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Continuing Invertebrate Taxonomy
Volume 16, 2002
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© CSIRO 2002 10.1071/IS02012 1445-5226/02/060839
Invertebrate Systematics, 2002, 16, 839–847
Molecular phylogeny of the mud lobsters and mud shrimps
(Crustacea : Decapoda : Thalassinidea) using nuclear 18S rDNA
and mitochondrial 16S rDNA
C. C. TudgeA,C and C. W. CunninghamB
ABiology Department, American University, 4400 Massachusetts Ave. NW, Washington, DC 20016-8007,
USA and Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution,
Washington, DC 20560-0163, USA.
BDepartment of Zoology, Duke University, Durham, NC 27708-0325, USA.
CTo whom correspondance should be addressed. Email: email@example.com
14 species of thalassinidean shrimp (families Callianassidae, Laomediidae, Strahlaxiidae, Thalassinidae and
Upogebiidae) and a further six species in related decapod infraorders (families Aeglidae, Astacidae, Lithodidae,
Palinuridae, Raninidae and Scyllaridae). Maximum-likelihood and Bayesian analyses show equivocal support for
the monophyly of the Thalassinidea, but show strong support for division of the infraorder into two major clades.
This dichotomy separates representatives in the Upogebiidae, Laomediidae and Thalassinidae from those in the
Strahlaxiidae and Callianassidae. The Laomediidae is shown to be paraphyletic, with the thalassinid species,
Thalassina squamifera, being placed on a branch between Axianassa and a clade comprising Jaxea and Laomedia,
the three current laomediid genera. For a monophyletic Laomediidae, the family Axianassidae should be resurrected
for the genus Axianassa.
Partial sequences of the 18S nuclear and 16S mitochondrial ribosomal genes were obtained for
Molecular phylogeny of the ThalassinideaC. C. Tudge and C. W. Cunningham
The decapod infraorder Thalassinidea is a group of cryptic,
marine, burrowing shrimp-like or lobster-like crustaceans
that occur worldwide (with the exception of the coldest polar
waters) in mostly shallow (<200 m) benthic habitats. Most
species form complex burrow systems in soft sand or mud
environments, but several taxa live in stony or coral-rubble
areas and some even excavate burrows in living coral and
sponge colonies (Dworschak 2000). There are currently
528 species in 84 genera spread across 11 recognised
families of the three superfamilies: Axioidea, Thalassinoidea
and Callianassoidea (Poore 1994; Dworschak 2000;
P. Dworschak, personal communication).
Borradaile (1903: 551), in his seminal work on the
classification of the Thalassinidea, presented a ‘genealogical
tree’ of proposed relationships among 12 known genera in the
four families recognised at that time. His tree (reproduced in
Fig. 1A with current family designations) separates these
genera into five major clades corresponding with the families
Laomediidae (Jaxea, Laomedia), Thalassinidae (Thalassina)
and the two subfamilies, Callianassinae (Callianassa,
Callianidea, Glypturus) and Upogebiinae (Gebicula,
Upogebia), of the Callianassidae. Later, Gurney (1938: 343)
used larval morphology to present an intuitive tree of
relationships among four thalassinidean families and the
Anomura (Fig. 1B). A period of 56 years elapsed before any
further publications on the phylogenetic relationships within
this obscure, but taxonomically large, infraorder emerged.
Poore (1994: 120) published a comprehensive revision of the
members of the Thalassinidea based on morphology, including
a new classification scheme, keys to families and genera and
a phylogenetic tree (reproduced in Fig. 1C to the family level
only). This tree was the first cladistic analysis of the infraorder
as a whole and separated the 22 selected genera into the
currently recognised 11 families and three superfamilies
(despite showing a basal dichotomy with only two major
clades). He also proposed a monophyletic origin for the
infraorder, with the Anomura being the closest sister-group.
Several papers have been published employing cladistic
analyses of relationships among or within genera of certain
thalassinidean families. These investigations cover the
families Axiidae and Calocarididae (Kensley 1989),
Callianideidae (Kensley and Heard 1991), Callianassidae
(Staton and Felder 1995; Staton et al. 2000; Tudge et al.
2000) and Ctenochelidae (Tudge et al. 2000).
