Crustacea: Decapoda)

Zootaxa 3640 (2): 224–241 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3640.2.6 http://zoobank.org/urn:lsid:zoobank.org:pub:77FB19E2-2C2D-4F16-A0A4-A04356270668 New records of processid shrimps from the Indo-West and East Pacific (Crustacea: Decapoda)

SAMMY DE GRAVE1 & ARTHUR ANKER2 1Oxford University Museum of Natural History, Parks Road, Oxford, OX1 3PW, United Kingdom. E-mail: [email protected] 2Department of Biological Sciences / TMSI, Ecology and Systematics Laboratory, National University of Singapore, 14 Science Drive 4 / Lower Kent Ridge Road, Singapore 119260, Republic of Singapore. E-mail: [email protected]

Abstract

New records are presented for the caridean shrimp family Processidae in Indo-West and East Pacific waters, based mainly on recently collected material. These records highlight that many processid species may be relatively widespread, although the distribution of some species remains imperfectly known. The colour pattern of several species is here documented for the first time, showing its potential use as an additional identification character. The available information on the ecology of the reported species is summarised.

Key words: Decapoda, Processidae, Indo-Pacific, new records, colour pattern

Introduction

The Processidae is a relatively species-poor family, with currently only 65 species placed in 6 genera (De Grave & Fransen 2011; Hendrickx 2012). However, they can be very abundant in certain habitats, notably tropical seagrass beds (e.g. Ledoyer 1969, 1970, 1984; Unsworth et al. 2007, 2011). A solid foundation for their taxonomy was laid down by Hayashi (1975), who reviewed all Indo-Pacific species and provided a worldwide key to all then known taxa. Since then only a handful of species have been described in the Indo-Pacific (e.g. Hayashi 1981; Noël 1986; Burukovsky 1990, 2007), with the last worldwide key provided by Noël (1986). Despite their apparent solid taxonomic framework, processid distribution and ecology in the Indo-Pacific remains poorly known, even for purported widespread taxa, quite probably linked to their nocturnal lifestyle (e.g. Unsworth et al. 2010) and general rarity in samples. In this contribution, accumulated material in several museums is reported upon, which sheds further light on their distribution. In recent years, much emphasis has been placed on colour patterns of caridean shrimps, notably Alpheidae and Pontoniinae. In contrast, the colour patterns in Processidae and indeed their potential use for species identification has been poorly documented and discussed. This was recently highlighted by De Grave & Felder (2012), who noted potential differences in colour pattern between two common Caribbean species, but at the same time drew attention to diurnal differences in pattern and discrepancies with older colour descriptions. Asides from photos published in several popular underwater guide books, for instance, Hayashidonus japonicus (De Haan, 1844) in Kuiter & Debelius (2009) and Nikoides maldivensis Borradaile, 1915 in Minemizu (2013), there appear to be no known specimen matched photos of Indo-Pacific Processidae at present in the scientific literature. This is here rectified for several taxa, which in some species shows promise as an aid in field identification. As however, the colour pattern in Processidae is highly influenced by time of the day, more specimen-matched photographic images need to be accumulated to fully ascertain its usefulness in species identification. The material reported herein primarily originates from three sources: (1) fieldwork carried out over the last decade in the Indo-Pacific by the Oxford University Museum of Natural History, Oxford, UK (OUMNH) and miscellaneous donations to that collection; (2) recent expeditions by the Florida Museum of Natural History, University of Florida, Gainesville, USA (FLMNH UF); and (3) unreported as well as recently collected material in

224 Accepted by J. Goy: 25 Feb. 2013; published: 17 Apr. 2013 the collections of the Zoological Reference Collection of the Raffles Museum of Biodiversity Research, National University of Singapore, Singapore (ZRC). The ecology for each species is summarised, based on present records as well as scattered literature information. Post-orbital carapace length (pocl, in mm) was measured from the edge of the orbit to the posterior margin of the carapace; latitude and longitude are reported in decimal annotation; fieldwork lot numbers are reported between brackets.

Taxonomy

Processidae Ortmann, 1896

Ambidexter Manning & Chace, 1971

Ambidexter swifti Abele, 1972 Fig. 1A–B

Ambidexter swifti Abele, 1972: 366, figs. 1–3.

Material examined. Panama: 2 ov. females (pocl 5.5, 6.0), OUMNH.ZC.2009-18-012, Chumical, mixed rock- sand intertidal exposed at night, tide pool, leg. A. Anker, 17.04.2007. Colour pattern. Cephalothorax and abdomen generally transparent, with numerous red-brown chromatophores, larger on cephalothorax than abdomen; third maxillipeds and first pereiopod also covered with chromatophores, posterior pereiopods transparent (Fig. 1A–B). Remarks. The specimens present no noteworthy features and correspond closely to the type description by Abele (1972). Ecology. The ecology of A. swifti is relatively poorly known. The majority of specimens reported in Abele (1972) were collected from burrows on a mid-tidal, sandy beach. The present specimens were collected in a similar habitat (rock-sand shore); however, they were dwelling in tide pools at night. Distribution. According to Wicksten & Hendrickx (2003) the species is distributed from San Diego, California southwards to Panama, also occurring in the Galapagos Islands. However, records are very sparse throughout its range.

