474 J. Paleont., 74(3), 2000, pp. 474–486 Copyright q 2000, The Paleontological Society 0022-3360/00/0074-0474$03.00 PECTINATE CLAWS IN DECAPOD CRUSTACEANS: CONVERGENCE IN FOUR LINEAGES DALE TSHUDY AND ULF SORHANNUS Department of Geosciences, Department of Biology, Edinboro University of Pennsylvania, Edinboro, 16444, ,[email protected]., ,[email protected]. ABSTRACT—Decapod crustaceans bearing major claws with long, slender fingers armed with pectinate (comblike) denticles have been described in six genera arrayed within three families (Polychelidae, Nephropidae, and Ctenochelidae) in three infraorders (Palinura, Astacidea, and Anomura, respectively). Only one or a few genera in each infraorder exhibit this claw form. The pectinate claw form is confidently interpreted as having evolved independently in four lineages: once in the Polychelidae, once in the Ctenochelidae, and twice in the Nephropidae. Three of the lineages are known from both the fossil record and modern seas; the polychelid form is known only from Jurassic rocks. Convergence in this claw form developed to the extent that isolated fossil claws (i.e., claws without associated bodies) have commonly been misidentified at high taxonomic levels. The fossil record confirms what seems intuitively reasonable: that claw morphology is prone to convergence and should not, by itself, be given a high degree of taxonomic importance. INTRODUCTION C ONVERGENT EVOLUTION of morphologic features, including some basic features, is well documented in the Arthropoda. Several of these convergences appear both in the Crustacea and in other arthropod groups. The very existence of claws in dis- tantly related arthropod groups (i.e., the Chelicerata and Crus- tacea) has been interpreted as the result of convergent evolution (Robison and Kaesler, 1987, p. 216). This paper documents that, within the Crustacea alone, a specific and distinctive ‘‘pectinate’’ (comblike) claw form has independently evolved four times. Pectinate claws have been described in six genera arrayed within three families (Polychelidae Wood-Mason, 1874; Ne- phropidae Dana, 1852, sensu Tshudy and Babcock, 1997; and Ctenochelidae Manning and Felder, 1991) belonging to three infraorders (Palinura Latreille, 1803; Astacidea Latreille, 1803; and Anomura H. Milne-Edwards, 1832, respectively) (note that Scholtz and Richter (1995), in their cladistic analysis of the Rep- tantian decapods, argue that the infraorders Anomura and Pal- inura are paraphyletic groups). Only one or a few genera in each infraorder exhibit this claw form. This claw form is confidently interpreted herein as having evolved independently in four lin- eages, once in the Polychelidae, once in the Ctenochelidae, and twice in the Nephropidae (Fig. 1). Three of the lineages are known from both the fossil record and modern seas; the poly- chelid form is known only from Jurassic rocks. The Polychelidae are represented here by Paleopentacheles von Knebel, 1907 (Fig. 2.4), the Ctenochelidae by Ctenocheles Kishinouye, 1926 (Fig. 2.3), the Nephropidae by Acanthacaris Bate, 1888 (Fig. 2.2) in one lineage, and Oncopareia Bosquet, 1854, sensu Tshu- dy (Figs. 3–5), Thaumastocheles Wood-Mason, 1874 (Fig. 2.1), and Thaumastochelopsis Bruce, 1988 in a second lineage. Convergence in this claw form developed to the extent that isolated fossil claws (i.e., claws without associated bodies) com- monly have been misidentified at high taxonomic levels (Tshu- dy, 1993, p. x). Isolated claw pairs or, more problematically, single claws are common in the fossil record because claws are generally more preserveable than the remainder of the body. The ctenochelid shrimp genus Ctenocheles is the best example of this preservational bias for claws. Ctenocheles has densely cal- cified claws but the remainder of the body is so poorly calcified that the fossil record of this genus consists only of claws. Among the four lineages, convergence in major claw form is more impressive in finger morphology than in palm morphology. The exception is in the genera Ctenocheles, Thaumastocheles, and Thaumastochelopsis which, as noted by many workers, have major claws of very similar finger and palm form. The similarity of fingers from fossils now mostly identified as Ctenocheles and Oncopareia has resulted in considerable tax- onomic confusion. Tshudy (1993), using the morphologic cri- teria given herein (Table 1), addressed the taxonomic confusion surrounding isolated fossilized claws. A list of recognized taxa with pectinate claws and their occurrences is included herein (Appendix 1). For a combination of reasons, it seems intuitively reasonable that, within the crustacean order Decapoda, certain specific claw forms might have evolved independently two or more times. Claw function is strongly and directly linked to environmental pressures (e.g., substratum, food sources, etc.) which, of course, are variable. There are only a limited number of claw designs that will fulfill these functions. Similar selection pressures un- doubtedly arose multiple times. It then follows that claw mor- phology should be prone to convergent evolution. The decapod fossil record herein documents this tendency for convergence. The purpose of this paper is to document the convergence of claw, and especially finger, morphology in decapod crustaceans. The paper includes an annotated list (Appendix 1) of fossil and recent decapod taxa bearing pectinate claws, an interpretation of the phylogenetic relationships of these taxa, a discussion of the evolutionary and bathymetric history of the four convergent lin- eages, and discussion of the adaptive significance of the pecti- nate claw form. MATERIALS AND METHOD The senior author examined specimens of as many of the fos- sil and recent taxa with pectinate claws as were available. This included all species of the Recent lobster genera Acanthacaris and Thaumastocheles, all known material of the fossil lobster Oncopareia, and several fossil and Recent species of Ctenoche- les. The Recent lobster Thaumastochelopsis and the fossil deca- pod Palaeopentacheles, both monospecific genera, were not ex- amined firsthand. Phylogenetic relationships between nephropid lobster genera bearing pectinate major claws were evaluated by the cladistic method. We wanted to determine if the pectinate claw form was also convergent within the lobster family Nephropidae. We an- alyzed a good representation of the overall diversity of the group (13 of 17 nephropid genera). Recent studies (e.g., Bremer et al., 1999) indicate that the percentage of supported nodes within a
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474
J. Paleont., 74(3), 2000, pp. 474–486Copyright q 2000, The Paleontological Society0022-3360/00/0074-0474$03.00
PECTINATE CLAWS IN DECAPOD CRUSTACEANS: CONVERGENCE INFOUR LINEAGES
DALE TSHUDY AND ULF SORHANNUSDepartment of Geosciences, Department of Biology, Edinboro University of Pennsylvania,
ABSTRACT—Decapod crustaceans bearing major claws with long, slender fingers armed with pectinate (comblike) denticles have beendescribed in six genera arrayed within three families (Polychelidae, Nephropidae, and Ctenochelidae) in three infraorders (Palinura,Astacidea, and Anomura, respectively). Only one or a few genera in each infraorder exhibit this claw form. The pectinate claw formis confidently interpreted as having evolved independently in four lineages: once in the Polychelidae, once in the Ctenochelidae, andtwice in the Nephropidae. Three of the lineages are known from both the fossil record and modern seas; the polychelid form is knownonly from Jurassic rocks. Convergence in this claw form developed to the extent that isolated fossil claws (i.e., claws without associatedbodies) have commonly been misidentified at high taxonomic levels. The fossil record confirms what seems intuitively reasonable: thatclaw morphology is prone to convergence and should not, by itself, be given a high degree of taxonomic importance.
