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New records of (: Brachyura) from the New Zealand region, including a new of Rochinia A. Milne-Edwards, 1875 (Majidae), and a revision of the Web...

Article in Zootaxa · May 2009 DOI: 10.11646/zootaxa.2111.1.1

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ZOOTAXA

2111

New records of crabs (Decapoda: Brachyura) from the New Zealand region, including a new species of Rochinia A. Milne-Edwards, 1875 (Majidae), and a revision of the genus Dromia Weber, 1795 ()

COLIN L. McLAY

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Colin L. Mclay New records of crabs (Decapoda: Brachyura) from the New Zealand region, including a new species of Rochinia A. Milne-Edwards, 1875 (Majidae), and a revision of the genus Dromia Weber, 1795 (Dromiidae) (Zootaxa 2111) 66 pp.; 30 cm. 18 May 2009 ISBN 978-1-86977-369-4 (paperback) ISBN 978-1-86977-370-0 (Online edition)

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Zootaxa 2111: 1–66 (2009) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2009 · Magnolia Press ISSN 1175-5334 (online edition)

New records of crabs (Decapoda: Brachyura) from the New Zealand region, including a new species of Rochinia A. Milne-Edwards, 1875 (Majidae), and a revision of the genus Dromia Weber, 1795 (Dromiidae)

COLIN L. McLAY School of Biological Sciences, Canterbury University, Christchurch 8004, New Zealand. E-mail: [email protected]

Table of contents

Abstract ...... 4 Introduction ...... 5 Taxonomic account ...... 6 Section De Haan, 1833 ...... 6 Family Alcock, 1899 ...... 6 Dicranodromia Milne-Edwards, 1880 ...... 6 Dicranodromia spinulata Guinot, 1995 ...... 6 Dicranodromia delli Ahyong, 2008 ...... 8 Family Ortmann, 1892 ...... 11 Metadynomene McLay, 1999 ...... 11 Metadynomene tanensis (Yokoya, 1933) ...... 11 Dynomene Desmarest, 1823 ...... 12 Dynomene pilumnoides Alcock, 1900 ...... 12 Family Dromiidae de Haan, 1833 ...... 12 Dromiinae de Haan, 1833 ...... 12 Dromia Weber, 1795 sensu stricto ...... 14 Key to genera of Dromiinae dealt with herein ...... 15 Metadromia gen. nov...... 15 Metadromia wilsoni (Fulton and Grant, 1902) new combination ...... 17 Tumidodromia gen. nov...... 19 Dromia McLay, 1993 (part) ...... 19 Tumidodromia dormia (Linnaeus, 1763) new cominaion ...... 19 Section Saint Laurent, 1980...... 23 Family Samouelle, 1819 ...... 23 Tanaoa Galil, 2003 ...... 23 Tanaoa pustulosus (Wood-Mason in Wood-Mason & Alcock, 1891) ...... 25 Family Macleay, 1838 ...... 26 Dorhynchus Wyville Thomson, 1873 ...... 26 Dorhynchus ramusculus (Baker, 1906) ...... 26 Family Inachoididae Dana, 1851 ...... 29 Pyromaia Stimpson, 1871 ...... 29 Pyromaia tuberculata (Lockington, 1877) ...... 29 Family Epialtidae MacLeay, 1838 ...... 30 Pisinae Dana, 1851...... 30 Rochinia A. Milne-Edwards, 1875 ...... 30 Rochinia ahyongi sp. nov...... 30 Family Palicidae Bouvier, 1898 ...... 33

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Pseudopalicus Moosa & Sèrene, 1981 ...... 33 Pseudopalicus undulatus Castro, 2000 ...... 33 Family Atelecyclidae Ortmann, 1893 ...... 34 Trichopeltarion A. Milne Edwards, 1880 ...... 34 Trichopeltarion janetae Ahyong, 2008 ...... 34 Family Calappidae de Haan, 1833 ...... 36 Mursia Desmarest, 1823 ...... 36 Mursia microspina Davie & Short, 1989 ...... 36 Mursia australiensis Campbell, 1971 ...... 40 Family Portunidae Rafinesque, 1815 ...... 41 Ovalipes Rathbun, 1898 ...... 41 Ovalipes elongatus Stephenson & Rees, 1968 ...... 41 Liocarcinus Stimpson, 1871 ...... 43 Liocarcinus corrugatus (Pennant, 1777) ...... 43 Thalamita Latreille, 1829 ...... 46 Thalamita danae Stimpson, 1858 ...... 46 Nectocarcinus A. Milne-Edwards, 1860 ...... 46 Family Plagusiidae Dana, 1851 ...... 49 Percnon Gistel, 1848 ...... 49 Percnon planissimum (Herbst, 1804) ...... 49 Family Ocypodidae Rafinesque, 1815 ...... 53 Ocypode Weber, 1795 ...... 53 Ocypode pallidula Jacquinot, in Hombron & Jacquinot, 1846 ...... 53 Acknowledgements ...... 57 References...... 57

Abstract

Seven species and two families previously unknown from New Zealand are recorded: Dynomene pilumnoides Alcock, 1900, Metadynomene tanensis (Yokoya, 1933) (Dynomenidae new to New Zealand); Pseudopalicus undulatus Castro, 2000 (Palicidae); Mursia microspina Davie & Short, 1989 (Calappidae new to New Zealand); Ovalipes elongatus Stephenson & Rees, 1968, Thalamita danae Stimpson, 1858, (Portunidae); Percnon planissimum (Herbst, 1804) (Plagusiidae); Ontogenetic morphological changes in Nectocarcinus antarcticus (Portunidae) are described. The southern limit for Pseudopalicus undulatus is extended from Fiji to New Zealand and the depth limit from 410 m to more than 560 m. A checklist of Brachyura recorded from the Kermadec Is is also updated. The genus Dromia Weber, 1795 (Dromiidae) is revised and restricted to the Atlantic Ocean (6 species) and Metadromia gen. nov. and Tumidodromia gen. nov., are erected for Indo-West Pacific species previously included in Dromia sensu lato. A new species of Rochinia (Epialtidae) is described from the Rumble V seamount. The stridulatory ridges on the chelipeds of Ocypode (Ocypodidae), important for identification, are shown to be sexually dimorphic: males have more striae than females. A single specimen of O. pallidula is reported from the Leigh Marine Reserve where it may have been accidentally released.

Keywords: Crustacea, Brachyura, Atelecyclidae, Calappidae, Dromiidae revision, Dynomenidae, Epialtidae, Grapsidae, Homolodromiidae, Inachidae, Inachoididae, Leucosiidae, Ocypodidae, Palicidae, Plagusiidae, Portunidae, New Zealand, Kermadec Is, Norfolk Is, Lord Howe Is

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Introduction

Additional research in the New Zealand EEZ (Exclusive Economic Zone) has seen knowledge about the number of species of marine Crustacea increase during recent years. New species have been discovered, especially from the Kermadec area northeast of New Zealand (Webber & Takeda 2005; Takeda & Webber 2006; Komatsu & Takeda 2007; McLay 2007; Ng & McLay 2007; Schnabel 2009) and from seamounts and chemosynthetic habitats on the eastern side of New Zealand (Schnabel & Bruce 2006; Ahyong 2008). The new material reported here comes from the NIWA collection at Greta Point, including additional material from seamounts and chemosynthetic habitats, and from the Canterbury Museum. The latter collection includes many specimens referred to by Chilton (1911) and also unidentified material collected subsequently. Much of this material is probably from the collections of E.W. Bennett and Professor Charles Chilton, but was not actually collected by them. Landmark papers on the New Zealand marine Crustacea were published by Chilton & Bennett (1929) and by Bennett (1964). The latter paper was in fact written prior to 1937, but brought up to date before publication in 1964. Besides the Canterbury Museum, Bennett’s sources of material included the Otago Museum, Dunedin, Dominion Museum (now Te Papa), Wellington, and the Memorial Museum, Auckland. It was not standard practice at that time to publish exact records of collection and station details, so it is not always possible to link specimens with registration numbers, added later, unless there was information on the accompanying label. Some records reported here came to light during identification of specimens from the Port Surveys, undertaken by NIWA on behalf of Biosecurity New Zealand (McLay 2004; Inglis et al. 2006). Other records are based on specimens held by Te Papa, Wellington and Museum National d’Histoire Naturelle, Paris. New material collected from Raoul I. by DOC volunteers, is also reported. The 20 species from 13 families dealt with herein are summarized in Table 1. One new species of Rochinia A. Milne Edwards, 1875 is described herein from the Rumble Seamounts near the Kermadec Is. This genus was already represented by R. riversandersoni (Alcock, 1895) specimens collected in the same vicinity. The other new taxa dealt with here are two new genera that allow a more natural arrangement of some of the -carrying dromiid species previously placed in Dromia Weber, 1795 sensu lato. New collections of the species previously known as “Dromia” wilsoni (Fulton & Grant, 1902) provided the opportunity to revise this genus and place it in Metadromia gen. nov. Consequently the other species of “Dromia,” D. dormia (Linnaeus, 1763), is transferred to a new monotypic genus Tumidodromia gen. nov. This restricts Dromia sensu stricto to the Atlantic Ocean and the Mediterranean Seas. Takeda & Webber (2006) made an important contribution to our knowledge of the fauna of the Kermadec Is (especially Raoul I.) north of New Zealand, listing 52 species from 17 families. This is a considerable increase from the 25 species reported by Chilton (1911). A re-examination of Chilton’s material, held by the Canterbury Museum, shows that Chilton made several identification errors so that their inclusion in the Kermadec checklist by Takeda & Webber (2006) needs to be revised. Measurements of carapace width (CW) and carapace length (CL) are given in mm; CL followed by CW in the Material Examined. The abbreviations P1–P5 are used for the chelipeds and four walking legs, respectively, and G1 and G2 for the first and second male gonopods respectively. Records beginning with “AQ” are from the Canterbury Museum, Christchurch. The following abbreviations are used for records resulting from the NIWA Port Surveys, listed here from north to south: “WRE–CB” (Whangarei Harbour and Marsden Point), “AKL–CB” (Waitemata Harbour), “TRG–CB” (Tauranga Harbour), “NPL–CB” & “TK–CB” (New Plymouth Harbour), “WT–CB” & “WLG–CB” (Wellington Harbour), “NSN–CB” (Nelson Harbour), “LYT–CB” (Lyttelton Harbour), and “DUD–CB) (Dunedin Harbour & Port Chalmers). “DOC” refers to the Department of Conservation; “NIWA” refers to the National Institute of Water and Atmosphere and “NZOI” refers to the New Zealand Oceanographic Institute, a precursor of NIWA.

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Taxonomic account

Section Dromiacea De Haan, 1833

Family Homolodromiidae Alcock, 1899

Dicranodromia Milne-Edwards, 1880

Dicranodromia spinulata Guinot, 1995 (Fig. 1a)

Dicranodromia spinulata Guinot, 1995; 225, figs 21 a–c, 22 A–B, 25 D. — Ahyong 2008: 11, fig.1C.

