Contributions to Zoology, 72 (2-3) III-I17 (2003)
SPB Academic Publishing bv, The Hague
Interpreting ecology and physiology of fossil decapod crustaceans
Rodney+M. Feldmann
Department of Geology, Kent State University, Kent, Ohio 44242, USA
Keywords:: Crustacea, Decapoda, Mesozoic, Cenozoic, behavior
Abstract that surface the rubbing against carapace margin
(stridulation), it is possible to identify similar struc-
Decapods are the most of in diverseand complex group crustaceans, tures some fossil remains and to assign to them for life in all adapted parts of the marine environment, many a similar function (Feldmann & Bearlin, 1988). aquatic habitats, and some terrestrial niches. With this diversity the function of of Similarly,, grooming appendages, life of of has styles, a vast range morphotypes decapods characteristic of and other evolved, almost in many shrimp decapods, exploiting every imaginable variation mor- be the phology of the complex exoskeleton that characterizes them. can inferred on basis of modern analogy.
of Many variants are a to Much of the work the morphological response exploiting of paleontologists is character- a particular niche in which the organisms live or an adaptation ized by this functional morphological approach. to particularbehavioral characteristics. Assessing the significance Form and function of the and carapace of append- morphological variation in the fossil record is challenging ages has been the subject of voluminous literature. because of the taphonomic overprint that results in loss ofsoft The tissue, two most works are those of preservation ofpartial remains of hard parts, and vastly comprehensive
reduced numbers Schafer and Manton and ofpreserved individuals as contrasted to the (1972) (1977) their ef-
once-living population. The of the is to purpose present paper forts will not be repeated here. Suffice it that feed- of that be useful in identify aspects morphology may interpreting ing behavior may be deduced from the of the shape behavioral of the its responses organism to environment, claws and the nature of their denticles, and life w,th style, primary emphasis on morphological features of the exo-
including swimming, free can skeleton that are burrowing, not expressed onall individualsbut that occur living, at low, and be from the of the unpredictable, frequencies. interpreted shape carapace. Nu-
merous examples are given within those familiar
works. Thus, form reflects behavior. Introduction: functional morphology Some crabs exhibit behavior that permits them
to live within other organisms. Perhaps the best When known are the assigning significance to a specific morpho- Pinnotheridae, tiny crabs which may logical attribute, neontologists are able to study be endosymbiotic within bivalve, gastropod, and the living organisms, observe their behavior, and polyplacophoran molluscs, brachiopods, echino-
interpret the functional of that mor- derms, and significance enteropneusts, ascidians, worms, bur-
phological attribute. Because rows of worms other paleontologists can and decapods (Ross, 1983).
observe behavior Their has been only by analogy with living or- presence noted since the time of ganisms, they rely on deductive reasoning to inter- Aristotle (Aristotle, 1862). Several free-living pin- pret the fossil record. Thus, by noting the crenulated notherids havebeen described from the fossil record
re gions the antennal & near base of some lobsters or (Schweitzer Feldmann, 2001) but there has been along the inner surface of the propodus of some only one report of pinnotherids within the host extant crabs, and observing the behavioral pattern organism (Zullo & drivers, 1969). Another endo-
111 living animals modern of generating a rasping sound by symbiotic association is typified by gall
Downloaded from Brill.com09/24/2021 08:38:58PM via free access I 12 R.M. Feldmann - Interpreting ecology and physiology
of crabs, the Haplocarcinidae. Some these tiny crabs purely nektonic life, or for other life styles have
induce form been the perform a specific behavior to corals to characterized on basis of morphological
a gall that surrounds and protects the animal (Ross, attributes. Many of these types of characters not
fossil should 1983). Their presence in the record only have functional significance but, because of
be assured because entombed their in the useful they are essentially ubiquitous presence population, are
coral and the within tissue; despite that, none has been in identifying classifying organism.
reported from the fossil record. However, there are Other characters, associated primarily with on-
several of crabs within exhibit that reports tiny, cryptic living togeny, may systematic variation can
and coral that of be as in of the sponge structures are note (Collins expressed changes morphometries
& Muller & allometric be definable Wicrzbowski, 1985; Collins, 1991; organism. Thus, growth may
Beschin ct ah, 1996, 2001). These crabs often form within a collection of fossil decapods. Schweitzer
diverse Fcldmann a very assemblage but are almost completely & (2000) recognized allometric growth
unknown because tend be in the Eocene Chaceon they are very small, to patterns geryonid, peruvianus
in that it preserved the same fashion as the surrounding (d’Orbigny) helped distinguish from a con-
coral, and are extremely difficult to detect. temporaneous portunid in which the proportions
of the adult portunids were similar to the propor-
tions of the juvenile geryonid.