In some cases, thalassinidean representatives have been
used as either ingroup or outgroup taxa for larger cladistic
analyses of various Decapoda; these include studies using
morphological characters by Martin and Abele (1986),
Scholtz and Richter (1995) and Tudge (1997). Several
840 C. C. Tudge and C. W. Cunningham
molecular phylogenetic analyses of various decapod
taxonomic groups have also included thalassinidean taxa for
comparison with anomurans (Spears and Abele 1988; Pérez-
Losada et al. 2002), astacideans (Crandall et al. 2000) and
crab-like decapods (Morrison et al. 2002). Also, see the
synopsis of recent research into decapod phylogenetics by
Schram (2001) for a review of the different approaches and
data sets being applied to the field.
The contribution of the small-subunit 18S ribosomal
(r)DNA nuclear gene to crustacean phylogeny is well known
and has been useful in investigating relationships across a
wide variety of groups (Spears and Abele 1988, 1997, 1998;
Abele et al. 1989, 1992; Kim and Abele 1990; Abele 1991;
Spears et al. 1992; Crandall et al. 2000; Morrison et al. 2002;
Pérez-Losada et al. 2002). Similarly, the mitochondrial 16S
rDNA gene has been regularly used to investigate decapod
relationships (Cunningham et al. 1992; Crandall and
Fitzpatrick 1996; Tam et al. 1996; Ovenden et al. 1997;
Kitaura et al. 1998; Tam and Kornfield 1998; Crandall et al.
1999, 2000; Schubart et al. 2000, 2001; Duffy et al. 2000;
Morrison et al. 2002). Different regions of ribosomal genes
evolve at varying rates, making these useful molecular
markers across a broad taxonomic spectrum.
This paper presents the results of a phylogenetic analysis
of partial sequences of mitochondrial 16S rDNA and nuclear
18S rDNA from 14 thalassinideans in five families
Borradaile (modified from Borradaile 1903). (B) Intuitive tree of relationships between some
thalassinideans based on larval characters (modified from Gurney 1938). (C) Cladogram of
phylogenetic relationships between 11 families of the Thalassinidea based on morphological
characters (modified from Poore 1994). Note: the trees of both Borradaile and Poore have
been modified to show taxonomic relationships at the family-level or higher.
(A) Genealogical relations between families in the Thalassinidea according to
Molecular phylogeny of the Thalassinidea841
(Callianassidae, Laomediidae, Strahlaxiidae, Thalassinidae
and Upogebiidae). Six other decapod taxa from the
Astacidea, Palinura, Brachyura and Anomura are included in
the analysis as outgroups. Similarities and differences to
previous evolutionary trees of the Thalassinidea, most
notably those of Borradaile (1903), Gurney (1938), Poore
(1994) (Figs 1A–C) and Tudge et al. (2000), are discussed.
Materials and methods
With the exception of the Raninoides specimen from 1991, animals for
the genetic analysis were mostly collected over a 5-year period
(1995–1999) from a variety of subtidal and intertidal locations
worldwide (Table 1), but with an emphasis on coastal Australia and the
United States. The intertidal specimens were collected by ‘yabby’ pump,
digging or by hand and the subtidal specimens were trawled or dredged.
Table 1. Specimens, collection, voucher and GenBank-sequence information
Higher-level classification from Martin and Davis (2001). Abbreviations: AMNH, American Museum of Natural History (New York); MV,
Museum Victoria; Qld, Queensland; USA, United States of America; USNM, National Museum of Natural History (Washington DC); Vic, Victoria.
Procambarus clarkii (Girard, 1852). Louisiana, USA, Jan. 1998. Voucher: AMNH 17874. GenBank sequences: AF436040 (16S), AF436001
Panulirus argus (Latreille, 1804). GenBank sequences: AF337966 (16S), U19182 (18S)
Thenus orientalis (Lund, 1793). Gulf of Carpentaria, Qld, Australia, 1997. Voucher: not available
Raninoides louisianensis Rathbun, 1933. Gulf of Mexico, USA, 1991. Voucher: T. Spears personal collection. GenBank sequences:
AF436044 (16S), AF436005 (18S)
Aegla uruguayana Schmitt, 1942. San Antonio de Areco, Buenos Aires, Argentina, 10 Mar. 1996. Voucher: C. Cunningham personal
collection. GenBank sequences: AF436051 (16S), AF436012 (18S).
