JOURNAL OF CRUSTACEAN BIOLOGY, 20, SPECIAL NUMBER 2: 129-149, 2000

PRELIMINARY PHYLOGENETIC ANALYSIS OF GENERIC RELATIONSHIPS WITHIN THE CALLIANASSIDAE AND CTENOCHELIDAE (DECAPODA: THALASSINIDEA: CALLIANASSOIDEA)

Christopher C. Judge, Gary C. B. Poore, and Rafael Lemaitre

(CCT, GCBP) Crustacea Laboratorj', Museum Victoria, P.O. Box 666E, Melbourne. Victoria 3001, Australia; (CCT, RL) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. (conesponding author (CCT) e-mail: [email protected])

ABSTRACT A phylogenetic analysis of 106 representatives in the ihala.<>$inidcan families Ctenochelidae and Callianassidae, based on 93 adult morphological characters, was performed to investigate the inter- relationships between the 2.5 currently recognized genera. The callianidcid. Callianidea typa H. Milne Edwards, was used as the out-group. The two in-group families, Ctenochelidae and Cal- lianassidae, were both shown to be monophyletic. Monophyly of (he four ctenochelid subfamilies (Anacalliacinae. Callianopsinae, Ctenochelinae, and the recently suggested Gourretinae) was not supported. Three callianassid subfamilies (Callianassinae, Cheraminae, and Hucalliacinae) were found to be monophyletic but with species compositions different from those presently understood. The fourth callianassid subfamily (Callichirinae) is paraphyletic and ancestral to the other subfam- ilies. Several species described as species of Callianassa s.l. were found to belong to other genera. Recent diagnosis of several small genera from the Americas and Japan has left larger traditional genera such as Callianassa .\.l. and Cheramus paraphyletic. Glypturus is apparently paraphyletic as presently diagnosed. The taxonomic position of some species remains enigmatic largely because of missing data. A listing of all valid genera and species of Ctenochelidae and C'allianassidae at the time of publication is provided.

Recent studies have greatly increased the lianassa" species should be assigned to other number of known species in the tha- genera. Poore (1994) presented a phylogeny lassinidean families Ctenochelidae (e.g., Mat- of all the families of the Thalassinidea, con- suzawa and Hayashi, 1997; Sakai, 1999), and firmed the monophyly of the infraorder, es- Callianassidae (e.g., Ngoc-Ho, 1991, 1994, tablished that it contained 11 families in three 1995; Sakai, 1992; Manning and Tamaki, superfamilies, and that the families Cal- 1998). The callianassids of the Americas in lianassidae and Ctenochelidae are in fact sis- particular have received much attention (e.g.. ter taxa. His notable changes in the families Manning and Felder, 1995; Lemaitre and Callianassidae and Ctenochelidae were the Felder, 1996; Heard and Manning, 1998; suggestion that the latter is paraphyletic and Hemindez-Aguilera, 1998). Various classifi- the reassignment of the genus Anacalliax cation systems have been proposed for the from the Ctenochelidae to the Callianassidae. decapod infraorder Thalassinidea. De Saint However, many carcinologists have experi- Laurent (1973) defined the Upogebiidae and enced difficulties in applying the classifica- Callianassidae and assigned those species tions of de Saint Laurent (1973) and Manning previously in Callianassa s.l. to six distinct and Felder (1991a) (e.g., Poore, 1975, 1994; genera, Callianassa, Callichirus. Gourretia, Poore and Griffin, 1979; Rabalais etal., 1981; Calliax, Callianopsis, Anacalliax, and created Williams, 1984; Ngoc-Ho, 1994; Sakai, 1999) a seventh new genus Calliapagurops. Later and their global application has been ques- Manning and Felder (1991a), focusing on the tioned. Clearly, additional studies of genera callianassids of the Americas, reviewed the in these two families are still required before genera in the Callianassidae, erected the new true relationships can be ascertained. family Ctenochelidae, and concluded that the A generic-level phylogenetic analysis of large and disparate genus Callianassa s.l. the Ctenochelidae and Callianassidae has not should be restricted to a few species in the previously been attempted although various eastern Atlantic, and that all other "Cal- representatives have been used in investiga-

129 130 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPEOAL NO. 2.2000 tions of relationships between other tha- imens examined (e.g., incomplete specimens or only one lassinidean taxa (Borradaile, 1903; Kensley sex available). and Heard, 1991; Poore, 1994), or between Phylogenetic Analysis thalassinideans and other decapods (Martin The phylogenetic analysis used PAUP version 3.1.1 and Abele, 1986; Scholtz and Richter, 1995). (Swofford, 1993), utilizing a data matrix originating in These analyses were based on morphologi- MacClade version 3 (Maddison and Maddison, 1992). cal characters, but others have used compar- Heuristic search analyses were performed with the fol- lowing options in effect: addition sequence, simple; one isons of 18s rRNA nucleotide sequences tree held at each step during stepwise addition; tree-bi- (Spears and Abele, 1988), neuroanatomy section-reconnection (TBR) branch-swapping performed; (Paul, 1989; Sandeman et ai, 1993), and ul- MULPARS option activated; steepest descent option in- trastructural morphology of spermatozoa and active; branches having maximum length zero collapsed to yield polytomies; topological constraints not enforced; spermatophores (Tudge, 1997). trees unrooted; multistate taxa interpreted as uncertainty; The objective of this paper is to subject character-state optimization, accelerated transformation the current taxonomy of the Callianassidae (ACCTRAN). All characters were unordered, unsealed, and Ctenochelidae to phylogenetic analysis and equally weighted. based on existing, and readily available, char- The large size of the data matrix excluded search strategies other than the heuristic method. acter information. The aim is to highlight taxa The callianideid Callianidea typa was selected as the where taxonomy may not be substantiated by out-group taxon because of its type species status and parsimonious analysis rather than to provide the undisputed, close sister-group relationship to the in- a well-defined alternative classification for group families Ctenochelidae and Callianassidae (Bor- these two families. The current analysis is not radaile. 1903; Kensley and Heard, 1991; Poore, 1994). intended for elucidation of species-level tax- onomic groupings, and associations at this RESULTS level should be treated with caution. The 50% majority rule consensus tree cal- culated from the 1,700 equally parsimonious MATERIALS AND METHODS trees obtained from an heuristic analysis of The taxa included in this analysis are indicated by an the data matrix is shown in Figs. 3 and 4 and asterisk (*) in the list of all valid species in the Iha- is the subject of our discussion. The tree lassinidean families Ctenochelidae and Callianassidae length = 838 steps, and the consistency in- (Appendix 1). This list includes all valid species known to date (April 1999) arranged according to their most re- dex (CI) = 0.905. Some higher taxonomic cent generic and subfamily assignments, and not accord- categories are shown with brackets and ar- ing to their original combinations. The taxa selected for rows (Fig. 3). Clades not supported by 100% the analysis have greater than 50% of their character slates of the trees are indicated (Fig. 4). able to be scored at this time (one exception being "Cat- For brevity we will mostly discuss the sys- lianassa" acutirostella with 55% missing data). The largest genus is Callianassa s.l., not because it is the tematic position of species that appear to lie largest monophyletic group of species but because many outside the genera with which they are cur- authors have used it as a default name until a more sta- rently classified and apparently poly- or para- ble systematic arrangement becomes generally acceptable. phyletic genera. Approximately half (51 %) of In the text we use "Callianassa" in quotes where we sus- pect the generic placement is incorrect. The out-group the species in this analysis have incomplete taxon, Callianidea typa H. Milne Edwards, 1837, (fam- data sets (Fig. 4, and Appendix 3), and there- ily Callianideidae) is also included in Appendix 1. For fore many of the contentious taxa also fall each genus, the type species is indicated, and the source into this category. The average percentage of of morphological information is indicated in square brack- missing data in the matrix is only 11 %, but ets. Appendix 2 is a list describing the 93 adult somatic morphological characters used in the analysis and their in some species this percentage is as high as suggested polarity. There are 56 binary characters and 30% or 50%. The placement of taxa with high 37 multistate characters. There were no nonapplicable proportions of missing data should be viewed characters. This character list is a modified version of that with some reservation. used by Poore (1994). Throughout this paper the terms "antenna I" and "antenna 2" refer to the antennule and antenna, respectively. Figures I and 2 illustrate some of DISCUSSION the characters used. The complete data matrix is provided in Appendix 3. Of the characters employed in this analy- The two families Ctenochelidae and Cal- sis, 13 were considered to be phylogenetically uninfor- lianassidae are both unambiguously mono- mative due to their being autapomorphies or consistent in phyletic, and contrary to Sakai's (1999) view, state across the included taxa. The missing character states (indicated by "?") indicate that the information was un- warrant family rather than subfamily status. able to be obtained from the literature or from the spec- Poore (1994) considered the Callianassidae TTJDGE ETAL: PHYLOCENY OF THE CALLIANASSIDAE AND CTENOTHELIDAE 131

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Fig. I. Diagrammatic representation of selected ctenochelid and callianassid morphological character states. A, B, Lateral view of carapace: A. Clenockeles serrifrons; B, Bijfarius hiformis (both after Manning and Felder, 1991a). C-E, Dorsal view of rostral region and eyestalks: C, Corallichirus xuthus (after Manning and Felder, 1991a); D, Callichirus seilacheri (after Manning and Felder, 1986); E, Lepidophlhalmus jamaicense (after Manning and Felder, 1991a). F-H, Maxilliped 1: F, Caliianidea typa (after Poore, 1997); G, Biffarius delUatulus (after Rodrigues and Man- ning, 1992a); H, Glypturus acamhocliirus (after Biffar, 1971b). I-K, Maxilliped 3: 1, Trypaea australiensis (after Poore and Griffin, 1979); J, "Callianassa" praedatrix (after Sakai, 1988); K, Dawsontus latispina (after Manning and Felder, 1991a). Not (o scale. Abbreviations: ap, anterolateral projections; c, carpus; cd. crista dentata; co, cornea; cp, cardiac prominence; d, dactylus; do, dorsal oval; e, eyestalk; en. endopod; ex, cxopod; i, ischium; It, linea tha- lassinica; m, merus; ms, meral spine: p. propodus; r, rostrum; re, median rostral curina. 132 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2.2000