Frequency of occurrence of morphological Characters that are gender-specific are anticipated
traits within a species to occur with a frequency equal to the gender dis-
tribution within the taxon. In decapods, that ratio
A different approach to understanding the relation- is quite variable and in the fossil record it may be
ship between morphology and behavioral adapta- difficult to discern. Schweitzer Hopkins & Feldmann
tions be taken the of (1997) studied a suite of of Eoce- may by assessing frequency large specimens
mud from State occurrence of a particular morphological trait within ne-Oligocene shrimp Washington
and determined that 65% of the claws a fossil population. Morphological traits may be (USA) were
in individuals within male and 35% were female. This is expressed all a species or may study particu-
be found in the ori- because it demonstrated that only a subset, depending upon larly significant two
named in gin of the trait. previously species were, fact, sexual di-
of a clallamensis Morphological attributes that are necessary for morphs single taxon, Callianopsis
In the functioning of all individuals within a taxon (Withers). a more recent study of Pleistocene-
should be discernable on all members of the taxon, Holocene decapods from Guam, Schweitzer et al.
if the appropriate region is preserved. Swimming (2002) noted that the ratio of males to females,
behaviour in nektobenthic crabs, for example, is determined on specimens exhibiting well-preserved
often facilitated by expansion and flattening of the pleons, ranged from 50% to 89%. Some shrimp,
distal elements of some of the pereiopods and trans- \ notably the Pandaloideawithin the Caridea, undergo
sexual reversal within their The verse elongation of the carapace (Schafer, 1972). ontogeny. shrimp
to Similarly, decapods adapted for burrowing, for a typically grow maturity as males, undergo sexual
- I. 1 - view of with bar Fig. dorsal extant Callinectes sp. Balanus epibionts (scale equals 10 mm); 2 dorsal view ofLobocarcimis pustulosus Feldmann & Fordyce, with arrows showing the position ofa large, straight serpulid and a small, coiled serpulid worm tube
bar - (scale equals 10 mm); 3 enlarged view of the concave surface of the counterpart of Trichopeltarion greggi Dell, from the
Miocene of New Zealand. Outer cuticular the broken material adheres to counterpart; where the cuticle is fortuitously away, a mould
of the interior of a balanid barnacle epibiont is revealed (scale bar equals 5 mm); 4 - ventral view of Tumidocarcinus giganleus
from the Miocene of New Glaessner, Zealand, showing straight-sided, exceptionally wide male abdomen ofa feminised individual
bar 10 - (scale equals mm); 5 dorsal view of Torynomma australis Feldmann et al., from the Cretaceous of Antarctica, showing a
severe bopyrid isopod swelling on the right branchial chamber (scale bar equals 10 mm); 6 - scanning electron micrograph of a
portion of the cuticle of an extant Cancer from and loss sp., Mexico, showing exfoliating exocuticle concomitant of an encrusting
bar bryozoan (scale 1 7 - malformed claw of extant equals mm); an Homarus americanus FI Milne Edwards, from Maine, USA (scale
bar equals 10 mm).
