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Brachyura) of Different Habitat Conditions

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Brachyura) of Different Habitat Conditions CRUSTACEAN RESEARCH，NO. 35・56-- 66，2006 Comparison of social behaviors among six grapsoid species (Brachyura) of different habitat conditions Yuuka Nara，Jun Kitaura and Keiji Wada Abstract.- Social behaviors were Klaassen，1975; Henning，1975; Lindberg， observed in the field for six grapsoid 1980; Seiple & Salmon，1982; Abele et al. ， crabs living under different habitat 1986; Beinlich & Polivka，1989) ，compared conditions from exposed to sheltered with ocypodoid crabs.Furthermore ，inter- conditions，i. e. Helice tridens，H. specific comparison of social behaviors was jα :ponicα，H.leαchi，Ch αsmαgnαthus only attempted by Wright (1968) addressing convexus of the family Varunidae，and taxonomic distribution of chela-using display Chiromαntes dehαα ni， and pattern ，and by Brockerhoff & McLay(2005) Perisesαrmα bidens of the family dealing with comparative analysis of mating Sesarmidae. Fighting behaviors in strategies. these six species，cheliped ・displays in Semi-terrestrial crabs that live in f1 at H. tridens，H. jα:p onicαa n d P. bidens， habitats and show diurnal activity are con- and copulatory behaviors in H. tridens， sidered to have advantage for the develop- C. dehαα ni and P. bidens were ment of visual signaling (Salmon & Atsaides， described. Elements of fighting behav- 1968). Grapsoid species living in open and ior varied among species irrespective f1 at habitats ，therefore ，are expected to have of taxonomic group. The three species developed social behavior for visual commu- that showed cheliped-displays occurred nication ，compared with grapsoid species liv- in more exposed habitats compared to ing in structured habitats. the other species，suggesting the impor- 1n this study，social behaviors such as tance of habitat conditions in the evo- aggressive behavior，ch eliped-displays and lution of such displays. Copulatory copulatory behavior are described for 6 behaviors were observed in three grapsoid species that occur in open to verti- species，but precopulatory courtship cal habitats: Helicetridens (de Haan，1835) was absent. and H. japonica K. Sakai & Yatsuzuka，1980 inhabiting open mud flat (Omori et al. ， 1997) H elice leachi Rathbun 1931 Introduction ， ， ， Chasmagnathus convexus de Haan，1833 and Brachyuran species of the superfamily Chiromantes dehaani (H.Milne Edward， Ocypodoidea and Grapsoidea mostly live in 1853) inhabiting structured habitats such as intertidal to supratidal areas，but they gener- salt marsh or mangrove (Nakasone，1977; ally differ in habitat conditions. Many ocy- Fukui & Wada，1983 )， and Perisesarma podoids occur in exposed habitats ，such as bidens (de Haan，1835) inhabiting transition- sand beaches and mud/sand f1 ats ，whereas al habitats from structured to open habitats many grapsoids occur in more structured (Nakasone，1977; Fukui & Wada，1983) . vertical habitats，such as rocky shores， 1nterspecific comparisons of the behavior beneath stones or in vegetation. Given such patterns are made to explore the relation- habitat conditions，social behaviors of grap- ships with taxonomic group and habitat con- soid crabs have been described for few ditions. species (Kramer，1962; Warner，1970; SOC凶L BEHAVIORS OF GRAPSOID CRABS 57 Table 1. List of species studied，observation sites and periods. Species Site Period Family Varunidae H elice japonica Doki Ri ver Estuary，Kagawa Pref. 24，25/ August，8- 10 /September / (34 0 18' N ，133 0 48' E) 2003; 16-18/ May，15 ，16/ June /2004; 19-21 /April ，2005 (13 days) Helice leachi Funaura，Iriomote Is. ，Okinawa Pref. 16-21 /April /2004 (6 days) (24 0 23' N ，123 0 49' E) H elice tridens Kushida Ri ver Estuary，Mie Pref. 16 ，28-29/ April ，1 5-17 /May ，12-14 ， (34 0 36' N ，136 0 35' E) 27-29/ June /2003 (13 days) Chasmagnathus convexus Kogi Ri ver Estuary，Osaka Pre f. 20-23/ June /2005 (3 days) (34 0 26' N ，135 0 21' E) Family Sesarmidae Chiromantes dehaani Kogi Ri ver Estuarγ ，Osaka Pref. 19-21/ July，14 ，1 8，19 ，August /2004; (34 0 26' N ，135 0 21' E) 19 ，22，23 /July ，21 ，22 /July /2005 (11 days) Perisesarma bidens Kogaura，Shirahama ，Wakayama Pref. 3，4/ July，3 ，4 / August ，18 ，19/ (33 0 40' N ，135 0 22' E) September /2004; 9-11 /June ，4， 5 / August /2005 (11 days) Table 2. Elements of fighting behaviors and the number of occasions observed for each element in 6 grapsoid species. Varunidae Sesarmidae Helice tridens Helice japonica Helice leachi Chasmagnathus Chiromantes Perisesarma bidens convexus deh ωnI 1) Grasping 12 33 3 7 46 26 2) Arm-flexed pushing 27 29 5 3) Arm-extended pushing 4 4) Leg-touching 2 1 6 3 52 5) Dash 3 20 14 2 11 89 6) Striking 4 12 37 63 7) Leg-tapping 72 Total 48 58 18 27 198 235 Materials and Methods of the behaviors were as follows. Fighting