Contributions to Zoology, 72 (2-3) 181-186 (2003)

SPB Academic Publishing bv, The Hague

Progress on the fossil Xanthoidea MacLeay, 1838 (Decapoda, Brachyura)

Carrie+E. Schweitzer

Kent State Stark Department of Geology, University, Campus, 6000 Frank Ave. NW, Canton, Ohio 44720, USA

Keywords: : Decapoda, Brachyura, Xanthoidea, Xanthidae, proxy characters

Abstract notable recent works by neontologists do provide dorsal information carapace (Ng, 1998; Davie, 2002)

Identification and utilization of characters in extant fa- proxy and are thus quite useful to the paleontologist as is has milies been accomplished with success for fossil members the work on the sternum, abdomen, and pereiopods of the Calappidae, Hepatidae, Necrocarcinidae, Matutidae, and by Guinot (1968a, b, 1978, 1979) and others. Hexapodidae. The fossil and extant Raninidaehave been revised Because many xanthoid families are using primarily hard-part morphology preserved in the fossil superficially record and similar in of dorsal although formal designation of “proxy characters” has amazingly terms carapace not occurred. Hard-part characters can be used to differentiate also morphology, it can be very difficult to place fossil among members of the Atelecyclidae and the Cheiragonidae. xanthoids within a family. In addition, the dorsal the method of Thus, using proxy characters is demonstrably carapace morphology of members of subfamilies suitable for like the Xanthoidea. application to a difficult group within the same family are often extremely diver-

in gent, as the Goneplacidae, Panopeidae, and Pi- Introduction lumnidae (Davie, 2002; Karasawa & Kato, in press).

Thus, the framing of a family-level diagnosis for

Fossil Xanthoidea have traditionally been referred these xanthoid families based upon features of the to the 1838 dorsal is Xanthidae MacLeay, senso lato, and carapace nearly impossible. occasionally arrayed within the various xanthid The application of the work of Guinot (1968a, subfamilies (Rathbun, 1930; Glaessner, 1969). Over b, 1978, 1979) on the sternal, abdominal, and first the past four decades, biologists have separated the pereiopod morphology of brachyurans has been of

in extant Xanthoidea into several families. At the time great assistance family placement of fossil xan- of this writing, twelve extant families are widely thoids. For example, members of the Carpiliidae

& & have been in the fossil record based recognised (Martin Davis, 2001; Ng Liao, 2002; recognized upon

in Schweitzer, press), and an additional two ex- the merus of the major cheliped being fused to the tinct families are known (Schweitzer, in press) basis-ischium and the direct articulation ofthe merus (Table 1). Extant xanthoids are diagnosed based and coxa (Guinot, 1968a, b). Differences in the

features of the of fusion of male abdominal somites upon eyes, antennae, mouthparts, degree has gonopores, gonopods, and the degree of fusion of been quite useful in placing fossils into the Hexapo- the articles of the first pereiopods, the abdomen, didae and Goneplacidae and promises to be useful and the sternum (Guinot, 1979; Davie, 2002). Most for the Panopeidae and Pilumnidae as well (Schweit- of these features rarely, if ever, fossilize. Thus, zer & Feldmann, 2001; Davie, 2002; Karasawa &

are forced Kato, in Paleontologists to rely primarily upon press). features of the dorsal In Even articles of the carapace. some serendipi- though pereiopods and ele- tous cases, the entire cheliped, sternum, and abdo- ments of the sternum and abdomenare useful, they men much are preserved and can be used as well. Some are less commonly preserved than the dorsal

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In often carapace. addition, they are incompletely not designate proxy characters. Hard-part charac-

such Thus, features, as whether ters can also be used to differentiate preserved. important among mem-

in the eighth sternite is visible dorsal view or not, bers of the Atelecyclidae Ortmann, 1893 and the

be may not observable. The latter feature is im- Cheiragonidae Ortmann, 1893 (Schweitzer & Salva,

portant for classifying goneplacids and pilumnids 2000). Thus, the method is demonstrably suitable