Downloaded from Brill.com09/24/2021 08:38:58PM via free access Contributions to 72 - 2003 Zoology, (2-3) 113
Downloaded from Brill.com09/24/2021 08:38:58PM via free access I 14 R.M. Feldmann - Interpreting ecology andphysiology
reversal, and live the final part of their lives as graphically, fossil bopyrids have been identified
females (Bliss, 1982). In this instance, secondary from as far north as Greenland to as far south as
sexual characteristics should be expressed in a 50/ Antarctica. A review of parasitism in extant crus-
50 ratio; however, the females should typically be taceans (Overstreet, 1983) noted that bopyrids typi-
larger than the males. Regardless of the ratio, it is cally were found in macrurans and anomurans. The
clear that gender differences will result in occur- only macrurans discussed were shrimp, a groupwith
char- fossil record. rences of both primary and secondary sexual a poor
than acteristics at a frequency less 100% for each Otherparasites, rhizocephalan barnacles, produce
gender. a condition called parasitic castration. The process
Finally, certain morphological attributes observ- involves introduction of the barnacle into the in- able fossils testinal and destruction of the in occur at very low, and unpredict- tract androgenic gland
able, frequencies. These characters reflect interaction in males, and the barnacle ultimately manifests it-
of the individual crab with its environment. Such self by suppressing male hormones and feminiz-
conditions as pathology and parasitism, infestation ing the males. Females that are infested take on a
by epibionts, predation, autotomy, and regenera- mature appearance at a prematurely early stage
tion of normal or abnormal appendages lie within (Overstreet, 1983). This condition has most fre-
this these conditions have been observed in it is category. In many cases, quently portunids, although
in little effect on the hard parts of the organism and, known other brachyurans as well. The only record
therefore, are not recognizable in the fossil record. of parasitic castration in the fossil record is that of
xanthid from In other cases, such as autotomy or regeneration, a Miocene New Zealand (Feldmann,
of this the evidence may be quite circumstantial. If an 1998). Recognition condition can be made
appendage is not preserved in the fossil record, it only by determining that the abdomen of infected
be attributed loss after death of the males is broadened simulate the form may to organ- unusually to
ism rather than to autotomy. However, some con- of the abdomen of females (Fig. 1.4); thus, it is
ditions do leave evidences that can be recognized necessary to observe the ventral morphology of a
of and interpreted, as discussed below. large number crabs to recognize the condition.
Most pathological and parasitic conditions in The exoskeleton of decapods provides a firm
have little effect the exoskeleton. substratum that base for attachment decapods on Two may serve as a
that of have been recognized are parasitization within of a host epibionts on extant taxa (Fig. 1.1),
the branchial chamber by bopyrid isopods and within including hydroid and anthozoan cnidarians, bryo-
the reproductive system by rhizocephalan barnacles. zoans, bivalves, barnacles, and annelid worms.
The former Almost condition has been summarized by For- any organism requiring a firm base of at-
ster (1969), who reported numerous swellings tachment might be anticipated. The incidence of attributed to in the Galatheidae and Proso- of the these isopods fouling decapod carapace by organ-
Late in Cretaceous isms is the ponidae of Jurassic age and \ quite variable, based upon age of host,
Raninidae. Straelen duration of the inter-molt Prior to his work, van (1928) period, location on the and Housa (1963) hadalso summarized occurrences organism, ecological setting, and life habit of the of bopyrids. Subsequent to Forster’s work, Muller host (Ross, 1983). Many of these epizoans have no
(1984) documented bopyrid swellings in Miocene hard parts and have only a slight chance of being
in decapods (Galatheidae, Porcellanidac Bishop (1986) preserved the fossil record. Others with hard parts
in Cretaceous Homolidae, Col- be lost after death of the host the recognized bopyrids may as waxy
lins & Rasmussen noted their in (1992) presence a epicuticle separates from the remainder of the cu-
LateCretaceous raninid, and Feldmann et al. (1993) ticle (Fig. 1.6) (Waugh & Feldmann, work under
recorded them from a Cretaceous species of the way). Still others are notrecognized because, when
this list thin Torynommidae (Fig. 1.5). Although may opening concretions, a layer of exocuticle may
not be it is exhaustive, interesting to note that all remain attached to the counterpart, obscuring the
the fossil of occurrences bopyrids are in galatheids, epibionts (Fig. 1.3) (Waugh & Feldmann, work
porcellanids, or in the so-called primitive crabs. Geo- under way). The result is that the known occur-
Downloaded from Brill.com09/24/2021 08:38:58PM via free access Contributions - to Zoology, 72 (2-3) 2003 115
rence of on fossil is considerab other epibionts decapods material as camouflage but, again, leave no ly lower than would be based death of the anticipated upon trace upon organism.