Four species of the family Varunidae， behavior is an agonistic interaction between Helice tridens ，H. jαponica，H. leachi， two individuals with physical contact，and Chasmagnathus convexus，and 2species of with one individual aggressively approach- the family Sesarmidae， Chiromantes ing the other. Waving displays are rhythmi- dehaani，Perisesarma bidens were observed cal motions of chelipeds. Cheliped-display is in the field for 3-13 days at each one loca- anon-waving display，with acrab moving its tion between 2003-2005 (Table 1). chelipeds repetitively in same locus. Observations were made only during day- Copulatory behavior is copulation and asso- time low tides，aided by adigital video cam- ciated behaviors occurring preceding or fol- era (SONY DCR・TRV30). 羽Tefocused on lowing copulation. When either of these three types of social behavior; fighting，dis- social behaviors was observed，sequential plays with chelipeds including waving dis- events were noted. The crabs that per- plays，and copulatory behavior. Definitions formed the social behaviors were sexed in 58 y. NARAET A L. case when the sex could be determined Three elements of fighting behavior could from the extemal feature. be distinguished as grasping，leg-touching ， and dash. All three elements were per- formed in the same way as in H. tridens. Results Successive occurrence of the elements was Fighting behαmor observed as dash to grasping. Among the three elements，dash occurred most com- Helice tridens :A total of 48 encounters monly. were observed during 13 days (Table 2). Five elements of fighting behavior could be Chαsmagnathus convexus :A total of 27 distinguished as follows. 1) Grasping (Fig encounters were observed during 3days 1-A): one crab extends its chelipeds to the ぐrable 2). Four elements of fighting behav- opponent and grasps it ，or both crabs grasp ior could be distinguished as grasping，leg- each other. They grasp any part of the body， touching，dash ，and striking. Grasping 1) ， induding the carapace ，walking legs ，and leg-touching 4) and dash 5) were made in chelae. 2) Arm-f1 exed pushing (Fig 1-B): the same way as in H. tridens. 6) Striking one or bo出 crabs push against the opponent was made in the same way as in H. japonica. using the outer faces of chelae. 3) Arm- Successive occurrence of the elements was extended pushing (Fig 1-C): two crabs observed as 4) to 1) or 6) ，6) to 1) ，and 5) to stand at ac1 0se distance facing each other， 6). Among the four elements，striking with their both chelipeds extending to the occurred most commonly，compared to the side ，so that their chelae sometimes touch. other three elements. 4) Leg-touching (Fig 1-E): one of walking legs is rapidly extended to make contact Chiromantes dehaani :A total of 198 with the opponen t. 5) Dash: one crab makes encounters were observed during 11 days adash for the opponent without any physical ぐfable 2). Six elements of fighting behavior contact. Successive occurrence of the ele- could be distinguished as grasping，arm- ments was observed as 4) to 1) or 2) ，and 2) flexed pushing，leg-touching ，dash ，striking ， to 3). Among the five elements，grasping and leg-tapping. Grasping 1) ，arm- f1 exed and arm-f1 exed pushing occurred more com- pushing 2) ，leg-touching 4) and dash 5) were monly，compared to other three elements. made in the same way as in H. tridens. Striking 6) was made in the same way as in Helice japonica :A total of 58 encounters H. japonica. 7) Leg-tapping (Fig 1-F): one were observed during 13 days (Table 2). crab touches the opponent by some walking Four elements of fighting behavior could be legs and vibrates them rapidly on the oppo- distinguished as grasping，leg-touching ， nen t. Successive occu汀 ence of the elements dash，and striking. Grasping 1) ，leg-touching was observed as 1) to 7) or 6) ，2) to 1) or 6) ， 4) and dash 5) were made in the same way 4) to 1) ，2) ，6) or 7) ，5) to 1) ，6) or 7) ，6)ω as in H. tridens. 6) Striking (Fig 1-D): one 1) ，and 7) t01). Among the six elements，leg- crab raises both chelipeds，and down toward tapping occurred most commonly，compared the opponent like scratching. Successive to other five elements. occurrence of the elements was observed as 4) to 1) or 6) ，5) to 1) ，and 6) to 1). Among Perisesarma bidens A total of 235 the four elements，grasping and dash encounters were observed during 11 days occurred more commonly，compared to σ'able 2). Five elements of fighting behavior other two elements. could be distinguished as grasping，arm- f1 exed pushing，leg-touching ，dash ，and Helice le αchi :A total of 18 encounters striking. Grasping 1) ，arm-flexed pushing 2) ， were observed during 6days (Table 2). leg-touching 4) and dash 5) were made in SOCIAL BEHAVIORS OF GRAPSOID CRABS 59 B C E F Fig. 1. Fighting elements observed in six grapsoid species.
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