Ecology and Systematics of Mangrove Crabs of the Genus Perisesarma (Crustacea: Brachyura: Sesarmidae) from East Africa
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Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin- nean Society of London, 2004? 2004 141? 435445 Original Article ECOLOGY AND SYSTEMATICS OF EAST AFRICAN PERISESARMA D. P. GILLIKIN and C. D. SCHUBART Zoological Journal of the Linnean Society, 2004, 141, 435–445. With 4 figures Ecology and systematics of mangrove crabs of the genus Perisesarma (Crustacea: Brachyura: Sesarmidae) from East Africa DAVID PAUL GILLIKIN1* and CHRISTOPH D. SCHUBART2 1Department of Analytical and Environmental Chemistry, Vrije Universiteit Brussel, Pleinlaan 2, B-1050 Brussels, Belgium 2Biologie 1, Institut für Zoologie, Universität Regensburg, 93040 Regensburg, Germany Received August 2003; accepted for publication February 2004 East Africa has a reduced mangrove crab species richness when compared to Asian mangroves. To date, only one spe- cies of Perisesarma de Man, 1895 has been reported in East Africa, despite more than 30 years of mangrove research in this region. Based on morphology, colour, mtDNA and behaviour, we describe a new species of Perisesarma from Kenya, P. samawati sp. nov. Surprisingly, when comparing molecular data from other species within this genus, P. samawati sp. nov. and the sympatric P. guttatum (A. Milne Edwards, 1869) are not sister species. Some aspects of the ecology of P. guttatum and P. samawati sp. nov. are compared and the differences discussed. Additionally, we compare P. samawati sp. nov. with the ecological literature of a possible sister species P. eumolpe de Man, 1895 from Malaysian mangroves. Our findings suggest that the new species is an ecologically important species in East African mangroves. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 435–445. ADDITIONAL KEYWORDS: 16S rRNA – Grapsoidea – Indian Ocean – Kenya – mangroves – morphology – mtDNA – new species – phylogeny – taxonomy. INTRODUCTION Grave & Tack, 2001). Perisesarma spp. are reported to have the highest biomass of mangrove crabs in certain It is continuously being reported that grapsoid crabs, forests (Ashton, Macintosh & Hogarth, 2003). especially those from the species-rich family Sesarmi- East Africa, in general, has a limited number of dae, play an important ecological role in mangrove brachyuran crabs associated with mangroves (about ecosystems (reviewed in Lee, 1998). For example, 35 species; cf. Hartnoll, 1975) compared with south- while many mangrove sesarmids do not assimilate east Asia (>100 in Malaysia alone; Tan & Ng, 1994). In much carbon from mangrove leaves and rely more on fact, there has not been a new grapsoid species mangrove sediments (Bouillon et al., 2002; Skov & described from East Africa since 1965, when Crosnier Hartnoll, 2002), it appears that Perisesarma spp. do distinguished Chiromantes ortmanni from the well supplement their diet with mangrove leaves (Leh & known C. eulimene de Man (1895), despite the large Sasekumar, 1985; Slim et al., 1997; Dahdouh-Guebas amount of ecological research concentrated in this et al., 1999; Ashton, 2002). The removal and process- area (e.g. Bright & Hogue, 1972; Hartnoll, 1975; Mich- ing of mangrove leaves by crabs helps to trap the eli, Gherardi & Vannini, 1991; Cannicci et al., 1996; energy stored in these leaves within the mangal before Ruwa, 1997; Slim et al., 1997; Dahdouh-Guebas et al., the tide can carry them away (see Lee, 1998; Skov & 1999; Fratini, Cannicci & Vannini, 2000; Flores, Paula Hartnoll, 2002). Furthermore, their faecal material & Dray, 2003). Currently, there is only one species of potentially contributes to secondary production via a Perisesarma, P. guttatum, attributed to East Africa coprophagous food chain (cf. Lee, 1997; Gillikin, De (Crosnier, 1965; Hartnoll, 1975; Vannini & Valmori, 1981), whereas there have been 19 species reported *Corresponding author. E-mail: [email protected] within the Indo-Pacific region as a whole (Davie, © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 435–445 435 436 D. P. GILLIKIN and C. D. SCHUBART 2003). Given the long planktonic phase of East African species was determined by visual inspection in 10 m species of Perisesarma (22–25 days, see Pereyra Lago, diameter plots along a transect perpendicular to the 1993), and the presence of strong oceanic currents in coastline, covering the full width of the forest. We the Indian Ocean (cf. Hartnoll, 1975), it is surprising investigated 15 plots along a 364 m long transect in that there are not more species of this genus present Gazi Bay and 23 plots along a 1298 m long transect in in East Africa. Perisesarma guttatum, endemic to East Mida Creek. Each plot was first inspected using Africa, is an important element of the Kenyan man- 10 ¥ 42 binoculars from a distance of approximately grove fauna, removing almost anything that falls to 7 m for 15 min and the crab species recognized the forest floor (Gillikin, 2000). These crabs are not at recorded. Afterwards, naked eye observations