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Fig. 2. Diagrammatic representation of selected ctenochelid and callianassid morphological character states. A-F, Uropods. lelson, and 6lh abdominal somite: A, Dawsonius latispina (after Biffar, 1971b); B, Ctenacheles maorianus (after Dworschak. 1992); C, Neocallichirus lemailrei (after Manning, 1993); D, Cheramus profundus (after Biffar, 1973); E, Necallianassa berylae (after Heard and Manning, 1998); F, Callichirus islagrande (after Manning and Felder, 1991a). G-I, Appendix intema on plcopods 3-5: G, Cheramus marginatus; H, Callichirus seilacheri; I, Notiax brachyophthalma (all after Manning and Felder, 1991a). J, Posterior margin of carapace and abdominal somites I and 2. K, Lateral view of abdominal segments 1-6 and tailfan. showing position of plumose setal rows on somites 2-6, and dense tufts of lateral setae on somites 3-5. Not to scale. Abbreviations: Al-6, abdominal somites; ai, ap- pendix intema; al, anterolateral lobes; ca, carina; dls, distolateral spine; dp, dorsal plate; en, endopod; ex, exopod; Ip, lateral projection; Is, lateral setal tufts; pi, pleuron; ps, plumose setal row; s, spine; t, telson. TUDOE ETAL: PHYLCXiENY OF THE CALLIANASSIDAE AND CTENOCHELIDAE 133

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Fig. 3. Phylogram of 50% majority rule consensus tree of 1,700 equally parsimonious trees obtained from an heuristic analysis of 107 thalassinidean taxa using 93 morphological characters and using the out-group method. Tree length = 838 steps; consistency index (CI) = 0.905. Some higher taxonomic categories are indicated (arrows and brackets), as are type species (boldface). 134 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000

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fCa|Uanass4 caleoopica CalTianassa joculalnx *Odlianasaapixii *Ca|uanasaa maicfaali Camanasaa ^rniena Gilvostiut setimanus N««rypaca«gas NQtiu DracDyophihalma ^Callianassa mSunuana Callianassa subterranca *Cal|ianassa ambpinensis *CaIuanassa loagicauda *CaIlianassapraedalrix 'CaUianassa propinqua *Callianaua redaneularis 'Chefamus profunda *Cal)ianasu sibogae *Pod ^aucno *Callianassa spinophthalma Qttfainus marnoatus. 'Caifiax novaenitanniae *Ca]liax buUrpba *Calliax aequimana galUax punica ucalliax cearaensis 'Eucalliax jonesi Jiax qui|dracul4 lax mcilhennyi ^CallUxlobi^ . ^Sergio guaiquen *CanVanSua jousseaumei NeocalUchJms rathbunae lima saasuidrensis igrandinuna scac^uale fissHiJlfL s Kmaitrei *Neoca]|idijnis caechabitaior *Neocailichinis nomen •Neoca jjchinis ipi *Neocallicnirus

.„.iichirusfofesti *Callidiinis balssi *Ca])ichin)s guineensis Sergio minm ""gio trilobaivs '^o guassutinga iomericeae ^losulnneus *CoralUanassa collaroy Corallianassa longivenlris Callianassa ranongensis *Caltichinu turneranus *LepidophD)ainiusDOCouni Lratdophthajmus jainaicense. Lepidophthaimus louiuanensu Lepidophthalmus rafai mus sinboia Lepidophthalmus ricfaardi CaTrianuiaairiaeniitaia

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Fig. 4. Cladogram of 50% majority rule consensus tree of 1,700 equally parsimonious trees obtained from an heuristic analysis of 107 thalassinidean taxa using 93 morphological characters and using the out-group method. Tree length = 838 steps; consistency index (CI) = 0.905. Species without a full complement of character states scored (missing data in the matrix) are indicated with an asterisk (*). Clades are supported by 100% of trees unless otherwise indicated. TUDGE ETAL: PHYLOf.ENY OF THE CALLlANASSIDAt AND (TENCX HELIDAE 133 paraphyletic, but the synapomorphies—pres- Callianassinae ence of a dorsal plate on the uropodal exopod (= anterodorsal thickening), absence of dor- The clade of nine species and including the sal carinae on the same uropodal exopod, and type species of Biffarius has 59% support. absence of a cardiac prominence—define the The genus may therefore include "Cal- family. lianassa'' lewtonae, "C." australis, "C." di- aphora, and "Neocallichirus" limosa as well Ctenochelidae as those originally assigned to it. Although The four subfamilies (Anacalliacinae, Cal- "Callianassa " lewtonae has less than half of lianopsinae, Ctenochelinae, Gourretinae) the characters unscored, it shares 45 synapo- (Manning and Felder, 1991a; Sakai, 1999) are morphies with the five Biffarius species. We not monophyletic in the present analysis. The consider at least five significant: absence of subfamily Anacalliacinae, containing the sin- the anterolateral projections on the frontal gle genus Anacalliax, is not a member of the margin of the carapace, equal or subequal Callianassidae, as suggested by Poore (1994). chelipeds in the female, denticulate ischium This analysis clearly shows that the genus of the minor cheliped, and a telson which is (and subfamily) is part of the Ctenochelidae, as long as broad but shorter than the uropods. supporting the placement of Manning and "Callianassa" lewtonae is almost certainly a Felder (1991a) and Sakai (1999). However, species of Biffarius. most of the taxa used to represent this fam- The analysis also suggests that "Neocal- ily in the present analysis have missing data, lichirus" limosa should not be in Neocal- and the intergeneric relationships shown thus lichirus, where it was placed by Sakai (1988) should be treated with caution. and Kazmi and Kazmi (1992), but perhaps The genus Gourretia is paraphyletic in this should be considered as a member of Biffar- analysis, and it has previously been suggested ius. This taxon has a full complement of char- that it comprises more than one genus (Man- acters and differs from the ten other species ning and Felder, 1991a). In the analysis G. of Neocallichirus in presence of a distinct denticulata, the type species of Gourretia, meral hook on the major cheliped, presence does not group with the two other species of of ventral denticles on the ischium of the mi- Gourretia (Figs. 3, 4). The type species dif- nor cheliped, a male first pleopod with the fers from them in having a flatly spinose ros- second segment triangular instead of ovate, trum instead of an upturned one; it has no dis- a male second pleopod vestigial and not cernible antero-lateral projections on the overtly biramous, pleopods 3-5 with the ap- frontal margin of the carapace (Fig. IE); the pendix intema a stubby projection and not endopod on maxilliped 1 is elongate (Fig. IF) embedded into the rami (Fig. 2H, I), and a and not minute (Fig. IG); there is a definite terminally rounded telson (Fig. 2A) and not heel on the propodus of pereiopod 3; the tel- flat or slightly indented (Fig. 2B, C). Place- son is longer than the uropods and has a ment of "Callianassa" limosa in Neocal- rounded distal edge (Fig. 2A) instead of a flat- lichirus is clearly erroneous, and it, "C." aus- tened one; and lastly, transverse setal rows on tralis, and C. diaphora are more probably abdominal somite 6 are absent (Fig. 2K). species of Biffarius. All share similarities in rostral and anterolateral projection morphol- ogy, presence of a distinct meral hook on the Callianassidae major cheliped, telson and uropod character- Three of the four subfamilies (Callianassi- istics, 2nd and 3rd maxilliped morphology nae, Cheraminae, Eucalliacinae) (Manning and male pleopodal morphology. and Felder, 1991a) are monophyletic in this Remaining clades within the (Callianassinae analysis only after reappraisal of their con- are well supported. stituent taxa. The fourth subfamily, Cal- Callianassa subterranea, type species of lichirinae, is paraphyletic. this genus, (currently grouping with C. mau- Restriction of Calliamissa (Manning and ritiana) is at the base of a clade that includes Felder, 1991a) to Callianassa subterranea is other .species of Callianassa s.l. plus the gen- supported by the current analysis and sug- era Notiax, Gilvossius, Neotrypaea, Necal- gests reassessment and reassignment of other lianassa, Nihonotrypaea, and Trypaea. The Callianassa s.l. presence of other species of Callianassa s.l. 136 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000 in other clades reflects the very broad way of multiple spines on the distal and distolat- this genus name has been applied rather than eral edges of the telson (Fig. 2D). Sakai misclassification. The smaller genera are spe- (1970b) placed "Callianassa" spinophthalma cialized derivatives of Callianassa s.s. The in the subgenus Cheramus, and this genus genus Neotrypaea is not monophyletic, with name may be appropriate for all species in N. californiensis the sister taxon of Trypaea this clade. australiensis and N. gigas the sister taxon of Gilvossius setimanus. Both associations are Eucalliacinae well supported. Neotrypaea californiensis dif- A monophyletic clade comprises all the in- fers from N. gigas in that abdominal somite vestigated members of the subfamily Eucal- 2 is greater in length than abdominal somite liacinae, with the exception of Calliax 6 (Fig. 2K) and not equal to it; abdominal tooradin. Although nearly a third of the char- somites 3-S have dense tufts of lateral setae acter states are missing for Calliax tooradin, (Fig. 2K); antenna 1 peduncle is slightly it differs from other members of its genus in longer than antenna 2 peduncle, rather than nine characters (see Appendix 2, characters subequal; antenna 1 also has a reasonably Ih, 5b, e, 7m-o, 8b-d). This species may not dense brush of ventrally directed setae; a belong in Calliax, as previously suggested by more prominent crista dentata is present (Fig. Poore and Griffin (1979) and de Saint Lau- 2J); ischium of the minor cheliped lacks ven- rent and Manning (1982). Calliax punica (the tral denticulation; propodus of pereiopod 3 only member of this genus without missing has a distinct heel on the proximal comer of data) appears as the sister taxon of Eucalliax the lower margin whereas the same in N. gi- cearaensis. The association has only 64% gas is more oval in shape; male pleopod 1 is support, and a review of characters indicates present, but is unusually absent in N. gigas; that C. punica has no special affinities with female pleopod 1 is 2-segmented rather than Eucalliax that it does not share with its con- 3-segmented; pleopods 3-5 have an embed- geners. It differs from E. cearaensis in eight ded appendix intema (Fig. 21), as opposed to characters associated with abdominal somite a stubby projecting one (Fig. 2H); and finally, setation, branchial formulae, and antennal, the telson has equal dimensions in N. cali- mouthpart, and pleopod morphology. How- forniensis and is broader than long in N. gigas. ever, Felder and Manning (1994) noted that "Callianassa" amboinensis sits in a clade C. punica shares the presence of one or two remote from other taxa and may warrant sep- elongate setae on the propodus of pereiopod arate generic status. 4 (not a character used in this analysis) with all known species of Eucalliax. The close re- Cheraminae lationship between E. jonesi and E. quadra- A well-supported clade (here suggested to cuta was previously documented by Heard constitute the subfamily Cheraminae) is com- (1989), as were their differences from Calliax posed of six species of "Callianassa," Poti lobata and others. Differentiation of these two gaucho, Cheramus profunda, and C mar- genera remain enigmatic. ginatus (Figs. 3, 4). All except the last are missing significant character information Callichirinae (13% to 39%). "Callianassa" longicauda, The paraphyletic Callichirinae are the least "C. " praedatrix, and "C. " propinqua (all resolved subfamily and contain the only three from the Southeast Asia region) share a sim- polytomies in the consensus tree. ilar rostral morphology, antennal proportions, Sergio appears to be a paraphyletic genus denticulation of the ischium and merus of the with its seven species occurring on four major cheliped, and a distinctive tooth on the clades. Character information is almost com- distal margin of the merus of the 3rd maxil- plete with only Sergio guara and 5. guaiqueri liped (Fig. 2J). The importance of this latter each missing three characters associated with character in linking these three taxa has pre- female pleopod morphology. The four-way viously been recognized (Sakai, 1988). Six of split of the genus may be caused by a large the nine species in this clade {"Callianassa " polytomy (77% supported) at this point in the praedatrix, "C." propinqua, "C." rectangu- tree obscuring any characters that may unite laris, "C." sibogae, Cheramus profunda, and the species. The three Brazilian species (5. C. marginatus) are linked by the possession guara, S. mirim, and S. guassutinga) occur TUDGE ETAL: PHYLOGENY OF THE CALLIANASSIDAE AND CTENOCHEUDAE 137