Karasawa & in (Davie, 2002; Kato, press). Conse- for application to a difficult group like the Xan-

quently, it is important to try to identify a suite of thoidea.

characters of the primarily dorsal carapace, but

of including some features the chclipeds, sternum,

and abdomen, that are diagnostic for a particular Examples

family. These suites of dorsal carapace characters

called characters are proxy (Schweitzer & Feldmann, The Carpiliidae are clearly recognizable if the el-

2000; Schweitzer, in The term char- press). ‘proxy ements of the first pereiopods are preserved; even

acter’ is used because these characters track with if the they are not, smooth, vaulted carapace; long, the that soft-part morphology neontologists typi- convex anterolateral margins; lobate front; narrow

use. These characters be found in cally proxy may posterior margin; posterolateral to posterior mar- both and fossil members extant of a taxon, but they gin angle of 25-30 degrees; and fused male abdo-

are seldom if noted ever by neontologists when minal somites 3-5 are diagnostic for the family

are described. in decapods (Schweitzer, press). Although there are certainly

Definition, choice, and use of proxy characters other smooth, xanthoid decapods in the fossil record,

is relatively straightforward. Members of extant of this entire suite characters is diagnostic only for defined decapod families, as by biologists based the Carpiliidae and not for other smooth xanthoids. lar- upon soft-part morphology, serology, genetics, Fossil Pseudoziidae are not easily distinguished

and are examined either in from vae, sperm, museum carpiliids; however, several features can be collections or from illustrations in biological lit- used to differentiate the two families. Pseudoziids

erature. Then, distinctive features of the dorsal have smooth carapaces; short anterolateral margins; be identified that characterize carapace can most a posterolateral to posterior margin angle of 40-45 or all members of the In group. many cases, sev- degrees; a broad posterior margin; an incompletely eral characters must be used as a suite. together fused merus and basis-ischium, and no fusion of

Often there is not one dorsal char- single carapace abdominal somites (Schweitzer, in press). that identifies members of if there acter a family; MembersofVia’s (1959) subfamily Zanthopsinae classification of the Xanthoideawould be was, not (= Xanthopsinae) have often been placed within so difficult! The of characters that have types proven the Carpiliidae (Schweitzer, 2000); however, they to be useful thus far are various length/width ra- are morphologically distinct. The merus is not fused tios; the shape, length, and ornament of the front to the and the basis-ischium, merus does not ar- and anterolateral margins; the angle of the poste- ticulate with the directly coxa; thus, they are easily rolateral to the the rela- If margin posterior margin; excluded from the Carpiliidae. the chelipeds are tive size of the and ornament carapace regions; shape not preserved, zanthopsines may be recognized of the orbits; and the of maximum width because have position they a relatively smooth carapace that of the carapace. is not separated into distinctive regions; shallow identification and utilization of characters proxy grooves; a tightly convex anterolateral margin; a in extant families has been with suc- male accomplished quadri-lobed front; and fused abdominal so- cess for fossil members of the mites Calappidae, Hepa- (Schweitzer, in press). They form a distinc- tidae, Necrocarcinidae, Matutidae, and tive Hexapodidae group united by these morphological features.

(Schweitzer & Feldmann, 2000, 2001). Tucker Several fossil genera cannot be accommodated

(1998) revised the fossil and extant Raninidae de known by any family; a new family has been erected

Haan, 1839, using primarily hard-part morphology to contain them (Schweitzer, in press). Palaeo- preservable in the fossil record although she did xanthopsis Beurlen, 1958, Verrucoides Vega et al.,

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and two all characterized to take into account these mor- 2001, new genera are by vastly divergent exhibiting a broad carapace; extremely deep, V- phologies.

