (Fig. 8A–C); and (3) epigynal teeth (Fig. 8D–E).
Wunderlich (2004b, figs 4–8) mentions a specialized
gland or ‘‘tarsal pit organ’’ of tarsi of legs 1–4 as
synapmorphic for ‘‘Borboropactidae’’. However, this
sensory organ appears to be autapomorphic for
Borboropactus, as it is absent in Geraesta (Benjamin
unpublished data). Furthermore, his ‘‘Borboropactidae’’
should include the genera Cebreninnus, Cupa, Epidius
and Stephanopis, which then is essentially Simon’s
‘‘Stephanopinae’’. Homann (1934) suggested that



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S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

Fig. 6. Majority rule consensus tree from the trees found by the implied weight analyses with K ranging from 4 to 100. Results with K-values below 4
were excluded from the consensus as suggested by Goloboff (1993b, 1995).

thomisids should be placed in Lycosoidea as they have a
grate-shaped tapetum. Interestingly, Borboropactus has
epigynal teeth as do some Lycosoidea (Fig. 7D,E;
Griswold, 1993). However, Borboropactus has a canoeshaped tapetum and not a grate-shaped tapetum
(Benjamin unpublished data). This apparent character
conflict needs to be addressed with a broader taxon
sample in future studies.
Monophyly of the Epidius and Stephanopis clades
No morphological characters supporting these two
clades are known. This is because Stephanopis and
related genera are poorly studied. It is clear that
Stephanopis is paraphyletic and badly in need of
revision. Preliminary data indicate that Cebrenninus
might be monophyletic. Epidius is the only genus that is
well defined, due to its elongated tibia (Benjamin, 2000).
Monophyly of the Thomisus clade
Our study revealed the Thomisus clade, morphologically the most homogeneous, to be monophyletic
(Figs 1–5). It is defined by several morphological
synapomorphies: (1) scopula hairs circular in crosssection (Homann, 1975); (2) bulbus subequal in length
and width (Benjamin, 2000); (3) tegulum disc shaped
(Ono, 1988); (4) sperm duct follows a circular peripheral

course through the tegulum (Schick, 1965; Ono, 1988;

Benjamin, 2000); (5) conductor absent (Benjamin, 2000;
Schick, 1965); and (6) median apophysis absent.
Within the Thomisus clade, although there is some
phylogenetic structure, none of the groupings are well
supported. This might be due to either an inadequate
taxon sample, a suboptimal character sample (shorter or
less informative gene fragments) or both. On the other
hand it might well be due to paraphyletic genera. This is
most likely the case. The apparent non-monophyly of
several thomisid genera is not surprising, and was
suspected earlier (Benjamin, 2001, 2002; Lehtinen,
2005). The study of generic relationships within the
Thomisus clade will require additional data.

Future studies
Although our hypothesis of thomisid relationships
forms the basis for future phylogenetic studies, it also
emphasizes that little progress has been made in
understanding the phylogeny of this key dionychan
family since Simon’s seminal work almost a century ago.
The confusion in thomisid systematics is probably due
to the emphasis placed on single character systems, such
as male genital morphology, relative size of eyes or the
presence of cheliceral teeth. Thomisid genitalia are
relatively simple and uniform, thus less informative.


S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

719


Fig. 7. Tree recovered from the Bayesian analysis. Clade posterior probabilities are reported. Values <95% are omitted.

Further, it also may be due to the placement of undue
emphasis on taxonomic work based on Holarctic fauna
as in Loerbroks (1984), ignoring the fact that thomisid
diversity is biased towards tropical habitats, following
the typical latitudinal gradient. Furthermore, cladistic
methodology was not used in previous studies.
Addition of more taxa and morphology to future
studies is vital for understand evolutionary relationships
within the four clades named here. Species of most
described genera could be collected easily for molecular

work, and many more are available in museums for
morphological studies. Unfortunately, some enigmatic
genera, such as Heterogriffus Platnick, 1976 and Stiphropella Lawrence, 1952 (Stiphropodinae) or Simorcus
Simon, 1895 (Strophiinae) are known only from a few
museum specimens. There is an immediate need to
include them in future molecular and morphological
studies.
Finally, thomisid outgroup relationships need attention. The phylogenetic relationships of Dionycha


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S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

A


B

C

D

E

Fig. 8. Borboropactus cinerascens. Male palp: (A) ventral view; (B) prolateral view; (C) retrolteral view. Female: (D) epigynum, ventral view; (E)
vulva, ventral view. Scale bars: 1 mm (D,E), 2 mm (A–C). Abbreviations: C conductor; E embolus; ET epigynal teeth; MA median apophysis; RTA
retrolateral tibial apophysis.