Ambidexter panamensis Abele, 1972 Fig. 1C

Ambidexter panamensis Abele, 1972: 373, figs. 4–5. —Hendrickx, 1988: 246.

Material examined. Panama: 1 female (pocl 3.3), OUMNH.ZC.2007-13-015, Chame Bay, exposed mud flat, burrow, suction pump, leg. A. Anker et al., 07.10.2006. Colour pattern. Body largely transparent, with sparse red chromatophores on cephalothorax and abdomen, pereiopods transparent (Fig. 1C). Remarks. The specimen presents no noteworthy features and corresponds closely to the type description by Abele (1972). Ecology. The ecology of A. panamensis is known rather insufficiently. Abele’s (1972) specimens were collected from sandy tide pools. Wicksten (1983) equally reported specimens from tidal pools, but also included a single specimen dredged from 65 m. The present specimen was collected with a suction pump on an intertidal mudflat, but it remains unknown if the shrimp was dwelling inside a burrow of an unidentified burrowing host or in its own shelter in the vicinity of the burrow. Distribution. According to Wicksten & Hendrickx (2003) the species is distributed from Baja California southwards to Panama, also occurring in the Galapagos Islands.

NEW RECORDS OF PROCESSIDAE Zootaxa 3640 (2) © 2013 Magnolia Press · 225 FIGURE 1. Eastern Pacific species of Ambidexter Manning & Chace, 1971: A, A. swifti, ov. female, Panama, lateral view (OUMNH.ZC.2009-18-012); B, same specimen, on white background; C, A. panamensis, female, Panama, lateral view (OUMNH.ZC.2007-13-015).

Clytomanningus coutierei (Nobili, 1904)

Processa coutierei Nobili, 1904: 234.—Nouvel, 1945: 395, figs. 1–8.—Hayashi, 1975: 95, figs. 17–18. Clytomanningus coutierei.—Chace, 1997: 34 (key).

Material examined. Papua New Guinea: 1 ov. female (pocl 3.0), OUMNH.ZC.2011-02-0026, New Ireland, Nusalik, seagrass bed, leg. M. Dowell, 28.08.2004. Republic of Palau: 1 ov. female (pocl 2.2), 2 females (pocl 1.5, 1,7), OUMNH.ZC.2012-01-076, Palau, outside of Malakal Harbour, 7.27065 134.46523, 55 m, light trap, leg. S. De Grave & C. Burras, 31.05.2002 [st 288]. Colour pattern. Not recorded. Remarks. All specimens agree with the re-description of the type material in Nouvel (1945) and the description of Kenyan material in Hayashi (1975). Ecology. Both Ledoyer (1984) and Unsworth et al. (2007, 2010) record C. coutierei from shallow water seagrass beds, 8 m and less than 2 m, respectively. Unsworth et al. (2010) showed that the species has a particularly pronounced diurnal change in abundance, which is likely to account for the paucity of records. Interestingly, the specimens from Palau were collected at 55 m depth, the deepest the species has been recorded so far. Distribution. This species was considered to be very rare by Hayashi (1975), as at that time it was only known from Djibouti and Kenya. Since then, it has been recorded from New Caledonia (Ledoyer 1984), Oahu, Hawaii (Titgen 1987), Sulawesi, Indonesia (Unsworth et al. 2007, 2010), and is herein recorded from Papua New Guinea and Palau.

Clytomanningus molaris (Chace, 1955) Fig. 2

Processa molaris Chace, 1955: 11, fig. 5.—Hayashi, 1975: 124, figs. 29–30. Clytomanningus molaris.—Chace, 1997: 34.

Material examined. French Polynesia: 1 female (pocl 1.7), FLMNH UF 23275, Moorea, back reef at west corner of Cook’s Bay Pass, -17.4815 -149.8255, backreef, 1–2 m, algal wash, leg. L. Watling, 12.12.2009 [BMOO-10719, BIZ-284]. Hawaii: 1 ov. female (pocl 1.8), FLMNH UF 15204, Maui, Wahikuli Beach Park, 20.9189 -156.6644, Halimeda bed, 6–12 m, algal wash, leg. P. Fiene, 30.11.2008 [G-08-087, Pittman-1081130]; 1 female (pocl 1.7), FLMNH UF 15205, Maui, Wahikuli Beach Park, 20.9189 -156.6644, Halimeda bed, 6–12 m, algal wash, leg. P. Fiene, 30.11.2008 [G-08-088, Pittman-1081130]. Colour pattern. Semitransparent with irregular large white areas on carapace and abdomen (some possibly covering internal organs); isolated patches of red chromatophores present on carapace, e.g., around orbits, and on abdomen, here forming faint red bands; antennular peduncles with small white patch distally; third maxilliped white proximally, greyish-white with red chromatophores distally; first pereiopods mostly white; remaining pereiopods transparent with some red chromatophores; tail fan with large white patches (Fig. 2). Remarks. The material from Hawaii and Moorea corresponds closely to the description by Hayashi (1975). Ecology. Although the ecology of this species is still poorly known, it appears to occur in reef-associated environments, e.g., Halimeda beds, lagoon reefs and pools, between low tide mark and 12 m (Hayashi, 1975; present records). Distribution. This species appears to be widespread in the Indo-Pacific, although it has been only formally recorded from a few scattered localities. Hayashi (1975) provided records of C. molaris from Rongelap and Bikini atolls (Marshall Islands), Paternoster Island (currently Kepulauan Tengah) (Indonesia), Eilat (Red Sea) and Kenya. Devaney & Bruce (1987) provided the only further record, from Enewetak Atoll (Marshall Islands). The present records constitute the first for the species from French Polynesia and Hawaii.