INTRODUCTION
CONVERGENT EVOLUTION of morphologic features, includingsome basic features, is well documented in the Arthropoda.
Several of these convergences appear both in the Crustacea andin other arthropod groups. The very existence of claws in dis-tantly related arthropod groups (i.e., the Chelicerata and Crus-tacea) has been interpreted as the result of convergent evolution(Robison and Kaesler, 1987, p. 216). This paper documents that,within the Crustacea alone, a specific and distinctive ‘‘pectinate’’(comblike) claw form has independently evolved four times.
Pectinate claws have been described in six genera arrayedwithin three families (Polychelidae Wood-Mason, 1874; Ne-phropidae Dana, 1852, sensu Tshudy and Babcock, 1997; andCtenochelidae Manning and Felder, 1991) belonging to threeinfraorders (Palinura Latreille, 1803; Astacidea Latreille, 1803;and Anomura H. Milne-Edwards, 1832, respectively) (note thatScholtz and Richter (1995), in their cladistic analysis of the Rep-tantian decapods, argue that the infraorders Anomura and Pal-inura are paraphyletic groups). Only one or a few genera in eachinfraorder exhibit this claw form. This claw form is confidentlyinterpreted herein as having evolved independently in four lin-eages, once in the Polychelidae, once in the Ctenochelidae, andtwice in the Nephropidae (Fig. 1). Three of the lineages areknown from both the fossil record and modern seas; the poly-chelid form is known only from Jurassic rocks. The Polychelidaeare represented here by Paleopentacheles von Knebel, 1907(Fig. 2.4), the Ctenochelidae by Ctenocheles Kishinouye, 1926(Fig. 2.3), the Nephropidae by Acanthacaris Bate, 1888 (Fig.2.2) in one lineage, and Oncopareia Bosquet, 1854, sensu Tshu-dy (Figs. 3–5), Thaumastocheles Wood-Mason, 1874 (Fig. 2.1),and Thaumastochelopsis Bruce, 1988 in a second lineage.
Convergence in this claw form developed to the extent thatisolated fossil claws (i.e., claws without associated bodies) com-monly have been misidentified at high taxonomic levels (Tshu-dy, 1993, p. x). Isolated claw pairs or, more problematically,single claws are common in the fossil record because claws aregenerally more preserveable than the remainder of the body. Thectenochelid shrimp genus Ctenocheles is the best example ofthis preservational bias for claws. Ctenocheles has densely cal-cified claws but the remainder of the body is so poorly calcifiedthat the fossil record of this genus consists only of claws.
Among the four lineages, convergence in major claw form ismore impressive in finger morphology than in palm morphology.
The exception is in the genera Ctenocheles, Thaumastocheles,and Thaumastochelopsis which, as noted by many workers, havemajor claws of very similar finger and palm form.
The similarity of fingers from fossils now mostly identifiedas Ctenocheles and Oncopareia has resulted in considerable tax-onomic confusion. Tshudy (1993), using the morphologic cri-teria given herein (Table 1), addressed the taxonomic confusionsurrounding isolated fossilized claws. A list of recognized taxawith pectinate claws and their occurrences is included herein(Appendix 1).
For a combination of reasons, it seems intuitively reasonablethat, within the crustacean order Decapoda, certain specific clawforms might have evolved independently two or more times.Claw function is strongly and directly linked to environmentalpressures (e.g., substratum, food sources, etc.) which, of course,are variable. There are only a limited number of claw designsthat will fulfill these functions. Similar selection pressures un-doubtedly arose multiple times. It then follows that claw mor-phology should be prone to convergent evolution. The decapodfossil record herein documents this tendency for convergence.
The purpose of this paper is to document the convergence ofclaw, and especially finger, morphology in decapod crustaceans.The paper includes an annotated list (Appendix 1) of fossil andrecent decapod taxa bearing pectinate claws, an interpretation ofthe phylogenetic relationships of these taxa, a discussion of theevolutionary and bathymetric history of the four convergent lin-eages, and discussion of the adaptive significance of the pecti-nate claw form.
MATERIALS AND METHOD
The senior author examined specimens of as many of the fos-sil and recent taxa with pectinate claws as were available. Thisincluded all species of the Recent lobster genera Acanthacarisand Thaumastocheles, all known material of the fossil lobsterOncopareia, and several fossil and Recent species of Ctenoche-les. The Recent lobster Thaumastochelopsis and the fossil deca-pod Palaeopentacheles, both monospecific genera, were not ex-amined firsthand.
Phylogenetic relationships between nephropid lobster generabearing pectinate major claws were evaluated by the cladisticmethod. We wanted to determine if the pectinate claw form wasalso convergent within the lobster family Nephropidae. We an-alyzed a good representation of the overall diversity of the group(13 of 17 nephropid genera). Recent studies (e.g., Bremer et al.,1999) indicate that the percentage of supported nodes within a
475TSHUDY AND SORHANNUS—CONVERGENCE IN DECAPOD CLAWS
FIGURE 1—Evolutionary relationships of taxa bearing pectinate claws. Relationships of higher taxa modified from Glaessner (1969). Relationshipsof nephropid lobster taxa from Tshudy and Babcock (1997) and herein. The claw form has independently evolved four times: twice in the nephropidlobsters and once each in the Anomura and Palinura.
cladogram is positively correlated with the number of characters,and negatively correlated with the number of taxa, and that thisjustifies removing closely related taxa. In this study, we omittedfour genera. Palaeonephrops and Paraclytia were omitted be-cause they consistently plot out with a clade including Nephropsand Metanephrops but are anatomically less well known thanthe latter two. Homarinus was omitted because, in external mor-phology, it is very similar to Homarus, and the two consistently
plot out together in our (external morphology based) cladisticanalyses. Additionally, Hoploparia was omitted because, in thesenior author’s opinion, some species presently referred to thegenus are of uncertain taxonomic affinity. Taxonomic assign-ments of species to the genera considered in the cladistic anal-ysis are largely summarized in Tshudy (1993). The 41-characterdata matrix includes exclusively morphologic characters; mostare external hardparts (Appendix 2).
