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The Behavior of Some Deep-Water Hermit Crabs (Decapoda: Paguridea) from the Straits of Florida!

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The Behavior of Some Deep-Water Hermit Crabs (Decapoda: Paguridea) from the Straits of Florida! THE BEHAVIOR OF SOME DEEP-WATER HERMIT CRABS (DECAPODA: PAGURIDEA) FROM THE STRAITS OF FLORIDA! BRIAN A. HAZLETT Institute of Marine Science, University of Miami and Biological Laboratories, Harvard University ABSTRACT The behavior of specimens of eleven species of hermit crabs, collected from depths of 110 to 713 meters, was observed in the laboratory (Dio- genidae: Clibanarius anomalus, Paguristes spinipes, Paguristes moorei, Paguristes "A", and Dardanus insignis; Paguridae: Benthopagurus cokeri [new combination], Pagurus rotundimanus, Pagurus politus, Pylopagurus discoidalis, Parapagurus piClUs, and Spiropagurus iris). The aggressive displays executed as individuals approached one another are described for each species. These movements of the chelipeds and ambulatory legs are similar to those of congeneric species from shallow water. Pagurid hermits exhibited extreme avoidance of conspecific individuals during diurnal observation periods. Specimens of Pagurus politus showed a peculiar "appeasement" behavior whereby individuals could come closer than usual to one another. The behavior patterns associated with shell fighting are described for four species (Clibanarius anomalus, Paguristes moorei, Dar- danus insignis and Pylopagurus discoidalis). Several adaptations of the behavior of Pylopagurus discoidal is affected by inhabiting scaphapod shells are described. INTRODUCTION A number of species of hermit crabs, especially in the family Paguridae, are found well below the littoral zone. The social behavior of the shallow- water hermit crabs of Cura<;ao, N.A. (Hazlett, In Press) and of the Miami area (Hazlett, MS) has been examined from a comparative viewpoint. Although each species observed has a characteristic repertoire of be- haviors, the patterns of all shallow-water species examined are markedly similar. The intraspecific interactions of some deep-water species were observed both for systematic purposes and to compare the behavior of these forms with that of shallow-water species. The 11 species considered in this paper were collected by bottom trawls and dredges between 60 and 390 fathoms (110-713 m) during cruises by University of Miami vessels, the RjV JOHN ELLIOTT PILLSBURY and the RjV GERDA. The range of these cruises extended from Key West to 'Contribution No. 669 from the Institute of Marine Science, University of Miami. This work was supported by fellowship F 2 MH-14, 274 from the National Institutes of Health and a Grant-in-Aid of Research from the Society of the Sigma Xi. Biological Ship time was supplied by Grant GB-1204 from the National Science Foundation. The material reported here was collected with the coopera- tion of the National Geographic Society-University of Miami Deep-Sea Biology Program. 1966] Hazlett: Deep-water Hermit Crab Behavior 77 Bermuda but the majority of the collecting was done in the Straits of Florida proper. When brought on deck, the specimens were quickly placed in individual containers of cooled Florida Current water in a refrigerator. Observations were carried out by placing the specimens of one species together in a cooled aquarium with sand on the bottom and watching them for periods of 30 to 45 minutes, except when prolonged interactions ex- tended beyond this time. All observations of specimens collected on the RjV GERDAwere carried out at the Institute of Marine Science in Miami. Animals were maintained at 10°, 15°, or 20° C depending on the depth and area they came from and were kept on a schedule of 12 hours illumi- nation and 12 hours dark. Except when placed together for observation, the specimens were kept in individual containers of Gulf Stream water. The change in water temperature during an observation period was 1°_2° C. Although the level of general activity sometimes appeared to increase slightly as the water temperature rose, the character of the move- ments executed during social interactions did not change. Observations at night were carried out with a "red light" (an ordinary low wattage red light bulb, not a monochromatic source) at some times and a low level white light at other times. The conditions of carrying out the observations and the limited number of specimens available make these descriptions preliminary in nature. With several exceptions, however, the behavior patterns observed were not at all unusual when compared with the laboratory and field behavior of related shallow-water hermit crabs (Hazlett, In Press and MS). Indeed, the similarity between the behavior of these deep-water species and that of congeneric shallow-water forms was surprising in view of the difference in light conditions in the field for the two groups. The author wishes to thank Mrs. Edith Marks and Talbot E. Murray, Jr. for their help in maintaining specimens in the laboratory and Dr. A. J. Provenzano, Jr. of the Institute of Marine Science, for his aid and advice. Species identifications were confirmed by