were all shy, have wide distribution across the intertidal made with two observers sitting on opposite sides of mangal [generally, from the Ceriops tagal (Perr.) C.B. the plot for at least 30 min (cf. Hartnoll et al., 2002; Rob. zone down to the seaward front of the mangal] Skov et al., 2002). On approaching the plot, a distur- and are usually the first of the mangrove fauna to bance was created, causing all crab species to imme- reach any potential food source (cf. Ruwa, 1997; Fra- diately take refuge. The time at which the first tini et al., 2000; Gillikin, 2000). individual of a species would re-emerge was recorded. During field surveys in 1998, it was noticed that All observations took place at low tide during the day; among the crabs that were referred to as P. guttatum, no distinction of lunar cycle was made. To further two forms with distinct colour patterns could be dis- assess the behavioural differences between the species tinguished. Further investigations also revealed of Perisesarma, one individual of each type was placed behavioural and ecological differences, the more in a 50 cm diameter container and stimulated by mov- colourful specimens not venturing out to forage as rap- ing a hand over the container. This experiment was idly as the darker form, and seemingly returning to a replicated with males and females, each three times. specific shelter more regularly when retreating. This At two plots, where numbers of both types appeared led us to believe that the crab was not in fact high, feeding experiments were conducted on three P. guttatum, but a different species of Perisesarma, separate days. First, a fresh green leaf fragment previously unrecorded in East Africa. After detailed (± 2 cm2) was placed on the forest floor and the first morphological examinations and careful comparisons species to reach it was recorded (for discussions of with the literature describing other species of Peris- crabs feeding on green leaves, see Ashton 2002; Gil- esarma, it became apparent that this species was likin, De Wachter & Tack, 2004). Secondly, ten or more indeed different from P. guttatum, as well as from all leaves were fragmented and spread on the forest floor other described members of the genus. In this study, to create an abundance of leaf fragments. The behav- we describe the new species of Perisesarma from Ken- iour following both of these experimental settings was yan mangroves, where it occurs sympatrically with recorded. P. guttatum. Further evidence is provided by mtDNA Type specimens have been deposited at the Senck- sequences to show that, compared with other mem- enberg Museum (SMF), Frankfurt a. M., Germany; bers of the genus, the new species is not the sister spe- the Royal Belgian Institute of Natural Sciences cies of P. guttatum. (KBIN), Brussels, Belgium; the Nationaal Natuurhis- torisch Museum (RMNH), Leiden, Netherlands; the Naturhistorisches Museum (NHMW), Vienna, Aus- MATERIAL AND METHODS tria; the British Museum of Natural History (NHM), All but one specimen of the studied material was col- London, UK; the Zoological Reference Collection Raf- lected from the eastern riverine mangroves of Mida fles Museum (ZRC), Singapore; and the National Creek, Kenya (near the village of Dabaso, 80 km north Museum of Natural History (USNM), Smithsonian of Mombasa) during September to November 1998 and Institute, Washington, D.C., USA. Measurements of 1999 by D. P. Gillikin and A. Verheyden. One female the studied material represent the carapace width and was found north of Kidogoweni Creek in the riverine length in millimetres. Abbreviations and measure- forest type of the Gazi Bay mangrove system (40 km ments used in this text are: cw, maximum carapace south of Mombasa) (forest types according to Lugo & width; cl, carapace length; bh, body height; iw, inter- Snedaker, 1974). Perisesarma guttatum was collected orbital width; el, extraorbital tooth length; 3mxl, third from both Mida Creek and Gazi Bay mangroves dur- maxilliped length; 3mxw, third maxilliped width; pah, ing the same time. All collections took place during chelar palm height; prp4, propodus length of fourth field surveys for the European Union Project ‘GRO- pereiopod; 4merl, merus length of fourth pereiopod; FLO’ (contract IC18-CT96-0065). 4merw, merus width of fourth pereiopod; 4dacl, dacty- Ecological studies took place in both Mida Creek lus length of fourth pereiopod; tell, telson length; telw, and Gazi Bay mangroves and are detailed in Gillikin telson width; 6abl, length of sixth abdominal segment; (2000). In both forests, presence or absence of crab 6abh, width of sixth abdominal segment. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 435–445 ECOLOGY AND SYSTEMATICS OF EAST AFRICAN PERISESARMA 437 The following material was examined: Holotype: using the software MEGA 2.1 (Kumar et al., 2001) and (SMF 29333) male (24.63 ¥ 20.00 mm), mangroves PAUP* (Swofford, 1998). Subsequently, significance near village of Dabaso, Watamu, Kenya (S03∞20.905¢ levels were evaluated with the bootstrap method E039∞59.498¢), Sept.–Nov.