in three clades, while three species from "Callianassa" ranongensis and "Cal- Florida (5. trilobate, S. guassutinga, and S. lichirus" turneranus belong in the clade of mericeae) occur in two clades. The individ- seven of the eight published species of Lep- ual species groups can be identified by sets idophthalmus, for they share a large propor- of characters, but these are not exclusive to tion of their characters with members of this each of the clades. Sergio guara and S. guai- genus. Both can confidently be transferred to queri can be linked by eight characters (but Lepidophthalmus. are not sister taxa in this analysis); 5. mirim The base of the C^allianassidae is poorly re- and 5. trilobata are linked by five characters; solved as most of its taxa have a high pro- and S. guassutinga, S. mericeae, and S. sul- portion of missing data (14% to 37%). "Cal- fureus are linked by two characters. The lat- lianassa " tridentata, with a full complement ter two Sergio clades have 100% support, and of character states, is sister taxon of the two the close association of S. guassutinga, S. investigated species of Corallichirus (C. mericeae, and S. sulfureus was supported by placidus and C xuthus). "C." tridentata Manning and Felder (1995) and Lemaitre and shares 71 characters with these two Coral- Felder (1996). Sergio may be a paraphyletic lichirus species, and it is possible that "C." genus, but it is possible that their uniting tridentata is a fourth species of Corallichirus. characters are outweighed by convergences Ba&ally associated with Corallichirus is elsewhere in this large cladogram. Glypturus martensi. With only 83% support A monophyletic clade (100% supported) for the association and with a significant pro- containing 10 species oi Neocallichirus also portion of the character states still missing, includes "Callianassa " jousseaumei. Nearly it is difficult to decide if this taxon is simply half of the character data are missing for this reflecting the lack of characters to link it with species. However, C. jousseaumei shares 39 its congeners. It shares many character states of the remaining characters with Neocal- with Corallichirus (including "Callianassa" lichirus, of which four, associated with max- tridentata) and differs in seven characters illiped 3 morphology and shape of the uropo- from the majority of Glypturus, of which only dai endopods, are not shared with Callianassa two are shared with Corallichirus. The seven S.S. Biffar (1971b) noted that "C." character states in which G. martensi differs jousseaumei showed numerous morphologi- from the remaining species in Glypturus are: cal similarities with Neocallichirus grandi- terminally rounded eyestalks (Fig. IC) in- mana (as Callianassa branneri) and ^V. indica stead of pointed (Fig. ID, E); antenna 1 pe- (as Callianassa indica), so its membership duncle is equal to or larger than antenna 2 pe- of this genus seems certain. duncle rather than shorter; exopod of maxil- Similarly, "Callianassa" kraussi shares a liped 1 appears to be 2 or 3 segmented (Fig. significant proportion of its characters with IG) and not consisting of a single segment the members of Callichirus (seven exclu- (Fig. IH); unlike the other species, the merus sively), and the cladogram places it in this of maxilliped 3 projects beyond the articula- genus. tion with the carpus (Fig. 21, J); ventral mar- De Saint Laurent and Le Loeuff (1979) gin of the merus of the minor cheliped has a subdivided the then large genus Callichirus convex margin (shared with G. mucronata) (15 species) into six categories, some upheld instead of a straight or toothed margin; fin- in the current analysis. Manning and Felder gers of the minor cheliped are longer than the (1986) redefined Callichirus and restricted its propodal palm and not shorter than or equal species to C. major, C. islagrande, C. to it; and finally, propodus of pereiopod 3 seilacheri, and C adamas. They stated ". . . would appear to lack the distinctive heel seen these four species comprise a distinctive in the other five species (although this char- group of species within the Callianassidae acter state is ambiguously scored as oval or recognizable as a distinct genus, Callichirus." heeled in the data matrix. Appendix 3). Glyp- (Manning and Felder, 1986: 439). This analy- turus appears polyphyletic with or without the sis supports their statement, showing a dis- inclusion of "G." martensi. Glypturus tinct clade of these four taxa within Cal- karumba and G. mucronata (both Australian lichirus. and Southeast Asian s]>ecies) form a small The two species of Corallianassa are sis- subclade separated from their congeners, a re- ter taxa, and the genus is supported. lationship supported by Poore and Griffin 138 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2.2000

(1979). These two species share a reduced but nosed, or as a large polyphyletic genus from spinose rostrum, absence of anterolateral pro- which specialized clades (e.g., Trypaea, jections on the carapace, and absence of a Neotrypaea) have derived. scaphocerite on antenna 2. All are characters Further progress on elucidating generic that differentiate them from the other Glyp- structure and relationships in these two fam- turus species studied (Poore and Suchanek, ilies will be made by completing the data ma- 1988). trix, addition of new characters and taxa, and "Callianassa " audax, at the base of the en- concentration on the better-defined clades in tire callianassid clade, may simply be a case the tree presented here. Examination of spec- of there being too much missing data to be imens of type species of many of the included effectively placed in the family. Of the 59 taxa should add most of the character data characters scored for "C." audax, only four that is currently missing from the matrix. This differ from the majority of the species cur- character-state information, with the addition rently in Callianassa s.l. Its basal position in of new characters and the inclusion of newly the familial clade is enigmatic. described taxa, would form the basis for a Paraglypturus calderus appears as the sis- more complete and rigorous analysis of these ter taxon to all other callianassids. It shares two thalassinidean families. Subsequent re- (among other characters) the presence of the analysis of the individual clades presented in anterodorsal thickening (dorsal plate) on the this consensus tree may lend support to the uropodal exopod with all other members of relationships shown and increase confidence the family but possesses the primitive char- in the systematic changes suggested. acter of a large exopod on the third maxil- We do not believe this cladogram is more liped. This latter character is observed in the than indicative of generic relationships, but out-group taxon, Callianidea typa, the hope it is a stimulus for further investigation. ctenochelids Gourretia and Anacalliax, and the callianassid Calliax tooradin. ACKNOWLEDGEMENTS CONCLUSION The authors wish to acknowledge the assistance of B. Kensley and the late R. B. Manning (National Museum In spite of the high frequency of missing of Natural History, Smithsonian Institution) and the late data (still to be obtained) the cladogram can A. B. Williams (National Oceanographic and Atmospheric be interpreted in a meaningful way which Administration, National Marine Fisheries Service) for supports the existing systematic arrange- providing access to literature and specimens and advice on character-state assignments. Thanks are also extended ments. to K. Fauchald (National Museum of Natural History, In the Ctenochelidae there is little justifi- Smithsonian Institution) for facilitating the senior author's cation for the subfamily arrangements re- access to the Department of Invertebrate Zoology and its cently proposed, and the genus Gourretia ap- collections as a visiting researcher on two separate oc- pears to be polyphyletic. casions in 1998 and 1999. The senior author was sup- ported by an Australian Research Council Postdoctoral The traditional subfamily structure of the Research Fellowship (1996-1999). This paper is dedi- Callianassidae is revealed in the cladogram, cated to R. B. Manning, in recognition of his many out- and it is no surprise that one of the four sub- standing contributions to the taxonomy and systematics families (defined in terms of plesiomorphies) of the Thalassinidea and to the callianassoids of the Amer- icas in particular. Credit is due also to his wife, Lilly King is paraphyletic. The genus name Callianassa Manning, for preparing the illustrations that accompany has been used loosely in the past, whereas many of her husband's papers. smaller genera remained poorly defined or es- pecially restricted to small, tightly diagnosed groups of species. There are two conse- LITERATURE CiTED quences of this. One is that our cladogram Barnard, K. H. 1950. Descriptive catalogue of South suggests the placement of several species of African decapod Crustacea (crabs and shrimps).—An- nals of the South African Museum 38: 1-864. Callianassa s.l. in alternative monophyletic Bate, C. S. 1888. Report on the Crustacea Macrura col- genera. This may necessitate the rediagnosis lected by H.M.S. Challenger during the years and/or subdivision of these genera. The other 1873-76.—Report on the Scientific Results of the Voy- consequence is that Callianassa can be age of H.M.S. Challenger.—Zoology 24: 1-192. Biffar, T. A. 1970. Three new species of callianassid treated either as a very small genus (e.g.. shrimp (Decapoda, Thalassinidea) from the Western At- Manning and Felder, 1991a), in which case lantic.—Proceedings of the Biological Society of Wash- several new smaller genera need to be diag- ington 83: 35-50. TUDGE ET AL: I'HYLOGENY OF THE CALLIANASSIDAK AND CTENOCHliLIDAE 139