Cryptolithodes typicus Brandt, 1849. British Columbia, Canada. Voucher: S. Zaklan personal collection. GenBank sequences: AF425325
(16S), AF436019 (18S)
Thalassina squamifera de Man, 1915. Town of 1770, Qld, Australia, 29 Dec. 1997. Voucher: MV J41662
Biffarius arenosus (Poore, 1975). Dunwich, Qld, Australia, 21 May 1997. Voucher: MV J40669
Biffarius delicatulus Rodrigues & Manning, 1992. Fort Pierce, Florida, USA, 9 Jun. 1999. Voucher: USNM 309754
Callianassa filholi A. Milne Edwards, 1878. Otago Harbour, New Zealand, 14 Nov. 1997. Voucher: MV J44818
Callichirus major (Say, 1818). Isles Dernieres, Louisiana, USA, 27 Mar. 1996. Voucher: MV J39044. GenBank sequences: AF436041 (16S),
Neocallichirus rathbunae (Schmitt, 1935). Fort Pierce, Florida, USA, 8 Jun. 1999. Voucher: USNM 309751
Neotrypaea californiensis (Dana, 1854). Marina del Rey, California, USA, 20 Jun. 1997. Voucher: A. Harvey personal collection. GenBank
sequences: AF436042 (16S), AF436003 (18S)
Sergio mericeae Manning & Felder, 1995. Fort Pierce, Florida, USA, 8 Jun. 1999. Voucher: USNM 309755
Axianassa australis Rodrigues & Shimizu, 1992. Sáo Sebastiáo, Sáo Paulo, Brazil, 20 Jul. 1997. Voucher: MV J44613
Jaxea nocturna Nardo, 1847. Loch Sween, Argyll, Scotland, 10 Nov. 1996. Voucher: MV J39045. GenBank sequences: AF436046 (16S),
Laomedia healyi Yaldwyn & Wear, 1970. Western Port Bay, Vic., Australia, 16 Apr. 1997. Voucher: MV J40697
Gebiacantha plantae (Sakai, 1982). Yorke Island, Qld, Australia, 26 Feb. 1998. Voucher: MV J44914
Upogebia affinis (Say, 1818). Fort Pierce, Florida, USA, 15 Nov. 1995. Voucher: MV J40668. GenBank sequences: AF436047 (16S),
Neaxius glyptocercus von Martens, 1868. Dunwich, Qld., Australia, 21 May 1997. Voucher: MV J39643
842C. C. Tudge and C. W. Cunningham
Tissue samples were preserved in 95% ethanol. DNA was isolated from
fresh, frozen or ethanol-preserved specimens by grinding small
fragments of muscle tissue in a buffer (0.1 M EDTA, 0.01 M Tris pH 7.5,
1% SDS, Palumbi et al. 1991), followed by phenol-chloroform-isoamyl
alcohol extraction and precipitation with 7.5 M ammonium acetate and
cold isopropanol as described by Palumbi et al. (1991). Partial
sequences of mitochondrial 16S rDNA were amplified using rDNA
primers (LR-N-13398, alias 16Sar, and LR-J-12887, alias 16 Sbr,
Simon et al. 1994). Most of the nuclear 18S rDNA gene was amplified
using 18E-F (5′-CTGGTTGATCCTGCCAGT-3′) and 18SR3 (5′-
TAATGATCCTTCCGCAGGTT-3′). The amplification and sequencing
of the two genes (16S and 18S) was identical (except where indicated)
and entailed the following regime. Polymerase chain reaction (PCR)
was carried out on a Perkin Elmer GeneAmp PCR System 9600
(www.pecorporation.com) using 50 µL reactions consisting of 1 µL
template DNA, 3 µL primers, 4.5 µL 10× reaction buffer, 5 µL Taq
polymerase, 4 µL deoxynucleoside triphosphates, 5 µL MgCl2 and 24.5
µL water. Amplification involved 20 sec denaturation at 94°C, 1.5 min
annealing at 50°C (40°C for 16S) and 2.5 min of extension at 72°C for
35 cycles. The PCR product (3 µL) was visualised, via electrophoresis,
on a 1% agarose gel using ethidium bromide staining. The remaining
product (45 µL) was purified using the Wizard PCR Preps
(www.promega.com) DNA Purification System. Applied Biosystems,
Inc. (ABI) (www.appliedbiosystems.com) Prism BigDye Terminator
Cycle Sequencing Ready Kits and an ABI Prism 3700 DNA Analyzer
were used for sequencing. Sequences are available from the authors
Sequence alignment and phylogenetic analysis
Sequences were aligned using Clustal X (Thompson et al. 1994, 1997)
with gap insertion and extension costs at 10 and 5, respectively, and
regions of uncertain homology removed before phylogenetic analysis.