477TSHUDY AND SORHANNUS—CONVERGENCE IN DECAPOD CLAWS
FIGURE 3—Oncopareia bredai Bosquet, 1854, sensu Tshudy, 1993; 1, left view of cephalothorax [Institut Royal des Sciences Naturelles de Belgique(IRScNB)] 90-32c; 2, 3, dorsal and right views of abdomen (IRScNB 90-35c); 4, right view of cephalothorax (IRScNB 90-15b); 5, left view ofcephalothorax (IRScNB 90-17); 6, right view of second abdominal somite (IRScNB 90-23b). Scale bars equal 1.0 cm.
The data matrix was subjected to the exact search algorithm(ie*) in the computer program Hennig86 (Farris, 1988). All char-acters were weighted equally (weight 5 1) and treated as un-ordered (cc-.). Polymorphic character states were coded asunique states. The non-pectinate lobster genus Eryma Von Mey-er, 1840, representing the family Erymidae, was selected as theoutgroup and used to root the trees.
We examined the relative clade stability in the most parsimo-nious trees by using jacknife monophyly analysis, ‘‘a simple, an-alytical tool designed to provide some insight into the effect ofhomoplasy on the relative stability of clades in most parsimonioustrees’’ (Siddall, 1995, p. 47). Jackknife monophyly analysis is a
data sampling technique that involves iterative cladistic analysisof the data matrix minus one taxon per iteration. This has certainadvantages over other methods, such as the bootstrap approach,in that it is not artificially biased by uninformative characters orthe relative number of synapomorphies (Siddall, 1995). However,Jackknife Monophyly Index (JMI) values are comparable onlywithin the data set (Siddall, 1995). The entire jackknife analysiswas performed by Option 3 of LANYON.EXE in the RandomCladistics software package (Siddall, 1997). The following Hen-nig86 (Farris, 1988) commands were used in the analysis: cc-.ie*.The program TREEVIEW (Page, 1996) was used to construct thecladogram.
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FIGURE 4—Oncopareia bredai Bosquet, 1854, sensu Tshudy, 1993 from the upper Maastrichtian Maastricht Formation, Kunrade, southeasternNetherlands; 1, lower view of right (major) and left (crusher) claws (IRScNB 90-28); 2, major claw fingers showing acicular dentition (IRScNB90-3b); 3, lower view of major claw (90-27b); 4, lower view of minor (crusher) claw (IRScNB 90-32a); 5–8, lower, inner, outer, upper views ofmajor claw (IRScNB 90-19f). Scale bar equals 1.0 cm.
479TSHUDY AND SORHANNUS—CONVERGENCE IN DECAPOD CLAWS
FIGURE 5—Oncopareia sp. from Miocene of Fiordo San Pablo, TresMontes, Chile (Tshudy, 1993). This Miocene specimen possesses bothmajor and minor claws very similar to those of recent Thaumastochelesand Thaumastochelopsis. Lower view of right (major) and left (minor)claws, including proximal limb segments [Museum National d’HistoireNaturelle (Paris) (MNHN) 4280]. Scale bar equals 1.0 cm.
EVOLUTIONARY AND BATHYMETRIC HISTORY OF FOUR LINEAGES
Pectinate claws are interpreted to have evolved independentlyin four lineages, once each in the infraorders Palinura and An-omura and twice in the Infraorder Astacidea. Both traditionalsystematic and cladistic (Fig. 6) approaches indicate that the
claw form evolved twice in the Astacidea within the family Ne-phropidae.
Lineage 1 of 4.—The palinurid Palaeopentacheles (Fig. 2.4)is a monospecific genus known by a single, well-preserved spec-imen from the Upper Jurassic of southern Germany [P. rotten-bacheri (Munster)]. Palaeopentacheles evolved from an eryonidancestor [inferred from Glaessner’s (1960, p. R411) phylogenyof the order Decapoda] with claws of similar proportions butwithout pectinate denticles (see Eryonidae in Glaessner, 1969,p. R469). No other genus in the Family Polychelidae (M. Jur.—Rec.) has pectinate dentition.
Lineages 2 and 3 of 4.—The nephropid Acanthacaris (Fig.2.2) has no fossil record, nor is there a fossil genus which bearsenough resemblance to Acanthacaris to be considered ancestralor relatively closely related. Glaessner (1932) erroneously alliedAcanthacaris (then Phoberus A. Milne-Edwards, 1881) with anew, non-pectinate, fossil genus, Palaeophoberus Glaessner,1932; this proposed lineage has been rejected by morphology-based intuitive and cladistic analyses (Tshudy, 1993; Tshudy andBabcock, 1997). Palaeophoberus now resides in the family Chi-lenophoberidae Tshudy and Babcock, 1997.
A variety of morphology-based cladistic analyses consistentlysupport the separateness of the Acanthacaris and the Oncopar-eia-Thaumastocheles/Thaumastochelopsis lineages (Tshudy,1993; Tshudy and Babcock, 1997; herein, Fig. 6). In this study,the exhaustive search for the shortest tree(s) yielded two mostparsimonious trees with a length (L) of 79 steps and consistency(CI) and retention (RI) indices of 75 and 68, respectively. Theonly difference between the two optimal phylogenies is seen inthe relationship within the Oncopareia, Thaumastocheles, Thau-mastochelopsis clade. In one of the most parsimonious trees(Fig. 6) the relationship is resolved while in the other optimaltree (not shown) the relative branching order between Oncopar-eia, Thaumastocheles, Thaumastochelopsis is unresolved.
The clade consisting of Oncopareia, Thaumastocheles andThaumastochelopsis is supported by two synaporphies, includ-ing (28-1) ‘‘abdominal pleura wider than long’’ and (40-2) ‘‘ma-jor claw palm shape bulbous.’’ The clade consisting of Homarusand nine other nephropid genera is supported by one synapo-morphy (33-1), the presence of ‘‘spines on posterolateral marginof telson.’’ The Thymops/Thymopsis clade is supported by twosynapomorphies, including (26-1) ‘‘parabranchial groove pres-ent’’ and (39-0) ‘‘exopod on maxilliped 3 absent or reduced.’’The Nephrops/Metanephrops clade is supported by five syna-pomorphies, including (5-2) ‘‘thoracic median carina present asdouble row,’’ (12-1) ‘‘cervical spine present,’’ (13-1) ‘‘postcer-vical spine present,’’ (18-2) ‘‘intermediate carina present, withintermediate spine,’’ and (27-2) ‘‘inferior groove present; curv-ing under mandibular insertion.’’
The JMI values are shown on the tree in Figure 6. The cladecomposed of Oncopareia, Thaumastocheles, and Thaumasto-chelopsis received a JMI value of 100. The other major clade,which consists of Homarus Weber, 1795, Thymopides Burukov-sky and Averin, 1976, Acanthacaris Bate, 1888, Thymops Hol-thuis, 1974, Thymopsis Holthuis, 1974, Nephropsis Wood-Ma-son, 1873, Nephropides Manning, 1969, Eunephrops Smith,1885, Nephrops Leach, 1814, and Metanephrops Jenkins, 1972,also shows a JMI value of 100. This clearly suggests that elon-gate, pectinate major claws have originated independently twicewithin the Nephropidae: once within the Oncopareia/Thaumas-tocheles/Thaumastochelopsis clade and once within the Acan-thacaris clade.