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. 1979. Sur la classification el la phylogdnie des Scholtz, G., and S. Richter. 1995. Phylogenetic system- Thalassinides: Definitions de la supeifamilles des Ax- atics of the reptantian Decapoda (Crustacea, Malacos- ioidea, de la sous-famille des Thomassiniinae et Deux traca).—Zoological Journal of the Linnean Society 113: genres nouveaux (Crustacea Dccapoda).—Comptes ren- 289-328. dus de I'Acad^mie de Science, Paris (D) 288:1395-1397. Spears, T. and L. G. Abele. 1988. Phylogenetic rela- , and B. Bozic. 1972. Diagnoses et tableau de tionships of crustacean anomurans based on partial 18s determination des callianasses de I'Atlantique notd ori- rRNA nucleotide sequences.—American Zoologist 29: ental et de Mediterranee (Crustacea, Decapoda, Cal- 25A [Abstract.] lianassidae).—Thalassia Jugoslavica 8: 15-40. Stimpson, W. 1866. Descriptions of new genera and . and P. Le Loeuff. 1979. Crustac^s D6capodes species of macrurous Crustacea from the coasts of Thalassinidea. I. Upogebiidae et Callianassidae.—R€- North America.—Proceedings of the Chicago Academy sultats Scientifiques des Campagncs de la CALYPSO of Science 1: 46-68. 11:29-101. Swofford, D. L. 1993. PAUP: Phylogenetic analysis us- -, and R. B. Manning. 1982. Calliax punica, es- ing parsimony, version 3.1.1 (computer program dis- pdce nouvelle de Callianassidae (Crustacea, Decapoda) tributed by the Illinois Natural History Survey, Cham- des eaux mediterraneennes.—QuadernI del Laborato- paign, Illinois, U.S.A). rio di Tecnologia della Pesca 3: 211-224. Tirmizi, N. M. 1977. A redescription of the holotype Sakai, K. 1966. On Callianassa (Callicliirus) novaebri- species of Callianassa mucronata Strahl, 1861 (Die- lanniae Borradaile (Thalassinidea, Crustacea) from capoda, Thalassinidea).—-Crustaceana 32: 21-26. Japan.—Journal of the Faculty of Agriculture, Kyushu Tudge, C. C. 1997 Phylogeny of the Anomura (De- University 14: 161-171. capoda, Crustacea): spermatozoa and spermatophore . 1967a. Three new species ofThalassinidea (De- morphological evidence.—Contributions to Zoology capoda, Crustacea) from Japan.—Records of the Crus- 67: 125-141. tacea of Japan 3: 39-51. TUrkay, M., and K. Sakai. 1995. Decapod crustaceans . 1967b. Three new species ofThalassinidea (De- from a volcanic hot spring in the Marianas.—Senck- capoda, Crustacea) from South—West Japan.—Publi- enbergiana Maritima 26: 25-35. cations of the Seto Marine Biological Laboratory 15: Williams, A. B. 1984. Shrimps, lobsters, and crabs of the 319-328. Atlantic coast of the eastern United States, Maine to . 1970a. Supplementary description of Cal- Florida. Smithsonian Institution Press, Washington, lianassa (Callichirus) tridentata von Martens (Crus- D.C. 550 pp. tacea, Thalassinidea). Noona Dan Papers No. 97.— Publications of the Seto Marine Biological Laboratory RECEIVED: 21 April 1999. 17: 393^01. ACCEPTED: 21 December 1999. . 1970b. A small collection of thalassinids from the waters around Tsushima Islands, Japan, including NOTE ADDED IN PROOF a new species of Callianassa (Crustacea, Anomura).— Publications of the Seto Marine Biological Laboratory Three new genera and eight new species have been added 18: 37^7. to the callianassid fauna since this paper was first sub- . 1983. On a new species of the genus Cal- mitted: Grynaminna Poore, 2000 (type species Gry- lianassa (Crustacea, Decapoda) from Thailand.—Re- naminna tamakii Poore, 2000); Podocallichirus Sakai, searches on Crustacea 12: 111-115. 1999a (type species Callianassa madagassa Lenz and . 1984. Some thalassinideans (Decapoda: Crus- Richters, 1881): Pseudobiffarius Heard and Manning, tacea) from Heron Is., Queensland, Eastern Australia, 2000 (type species Pseudobiffarius caesari Heard and and a new species of Gourretia from East Africa.—The Manning, 2000); Callianassa gruneri Sakai. 1999a; Cal- Beagle 1:95-108. lianassa ngochoae Sakai, 1999a; Callianassa poorei . 1987a. On Callianassa ranongensis (Tha- Sakai, 1999b (probably a species of Biffarius); Cal- lassinidea: Decapoda: Crustacea) from Halmahera, In- lianassa whitei Sakai. 1999a; Calliax doerjesli Sakai, donesia.—Reports of the Usa Marine Biological Insti- 1999a; Grynaminna tamakii Poore, 2000; Neocallianassa tute, Kochi University 9: 45-49. kempi Sakai, 1999a: Neocallichirus raymanningi Blanco . 1987b. Two new Thalassinidea (Crustacea: De- Rambla and Lemaiire, 1999. capoda) from Japan, with the biogeographic distribu- Sakai's (1999a) synopsis of the Callianassidae has tion of the Japanese Thalassinidea —Bulletin of Ma- reached conclusions about the taxonomy of the family rine Science 41: 296-308. very different from ihose of our phylogenetic analysis and 1988. A new genus and five new species of Cal- from those recently proposed (e.g.. Manning and Felder, lianassidae (Crustacea: Decapoda: Thalassinidea) from 1991a: Poore, 1994). Sakai recognizes four subfamilies Northern Australia.—The Beagle 5: 51-69. of which only the Eucalliacinae is similar to that sub- . 1992. Notes on some species of Thalassinidea family in our analysis. Within this subfamily he has syn- from French Polynesia (Crustacea: Decapoda).— onymised Eucalliax with Calliax and included Paraglyp- Senckenbergiana Maritima 22: 211-216. lurus. The four species assigned to this genus include two . 1999. Redescription of Ctenocheles balssi (Calliax looradin and Paraglypturus calderus) which our Kishinouye, 1926, with comments on its systematic po- analysis places basally within the Callianassidae clade sition and establishment of a new subfamily Courreti- and a third (C. novaehntlaniae) which clusters with other inae (Decapoda, Callianassidae).—Crustaceana 72: species of Calliax. 85-97. The second of Sakai's subfamilies Calliapaguropsinae, Sandeman, D. C, G. Scholtz. and R. E. Sandeman. 1993. for Calliapaguropsis i.han-oti only, is an unusual cal- Brain evolution in decapod Crustacea.—The Journal of lianassid, hitherto poorly described and not included in Experimental Zoology 265: 112-133. our analysis. 142 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000

The third subfamily, Anacalliacinae, was erected in Appendix I. List of valid species in the Ctenochelidae Ctenochelidae by Manning and Feldcr and treated within and Callianassidae, arranged according to their most re- Callianassidae by Poore (1994). It contains a single genus cent revisions. * indicates taxa used in this analysis; which our analysis suggests is a highly derived clade of sources of information, reference and/or musuem collec- Ctenochelidae. tions are given in square brackets. Species in new com- Sakai (1999a) treats all remaining callianassids as binations are in curly brackets. Collections are from members of the Callianassinae. He did not recognise USNM (National Museum of Natural History, Washing- Cheraminae or Callichirinae. The former is a small, ton, D.C., U.S.A.) and NMV (Museum Victoria, Mel- clearly monophyletic clade. The latter is probably para- bourne, Australia). phyletic but clearly defined (Poore, 2000). Sakai's treatment of genera diverges from the results Family: Callianideidae Kossmann, 1880 of our cladistic analysis. We agree on the monophyly of Genus Callianidea Kossmann, 1880 Neocalitchirus but not on the assignment of some species *Callianidea typa Milne Edwards, 1837 TYPE SPECIES of Glypturus to it. His synonymy of Corallianassa with [Poore, 1994, 1997; Kensley and Heard, 1991; USNM] Glypturus i.s not supported in our phylogeny. We agree too on the monophyly of Lepidophthalmus. Sakai's divi- Family: Ctenochelidae Manning and Felder, 1991a sion of Callichirus into two genera, Callichirus and Subfamily Anacalliacinae Manning and Felder, 1991a (as Podocallichirus, is not inconsistent with our analysis but Anacalliinae) is not supported by a clear dichotomy between species Genus Anacalliax de Saint Laurent, 1973 groups. *Anacalliax agassizi (Biffar, 1971) [Biffar, 1971a) All remaining ten genera were synonymised by Sakai *Anacalliax argentinensis (Biffar, 1971) TYPE SPECIES into Callianassa. While he recognised that some genera [Biffar, 1971a; Manning and Felder, 1991a; USNM] are "fundamentally different" from the type species of Callianassa or cited "the traditional generic criterion" to Subfamily Callianopsinae Manning and Felder, 1991a define this genus and highlighted differences in some Genus Callianopsis de Saint Laurent, 1973 characters, he appears so influenced by similarities in oth- *CalUanopsis goniophthalma Rathbun, 1902 TYPE ers and the presence of "intermediate forms" that syn- SPECIES [Manning and Felder, 1991a; USNM) onymy seemed the simplest solution. Results of our phy- logenetic analysis do not indicate monophyly of a large Subfamily Ctenochelinae Manning and Felder, 1991a genus Callianassa but rather numerous monophyletic Genus Ctenocheles Kishinouye, 1926 clades of few or several species, each supported by one *Clenncheles halssi Kishinouye, 1926 TYPE SPECIES or more synapomorphics. Generic names must be found [Kishinouye, 1926; Matsuzawa and Hayashi, 1997; for these, and we conclude that those erected so far should Sakai, 1999] stand for the time being. Ctenocheles collini Ward, 1945 *Ctenocheles holthuisi Rodrigues, 1978 [Rodrigues, Blanco Rambia, J. P, and R. Lemaitre. 1999. Neocal- 1978] lichirus raymanningi, a new species of ghost shrimp Ctenocheles leviceps Rabalais, 1979 from the northeastern coast of Venezuela (Crustacea: *Ctenocheles maorianus Powell, 1949 [Powell, 1949; Decapoda: Callianassidae).—Proceedings of the Bio- Dworschak, 1992; USNM] logical Society of Washington 112: 768-777. Ctenocheles serrifrons Le Loeuff and Intes, 1974 Heard, R., and R. B. Manning. 2000. A new genus and Genus Dawsonius Manning and Felder, 1991a species of ghost shrimp from Tobago, West Indies *Dawsonius latispina (Dawson, 1967) TYPE SPECIES (Crustacea: Decapoda: Callianassidae).—Proceedings [Biffar, 1971b; Manning and Felder, 1991a; USNM] of the Biological Society of Washington 113: 71-77. Genus Gourretia de Saint Laurent, 1973 Poore, G. C. B. 2000. A new genus and species of cal- Gourretia barracuda Le Loeuff and Intfes, 1974 lianassid ghost shrimp from Kyushu, Japan (Decapoda: *Gourretia hiffari Blanco Rambia and Linero Arana, 1994 Thalassinidea).—Journal of Crustacean Biology 20 [Blanco Rambia and Linero Arana, 1994; USNM] (Special no. 2): 150-156. Gourretia coolihah Poore and Griffin, 1979 Sakai, K. 1999a. Synopsis of the family Callianassidae, Gourretia crosnieri Ngoc-Ho, 1991 with keys subfamilies, genera and species, and the de- *Gourretia denticulata (Lutze, 1937) TYPE SPECIES [de scription of new taxa (Crustacea: Decapoda: Tha- Saint Laurent and Bozic, 1972; Le Loeuff and Int^s, lassinidea).—Zoologische Verhandelingen, Leiden 326: 1974; USNM] 1-152. Gourretia lahouensis Le Loeuff and Intfes, 1974 . 1999b. A new species, Callianassa poorei, sp. *Gourretia laresi Blanco Rambia and Linero Arana, 1994 nov. (Decapoda: Crustacea: Callianassidae) from Tas- [Blanco Rambia and Linero Arana. 1994; USNM] mania.—Journal of the Marine Biological Association Gourretia manihinae Sakai, 1984 of the United Kingdom 79: 373, 374. Genus Paracalliax de Saint Laurent, 1979 *Paracalliax hollorei de Saint Laurent, 1979 TYPE SPECIES [de Saint Laurent and Le Loeuff, 1979]