Material Examined. Diabolical Seamount (Graveyard Complex), Chatham Rise: NIWA stn Z10697, 1 female, 10.9 mm x 14.1 mm, 42°47.57’S, 179°58.86’W, 950–900 m, Tangaroa, stn TAN0104/47, 16 Apr. 2001 (NIWA 42394). Zombie Seamount (Graveyard Complex), Chatham Rise: NIWA, stn Z10727, 1 male, 6.9 mm x 9.3 mm, 42°46.07’S, 179°55.31’W, 955–890 m, Tangaroa, stn TAN0104/336, 20 Apr. 2001 (NIWA 42395); NIWA stn Z10728, 1 male, 6.5 mm x 8.9 mm, 42°46.00’S, 179°55.36’W, 970–900 m, Tangaroa, stn TAN0104/337, 20 Apr. 2001 (NIWA 42396); NIWA stn Z10793, 1 damaged female, 6.7 mm x 7.8 mm, 42°45.81’S, 179°59.25’W, 927–886 m, Tangaroa, stn TAN0104/395, 21 Apr 2001 (NIWA 42397); NIWA stn Z10743, 1 female, 7.5 mm x 10.3 mm, 1 ovigerous female, 8.8 mm x 11.9 mm, 1 male, 5.8 mm x 8.9 mm, 42°43.20’S, 179°57.63’W, 1012–890 m, Tangaroa, stn TAN0104/399, 21 Apr. 2001 (NIWA 42398). East of North Cape: NIWA stn Z11063, 1 male, 5.6 mm x 7.4 mm, 34°09.85’S, 173°57.84’E, 820–805 m, Kaharoa, stn KAH0204/40, 18 Apr. 2002 (NIWA 42399); NIWA stn Z11066, 2 males 5.5 mm x 7.3 mm, 9.2 mm x 11.5 mm, 34°02.55’S, 174°49.02’E, 830–792 m, Kaharoa, stn KAH0204/47, 19 Apr. 2002 (NIWA 48725). Remarks. The two genera in the family, Dicranodromia and Homolodromia A. Milne Edwards, 1880, are represented in New Zealand. Guinot (1995) revised the family. Dicranodromia spinulata Guinot, 1995 was first reported from New Caledonia at depths of 535–680 m. Subsequently it was taken from samples east of New Zealand, with the first report by Ahyong (2008) of one of the specimens from the Rock Garden cold seep on the Hikurangi Plateau examined here. Most of the specimens had in fact already been sent to the present author by Steve O’Shea for identification and so the remainder of the collection is reported herein. Most of the material comes from the Chatham Rise, which extends the distribution of the species further south to 42°47.57’S and increases the depth range from 535–747 m to 535–1012 m. The maximum size for the New Zealand specimens is 9.2 mm x 11.5 mm for males and 10.9 mm x 14.1 mm for females. These are considerably larger than specimens from New Caledonia. An ovigerous female, 8.8 mm x 11.9 mm, from NZOI, stn Z10743, near the maximum depth carried approximately 20 eggs 2.0 mm in diameter, much larger than reported for a female from New Caledonia (CL 5.5 mm) carrying oval-shaped eggs 1.2 mm x 1.3 mm diameter (Guinot 1995). Guinot did not give the number of eggs but it may well be that, given the relationship between egg size and egg number, the brood size was greater than for the Chatham Rise female. A female D. nagai Guinot, 1995 from Japan was recorded with less than 20 eggs (exact size unknown) under its abdomen, some of which had hatched into megalopae, thereby showing that this species has direct development. It may well be that D. spinulata also has direct development in the southern part of its range. Distribution Dicranodromia spinulata is known from New Caledonia, and in New Zealand waters, east of North Cape and the Chatham Rise. Depth range 535–1012 m.

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TABLE 1. Summary of the New records and additional information on the Brachyura of New Zealand and neighbouring regions presented herein. Family/Species New Information Homolodromiidae Dicranodromia spinulata Guinot, 1995 Range extension Dicranodromia delli Ahyong, 2008 Range extension in NZ waters; male characters Dynomenidae New family record Dynomene pilumnoides Alcock, 1900 New record Metadynomene tanensis (Yokoya, 1933) New record Dromiidae Metadromia wilsoni (Fulton & Grant, 1902) New genus Tumidodromia dormia (Linnaeus, 1763) New genus (not recorded from New Zealand) Leucosiidae Tanaoa pustulosus (Wood-Mason in Wood-Mason & Alcock, Additional specimens collected 1891) Inachidae Dorhynchus ramusculus (Baker, 1906) Range extension Inachoididae Pyromaia tuberculata (Lockington, 1877) Range extension Epialtidae Rochinia ahyongi sp. nov. New species Palicidae Pseudopalicus undulatus Castro, 2000 New record Atelecyclidae Trichopeltarion janetae Ahyong, 2008 Range extension Calappidae New family record Mursia australiensis Campbell, 1971 New record (Lord Howe Rise) Mursia microspina Davie & Short, 1989 New record; female characters Portunidae Ovalipes elongatus Stephenson & Rees, 1968 New record Thalamita danae Stimpson, 1858 New record (Kermadec Is) Liocarcinus corrugatus (Pennant, 1777) Additional records (Kermadec Is, mainland New Zealand) Nectocarcinus antarcticus (Hombron & Jacquinot, 1846) Ontogenetic morphological changes described Plagusiidae Percnon planissimum (Herbst, 1804) Additional specimens (Kermadec Is); new record (mainland New Zealand) Ocypodidae Ocypode pallidula Jacquinot, in Hombron & Jacquinot, 1846 Additional specimens (Kermadec Is, Norfolk Is); probable accidental introduction (Leigh Marine Reserve)

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FIGURE 1. a. Dicranodromia delli Ahyong, 2008, male 9.0 x 13.0 mm (stn KAH9901/53) (left). b. Dicranodromia spinulata Guinot, 1995, male 9.2 mm x 11.5 mm (stn KAH0204/47) (right).

Dicranodromia delli Ahyong, 2008 (Fig. 1b)

Dicranodromia delli Ahyong, 2008: 7, figs 2–4.

Material Examined. East of Coromandel Peninsula: NIWA stn Z9840, 1 male, 9.0 mm x 13.0 mm, 36°30.27’S, 176°30.45’E, 990–1100 m, Kaharoa, stn KAH9901/53, 05 June 1999 (NIWA 48726). Gothic Seamount (Graveyard Complex), Chatham Rise: NIWA stn Z19714, 1 male, 6.5 mm x 9.2 mm, 42°43.95’S, 179°53.91’W, 1076–990 m, Tangaroa, stn TAN0104/153, 18 Apr. 2001 (NIWA 48727). Description of male. P1 similar to female (see Ahyong 2008, p7), sparse conical granules, spinules especially dorsally, ventrally; fingers slightly shorter than dorsal margin of propodus, occlusal margins armed with 5 or 6 almost obsolete teeth, only distal 2 teeth demarcated, inter-locked; fingers with narrow gape proximally when closed. P3 longest (2.2CL); P2, P3 dactyli with 12–15 well developed corneous spines along inner margins. P4 dactylus short curved opposed by 3 longer corneous spines with 1or 2 spines on outer propodal margin, inner margin of dactylus armed with 3 or 4 small spines. P5 dactylus, short, curved, opposed by 3 corneous spines with 4 spines on outer propodal margin, small corneous spine on outer margin of dactylus. Gonopods as for all species of the Homolodromiidae (Guinot 1995: 177, figs.4A–C): G1 stout, straight, spiralling to densely setose apex containing aperture into which needle-like G2 fits. G2 longer than G1. Telson

8 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. triangular, posterior margin sub-acute, 1.4 as long as wide, fused with somites 5 and, 6] so that no flexibility allowed, but all sutures evident. No abdominal locking mechanism, but abdomen is folded ventrally at joint of somites 4 and 5 where limited flexibility eliminates need for abdominal retention against sternum. No vestigial pleopods on somites 3 – 5. Uropods ventral, reduced (length 0.12 telson width), oblong, calcified, rigid. Remarks. Only a female (15.5 mm x 19.0 mm) from the Nukuhou Seamount, Bay of Plenty, was available when Ahyong (2008) described D. delli. Two males are reported here from off the Coromandel Peninsula and Chatham Rise, both much smaller than the type. The known depth range is now 990–1134 m, slightly deeper than D. spinulata. Guinot (1995) provided a key to the identification of the 16 Atlantic and Indo-West Pacific species of Dicranodromia then known. Ng & McLay (2005) described a new species, D. danielae Ng & McLay, 2005, from the Philippines and the new species, D. delli, make a total of 18 species. The two New Zealand species, D. delli from D. spinulata, can be distinguished from each other by the density of spinules on the carapace, but the best way to separate them is by comparing the proportions of the P2 and P3 articles (Fig. 1a, b): for female D. delli merus length is around six times the merus height and propodus length is around 10 times its height (eight and nine times respectively in males) and twice the length of the dactylus (2.2 times and 2.4 times respectively in males). In D. spinulata the P2 and P3 meri length is around 5 times height, propodus length less than 7 times height and propodi are only 1.6 times dactylus length. In D. delli the P2 and P3 are therefore longer and more slender. The reduced camouflage-carrying P4 and P5 are also longer than in D. spinulata. An additional difference between the two species is that the inner margins of the P2 and P3 dactyli of D. delli are armed with 12–15 corneous spines whereas in D. spinulata there are only seven or eight. Ahyong (2008) also reported Homolodromia kai Guinot, 1993 from the Bay of Plenty at a depth of 632–458 m. This species is easily distinguished from the two species of Dicranodromia by a longer pair of walking legs than Dicranodromia and by the last pair of prehensile legs having a distal prolongation on the propodus, which is similar to the fixed finger of the chelipeds. There are several secondary sexual characters between male and female D. delli as follows: suture between last abdominal somite and telson is faint or absent in females (clearly evident in males); P3 of females is 1.9CL (2.2CL in males); P4 in females has two corneous spines on outer margin and three or four opposing spines (males have one or two corneous spines on outer margin and 3 opposing); no corneous spines on the inner margin of dactylus were reported by Ahyong (2008) (inner margin of male dactylus is armed with 3 or 4 inclined small spines). The female P5 has two corneous spines on the outer margin and 7 spines opposing the dactylus (Ahyong 2008) (male has a spine on the outer propodal margin and three opposing spines); female has 3 or 4 obliquely inclined spines on the inner margin of the dactylus (male has a spine on the outer propodal margin and 3 spines opposing); no spines on outer margin of the P5 dactylus (one small spine in male).

TABLE 2. Arrangement of corneous spines on the distal margins of the P4 and P5 propodi and dactyli of Dicranodromia delli and D. spinulata. Number of Number of spines Number of spines Number of spines Number of spines Species spines opposing on Lateral Propodal on Outer Propodal on Inner Margin of on Outer Margin of P4 (P5) dactylus Margin of P4 (P5) Margin of P4 (P5) Dactyl of P4 (P5) Dactyl of P4 (P5) D. delli 3, 4 (4-7?) 0 (0) 1, 2 (4) 3, 4 (0) 0 (0, 1) D. spinulata 2, 3 (4) 0 (1, 2) 1 (2–4) 6 (0–6) 0 (0, 1)

The number and arrangement of spines on the P4 and P5 propodi and dactyli for D. delli and D. spinulata is summarized in Table 2. There is considerable variation in these spines both within and between species. Some of this variation is caused by spines being broken off so it is probably best to take the larger number for each category as being representative of the species. Guinot (1995) provides detailed information for several species and the present two species fall within this range. Homolodromiids have comparatively large numbers

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FIGURE 2. Metadynomene tanensis (Yokoya, 1933), ovigerous female 19.2 mm x 18.0 mm (stn F867): a. dorsal view, b. frontal view.

10 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. of spines on their reduced last two pairs of pereopods and these are used for hold pieces of camouflage that are carried above the crab. Such large numbers of corneous spines are also found in the primitive Dromiidae such as in the species of Sphaerodromia Alcock, 1899. Large numbers of corneous spines on camouflage-carrying pereopods is a plesiomorphic character state. The two homolodromiid genera differ in Homolodromia the propodal spines are on an extension of the propodus with resulting longer dactyli, whereas in Dicranodromia the ventral distal margin of the propodus is not extended so that the dactylus is shorter and more strongly curved. Species of both genera can have spines on the inner margin of the dactyli as well as distal propodal spines surrounding the base of the dactyli. In several species of both Dicranodromia and Homolodromia, males have vestigial pleopods on somites 3, 5, in addition to the gonopods of somites 1, 2 (Guinot, 1995, 179). D. delli males do not have any vestigial pleopods other than the uropods and in this respect they are the same as the D. spinulata males listed above. No evidence of the “rudiments” mentioned in the description of D. spinulata by Guinot (1995, 225) could not be found in the New Zealand specimens. The uropod length in male D. spinulata (0.13 telson width) is also similar to those in male D. delli. Distribution. Dicranodromia delli is a New Zealand endemic originally reported from the Nukuhou Seamount, Bay of Plenty, but now known from East of Coromandel and the Gothic Seamount (Graveyard Complex), Chatham Rise Peninsula. Depth range is 990–1134m.