Acanthacaris is an eyeless lobster known only known fromdeep water. It has been collected in depths ranging from 293–1,463 m; most occurrences are 500 m and deeper (Holthuis,
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TABLE 1—Comparative morphology of pectinate claws in six decapod genera.
FIGURE 6—Relationships of several clawed lobster genera determined bythe cladistic method (exact search (ie*) algorithm of Hennig86). Clad-ogram shows synapomorphies, Jacknife Monophyly indices and indi-cates separateness of the Acanthacaris and the Oncopareia-Thaumas-tocheles/Thaumastochelopsis lineages.
1974). Burukovsky and Ckreko (1986) proposed an onshore-offshore migration for a lineage including the Upper Jurassic,shelf-dwelling Palaeophoberus and the recent Acanthacaris but,as mentioned above, this is not a valid lineage.
The existence of a lineage including, in evolutionary order,Hoploparia, Oncopareia, and Thaumastocheles (and now Thau-mastochelopsis) was proposed by Mertin (1941, p. 195) and issupported herein. A variety of cladistic analyses of external mor-phology are unable to reject this hypothesized lineage (Tshudy,1993; Tshudy and Babcock, 1997; herein). Oncopareia is mor-phologically intermediate between Hoploparia and Thaumasto-cheles; it has a cephalothorax like Hoploparia but an abdomenand claws like Thaumastocheles. The claws of Oncopareia, andits descendant (or at least closely related) genera, Thaumasto-cheles and Thaumastochelopsis, are pectinate.
The interpreted bathymetric history of the Hoploparia-On-copareia-Thaumastocheles/Thaumastochelopsis lineage is char-acterized by an onshore-offshore radiation similar to that inter-preted for diverse other animal groups (eryonid lobsters: Beur-len, 1931; Cambro-Ordovician invertebrate communities:Jablonski et al., 1984; crabs: Feldmann and Wilson, 1988; theclawed lobster Metanephrops: Feldmann and Tshudy, 1989;many others). Hoploparia is known widely from Cretaceousshallow-water (continental shelf depth) deposits. Hoploparia hasa ‘‘normal’’ clawed-lobster appearance and is considered ances-tral to several other shelf-dwelling lobsters, including the Maine
481TSHUDY AND SORHANNUS—CONVERGENCE IN DECAPOD CLAWS
FIGURE 7—Stratigraphic and bathymetric distribution of the Oncopareia-Thaumastochles/Thaumastochelopsis lineage, showing loss of shelf-dwelling tendancy in this lineage. a, O. esocinus; b, O.? lunatus; c, O.coesfeldiensis; d, O.? macrodactyla; e. Oncopareia sp. Mertin; f, g,Oncopareia sp. Tshudy; h, O. klintebjergensis; i, Oncopareia sp. Tshu-dy; j, Thaumastocheles japonicus; k, T. zaleucus; l, Thaumastochel-opsis wardi. Detailed list of occurrences in Appendix 1.
FIGURE 8—Stratigraphic and bathymetric distribution of the anomuranCtenocheles. a, C. inaequidens; b, Ctenocheles sp. Rasmussen; c, C.cultellus; d, e, Ctenocheles sp. Feldmann; f, Ctenocheles sp. Rathbun;g, C. cookei; h, C. rupeliensis; i, C. fragillis; j, Ctenocheles sp. Secre-tan; k, C. sclephros; l, C. compressus; m, Ctenocheles sp. B Holthuis;n, C. maorianus; o, C. leviceps; p, Ctenocheles sp. A Holthuis; q, C.balssi, r, C. serrifrons. Detailed list of occurrences in Appendix 1.
Lobster, Homarus americanus H. Milne-Edwards, 1837 (Glaes-sner, 1947; Secretan, 1964; Tshudy, 1993; others) and the fossilgenus Oncopareia. Oncopareia occurs in fine-grained sedi-ments, including chalks, marls and fine siliciclastics, which weredeposited in continental shelf environments. The geologicallyyoungest and only Tertiary Oncopareia, Oncopareia sp. Tshudyfrom the Miocene Tres Montes Formation in Chile (Fig. 5) andO. klintebjergensis from the Paleocene of Denmark, were bothshelf-dwellers. [Beurlen (1939) published a report of a shelf-dwelling Oligocene form, but this claw belongs to Ctenocheles(C. rupeliensis)]. It is uncertain whether or not Oncopareia in-habited the continental slope or rise because slope and rise sed-iments are rarely preserved. However, it is clear that the Onco-pareia-Thaumastocheles lineage has lost its shelf-dwelling ten-dancy because Thaumastochelopsis and Thaumastocheles areexclusively deep-dwellers (Fig. 7). Thaumastochelopsis isknown from 425 m (Bruce, 1988) and have reduced eyes. Thau-mastocheles japonicus is known from 200 and 400 meters andT. zaleucus is known from 720–825 meters; both are eyeless.While we are unable to absolutely reject the possibility that thelineage originated in deep water during or before the Late Cre-taceous, it seems most parsimonious to conclude that the eyedOncopareia invaded deeper depths and then was ultimately an-cestral to the eyeless Thaumastocheles.
The cause of the apparent radiation of this lineage into deepwater, and abandonment of the shelf, is unknown. Radiation intodeep water could simply have been a passive dispersal from sitesof origin in shallow water, but why abandonment of the shelf?Abandonment of the shelf could have been the result of global
marine regressions (Beurlen, 1931) or competition from the rap-idly-diversifying crabs (Glaessner, 1969, p.R 425), or both, butcertainly there are other possibilities.
Lineage 4 of 4.—Ctenocheles belongs to the Ctenochelidae,a family which includes five other genera (Manning and Felder,1991). Ctenocheles is the only genus in family Ctenochelidae,or even the Infraorder Anomura (L. Jur.—Rec.) (Glaessner,1969, p.R475), to have a pectinate major claw.
Whereas the Oncopareia-Thaumastocheles/Thaumastochel-opsis lineage shows an onshore-offshore trend, the record ofCtenocheles (Maast.—Rec.) does not (Fig. 8). The Cretaceousand Tertiary fossil record shows many inner shelf occurrences;any ancient slope and rise dwellers may never be known due tothe preservational biases against these forms. Recent Ctenoche-les species inhabit depths ranging from 35 to 800 meters.
ADAPTIVE SIGNIFICANCE OF CLAW FORM
Direct observations of decapods bearing pectinate claws havebeen few. Consequently, the adaptive significance of this clawform can be addressed only by speculation. Because Recentdecapods with pectinate claws commonly inhabit aphotic depthsand, moreover, that several of these taxa are totally eyeless, theinterpreted function(s) of these claws must accommodate livingblindly and on very fine-grained substrata.