(a clade that includes 15 families in addition to Thomisidae) have got to be resolved. Understanding the
placement of Thomisidae within Dionycha is a necessary
first step to resolve this major issue in spider evolution.

Acknowledgments
Thanks to the following individuals for spider specimens: Fernando Alvarez, Cristian Grismado, Charles
Griswold, Charles Haddad, Stephen Lew, Lara Lopardo, Martin Ramirez and Peter Schwendinger. Jonathan Coddington improved earlier drafts of this
manuscript. We thank Fernando Alvarez for analytical
support and Charles Griswold and Pekka Lethinen for
discussion. A. H. Sumanasena (Department of Wild Life
Conservation, Colombo) provided a research permit to
collect in Sri Lanka. Funding for this research has been
provided by grants from the US National Science
Foundation (DEB-0328644 to G. Hormiga and
G. Giribet and EAR-0228699 to W. Wheeler, J.
Coddington, G. Hormiga, L. Prendini and P. Sierwald)
and by a REF grant from The George Washington
University to GH. Additional funding to SPB came

from the Schlinger Foundation, the University of
California, Berkeley and a Smithsonian Institution
postdoctoral fellowship.

References
Aldrich, J.R., Barros, T.M., 1995. Chemical attraction of male crab
spiders (Araneae, Thomisidae) and kleptoparasitic flies (Diptera,
Milichiidae and Chloropidae). J. Arachnol. 23, 212–214.
Arnedo, M.A., Coddington, J., Agnarsson, I., Gillespie, R.G., 2004.
From a comb to a tree: phylogenetic relationships of the combfooted spiders (Araneae, Theridiidae) inferred from nuclear and
mitochondrial genes. Mol. Phylogenet. Evol. 31, 225–245.
Barrion, A.T., Litsinger J.A., 1995. Riceland Spiders of South and
Southeast Asia. CAB International, Wallingford, UK.

Benjamin, S.P., 2000. Epidius parvati sp. n., a new species of the genus
Epidius from Sri Lanka (Araneae: Thomisidae). Bull. Br. Arachnol.
Soc. 11, 284–288.
Benjamin, S.P., 2001. The genus Oxytate L. Koch 1878 from Sri
Lanka, with description of Oxytate taprobana sp. n. (Araneae:
Thomisidae). J. South Asian Nat. Hist. 5, 153–158.
Benjamin, S.P., 2002. Smodicinodes schwendingeri sp. n. from Thailand
and the first male of Smodicinodes Ono, 1993, with notes on the
phylogenetic relationships in the tribe Smodicini (Araneae: Thomisidae). Rev. Suisse Zool. 109, 3–8.
Coddington, J.A., Levi, H.W., 1991. Systematics and evolution of
spiders (Araneae). Annu. Rev. Ecol. Syst. 22, 565–592.
Colgan, D.J., MacLauchlan, A., Wilson, G.D.F., Livingston, S.P.,
Edgecombe, G.D., Macaranas, J., Cassis, G., Gray, M.R. 1998.
Histone H3 and U2 snRNA DNA sequences and arthropodmolecular evolution. Aust. J. Zool. 46, 419–437.
Comstock, J.H., 1948. The Spider Book. Comstock Publishing Co.,
Inc., Ithaca, New York.