Family Dynomenidae Ortmann, 1892

Metadynomene McLay, 1999

Metadynomene tanensis (Yokoya, 1933) (Fig. 2a, b)

Dynomene tanensis Yokoya, 1933: 96, fig. 38. Dynomene praedator — Sakai 1976: 30, fig. 17. – Naga & Tsuchida 1995: 108, pl. 1, fig. 2. Not Dynomene praedator A. Milne-Edwards, 1879. Metadynomene tanensis — McLay 1999: 521, figs 4d, 6c, 7f, 9d, e, 11, 13c, e, f, 14e, 25b, 27, 28. — McLay et al. 2001: 964. — Ng et al. 2001: 6. — Guinot & Quenette 2005: 284, fig. 4A, B. — McLay & Ng 2005: 24.

Material Examined. East Cape: 1 ovigerous female, 19.2 mm x 18.0 mm, 37°30’S, 179°00E, 307–340 m, Taranui, F867, TAM, 2 Oct. 1968. (NIWA 48580) Remarks. The discovery of M. tanensis is the first record of the Dynomenidae in New Zealand. Dynomene pilumnoides is also being reported from New Zealand for the first time (see below). The systematics of this family has been reviewed by McLay (1999). The specimen was discovered in the NIWA collection, where it had been in storage for more than 30 years. The type species of Metadynomene McLay, 1999 is Dynomene tanensis Yokoya, 1933, collected off the eastern coast of Japan. The geographical distribution of this species includes Taiwan, Indonesia, New Caledonia, Loyalty Is, Chesterfield Is and Vanuatu, to which we can now add New Zealand. A large male specimen reported as “M. tanensis” by McLay (1999: 522) from the Tuamotu Archipelago, , probably belongs to an undescribed species. There are two other species: M. crosnieri McLay, 1999, from the Glorieuse Is, Indian Ocean and M. devaneyi (Takeda, 1977) from the Hawaiian Is and the Marquesas Is, French Polynesia. Species of this genus inhabit for the most part water between 200 and 500 m and some are associated with precious corals (Corallium sp.). They seem to live on rocky bottoms. Coarse sand, broken shell, rocks, Glycymeris, Venericardia, Pecten (Bivalvia), Munida(), pagurids, electric ray, and ophiuroids were also collected along with the New Zealand specimen of M. tanensis. There is a superficial resemblance between this species and Metadromia wilsoni nov. comb. because both species have a dense, short, uneven tomentum covering the carapace, but they are easily distinguished because

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M. tanensis only has the last pair of legs (P5) reduced, whereas M. wilsoni has the last two pairs (P4 and P5) reduced, although not as much as for M. tanensis P5. Distribution Metadynomene tanensis is known from Taiwan, Indonesia, New Caledonia, Loyalty Is, Chesterfield Is and Vanuatu, and now from East Cape, New Zealand. Depth range is 205–520m.

Dynomene Desmarest, 1823

Dynomene pilumnoides Alcock, 1900 (Figs. 3a, b)

Dynomene pilumnoides Alcock, 1900:133; l90l: 35, pl. 1, fig. 2. — Barnard 1947: 371; 1950: 337, f ig.65 a–c. — Sakai 1965: 12, pl. 6, fig. 2; 1976: 29, pl. 6, fig. 3. — Serène & Vadon l98l:121. — Miyake 1983: 11, pl.4, fig. 2. — Baba 1986: 310, fig. 163. — Garth et al. 1987: 241. — Nagai 1989: 43. – McLay 1999: 494, figs 3c, d, 8d, e, 11, 12e, f, 14c, 17d, 21a–g. — McLay & Ng 2005: 20. — Guinot & Quenette 2005: 282, fig. 4C, D Maxillothrix actaeiformis Stebbing, 1921: 457, pl. 14 (Crust. Pl. 109).

Material Examined. Monowai Seamount: 1 female, 10.0 mm x 8.4 mm, 27°12.51'S 177°26.02'E, rock dredge, Tangaroa, stn TAN0411/32, 252–403m, 7 Oct. 2004. (NIWA 48574) Comparative Material. Fiji: 1 male, 13.5 mm x 11.4 mm, 18°18.4' S, 178°02.5' E, 233–248 m, MUSORSTOM 10, DW1377, 17 Aug. 1998. Remarks. Dynomene pilumnoides is the second dynomenid species discovered in New Zealand. D. pilumnoides can be separated from the other dynomenid species, Metadynomene tanensis, by virtue of its 4 sub-acute anterolateral teeth (versus 3 blunt teeth) and the short plumose setae, interspersed with clumps of longer filiform setae, on its carapace (versus short soft setae all about the same length). Species of Dynomene are for the most part found in shallow water, occurring from the intertidal to 100 m. D. pilumnoides has the greatest depth range, 18–400 m, and it has been collected from rocky bottoms covered in Lithothamnion (Rhodophyta), and corals. The specimen of D. pilumnoides was taken with a rock dredge that also caught Cheiraster sp. (Asteroides), Bathymodiolus sp. and Branchipolynoe sp. (Polychaeta) in other nearby samples. The Monowai Seamount is part of the Kermadec volcanic system. The specimen (Fig. 3b) collected during the MUSORSTOM 10 expedition is a new record for Fiji. The seven species of Dynomene can be identified using the key provided by McLay (2001: 810). Geographical Distribution D. pilumnoides has a wide Indo-West Pacific distribution that includes Madagascar, Réunion, , Indonesia, Philippines, Japan, Hawaiian Is, New Caledonia, and now Fiji and the Monowai Seamount, New Zealand. Depth range is 18–403m although most records come from 100–300m.

Family Dromiidae de Haan, 1833

Dromiinae de Haan, 1833

Dromia Weber, 1795 is the type genus of this subfamily created by Guinot and Tavares (2003). In his revision of the genus McLay (1993) synonymised Sternodromia Forest, 1974 with Dromia and included ten species from the Atlantic Ocean, Mediterranean Sea and Pacific Ocean. Guinot & Tavares (2003) resurrected Sternodromia for S. spinirostris (Miers, 1881) and transferred Dromia monodi Forest & Guinot, 1966 to the same genus. Dromia foresti McLay, 1993 should be transferred to Stebbingdromia Guinot & Tavares, 2003. On the basis of larval and adult morphological evidence, “Dromia” wilsoni (Fulton & Grant, 1902) is herein transferred to Metadromia gen. nov., “Dromia” dormia (Linnaeus, 1763) to Tumidodromia gen. nov., and

12 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. restrict Sternodromia to its type species (see below). This leaves six species, which at least in adult morphological terms, form a much more natural group of Atlantic species than when the other five species were included. The species of Dromia are: D. bollorei Forest, 1974, D. erythropus (George-Edwards, 1771), D. marmorea Forest, 1974, D. monodi Forest & Guinot, 1966, D. nodosa A. Milne-Edwards & Bouvier, 1898 and D. personata (Linnaeus, 1758).

FIGURE 3. Dynomene pilumnoides Alcock, 1900: a. female 10.0 x 8.4 mm (stn TAN0411/32, New Zealand): b. male, 13.5 x 11.4 mm, MUSORSTOM 10 (DW1377, Fiji).

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The only larvae of Dromia known are those of D. personata and D. erythropus and these share four unique characters (McLay et al. 2001). Note that Dromia gouveai Melo & Campos, 1999, which was included in the list of Ng et al. (2008), is a of D. erythropus (George-Edwards, 1771) (Marcos Tavares, pers. com). All three of the Pacific species, previously included in Dromia, should therefore be transferred to two new genera, making Dromia sensu stricto an exclusively Atlantic/Mediterranean genus. The type species of the family, D. personata, occurs in both the Atlantic and Mediterranean. The only other large sponge-carrying dromiids in the Atlantic/Mediterranean/Caribbean regions are Sternodromia spinirostris, Moreiradromia antillensis (Stimpson, 1858), as well as Metadromia wilsoni, which has been recorded once from Saint Helena, South Atlantic (Forest, 1974: footnote on p. 89). The relationships between these Atlantic and Pacific dromiids should be explored using molecular characters. Definitions of the new genera are given below. As a result of the removal of some species, a revised definition (after McLay, 1993) of Dromia Weber, 1795 sensu stricto follows.

Dromia Weber, 1795 sensu stricto

Definition Carapace wider than long (1.2 – 1.4), surface sculptured. Rostrum tridentate, median tooth less than or equal in length to lateral teeth. Female sternal grooves normally end apart in pits between bases of P2. Cheliped with epipod. Sternite 4 T-shaped (inverted), episternites extend laterally between coxae of chelipeds, P2 to form cross-piece of “T”, stem of “T” formed by sternite narrowing anteriorly, tip truncate, ratio of distance across episternites to sternite length 1.0–0.9. Walking legs (P2–P4) not knobbed or ridged. Length of propodi, dactyli of first two pairs of legs (P2, P3) usually equal, inner margins of dactyli armed with 5–7 small spines. P4 dactylus opposed by single propodal spine, no spine on the outer propodal margin. Fourth leg (P5) shorter than second (P3), dactylus usually opposed by two propodal spines, with another spine on the outer propodal margin. Margin of telson rounded. Uropod plates well developed, visible externally, used in male abdominal locking mechanism by fitting in front of serrated flanges on the bases of first legs (P2). Joint between last two abdominal somites freely movable. Remarks. One of the enduring difficulties in deciding the limits of Dromia has been the importance attached to the female sternal groove character. These grooves, which mark the suture between sternite 7 and 8, are of more than passing interest because they encompass the apertures to the spermathecae and are intimately involved in copulation, sperm transfer and fertilization. They may be short so that the apertures are posterior to the female gonopores or long so that the apertures are anterior to the gonopores in the coxae of the second walking legs (P3). In addition the grooves may terminate wide apart or very close together. In the type species, , the sternal grooves are somewhat intermediate, ending apart in front of the gonopores, between the bases of the first pair of walking legs (P2). This condition is found in all the species currently included in the genus, except for D. bollorei, in which the sternal grooves end close together in pits on an elevated protuberance between bases of the first walking legs (P2). However, sternite 4 in this species is similar to that of the other Dromia species, having wing-like lateral projections between the coxae of the cheliped and first leg (P2). The only problem species is D. monodi, which has all the characters of Dromia, but whose body shape is remarkably similar to Sternodromia spinirostris. Forest (1974) commented that immature specimens of D. monodi and S. spinirostris are difficult to separate so that Guinot & Tavares (2003) transferred Dromia monodi Forest & Guinot, 1966 to Sternodromia Forest, 1974. McLay (1993) synonymised Sternodromia with Dromia because of the similarity of the modifications to the two camouflage-carrying limbs (last two pairs of legs, P4 and P5) as well as in body shape, and chose to overlook the sternal characters. This was ill-advised because we should regard the sternum as a more conservative feature than the arrangement of propodal spines on the carrying legs. The truncated sternite 4, the slit-like apertures of the spermathecae and sternal grooves terminating close together at the base of a large conical swelling between bases of first legs are sufficiently important as to warrant a separate genus. Also the abdominal locking mechanism in S. spinirostris differs from that typical of Dromia: the uropods are smaller

14 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. and the coxae, with which they engage, are only shallowly indented. The same argument applies to D. monodi and therefore this species should not be placed in Sternodromia. Given the importance of the sternal characters it seems wiser to keep D. monodi in Dromia and regard conformation of the carapace as being convergent. If we use sternal differences to justify Sternodromia as a genus, then we must use the same argument to exclude D. monodi. The following key for identifying the genera resulting from the revision of Dromia sensu lato uses the shape of the sternite 4 between the bases of the cheliped. This sternite is T-shaped with the episternites forming the top of the “T,” articulating with the cheliped coxae on each side, and the long axis of the “T” extending forward between the coxae of chelipeds.