the and branchial Within those families that have dorsal shaped grooves separating gastric divergent

nodes the members with each regions; large on carapaceregions; a long carapace morphologies, usually anterolateral 4 5 distinctive within the margin with or pronounced spines; morphology are placed same and a last anterolateralspine that is especially long subfamily of the family. Thus, at least for the Gone-

directed These and it and laterally or posterolaterally. placidae, Panopeidae, Pilumnidae, appears features are quite distinct from any other known that proxy characters will need to be developed at xanthoid; thus, a new family has been erected to the subfamily, not family, level. Each subfamily

would then have contain these genera (Schweitzer, in press). to be considered when working

the Members of the Hexapodidae are recognized by with these families. This greatly complicates their that is widest a as the number of subfamilies rectangular carapace distally; work, recognized

bi-lobed rostrum that extends beyond the orbits; a within a family varies widely, depending on the

and because consider fronto-orbital width that partially occupies the en- author, many authors some

ab- subfamilies to be distinct families while others tire frontal margin of the carapace; fused male

dominal somites; and sternite and pereiopod 5 lost maintain them as subfamilies. For example, Ng &

(Schweitzer & Feldmann, 2001). These features can Liao (2002) considered the Pseudoziidae to be a

permit hexapodids to be distinguished from pin- distinct family, while Davie (2002) considered it

the notherids and some goneplacids which also have to be a subfamily of Goneplacidae. Davie (2002)

considered the Eumedonidaeto be of small, rectangular carapaces. a subfamily

& Those are the success stories. However, there the Pilumnidae, while Martin Davis (2001) main-

are other families that pose major problems for tained it as a separate family. Clearly, working with

palaeontologists. Within the Goneplacidae, there xanthoids is anything but simple.

several all of are dorsal carapace morphologies,

which are quite different from one another (Schweit-

zer, 2000; Davie, 2002; Karasawa & Kato, in press). Implications

Currently, Karasawa and Kato are working on the

phylogeny of this family as well as attempting to With all of these complications and difficulties,

discern features that can be used to distinguish why not just refer problematic xanthoids to the

members of each constituent subfamily. It appears Goneplacidae or Xanfhidae sensu lato as has been

that within the Goneplacidae, the nature of the male done in the past? I have done this myself, for

abdomen and whether the male abdomen with the or not example, problematic genera Amydrocar-

covers sternite 8 will be important at the subfamily cinus Schweitzer et al., 2002 and LobonotusA. Milne

level (Tucker & Feldmann, 1990; Davie, 2002; Edwards, 1864. I think, however, that it is time to

Karasawa & Kato, in press). Within the Panopeidae, tackle these difficult fossil xanthoids, because these

there is vast of dorsal the similarly a range carapace fossils have manifold implications for history

morphology, and within that family, the position of brachyuran evolution, not to mention for the

of the male gonopore ranges from coxal to coxal- ongoing revision of the Decapoda volume of the

In of Treatise sternal (Davie, 2002). addition, the features on Invertebrate Paleontology.

the dorsal and the in the carapace gonopores pano- Currently, we do not have good constraints on

peids overlap with some other families of xanthids. the origin of the Xanthoidea and the subsequent

Thus, placement of fossils into the Panopeidae is radiation of this robust superfamily. As of this writ-

currently extremely difficult. Similarly, the Pilum- ing, only the Goneplacidae and possibly the Hexa-

nidae is a difficult family for palaeontologists. The podidae have confirmed Cretaceous records; two

dorsal from xanthid-like (sensu referred to the carapace ranges ( genera formerly Xanthidae, Etyus

stricto) to ovate to bizarrely produced rostrally into Mantell, 1822, and Xanthosia Bell, 1863, havebeen

a bi-pronged extension (Davie, 2002). Any dorsal referred to the Etyidae Guinot & Tavares, 2001,

carapace diagnosis for the pilumnids would need and are considered to be primitive podotrematous

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Table I. List ofxanthoid families, recognized at the time ofthis writing (March, 2003)