Davila, D.S., 2003. Higher-level relationships of the spider family
Ctenidae (Araneae: Ctenoidea). Bull. Am. Mus. Nat. Hist. 274, 1–
86.
Dippenaar-Schoeman, A.S., 1989. The crab-spiders of Africa (Araneae: Thomisidae). 8. The genus Thomisops Karsch, 1879. Phytophylactica, 21, 319–330.
Evans, T.A., 1995. Two new species of social crab spiders of the genus
Diaea from Eastern Australia, their natural history and distribution. Rec. West. Aust. Mus. 52(Suppl.), 151–158.
Evans, T.A., 1998. Offspring recognition by mother crab spiders
with extreme maternal care. Proc. R. Soc. Lond. B, 265, 129–
134.
Farris, J.S., 2001. Support weighting. Cladistics 17, 389–394.
Farris, J.S., Albert, V.A., Kallersjo, M., Lipscomb, D., Kluge, A.,
1996. Parsimony jackknifing outperforms neighbor-joining. Cladistics 12, 99–124.
Farris, J.S., Ka¨llersjo¨, M., Kluge, A.G., Bult, C., 1994. Testing
significance of incongruence. Cladistics 10, 315–319.
Felsenstein, J., 1985. Confidence limits on phylogenies: an approach
using the bootstrap. Evolution 39, 783–791.
Fitch, W.M., 1971. Towards defining the course of evolution:
minimal change for a specific tree topology. Syst. Zool. 20, 406–
416.
Gabritschevsky, E., 1927. Experiments on color changes and regeneration in the crab-Spider, Misumena vatia. J. Exp. Zool. 47, 251–
266.
Giribet, G., 2001. Exploring the behavior of POY, a program for direct
optimization of molecular data. Cladistics 17, 60–70.


S. P. Benjamin et al. / Cladistics 24 (2008) 708–722
Goloboff, P.A., 1993a. Estimating character weights during tree
search. Cladistics 9, 83–91.
Goloboff, P.A., 1993b. Pee-Wee, ver. 3.0. Program available at http://
www.cladistics.com.

Goloboff, P.A., 1995. Parsimony and weighting: a reply to Turner and
Zandee. Cladistics 11, 91–104.
Goloboff, P.A., Farris, J.S., Nixon, K., 2003a. T.N.T. Tree Analysis
Using New Technology, Version 1.0. Program and documentation
available at />Goloboff, P.A., Farris, J.S., Ka¨llersjo¨, M., Oxelman, B., Ramı´ rez,
M.J., Szumik, C.A., 2003b. Improvements to resampling measures
of group support. Cladistics 19, 324–332.
Greco, C.F., Kevan, P.G., 1994. Contrasting patch choosing by
anthophilous ambush predators: vegetation and floral cues for
decisions by a crab spider (Misumena vatia) and males and females
of an ambush bug (Phymata americana). Can. J. Zool. 72, 1583–
1588.
Griswold, C.E., 1993. Investigations into the phylogeny of the
lycosid spiders and their kin (Arachnida: Araneae: Lycosoidea).
Smithson. Contrib. Zool. 539, 1–39.
Harvey, P.H., Pagel, M.D., 1991. The Comparative Method in
Evolutionary Biology. Oxford University Press, Oxford.
Hedin, M.C., Maddison, W.P., 2001. A combined molecular approach
to phylogeny of the jumping spider subfamily Dendryphantinae
(Araneae: Salticidae). Mol. Phylogenet. Evol. 18, 386–403.
Holm, A., 1940. Studien u¨ber die Entwicklung und Entwicklungsbiologie der Spinnen. Zool. Bidr. Uppsala 19, 1–214.
Homann, H., 1934. Beitra¨ge zur Physiologie der Spinnenaugen. IV.
Das Sehvermo¨gen Thomisiden. Z. Vergl. Physiol. 20, 420–429.
Homann, H., 1975. Die Stellung der Thomisidae und der Philodromidae im System der Araneae (Chelicerata, Arachnida). Z. Morph.
O¨kol. Tiere 80, 181–202.
Jocque´, R., Dippenaar-Schoeman, A.S., 2006. Spider Families of the
World. Royal Museum of Central Africa, Tervuren.
Krell, F., Kraemer, F., 1998. Chemical attraction of crab spider
(Araneae, Thomisidae) to a flower fragrance component. J.
Arachnol. 26, 117–119.