Key to genera of Dromiinae dealt with herein

1. Width to length ratio of sternite 4 approximately 1.0...... 2 - Width to length ratio of sternite 4 much greater or much less than 1.0...... 3 2. Lateral margins of sternite 4 narrowing anteriorly, tip rounded ...Dromia sensu stricto (Atlantic and Mediterranean). - Lateral margins of sternite 4 parallel, truncated tip..... Metadromia gen. nov. (Indo-West Pacific and South Atlantic) 3. Episternites of sternite 4 extend laterally between coxae of cheliped (P1) and first walking leg (P2), sternite extends anteriorly between coxae of chelipeds, margins narrowing anteriorly then parallel for last half, truncated tip, width to length ratio of anterior extension approximately 1.3 ...... Tumidodromia gen. nov. (Indo-West Pacific) - Episternites of sternite 4 truncated, not extending laterally between coxae of cheliped (P1) and first walking leg (P2), margins of anterior extension of sternite parallel, tip rounded, ratio of width to the length of the anterior exten- sion less than 0.65 ...... Sternodromia Forest, 1974 (Atlantic and Mediterranean)

Metadromia gen. nov.

Dromia McLay, 1993: 149 (part).

Definition Carapace oval, wider than long (CW/CL > 1.2), surface smooth or sculptured, covered by fine, rippled tomentum. Rostrum tridentate. Female sternal grooves end apart between or behind cheliped bases. Prominent subhepatic tooth at or below level of suborbital margin, 3 well developed anterolateral teeth. Branchial groove well marked with small tubercle near base of posterolateral tooth. Cheliped with epipod, well developed, blunt teeth, fingers not gaping. Anterior extension of sternite 4 short, margins parallel, truncated anteriorly, ratio of width of sternite 4 to length approximately 1.0, episternites extending laterally between coxae of cheliped (P1), first walking leg (P2). Walking legs (P2–P5) knobbed, length of propodi, dactyli usually equal, inner margins of dactyli with 5–7 small spines. P4 dactylus opposed by propodal spine, no spine on the outer propodal margin (there may be some small spines on margin of inner dactylus). Fourth leg (P5) shorter than second leg (P3), dactylus opposed by a single spine, with sometimes another spine on the outer propodal margin. External surface of abdomen sculptured, margin of telson rounded. Uropod plates well developed, visible externally, used in male abdominal locking mechanism by fitting in front of serrated flange on bases of first legs (P2). All somites of abdomen freely movable in both sexes. Vestigial pleopods may be present on third-fifth somites in males. Type Species: Cryptodromia wilsoni Fulton & Grant, 1902. Included Species: Metadromia wilsoni (Fulton & Grant, 1902). Etymology. The generic name Metadromia is a combination of meta, meaning “after” or “beyond” and the genus Dromia. Gender is feminine. Distribution Metadromia is the most widespread genus of all the dromiids as its distribution includes the Indo-West Pacific and South Atlantic. The only species, M. wilsoni, occurs in cooler waters: in tropical areas it is found in greater depths whereas in temperate areas it can be found in much shallower depths, even in the intertidal.

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FIGURE 4. Metadromia wilsoni (Fulton & Grant, 1902) nov. comb., male 19.3 x 15.2 mm, CB2TRG032: a. dorsal view, b. frontal view.

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Discussion. Fulton & Grant (1902) placed their new species in Cryptodromia Stimpson, 1858, before the generic definition was revised. Borradaile (1903) made an important addition to generic characters in the Dromiidae when he stated that a cheliped epipod is absent in Cryptodromia but present in Petalomera Stimpson, 1858. Applying this new definition Rathbun (1923) changed the Australian species to Petalomera, but this was never correct because it lacked the essential defining feature of this genus, having petaloid meri on the first two pairs of pereopods. Finally in his revision of Petalomera, McLay (1993) transferred it to Dromia, but not without some misgivings. The comparison of larval characters by Lim and Ng (2001), which was based on groupings of species on the basis of adult characters by McLay (1993), showed that it was apparent that Cryptodromia wilsoni did not belong in Dromia and thus deserved a new genus.

Metadromia wilsoni (Fulton and Grant, 1902) new combination (Figs. 4a, b)

Cryptodromia wilsoni Fulton & Grant, 1902: 61, pl. 9. — Ihle l9l3: 91 (in list). Cryptodromia lateralis Heller, 1865: 71. — Miers 1876: 57. — Thomson, 1899: 170, pl.20, figs l, 2. — Chilton 1911: 49. Not Dromia lateralis Gray, 1831. Dromia pseudogibbosa Parisi, l9l5: 102, pl. 2, figs l, 2. — Balss 1922: 106. — Yokoya 1933: 97. Petalomera lateralis — Richardson 1949b: 60, fig. 51. Not Dromia lateralis Gray, 1831. Petalomera wilsoni — Rathbun 1923:154, pl. 42, fig. 1. — Hale 1927: 113, fig. 111. —Sakai 1936a: 33, pl. l, fig. l; 1936b: 34, pl. l, fig. 4, text fig. 9; 1965: 9, pl. 4, fig. 2; 1976: 24, pl. 6, fig. 1. — Barnard 1947: 368; 1950: 313, fig. 59e. — Guiler 1952: 37. — Dell 1963: 22; 1968: 14, pl. 2. — Bennett 1964: 27, fig. l4l. — Suzuki & Kurata 1967: 95 (list). — Kim 1970: 9, pl. 1, fig. 3; 1973: 608. — Wear 1979: l; 1977: 572. — Griffin 1972: 56. — Forest 1974: 89 (footnote). — Kensley 1978: 251; 1981: 37 (list). — Kim & Kim 1982: 136. — Miyake 1983: 6, pl. 3, fig. l. — Wear & Fielder 1985: 22, figs 54–57. — Baba in Baba, Hayashi & Toriyama 1986: 309, pl. 162. — McLay 1988: 68, fig. 10a–f; 1991: 470, pl. 1B, figs 6a–d, 7a–c, 8a–c. Dromia wilsoni — McLay, 1993: 156, fig. 16e; 2001: 84. — McLay, Lim & Ng 2001: 737. — Ng et al. 2000: 159, fig. 2b. — Ng et al. 2001: 5 (list). — Brösing et al. 2002: 330, figs 1, 2. — Davie 2002: 162. — Takeda & Ueshima 2002: 163 (list). — Poore 2004: 304. “Dromia” wilsoni — McLay & Ng 2005: 13. [Dromia] wilsoni — Guinot & Quenette 2005: 288, figs 10A, B.

Material Examined. Tauranga Harbour: female, 16.2 mm x 13.0 mm, 37° 39’33.5” S, 176° 10’52.25” E, 2.6 m, wharf pile, TG017CB, 5 Mar. 2002 (NIWA 48745); female, 20.1 mm x 15.3 mm, 37° 38’34.25” S, 176° 10’53.26” E, 6.7 m, wharf pile, TG105CB, 7 Mar. 2002 (NIWA 48749); female, 19.3 mm x 14.6 mm, 37° 38’ 42.11” S, 176° 10’55.21” E, 6.4 m, wharf pile, TG113CB, 7 Mar. 2002 (NIWA 48750); male, 19.3 mm x 15.2 mm, 37° 38’37.50” S, 176° 10’54.55” E, 9 m, wharf pile, 2TRG032CB, 5 Apr., 2005 (NIWA 48751). Description (modified after McLay, 1993). Carapace distinctly wider than long, moderately convex, surface smooth, gently undulating under thick cover of soft, long setae giving surface an areolate appearance. Cardiac, branchial grooves well marked by depressions, one pair of medial cardiac pits, another one more posterior. Rostrum tridentate, median tooth small, blunt, on lower level, projecting as far forwards as lateral teeth separated by a U-shaped sinus, from which extends distinct frontal groove separating 2 rounded protuberances. Three strong anterolateral teeth extend back from level of suborbital tooth: first tooth directed forwards, last 2 upwardly directed. Posterolateral tooth large, also projecting upward. Small tubercle on the ridge behind branchial groove close to base of posterolateral tooth. Posterolateral carapace margins convergent, posterior margin concave. Lateral rostral teeth continuous with supraorbital margin, with broad, blunt supraorbital tooth. External orbital corner not produced, with small fissure separating it from strong suborbital tooth visible dorsally. In dorsal part of orbit, beneath supraorbital margin, vestige of parallel ridge meeting weak vertical ridge (an extension of the supraorbital tooth) at lateral end of ridge, which tends to divide off corneal region of orbit. First article of antenna much wider than long, medially beaked, gaping, twisted. Antennal flagella length 0.43CW.

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Subhepatic area of carapace convex with small, blunt tubercle beneath suborbital tooth, larger tubercle, lower, between it, first anterolateral tooth. Female sternal grooves ending wide apart on small raised tubercles between bases of P2, P3. Chelipeds (P1) large, especially in male. Merus triangular in section, all 3 borders with small rounded granules. Carpus with 2 large distal nodules, inner angle with sharp tooth. Propodus smooth, upper border in male sparsely covered in rounded nodules, nodules rudimentary in females. Inner, outer surfaces of fingers longitudinally grooved, covered with tomentum, distal surface alone naked, glabrous. Fingers pink, hollowed out internally, armed with seven well developed teeth, gaping when closed; long, silky hairs on inner surface of propodus, fingers. Ratio of width of sternite 4 to length approximately 1.0. P2, P3 shorter than chelipeds, first slightly longer than second. Carpi, propodi with tuberculiform nodules at distal ends of anterior borders. Dactyli approximately as long as propodi, inner margins with 5–7 small spines increasing in size distally. P4, P5 much reduced, similar in size. One propodal spine opposing the P4 dactylus whose inner margin has 3 or 4 minute spines. P5 dactylus opposed by propodal spine. No spines on outer propodal margins of either leg. Abdomen with 6 free somites. Telson much wider than long, male telson trigonal (width= 1.5length), female telson subtruncate (width = 1.7length). Uropod plates well developed, visible externally. Abdominal locking mechanism involves uropods fitting in front of well developed serrated flange on bases of P2. G1 as partially rolled tube with densely setose, broadly rounded tip armed with sharp horny tubercle. G2 simple, needle-like. Vestigial pleopods on somites 3–5 in males. Remarks. Both Fulton and Grant (1902b) and Rathbun (1923a) stated that there are four anterolateral teeth, but the first tooth is clearly subhepatic in position and only three teeth are actually on the anterolateral border. Vestigial pleopods are found on the abdominal somites 3–5 in males of G. wilsoni. Another feature that may well be plesiomorphic is the presence of small spines on the inner margin of the P4 dactylus. These are also found on the P4 and P5 dactyli in the species of Sphaerodromia (McLay, 1991, Table 1), which have a number of ancestral character states. McLay (1991, 465) presented a hypothesis about the origin of the dromiid camouflage-carrying limbs from walking legs that normally have spines on their inner margins, and so according to this view the condition found in M. wilsoni is transitional between a walking limb and a limb solely adapted for carrying. M. wilsoni occupies a wide range of depths and it is the only dromiid known in southern cooler temperate waters (New Zealand). In higher latitudes it occurs intertidally, but in tropical waters it is only found in deeper water, presumably where temperatures are colder. Depth range for this species is 0–520 m. Compared to crabs of similar size from other families, the reproductive strategy of M. wilsoni is not directed at producing large numbers of larvae: two ovigerous females from the Marquesas Is (CW = 46.5mm), carried approximately 3300 eggs (0.7mm diam.), and another (CW = 44.6mm), approximately 3500 eggs (0.65mm diam.) (McLay, 1991). Wear (1970) reported slightly larger eggs ready to hatch (1.1 mm) from New Zealand. M. wilsoni has three or possibly four zoeal stages and a large megalopa (Wear, 1970, 1977; Wear and Fielder, 1985), fewer zoeal stages than found in some other dromiids, but the length of the larval period is as yet unknown. Thus the exact reasons for its extraordinarily wide distribution are not clear: maybe some combination of longer larval duration, larger size and better survival. Camouflage: small crabs (CW<15 mm) carry sponge or ascidian caps, but larger crabs often do not carry any camouflage. The very hirsute tomentum may be sufficient concealment. Dell (1968) suggested that large crabs may not camouflage and subsequent observations seem to support his hypothesis. His statement that the camouflage may be attached to the body in some sponge crabs is not correct since camouflage is always carried by the last two pairs of pereopods and periodically replaced. Maximum size is 61.0 mm x 45.4 mm for males and 49.1 mm x 34.7 mm for females. Distribution Metadromia wilsoni was first described from Port Phillip Heads, Melbourne, Australia but it has perhaps the widest distribution of any dromiid crab, including the coasts of Australia from southern Western Australia around the Great Australian Bight to New South Wales, including Tasmania. Its

18 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. southernmost limit is off the Kaikoura coast, South I., New Zealand (42°25'S, 173°41'E, McLay, 1988). In the Indian Ocean M. wilsoni is known from the east coast of South Africa and even extending to St. Helena (15°58'S, 5°43'W) in the Atlantic Ocean (Forest, 1974). In the Pacific Ocean it is known as far north as Sagami Bay, Japan, Taiwan and the Hawaiian Is, and as far west as the Tubuai Is, in French Polynesia (23°51.4'S, 147°44.5'W). Its distribution thus includes three oceans as well as both sides of equator in the Pacific. Depth range is 0–520m.