Dr. Provenzano. Most of the specimens observed in this study have been deposited in the Institute of Marine Science Museum. The patterns described in this paper belong to two categories of aggres- sive behavior-general aggressive behavior and shell fighting behavior. The latter is an exchange of signals between two crabs which effects an exchange of shells without physical damage to the interacting crabs. In the twelve shallow-water species observed in Cura<;ao (Hazlett, In Press), the general outline of events was the same in all species although there were distinct familial, generic and specific characteristics in the execution of most acts. The attacking crab approaches another crab and, after an exchange of visual displays (see below), gets into an opposed position with respect to the other crab. In this position, the apertures of the two shells are facing, 78 Bulletin of Marine Science [16(1) FIGURE 1. Schematic diagrams of the aggressive displays of hermit crabs. The dotted outlines indicate the resting position of the limb. Appendages other than those involved in the display illustrated have been omitted. A. Side view of a Pagurus species, showing the major cheliped presentation (manus moved through almost 100° from its resting position to 10° beyond the vertical) and the major cheliped extension (moving the manus another 80° upward to a horizontal position). B. Front view, showing a single ambulatory raise (45° of movement to a position 20° below the horizontal). 1966] Hazlett: Deep-water Hermit Crab Behavior 79 forming an angle of about 30° with one another. The attacking crab has its ambulatory legs around the aperture of the defender's shell and is usually above the defender. The latter is withdrawn in its shell. From this position, the attacker carries out positioning movements (basically a rock- ing from side to side of the attacker's body-shell) and then rapping move- ments (a rapid bringing together of the two shells by the attacker). After the execution of a number of these movements, the defending crab may come competely out of its shell and move onto the back of the shell of the attacker. The two crabs then exchange shells. In addition to this form of interaction, individuals of all species also interact in a more general way. Hermit crabs are not territorial but like most animals have an individual distance. When another crab moves too close to an individual, it usually responds to the presence and/or move- ments of the other either by retreat or the execution of a stereotyped visual display. In most species of hermits, physical damage between con- specific individuals almost never occurs but rather, exchanges of these display movements result in one crab moving away or ducking back into its shell. The three most common displays have been termed the ambula- tory raise, the cheliped presentation and the cheliped extension. The speed and extent of movement of the respective limbs in these displays vary from species to species. When one crab executes a display, the other usually responds by either retreating or executing a display. Thus long series of exchanges of displays may occur before one or both crabs move away. Smaller crabs usually retreat from just the approach of a larger crab or from the execution of a cheliped presentation movement. The ambulatory raise and cheliped extension displays are stronger aggressive signals and in extended en- counters, the simultaneous movement of several ambulatory legs and both chelipeds is common. It is difficult to clearly describe the limb movements involved in these displays in a few words. The terminology used in these descriptions is illustrated in Figure 1. The cheliped presentation display (Fig. lA) brings the limb(s) forward (with little lateral movement) from the normal resting position-some degrees posterior to the D-V axis-until the manus is approximately vertical (parallel to the D-V axis). In the extension display, the limb is moved forward farther. In both movements the proximal segments of the chelipeds are moved almost to the horizontal (parallel to the A-P [anterior- posterior] axis). In each species, these movements are described in terms of the movement of the manus (number of degrees in the angle formed by lines connecting the tip of the dactyl of the manus and the carpus-manus articulation in the two positions) and the final position of the manus (degrees anterior or posterior to the vertical or degrees below the horizon- 80 Bulletin of Marine Science [16(1) tal). The ambulatory raise movement (Fig. IB) brings the limb(s) from the normal walking position, laterally upward, to a position in which some or all segments of the limb are almost horizontal (at right angles to the D-V [dorso-ventral] axis of the animal). In some species, the segments are approximately straight in the raised position. In others the dactyl may be declined below the line of orientation of the other segments. In the simplest case, only the degrees of movement of the dactyl (the number of degrees in the angle formed by lines connecting the dactyl and coxa of the limb in the two positions) and the final position (in degrees below the hori- zontal) are given.
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