Family: Callianassidae Dana, 1852 Subfamily Callianassinae Dana, 1852 Genus Biffarius Manning and Felder, 1991a *Biffarius arenosa (Poore, 1975) [Poore, 1975; Poore and Griffin, 1979; NMV; USNM] *Biffarius australis (Kensley, 1974), new combination TUDGE ETAL: PH YLOCiENY OF THE CALLIANASSIDAI; AND CTENOCHELIDAE 143

*Biffarius bifomis (Biffar, 1971) TYPE SPECIES [Bif- Callianassa maxima A. Milne-Edwards, 1870 far, 1971a; Williams, 1984; Manning and Felder, 1991a; Callianassa modesta de Man, 1905 USNM) Callianassa nakasonei Sakai, 1967 [Sakai, l%7a] *Biffarius ceramica (Fullon and Grant, 1906) [Fulton and Callianassa par\'a Edmondson, 1944 Grant, 1906; Poore and Griffin, 1979; NMV; USNM] Callianassa par\>ula Sakai, 1988 Biffarius dehilis Hemindez-Aguilera, 1998 'Callianassa pixii Kensley, )975 [Kensley, 1975] *Bijfarius delkatulus Rodrigues and Manning, 1992 [Ro- Callianassa pontica Czemiavsky, 1884 drigues and Manning, 1992a; USNM] {'Callianassa praedatrix de Man, 1905 [de Man, 1928; *Biffarius diaphora (Le Loeuff and Intes, 1974), new Sakai. 1988; Ngoc-Ho, 1994]-removed to Cheramus) combination {'Callianassa pwpinqua de Man, 1905 [de Man, 1928; * Biffarius fragilis (Biffar, 1970) [Biffar, 1970, 1971b; Ngoc-Ho, l991]-removed to C/i^ramui} USNMI Callianassa pugnatrix de Man, 1905 *Biffarius lewtonae (Ngoc-Ho, 1994), new combination Callianassi, pygmaea dc Man, 1928 *Biffarius limosa (Poore, 1975). new combination {'Callianassa rammgensis Sakai, 1983 [Sakai, 1983, Genus Callianassa Leach, 1814 1987a; USNM]-removed to Lepidophthalmus) Callianassa ahdominalix White, 1847 {'Callianassa rectangularis Ngoc-Ho, 1991 [Ngoc-Ho, 'Callianassa acutimslella Sakai. 1988 [Sakai, 1988] 1991 [-removed to Cheramus) *CalUanassa amhoinensis de Man. 1888 [de Man, 1928; Callianassa nisae Nobili, 1906 Poore and Griffin, 1979; Sakai, 1984; Ngoc-Ho, 1991] Callianassa nnundicauikita Stebbing, 1902 Callianassa assimilis (de Man, 1928) {'Callianassa sihogae de Man, 1905 [de Man, 1928; 'Callianassa audax de Man, 1911 [de Man, 1928; Ngoc-Ho, 19941-remiived to Cheramus) Dworschak, 1992] {'Callianassa spinophthalma Sakai, 1970 [Sakai, {'Callianassa auslralis Kensley, 1974 [Kensley, 1974; 1970b]-removed to Cheramus) de Saint Laurent and Le Loeuff, 1979; USNM]-re- 'Callianassa suhlerranea (Montagu, 1808) TYPE moved to Biffarius) SPECIES [de Man, 1928; de Saint Laurent and Bozic, Callianassa houvieri Nobili, 1904 1972; de Saint Laurent and Le Loeuff, 1979; Manning Callianassa hrevicaudata (A. Milne-Edwards. 1870) and Felder. 1991a; USNM] 'Callianassa caledonica Ngoc-Ho, 1991 [Ngoc-Ho, Callianassa tonkinae Grebenjuk, 1975 1991] ('Callianassa tridentaki Von Martens, 1868 [de Man, Callianassa calmani Nobili, 1904 1928; Sakai, 1970a; USNM]-removed to Coral- Callianassa Candida (Olivi, 1792) lichirus) Callianassa carinaedoris While, 1847 'Callianassa ryrrhena (Petagna, 1792) [de Saint Laurent Callianassa chilensis A. Milne-Eklwards, I860 and Bozic, i972; Hollhuis, 1991; USNM] Callianassa convexa de Saint Laurent and Le Loeuff, Callianassa variahilis Hdmondson, 1944 1979 Callianassa' vigilax de Man, 1916 Callianassa coutierei (Nobili. 1904) Callianassa winslowi (Edmondson, 1944) Callianassa crislala (Borradaile, 1910) Genus Calliapagurops de Saint Laurent, 1973 {'Callianassa diaphora Le Loeuff and Int^s. 1974 [Le Calliapagurops chanoti de Saint Laurent, 1973 TYPE Loeuff and Intfes, 1974; de Saint Laurent and Le Loeuff, SPECIES 1979]-removed to Biffarius) Genus Gilvossius Manning and Felder, 1991b 'Callianassa filholi A. Milne-Edwards. 1878 [Chilton, 'Gilvossius setinuinus (DeKay, 1844) TYPE SPECIES 1906; Devine, 1966; NMV; USNMj [Manning and Felder. 1991b; USNM] Callianassa gilchristi Barnard, 1947 Genus Necallianassa Hoard and Manning, 1998 Callianassa grandidieri Coutifire. 1899 Necallianassa acanthura (Ciu'oli, 1946) Callianassa gravieri Nobili, 1906 'Necallianassa hervlae Heard and Manning, 1998 TYPE Callianassa intermedia de Man, 190.5 SPECIES [Heard and Manning, 1998] 'Callianassa joculatrix de Man, 190.*) [Poore and Grif- Necallianassa truncata (Giard and Bonnier, 1890) fin, 1979; Ngoc-Ho, 1991; USNM] Genus Neotrypaea Manning and Felder, 1991a {'Callianassa jousseaumei Nobili, 1904 (de Man, 1928; Neotrypaea hiffari (Viohhuis. 1991) Dworschak, 1992]-removed to SeocalUchirus) 'Neotrypaea taliforniensis (Dana, 1854)TYPE SPECIES Callianassa kewalramanii (Sankolli, 1971) [Holthuis. 1991; Manning and Felder, 1991a; {'Callianassa kraussi Stebbing, 1900 [Barnard, 1950; Dworschak. 1992; USNMj Holthuis, 1991; USNM|-removed to Callichirus) 'Neotnpaea gigas (Dana. 1852) [Holthuis, 1991; {'Callianassa lewtonae Ngoc-Ho. 1994 [Ngoc-Ho, Dworschak, 1992; USNMj 1994]-removed to Biffarius) Neotrypaea rochei (Bouvier, 1895) Callianassa lignicola Alcock and Anderson. 1899 Neotrypaea uncinala (H. Milne Edwards, 1837) Callianassa lohetohensis de Man, 1903 Genus Nihonotrypaea Manning and Tamaki, 1998 {'Callianassa longicauda Sakai, 1967 [Sakai, 1967b]-re- 'Nihonotrypaea harmandi (Bouvier, 1901) [de Man, moved to Cheramus) 1928; Manning and Tamaki, 1998; USNM] Callianassa madagassa Lenz and Richters, 1881 'Nihonotrypaea japonica (Ortmann, 1891) TYPE Callianassa maldivensis Borradaile, 1904 SPECIES (de Man, 1928; Manning and Tamaki, 1998; 'Callianassa marchali Le Loeuff and Inlds. 1974 [Le USNM] Loeuff and Intes, 1974; de Saint Laurent and Le Ixjeuff, 'Nihonotrypaea petalura (Stimpson, 1860) [Holthuis, 1979] 1991; USNM] Callianassa masoomi (Tirmizi, 1970) Genus Notiax Manning and Felder, 1991a 'Callianassa mauriliana Miers, 1882 [de Man, 1928; 'Notiax brachyophthalma "A. Milne-Edwards, 1870) Kensley, 1975] TYPE SPECIES [Manning and Felder, 1991a; USNM) 144 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2. 2000