Tumidodromia gen. nov.

Dromia McLay, 1993 (part)

Definition Carapace wider than long (ratio = 1.1–1.2), surface smooth, covered by short, even velvety tomentum, not hirsute. Rostrum tridentate, median tooth longer than lateral teeth, clearly visible dorsally. Female sternal grooves converge gradually anteriorly, ending with divergent tips between bases of chelipeds, separated by a prominent smooth ridge. Chelipeds (P1) massive, inner superior margin with 4 tubercles, epipod present. Sternite 4 T-shaped (inverted), episternites extend laterally between coxae of cheliped, P2 to form cross-piece of “T”, stem of “T” formed by sternite narrowing anteriorly, tip truncate, ratio of distance across episternites to sternite length 1.9 (males) to 2.2 (females). Walking legs (P2–P5) not knobbed or ridged, dactyli much shorter than propodi, inner margins armed with 4 or 5 strong spines. Carrying legs (P4 & P5) shorter. P4 dactylus opposed by a single propodal spine, no spine on outer propodal margin. P5 slightly shorter than third, dactylus opposed by 2 spines with another on the outer propodal margin. Margin of telson rounded. Uropod plates well developed, visible externally, used in male abdominal locking mechanism by fitting in front of serrated flange on bases of first legs. Joint between last two abdominal somites freely movable. Etymology. The new generic name is derived from the Latin tumidus and the name of the genus Dromia, the type species of the family, alluding to the swollen inflated shape of the carapace. Distribution Tumidodromia has a wide spread Indo-West Pacific distribution, associated with coral reefs, inhabiting shallow depths down to around 100 m. Its sole species, T. dormia, is the largest known dromiid crab, growing to around CW = 200mm.

Tumidodromia dormia (Linnaeus, 1763) (Figs 5a, b, 6a, b)

Cancer lanosus Rumphius, 1705: 19, pl. 11, fig. 1. — Seba 1759: 42, pl. 18, fig. 1. Cancer dormia Linnaeus, 1763: 413: 1769: l043. — Fabricius 1775: 405. Cancer dromia — Fabricius 1781: 501; 1787: 320: l793: 451 (erroneous spelling for dormia). Cancer dormitator Herbst, l790: 250, pl. 18, fig. 103. Dromia rumphii Weber, 1795: 92. — Fabricius 1798: 359. — Latreille 1803 : 386; 1806 : 27; 1818 : 278, fig. 1. — Lamarck l818: 264. — Hilgendorf 1879: 812 (part, Mozambique). — Lenz 1901: 450. — de Man l902: 687. — Nobili 1906a: 144. — Edmondson 1922: 33, pl. l. Dromia hirsutissima Dana, 1852: 403 (part). — Borradaile 1903: 298. — Macnae & Kalk 1958: 7l, ll7, 125. —McLay 1993: 151, fig. 16c; 2001: 82.—Ng et al. 2000: 159, fig. 2a. —Ng et al. 2001: 5 (list). — Ng & Davie 2002: 370 (list). — Takeda & Manuel- Santos 2006: 84, fig. 1D. Dromidiopsis dormia — Rathbun 1923b: 67. — Sakai 1936: 11, pl. 5, fig. 2. — Buitendjik 1939: 223. — Ward 1942: 70. — Tinker 1965: 66. — Holthuis 1968: 220. — Takeda 1973: 79. — Alcala 1974: 174, figs la, b. — Sakai 1976:9, pl. 3. — Dai, Yang, Song & Chen 1981: 131, figs l, 2, pl. 1 (1). — Lewinsohn 1984: 95, pl.2. — Dai & Yang 1991: 18, figs 4 (2, 3), pl. 1 (2). [Dromia] dormia — Guinot & Quenette 2005: 287, figs. 8 A, B, 9 A–C. Not Dromia rumphii — H. Milne Edwards 1837: 174. — de Haan 1839: 107. — Stimpson 1858: 240; 1907: 177, pl.21,

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fig. 7. — Targioni-Tozzetti 1877: 207. — Ortmann 1892: 548. — Alcock 1900:137; l90l: 44, pl.2, fig. 4. — Borradaile 1903b: 576, pl. 33, fig. 1 [= Dromidiopsis dehaani (Rathbun, 1923b)]. Not Dromia dormia – Rathbun 1902: 32. — Stebbing 1905: 61; l9l0: 342. — Ihle l9l3: 22. — Shen l93l: 96, figs 3a, b, 4 a, b. — Barnard 1950: 310, fig. 58c–e. [= Dromidiopsis dehaani (Rathbun, 1923b)]. Not Dromia dornica — Balss 1913: 109 (erroneous spelling for dormia) (= Dromidia aegibotus Barnard, 1947). Not Dromia dormia — Barnard 1947: 366 (= Dromidia aegibotus Barnard, 1947).

Material Examined. Taiwan, Ta-chi: ZRC2000.1819, male, 130.2 mm x 103.3 mm, collected by P.K.L. Ng from local fishermen, Apr. 2000. Philippines, Pangalao, Balicasa I.: ZRC2002.0636, female, 127.3 mm x 106.2 mm, collected in tangle nets by local shell fishermen, June 2002. Description (modified after McLay, 1993). Carapace wider than long (CW/CL = 1.1–1.2), strongly convex, rising steeply behind front and from anterolateral margins, covered by short velvety tomentum. Cardiac, branchial grooves shallow, as is frontal groove which extends back from median rostral tooth, separating two rounded protuberances. Frontal area narrowed, rostrum tridentate, median tooth large, blunt and extending further forward than lateral teeth, clearly visible dorsally. All 3 rostral teeth directed horizontally. Anterolateral carapace margin begins beneath suborbital level and bears 4 unequal teeth. The first tooth is by far the largest, second much smaller and more acute, third very small, close to the second and the fourth intermediate in size between the first and second narrow and more acute, directed slightly upward. Anterolateral teeth are arranged along an almost straight line connecting the rostrum and posterolateral tooth which is large, broad based, narrowing apically and directed anteriorly. Posterolateral carapace margin convergent and posterior margin almost straight. Supraorbital margin extends uninterrupted from lateral rostral tooth, concave to postorbital corner where there is a narrow fissure. Suborbital margin has small rounded lobe which is almost vertical rather than horizontal. Immediately beneath suborbital margin is large, prominent suborbital tooth which is clearly visible dorsally, and beneath this again is more acute tooth at corner of buccal frame, also visible dorsally. Sternal grooves in female gradually convergent, ending with divergent tips between bases of chelipeds, separated by prominent smooth ridge. First article of antenna much wider than long, beaked medially, gaping narrowly, not twisted. Epistome triangular with smooth convex surface. Chelipeds (P1) massive. Merus trigonal, posterior margin with 7–8 small tubercles, inferior margin with 4–5 larger tubercles, anterior margin smooth. Outer surface of carpus sculptured, distal margin with two blunt extensions, superior margin with strong, acute distal tooth. Outer face of propodus inflated, inner superior margin with 4 tubercles, inner face covered with shaggy tomentum. Fixed finger armed with 7–8 large conical teeth. Dactylus with 8 teeth, the first large, blunt, second to fourth much smaller, fifth large, more acute, the last three much smaller. Fingers down curved, only last four teeth interlocking. Ratio of width to length of sternite 4 of approximately 1.9 (male) to 2.2 (female). First two pairs of legs (P2 & P3) shorter than chelipeds, distal borders of carpi and propodi produced as rounded lobes. Dactyli much shorter than propodi, inner margins armed with 4–5 strong spines, set at an angle close to the dactylus and increasing in size distally. Last 2 pairs of legs (P4 & P5) reduced, fourth pair slightly shorter and stouter. Dactylus of third leg (P4) opposed by strong propodal spine, no spine on outer propodal margin. Dactylus of fourth leg (P5) opposed by two similar spines with another small spine on outer propodal margin. Telson about as wide as long, central longitudinal furrow present distally, central region distally convex which continues along adominal somites. Abdominal locking mechanism in male consists of large serrated boss on bases of first legs against convergent margin of penultimate somite with well developed uropods in front of bosses. All somites of abdomen freely movable in both sexes. G1 a semi-rolled tube, bluntly tipped and setose, G2, needle-like, without exopod.

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FIGURE 5. Tumidodromia dormia (Linnaeus, 1763) nov. comb., male, 130.2 x 103.3 mm (ZRC 2000.1819), Taiwan: a. dorsal view; b. frontal view.

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FIGURE 6. Tumidodromia dormia (Linnaeus, 1763) nov. comb., male 130.2 x 103.3 mm (ZRC 2000.1819, Taiwan) a. male sternite 4; female 127.3 x 106.2 mm (ZRC 2002.0636, Philippine Is) b. female sternite 4.

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Remarks. Because of its large size and occurrence in shallow waters, T.dormia was one of the earliest dromiids collected. Because there are several large Indo-West Pacific dromiids, however, there has been much confusion about the identity of these species. The main problem was the separation of T. dormia (Linnaeus, 1763) from “Dromia rumphii” Weber, 1795. Rathbun (1923) pointed out that these two names had been used rather indiscriminately for two different species and provided a partial answer to the difficulties by erecting “Dromidiopsis dehaani” Rathbun, 1923 for some of the specimens that had been called “Dromia rumphii”. Lewinsohn (1984) also helped to clarify the situation, but placed T. dormia in “Dromidiopsis dormia”. The revision of the Dromiidae by McLay (1993) showed that the genus Dromidiopsis in fact contains only rather small-size dromiids and should not include either of the two above species. McLay (1993) used “Dromia dormia” (Linnaeus, 1763) and erected a new genus, McLay, 1993. This allowed “Dromia rumphii” and “Dromidiopsis dehaani” to be placed in (Rathbun, 1923). T. dormia and L. dehaani can be easily separated using some simple characters. In T. dormia, the anterolateral carapace margin is armed with 4 unequal teeth, first largest (anterolateral margin in L. dehaani armed with 3 teeth); the posterolateral carapace tooth is large, broad-based, blunt and directed anteriorly (posterolateral tooth similar to anterolateral teeth, pointed, directed laterally in L. dehaani); the inner margins of the dactyli of the first two pairs of walking legs (P2 and P3) are armed with 4 or 5 strong spines (16–20 small spines in L. dehaani); the female sternal grooves are convergent, ending between the cheliped bases with the spermathecal openings separated by a ridge (7/8 sutures end apart with spermathecal openings at base of well developed tubes behind chelipeds in L. dehaani); all the abdominal somites are freely moveable (joint between fifth and sixth somites partially fused in L. dehaani). T. dormia is covered in an evenly short, dense, light brown tomentum whereas in L. dehaani the tomentum is longer and less dense. Lewinsohn (1984) also lists differences between these two species as well as L. intermedia Laurie, 1906. Specimens are often caught in shallow depths by fishermen and SCUBA divers. Maximum recorded sizes are male 200 mm x 160 mm, female 172 mm x 136.5 mm. T. dormia is the largest known sponge crab and as such it is often collected, but the whereabouts of small specimens (CW<50 mm) remains a mystery. Perhaps they take shelter in coral reef crevices, but they should then be occasionally taken by divers. Only two ovigerous T. dormia females have been collected: the first a female 112.2 mm x 95.6 mm, from New Caledonia, carrying approximately 24,000 eggs (0.5 mm diam.) (McLay, 1993), and the second a female 137.3 mm x  104.0 mm from the Marquesas Is, carrying approximately 129, 600 new eggs, 0.5 mm diam.). T. dormia is a dromiid that produces large numbers of relatively small eggs, especially for a dromiid crab, and is similar to L. dehaani. T. dormia has a very wide geographic distribution, like G. w i ls on i , but has much larger brood sizes. Larvae are not known for this species. Tumidodromia dormia has been recorded carrying sponges (Hyattella sp.), sometimes clearly insufficient for the purpose of total concealment and covering only 40% of the body, the posterior third of the carapace (McLay, 2001). Edmondson (1946) provides several records and some interesting examples of the use of diverse varieties of camouflage carried by Hawaiian T. dormia: in one case a hollowed out piece of wood and in another an old shoe sole. Clearly, individual improvisation is possible in this species. Distribution Tumidodromia dormia is found in the Indo-West Pacific region, from the Red Sea, East Africa, Madagascar, Seychelles, Mauritius, China, Japan, Ambon (Indonesia), Philippines, New Caledonia, Hawaiian Is, and Marquesas Is (French Polynesia). Depth range is 8–112 m.

