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Ethology and Distribution of Pylochelidae (Crustacea Decapoda Coenobitoidea)

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Ethology and Distribution of Pylochelidae (Crustacea Decapoda Coenobitoidea) BULLETIN OF MARINE SCIENCE, 41(2): 309-321, 1987 ETHOLOGY AND DISTRIBUTION OF PYLOCHELIDAE (CRUSTACEA DECAPODA COENOBITOIDEA) Jacques Forest ABSTRACT The Pylochelidae differ from the other hermit crabs by the complete segmentation of the abdomen and the presence of paired appendages on each of its segments. They do not usually inhabit gastropod shells, but dwell in decayed pieces of wood, stones, tusk-shells, or living sponges. A recent revision, founded on most of the previously recorded specimens and on a large unidentified collection, increased the number of known species from 16 to 39, and the genera from 5 to 7. Two new subgenera have been established, and the family divided into six subfamilies. This paper deals first with the eco-ethological characteristics of the different taxa. According to their dwelling, genera and subgenera can be classified, as a whole, as xylicolous, petricolous, tusk-dwellers, spongicolous, with a few specifical or individual ex- ceptions. In connection with the habitat, adaptive features have been described: opercular structures, boring "rasp," stridulating apparatus ... The Pylochelidae are present in the Indo- West Pacific (36 species or subspecies in 6 genera), and in the NW Atlantic (4 species in 3 genera). Two genera only, belonging to the sole non monotypic subfamily, provide a biogeo- graphical link between the two areas. In I-W.P., the family is known from the SW Indian Ocean to Japan, Kermadec Islands and New Zealand. Indonesia, with 14 species and 5 genera appears as a center of dispersion and diversification. Japanese endemism is noteworthy: one genus and six of the seven species have not been reported elsewhere. The probable relation between the availability of dwelling material and the geographical distribution is also dis- cussed. The vertical distribution extends from 30 to 1,570 m, but the group is mostly rep- resented between 200 and 500 m, where 28 species are living. The Pylochelidae differ from all other hermit crabs in having a well developed abdomen, all segments of which are articulated and provided with a pair of appendages, similar to the normal abdomen of other reptant decapods. Pylochelids are commonly called "symmetrical" hermit crabs, but this is not entirely correct, because in one genus the abdomen, telson and pleopods are noticeably asym- metrical. The group has been considered as rare, only 16 species being recorded from a small number of rather deep water stations in Indo-West Pacific (I-W.P.) and western Atlantic, and the most being known only from their type localities. The abundance of a new material, originating mainly from ALBATROSSdredgings and from recent French explorations in the I-W.P. has led to a systematic revision (Forest, 1987). As a result, 24 new species-group taxa have been added to the 16 previously established valid species, the five known genera have been redefined, the genus Pylocheles has been divided into three subgenera, and two new genera have been proposed. Until now the Pylochelidae has been considered as a relatively restricted family of infrequently encountered species; aside from three forms captured on several occasions in Japanese waters, the total number of specimens recorded in literature did not exceed 60, caught at about 30 stations. The present revision includes more than 400 specimens, collected at some 200 stations! The importance of pylochelid fauna in tropical and subtropical waters has been, therefore, much underestimated and, most probably, new taxa and localities will be added in the future. This research, however, has not been restricted to the description of new forms. 309 310 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.2, 1987 Investigations on relationships between the various genera have shown that the whole group is made up of several distinct phyletic lines whose respective affinities are not entirely clear, and is likely polyphyletic. Therefore, the family has been divided, provisionally, into six subfamilies. The classification and content of the family Pylochelidae is given in Appendix 1. The Pylochelidae, within the section Paguridea of the infraorder Anomura, are classified in the superfamily Coenobitoidea. A comparative study of their main characters allows us to suggest that they are close to the Diogenidae. They cannot however be regarded as primitive representatives of that family; both Diogenidae and Pylochelidae (if monophyletic) probably have a common ancestor, but they apparently evolved independently along various phyletic lines. The richness of the new material upon which the systematic revision of the family has been based, has also provided considerable information on the ecology or the habitat of many forms, and allows one to interpret the adaptive value of some morphological structures. The first part of the present communication is devoted to this aspect of the pylochelid biology. The second and third parts give an account of the biogeographical and bathymetrical distributions of these