Genus Poti Rodrigues and Manning, 1992 *Lepidophlhalmus bocourti (A. Milne-Edwards, 1870) {*Poii gaucho Rodrigues and Manning, 1992 TYPE TYPE SPECIES [Lemaitre and Ramos, 1992; USNM] SPECIES [Rodrigues and Manning, 1992b; Lepidophthalmus eiseni (Holmes, 1904) USNMl-removed to Cheraminae} 'Lepidophthalmus jamaicense (Schmitt, 1935) [Ro- Genus Trypaea Dana, 1852 drigues, 1971; Manning and Felder, 1991a; Felder and *Trypaea australiensis (Dana, 1852) TYPE SPECIES Manning, 1997; USNM] [Poore and Griffin, 1979; Holthuis, 1991; Poore, 1994; 'Lepidophthalmus louisianensis (Schmitt, 1935) NMV; USNM) [Dworschak, 1992; Felder and Rodrigues, 1993; USNM] 'Lepidophthalmus rafai Felder and Manning, 1998 Subfamily Callichirinae Manning and Felder, 1991a [Felder and Manning, 1998; USNM] Genus Callichirus Stimpson, 1866 'Lepidophthalmus ranongensis (Sakai, 1983), new com- *Callichirus adamas (Kensley, 1974) [Kensley, 1974; de bination Saint Laurent and Le Locuff, 1979] 'Lepidophthalmus richardi Felder and Manning, 1997 *Callichirus balssi (Monod, 1933) [Le Loeuff and Intis, [Felder and Manning, 1997; USNM] 1974] 'Lepidophthalmus sinuensis Lemaitre and Rodrigues, *CaUichirusforesli Le Loeuff and Intfes, 1974 [Le Loeuff 1991 [Lemaitre and Rodrigues, 1991; Felder and Man- and Intds, 1974] ning, 1997; USNM] Callichirus garthi (Retamal, 1975) 'Lepidophthalmus siriboia Felder and Rodrigues, 1993 *Callichirus guineensis (de Man, 1928) [de Man, 1928; [Felder and Rodrigues, 1993; USNM] Le Loeuff and Int^s, 1974] 'Lepidophthalmus turneranus (White, 1861), new com- Callichirus iniesi de Saint Laurent and Le Loeuff, 1979 bination *Callichirus islagrande (Schmitt, 1935) [Manning and Genus Neocallichirus Sakai, 1988 Felder, 1986; NMV; USNM] 'NeocalUchirus cacahuate Felder and Manning, 1995 *Callichirus kraussi (Stebbing, 1910), new combination [Felder and Manning, 1995; USNM] *Catlichirus major (Say, 1818) TYPE SPECIES [de Man, 'Neocallichirus caechabitator Sakai, 1988 [Sakai, 1988] 1928; Rodrigues, 1971; Williams, 1984; Manning and 'Neocallichirus darwinensis Sakai, 1988 [Sakai, 1988] Felder, 1986; NMV; USNM] Neocallichirus denticulalus Ngoc-Ho, 1994 Callichirus monodi de Saint Laurent and Le Loeuff, 1979 'Neocallichirus grandimana (Gibbes, 1850) [Manning Callichirus pentagonocephala (Rossignol, 1962) and Felder, 1991a; Lemaitre and Ramos, 1992; Felder *Callichirus seilacheri (Bott, 1955) [Manning and Felder, and Manning, 1995; USNM] 1986, 1991a; USNM] 'Neocallichirus horneri Sakai, 1988 TYPE SPECIES Callichirus tenuimanus de Saint Laurent and Le Loeuff, [Sakai, 1988] 1979 'Neocallichirus indica de Man, 1905 [de Man, 1928; {*Callichirus tumeranus (White, 1861) [de Man, 1928; Kensley, 1975] Le Loeuff and Int^s, 1974; de Saint Laurent and Le 'Neocallichirus jousseaumei (Nobili, 1904), new com- Loeuff, 1979; Holthuis, 1991]-removed to Lepidoph- bination thalmus } 'Neocallichirus lemaitrei Manning, 1993 [Manning, Genus Corallianassa Manning, 1987 1993; Felder and Manning, 1995; USNM] Corallianassa articulata (Rathbun, 1906) {'Neocallichirus limosa (Poore, 1975) [Poore, 1975; Corallianassa borradailei (de Man, 1928) Poore and Griffin, 1979; NMV; USNM]-removed to *Corallianassa collaroy (Poore and Griffin, 1979) JPoore Biffarius] and Griffin, 1979; Sakai, 1992] Neocallichirus mamingi Kazmi and Kazmi, 1992 *Coraltianassa longiventris (A. Milne-Edwards, 1870) Neocallichirus motuccensis (de Man, 1905) TYPE SPECIES [de Man, 1928; Biffar, 1971b; USNM] Neocallichirus natalensis (Barnard, 1947) Genus Corallichirus Manning, 1992 'Neocallichirus nickellae Manning, 1993 [Manning, Corallichirus hartmeyeri (Schmitt, 1935) 1993; USNM] *Corallichirus placidus (de Man, 1905) [de Man, 1928; Neocallichirus pachydactylus {A. Milne-Edwards, 1870) Manning, 1988] 'Neocallichirus rathbunae (Schmitt, 1935) [Biffar, 'Corallichirus tridentatus (Von Martens, 1868), new 1971b; Manning and Heard, 1986; USNM] combination 'Neocallichirus sassandrensis (Le Loeuff and Int^s, *Corallichirus xuthus (Manning, 1988) TYPE SPECIES 1974) [Le Loeuff and Intes, 1974] [Manning, 1988, 1992; USNM] Neocallichirus taiaro Ngoc-Ho, 1995 Genus Glypturus Stimpson, 1866 Genus Paraglypturus TUrkay and Sakai, 1995 *Glyplurus acanthochirus Stimpson, 1866 TYPE 'Paraglypturus calderus TUrkay and Sakai, 1995 TYPE SPECIES [Biffar. 1971b; Manning, 1987; USNM] SPECIES [TUrkay and Sakai, 1995] *Glypturus armatus (A. Milne-Edwards, 1870) [Kens- Genus Sergio Manning and Lemaitre, 1993 ley. 1975; Poore and Suchanek, 1988] 'Sergio guaiqueri Blanco Rambla, Linero Arana, and 'Glypturus karumba (Poore and Griffin, 1979) [Poore and Lares M., 1995 [Blanco Rambla et ai. 1995; USNM] Griffin, 1979] 'Sergio guara{KoAtipxti, 1971) [Rodrigues, 1971; Man- Glypturus laurae (de Saint Laurent, 1984) ning and Lemaitre, 1993; USNM] *Glyplurus martensi (Miers, 1884) [Poore and Griffin, 'Sergio guassutinga (Rodrigues, 1971) TYPE SPECIES 1979; Sakai, 1984; Dworschak, 1992] [Rodrigues, 1971; Biffar, 1971b; Manning and *Glypturus motupore Poore and Suchanek, 1988 [Poore Lemaitre, 1993; Manning and Felder, 1995; USNM] and Suchanek. 1988; NMV; USNM] 'Sergio mericeae Manning and Felder, 1995 [Manning *Glyplurus mucroruila (Strahl, 1861) [de Man, 1928; Tir- and Felder, 1995; USNM] mizi, 1977; Poore and Griffin, 1979; Dworschak, 1992] *5ergio mirim (Rodrigues, 1971) [Rodrigues, 1971; Man- Genus Lepidophthalmus Holmes, 1904 ning and Lemaitre, 1993; USNM] TUDGE ETAL.: PHYLOGENY OF THE CALLIANASSIDAE AND CTENIXTHELIDAE 145

*Sergio sulfureus Lemaitre and Feldcr, 1996 [Lemaitre b. Relative length of cephalothorax to total body length andFelder. 1996; USNM| (cl/tl): approximately half (0); a third or less (I). *Sergio trilobatus (Biffar, 1970) [BIffar. 1970, 1971b; c. Cardiac prominence (Fig. 1 A): present (0); absent (I). Manning and Lemaitre, 1993; NMV; USNM] d. Dorsal oval on carapace (Fig. IB, D): ab.<:ent (0); pres- ent (I). Subfamily Cheraminae Manning and Felder. 1991a e. Rostrum: present, strongly spinose (Fig. IA-C, E) (0); Genus Cheramus Bate, 1888 pre'ten;, weakly spinose (I); present, nonspinose and *Cheramus longicaudatus (Sakai. 1%7), new combination broad 'Fig. ID) (2y. absent (3). *Cheramus marginalus (Rathbun, 1901) (Biffar, 1971b; f. Rostrum: present, flatly spinose (Fig. I A) (0); present, Manning and Felder, 1991a: Blanco Ranibia and Linero spinose but downturncd < Fig. IB) (I); present, spinose Arana, 1994; USNM] but upturned (2): present, nonspinose and broad (3); Cheramus oblongus (Le Loeuff and Intds, 1974) absent (4). Cheramus orienialis Bate, 1888 g. Rostrum: dorsally urmed with spines (Fig. lA) (0); *Cheramuspraedatrix (de Man, 1905), new combination dorsally unarmed (1). *Cheramus pmfundus Biffar, 1973 TYPE SPECIES [Bif- h. Anterolaieral projectioni on the frontal margin of the far, 1973) carapace: present, strongly spinose (Fig. IC) (0); pres- *Cheramus propinquus (de Man. I9()S). new combination ent, but poorly developed as blunt lobes (Fig. IB, D) *Cheramus rectangularis (Ngoc-Ho. 19911, new combi- (1): absent (2). nation i. Median mstral carina: pn;sent (Fig. I A) (0); absent (I). *Cheramus sibogae (de Man, 1905), new combination *Cheramus spinophthalmus (Sakai, 1970), new combi- 2. Eyestalks and corneas nation a. Eyestalks: cylindrical ((i); flattened (I); flattened but Genus Poti Rodrigues and Manning. 1992 concave on dorsal surface (2). *Poti gaucho Rodrigues and Manning, 1992 b. Eyestalks: terminally rounded (Fig. IC) (0); termi- Genus Scallasis Bate, 1888 nally tapered to a blunt point (Fig. IE) (1); termi- Scallasis amhoinae Bate, 1888 TYPK SPECIES nally elongate and pointed (Fig. ID) (2). c. Corneas: terminal on eyestalk (Fig. IC) (0); subter- Subfamily Eucalliacinae Manning and Felder, 1991a (as minal on eyestalk (dorsally situated) (Fig. ID, E) (I); Eucalliinae) absent (2). Genus Calliax de Saint Laurent. 1973 d. Corneas: equal to eyestalk width (Fig. IC) (0); le.ss *Calliax aequimana (Baker, 1907) |Poore and Griffin, than eyestalk width (Fig. ID, E) (1); absent (2). 1979) e. Corneas: subglobular (Fig. IC) (0); disc-shaped (Fig. *CalUax bulimba (Pooie and Griffin, 1979) IPoore and IB, E) (1); indistinct or absent (2). Griffin, 1979) *CalUax lobata (de Gaillande and Lagarddre. 1966) 3. Abdominal somites TYPE SPECIES (de Gaillande and Lagardere. 1966; a. Abdominal somite I. anterolateral lobes (Fig. 21): de Saint Laurent and Bozic, 1972| present or indicated (0): absent (I). *Calliax novaebriianniae (Borradaile. 1899) (de Saint b. Abdominal somite I. pleuron (Fig. 2J): acute and pro- Laurent and Manning, 1982) jecting (0); blunt and obsolete (I). *Calliax punica de Saint Laurent and Manning, 1982 (de c. Anterolateral margm of abdominal somite 2: does not Man, 1928; de Saint Laurent and Manning, 1982; overlap abdominal somite I (0); overlaps abdominal USNM) somite 1(1). Calliax sakai de Saint Laurent, 1979 [dc Saint Laurent d. Abdominal somite 2: equal to length of somite 6 (0); and Manning. 1982) greater than length of somite 6 (Fig. 2K) (I); less than 'Calliax tooradin (Poore and Griffin. 1979) (Poore and length of somite 6 (2). Griffin, 1979; de Saint Laurent and Manning, 1982) e. Abdominal somites 3-.'': without dense tufts of lat- Genus Eucalliax Manning and Felder, 1991a eral setae or at most sparse vertical rows (0); with *Eucaltiax cearaensis Rodrigues and Manning, 1992 (Ro- dense tufts of lateral setae (Fig. 2K) (I). drigues and Manning, 1992a; USNM) f. Abdominal somites .V.S: not dorsally ornamented with *Eucalliaxjonesi(\icwd. 1989) (Heard. 1989; Felder and distinct patterns of grooves and integumental glands Manning, 1994; USNM] (0): dorsally ornamented with strong symmetrical pat- *Eucalliax mcilhennyi Felder and Manning, 1994 (Felder terning of grooves and integumental glands (1). and Manning, 1994; USNM] g. Abdominal somite 6: without lateral projections (Fig. *Eucalliax quadracula (Biffar, 1970) TYPE SPECIES 2B -F, K) (0); with lateral projections (Fig. 2A) (I). (Biffar, 1970, 1971b] h. Coxa of perciopod 4: rectangular, without anterome- sial lolx- (0): flattened, with anteromesial lobe (I).