of rates fouling on living decapods. Despite these Epibionts have not been used often by paleon- conditions, a of attached have variety epibionts tologists to infer aspects of the depositional envi- been recorded (Fig. 1.2), including bryozoans and ronment, and their occurrence is all too frequently
nested into brachiopods (S.L. Jakobsen, pers. comm.), oysters systematic papers; thus, the informa-
& (Bishop, 1981, 1983; Tshudy Feldmann, 1988), tion is difficult to extract. Because some of the barnacles Feldmann & (Glaessner, 1969; Fordyce, epibionts may be ecologically sensitive, it is quite and 1996), worms & Feldmann, that we can serpulid (Tshudy possible learn more about the setting
1988; Feldmann & The most in Fordyce, 1996). prom- which the decapods lived by studying their inti- for ising possibility discovering more, and differ- mate associates. ent, is under now. Sten L. Jakobsen Predation of epibionts study is, course, a daily occurrence for
is in (Geological Museum, Copenhagen), using clever, decapods modern settings, because they form a novel to diverse food preparation techniques expose a resource for many organisms. Flowever, in of from assemblage organisms the Middle Danian the fossil record, evidence of predation is limited,
Fakse Beds in Denmark. This into may develop partly because the effects are unrecognized and the most important single locality for studying epi- partly because the effects include total destruction bionts, both in terms of and of the remains. prevalence diversity Bishop (1975) described a partial of occurrence. crab specimen within a phosphatic nodule and
Hermit crabs it present a quite different combina- interpreted to be a regurgitate. A similar inter- tion of interaction host-epibiont because a typical pretation was given for a specimen collected from hermit crab the shell the occupies empty of a gastro- Lower Cretaceous of Mexico (Feldmann et al., pod and a of Other variety organisms, including hydro- 1998). occurrences have been noted but prob- zoan cnidarians and known invest bryozoans are to ably represent hydraulic accumulations. Tshudy et the shell (Taylor, 1994). Specimens from Argentina, al. (1989) described the feeding habits ofnautiloid
under are because the her- on exuvia of currently study, typical cephalopods lobsters, noted an in- mit crab initially occupies a small stinctive of the remains from the relatively gas- pattern eating pos- tropod shell and, instead ofreplacing the shell with terior of the abdomen towards the anterior, and a larger one periodically, the hermit crab relies on postulated that this selective ingestion of the abdo- the to continue and men the incrusting bryozoans growing might explain larger percentage of cara- developing a larger, coiled, protective sheath mim- paces than pleons found in the marine fossil record. icking the form of the original shell. Until very Interestingly, examination of a large collection of recently, shells to be freshwater gastropod have been taken crayfish from the Pliocene of the western 'he domiciles of hermit United primary choice for crabs, States showed no difference in the number bat Fraaije described in situ of (2003) an occurrence carapaces and pleons. Nothing in that environment °1 a hermit crab within an Early Cretaceous am- was utilizing the crayfish skeletons in the fashion monite. of the cephalopods. One type of facultative that have been documented epibiotic relationship Crayfish as prey species does not seem to manifest itself in the fossil record in one instance where it was concluded (Feldmann ls drat of de- carrying, or snagging, epibionts as a & May, 1991) that the systematic removal of the fensive or The dorsal of the camouflaging technique. sponge part carapace of Pleistocene crayfish crabs, Dromiidae, a of a an ane- was the result carry cap sponge, of predation either by a small mam- mone, or a of shell the that is mal piece over carapace or by man. Bishop (1972) noted the only oc- Sapped the fifth The does not known by pereiopods. cap currence to me of a crab that was attacked adhere to the carapace and, when released by the by a toothed animal, presumably a fish, and es- crab, leaves no trace. the Similarly, spider crabs, caped. Large puncture marks document the unsuc- Majoidea, have setal hairs like the cessful typically shaped interaction - unsuccessful at least in terms rooks on Velcro©. The hairs trap vegetation and of the predator.
