Crustacean Research 49 KARSTARMA UMBRA, a NEW SPECIES of CAVERNICOLOUS CRAB Which Is Clearly Trapezoidal in Shape with the of K

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Crustacean Research 49 KARSTARMA UMBRA, a NEW SPECIES of CAVERNICOLOUS CRAB Which Is Clearly Trapezoidal in Shape with the of K Crustacean Research 2020 Vol.49: 73–88 ©Carcinological Society of Japan. doi: 10.18353/crustacea.49.0_73 Karstarma umbra, a new species of cavernicolous crab (Brachyura: Sesarmidae) from Vanuatu, with a key to the genus Peter K. L. Ng Abstract.―A new species of cavernicolous sesarmid crab is described from Vanuatu. Karstarma umbra sp. nov. belongs to a group of three species that has the chitinous distal part of the male first gonopod slender and elongate. The new species can easily be distinguished from K. balicum (from Bali, Indonesia) and K. loyalty (from Loyalty Islands) by its distinctly longer ambulatory legs and its relatively more slender male first gonopod. Karstarma umbra is morphologically closest to K. waigeo (from Indo- nesian Papua) but has proportionately longer legs and the outer margin of the male first gonopod is distinctly concave. The taxonomy of the genus is discussed and a re- vised key to the 18 species is provided. LSIDurn:lsid:zoobank.org:pub:D54659BC-D9B8-43FA-ADBE-5A9324F52675 Key words: Taxonomy, Grapsoidea, South Pacific, new species, key, Karstarma, species-groups ■ Introduction zontally on a level surface; the length being taken from the proximal angle with the basis- Seventeen species of the cave sesarmid ge- ischium to the distalmost margin where it artic- nus Karstarma Davie & Ng, 2007, are known ulates with the carpus; while the width is ob- from the Indian Ocean and western Pacific (Ng tained at the widest point, which is usually at et al., 2008; Wowor & Ng, 2009, 2018; Husana the median point (cf. Maenosono & Naruse, et al., 2010; Poupin et al., 2018) (Table 1). A 2016). The following abbreviations are used: new species is here described from caves in G1=male first gonopod; G2=male second Santo Island in Vanuatu. The taxonomy of the gonopod; P2–P5=pereopods 2–5 (ambulatory genus is discussed and a revised key is con- legs 1–4, respectively). structed to help identify the species. Specimens are deposited in the Zoological ■ Taxonomy Reference Collection (ZRC) of the Lee Kong Chian Natural History Museum, Nation- Family Sesarmidae Dana, 1851 al University of Singapore; and the Naturalis Biodiversity Centre (ex Rijksmuseum van Karstarma Davie & Ng, 2007 Natuurlijke Historie, RMNH), Leiden, The Netherlands. Type species The terminology used follows Davie et al. Sesarmoides boholano Ng, 2002, by original (2015). Measurements provided, in millime- designation. tres, are of the maximum carapace width and length, respectively. The ambulatory merus is Remarks measured with the structure positioned hori- Although Wowor & Ng (2009) recognised Received: 29 Feb 2020. Accepted: 14 Apr 2020. Published online: 12 June 2020. 73 PETER K. L. NG Table 1. List of known Karstarma species Species Distribution References Karstarma ardea Wowor & Ng, 2009 Raja Ampat, Indonesian Papua Wowor & Ng (2009) Karstarma balicum (Ng, 2002) Nusa Penida, Bali, Indonesia Ng (2002) Karstarma boholano (Ng, 2002) Bohol, Philippines; Ng (2002), Ishigaki Island, Ryukyus, Japan; Naruse et al. (2005), Green Island, Taiwan Husana et al. (2010), Li et al. (2019) Karstarma cerberus (Holthuis, 1964) Nusa Lain, west of Ambon, Indonesia Holthuis (1964), Fransen et al. (1997), present study Karstarma emdi (Ng & Whitten, 1995) Nusa Penida, Bali, Indonesia Ng & Whitten (1995), Ng (2002) Karstarma guamense (Ng, 2002) Tumon Bay, Guam Ng (2002) Karstarma jacksoni (Balss, 1934) Christmas Island, Australian Indian Ocean Balss (1934), Territory Orchard (2012), Poupin et al. (2018) Karstarma jacobsoni (Ihle, 1912) Gunung Sewu, Central Java, Indonesia Ilhe (1912), Holthuis (1964), Naruse & Ng (2007), Wowor & Ng (2018) Karstarma loyalty (Ng, 2002) Lifou Island, Loyalty Islands, New Caledonia Ng (2002) Karstarma malang Wowor & Ng, 2018 Malang, East Java, Indonesia Wowor & Ng (2018) Karstarma microphthalmus (Naruse & Ng, 2007) Gua Marapetang, south Sulawesi, Indonesia Naruse & Ng (2007) Karstarma novabritannia (Ng, 1988) Arawe Island, New Britain Ng (1988), present study Karstarma philippinarum Husana, Naruse & Kase, 2010 Boracay and Samal Islands, Philippines Husana et al. (2010) Karstarma sulu (Ng, 2002) Palawan, Philippines Ng (2002) Karstarma ultrapes (Ng, Guinot & Iliffe, 1994) Florida and Malaita Islands, Solomon Islands Ng et al. (1994), Wowor & Ng (2009), Husana et al. (2010) Karstarma umbra sp. nov. Santo, Vanuatu Present study Karstarma vulcan Poupin, Crestey & Le Guelte, 2018 Réunion Island Poupin et al. (2018) Karstarma waigeo Wowor & Ng, 2009 Raja Ampat, Indonesian Papua Wowor & Ng (2009) two groups of Karstarma which they defined peared to be closer to species of Geosesarma by the shape of the carapace, proportions of the De Man, 1892 (see Ng & Wowor, 2019) and P3 and P4, as well as structures of the G1 and they form a third group (see also Poupin et al., vulva; three groups are actually discernible. 