Section Eubrachyura Saint Laurent, 1980

Family Leucosiidae Samouelle, 1819

Tanaoa Galil, 2003

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FIGURE 7. Tanaoa pustulosus (Wood-Mason in Wood-Mason & Alcock, 1891), male31.5 x 32.3 mm (stn KAH0204/ 02): a. dorsal view, b. frontal view.

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FIGURE 8. Tanaoa pustulosus (Wood-Mason in Wood-Mason & Alcock, 1891), male 31.5 x 32.3 mm (stn KAH0204/ 02) a. frontal view; ovigerous female, 43.9 x 43.0 mm (stn KAH0204/03) b. dorsal view.

Tanaoa pustulosus (Wood-Mason in Wood-Mason & Alcock, 1891) (Figs 7a, b, 8a, b, 9a, b)

Randallia pustulosa Wood-Mason in Wood-Mason & Alcock, 1891: 266. — Alcock 1896: 196. — Doflein 1904: 42, pl. XIV figs 1–5. — Ihle 1918: 246. — Yaldwyn & Dawson 1976: 95, figs 2–5. — McLay 1988: 100, fig. 20. Tanaoa pustulosus — Galil 2003: 404, figs 1D, 3G, H.

Material Examined: East of North Cape: NIWA stn Z11043, 1 male, 31.9 mm x 31.5 mm, 34°57.81’S, 175°12.64’E, 614–602 m, Kaharoa, stn KAH0204/01, 13 Apr. 2002 (NIWA 48569); NIWA stn Z11044, 1 male, 31.5 mm x 32.3 mm, 34°57.62’S, 175°10.79’E, 581–553 m, RV Kaharoa, stn KAH0204/02, 13 Apr

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2002 (NIWA 4873); NIWA stn Z11045, 1 ovigerous female, 43.9 mm x 43.0 mm, 34°57.63’S, 175°10.44’E, 554–518 m, Kaharoa, stn KAH0204/03, 13 Apr. 2002 (NIWA 48575). Remarks. With their almost rounded body and long chelipeds, the two species of Tanaoa Galil, 2003, are very distinctive species that are unlikely to be confused with any other New Zealand crabs. Besides T. pustulosus, a second species, T. distinctus (Rathbun, 1893) has been collected at 472 m depth north of North Cape. These two species can be distinguished by the anterolateral margin with 5 or 6 prominent tubercles larger than the rest (margins bear many low tubercles in T. distinctus) and intestinal region with a prominent tubercle anteriorly and upcurved spur-like tubercle posteriorly (intestinal region only has low tubercle posteriorly). Tanaoa pustulosus has been reported from the Kermadec Is and Bay of Plenty, east North I., but at much shallower depths (330–475 m). The other leucosiids recorded from New Zealand are Bellidilia cheesmani (Filhol, 1886), Ebalia tuberculosa (A. Milne Edwards, 1873), and Merocryptus lambriformis A. Milne Edwards, 1873. As ambush predators, these crabs spend much of their lives buried in soft sand, with only the eyes exposed. The ovigerous female T. pustulosus carried a large number of small eggs (diameter = 0.38–0.4mm) that were well protected by the tightly sealed brood chamber formed by the edges of the abdomen fitting into the elevated sternal rim as is typical in leucosiids. Fusion of abdominal somites 4–6 aids the formation of the brood chamber. The small egg size indicates that this species has indirect planktotrophic development and helps to explain the wide distribution of this species (see below). Distribution. Tanaoa pustulosus inhabits the Indo-West Pacific region: East Africa, Seychelles, Mozambique Channel, Comoro Is, Madagascar, Reunion, Laccadive Sea, Andaman Sea, Japan, Taiwan, Philippines, Indonesia, New Caledonia, New Zealand, Fiji, Caroline Is, Agalega Is, and Geyser Reef, The specimens reported here fall within the known depth range 85–977 m (Galil, 2003).

Family Inachidae Macleay, 1838

Dorhynchus Wyville Thomson, 1873

Dorhynchus ramusculus (Baker, 1906) (Figs. 10a, b)

Stenorhynchus ramusculus Baker, 1906: 104, pl. 1 figs 1, 1a. Achaeopsis ramusculus — Hale 1927: 124, fig. 121. — Griffin 1966: 35, fig. 4. — Griffin & Tranter 1986a: 21; 1986b: 352. — McLay 1988: 116, fig. 25a–d. Dorhynchus ramusculus — Davie 2002: 293. — Ahyong 2008: 36, figs. 16B, C

Material Examined. Chatham Rise: 1 male, 11.9 mm x 19.7 mm, another similarly-sized male too damaged to measure, 42°45.93’S, 179°59.34’W, 875–757 m, Tangaroa, stn TAN0104/02, 15 Apr. 2001 (NIWA 48576); 1 ovigerous female, 10.1 mm x 15.2 mm, 42°47.17’S, 179°59.11’W, 993–900 m, Tangaroa, TAN0104/48, 16 Apr. 2001 (NIWA 48570). Remarks. The genera Dorhynchus Thomson, 1873 and Achaeopsis Stimpson, 1857 have had a complex history that has been summarized by Griffin (1966), Manning & Holthuis (1981) and Davie (2002). The species has been known until now New Zealand as Achaeopsis ramusculus following the arguments put forward by Griffin (1966). However, Manning & Holthuis (1981) argued that the two genera should not be synonymised as had been proposed by Griffin (1966). Following Davie (2002) the species should now be known as Dorhynchus ramusculus. Dorhynchus ramusculus is most likely to be confused with Achaeus curvirostris (A. Milne Edwards, 1873), A. akanensis Sakai, 1937, and A. kermadecensis Webber & Takeda, 2005, but they are easily separated because they lack the long, divergent rostral spines.

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FIGURE 9. Tanaoa pustulosus (Wood-Mason in Wood-Mason & Alcock, 1891), ovigerous female, 43.9 x 43.0 mm (stn KAH0204/03): a. ventral view; b. frontal view.

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FIGURE 10. Dorhynchus ramusculus (Baker, 1906), male 11.9 x 19.7 mm (stn TAN0104/02): a. dorsal view, b. frontal view.

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Distribution. Dorhynchus ramusculus is restricted to New Zealand and Australia (from Western Australia to New South Wales). Its reported depth range is 150–740 m, but it should be noted that the specimens examined here are from 757–993 m, extending the lower depth limit. The species is now reliably known from 140–993 m. Its southern limit now extends to the Chatham Rise, 42°47’S.

Family Inachoididae Dana, 1851

Pyromaia Stimpson, 1871

Pyromaia tuberculata (Lockington, 1877) (Fig. 11)

Inachus tuberculatus Lockington, 1877: 30. Pyromaia tuberculata — Rathbun 1925: 133, pl. 40, fig. 3, pl. 218, figs 1–4. — Webber & Wear 1981: 370, figs 209–217 (larval stages). — Wear & Fielder 1985: 28, figs 71–73 (larval stages). — Fransozo & Negreiros-Fransozo 1997: 308, figs 2–7 (larval stages). — McLay 1988: 112, fig. 24. — Lemaitre, Campos & Bermudez 2001: 771 (key). —Davie 2002: 300, 299, unnumbered fig. (pl. 218, fig. 2 from Rathbun, 1925). — Poore 2004: 366.

Material Examined. Whangarei Harbour: 1 juv., 1.1 mm x 1.3 mm, 35° 47′S, 174° 24′E, pile scraping, WRE134CB; 1 male, 7.3 mm x 9.9 mm, 35° 47′S, 174° 24′E, benthic sled, WRE603CB. Auckland Harbour: 1 ovigerous female, 6.6 mm x 8.7 mm, 36° 50′S, 174° 52′E, benthic sled, AKL512CB, 4 Apr. 2003 (NIWA 48748); 1 male, 6.0 mm x 8.1 mm, 36° 50′S, 174° 52′E, benthic sled, AKL514CB, 4 Apr. 2003 (NIWA 48746); 1 male, 6.0 mm x 7.5 mm, 36° 50′S, 174° 52′E, benthic sled, AKL516CB, 7 Apr. 2003.

FIGURE 11. Pyromaia tuberculata (Lockington, 1877), ovigerous female 6.6 x 8.7 mm (AKL512): dorsal view.

Remarks. Pyromaia tuberculata is spreading northwards from Auckland, North I. because it now occurs in Whangarei Harbour. It is distinctive because it is the only shallow-water majid with a simple rostrum consisting of only a single projection.

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The ovigerous female from Auckland Harbour carried around 1200 small eggs (diameter = 0.4 mm) in a swollen brood chamber that is large in relation to the rest of the body. A raised rim around the sternum fits tightly with the edges of the abdomen to protect the eggs. istribution. Pyromaia tuberculata is found in the Eastern Pacific Ocean (San Francisco, California to Colombia) and Brazil (Lemaitre et al. 2001); introduced by shipping to Western Australia, Japan (Sagami Bay), New Zealand (Firth of Thames, Tamaki Estuary and now Whangarei Harbour). Depth range is intertidal to 650m.

Family Epialtidae MacLeay, 1838

Pisinae Dana, 1851

Rochinia A. Milne-Edwards, 1875

Rochinia ahyongi sp. nov. (Figs 12a, b, 13a–g)

? Sphenocarcinus. — Clark & O’Shea 2001: 15

Material Examined. Rumble V Seamount: NIWA stn Z10806, 1 male, 8.7 mm x 11.7 mm (incl. spines and rostrum), 36°08.84–08.59’S, 178°12.24–11.99’E, 951–772 m, Tangaroa, stn TAN0107/225, 23 May 2001 (NIWA 42391. Holotype. Male, 8.7 mm x 11.7 mm, NIWA stn Z10806, 36°08.84–08.59’S, 178°12.24–11.99’E, 951–772 m, Tangaroa, stn TAN0107/225, 23 May 2001 (NIWA 42391). Etymology. The specific name of the new species acknowledges the valuable contribution of Shane Ahyong (NIWA) to our knowledge of New Zealand Brachyura, especially deep-sea species. Description. Carapace pyriform, much longer than wide, surface convex, finely granular with sparse, scattered, long, fine, straight or hooked setae. Rostral spines long (making up approximately 32% of total length), strongly divergent (distance between tips 2.4 width), 2 pairs of lateral spines, shorter anterior pair at mid-point of carapace (making up 30% of width), longer posterior pair further back (making up approximately 38% of width), all spines narrowing to blunt tip, cardiac region with low swelling in mid-line. Posterior carapace margin with rounded protuberance. Orbit incomplete dorsally, and ventrally, with small preorbital tubercle, postorbital corner slightly produced, blunt, remainder of orbital margin unarmed. Basal article of antenna fixed, lateral margin straight, anterolateral corner slightly produced, articles 2, 3 successively shorter, flagella not reaching tip of rostral spine. Basal article separated from post-orbital lobe by narrow U-shaped hiatus. Chelipeds (P1) same length as CW (including spines), narrow, palm about 5 times as long as wide, fingers elongate margins approximated; cutting edges with 3, 4 very small teeth on both chelipeds. Moveable finger approximately 0.5 propodus length. Walking legs (P2–P5) narrow, smooth, first pair longest 1.25x carapace length (decreasing in length posteriorly), merus about same length as carpus + propodus length, margins unarmed; dactylus about same length as propodus. Abdomen of 6 free somites, second somite widest decreasing in width posteriorly, penultimate somite much wider than long, telson as wide as long, posterior margin rounded. First male gonopod (G1) straight, flattened, tip bilobed, medial lobe angled and longer. Remarks. One other species of this genus is known from the vicinity of the Kermadec Is, Rochinia riversandersoni (Alcock, 1895), but the two species are easily distinguished by the fact that this species has 12 long carapace spines while R. ahyongi sp. nov. only has four spines. Both species do have long divergent rostral spines. In the Australasian area the only spider crab resembling R. ahyongi sp. nov. is Rochinia fultoni (Grant, 1906) from New South Wales and Tasmania, but it has anterior lateral spines that are short and

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FIGURE 12. Rochinia ahyongi sp. nov. holotype male 8.7 x 11.7 mm (stn TAN0107/225) Registration Code NIWA 42391: a, dorsal view, b, ventral view.