Some or all of the taxa of interest are burrow-dwellers. Acan-thacaris has been photographed protruding from burrows at 800m (Burukovsky and Ckreko, 1986). Ctenocheles has been fos-silized in burrows (Jenkins, 1972). Isolated claws of RecentCtenocheles (Powell, 1949) and Thaumastocheles (Holthius,1974) have come up without their associated bodies in dredges,
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TABLE 2—Morphology-based character matrix analyzed cladistically. ? 5 data for character unavailable or indeterminable. For character descriptions, seeAppendix 2.
which suggests that the body was held in a burrow. Nevertheless,the major claw seems poorly adapted for excavating burrows.Various food-gathering functions seem more reasonable.
A wide variety of food-gathering functions seem plausible—even for decapods which are blind or inhabit aphotic depths.Predation and scavenging on benthic, soft-bodied animals (asnoted by Mertin, 1941, p.181) seems reasonable for all taxaconsidered, and could, conceivably, be accomplished by ‘‘grop-ing,’’ ‘‘raking’’ or ‘‘sieving.’’ A pectinate major claw, whenopened and held horizontally, would enable the animal to gropeover the seafloor in search of live or dead megafauna. Thisseems especially reasonable for Thaumastocheles, which have adactylus that opens to well beyond perpendicular to propodus(to nearly 180 degrees in T. zaleucus). The tremendous range ofmotion of the dactylus (moveable finger) in Thaumastochelesmight even permit the animal to rake across the bottom (Feld-mann and Tshudy, 1990). The same claw, when closed, wouldenable the animal to sieve through soft, very fine sediment (the‘‘raking’’ of Green, 1961, p.126) in order to collect organismsand detritus directly, or to resuspend these materials for collec-tion by the other feeding appendages. Variably-sized denticleswould enable the animal to change ‘‘mesh-size’’ by simplychanging the gape between the fingers.
Predation on swimming prey, particularly via ambush fromthe burrow opening, as described by Jenkins (1972, p.133) andmany others, also seems reasonable—even for blind taxa, which
could perhaps detect the movement or odor (J. Williams, 1997,personal commun.) of nearby swimming prey. Filter feeding asobserved in the thalassinid Upogebia Leach, 1814 has been sug-gested for Ctenocheles (C. McLay, 1997, personal commun.)and perhaps the other taxa.
ACKNOWLEDGMENTS
Specimens examined in this study were made available by R.Berglund, Bainbridge Island, Washington, A. Dhondt, InstitutRoyal des Sciences Naturelles de Belgique, R. Feldmann, KentState University, and J. Jagt, Natuurhistorisch Museum Maas-tricht. Anonymous reviewers significantly improved the originalsubmission.
REFERENCES
BATE, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S.‘‘Challenger’’ during the years 1873–1876. In Report on the ScientificResults of the Voyage of the H.M.S. Challenger. Zoology, 24, 942 p.
BERGLUND, R. E., AND R. M. FELDMANN. 1989. A new crab, Rogueusorri n. gen. and sp. (Decapoda: Brachyura), from the LookingglassFormation (Ulatisian Stage: lower middle Eocene) of southwesternOregon. Journal of Paleontology, 63(1):69–73.
BEURLEN, K. 1931. Die Besiedelung der Tiefsee. Naturund Museum 61:269–279.
. 1939. Neue Decapoden-Krebse aus dem ungarischen Tertiar. Pa-laeontologische Zeitschrift, 21(2):135–160.
BOSQUET, J. 1854. Les crustaces fossiles du terrain Cretace du Duche
483TSHUDY AND SORHANNUS—CONVERGENCE IN DECAPOD CLAWS
Limbourg. Verhandelingen van de Commission van de GeologischeBeschrijving en Kaart van Nederland, Deel II:10–371.
BREMER, B., R. K. JANSEN, B. OXELMAN, M. BACKLUND, H. LANTZ,AND K. KIM. 1999. More characters or more taxa for a robust phy-logeny—case study from the coffee family (Rubiaceae). SystematicBiology, 48(3):413–435.
BRUCE, A. J. 1988. Thaumastochelopsis wardi, gen. et sp. nov., a newblind deep-sea lobster from the Coral Sea (Crustacea:Decapoda:Ne-phropidea). Invertebrate Taxonomy, 2:903–914.
BURUKOVSKY, P. H., AND B. S. AVERIN. 1976. Bellator grobovi gen. etsp. n., a representative of the Family Nephropidae (Decapoda, Crus-tacea) from the Herd Island region in the subantarctic. ZoologicheskifiZhurnal, Moscow, 55:296–299.
, AND B. I. CKREKO, 1986. Arcaic lobsters. Priroda, 12:93–95. (InRussian)
CALMAN, W. T. 1913. A new species of crustacean genus Thaumasto-cheles. Annales and Magazine of Natural History, 8(12):229–233.
FARRIS, J. S. 1988. Hennig86, Version 1.5. Distributed by the author,Port Jefferson Station, New York.
FELDMANN, R. M. 1991. Decapod Crustacea from the Tapui GlauconiticSandstone (Bortonian: Middle Eocene) in the Waitaki valley, SouthIsland, New Zealand. New Zealand Journal of Geology and Geo-physics, 34:17–22.
, AND P. W. DUNCAN. 1992. Eocene decapod crustaceans fromSnowdrift Quarry. South Otago, New Zealand. New Zealand Journalof Geology and Geophysics, 35:455–461.
, AND D. M. TSHUDY. 1989. Evolutionary patterns in macrurousdecapod crustaceans from Cretaceous to early Cenozoic rocks of theJames Ross Island region, Antarctica, p. 183–195. In J. A. Crame(ed.), Origins and Evolution of the Antarctic Biota, Geological So-ciety Special Publication 47.
. 1990. Elongate claws in decapod crustacean taxa: ‘‘preadapta-tion’’ to scavenging in aphotic habitats. Geological Society of Amer-ica, Abstracts with Programs 22(5):10.
, AND M. T. WILSON. 1988. Eocene decapod crustaceans from Ant-arctica, p. 465–488. In R. M. Feldmann and M. O. Woodburne (eds.),Geological Society of America Memoir 169.
, J. J. JAGT, AND D. M. TSHUDY. 1990. Mededelingen rijks geo-logische dienst, 44(1):23–35.
FISCHER, W. (ed.). 1978. FAO Species Identification Sheets for FisheryPurposes; Western Central Atlantic (Fishing Area 31). Food and Ag-riculture Organization of the United Nations, Rome.
FRAZIER, W. J., AND D. R. SCHWIMMER. 1987. Regional Stratigraphy ofNorth America. Plenum Press, New York, 719 p.
FRITSCH, A., AND J. KAFKA. 1887. Die Crustaceen der bohmischen Krei-deformation. Prag, Selbstverlag, 53 p.