Lehtinen, P.T., 2005. Taxonomic notes on the Misumenini (Araneae:
Thomisidae: Thomisinae), primarily from the Palaearctic and
Oriental regions. In: Logunov, D.V., Penney, D. (Eds.), European
Arachnology 2003. Arthropoda Selecta (Special Issue no. 1, 2004).
Levi, H.W., 1982. Araneae. In: Parker, S.P. (Ed.), Araneae. McGraw
Hill, New York, pp. 77–95.
Loerbroks, A., 1984. Mechanik der Kopulationsorgane von Misumena
Vatia. Clerck, 1757) (Arachida: Araneae: Thomisidae). Abh.
Naturw, Ver. Hamburg (NF) 27, 383–403.
Maddison, W.P., Maddison, D.R., 2001. MacClade: Analysis of
Phylogeny and Character Evolution, ver. 4.0. Sinauer Associates,
Sunderland, MA.
Main, B.Y., 1988. The biology of a social thomisid spider. In: Austin,
A.D., Heather, N.W. (Eds.), Australian Arachnology. Australian
Entomological Society, Brisbane, pp. 55–73.
Mickevich, E., Farris, J.S., 1981. The implications of congruence in
Menidia. Syst. Zool. 30, 351–370.
Morse, D.H., 1988. Cues associated with patch-choice decisions by
foraging crab spiders Misumena vatia. Behaviour 107, 297–313.
Nixon, K.C., 2002. WinClada. Published by the Author.
Nyffeler, M., Benz, G., 1987. Spiders in natural pest control: a review.
J. Appl. Entomol. 103, 321–339.
Oliveira, P.S., Sazima, I., 1984. The adaptive bases of ant-mimicry in a
neotropical aphantochilid spider (Araneae: Aphantochilidae). Biol.
J. Linn. Soc. 22, 145–155.
Ono, H., 1988. A Revisional Study of the Spider Family Thomisidae
(Arachnida, Araneae) of Japan. National Science Museum, Tokyo.
Packard, A.S., 1905. Change of color and protective coloration in a
flower-spider (Misumena vatia Thorell). J. N.Y. Entomol. Soc. 13,
85–96.


721

Petrunkevitch, A., 1928. Systema Araneorum. Trans. Connect. Acad.
Arts Sci. 29, 1–270.
Pickard-Cambridge, O., 1871. Arachnida. Zool. Rec. 7, 207–224.
Platnick, N.I., 2007. The World Spiders Catalog, ver. 7.5. American
Museum of Natural History, New York. />entomology/spiders/catalog81-87/index.html/.
Posada, D., Crandall, K.A., 1998. MODELTEST: testing the model of
DNA substitution. Bioinformatics 14, 817–818.
Riechert, S.E., 1974. Thoughts on the ecological significance of spiders.
Bioscience 24, 352–356.
Riechert, S.E., Lockley, T.C., 1984. Spiders as biological control
agents. Annu. Rev. Entomol. 29, 299–320.
Roberts, M.J., 1995. Spiders of Britain and Northern Europe. Harper
Collins, London.
Roewer, C.F., 1954. Katalog der Araneae von 1758 bis 1940, bzw
1954. Institut Royal des Sciences Naturelles de Belgique, Bruxelles.
Ronquist, F., Huelsenbeck, J.P., 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572.
Schick, R.X., 1965. The crab spiders of California (Araneida:
Thomisidae). Bull. Am. Mus. Nat. Hist. 129, 1–180.
Shelford, R., 1902. Observations on some mimetic insects and spiders
from Borneo and Singapore. Proc. Zool. Soc. Lond. 1902, 230–284.
Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H., Flook, P.,
1994. Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase
chain reaction primers. Ann. Entomol. Soc. Am. 87, 651–701.
Simon, E., 1892 (1892–1895). Histoire naturelle des Araigne´es. Roret,
Paris. pp. 949–1066.
Steel, M.A., 1994. The maximum likelihood point for a phylogenetic
tree is not unique. Syst. Biol. 43, 560–564.