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FIGURE 13. Rochinia ahyongi sp. nov. holotype male 8.7 x 11.7 mm (stn TAN0107/225) Registration Code NIWA 42391: a. dorsal view (scale bar = 2 mm), b. dorsal view of orbit (scale bar = 0.5 mm), c. abdomen (scale bar = 1 mm), d. first gonopod (scale bar = 0.5 mm), e. outer face of cheliped palm (scale bar = 1 mm), f. dorsal view of cheliped (scale bar = 1 mm), g. dorsal view left fourth pereopod (scale bar = 1 mm).

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Family Palicidae Bouvier, 1898

Pseudopalicus Moosa & Sèrene, 1981

Pseudopalicus undulatus Castro, 2000 (Figs. 14a, b)

Pseudopalicus undulatus Castro, 2000: 483, figs 12f, 14, 53.

Material Examined. East of North Cape: 1 ovigerous female, 11.0 mm x 9.1 mm, 34°06 34.22’S, 174°08 18.05’E, 600–562 m, Kaharoa, stn KAH0204/09, 14 Apr. 2002 (NIWA 48577). Remarks. The genus Pseudopalicus contains 11 species found in the Indo-West Pacific region, from the coast of Africa to French Polynesia and as far north as Japan and as far south as New Zealand. The first reported palicid from New Zealand waters was P. oahuensis (Rathbun, 1906) by Takeda & Webber (2006) collected in 1975, from a depth of 530–585 m off Raoul I., northeast of New Zealand. P. undulatus Castro, 2000 is the second species recorded and comes from a similar depth East of North Cape, so is much closer to the New Zealand mainland. This record considerably extends the southernmost limit for this species from around 18°S (Fiji) to 34°S (New Zealand). It has also been recorded from the Coral Sea, Indonesia, South China Sea and Japan. Castro (2000) gave the reliably known depth as 206–410 m, but the New Zealand specimen came from 600–562 m, thereby extending the lower depth limit. These species can be distinguished the number of anterolateral teeth, 3 for P. undulatus, 4 for P. oahuensis. Three other species, P. acanthodactylus Castro, 2000, P. declivis Castro, 2000 and P. glaber Castro, 2000 are known from the Norfolk Ridge. Thus 5 of the 11 species in the genus are known from the warmer waters of New Zealand and nearby areas to the north. Other palicid species occurring near New Zealand include Crossotonotus spinipes (De Man, 1888) also from the Norfolk Ridge. The excellent revision by Castro (2000) provides keys to identify the genera and species of the Indo-West Pacific Palicidae. The ovigerous female P. undulatus carried around 800 small eggs (diameter = 0.45 mm) that are protected by a brood chamber formed by the swollen abdomen fitting tightly with the elevated sternal rim. In this respect palicids are similar to leucosiids. The small egg size indicates that this species has indirect

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FIGURE 14. Pseudopalicus undulatus Castro, 2000, ovigerous female 11.0 x 9.1 mm (stn KAH0204/09): a. dorsal view, b. frontal view.

Family Atelecyclidae Ortmann, 1893

Trichopeltarion A. Milne Edwards, 1880

Trichopeltarion janetae Ahyong, 2008 Fig. 15

Trichopeltarion janetae Ahyong, 2008: 17, figs 7–11, 28 D)

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Material Examined. Chatham Rise (Graveyard Complex): 2 females, 13.9 mm x 12.3 mm, 16.1 mm x 14.9 mm, 42°43.03’S, 179°57.55’W, 1051–964 m, Tangaroa, stn TAN0104/82, 17 Apr. 2001 (NIWA 48583); 5 females, 12.5 mm x 11.3 mm, 14.1 mm x 12.7 mm, 18.9 mm x 16.9 mm, 20.2 mm x 17.0 mm, 23.7 mm x 21.0 mm, 1 ovigerous female, 20.5 mm x 18.2 mm, 1 male, 15.9 mm x 13.8 mm, 42°43.10’ S, 180°5.43’E, 1075–1008 m, Tangaroa, stn TAN0104/333, 20 Apr. 2001 (NIWA 48582). Colville Ridge: 1 female, 11.6 mm x 11.0 mm, 32°36.88’S, 179°38.17’W, 888–741 m, Tangaroa, stn TAN0205/34, 17 Apr. 2002 (NIWA 48728).

FIGURE 15. Trichopeltarion janetae Ahyong, 2008, 2000, female 16.1 x 13.9 mm (stn TAN0104/82): dorsal view.

Remarks. This species was described using a large sample of the new species from eastern New Zealand, from Bay of Plenty south to the Chatham Rise and Bollons Seamount at depths of 830–1506 m. The species has also been reported from seamounts off Tasmania (Ahyong, 2008) so the female from the Colville Ridge bridges the gap between NZ and Australia. An ovigerous T. janetae female, 20.5 mm x 18.2 mm, carried around 1600 eggs of diameter 0.95 mm. The smallest ovigerous female reported by Ahyong (2008) was 16.7 mm x 17.3 mm. This species has a relatively small abdomen that is totally inadequate for protecting the brood of eggs (cf. the high level of protection provided by the brood chamber in Pyromaia tuberculata, Pseudopalicus undulatus and Tanaoa pustulosus see herein). Thompson & McLay (2005) suggested that in Trichopeltarion, and some other atelecyclid genera, brood protection of the relatively small number of large eggs may be provided by living in a burrow. T. janetae is readily distinguished from the only other species in New Zealand waters, T. fantasticum Richardson & Dell, 1964 because it is much smaller, the carapace is densely setose rather than sparsely setose and the last anterolateral tooth is small compared to the large tooth in T. fantasticum. Males differ in that T. janetae do not have the enormously developed right cheliped; sexual dimorphism is much more modest.

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Ahyong (2008) provides excellent photos of the two NZ species of Trichopeltarion. Distribution The distribution of Trichopeltarion janetae includes Tasmania, Colville Ridge and New Zealand. Depth range is 830–1700 m.

Family Calappidae de Haan, 1833

Mursia Desmarest, 1823

Mursia microspina Davie & Short, 1989 (Figs 16a, b, 17a, b, 18a, b)

Mursia microspina Davie & Short, 1989: 172, figs 9a–g, 10 [type locality: SE Queensland, Australia]. — Galil 1993: 365, figs 4e, 6h, i, 8e, f, 12 (colour). — Davie 2002: 128. — Takeda & Komatsu 2005: 277. Mursia aspersa — Baba, Hayashi & Toriyama 1986: 221, pl. 165 [not Mursia aspersa Alcock, 1899].

Material Examined. Bay of Plenty: 1 female, 19.0 mm (with spines 20.8 mm) x 15.9 mm, 1 male, 30.8 mm (with spines 33.2 mm) x 26.8 mm, 37°28.14’S, 177°6.97’E – 37°28.11’S, 177°6.51’E, 230–318 m, Kaharoa, stn KAH9907/50, 5 June 1999 (NIWA 48578); 1 female, 27.7 mm (with spines 32.6 mm) x 24.0 mm, 37°32.98’S, 176°58.44’E – 37°32.97’S, 176°58.63’E, 280–155 m, Kaharoa, stn KAH0011/40, 5 Nov. 2000 (NIWA 48581). Description. (only male characters unavailable for the type specimen and some additional female characters are given here): male abdominal somites 3–5 fused, but sutures still evident; somite 1 hidden beneath posterior margin of carapace; somite 2 with 3 large flattened, blunt transverse tubercles (“tri-lobate carina” of Galil, 1993); somite 3 with 2 low swellings on posterolateral corners; abdominal locking mechanism on somite 6 uses excavated posterolateral corners of to lock on to blunt tubercles on P2 sternite; telson triangular, longer than wide tip rounded. All somites of female abdomen free, margins setose; telson as in male; abdominal locking mechanism tubercles still present on P2 sternite of mature female. Gonopore simple, not operculate. Cheliped superior margin in both sexes setose; outer face of propodus has 8 larger blunt tubercles adorning finely granulate surface, inferior margin of propodus with 12–14 small acute spines increasing in size distally; fingers down-curved, stout, tips crossed when closed; left cheliped fingers with 4, 5 interlocking teeth; right crusher cheliped fingers differ markedly, dactylus with large proximal molariform tooth fitting into proximal slot behind ridge on fixed finger and with 3, 4 distal teeth; margin of fixed finger divided into 2 parts, proximally there are 2 stout blunt teeth corresponding to molariform tooth on dactylus, while distal part has 3, 4 teeth increasing in size distally. Inner face of both male and female dactylus bears stridulatory ridge of about 25 fine transverse striae as is typical for Mursia. Row of about 20 small blunt tubercles lie along distal margin of third maxilliped ischium. Walking legs (P2–P5) articles flattened, smooth, dactyli long, styliform, dorsal margin of meri setose. P2 longest, other pereopods decreasing in size posteriorly in both sexes. Remarks. Davie & Short (1989) only had a male specimen that was missing its abdomen all the walking legs. The present collection includes intact specimens as well as females so it is now possible to complete the description of M. microspina. The male abdomen conforms to the generic pattern. Galil (1993) has already recorded several females from New Caledonia and the Loyalty Is, but she did not comment on the female characters. The female of M. microspina is typical of others in this genus. Compared to the type specimen described by Davie & Short (1989) the present specimens show a few small differences: the seven rows of tubercles radiating out from the front in the type are less distinct, being more scattered than aligned; the rostrum is evenly rounded, rather than sculptured as in the type; the outer distal margin of cheliped merus has only two (rather than three) spines increasing in size, but there is a low proximal tubercle that may represent the first spine; the projections on the posterior carapace margin are more

36 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. pronounced than as shown in Davie & Short (1989: fig. 9g). In other respects the New Zealand specimens agree closely with those from Australia. One feature that may be diagnostic of this species is the presence of a group of four blunt tubercles, forming the corners of an oblong, behind the orbits: the mesial pair is larger and further apart than the lateral pair. The supraorbital margins are eroded, allowing the eyes to view upwards, an appropriate adaptation for a sit-and-wait ambush predator that buries itself in the sand.

FIGURE 16. Mursia microspina Davie & Short, 1989, female 32.6 x 24.0 mm (stn KAH0011/40); a. dorsal view, b. frontal view.

Galil (1993: fig. 12) provided a colour photo of a male from New Caledonia and described it as having “Branchial regions tinged red. Distal margin of buccal cavity [i.e. anterior face of epistome, just below antennules] with two red spots. Chelipeds pale coral, fingers white. Inner palmar face with a large oculus- shaped coral-coloured patch.” The specimens from New Zealand show the same patterns except that the patch

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FIGURE 17. Mursia microspina Davie & Short, 1989, a. female 32.6 x 24.0 mm (stn KAH0011/40): a, ventral view; male 33.2 x 26.8 mm (stn KAH9907/50): b, dorsal view.