GLAESSNER, M. F. 1932. Zwei ungenugend bekannte mesozoische De-kapodenkrebse. Palaeotogische Zeitschrift, 14(1/2):108–121.
. 1947. Decapod Crustacea (Callianassidae) from the Eocene ofVictoria. Proceedings of the Royal Society of Victoria, 59:1–7.
. 1960. The fossil decapod Crustacea of New Zealand and theevolution of the Order Decapoda. Paleontological Bulletin, 31:1–79.
. 1969. Decapoda, p. R399–R651. In R.C. Moore (ed.), Treatiseon Invertebrate Paleontology, Pt. R, Arthropoda 4(2). Geological So-ciety of America and University of Kansas Press, Lawrence.
GREEN, J. 1961. A Biology of the Crustacea. H. F. and G. Witherby,Ltd., London, 180 p.
HOLTHUIS, L. B. 1967. A survey of the genus Ctenocheles (Crustacea:Decapoda, Callianassidae), with a discussion of its zoogeography andits occurrence in the Atlantic Ocean. Bulletin of Marine Science17(2):376–385.
. 1974. The lobsters of the Superfamily Nephropidae of the Atlan-tic Ocean (Crustacea:Decapoda). Bulletin of Marine Science 24(4):723–884.
. 1991. FAO Species Catalogue, 13, Marine Lobsters of the World.FAO Fisheries Synopsis 125(13), 292 p.
IMAIZUMI, R. 1957. An interesting crustacean remain, Ctenocheles su-jakui n. sp. from the Paleogene of Kyushu, Japan. Transactions of thePaleontological Society of Japan, 32:299–304.
JABLONSKI, D., J. J. SEPKOSKI, JR., D. J. BOTTJER, AND P. M. SHEEHAN.1984. Onshore-offshore patterns in the evolution of Phanerozoic shelfcommunities. Science, 222:1123–1125.
JAKOBSEN, S. L., AND J. S. L. COLLINS. 1979. Decapod Crustacea from
the Palaeocene of Zealand, Denmark. Proceedings of the GeologicalAssociation, 90(2):61–64.
JENKINS, R. J. F. 1972. Australian Fossil Decapod Crustacea: Faunaland Environmental Changes, Volume 1. Unpublished dissertation,University of Adelaide, 392 p.
KISHINOUYE, K. 1926. Two rare and remarkable forms of macrurousCrustacea from Japan. Japanese Journal of Zoology, 11:63–70.
LATREILLE, P. A. 1802–1803. Histoire naturelle, generales et particuli-ere, des crustaces et des insectes, Volume 4, 468 p.
LEACH, W. E. 1814. Crustaceology. In Brewster, D. The Edinburgh En-cyclopaedia, vol. 7:383–437 (not seen).
LE LOEFF, P., AND A. INTES. 1974. Les Thalassinidea (Crustacea, De-capoda) du Golfe de Guinee systematique ecologie. CahierO.R.S.T.O.M., serie Oceanographique, 12(1):17–69 (not seen).
MANNING, R. M. 1969. A new genus and species of lobster (Decapoda,Nephropidae) from the Caribbean Sea. Crustaceana, 17(3):303–309.
, AND D. L. FELDER, 1991. Revision of the American Callianas-sidae (Crustacea: Decapoda: Thalassinidea). Proceedings of the Bio-logical Society of Washington, 104(4):764–792.
MERTIN, H. 1941. Decapode Krebse aus dem subhercynen und Braun-schweiger Emscher und Untersenon sowie Bemerkungen uber einageverwandte Formen un der Oberkreide. Nova Acta Leopoldina, N.F.10, 264 p.
MILNE-EDWARDS, A. 1881. Description de quelques crustaces macrou-res provenant des grandes profondeurs de la mer des Antilles. Annalesdes Sciences Naturelles. Zoologie, Series 6, 11(4):1–16.
MILNE-EDWARDS, H. 1832. Recherches sur l’organisation et la classi-fication naturelle des Crustaces Decapodes. Annales des Sciences Na-turelles. Zoologie, 25:298–332 (not seen).
MUNSTER, G. G. ZU. 1839. Abbildung und Bescreibung der fossilenlangschwanzigen Krebse in den Kallschiefern von Bayern (not seen).
PAGE, R. D. M. 1996. TREEVIEW: An application to display phylo-genetic trees on personal computers. Computer Applications in theBiosciences 12:357–358.
PELSENEER, P. 1886. Notice sur les crustaces decapodes du Maestri-chtien du Limbourg. Bulletin du Musee Royal d’Histoire Naturellede Belgique, 4(3):161–175.
PHILIPPE, M., AND S. SECRETAN. 1971. Crustaces decapodes du Burdi-galien des Courennes (Vaucluse). Annales de Paleontologie Inverte-bres 57(2):115–141 (not seen).
POWELL, A. W. B. 1949. New species of Crustacea from New Zealandof the genera Scyllarus and Ctenocheles with notes on Lyreidus tri-dentatus. Records of the Auckland Institute and Museum 3(6):368–371.
RABALAIS, N. N. 1979. A new species of Ctencocheles (Crustacea: De-capoda: Thallassinidae) from the northwestern Gulf of Mexico. Pro-ceedings of the Biological Society of Washington, 92(2):294–306.
RASMUSSEN, H. W. 1971. Echinoid and crustacean burrows and theirdiagenetic significance in the Maastrichtian-Danian of Stevns Klint,Denmark. Lethaia 4:191–216.
RATHBUN, M. J. 1935. Fossil Crustacea of the Atlantic and Gulf CoastalPlain. Geological Society of America Special Papers, 2, 160 p.
ROBISON, R. A. AND R. L. KAESLER. 1987. Phylum Arthropoda, p. 205–269. In R. S. Boardman, A. H. Cheetham and A. J. Rowell (eds.),Fossil Invertebrates, Blackwell Scientific, Palo Alto, CA, 713 p.
RODRIQUES, SERGIO DE A. 1978. Ctenocheles holthuisi (Decapoda,Thalassinidea), a new remarkable mud shrimp from the AtlanticOcean. Crustaceana 34(2):113–120.
SCHLUTER, C. 1862. Die Macruren decapoden der Senon- und Ceno-manbildungen Westphalens. Zeitschrift der deutschen GeologischenGesellschaft, 14:702–749.
, AND W. V. D. MARCK, 1868. Neue Fische und Krebse aus derKreide v. Westfalen. Paleontolographica, 15:269–304.
SCHOLTZ, G., AND RICHTER, S. 1995. Phylogenetic systematics of thereptantian Decapoda (Crustacea, Malacostraca). Zoological Journal ofthe Linnean Society, 113:289–328.
SCHOPF, T. J. M.1980. Paleoceanography. Harvard University Press,Cambridge, Massachusetts, 341 p.