Maximum-likelihood (ML) analyses using heuristic searches and TBR
branch swapping were applied to the aligned sequences using PAUP*
4.0 (Swofford 2001). For both maximum-likelihood and Bayesian
analyses, we used the general-time-reversible (GTR) model to estimate
the proportion of invariant sites and the alpha parameter of the gamma
distribution from the data (the best-fit model using ModelTest, Posada
and Crandall 1998). Incongruence testing was carried out under the
parsimony criterion using the incongruence length difference (ILD) test
(Farris et al. 1995), as implemented in PAUP* 4.0.
The Bayesian analyses were performed using the program
MRBAYES 1.11 (Huelsenbeck and Ronquist 2001). MRBAYES uses a
metropolis-coupled Markov chain Monte Carlo (MCMCMC) algorithm
to sample from the posterior distribution. Each Markov chain was
started from a random tree and run for 5 million cycles. The chain was
sampled every 500 cycles in order to minimise the size of the output
files and to ensure that these samples were independent. Four chains
were run simultaneously with a ‘temperature’ of 0.2. The initial 40% of
cycles were discarded as burn-in and convergence was checked by
ensuring that the likelihood values of sampled trees had approached a
level distribution. Each analysis was performed twice to ensure that
convergence was repeatable.
Tests of alternative topologies were performed using the
Shimodaira–Hasegawa (SH) test (Shimodaira and Hasegawa 1999),
which does not require that the hypotheses be designated a priori.
Probability values were calculated in PAUP* 4.0 using 1000 RELL
replicates to obtain a distribution.
After removing regions of uncertain alignment, our aligned
sequences consisted of 1731 base pairs (bp) of nuclear 18S
rDNA and 327 bp of mitochondrial 16S rDNA for a total of
2058 bp. Our final alignments are available from the authors
(CWC). An ILD test (Farris et al. 1995) showed no
significant incongruence (P > 0.05) and the two genes were
analysed separately and together.
For both genes, the best-fit model found using ModelTest
under maximum-likelihood was GTR + gamma + inv.
Bootstrap consensus trees for both genes, analysed separately
for the maximum-likelihood bootstrap and Bayesian analyses,
are presented in Fig. 2. There is weak support in the 18S
analysis for a monophyletic Thalassinidea (70% Bayes, 42%
bootstrap). Although Bayesian analysis of the 16S gene
showed rather strong support for a paraphyletic Thalassinidea
(85%), there is little ML bootstrap support for a paraphyletic
Thalassinidea (33%). The conflict between the 18S and 16S
genes is not strong (also reflected by a non-significant ILD
test of P > 0.05) and they were combined for further analysis.
The ML analysis found a tree (Fig. 3) with Ln likelihood
= –8184.94 that weakly supports a paraphyletic
Thalassinidea (76% Bayes, 56% bootstrap). A monophyletic
Thalassinidea could not be rejected by an SH test.
In these analyses, monophyly of the Thalassinidea was
equivocal, being weakly supported by the 18S dataset only
(Fig. 2, left side) and not by the 16S dataset (Fig. 2, right side)
or the combined analysis (Fig. 3). Poore (1994; Fig. 1C) found
a monophyletic Thalassinidea based on two morphological
synapomorphies (reduction of pleurobranchs on gills to seven
or less and presence of a setose lower margin on the propodus
and carpus of pereopod 2), but a molecular study by Morrison
et al. (2002) found strong support for non-monophyly of the
Thalassinidea (89% ML bootstrap support). The latter study
included more sequences (16S and 18S, as well as partial
sequences of mitochondrial COII and nuclear 28S ribosomal
DNA), but fewer thalassinidean taxa than the present study.