NEW RECORDS OF PROCESSIDAE Zootaxa 3640 (2) © 2013 Magnolia Press · 227 FIGURE 2. Clytomanningus molaris (Chace, 1955). A, female, dorsal view, French Polynesia (FLMNH UF 23275); B, same specimen, lateral view.

Hayashidonus Chace, 1997

Hayashidonus japonicus (De Haan, 1844)

Nika japonica De Haan, 1844 [in De Haan, 1833–1850]: pl. 46, fig. 6. Processa japonica.—De Man, 1920: 208; pl. 18, fig. 53.– Hayashi, 1975: 110, fig. 24.—Noël, 1986: 287. Hayashidonus japonicus.—Chace, 1997: 35, fig. 18.—Kim et al., 2011: 262, fig. 1.

228 · Zootaxa 3640 (2) © 2013 Magnolia Press GRAVE & ANKER Material examined. Republic of Palau: 1 female (pocl 7.4), OUMNH.ZC.2012-01-075, Palau, outside of Malakal Harbour, 7.27065 134.46523, 55m, light trap, leg. S. De Grave & C. Burras, 31.05.2002 [st 288]. Colour pattern. Not recorded. Remarks. The single specimen presents no noteworthy features and corresponds closely to the descriptions in De Man (1920) and Hayashi (1975), with the broadly triangular rostrum being highly characteristic for the species. Ecology. Ledoyer (1984) and Unsworth et al. (2010) found H. japonicus to be particularly abundant in nocturnal seagrass bed samples in New Caledonia and Sulawesi, respectively. However, the depth range of the species (0–150 m) indicates that it is not exclusively associated with seagrass beds. Distribution. This species is widely distributed in the Indo-Pacific, from East Africa through Indonesia and the Philippines to Japan (Hayashi 1975; Noël 1986; Chace 1997) and Korea (Kim et al. 2011), and has been recorded from almost intertidal depths down to 150 m (Noël 1986). The present new record from Palau constitutes the eastern-most record for the species.

Nikoides Paulson, 1875

Nikoides danae Paulson, 1875 Fig. 3

Nikoides danae Paulson, 1875: 98; pl. 14, figs. 5-5d.– Hayashi, 1975: 53, figs. 1–2.– Noël, 1986: 263.—Chace, 1997: 36.

Material examined. Madagascar: 1 male (pocl 4.2), FLMNH UF 14153, Nosy-Bé, west of Hellville, -13.4156 48.2465, sand flat with seagrass bed, 0–3 m, leg. A. Anker, G. Bakary, F. Michonneau, G. Paulay & T. Werner, 25.05.2008 [BMADA-0487, MGNW-50]. Mariana Islands: 1 ov. female (pocl 7.5), FLMNH UF 900, Guam, Tepungan Channel, 2–3 m, leg. Rittson-Williams, 01.03.2000 [ZZZ-015510]; 1 ov. female (pocl 7.1), FLMNH UF 23185, Guam, Tepungan Channel tunnel, 13.4646 144.688, 0–4 m, leg. G. Paulay, 16.06.2003 [BGUAM-106]. Australia: 1 female (pocl 4.6), FLMNH UF 27713, Western Australia, Ningaloo Reef, off “Ningaloo base camp”, - 22.67 113.65, backreef near reef breakers, sand with coral bommies and reef platform, in coral rubble crevices, 2–3 m, leg. A. Anker, 23.10.2010 [AUST-7166]. Singapore: 1 post-ov. female (pocl 6.4), ZRC 2013.0398, Pulau Sekudu off Check Jawa (Pulau Ubin), sand-mud flat with seagrass and coral rubble exposed at low tide, burrow, suction pump, leg. A. Anker, 18.10.2012 [JS-1448]. Colour pattern. Semitransparent with pale orange tinge and numerous white or pale-yellow, irregular spots of various sizes on carapace and abdomen; spots most abundant on posterior half of abdomen; first to fourth abdominal somites with diffuse red transverse bands dorsally; isolated patches of red chromatophores present on carapace, e.g., posterior to orbits; antennular peduncles with alternating pale orange and white patches, and some red chromatophores; third maxilliped and first pereiopods reddish with conspicuous white patches; remaining pereiopods mostly transparent; tail fan densely covered with pale yellow and whitish spots and reddish areas (Fig. 3). Remarks. The present material confirms the variability of the characters previously noted by Noël (1986), with some specimens being intermediate between N. danae and the closely related N. gurneyi Hayashi, 1975 (see below). Ecology. Nikoides danae is found in high abundances on seagrass beds, primarily at night (Ledoyer, 1984; Unsworth et al., 2010). However, the present records concur with previous observations (see Hayashi 1975) that this species has a much wider habitat preference than just seagrass beds, as some specimens were collected from reef habitats with corals and coral rubble. Although primarily a shallow water species (up to intertidal), N. danae has also been recorded from a depth of 32 m (De Man, 1920). Distribution. The species is known from the Red Sea, East Africa and Madagascar, eastwards to Hawaii and Great Barrier Reef (Hayashi 1975); it is herewith newly recorded from Guam, Singapore and Western Australia.

