Assemblage structure, distribution and habitat type of the grapsoid crabs (Brachyura: Grapsoidea) of the coastal forested swamps of northern Borneo

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Abstract

Several intertidal and supratidal systems were investigated in the Brunei Bay to assess the diversity and assemblage structure of grapsoid crabs (Brachyura: Grapsoidea) in different salinity and substrate conditions, type of vegetation, and distance from water bodies. The structure of the assemblages differed remarkably across and within sites, and several taxa were not previously recorded from this region. Diversity decreased towards more terrestrial conditions, with systems closer to the sea supporting more diversified assemblages. High and low intertidal areas also hosted different assemblages in terms of taxonomic composition. Most of these species can be considered habitat specialists, and only occurred in a small number of the investigated areas. A small number of habitat generalists were found in several areas, exhibiting eurytopic ecological traits. A time-based sampling method was used to estimate population densities. A few habitat specialists were recorded in high densities, suggesting adaptations to specific environmental conditions in which these species can bloom. These results highlight how the diversity of these assemblages needs to be considered in management plans, to prevent local extinctions and loss of diversity in the area.

Introduction

Grapsoid crabs (Brachyura: Grapsoidea) are dominant components of the intertidal macrofauna of mangrove forests and other wetland ecosystems of the Indo West Pacific region (IWP; Lee, 1998). Grapsoid crabs are ancestrally marine organisms, that successfully colonised semi-terrestrial and terrestrial systems, such as mangrove and peat swamp forests, limestone caves, and rain forests (e.g. Ng and Lim, 1987, Lee, 1998, Davie and Ng, 2007). Despite the key role of these species in ecosystem functions and processes, their ecology and diversity are poorly known (Hogarth, 2007, Lee et al., 2017). The taxonomy of the large family Sesarmidae, which dominates these assemblages, is in a state of flux (e.g. Ng and Schubart, 2002, Davie and Ng, 2007, Ng et al., 2008, Rahayu and Ng, 2009, Naderloo and Schubart, 2010, Rahayu and Li, 2013, Cannicci and Ng, 2017). The autecology of wetland grapsoid crabs has been investigated only in a few species, mainly in Neotropical (e.g. Warner, 1969, Erickson et al., 2008, Diesel et al., 2000) and African systems (e.g. Cannicci et al., 1999, Gillikin and Schubart, 2004, Emmerson and Ndenze, 2007). Data on the habitat environmental conditions of many species are only available from anecdotal information in taxonomic descriptions (e.g. Rahayu and Ng, 2009, Tweedie, 1940, Tweedie, 1950a). There is also a scarcity of studies on the distribution and abundance of grapsoid crabs in mangrove and other coastal ecosystems (Salgado Kent and McGuinness, 2010). This lack of ecological information has been attributed to taxonomic uncertainty and complexity, practical field work limitations, and the challenges posed by population density measurements (e.g. Salgado Kent and McGuinness, 2006, Hogarth, 2007, Lee et al., 2017).

In northern Borneo, the only quantitative study on the community ecology of grapsoid crabs living in forested coastal wetlands is Ashton et al. (2003b). Other reports from this region include annotated checklists (e.g. Choy, 1991, Tan and Ng, 1994, Tweedie, 1950b), qualitative reports (Choy and Booth, 1994), and species descriptions (e.g. Ng, 1995a, Ng, 1995b, Ng, 2013).

The present study investigates the grapsoid crabs from mangrove and other peritidal soft-bottomed ecosystems of the Brunei Bay (northern Borneo), and in particular (i) assemblage structure and diversity, (ii) species spatial distribution and their association with selected environmental variables, and (iii) level of association of the species within and among assemblages.

Section snippets

Study sites and sampling

Brunei Darussalam (Fig. 1) is characterised by a tropical climate, with an average annual temperature of 27 °C, and an average annual rainfall of 2880 mm (average value from 1966 to 2006), with two peaks in correspondence of the southwest monsoon (May), and the northeast monsoon (December) (Brunei Meteorological Service, Department of Civil Aviation). The study sites include several coastal wetlands in the Brunei Bay and along the coast of the South China Sea (Fig. 1), influenced by both marine

Structure and diversity of the assemblages

A total of 306 individuals were collected, including 13 grapsoid genera, 25 species, two morphospecies (i.e. Episesarma sp.1 and Parasesarma sp.1), and 25 immature specimens, which could only be discriminated to the genus level (Table 3). Fourteen species were previously not recorded in the Brunei Bay, and eight of these are also new records for Borneo (Ashton et al., 2003b, Choy, 1991, Choy and Booth, 1994, Tan and Ng, 1994; Table 3, Fig. 2).

The highest species richness was measured in site

Structure and diversity of the assemblages

When considering only the mangrove sites (i.e. sites 1–4), the number of recorded taxa (n = 9–14) is comparable with previous studies from other South East Asian mangrove sites [e.g. Sasekumar, 1974 (n = 12); Frith et al., 1976 (n = 15); Ashton et al., 2003b (n = 17); Ashton et al., 2003a (n = 12); Diele et al., 2013 (n = 12)]. For these ecosystems, our data confirm this biogeographic region as a biodiversity hotspot (Briggs, 1999, Myers et al., 2000), hosting assemblages of higher diversity

Conclusions

The present study investigates the diversity of several grapsoid assemblages in different peritidal systems in northern Borneo. The assemblages differ among sites with different influence of freshwater, type of substrate and vegetation, and at different distances from water bodies. The results suggested that the diversity of these systems is currently underestimated, prompting for further investigations. Several species have stenotopic ecological traits, being associated with peculiar habitat

CRediT authorship contribution statement

Laura Ribero: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, Writing - review & editing. Phaik Eem Lim: Funding acquisition, Project administration, Resources, Supervision, Validation, Writing - review & editing. Rosli Ramli: Project administration, Resources, Software, Supervision, Validation, Writing - review & editing. Gianluca

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgements

Thanks to Peter K. L. Ng (National University of Singapore), Ngan Kee Ng (National University of Singapore) and Dwi Listyo Rahayu (Indonesian Institute of Science, Jakarta), for their help with taxonomic discriminations. Thanks to Tan Siong Kiat and Jose’ Christopher E. Mendoza for curatorial assistance during L.R. visits to the Lee Kong Chian Natural History Museum in Singapore. Thanks to Claas Damken for the specimens he collected during his entomological surveys in Sungai Belayang, Brunei.

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