It is possible that the more rapidly evolving sequences used
in the Morrison et al. (2002) study give a more reliable
indication of relationships than just the two genes used in the
Between the Bayesian and the ML analyses there is no clear
candidate for a sister-group to the Thalassinidea as a whole
(Figs 2, 3). Interestingly though, the Bayesian analysis found
strong (90%) support for a monophyletic sister-group to some
taxa in the Thalassinidea, including the spiny and slipper
lobsters (Panulirus and Thenus), the anomurans (Aegla and
Cryptolithodes) and the brachyuran (Raninoides). Although
this is intriguing, the weak support for this clade in
bootstrapping (59%) suggests that further data may be
necessary to test this hypothesis.
Within the Thalassinidea, we found two strongly supported
major clades (Figs 2, 3). The first clade includes a
monophyletic Callianassidae as the sister-group to a
representative in the axioid family, Strahlaxiidae (Neaxius
glyptocercus). Within the subfamily Callianassinae, the genus
Molecular phylogeny of the Thalassinidea 843
Biffarius is paraphyletic, with representatives of the genus
apparently grouping according to biogeography, uniting the
American taxa Biffarius delicatulus and Neotrypaea
californiensis, and uniting the antipodean taxa, Biffarius
arenosus and Callianassa filholi. These associations, although
at first appearing enigmatic, should be viewed in light of
comments by Tudge et al. (2000) that generic relationships
within the family Callianassidae should be treated with some
caution pending a re-diagnosis of the inclusive taxa, the
subfamily Callianassinae and the nebulous genus Callianassa
in particular. The apparent biogeographic associations
between the four species mentioned above are contradictory
to those proposed for the same taxa in the morphological
analysis (Tudge et al. 2000), where the two Biffarius species
are included in a monophyletic clade (only 59% supported)
and C. filholi and N. californiensis are in another smaller
monophyletic clade (100% supported). The same
morphological analysis by Tudge et al. (2000) grouped these
four callianassine taxa into a monophyletic clade and indicated
a comparable lack of resolution between this clade and the
callichirine taxa, Callichirus, Sergio and Neocallichirus. The
monophyletic Callianassidae in this molecular analysis (Figs
2, 3) is supported by at least 10 morphological characters (see
family diagnosis in Poore 1994). In addition to Tudge et al.
(2000), a recent synopsis of the family Callianassidae has been
provided by Sakai (1999). His paper indicates a polyphyletic
family, extensively synonymises
Callianassoidea and has been considered of limited value in
helping to elucidate the relationships in this group of
thalassinideans (Tudge et al. 2000; Poore 2000).
The large clade constituted by the Callianassidae and
Strahlaxiidae in this molecular analysis (Figs 2, 3) is also
supported by six morphological synapomorphies. Three of
these synapomorphies can be found in the analysis of Poore
(1994), but are also shared by representatives in several other
thalassinoid families (Callianideidae,
Micheleidae and Thomassiniidae) for which tissue was not
available for this molecular analysis. These characters are (1)
the presence of dense tufts of setae on abdominal somites 3–5
(2) the absence of the appendix interna on male pleopod 1 and
(3) a similar absence on male pleopod 2. Three additional
morphological synapomorphies were found to support the
members of the
side) and the 16S data set (right side). Percentage support for Bayesian posterior probabilities (upper) and maximum-likelihood bootstrap values
(lower) are given for each node. Best-fit model = GTR + gamma + inv.