NEW RECORDS OF PROCESSIDAE Zootaxa 3640 (2) © 2013 Magnolia Press · 229 FIGURE 3. Nikoides danae Paulson, 1875. A, female, dorsal view, Western Australia (FLMNH UF 27713); B, same specimen, lateral view; C, male, lateral view, Madagascar (FLMNH UF 14153).

Nikoides gurneyi Hayashi, 1975: 58, figs. 3–4.—Chace, 1997: 36.

Material examined. Singapore: 4 ov. females (pocl 8.6-8.7), ZRC 2012.0974, Pulau Ubin, Chek Jawa, leg. N. Sivasothi et al., 24.07.2001. Malaysia: 1 ov. female (pocl 5.0), ZRC 2012.0975, Pulau Tioman, Paya Beach, leg. NUS Honours Field Class, 07–12.09.2002. Taiwan: 2 females (pocl 4.2, 4.5), OUMNH.ZC.2010-02-072, Houpihu Village, Pingtung County, 21.935843 120.746527, night dive, leg. C.W Lin & T. Naruse, 25.07.2009. Colour pattern. Not recorded. Remarks. The specimens assigned herein to N. gurneyi have the pleuron of the fifth abdominal somite acutely pointed and the stylocerite mesially and laterally pointed, thus fitting the diagnosis of the species in Hayashi (1975). It is acknowledged, however, that Noël (1986) suggested that N. gurneyi may be synonymous with N. danae, a taxonomic subject beyond the scope of the present study. Ecology. According to Ledoyer (1984), N. gurneyi co-occurs with N. danae in shallow seagrass beds. As with that species, other records (see Hayashi 1975) suggest the species is not restricted to seagrass beds, and equally deeper water records are known, down to 27 m (Hayashi 1975). Distribution. The distribution of the species is unclear, due to confusion in older literature with the closely related N. danae. According to Hayashi (1975), the species is known from Sumatra (Indonesia), Egypt (Red Sea), Kenya, Tanzania and Mozambique. The only further records are by Ledoyer (1984) from New Caledonia and the present records from Singapore, Malaysia and Taiwan.

Nikoides maldivensis Borradaile, 1915 Fig. 4

Nikoides maldivensis Borradaile, 1915: 209.—Borradaile, 1917: 411, pl. 58, fig. 11.—Hayashi, 1975: 62, fig. 5—Chace, 1997: 36. Processa jacobsoni De Man, 1921: 95.

Material examined. Mariana Islands: 1 ov. female (pocl 6.1), FLMNH UF 4125, Guam, Tepungan Channel tunnel, 13.4646 144.688, 0–4 m, leg. G. Paulay, 16.06.2003 [BGUAM-106]; 1 ov. female (pocl 8.2), FLMNH UF 4194, Guam, Tepungan Channel tunnel, 13.4646 144.688, 0–4 m, leg. G. Paulay, 16.06.2003 [BGUAM-106]. French Polynesia: 1 male (pocl 4.5), FLMNH UF 23270, Moorea, Atiha Bay, inside barrier reef, near pass channel, -17.5957 -149.8456, under and on rocks and rubble, hand collecting, leg. S. McPherson, G. Paulay & C. Meyer, 08.12.2009 [BMOO-9974]; 1 female (pocl 5.0), FLMNH UF 23274, Moorea, NW Motu Islands, in channel between two islands, rubble field in shallow lagoon, -17.4893 -149.9132, under large boulders, 0.5–2 m, hand collecting, leg. S. McPherson, J. Houder & A. Crowther, 25.11.2009 [BMOO-9278]; 1 ov. female (pocl 7.6), FLMNH UF 23276, Moorea, lagoon in front and west of Hilton Hotel, -17.4823 -149.825, under rocks and on ledges, 10–15 m, leg. G. Paulay, C. Meyer, S. McPherson, S. Haddock, M. Parker & S. McKeon, 22.10.2009 [BMOO-6503]. Australia: 1 female (pocl 3.6), ZRC 2012.0976, Christmas Island, Flying Fish Cove, -10.429152 105.669210, reef slope, night dive, leg. H.H. Tan et al., 01.02.2010 [CI-D17(152)]. Colour pattern. Bright yellow-orange, resulting from small reddish chromatophores on semitransparent yellow background, irregular white chromatophores forming short, dorsal, transverse bands on abdomen, sometimes connected by thin longitudinal line of white chromatophores, and larger or smaller patches and spots elsewhere on abdomen and on carapace; antennular peduncles yellow with bright white spots; third maxilliped reddish-orange with conspicuous white patches; pereiopods mostly uniform yellow; tail fan yellow densely covered with large areas of white chromatophores (Fig. 4). Remarks. Chace (1997) clarified the type locality of N. maldivensis as Amirante Island, Seychelles (as stated in Borradaile 1917), with the “Maldive Islands” as cited in Borradaile (1915) being in error. The present material presents no particular features, with the large dorsal tooth separated from the rostral apex being highly diagnostic for the species. The colour pattern of N. maldivensis also seems to be very characteristic, as the Moorea specimens (Fig. 4) match the colour photo in Minemizu (2013).