Downloaded from Brill.com09/24/2021 08:38:58PM via free access 116 RM. Feldmann - Interpreting ecology andphysiology
Autotomy, casting off an appendage in the face Beschin C, De Angeli A, Checchi A. 2001. Crostacei decapodi
and associati a coralli della “Formazione di Cas-telgomberto” ofpredation regeneration of the lost limb, oc-
(Oligocene) (Vicenza-Italia Settentrionale). Studi e Ricerche, the curs frequently in modern world but is difficult Assoc. Amici Mus. civ. 'G. Zannato’ 11: 13-30. in to document the fossil record. Occasionally, a GA. 1972. Crab bitten from the Bishop by a fish upper fossil will be illustrated that to specimen appears Cretaceous Pierre Shale of South Dakota. Bull. Geol. Soc.
have an unusually small first pereiopod; however, Amer. 83: 3823-3826. this is only circumstantial evidence of autotomy. Bishop GA. 1975. Traces of predation. In: Frey RW. (ed,).
stated The study of trace fossils: 261-281. New York: Springer. As earlier, absence of the appendage on a GA. 1981. The lobster fossil could result Bishop Linuparus preserved as an arise as a of many factors. The attachment scar on the oyster Exogyra_costata, Ripley one unequivocal example ofregeneration would be Formation (Late Cretaceous), Union County, Mississippi. the of a deformed as a result of growth appendage Mississippi Geol. 2: 2-5. the The damage during growth process. presence Bishop GA. 1983. Oyster predation by decapod crustaceans of not only deformed, regenerated claws, but also in the Late Cretaceous of the Mississippi Embayment. deformed Georgia Jour. Sci. 41: 24. carapaces, has been well documented in Bishop GA. 1986. Occurrence, preservation, and biogeography living Homarus americanus H. Milne Edwards (Fig. of the Cretaceous crabs of North America. In: Gore RH, I of 1.7). am not aware of any demonstration the Heck KL. (eds). Crustacean Biogeography: 111-142. Rotter- in the fossil record, I have phenomenon although dam; A.A. Balkema. always been curious about the bizarre claw depicted Bliss DE. 1982. Shrimps, lobsters and crabs. Piscataway, on Schlueteria tetracheles Fritsch & Kafka (1887, NJ: New Century Publishers Inc.
Collins Rasmussen HW. 1992. Cretaceous-Lower fig. 53). JSH, Upper Tertiary decapod crustaceans from west Greenland. Gronl.
geol. Unders. Bull. 162: 1-46.
Collins JSII, Wicrzbowski A. 1985. Crabs from the Oxfordian Summary sponge megafacies of Poland. Acta geol. pol. 35: 73-88.
Feldmann RM. 1998 Parasitic castration of the crab, Tumi-
docarcinus Ecological and physiological characteristics of fossil giganteus Glaessner, from the Miocene of New
Zealand: coevolution within the Crustacea. Jour. Paleo. decapod Crustacea can be inferred by using a va- 72: 493-498. riety of functional morphological approaches. Ad- Feltlmanii RM, Bearlin RK. 1988. korura Linuparus n. sp. ditionally, considering the frequency ofoccurrence (Decapoda: Palinura) from the Bortonian (Eocene) of New of in of fossil crabs a morphotype a population may Zealand. Jour. Paleo. 62: 245-250. reveal features unique to the individual as an indi- Fcldmann RM, Fordyce RE. 1996. A new cancrid crab cation of its interaction with the environment. Al- from new Zealand. N.Z. Jour Geol. Geophys. 39: 509-
513. though much is known about the behavioral patterns Feldmann RM, May W. 1991. Remarkable crayfish remains of living crabs and, by analogy, fossil forms, many (Decapoda: Cambaridae) from Oklahoma - Evidence of significant observations are within de- presented predation. Jour. Paleo.. 65: 884-886. tailed works and difficult to locate. systematic are Feldmann RM, Tshudy DM & Thomson MRA. 1993. Late
This summary is, in part, a notice of the types of Cretaceous and Paleocene decapod crustaceans from James
Ross Basin, Antarctic Peninsula. Paleont. interpretations that can be made and a plea to call Soc. Mem. 28: 1-41. specific attention to low-frequency, unpredictable Fcldmann RM, Vega FJ, Applegate SP, Bishop GA. 1998. morphological characters. Early Cretaceous arthropods from the Tlaytia Formation
at Tepexi de Rodriguez, Puebla, Mexico. Jour. Paleo.
72: 79-90.
References Forster R. 1969. Epokie, Entokie, Parasitismus und Regenera-
tion bei fossilen Dekapoden. Mitt. Bayer. Staatssamml.