4. Gills Appendix 2. List of characters and their states used in a. Epipotis: 1-7 (rarely 2-7) present (0); 4-7 vestigial analysis of genera of Ctenochelidae and Callianassidae. or absent (I): 3-7 absent (2); 2-7 absent (3). Alphanumeric character labels correspond with those in b. Podobranch 2: present (0); vestigial or absent (I). Appendix 3. c. Podobranchs 3-7: present (0); 3-7 absent or some rudimentary (I). I. Carapace and rostrum d. Arthrobranchs on somites 1-7: 022 2222 (0); 012 a. Linea thalassinica: present and typically extending 2222 (I); 002 2222 (2); 001 2222 (3). back to posterior edge of carapace (Fig. I A) (0); pres- e. Pleurobranchs: 5-7 absent, 8 rudimenury (0); 5-8 ab- ent but incomplete (I); absent (2). sent (I). 146 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2.2000

5. Epistome and antennae g. Pereiopod 1, major cheliped, propodus: tapering dis- a. Epistome: without setae (0): with long setae (I). tally (0); not tapering distally (I), b. Antenna I (antennule), segment 3: longer than seg- h. Pereiopod I, major cheliped; angle of fixed finger to ment 2 (0): shorter than or about as long as segment palm: 90° or less (0); greater than 90° (I), 2(1). i. Pereiopod I, major cheliped, chelae fingers: shorter c. Antenna 1 peduncle: longer than antenna 2 (antenna) than, or approximately equal to palm (0): longer than peduncle (()); shorter than antenna 2 peduncle (I): as palm (I), long as antenna 2 peduncle (2). j. Pereiopod I, minor cheliped, chelae fingers: shorter d. Antenna I, dense brush of ventrally directed long se- than or approximately equal to palm (0); longer than tae: absent (0); present (I). palm (I), e. Antenna 2 scaphocerite: prominent, much longer than k. Pereiopod I, major cheliped, chelae fingers: without wide (0); reduced but articulating, about as long as comb of fine teeth (0); with comb of fine teeth (I). wide (I); absent (2). I. Pereiopod I, major cheliped, length-to-width ratio of propodus: 1.0-2.0 (0); 2.1-3.0 (I); 3.1-4.0 (2); 6. Mouthparts 4.1-5.0(3). a. Maxilla 2 scaphognathitc: without long setae (0); with m. Pereiopod 1, major cheliped denticulation: even (0); one or more long seta(e) (1). uneven (with occasional larger teeth) (1). b. Maxilliped I endopod: 2- or 3-segmented (or elongate n. Pereiopod 1, major cheliped, ventral teeth on ischium: and tapering) (Fig. IF) (0); minute (Fig. IG) (I); ab- present (0); absent (I), sent (Fig. IH) (2). o. Pereiopod I, minor cheliped, ventral teeth on ischium: c. Maxilliped 1 exopod: 2- or 3-segmented (sometimes present (0); absent (I), with flagellum) (Fig. IG) (0): I-segmented (Fig. IF, p. Pereiopod 2, row of setae on lower margin of ischium- H)(l). propodus: absent (0); present (I). d. Maxilliped 2 exopod: equal to or longer than merus q. Pereiopod 2: simple (0); chelate, with dactylus longer of endopod (0); shorter than merus of endopod (1); than fixed finger (1); chelate, with dactylus as long absent (2). as fixed finger (2). e. Maxilliped 3: pediform, ischium-merus length more r. Pereiopod 3 propodus: linear (more than 3 times as than 3 times merus width (Fig. IK) (0); subpediform, long as wide) (0); oval (approximately twice as long ischium-merus length about twice merus width (Fig. as wide) (I); oval, but with prominent heel on prox- IJ) (I); operculiform, ischium-merus length less than imal corner of lower margin (2). 2 times merus width (Fig. II) (2). s. Pereiopod 4 propodus: linear (more than 3 times as f. Maxilliped 3, exopod: reaching almost to end of merus long as wide) (0); oval and flattened (twice as long of endopod (0); reduced or vestigial (1); absent (Fig. as wide or less) (I), II-K) (2). t. Pereiopod S: simple (0); chelate or subchelate (I). g. Maxilliped 3, crista dentata: as prominent toothed ridge (Fig. IJ, K) (0); obsolete or absent (Fig. II) (I). h. Maxilliped 3. meral spine (Fig. IJ, K): present (0); ab- 8. Pleopods sent (1). a. Pleopods: I absent or reduced, 2-5 similar and lamel- t. Maxilliped 3, merus: with a denticulate distal border lar (0); 1 and 2 variously absent, reduced or sexually (Fig. IJ) (0); without a denticulate distal border (Fig. modified, 3-5 similar and lamellar (1). II) (I). b. Male pleopod I: absent (0); present, I-segmented (I); j. Maxilliped 3, merus: not projecting beyond articula- present, 2-segmented, with the 2nd segment more or tion with carpus (Fig. IK) (0); barely projecting be- less triangular (2); present, 2-segmented, with the 2nd yond articulation with carpus (Fig. IJ) (I); strongly segment ovate (3); present, 3- or more segmented (4). projecting beyond articulation with carpus (Fig. II) (2). c. Male pleopod I: absent (0); pre.sent, with obvious ap- k. Maxilliped 3, propodus: slender, longer than broad, pendix intema (I); present, appendix intema as minute at most slightly wider than dactylus (Fig. II, K) (0): hooks on ramus (2); present, appendix intema absent oval, as broad as long, or at least twice dactylus width (3). (Fig. IJ)(I). d. Male pleopod I: present, uniramus (0); present, but I. Maxilliped 3, dactylus: digitiform and slender, longer vestigial (I); ab.sent (2). than broad (Fig. II-K) (0); oval, as broad as long (t). e. Male pleopod 2: present, biramous(O); present, uni- ramous (1); present, but vestigial (2); absent (3). 7. Pereiopods f. Male pleopod 2: present, appendix masculina present a. Pereiopod I (chela) in male: equal or subequal (0); (0); present, appendix masculina pre.sent and fused unequal (I). to appendix intema (I); present, appendix masculina b. Pereiopod I in female: equal or subequal (0); unequal absent (2); absent (3). (I). g. Female pleopod I: present, biramous (0); present, uni- c. Pereiopod I, merus of major cheliped: with straight ramus (I). lower margin (0); with convex lower margin (I); with h. Female pleopod I: present, I -segmented (0); present, toothed lower margin (2): with distinct meral hook (3). 2-segmented, with the 2nd segment more or less tri- d. Pereiopod I. merus of minor cheliped: with straight angular (I): present, 2-scgmcntcd, with the 2nd seg- lower margin (0); with convex lower margin (I); with ment ovate (2); present, 3- or more segmented (3). toothed lower margin (2); with distinct meral hook (3). i. Female pleopod 2: present, biramous(O): present, uni- e. Pereiopod I: simple or subchelate (0); chelate (I). ramous (I). f. Pereiopod I, major cheliped, carpus and propodus: j. Pleopods 3-5: rami lanceolate (0); rami broad (I). without three spines on upper margin (0); with three k. Pleopods 3-5, endopod and exopod: not a .semisphere spines on upper margin (I). when combined (0); semispherical when combined (I). TUDGE ETAL: PHYLOGKNY OF THE CALLIANASSIDAE AND CTENtXTHELIDAE 147

1. Pleopods 3-5: with appendix Intema (Fig. 2G-I) (0); h. Telson: longer than uropods (Fig. 2A, D) (0); shorter without appendix intema (I). than uropods (Fig 2B. F, K) (I); equal or subequal m. Pleopods 3-5, appendix inicmu: present and digiti- to uropods (Fig 2E, Ci (2). form (Fig. 2G) (0); present but reduced and stubby i. Telson: without one or two lateral spines (Fig. 2A-C, (Fig. 2H) (I); present but embedded (Fig. 21) (2); ab- F. K) (0); with one or two lateral spines (Fig. 2D, E) sent (3). (I), j. Posterior margin of telson: rounded (Fig. 2A, E) (0); 9. Uropods and lelson flattened (Fig. 2D. F) (1), slightly indented (Fig. 2C) a. Uropodal exopod: simply ovate (Fig. 2A) (0); with an- (2); strongly excavate iFig. 2B) (3). terodorsal setose thickening (= dorsal plate) (Fig. 2C-F, k. Posterior margin of telson: unarmed (Fig. 2A-C, F) K) (I); bilobed. or markedly bipartite (Fig. 2B) (2). (0); w iih one spine (1); with more than one spine (Fig. b. Uropodal exopod: longitudinally carinate dorsally 2D. E) (2) (Fig. 2A, B) (0): acarinate dorsally (Fig. 2C. E, F) (I). c. Uropodal exopod, lateral notch or incision: pre.sent lO.Setal rows (Fig. 2B)(0);ab.sent(l). a. Abdominal somite 2, lateral setal row of plumose se- d. Uropodal endopod: oval, distally rounded, with sim- tae (Fig. 2K): absent (»); present (I). ilar length to width (Fig. 2A, D) (0); distally trun- b. Abdominal somites 3-.i, lateral .setal row of plumose cate, distolateral margin subacutu, wider distally than setae (Fig. 2K): absent (0); present (1). proximally (Fig. 2C) (I); longer than broad, tapering c. Abdominal somite 6, lateral .setal row of plumose se- distally (Fig. 2B, K) (2); much longer than broad, not tae (Fig 2K): absent (0); present (I). tapering distally, strap-like (Fig. 2F) (3). d. Abdominal somite 6. tiansverse setal row of plumose e. Uropodal endopod: longer than broad, tapering, or not setae on posterior dorsolateral margin (Fig. 2K): ab- tapering, distally (Fig. 2B, E. F) (()); broader than sent (0); present (I). long, flattened distally (Fig. 2C) (1). e. Abdominal somite 6, oblique setal row of plumose se- f. Uropodal endopod: without distinct distolateral spine tae bi;tween lateral and transverse setal rows (above) (Fig. 2A-D, F, K) (0); with distinct distolateral spine (Fig. 2K): absent (0); present (I). (Fig. 2E)(I). g. Telson: longer than broad (Fig. 2A. D) (0): as long as broad (Fig. 2B, E) (I); broader than long (Fig. 2C, F) (2). 148 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2.2000

Appendix 3. Data matrix. ( j = 0/1, an equivocal character state assignment)

Cb*ract«r nuatoer lllllllII22222313}3)3344444555SS«««««e<«77777117177777717777a<8S8aa8SB88899999999»«00000 T«x« ABCDBroHIABCDSABCOBFGHABCOSABCDEUCDEFCKlJKLABCDEFGHIJIUJINOPORSTABCDBPGHIJKLMAfiCDErGHlJKABCDe