NEW RECORDS OF PROCESSIDAE Zootaxa 3640 (2) © 2013 Magnolia Press · 231 FIGURE 4. Nikoides maldivensis Borradaile, 1915. A, ov. female, dorsal view, Moorea (FLMNH UF 23276); B, same specimen, lateral view; C, male, dorsal view, Moorea (FLMNH UF 23270).

232 · Zootaxa 3640 (2) © 2013 Magnolia Press GRAVE & ANKER Ecology. This species appears to be restricted to coral reefs and adjacent rubble fields, usually dwelling in crevices of coral rubble and dead coral heads or under large rocks, and roaming on the reef or dock walls and ledges at night. It appears to prefer very shallow water, with a depth range of 0–4 m. Distribution. The species has a wide distribution in the Indo-West Pacific, based on previous records from Kenya, Seychelles (Amirante, Farquhar Atoll), Indonesia (Sumatra), Micronesia (Pohnpei), Samoa, and Hawaii (Hayashi 1975; Chace 1997) and is here recorded for the first time from Guam, Christmas Island and French Polynesia. Chace (1997) also mentioned Fiji and Caroline Islands as part of the distribution of N. maldivensis, but it is unclear on which material these records were based.

Nikoides multispinatus Hayashi, 1981 Fig. 5

Nikoides multispinatus Hayashi, 1981: 53; figs. 1–3.—Hayashi, 1984: 221, fig. 1.

Material examined. Republic of Palau: 1 ov. female (pocl 5.0) FLMNH UF 3855, Palau, between Malakal and Koror, near causeway, 5–10 m, leg. G. Paulay, 22.07.1999 [ZZZ-088941]. Australia: 1 female (pocl 2.6), FLMNH UF 18243, Queensland, Great Barrier Reef, Lizard Island, “Washing Machine” dive site, in dead Pocillopora sp., - 14.6504 145.4621, leg. M. Timmers, 18.02.2009 [AUST-2218, AUST-St-105MT]. Mariana Islands: 1 female (pocl 4.6), FLMNH UF 900, Guam, Tepungan Channel, on wall, 2–3 m, leg. J. Starmer, 03.12.1998 [ZZZ-003627]; 1 ov. female (pocl 4.7), FLMNH UF 23185, Guam, Tepungan Channel tunnel, 0–4 m, 13.4646 144.688, leg. G. Paulay, 16.06.2003 [BGUAM-106]. French Polynesia: 1 male (pocl 4.5), FLMNH UF 9887, Moorea, NE of entrance to Opunohu Bay, -17.4915 -149.8502, fringing reef and sandy slope of lagoon, on surface of Porites sp. at night, 1–3 m, hand collecting, leg. G. Paulay, 19.06.2006 [BMOO-1144, GP-Loc-861]; 1 male (pocl 3.5), FLMNH UF 23273, Moorea, Opunohu Pinnacle, -149.8624 -17.4952, silty wall, on/in dead piece of wood, leg. C. Meyer, 26.10.2009 [BMOO-07135, BIZ-050]; 1 male (pocl 3.0), FLMNH UF 16346, Moorea, between Cook’s Bay and Sheraton Hotel, -17.4767 -149.8300, outer reef slope, in large piece of dead coral, leg. S. McKeon, J. Moore & V. Ivanenko, 14.11.2008 [BMOO-5438, MIB-209]; 1 male (pocl 4.0), FLMNH UF 23271, Moorea, off Gump Station dock, - 17.4908 -149.8261, on wall, 0.2–0.7 m, leg. A. Anker, 23.10.2009 [BMOO-07132, BIZ-033]. Papua New Guinea: 1 ov female (pocl 7.5), OUMNH.ZC.2010-04-0025, Kimbe Bay, close to hole in reef wall, 25–30 m, hand collecting at night, leg. J.A Baeza, 07.2009. Taiwan: 1 female (pocl 6.0), OUMNH.ZC.201-02-074, Green Island, Chaikou Village, 22.677400 121. 482137, night dive, leg. C.W. Lin & T. Naruse, 21.07.2009; 2 females (pocl 4.0, 4.2), OUMNH.ZC.201-02-069, Green Island, Chaikou Village, 22.677400 121. 482137, night dive, leg. C.W. Lin & T. Naruse, 21.07.2009. Colour pattern. Semitransparent with five narrow red transverse bands on abdomen; appendages mostly colourless with some red chromatophores, especially on third maxilliped and first pereiopods; tail fan with pale reddish tinge (Fig. 5). Remarks. Hayashi (1984) considered the presence/absence of the posterior spine on the lobe of the sixth pleonite of N. multispinatus to be sexual dimorphism (present in females, absent in males). However, two of the males examined during the present study also have such a spine. More material of this species will need to be examined to clarify whether this is indeed a sexually dimorphic character or a variable one. The present material is also variable in the number of dorsal spines on the telson, which varies from three to six, with the spines not always arranged in lateral pairs. Ecology. On the basis of the present records and scattered information in the literature (Hayashi 1981, 1984), N. multispinatus seems to frequent coral reef lagoons and slopes, usually dwelling in coral rubble and dead coral heads; the known depth range is around 0–30 m. Nocturnal records suggest that shrimps roam on the reefs at night. Distribution. The species was previously known from the type series from Heron Island, Great Barrier Reef, Australia and Ishigaki-jima, Ryukyu Archipelago, Japan (Hayashi 1981), and additional specimens from Enewetak Island, Marshall Islands (Hayashi 1984) and Abaiang Atoll, Gilbert Islands (Hayashi 1986). The present records from Guam, Republic of Palau, Society Islands (French Polynesia), Papua New Guinea and Taiwan demonstrate that N. multispinatus is widespread in the western Pacific Ocean.