2018). Wowor & Ng (2018) noted that the three spe- Members of the K. ultrapes species-group cies of Karstarma from Java and Sulawesi ap- are the most distinctive, possessing a carapace 74 Crustacean Research 49 KARSTARMA UMBRA, A NEW SPECIES OF CAVERNICOLOUS CRAB which is clearly trapezoidal in shape with the of K. microphthalmus is also large, so this spe- posterior part very wide; the epigastric and cies probably has large eggs as well. This is the protogastric cristae are fused, forming a large group Wowor & Ng (2018) suggested was re- swelling that protrudes anteriorly, intruding to lated to Geosesarma as all the known Javanese the level of the frontal margin, with the frontal species in this genus have direct development region just or barely visible in dorsal view; the with the large eggs hatching into juvenile crabs cornea of the eye is not reduced; the P4 is very (Ng et al., 2015; Ng & Wowor, 2019). elongate (more than four times the carapace Species in the K. boholano species-group length); there are no prominent long tufts of se- have a subtrapezoidal to quadrate carapace tae between the coxae of P2–P4; the G1 has the with the posterior part slightly wider; the epi- chitinous distal part straight, cylindrical and gastric and protogastric cristae are separate and gently bent outwards at an angle of about distinct but do not protrude anteriorly with the 30–40° along the longitudinal axis; and the frontal region and margin distinct in dorsal vulva is formed by a large U-shaped posterior view; the cornea of the eye is not reduced; the sternal vulval cover with a prominent anterior P4 is long (less than four times the carapace cover, with both projections partially covering length); there are tufts of setae between the the prominent rounded operculum. Three spe- coxae of P2–P4 which are usually dense, long cies are in this group: K. ardea, K. philippina- and prominent; the G1 has the chitinous distal rum and K. ultrapes. No ovigerous females are part short or elongate and sharply bent out- known but on the basis of their small vulvae, wards at an angle of about 90° along the longi- their eggs are probably small and less than a tudinal axis; and the vulva has a prominent millimetre in diameter. posterior sternal vulval cover with a small an- Members of the K. jacobsoni species-group terior one, the whole structure being distinctly also have a trapezoidal carapace but the poste- elevated. Eleven species are in this group: K. rior part is proportionately less wide and the balicum, K. boholano (type species), K. cer- carapace appears more subquadrate; the epi- berus, K. emdi, K. guamense, K. jacksoni, K. gastric and protogastric cristae are lower, with loyalty, K. novabritannia, K. sulu, K. vulcan, the sharp epigastric crista protruding anteriorly and K. waigeo. The egg size of several species to some degree but not completely obscuring are known and they are all small, less than a the frontal region in dorsal view; the cornea of millimetre in diameter (see Ng, 2002; Orchard, the eye is distinctly reduced; the P4 is elongate 2012; Poupin et al., 2018; unpublished data), (less than four times the carapace length); there with Li et al. (2019) recently obtaining plank- are prominent tufts of dense, long setae be- tonic zoeae for K. boholano. Most of the spe- tween the coxae of P2–P4; the G1 has the chi- cies have a short and truncate chitinous distal tinous distal part straight, cylindrical and gen- part of the G1; only in three species is the chi- tly bent outwards at an angle of about 30–40° tinous distal part elongate and tapering̶K. along the longitudinal axis; and the vulva has a balicum, K. loyalty and K. waigeo, like in K. simple broad lateral sternal vulval cover with umbra sp. nov. the structure almost level with the sternal sur- The figures of K. cerberus and K. novabri- face. Three species are in this group: K. jacob- tannia appear to depict the two species pos- soni, K. malang and K. microphthalmus. The sessing epigastric cristae which are directed eggs of the first two species are large (ca. anteriorly and reaching the frontal margin 1.8 mm in diameter) and they almost certainly (Holthuis 1964: fig. 1, 2a; Ng 1988: pl. 1) like practice abbreviated or semi-abbreviated de- those in the K. ultrapes species-group (Fig. 6A, velopment (Wowor & Ng, 2018). The vulvae B). This is, however, not accurate as the cara- Crustacean Research 49 75 PETER K. L. NG pace had been slightly tilted anteriorly when be regarded as stygobites. All the other species, the structure was originally drawn or photo- including K. umbra, are best described as sty- graphed. Both species actually have normal gophiles or stygoxenes, and while mainly epigastric cristae that do not protrude anteriorly found in caves, are probably found in habitats (Fig. 5A–D). These two species also have setae outside the caves as well (see Ng, 2013; Ng & between the coxae of P2–P4 but the setae are Guinot, 2014).
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