most interesting pagurids. Etho-ecology of Pylochelidae. - The abdominal tergites in the Pylochelidae are better calcified and accordingly less vulnerable than in most other marine pagurids. However, these animals seek protection by dwelling in hollow objects. From lit- erature it was known that gastropod shells are infrequently used. Rather, pylo- chelids use wood or stone fragments, tusk-shells or sponges. Examination of numerous specimens still in their dwellings, sometimes observed at the time of their capture, provides the basis for these statements, and allows in many cases the accurate determination of the exclusive or preferential habitat of species be- longing to at least eight of the nine genera or subgenera. These dwellings are discussed below: WOOD DWELLINGS. These come from the shore or rivers and consist of decayed fragments oftrees excavated by wood-boring organisms, or hollow stems such as bamboo. All species of the subgenera Xylocheles and Bathycheles. in the genus Pylocheles. are wood-dwellers, or xylicolous. The size of the piece of wood is usually in relation with that of the inhabitant; small animals are found in stems of about I cm in diameter, middle-sized or large ones in much larger fragments weighing sometimes more than I kg. The cavity is subcylindrical, with a diameter of more than 2 cm for large animals, and always longer than the inhabitant. The walls of the lodging are extremely regular, owing to the activity of the animal, using rasps of tubercles on the carpus of the chelipeds. Presumably, the hermit first selects a piece of wood already hollowed by wood-borers, smoothens the walls with its rasps, and widens the cavity as it grows. The back of the lodge is rounded but remains often open to the exterior through the small primitive gallery. On the schematic illustration of Pylocheles (Xylocheles) macrops in its dwelling one can see the cylindrical main chamber, and the small primitive gallery (Fig. lb). All species of Xylocheles and Bathycheles have rasps and, as a rule, are xylic- olous. However, of 20 specimens of B. incisus collected at one station, about half dwelled in large tusk-shells of Fissidentalium magnificum, Parapylocheles scorpio, a monotypic genus, is also xylicolous and is usually found in bamboos. However, we were surprised to discover in the Philippines FOREST: SYMMETRICAL HERMIT CRABS 311 (; ,\ , ~~ .. I ,I~!!' I I, .! .:r- I' ), '~~' ,~: I ": I k,': \ , ~' J ,; \ , ; ,r,", 'I > \' ',,\, " :', ¥, I 1 ',J ft a }, III b , ~ l Figure I, a. Pylocheles (pylocheles) mortensenii Boas in a pumice stone; b, pylocheles (Xylocheles) macrops Forest in a piece of wood. one specimen settled in a corn cob! Apparently, there is no rasping apparatus in the genus Parapylocheles. STONE DWELLINGS. These are found mainly in broken pieces of soft stones such as sand conglomerates, limestone and volcanic rock, especially pumice. The two species of the nominotypic subgenus Pylocheles, P. (P.) agassizii and P. (P.) mortenseni, live most frequently in that last type of habitat; many specimens of the latter have been found in pumices weighing from a few to 80 g. Here again, the animal is located inside a more or less deeply hollowed subcylindrical cavity 312 BULLETIN OF MARINE SCIENCE, VOL. 41, NO.2, 1987 (Fig. la). Species of the genus Cheiroplatea also live in stone dwellings, and have similar opercular chelipeds. In these petricolous pylochelids, the chelipeds are narrowly coadapted, the hands perpendicular to the axis of the cavity, closing it perfectly. This is not the case in xylicolous forms where the appendages remain more or less parallel to the axis of the cavity and where both hands are not perfectly coadapted. On the other hand, there is no rasping apparatus on the carpus of the chelipeds in Pylocheles (pylocheles) and Cheiroplatea. Strong teeth at the edge ofthe carpus and propodus may be instrumental in fitting out the habitat. Some specimens have been collected in dead bryozoans or in living sponges. Presumably these latter organisms were epibiont on the primitive mineral dwelling of the hermit. The new genus, Cancellocheles, is probably also petricolous. The four known individuals of C. sculptipes have been described without any information on their habitat, but, from a strong convergent similarity with Cancellus (the known species of which live in stones, dead corals or calcareous algae), it can be presumed that they have a similar way oflife. In both genera, chelipeds and second thoracic legs are coadapted in the formation of a perfect operculum. SHELLDwELLINGS.Apart from a small number of Trizocheles, and from Mix- topagurus paradoxus A. Milne Edwards, from the Western Atlantic, pylochelids do not use gastropod shells. There are 12 known specimens of Mixtopagurus, only one of which has been collected with its dwelling, a shell of Xenophora sp. All specimens however have a slightly twisted, asymmetrical abdomen, and it can be presumed that the species is adapted to life in the same sort of wide, low-spiralled shell. The tendency of pylochelids to seek shelter in a more or less straight tubular cavity explains why they often inhabit tusk-shells, since they are elongated, conical and slightly arched.
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