NEW RECORDS OF PROCESSIDAE Zootaxa 3640 (2) © 2013 Magnolia Press · 233 FIGURE 5. Nikoides multispinatus Hayashi, 1981. A, male, dorsal view, Moorea (FLMNH UF 23271); B, same specimen, oblique view; C, male, dorsal view, Moorea (FLMNH UF 16346).

Processa steinii Edmondson, 1935: 3, fig. 1. Processa nanus Chace, 1955: 8, fig. 4. Nikoides steinii Hayashi, 1975: 69, figs. 8–9.—Chace, 1997: 37.

Material examined. French Polynesia: 1 ov. female (pocl 3.6), FLMNH UF 23278, Moorea, W of Cook Bay Pass, in front of Hilton Hotel, -149.8442 -17.4771, reef front, on or near reef crest, 3–6 m, hand collecting at night, leg. G. Paulay, S. McKeon, S. McPherson, S. Haddock & A. Collins, 23.10.2009 [BMOO-06746, BIZ-029]. Australia: 1 female (pocl 3.4), FLMNH UF 24679, Queensland, Great Barrier Reef, Heron Island, Wistari Reef, -23.452 151.8671, lagoon, patch reefs, rubble, 11–13 m, hand collecting, leg. J. Reimer, 11.11.2009 [AUST-4634, HI09- 002]; 1 ov. female (pocl 4.5), FLMNH UF 17488, Queensland, Great Barrier Reef, Lizard Island, Snake Pit, sta. 2, forereef, coral rubble, 5–15 m, leg. S. Smith, K. Mills & G. Cranitch, 24.02.2009 [AUST-1765, AUST-ST-089]; 1 female (pocl 3.2), FLMNH UF 17190, Queensland, Great Barrier Reef, Lizard Island, Day Reef, forereef, rubble wash, leg. N. Bruce, 19.02.2009 [AUST-1295, AUST-ST-061]; 1 female (pocl 3.1), FLMNH UF 27705, Western Australia, Ningaloo Reef, off “Ningaloo base camp”, -22.67 113.65, backreef near reef breakers, sand with coral bommies, rubble cracking, 2–3 m, leg. A. Anker, 23.05.2010 [AUST-7156]. Republic of Palau: 1 ov female (pocl 4.1), OUMNH.ZC.2012-01-077, Malakal Channel, 7.317 134.4607, from loose rubble, 2 m, leg. S. De Grave & C. Burras, 22.05.2002 [st 95]. Taiwan: 1 ov. female (pocl 4.5), 1 female (pocl 4.6), OUMNH.ZC.2010-02-070, Green Island, General Rock, 22.676964 121.495437, night dive, leg. C.W. Lin & T. Naruse, 20.07.2009; 1 female (pocl 4.2), OUMNH.ZC.201-02-071, Green Island, Chaikou Village, 22.677400 121. 482137, night dive, leg. C.W. Lin & T. Naruse, 21.07.2009. Colour pattern. Semitransparent with large dorsal patch of red chromatophores on carapace; isolated patches of red chromatophores also on abdomen and appendages, e.g., on pleura, antennular peduncles, third maxilliped, first pereiopods and tail fan; patches of white chromatophores around frontal area of carapace, on abdomen (sometimes absent or internal), third maxillipeds and tail fan; posterior abdominal somites occasionally speckled with smaller white spots (Fig. 6). Remarks. The material is typical for this species and presents no noteworthy features. Ecology. On the basis of the present records and scattered information in the literature (summarised in Hayashi 1975), N. steinii seems to frequent coral reef lagoons and reef fronts, dwelling in coral rubble and dead coral heads by day and roaming on reef walls and rubble fields by night. The depth range appears to be around 0–13 m. Distribution. This is a widely distributed species, known from Kenya, Tanzania (Zanzibar), Japan, eastern Indonesia, Australia, Palau, the Marshall Islands and Hawaii (Bruce 1993; Chace 1997). The present records of N. steinii from Ningaloo Reef and northern Great Barrier Reef complete Bruce’s (1993) previous Australian records from the Territory of Ashmore and Cartier Islands. The species is also recorded for the first time from French Polynesia and Taiwan.