Family Geologic Range

*Carpiliidae Ortmann, 18931893 Eocene-RecentEocene-Recent

Eriphiidae MacLeay, 1838 (= MenippidaeOrtmann, 1893) unresolved

Eumedonidae Dana, 1853 (considered by Ng & Clark (2000)

the Pilumnidae unknown and Davie (2002) as a subfamily ofthe Pilumnidae

*Goneplacidae MacLeay, 1838 Cretaceous - Recent

- *Hexapodidae Miers, 1886 Cretaceous?; Paleocene - Recent

*Panopeidae Ortmann, 1893 Eocene - Recent

*PilumnidaeSamouelle, 1819 Eocene - Recent

Platyxanthidae Guinot, 19771977 unknown

*Pseudoziidae Alcock, 1898 Eocene - Recent

PseudorhombilidaeAlcock, 1900"1900 unknown

TrapcziidaeTrapeziidae Miers, 1886 Miocene - recent

*Xanthidae MacLeay, 1838 unresolved fZanthopsidaetZanthopsidae Via, 1959 Paleocene - Miocene

{NewfNew family including Palaeoxanthopsis Beurlen, 1958 Cretaceous (Maastrichtian)-Eocene

* Schweitzer indicates families with a fossil record; f indicates extinct families. {See Glaessner, 1969;Muller, 1984; Schweitzer, 2000;

in & Feldmann, 2001; Schweitzer, press.

The it have been earliest for which crabs (Guinot & Tavares, 2001). Hexapodidae occurred; may Eocene,

and was formerly a subfamily of the Goneplacidae; these we have few fossils, it may have been Pale- two families are closely related and they appear to ocene. The paucity of Paleocene rocks and fossils have diverged by the Paleocene at the latest. Cre- makes it difficult to test this hypothesis, but work taceous rocks have also yielded a new family that is planned by colleagues and myself to investigate

affini- embraces Palaeoxanthopsis; however, the Paleocene occurrences in Europe. The timing of

of and its ties this family relationship to other xan- appearanceand radiations among the other xanthoid thoid families are not clear. Interestingly, taxa within families are similarly unknown. Only by placing this new family resemble some members of the fossil xanthoids within the appropriate family, us-

Rhachiosoma this of of Portunidae Rafmesque, 1815, e.g., ing proxy characters, can problem timing

well the and radiation these families be Woodward, 1871, as as Xanthoidea; per- divergence among

the of the and haps common ancestor Xanthoidea addressed. Providing a well-documented fossil re-

Portunoidea havebeen simi- cord for the various xanthoid families will may morphologically provide

which lar to members of this new family. a means by to test biological hypotheses about

the well of based By the Paleocene, Zanthopsidae was timing divergences upon molecular clock established and by the Eocene, four additional fami- evidence. lies abundant (Table 1) were sufficiently to pro-

robust duce a fossil record. Based uponpreliminary work, it appears that the Eriphiidae and Panopeidae Future studies also have Eocene records (Glaessner, 1969). Clearly, then, of the extant Xanthoideahad The of many appeared use proxy characters and other biologically the that the radiation by Eocene, so events must important hard-part characters to evaluate fossil have occurred before that time. and xanthoids to the and Biological pale- may help clarify nature extent ontological evidence suggests that the Carpiliidae, of the end-Cretaceous extinction event and the sub-

and are It Eriphiidae, Pseudoziidae, Zanthopsidae sequent recovery. is known already that at least

related (Schweitzer, in Eo- closely press); all have one extant xanthoid family, and possibly two, has cene records. Thus, share a common they must a Cretaceous record, suggesting that these animals ancestor and must have before that time. diverged were minimally affected by the K/T event. In addi- it is However, not known when that divergence tion, several xanthoid families have robust fossil

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records by Eocene time. The question is, when did References these xanthoid families appear? Was it during the Paleocene or Eocene? And is the Goneplacidae, or Alcock A. 1898. The family Xanthidae: the Brachyura Cyclo-

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