A combined figure showing the opposed consensus trees for both Bayesian and maximum-likelihood analyses of the 18S data set (left
844 C. C. Tudge and C. W. Cunningham
association of the Callianassidae with the Strahlaxiidae in an
unpublished analysis by the senior author, which did not use
the exact corresponding taxa. These characters are (4) a
chelate pereopod 2 with the dactylus as long as the fixed finger
(5) the absence of spiniform setae on the dactylus of pereopods
3 and 4 and (6) a row of lateral plumose setae on abdominal
The second major clade in Fig. 2 (left side only) and Fig. 3
includes three genera currently in the Laomediidae (Jaxea,
Laomedia, Axianassa), as well as representatives of the
Thalassinidae and Upogebiidae. Within this clade, the
placement of the thalassinid, Thalassina squamifera, makes
the Laomediidae paraphyletic, or monophyletic with the
inclusion of Thalassina. This latter, alternative hypothesis,
although lacking any convincing morphological support (see
below), could not be rejected by an SH test of the molecular
data. The paraphyly of the laomediids, suggested in the
present analysis, resurrects an interesting issue on the
validity of the monotypic
Axianassidae Schmitt, 1924. Kensley and Heard (1990)
succinctly summarised the history of the debate on whether
the Axianassidae is a valid family in the Thalassinidea, and
listed the supporters of the Axianassidae (Gurney 1938;
Wear and Yaldwyn 1966; Goy and Provenzano 1979; Poore
and Griffin 1979) and those who believe the genus
Axianassa is one of five in the Laomediidae (de Man 1928;
single maximum-likelihood tree for the combined 16S and 18S data sets. Percentage
support for Bayesian posterior probabilities (upper) and maximum-likelihood bootstrap
values (lower) are given for each node. Best-fit model = GTR + gamma + inv. Some higher
taxonomic categories are provided on specific clades and the six outgroup taxa are
indicated. Decapod images are modified from (top to bottom): Hobbs 1981; Rathbun 1937;
Jara and Palacios 1999; Bliss 1982; Holthuis 1991; Rodrigues and Shimizu 1992; Sakai
1992; Yaldwyn and Wear 1970; Kensley et al. 2000; Felder and Felgenhauer 1993; Hart
1982; Holthuis 1991. Note: images are not necessarily the same species as used in the
analysis, but are representative of the genus, or a closely related genus.
A composite of the consensus trees obtained from the Bayesian analysis and the
Molecular phylogeny of the Thalassinidea 845
Balss 1957; Le Loeuff and Intes 1974; Ngoc-Ho 1981). Our
data support retention of the family Axianassidae for the
genus Axianassa, with Jaxea and Laomedia remaining in the
Laomediidae. As well as the molecular evidence for a valid
Axianassidae presented here (Figs 2, 3), we have identified
10 morphological characters (six of them apomorphies) in
which Axianassa differs from Jaxea and Laomedia. These
apomorphic characters are: (1) the linea thalassinica
displaced dorsally (possible autapomorphy); (2) absence of
lateral lobes on abdominal somite 1; (3) linear epipods on the
gills; (4) reduction (or absence) of an exopod on maxilliped
3; (5) unequal chelae on pereopod 1; and (6) dense tufts of
lateral setae on abdominal somites 3–5.
The association of the Thalassinidae and the Laomediidae
(the 76% ML bootstrap supported node with Thalassina,
Jaxea, and Laomedia in Fig. 3) is interesting, but lacks
strong morphological support. In fact, only two
synapomorphies(?), the posterior margin of the carapace
with strong lateral lobes and both anterior and posterior teeth
on the mandibular incisor, can be gleaned from the cladistic
analysis by Poore (1994). The former is shared with
representatives of the Axioidea, whereas the latter character
is shared with members of the Thomassiniidae (once again
taxa missing from the current molecular analysis). Some
support, however, for this laomediid–thalassinid association,
is provided by studies of larval characters (Sankolli and
Shenoy 1979) and gill-cleaning structures and mechanisms
(Batang and Suzuki 1999; Batang et al. 2001). Although the
latter authors found gill-cleaning characters to be
conservative at the family level, there is still debate over their
utility in phylogenetic studies (Poore 1994; Suzuki and
No unique morphological synapomorphies support the
major monophyletic clade containing the Upogebiidae,
Axianassidae, Thalassinidae and Laomediidae, seen in
Figs 2, 3, even though there is strong molecular support in
both Bayesian and ML analyses. In the morphological
analysis of Poore (1994; Fig. 1C), Upogebia, Thalassina and
Laomedia do not form a distinct clade, but three
morphological characters variously ally these three taxa,
which are part of a monophyletic clade (with other taxa) in
the current molecular analysis. These are (1) the rostrum
augmented with ridges (with the exception of the
Laomediidae), (2) a cylindrical carpus and propodus on
pereopod 1 (a symplesiomorphy shared with some axioids
and some outgroup representatives) and (3) the loss or
reduction of the male first pleopod (except in Thalassinidae).