SECRETAN, S. 1964. Les crustaces decapodes du Jurasique Superieur etdu Cretace de Madagascar. Memoires du Museum National d’HistoireNaturelle, 19, 223p.
SIDDALL, M. E. 1995. Another monophyly index: revisiting the jack-knife. Cladistics, 11:35–56.
484 JOURNAL OF PALEONTOLOGY, V. 74, NO. 3, 2000
.1997. Randon cladistics. Version 4.0.3. (Ohio edition). Softwareavailable via ftp://zoo.toronto.edu.pub.
SMITH, S. I. 1885. Description of a new crustacean allied to Homarusand Nephrops. Proceedings of the United States National Museum,8:167–170.
THOMSON, C. W. 1873. Notes from the ‘‘Challenger’’ IV. Nature, 8:246–249.
TSHUDY, D. M. 1993. Taxonomy and Evolution of the Clawed LobsterFamilies Chilenophoberidae and Nephropidae. Unpublished disserta-tion, Kent State University, 472 p.
, AND L. E. BABCOCK. 1997. Morphology-based phylogenetic anal-ysis of the clawed lobsters (family Nephropidae and the new familyChilenophoberidae). Journal of Crustacean Biology, 17(2):253–263.
VON KNEBEL, W. 1907. Die Erymiden des Oberen Wiessen Jura VonSuddeutschland. Arhcive fur biontologie, 2:195–233.
WARD, M. 1945. A new crustacean. Memoirs of the Queensland Mu-seum, 12:134–135.
WEBER, F. 1795. Nomenclator entomologicus secundum Entomologiamsystematicum ill. Fabricii adjectis speciebus recens detectis et varie-tatibus, 171 p. (not seen).
WOOD-MASON, J. 1873. On Nephropsis stewarti, a new genus and spe-cies of macrurous crustaceans, dredged in deep water off the easterncoast of the Andaman Islands. Annals and Magazine of Natural His-tory, 4(12):59–64.
. 1874. Blind Crustacea. Proceedings of the Asiatic Society of Ben-gal, p. 180–181.
ACCEPTED 11 JANUARY 2000
APPENDIX 1
Annotated list of decapod taxa with pectinate claws
Infraorder Astacidea Latreille, 1803Family Nephropidae Dana, 1852, sensu Tshudy and Babcock, 1997
Genus Acanthacaris Bate, 1888 (Recent)Figure 2.2
A. caeca (A. Milne-Edwards,1881); Recent, Gulf of Mexico and Ca-ribbean Sea, including Straits of Florida (293–878 m, with most recordsfrom 550–825 m) (Holthuis, 1991, p. 27). A burrower, but nothing ofits morphology would suggest a burrowing mode of life (Holthuis,1974, p. 752).
A. tenuimana Bate, 1888; Recent, Indo-West Pacific (600–2,161 m)(Holthuis, 1991, p. 28).
Genus Oncopareia Bosquet, 1854, sensu Tshudyredefinition of genus in Tshudy (1993, p. 275); remarks on
synonomies in Tshudy (1993, p. 275–286) (Turonian-Miocene)Figures 3–5
O. bredai Bosquet, 1854, sensu Tshudy; Maastrichtian white chalkof the Netherlands and Belgium (Feldmann et al., 1990); redescriptionof type species O. bredai in Tshudy (1993, p. 286–298).
O. coesfeldiensis Schluter, 1862; lower and upper Senonian marls ofnorthcentral and northwestern Germany [e.g., yellowish marl with abun-dant glauconite (Schluter, 1862, p. 730)]; glauconite indicates diagenesisin 125–250 m (Schopf, 1980, p. 56). See also reports in Mertin (1941,p. 182).
O. esocinus (Fritsch and Kafka, 1887); Upper Turonian chalk of Po-debrady, north-central Czechoslovakia; holotype is an abdomen and as-sociated major claw. See also reports of O. cf. esocinus from SalzbergMarl (lower-upper Senonian boundary) in Mertin (1941, p. 181).
O. klintebjergensis Jakobsen and Collins, 1979; Paleocene LellingeGreensand Limestone (‘‘a fine grained sand with a dense conglomera-tion of comminuted shelly fragments some of which are glauconitized’’)(Jakobsen and Collins, p. 61) of Klintebjerg, east Zealand Island, south-eastern Denmark; known by anterior portion of one cephalothorax andincomplete fingers of an associated claw.
O.? lunatus Fritsch and Kafka, 1887; Turonian of Weissen Berge,near Prague, Czechoslovakia; claws unknown.
O.? macrodactyla (Schluter and Marck, 1868); upper lower Senonian,Dulman, northwestern Germany; known by a cephalothorax and asso-ciated claws.
Oncopareia sp. Mertin, 1941; upper Senonian of Konigslutter, eastof Brauschweiger, northcentral Germany; known by incomplete majorand minor claws (Mertin, 1941, p. 187).
Oncopareia sp. Tshudy, 1993; upper Maastrichtian chalk of Kunrade,southeastern Netherlands; known by three palms (mani), one complete(Tshudy, 1993, p. 303).
Oncopareia sp. Tshudy, 1993; Miocene (gray, very fine sandstone)of Fiordo San Pablo, Tres Montes, Chile; known by partial sternum andleft and right claws (Tshudy, 1993, p. 303).
Genus Thaumastocheles Wood-Mason, 1874 (Recent)Figure 2.1
T. japonicus Calman, 1913; Recent, Japanese waters (366–700 m)(Holthuis, 1991, p. 23).
T. zaleucus (Thomson, 1873); Recent, ‘‘West Indian region’’ (640–1,054 m, soft muds/oozes; ‘‘possibly a burrowing species’’) (Holthuis,1991, p. 24).
Genus Thaumastochelopsis Bruce, 1988 (Recent)T. wardi Bruce, 1988; Recent, Marian Plateau, off Townsville,
northeast Australia (425 m)Infraorder Anomura H. Milne-Edwards, 1832
Family Ctenochelidae Manning and Felder, 1991Genus Ctenocheles Kishinouye, 1926 (Upper Cretaceous-Recent)
Figure 2.3
C. bakeri (Glaessner, 1947); Middle Paleocene Pebble Point beds ofVictoria, Australia (Jenkins, 1972, p. 61, 117). Known by many claws.
C. balssi Kishinouye, 1926; Recent, Japan: Sagami Bay and westcoast of Honshu (119–800 m) (Holthuis, 1967, p. 377).
C. collini Ward, 1945; Recent, Mud Island, Moreton Bay, Queens-land, Australia (depth unknown); known by three complete specimensand two fragments (Holthuis, 1967, p. 377).
C. compressus Jenkins, 1972, p. 122; upper lower Miocene–lowermiddle Miocene of Murray river basin, South Australia: Cadell MarlLens of Morgan Limestone at Morgan [inner shelf deposit (Jenkins,1972, p. 41)], Upper Member of Morgan Limestone at Morgan [innershelf deposit (Jenkins, 1972, p. 41–42)]. Known by numerous clawfingers.