Processa Leach, 1815

Processa hawaiensis (Dana, 1852) Fig. 7A

Nika hawaiensis Dana, 1852: 20. Processa hawaiensis.—Rathbun, 1906: 912.—Chace, 1962: 616.—Hayashi, 1975: 106, figs. 22–23.—Bruce, 1976: 60 (sic hawaiiensis).—Alvarez et al., 1996: 721, fig. 5.—Ayón-Parente et al., 2012: 65, figs. 1–3. Processa paucirostris Edmondson, 1930: 3, fig. 1.

Material examined. French Polynesia: 1 male (pocl 1.6), FLMNH UF 23279, Moorea, Papetoai, fringing reef with sand flat, 0–3 m, night snorkel, -17.4911 -149.886, leg. S. McPherson, M. Wright, C. Meyer & A. Anker, 06.12.2009 [BMOO-10183, BIZ-243]; 1 male (pocl 1.7), FLMNH UF 24375, Moorea, north of Motu Ahe, sandflat with live coral heads, 1–3 m, hand collecting, -17.5496 -149.776, leg. C. Meyer, K. White, S. El-Tourky & J.

NEW RECORDS OF PROCESSIDAE Zootaxa 3640 (2) © 2013 Magnolia Press · 235 Thomas, 09.12.2009 [BMOO-10327, BIZ-262]; 1 male (pocl 1.4), FLMNH UF 23280, Moorea, Atiha Bay, inside barrier reef, under and on rocks and rubble, 0.5–1 m, hand collecting, -17.5957 -149.8456, leg. S. McPherson, G. Paulay & C. Meyer, 08.12.2009 [BMOO-10096, BIZ-257].

FIGURE 6. Nikoides steinii (Edmondson, 1935). A, ov. female, dorsal view, Moorea (FLMNH UF 23278); B, same specimen, lateral view; C, ov. female, lateral view, Lizard island (FLMNH UF 17488); D, female, dorsal view, Heron Island (FLMNH UF 24679).

236 · Zootaxa 3640 (2) © 2013 Magnolia Press GRAVE & ANKER FIGURE 7. Processa spp. A, Processa hawaiensis (Dana, 1852), male, dorsal view, Moorea (FLMNH UF 23279); B, Processa sulcata Hayashi, 1975, female, lateral view, Heron Island (FLMNH UF 25889).

Colour pattern. Semitransparent with small white spots on posterior half of abdomen; red chromatophores on frontal margin of carapace near rostral base; small patch of white chromatophores on each eyestalk; walking legs mostly transparent (Fig. 7A). Remarks. The French Polynesian material closely adheres to the description of Hayashi (1975), with the rudimental exopod on the third maxilliped and the short, simple rostrum being diagnostic for the species. Ecology. On the basis of the present records and scattered information in the literature (summarised in Hayashi, 1975), P. hawaiensis seems to frequent coral reefs rather than seagrass beds, usually in coral rubble. The depth range appears to be around 0–5 m. Distribution. This species is currently known from a widely disjunct distribution in the Indo-Pacific, being primarily recorded from the central and eastern Pacific: Maui and Oahu (Hawaii), Clipperton Island (Hayashi 1975), Nayarit and Jalisco (Mexico) (Alvarez et al. 1996; Ayón-Parente et al. 2012) and now also French Polynesia (present study). However, the species is also known from the western Indian Ocean, e.g., Kenya and Zanzibar (Hayashi 1975; Bruce 1976).

Processa sulcata Hayashi, 1975: 134, fig. 34.—Noël, 1986: 291, fig. 19.—Chace, 1997: 39.—Kim et al., 2011: 265, fig. 2.

Material examined. Australia: 1 female (pocl 2.4), FLMNH UF 25889, Great Barrier Reef, Heron Island, - 23.4284 151.9546, leg. J. Reimer, 25.11.2006 [AUST-6507, HI09-104]. Colour pattern. Semitransparent with patches of white chromatophores (some on internal organs), especially dense on and near tail fan, and patches of red chromatophores, mainly on carapace, third maxilliped, first pereiopods, bases of fifth pereiopods and pleopods, and tail fan (Fig. 7B). Remarks. The single individual from Heron Island presents no special morphological features, fitting the diagnosis of P. sulcata in Hayashi (1975). Ecology. The ecology of P. s ul c at a is poorly known, with the present record not alleviating this situation. Most of the literature records are from dredged material, with no information on bottom characteristics. Hayashi (1975) states that P. s u lc a ta was collected from “seaweed and sessile marine invertebrates, which get entangled in the gill nets used for fishing spiny lobsters”, and also indicates it to be present in seagrass beds. However, the species does appear to have a considerable depth range, 0–123 m (Chace, 1997). Distribution. The species is widespread in the Indo-Pacific, with records from South Africa, Arabian Sea, Java, Vietnam, Japan (Hayashi 1975), Madagascar (Noël 1986), Hong Kong (Bruce 1990), Queensland (Davie 2002), the Persian Gulf (De Grave 2007; De Grave & Ashelby 2011) and Korea (Kim et al. 2011). The inclusion of “South America” in the distribution for P. s u lc a ta by Chace (1997) is considered a lapsus for South Africa.