This last morphological
synapomorphy shared with some callianassids (Poore 1994)
or a symplesiomorphy also shared with some members of the
In summary, the monophyly of the Thalassinidea
suggested from a cladistic analysis of morphological
characters (Poore 1994; Fig. 1C) is only weakly supported by
character is a possible
the 18S gene sequence data (Fig. 2, left side) and
unsupported by the 16S data (Fig. 2, right side) and the
combined analysis of both genes (Fig. 3). Borradaile (1903),
in his paper on thalassinidean classification, did not
implicitly state that the group is monophyletic, but inferred
this in his intuitive genealogical tree (Fig. 1A) and Gurney
(1938; Fig. 1B) advocates paraphyly for the Thalassinidea he
studied. Although a paraphyletic Thalassinidea has some
support in the present molecular analysis, the question of
monophyly for this enigmatic group will require more
analysis of both morphological and molecular data sets to
A dichotomy within the Thalassinidea was previously
suggested by Poore (1994) based on morphological data, but
neither of his major clades corresponds to ours. In contrast,
the dichotomy described by Gurney (1938; Fig. 1B) based on
larval characteristics is reminiscent of the relationships seen
in our 16S tree and combined gene tree (Figs 2 (right side),
3). This closer relationship of the thalassinidean families
Laomediidae and Upogebiidae with the Anomura in
Gurney’s proposed classification (Fig. 1B) is only weakly
supported (under both Bayesian and ML analyses) here and
the anomuran representatives are always part of a larger
decapod sister-clade (Figs 2, 3). Since our analysis only
includes half of the families in the Thalassinidea, further
sampling will be necessary to determine the composition of
these major clades, which appear with regularity in cladistic
analyses of this group.
The three monophyletic superfamilies (Axioidea,
Thalassinoidea and Callianassoidea) indicated in the
classification of Poore (1994) are not maintained as
monophyletic entities in our analysis. In fact, in Poore’s
phylogeny (Fig. 1C), these superfamilies are not adequately
represented by monophyletic clades at the same level either.
This discrepancy was noted by Martin and Davis (2001),
who followed Poore’s revision in their recent work. Until
better taxonomic congruence is achieved, and more of the
11 families are represented in molecular analyses such as the
present study, direct comparisons will remain challenging.
This project was initiated when the senior author (CCT) was
an Australian Research Council Postdoctoral Research
Fellow (1996–1999) at Museum Victoria, Melbourne,
Australia, and the authors acknowledge the assistance of
Gary Poore, Les Christidis, Janette Norman and Megan
Osborne from this institution for project support, access to
laboratory facilities (CCT) and training in molecular
techniques (CCT). The authors thank Cheryl Morrison, Bo
Kong and Todd Oakley for technical support within the
Department of Zoology at Duke University. Also, Kristian
Fauchald and Rafael Lemaitre (Department of Systematic
Biology, National Museum of Natural History, Smithsonian
Institution) are thanked for supporting the Research
846C. C. Tudge and C. W. Cunningham
Associate status of CCT during the completion of the
project. The following colleagues are thanked for collecting,
assistance in collecting, or supplying specimens for this
study: Katrin Berkenbush, David Brewer, Vania Coelho,
Darryl Felder, Alan Harvey, Woody Lee, Rafael Lemaitre,
Cheryl Morrison, Louise Nickel, Philip O’Neil, Gary Poore,
John Salini, Jo Taylor and Stef Zaklan. The improvement of
the manuscript by three anonymous reviewers is gratefully
acknowledged. This is contribution no. 548 of the
Smithsonian Marine Station at Fort Pierce, Florida. The
project was funded by NSF DEB–9615461 (to CWC).
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Manuscript received 30 April 2002; revised and accepted 15 October