Anacalliax agassizi 01710011011112777700I7211110010101110201100011221011010000072271117001110770120010021000????? Anacalliax argentinensis 01010111011222010200112111110101II111021110001122100101001001221112200112I>11012000002102CI10000 Blffariua arenosa 01112312111111110010013112110ai>2011122011100113310100001101122011]103311011001111101100010100 1 2 Biffariua bifomis 011123121111111100000121121101020111221111t010331010000110112211122l>331101ia01111101100010100 BiCfarius ceramica 011111121111111100100131121101020111220111001132101000001011220113202211011011111100100010100 3 Biffariua delicatulus 011123121111111101000121121101010111221111101033101100000111220113201211011011110101100010100 1 2 Biffariua fragilis 0111231111111011010001311211000201112201111101030101000011011221112203311011011112002101010100 1 Ca21ianaaaa acutiroatalla O71101121111117?77?7017777770001777722011200?7??77????????777?7?17?777130?7771112002102177??7 Callianaaaa aoiboinenala 0?112112111111777177077777770001011122011200112ll011000000012271122ai211077771112002102177777 34 12 Callianaaa audax 01112312111111777?7707777?7?0101777712?111101120101100001117?27717?77711011??1112002102077?7? Callianaaaa auatralla 0111231211111211O010O1211111O1O1O11112011OOO11311O1OO00O10112201132O1211O11O1111110110221O11O 2 Callianaaaa caledonica 01110011111111?77177077?77??0000?117a201100011331011000110012277l7?77?110??7711120021021?7777 1 Callianaaaa diaphora 0111231111111177707707211H?0l0l010102O11000113HO0O0OO010112201122fl771101101iai07021001?7777 2 3 11 Callianaaaa filhali 011111111111111001100121121100010111220111001132100000001011220113203313011011110000001010100 Callianaaaa joculacrix 0111001111122211011001211211020101100201100011311000000011101220111213312011011111002101110100 2 1 Callianaaaa jouaaeaujaai 071101111121117777770777777700027??7120112101120101100001?7?7277777777?7777??7?111021000777?7 Callianaaaa krauaai 011123111101111101100121121100020110221110101U1101000001011221113203312011021112002102010100 2 2 Callianaaaa lawtonae 011110121101117772?70737??770002?117220112007011101100010007227?7??7?777?110017120011010777?7 Callianaaaa longicauda 01110210111110777777??3112170101777712100200?1731????1?»??077177122011130?77701120001011777?7 callianaaaa marchali 011100111211117771100721111701010101020110001131100100000011220110023313011011S10700100177777 1 Callianaaaa aiauritiana 01112312llllll7?77777777?77?7??l???7120110001131100?O0O107177277142O0113O7?7?11110021010777?? Callianaaaa pixii 0111101101222277??777?31171?0110702I10211100000iai00100011001220112300?1001113111200210307??77 2 1 Callianaaaa prajdatrix 011100111111107??0?701777?7?0101???722000200772710110?0010?7727?123002130?77?1112000112177?7? Callianaaaa propinqua 01110011111111777777O7211717O1O27111120O0200112210110000000121771230O211O110O1112O0011O2777?? Callianaaaa ranongenaia 011100111111111100000121111101010110121101111131101101001001221112300113011021111002101010010 1 Callianaaaa rectangularia 0111001111111177?07?0721??7?0001711122011200172210100000100122?77?7?77777777711120001122777?7 Callianaaaa aibogaa 011102110111117770770777777?0101777?020110007122101100010007?17?17?0071701100111100010217777? 2 Callianaaaa apinophthalma 011102111111101100000T7?777?0201777?12011100712110100000100771?7?7?777777777?11120021021777?? 3 2 Callianaaaa subcerranea 011123111111101101100121121101010101120110001131100100000001220112301210011011111001101210110 2 23 2 1 Callianaaaa cridencata 011102101111111101100131121100120100121111101131101000001001221112300211011001112002100010110 Callianaaaa tyrrhena 011123111111111111100131121100010101220112001131100100001111220110023313011011112001100010110 2 Callianidfta typa 211023121101121101000001101001011011000110001111101100011111110103200211000010112000100000010 Callianopais goniophthalma 011100110211120111001121131111000010020010111101101100001000211112300177011000012001101000011 1 Calliax aaquimana 0711231111111177777007777777110101002111111100111001000010072J11123001110117711120021020777?? 2 Calliax bulimba O711231111111177777707777777110277772211111107221O11O0OO1OO7777712300177777771112002101077777 Calliax lobata O7172311111112777777077?????llO1777?1201101111211011O00OlOO12i?71230O113OllO11102O02100177777 2 2 Calliax navaabritanniaa O111OO1O1O11117771770727777711O1O110211112110O221010O0OO1O0122011410O113011OO11120021O2077777 Calliax punica 0111231111111111010O012O12111101O11010O11O1100111811O0OO1OO122111110O211011O21112O021O101011O 1 Calliax cooradin 0?112311111111?77777072?????01000110101110111111100100000117?77?100202110777711120021000????? 2 Callichirua 011129111121117771110721111700110010211111101021101001011011221113300213011021113002102077777 222 12 2 1 1 calllchirua balaal 01111111111111777777072111170001010002111110113110010000101772771330221107777111200210207777? Callichirua foraaci 011123121111117771100721111700110110021110001121101000001011221111301211111021112002102077777 2 Callichirua guineenaia 0711231111111177777707211117OOO101100211100011211001OO0O1O11221711312213O11O21112002102077777 Callichirua ialagranda 0111231111211111O111O1211111OOO1010O221111101021101O01O210112211123O0211O11O211130O2102O10111 1 Callichirua major 0111231111111111S1110121111100U.0100221111101I3110«000001011231112>00211011021113002102010111 2 Callichirua aailachari 011123111121111100110121111100110100221111101021100100010011221112300211011021113002102010111 3 Callichirua turneranua 01H00111111111100100?21111?00010110221110101133101000001011721712300211017??1112002103077777 Charamua marginatus 01110111111111010000013113100101011112011000111110O0O10110011101113O33110111O1112O01102110000 1 2 1 CharaoUia profundua 011100111111117777000131121O01O2O11112O110000722101100O0100711711230O277711O01112O00211277777 2 Corallianaaaa collaroy O7110O10111ia077777O0?311717OlO0OlOO2200011O1122101100000001221113100213O77??1112O021O20?7?7? 1 1 Corallianaaaa longivancria 011102101111101101100131111111020110220110101122101100001001221111300212011021112002102010100 1 Corallichirua placidua 071100101000107777700777777701127777127110107722101100007007727777777777777771112002101077777 CorallichiruB xuchua 011100101000101100100131121100120200120110107122101100001001221117777711011011112002100000000 Ctanochalaa balaai O10002020O11121010001130121O110O0O1102O0100O110O1O0010131111210114300113011O020O100100210O00O 2 Ctanochalea holthuiai 010002100111120770001731121010000102020010001132100110131001211102200177000000000002000000000 Ctanochalaa maorianua 01000011011222110010013012101100000002011000110010011012111121011220017771100200200210200077? Dawaonlua laciaplna 011001110211120000001131111001010010020010001133100100011001211113200111011000010001000010011 2 Buealliax eaaraanaia OllO231111111111010OO121111112O201101201101100111O110OOaioai2211133O01110110«1112O021O101011O 1 2 TUDOE ETAL: PHYLOGENY OF THE CALLIANASSIDAE AND CTENOCHEUDAE 149

Appendix. 3. Continued character number llKUll:i222;'33)333)]44444SSSS5«(e(«et

Neocalliehlrua caechabitator 01112311U111277?77001????770101??7?122110107121100100001fl))??2??l??7??l]0111311111022020?77?7 22 1 11 Kaocallichirua darwinanaia 01112311111111777777012177770101011112011010•77ai?7'717?771?72771777?7130110711111021010??777 2 1 Haocallichirua yrandimana 011111111111111100100121111101010110120110101121101100001011221113300211011031111102102010101 2 3 Heocallichirus horneri 01112311111111777777017777770101777712011010>121100100000117727713300213077771111102202077777 Neocallichirua indica 01112311111 111?7?'770777?77701117?7712011010?1211C010C001C??72771330777?7777711111011020?7?7? 2 Heocallichirus lemaitrei 011129111111111101100121111111010110120110101121100100001011221113300111011021111102202010100 2 2 Neocallichirua linosa 0111231111111211011001311111010201111201101011I11011000010012311121022110110111U102200010100 2 2 1 1 Heocallichirus niekellae 01112311111 111 110010012111110101011012011010.1221011010010L1221113300277711011111102102010100 2 2 Neocallichirua rathbunae 0111021011111111020001311111011201101201101011211101100001011220113300211011021111102101010100 1 Neocalliehirus sassandrenais 01111312111lll?'>01000121111?0111011112ail01O>7?3177?70????11221117?777100110311111O110217???? 1 2 Neotrypaea californiensia 01112)111121111001100121121100120111122011200.112100010001011220112303311011021111001102110100 1 1 Neotrypaea givas 01112)11112111100000012112110202011022111200.l]21011000]100121011002331301101111Ut03102110100 1 Hihonotrypaea harmandi 0111341211111111011001311211030101103301130O1I331O1001000O11221113303311011011110001103110100 3 Nihonotrypaea japonica 0111)412111 111 110110013112110201011022011200.113101001001011220112303313011011111001102010100 1 I Hihonotrypaea petalura 0111331211111111001001)112110301011022011200-1)1101101010011221112303313011011111001101010100 2 1 Notiax brachyophthalma 011100121111101101100121121101010110221110001111101100001001220113301311011011111000101110100 1 2 Paracalliax bollorei 010023120;:12220010001721111001000010000010001122101100C01007200107770012011000010001000177777 1 Paraglyptuxua calderus 01102312111111100000072017771111001000011011111110110000111122111330011101101111200210007777? 1 Poti gaucho 1111021111111311030001211110010101111301100a??711777717'>77012107133a0211011001112000t00010100 3 2 Sergio guaiqueri 0111231111 nil 11011001)111110102021102011010112110110OCaiO112201123O027?7110211120O21O201010O Sergio guara 011123121111111101100121111101010111120110101121101100001011221113300277711021112002102010110 3 2 Sergio guassutinga 011100101111111101100121111111010110121110101121100100001111221113300211011011112002102010100 1 3 Sergio mericeae 011100101111111101100121111101010211221110101121100I00001C11221113300011011021112002102010100 1 3 Sergio siiria 011123111111111I021001311111O101O11O321110101122101100011C11320113300311011021113003103110110 1 1 Sergio aulfureus 011100101 Ul 11 UOl 10012111111111010112011010) IMIOOI00001011220113300011011021112003102010100 1 Sergio trilobatus 01112311111111010110012111111101011022111010I121100100011C11221113300111011021112002102010100 Trypaea australienaia 011111111111111101100121111100120111221112001132100000001011220113303311011011112001102010100 1