C. cookei (Rathbun, 1935); lower Eocene Sucarnoochee beds (nowPorters Creek Clay) (inner shelf deposits; Frazier and Schwimmer, 1987,p. 546) of Midway Fm., Alabama; known by several claw fragments.
C. cultellus (Rathbun, 1935); Paleocene and/or Eocene sediments (in-terpreted as shallow shelf deposits) of several Atlantic Coastal Plainand Gulf Coastal Plain localities; known only by claw finger fragments.
C. fragilis Jenkins, 1972, p. 118; late Upper Oligocene or early LowerMiocene to middle Lower Miocene; Jan Juc Marl and Puebla Clay(inner shelf deposits) of South Australia. Known by several incompleteclaws.
C. holthuisi Rodrigues, 1978; off mouth of Rio Sao Franscisco, Brazil(75 m).
C. inaequidens (Pelseneer, 1886); Lower Maastrichtian KunradeLimestone facies of the Maastricht Formation, Limburg, The Nether-lands; known by one major claw.
C. leviceps Rabalais, 1979; Recent, Gulf of Mexico (10–49 m) byboth bottom grab (three specimens) and otter trawl (two specimens).
C. madagascariensis Secretan, 1964, p. 149; Cretaceous: Cenoman-ian-Campanian (calcareous ferruginous sandstone) of Madagascar;known by several isolated claws.
C. maorianus Powell, 1949; Recent (35–73 m in Holthuis, 1967, p.379) and upper Pleistocene (C. cf. maorianus Glaessner, 1960) of north-ern New Zealand; Recent ones probably burrowers in soft mud (Hol-thuis, 1967, p. 378). Completely known.
C. rupeliensis (Beurlen, 1939); middle Oligocene Kisceller Tegel(clay) (interpreted as a deepwater deposit), Obuda (near Budapest) Hun-gary; a few claws described.
C. sclephros Jenkins, 1972, p. 124; middle lower Miocene-late lower(?early middle) Miocene; various horizons in Morgan Limestone, South
485TSHUDY AND SORHANNUS—CONVERGENCE IN DECAPOD CLAWS
Australia (interpreted as inner shelf deposits). Known by numerous in-complete claws.
C. serrifrons Le Loeuff and Intes, 1974; Recent, continental shelf offIvory Coast (Rabalais, 1979).
C. sujakui imaizumi, 1957, Eocene (Jenkins, 1972) beds of KishimaFormation of Kyushu, Japan; known by claws.
C. victor Glaessner, 1947; Upper Paleocene Rivernook Member ofDilwyn Formation, southern Victoria, Australia (Jenkins, 1972, p. 117).Known by claw fingers only.
Ctenocheles sp. Rasmussen, 1971; lowermost Danian of Stevns Klint(chalks and marls interpreted as inner shelf deposits); known by fingersof the major claw.
Ctenocheles sp. (Rathbun, 1935); lower Eocene Sucarnoochee bedsof Midway Fm., Alabama; known by one right manus with stump ofpropodus.
Ctenocheles sp. A Holthuis, 1967; Recent, Straits of Florida nearBimini (297–406 m) and north coast of Panama (109–295 m); knownby one left claw from each location.
Ctenocheles sp. B Holthuis, 1967, Recent, Atlantic Ocean off Colom-bia (62–100 m); known by one claw each from two sites.
Ctenocheles sp. Feldmann, 1991; middle Eocene Tapui GlauconiticSandstone (interpreted as inner shelf deposit) of North Otago, SouthIsland, New Zealand; known by two specimens, including a nearly com-plete propodus.
Ctenocheles sp. Feldmann and Duncan, 1992; middle Eocene fine,tan sandstone concretion (interpreted as inner shelf deposit); known bythree separate, incomplete fingers preserved together in matrix.
Ctenocheles sp. Philippe and Secretan, 1971; lower Miocene (Bur-digalien) of S. France (shallow marine sands).
Ctenocheles sp. (unpublished); lower Eocene Elkton Fm., DouglasCo., SW Oregon; minor claw and mold of second (major?) claw in darkgray coarse siltstone [part of a downslope-mixed fauna (Tucker, 1997,personal commun.); bathymetry of specimen unknown].
Ctenocheles sp. (unpublished); lower Eocene Lookingglass Fm. (Ten-mile Mbr.), Coos Co., SW Oregon; major claw in gray coarse siltsone[part of a downslope-mixed fauna (Baldwin, 1974, in Berglund andFeldmann, 1989)]; bathymetry of specimen unknown.
APPENDIX 2
Characters used in cladistic analysis. The reader is referred to Tshudy(1993:453–472) for discussions of these characters (some states recodedherein).1. Rostrum.
0—dorsoventrally compressed1—laterally compressed
2. Lateral rostral spines0—absent1—present
3. Dorsal rostral spines0—absent1—present
4. Cephalic median carina0—absent1—present as single row2—present as double row3—0/24—0/1
5. Thoracic median carina0—absent1—present as single row2—present as double row
6. Subdorsal carina0—absent1—present
7. Gastric tubercle0—absent1—present2—0/1
8. Supraorbital ornamentation0—absent1—present as spine only2—present as spine and carina3—1/2
9. Supraorbital carina0—absent or without very large spines and not extending to
postcervical groove1—with very large spines and extending to postcervical groove
10. Postorbital spine0–absent1—present2—0/1
11. Gastrolateral spine0—absent1—present2—0/1
12. Cervical spine0—absent1—present
13. Postcervical spine0—absent1—present2—0/1
14. Antennal carina0—absent1—short (extends to less than or equal to one-half the distance to
cervical groove)2—long (extends greater than one-half the distance to cervical
groove)3—0/1/2
15. Hepatic spine0—absent1—present2—0/1
16. Lateral carina0—absent1—present
17. Branchial carina0—absent1—present
18. Intermediate carina0—absent1—present, without intermediate spine2—present, with intermediate spine
19. Inferior carina0—absent1—present
20. Carapace rigidity0—not rigid1—rigid
21. Postcervical groove0—originating near dorsomedian; extending anteroventrally toward
cervical groove1—originating near dorsomedian; extending anteroventrally to cer-
vical groove2—originating on dorsomedian; joined by branchiocardiac groove;
combined groove bifurcating at midheight; postcervical groove extend-ing toward/to cervical groove as intercervical groove sensu Holthuis(1974); branchiocardiac groove, postcervical groove sensu Holthuis(1974) extending to hepatic groove
3—originating on dorsomedian; joined by branchiocardiac groove;postcervical groove extending toward cervical groove as intercervicalgroove sensu Holthuis (1974); no ventral extension of branchiocardiacgroove, postcervical groove sensu Holthuis (1974)
4—originating near dorsomedian; extending to hepatic groove5—2/3