Paldont. hist. Geol. 9: 45-59. Aristotle. 1862. Aristotle’s History ofAnimals: in Ten Books, Fraaije RHB. 2003. The oldest in situ hermit crab from the translated by Richard Cresswell. London: Bohn. Lower Cretaceous of Speeton, UK. Palaeontology 46: 53- Beschin C, Checchi A, Ungaro S. 1996. Crostacei brachiuri 57. dell’Oligocene di Castelgomberto (Lessini Orientali). Studi Fritsch A, Kafka J. 1887. Die Crustaceen der Bohmischen e Ricerche, Assoc. Amici Mus. civ. 'G. Zannato’ 6: 11-
20. Kreideformation. Praha: Selbstverlag.
Downloaded from Brill.com09/24/2021 08:38:58PM via free access Contributions to Zoology, 72 (2-3) - 2003 117
Glaessuer MF. 1969. Decapoda.In: Moore RC. (ed.). Treatise from similar forms. Jour. Paleo. 75: 330-345.
on Invertebrate Paleontology, Part R, Arthropoda 4(2): Schweitzer CE, Scott-Smith PR, Ng PKL. 2002. New R400-R533. Boulder: Geological Society of America/ occurrences of fossil decapod crustaceans (Thalassinidea,
Lawrence: University of Kansas Press. Brachyura) from late Pleistocene deposits ofGuam, United
Housa V. 1963. Parasites of Tithonian decapod Crustacea States Territory. Bull. Mizunami Fossil Mus. 29: 25-49.
(Stramberk, Moravia,). Shornlk Ustr. Ust. Geol. 38:101- Schweitzer Hopkins CE, Feldmann RM. 1997. Sexual
116. dimorphism in fossil and extant species of Callianopsis
Manton SM. 1977. The Arthropoda. Oxford: Clarendon Press. de Saint Laurent. Jour. Crust. Biot. 17: 236-252.
Muller P. 1984. Crustacea of the Badenian. Geol. Straelen V 1928 Contribution 1’etude Decapod van. a des isopods
Ser. Palaeont. 42: Hung., 1-317. meso- et cenozoiques. Mem. Acad. r. Belg., Cl. Sci. (2)9: Muller P, Collins JSH. 1991. Late Eocene coral-associated !-66.
decapods (Crustacea) from Hungary. Contr. Tert. Quatern. Taylor PD. 1994 Evolutionary palaeoecology of symbioses
Geol. 28: 47-92. between bryozoans and hermit crabs. Hist. Biol. 9: 157-
Overstreet, RM. 1983. Metazoan symbionts of crustaceans. 205.
In: Provenzano Jr AJ. (ed.). Pathobiology 6, The biology Tshudy DM, Feldmann RM. 1988. Macruran decapods, and Crustacea : of 156-250. New York: Academic Press. their epibionts, from the Lopez de Bertodano Formation
Ross DM. 1983. Symbiotic relations. In: Vernberg FJ, Vern- (Upper Cretaceous), Seymour Island, Antarctica. In; Feld-
berg WB. (eds). Behavior and ecology, 7, The biology of mann RM, Woodburne MO. (eds). Geology and Paleonto- Crustacea: 163-212. New York: Academic Press. logy of Seymour Island, Antarctic Peninsula. Mem. Geol. Schafer W. 1972. Ecology and palaeoecology of marine Soc. Amer. 169: 291-301.
environments (edited by G. Y. Craig). Chicago: University Tshudy DM, Feldmann RM, Ward PD. 1989. Cephalo-
of Press. Chicago poda; biasing agents in the preservation of lobsters. Jour. Schweitzer CE, Feldmann RM. 2000. New fossil portunids Paleo. 63: 621-626.
from Washington, USA, and Argentina, and a re-evaluation Zullo VA, Chivers DD. 1969. Pleistocene symbiosis: Pinno- of and within Portunoidea therid generic family relationships the crabs in pelecypods from Cape Blanco, Oregon.
Rafinesque, 1815 (Decapoda: Brachyura). Jour. Paleo. Veliger 12: 72-73, pi, 5.
74: 636-653.
Schweitzer CE, Feldmann RM. 2001. Differentiation of Received: 12 March 2003
the fossil Hexapodidae Miers, 1886 (Decapoda: Brachyura)
Downloaded from Brill.com09/24/2021 08:38:58PM via free access