Suzuki, Y., Glazko, G.V., Nei, M., 2002. Overcredibility of molecular
phylogenies obtained by Bayesian phylogenetics. Proc. Acad. Nat.
Sci. Phila. 99, 16138–16143.
Swofford, D.L., 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods), ver. 4. Sinauer Associates, Sunderland, MA.
Swofford, D.L., Olsen, 1991. Phylogeny reconstruction. In: Hillis,
D.M., Moritz, C. (Eds.), Phylogeny Reconstruction. Sinauer,
Sunderland, MA, pp. 407–514.
Thompson, J.D., Higgins, D.G., Gibson, T.J., 1994. CLUSTAL W:
improving the sensitivity of progressive multiple sequence alignment through sequence weighting, positions-specific gap penalties
and weigh matrix choice. Nucleic Acids Res. 22, 4673–4680.
Thorell, T., 1873. Remarks on synonyms of European spiders. Part IV,
C.J. Lundstro¨m, Uppsala, pp. 375–645.
Tikader, B.K., 1980. Thomisidae (Crab-spiders). Zoological Survey of
India, Calcutta.
Uetz, G.W., Halaj, J., Cady, A.B., 1999. Guild structure of spiders in
major crops. J. Arachnol. 27, 270–280.
Wheeler, W.C., 1996. Optimization alignment: the end of multiple
sequence alignment in phylogenetics? Cladistics 12, 1–9.
Wheeler, W.C., Hayashi, C.Y., 1998. The phylogeny of the extant
chelicerate orders. Cladistics 14, 173–192.
Wheeler, W.C., Gatesy, J., DeSalle, R., 1995. Elision: a method for
accommodating multiple molecular sequence alignments with
alignment-ambiguous sites. Mol. Phylogenet. Evol. 4, 1–9.
Wheeler, W.C., Aagesen, L., Arango, C.P., Faivovich, J., Grant, T.,
D’Haese, C., Janies, D., Smith, W.L., Varo´n, A., Giribet, G., 2006.
Dynamic Homology and Phylogenetic Systematics: A Unified
Approach Using POY. American Museum of Natural History,
New York.
Wilcox, T.P., Zwickl, D.J., Heath, T.A., Hillis, D.M., 2002. Phylogenetic relationships of the dwarf boas and a comparison of Bayesian
and bootstrap measures of phylogenetic support. Mol. Phylogenet.

Evol. 25, 361–371.
Wunderlich, J., 2004a. Fossil crab spiders (Araneae: Thomisidae) in
Baltic and Dominican amber. Beitr. Araneol. 3, 1747–1760.


722

S. P. Benjamin et al. / Cladistics 24 (2008) 708–722

Wunderlich, J., 2004b. The new spider (Araneae) family Borboropactidae from the tropics and fossil in baltic amber. Beitr. Araneol. 3,
1737–1746.

Appendix 1
Taxonomic changes, Fig. 7(A–E).
Genus: Borboropactus Simon, 1884.
Borboropactus cinerascens (Doleschall, 1859).
B. bangkongeus Barrion and Litsinger (1995). New synonomy.
B. mindoroensis Barrion and Litsinger (1995). New synonomy.
B. umaasaeus Barrion and Litsinger (1995). New synonomy.
B. mindoroensis is a juvenile female; penultimate female thomisids
some times have partly sclerotized genitalia. B. bangkongeus and
B. umaasaeus are one and the same species. Their illustrations (Barrion
and Litsinger, 1995) leave no doubt that they are synonymous with
B. cinerascens. Vouchers are deposited at the Natural History Museum
of Geneva.

Appendix 2
Glossary of terms
Terms used in the discussion taken from Jocque´ and DippenaarSchoeman (2006).


Conductor: a sclerite of the male copulatory palp that accompanies
and supports the embolus; a hyaline conductor is a conductor that is
transparent and very thin (Fig. 8C).
Colulus: short median protuberance in front of spinnerets, considered a modification of the spinning plate or cribelum.
Embolus: embolus is the intromitted part of the male copulatory
palp (Fig. 8B).
Epigynum: a usually chitinous section on the ventral side of the
abdomen of adult female spiders on which the genital openings are
found. Epigynes of some thomisids have lateral sclerotized projections
termed teeth (Fig. 7D,E).
Male palp: modified tarsus of the palp in male spiders. It is a
copulatory organ of great complexity and is probably the most
important character system in spiders (Fig. 7A,B).
Median apophysis: an apophysis of the male palpal bulb (Fig. 8B).
Scopula hairs: specialized hairs found on the tip of legs in some
thomisids. Thought to improve grip.
Tegulum: part of the male copulatory palp that houses the sperm
duct.