Processa zostericola Hayashi, 1975

Processa zostericola Hayashi, 1975: 137, fig. 35.– Ledoyer, 1984: 43, fig. 21.—Kim et al., 2011: 266, fig. 3.

Material examined. Singapore: 1 ov. female (pocl 4.3), ZRC 2012.0977, Pulau Busing, fouling community, leg. J.B. Sigurdsson, 01.05.1987; 1 female (pocl 4.2), ZRC 2012.0978, Sultan Shoal, leg. R. Teo & T.M. Sin, 26.02.1994. Colour pattern. Unknown. Remarks. Johnson (1962) recorded the closely related P. australiensis Baker, 1907 from Halophyllum pools at Changi Beach, but the present material is assigned to P. zostericola on account of the stylocerite having no lateral tooth, an antennal spine being present and the presence of a blunt process on the basicerite (see Hayashi 1975). It is currently not clear if both species occur in Singapore or whether the material identified as P. australiensis in Johnson (1962), which is now in very poor condition, refers to the present species. Ecology. The species typically lives in littoral seagrass beds (Hayashi 1975; Ledoyer 1984; Unsworth et al. 2007, 2010), the habitat from which the present specimens were obtained was not detailed. Distribution. This species was previously only known with certainty from Japan (Hayashi 1975), Noumea (Ledoyer 1984), southern Sulawesi (Unsworth et al. 2007, 2010) and Korea (Kim et al. 2011) and is now newly recorded from Singapore.

Discussion

The present records amply demonstrate that the distribution of most Processidae in the Indo-Pacific is not overly well documented, as a total of 21 new country/area records are included in the present contribution. The emerging picture is though that most species are widely distributed across the entire Indo-West Pacific. However, some species (e.g., P. z os te r ic o la ) remain known from only a few locations and the disjunct distribution of P. hawaiensis continues to draw attention. Doubtless, with further fieldwork in intermediate areas and the continued identification of further museum material this situation may change. Regarding the ecology of Indo-Pacific Processidae, it is becoming clear that two different habitats are involved. Some species, e.g. Hayashidonus japonicus and Nikoides danae frequent seagrass habitats, where they

238 · Zootaxa 3640 (2) © 2013 Magnolia Press GRAVE & ANKER may represent dominant forms of the decapod fauna. Others, e.g. Nikoides maldivensis and N. multispinatus, appear to frequent reef environments, where they are usually associated with rubble or dead corals. However, the recorded depth range of some species confounds this distinction, as for instance H. japonicus is known to occur as deep as 150 m (Noël 1986), far beyond the lower depth limit of marine angiosperms. This can only be clarified by further, targeted sampling. Although specimens of both eastern Pacific species of Ambidexter are routinely encountered in seagrass beds or intertidal beaches, it seems mosty likely that they dwell in or near burrows of larger burrowing decapods by day and only emerge and roam at night. The specimen-matched colour photographs included in the present report do strongly suggest that colour patterns, as with numerous other carideans, are to some extent species specific and can aid in species identification. However, it must be acknowledged that further work is required on this aspect, as diurnal vs nocturnal differences in the extent and variability of chromatophores have not been well documented (see also De Grave & Felder 2012).

Acknowledgements

The present material was collected during several marine surveys, including Biotas 2008 (PIs: Henrich Bruggeman, Université de la Réunion, and Gustav Paulay, FLMNH), Biocode Moorea 2008–2010 (PI for marine invertebrates: G. Paulay) and CReefs Australia 2009–2010 (PI: Julian Caley, logistics: Shawn Smith, AIMS). Fieldwork in Taiwan was facilitated and sponsored by Tin-Yam Chan (National Taiwan Ocean University, Keelung). The specimen from Christmas Island was collected by P.K.L. Ng et al. during an expedition there by the Raffles Museum of Biodiversity Research, National University of Singapore (NUS), operating under permit PS2009/04 from the Australian Department of the Environment, Water, Heritage and the Arts; and Australian National Parks. The Johor Strait marine biodiversity workshop held on Pulau Ubin, Singapore was organised by the National Parks Board and National University of Singapore. The workshop, as part of the Comprehensive Marine Biodiversity Survey (CMBS), was supported by general contributions from Shell Eastern Petroleum, Hong Kong and Shanghai Banking Cooperation 'Care for Nature' Trust Fund, and Asia-Pacific Breweries.

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