Geosesarmamirum, a new species of semi-terrestrial sesarmid crab (Crustacea, Decapoda, Brachyura) from central Taiwan

Abstract A new species of semi-terrestrial sesarmid crab of the genus Geosesarma De Man, 1892, is described from central Taiwan. Geosesarmamirumsp. nov. is distinct in possessing a strong transverse crest on the inner surface of the male chela and a diagnostic male first gonopod which is relatively long and stout, with the distal chitinous part broad and spatuliform. Like most Geosesarma species, G.mirumsp. nov. has large eggs and direct development, contrasting with the only other species known from Taiwan, G.hednon Ng, Liu and Schubart, 2004, which has small eggs and planktotrophic larvae.


Introduction
Geosesarma De Man, 1892, is a large genus of semi-terrestrial and terrestrial crabs occurring in many freshwater habitats in Southeast Asia, Andamans, and western Pacific. At the moment, 64 species have been recorded, with the majority from Indonesia (Ng et al. 2008;Ng 2015Ng , 2017Ng et al. 2015;Manuel-Santos et al. 2016;Ng and Gri-nang 2018;Ng and Wowor 2019;. Thus far, the most northerly record of the genus is Taiwan and Luzon in the Philippines, with two known species: G. vicentense (Rathbun, 1914) and G. hednon Ng, Liu & Schubart, 2004(Ng et al. 2004Ng and Lemaitre 2017). Both species, however, are atypical members of Geosesarma as they have small eggs and planktotrophic larvae (see Ng et al. 2004;Ng and Lemaitre 2017). Most species of Geosesarma (when egg sizes are known) (over 90%) have large eggs and direct development.
In late 2018, the authors became aware of a semi-terrestrial freshwater sesarmid living in the lowlands of Chiayi County in central-western Taiwan with large eggs. Specimens were eventually obtained and studies showed that they belonged to a new species of Geosesarma. This is the first record of a large-egged Geosesarma from such a high latitude. The description of the new species and comparisons with congeners form the basis of the present paper.

Materials and methods
The terminology used here follows Ng (1988) and Davie et al. (2015). Measurements provided are the carapace width and length, respectively. The abbreviations G1 and G2 are used for the male first and second gonopods, respectively. Specimens examined are deposited in the zoology collections of the National Taiwan Ocean University (NTOU), Keelung, Taiwan; and the Zoological Reference Collection (ZRC) of the Lee Kong Chian Natural History Museum, National University of Singapore. Specimens with eggs or young crabs were not collected and released after study.
Etymology. The name is derived from the Latin for "surprise", alluding to the unexpected discovery of this species in Taiwan. Remarks. Geosesarma mirum sp. nov. belongs to the group of species which have large eggs (ca. 1.0 mm or greater in diameter, measured in situ), the carapace is trapezoidal to subquadrate, the ambulatory meri are relatively short and stout, the exopod of the third maxilliped has a long flagellum, the inner surface of the male chela has a strong granulated transverse ridge and the G1 is relatively stout with the distal chitinised part spatuliform. The species in this group are: G. amphinome (De Man, 1899) [western Borneo], G. peraccae (Nobili, 1903) [Singapore and Peninsular Malaysia], G. penangense (Tweedie, 1940) [Penang, Peninsular Malaysia], G. sarawakense (Serène, 1968) [Sarawak, Borneo] and G. pylaemenes Ng, 2015 [western Borneo]. Geosesarma mirum can be distinguished from these species mainly by the distinctive form of its G1. Compared to G. amphinome, the distal chitinised part of the G1 of G. mirum is distinctly shorter and the tip is not bilobed (cf. Ng 2015: fig. 1A, B, 2A, G-K).   fig. 5A, B). The G1 of G. mirum differs from that of G. penangense in having the distal part gently curved rather than strongly bent, with the ambulatory leg merus proportionately stouter (cf. Ng 1988: fig. 58A, D, E). When compared to G. sarawakense, the carapace of G. mirum is distinctly more granulated and rugose, with the G1 proportionately stouter and shorter (Ng 2015: figs 6A, B, 7D-F). In contrast to G. pylaemenes, the external orbital tooth of G. mirum is more acute and the G1 is relatively stouter (cf. Ng 2015: fig. 3A, B, 4D-G).
In Taiwan and other parts of the Indo-West Pacific, Geosesarma mirum can be confused with species of Scandarma Schubart, Liu & Cuesta, 2003 (type species Scandarma lintou Schubart, Liu & Cuesta, 2003), and Pseudosesarma Serène & Soh, 1970(type species Sesarma edwardsii De Man, 1887 because in these genera, the male chelipeds do not have pectinated ridges on their chelae and there are no stridulatory granules on the dorsal margin of the dactylus. Geosesarma mirum can be easily distinguished from species of Scandarma as the outer surface of the male chela does not have a distinct swelling and the G1 is very short and stout (see Schubart et al. 2003;Naruse and Ng 2007;Ng 2013;Naruse and Ng 2019). From Pseudosesarma, many members of which live in freshwaters, G. mirum can be distinguished by the G1 morphology, with those of Pseudosesarma species short, very stout with the median or distal parts prominently dilated and with sharp chitinised "beaks" (see Ng and Schubart 2017).
Ecology. Geosesarma mirum sp. nov. has a semi-terrestrial habit and has been found in small streams in lowlands, with the crabs digging burrows under stones near the edge of the water. The first author investigated 28 sites in and around the type locality of species, in the Ba-Jhang River region (total area of ca. 200 km 2 ). Most of the sites examined were badly polluted, cemented, built over or no longer had permanent water. There were some sites with clean water but these were very close to the hills (altitude higher than ca. 100 m) but these sites only had the primary freshwater crab, Geothelphusa olea Shy, Ng & Yu, 1994 (Potamidae). Of the 28 sites surveyed, Geosesarma mirum was only found in six sites (in an area of ca. 4 km 2 ). These six sites were from different branches of the river and the crabs were relatively abundant in each of these locations (sometimes more than 50 individuals/m 2 ). From the surveys done, the species seems to have a relatively localised distribution, with some of the sites where they were found only a few dozen square metres in area, and the crabs absent from sites further upstream or downstream. This may not be the natural situation as the sites where the crabs were absent were invariably badly polluted or extensively concreted.
The development of G. mirum is direct (i.e., abbreviated), with the eggs measuring ca. 1.0 mm in diameter (specimen not preserved) (Fig. 1C). The egg sizes of Geosesarma species average between 1.2 and 1.8 mm (Ng 2017;Ng and Wowor 2019). Despite the slightly smaller egg size, we have observed females of G. mirum brooding young crabs under the pleon (Fig. 1D) and it is clear that the development is completely abbreviated (see Ng and Tan 1995). This contrasts with catadromous grapsoid species like Eriocheir japonica (De Haan, 1835) (Varunidae) which have much smaller eggs (ca. 0.34 mm; cf. Lai et al. 1986). Their eggs, however, are still smaller than those of primary freshwater species like Nanhaipotamon formosanum (Parisi, 1916) (Potamidae) which have egg diameters of up to 4.0 mm (cf. Lai et al. 2012).

Acknowledgements
The first author is most grateful to Dr H-T Lai who accompanied him to collect specimens of this interesting species. Thanks are also due to Mr Peter Wang for his important photographs of the crabs. The first author is also grateful to Dr Shih Hsi-Te for kindly passing the specimens to the second author for study during his recent visit to Singapore, and his many helpful comments on the manuscript. Constructive suggestions by Daisy Wowor and Shih Hsi-Te are also much appreciated.