Sound production in M. microspina involves a stridulatory ridge of around 25 fine transverse striae on the inner surface of the cheliped dactyli that a row of around 20 small rounded tubercles on the distal margin of the third maxilliped ischium rub against. These crabs normally hold their chelipeds close to their “face” as they lie semi-buried in sand, so sound production involves minimal movement of these appendages, thereby maintaining the cryptic position of the crab. This species can be compared with another sound producing crab

38 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. that is part of the NZ fauna (see below). Ocypode pallidula has a similar row of striae on the inner face of the cheliped palm and to produce sound these are rubbed against a plectrum on a pair of closely spaced ridges on the ventral margin of the cheliped ischium. O. pallidula normally produces sound by vertical movements of the chelipeds while hiding in its burrow. In this species only one appendage is used to make the sound whereas in M. microspina both the chelipeds and third maxillipeds are employed. Sound is also produced by Ovalipes elongatus and O. catharus, but they employ different appendages (see O. elongatus below). M. microspina is easily distinguished from M. australiensis by the short lateral carapace spines and the short meral spines on the chelipeds.

FIGURE 18. Mursia microspina Davie & Short, 1989, male 33.2 x 26.8 mm (stn KAH9907/50): a. ventral view, b. frontal view.

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Distribution. Southeast Queensland, New Caledonia, Loyalty Is, and New Zealand. M. microspina lives on the continental shelf and continental slope. The New Zealand specimens came from samples taken between 155–318 m, but the exact upper limit of M. microspina in the shallowest sample is uncertain and could be anywhere from 155–280 m. Depth range is 200–420 m.

Mursia australiensis Campbell, 1971 Fig. 19

Mursia australiensis Campbell, 1971: 31, pl. 2a, b, fig. 1. — Galil 1993: 355,figs 1d, 2g, h, 3e, f, 11 (colour). — Davie 2002: 128.

Material Examined. Lord Howe Rise, Capel Bank: NZOI stn I730, 1 specimen (sex unknown) 24.4 mm x 20.0 mm (CW incl. spines 38.6 mm), 25°32.50’S, 159°38.99’E, 300 m, 10 May 1979. This record is based on a photo of a live specimen at the time of capture: whereabouts of the specimen unknown. Remarks. M. australiensis is easily distinguished from M. microspina by the longer lateral carapace spines (about 40% of CW, about 10% of CW in M. microspina), the longer meral spines on the chelipeds (about 35% of CW, about 5% of CW in M. microspina) and colour of patch on inner palm surface of chelipeds (small light red patch vs large dark red-purple patch). The present specimen came from 300 m, within the known depth range of 40–320 m. Distribution. Australia (Queensland, New South Wales), New Caledonia and now Lord Howe Rise, Tasman Sea. Depth range 40–320 m.

FIGURE 19. Mursia australiensis Campbell, 1971: Lord Howe Rise, Capel Bank: sex unknown, 24.4 x 20.0 mm (NZOI stn I730): dorsal view (scanned photo).

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Family Portunidae Rafinesque, 1815

Ovalipes Rathbun, 1898

Ovalipes elongatus Stephenson & Rees, 1968 Fig. 20a

Ovalipes elongatus Stephenson & Rees, 1968: 232, fig. 1E, pls 36B, 39D, 42E. — Stephenson 1972: 23. — Davie 2002: 460. — Takeda & Webber 2006: 200, fig. 5D. Ovalipes bipustulatus — Chilton 1911: 554.

Not Ovalipes bipustulatus — Chilton & Bennett 1929: 755. [= Ovalipes catharus White, in White & Doubleday, 1843].

Material Examined. Kermadec Is: AQ489, 1 male, 13.3 mm x 11.9 mm, 1 female, 27.6 mm x 23.5 mm, Canterbury Museum, coll. W.R.B. Oliver, 1908. [Handwritten label, presumably by Chilton, shows “Ovalipes bipustulatus M. Edw.”, but also labelled as “Ovalipes elongatus identified by Stephenson & Rees, 1966”.]; AQ2799, 1 male 9.0 mm x 8.0 mm, Canterbury Museum, coll. R. Bell 1909–10; AQ2445, 2 females, 29.1 mm x 24.5 mm, 30.1 mm x 25.7 mm, cast ashore Raoul I., Canterbury Museum, coll. J.H. Sorenson 27 July, 1944. Whangarei Harbour: 1 male, 10.3 mm x 8.3 mm, 35° 43′ S, 174° 19′E, benthic sled, WRE621CB, 21 Nov. 2002 (NIWA 48729). Tauranga Harbour: 2 males, 6.3 mm x 5.3 mm, 6.5 mm x 5.4 mm, 37° 39′S, 176° 10′E, TRG801CB, 24 Feb. 2004 (NIWA 48730). New Plymouth Harbour: 1 female, 8.6 mm x 7.0 mm, 39° 04′S, 174° 04′E, benthic sled, TK022CB, 16 Apr. 2002 (NIWA 48731); 1 juv., 6.0 mm x 5.4 mm, 39° 04′S, 174° 04′E, 2NPL068, 15 Mar. 2005 (NIWA 48732); 3 juveniles, 5.5 mm x 5.2 mm, 5.6 mm x 5.3 mm, 6.0 mm x 5.4 mm, 39° 04′S, 174° 04′E, 2NPL076, 15 Mar. 2005 (NIWA 48733). Comparative Material. Ovalipes catharus (White, in White & Doubleday, 1843): Little Akaloa Bay, Banks Peninsula: female, 12.5 mm x 10.3 mm, 43° 40′S, 173° 00′E, 3 m, Apr. 1, 2008, personal collection (see Fig. 20b). Remarks. Chilton (1911) first reported this species from the Kermadec Is as “Ovalipes bipustulatus M.- Edw.” stating that he had seen “Several small specimens from Sunday Is collected by Mr Oliver and one by Captain Bollons”. These specimens are presumably the ones now in the Canterbury Museum collection as AQ489 (see above). A label with these specimens states that they were verified as O. elongatus by Stephenson & Rees in 1966. The specimen, said by Chilton to be collected by Captain Bollons, may be the small male in AQ2799, but the collection is attributed on the label (not written by Chilton) to “R. Bell, 1909-10” (see above). This collection also includes 18 megalopae (3.7 mm x 4.9 – 4.6 mm x 6.4 mm) that may or may not belong to the same species, but the chelipeds are certainly close to those of O. elongatus. Takeda & Webber (2006) reported a further 33 specimens of O. elongatus in collections from the Kermadec Is held by Te Papa. Thus it seems that O. elongatus is quite common in the Kermadec Is The specimens reported here from Whangarei Harbour (35° 43′ S, 174° 19′E), Tauranga Harbour (37° 39′S, 176° 10′E) and New Plymouth Harbour (39° 04′S, 174° 04′E) are the first specimens of this species reported from mainland New Zealand, extending the range much further south than Lord Howe Is (31° 30′ S, 159° 00′E) and the Kermadec Is (30°S, 178° 30′E). Chilton & Bennett (1929) also use O. bipustulatus (H. Milne Edwards, 1834) for Ovalipes specimens from NZ, but this is O. catharus (White, in White & Doubleday, 1843). Takeda & Webber (2006) included O. catharus in their list of crabs recorded from the Kermadec Is assuming that Chilton’s use of “Ovalipes bipustulatus” indicated the same species as found on the NZ mainland, but Stephenson & Rees showed that two different species were involved and as a result established the new species. O. elongatus Stephenson & Rees, 1968 is the only species of Ovalipes on the Kermadec Is

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FIGURE 20. Ovalipes elongatus Stephenson & Rees, 1968, male 10.3 x 8.3 mm (stn TK022CB): a. dorsal view; Ovalipes catharus (White, in White & Doubleday, 1843) female 12.5 x 10.3 mm (Little Akaloa Bay, Banks Peninsula): b. dorsal view.

O. elongatus (Fig. 20a) and O. catharus (Fig. 20b) are easily separated by the following differences: the tips of the 4 teeth on the frontal carapace margin in O. elongatus project further than a line drawn across the front, connecting the tips of the post-orbital teeth on either side (in O. catharus these 4 teeth do not reach this line), the CW/CL = 1.19 – 1.25 (versus 1.30 – 1.37) and the anterolateral teeth are long, acute and directed anteriorly (versus short, triangular and not breaking up the even curve of the anterolateral margin). The

42 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. maximum sizes reported for O. elongatus are CW = 38 mm (females), 15 mm males mm (Stephenson & Rees, 1968) compared with 115 mm (females) and 150 mm (males) for O. catharus (McLay 1988). No large males have been reported for O. elongatus, but presumably they grow at least as large as the females. O. catharus, from the New Zealand mainland (and South Australia and Victoria), clearly grow to a much larger size than O. elongatus from Lord Howe Is and the Kermadec Is. Ovalipes elongatus is another New Zealand crab that can produce sound (see Mursia microspina and Ocypode pallidula herein) by rubbing the ventral surface of the cheliped propodus across a plectrum on the dorsal margin of the second pereopods. Also Ovalipes catharus from the mainland can also produce sound (see Stephenson, 1969). These three genera each use different mechanisms to make sound and as they each belong to different families (Portunidae, Calappidae and Ocypodidae respectively), it shows that sound production is the result of parallel evolution. Distribution Ovalipes elongatus was formerly only known from Lord Howe I. and the Kermadec Is, but is now known to occur around the northern half of the North I. of New Zealand. O. elongatus is a species endemic to the Tasman Sea. Depth range is intertidal to 55 m.

Liocarcinus Stimpson, 1871

Liocarcinus corrugatus (Pennant, 1777) Fig. 21a, b

Cancer corrugatus Pennant, 1777: 5, pl. 5, fig. 9. Portunus strigilis Stimpson, 1858: 38. Portunus pusillus — Kirk 1878: 402 (not Portunus pusillus Leach, 1815 = Liocarcinus pusillus (Leach, 1815)) Portunus borradailei — Bennett 1930: 256, figs 1–4. Portunus corrugatus — Borradaile 1916: 97, fig. 9. — Chilton & Bennett 1929: 753. Macropipus corrugatus — Stephenson & Campbell 1960: 92, figs 1E, 2G, pl 2, fig. 4, pl. 5G. — Dell 1968: 20. — Stephenson 1972: 23. — Wear & Fielder 1985: 55, figs 142, 143. Liocarcinus corrugatus — Hale 1927: 148, fig. 149. — McNeill & Ward 1930: 379. — Powell 1937: 382. — Bennett 1964: 65. — Manning & Holthuis 1981: 84. — McLay 1988: 216, fig. 48a–d. — Davie 2002: 459. — Takeda & Webber 2006: 200. fig. 5C. — Ng et al. 2008: 149 (list).

Material Examined. Whangarei: 1 male, 33.9 mm x 27.2 mm, 35°47’S, 174°24’E, WRE507CB, 23 Nov. 2002 (NIWA 48747); 1 female, 15.4 mm x 13.2 mm, WRE607CB, 19 Nov. 2002 (NIWA 48734); 1 ovigerous female, 21.6 mm x 18.0 mm, 35°47’S, 174°24’E, WRE608CB, 19 Nov 2002 (NIWA 48735); 1 male, 16.3 mm x 13.6 mm, 35°47’S, 174°24’E, WRE622CB, 21 Nov. 2002 (NIWA 48736). Little Barrier Is: AQ2377, 1 male, 12.6 mm x 10.5 mm, 36°12’S, 175°05’E, 64 m, Hinemoa, 29 Dec. 1914. Tauranga: 1 male, 6.7 mm x 6.0 mm, 37°39’S, 176°10’E, TG142CB, 8 Mar. 2002. Chatham Rise, Giggenbach Seamount: NIWA 18022, 1 female, 19.2 mm x 16.1 mm, 42°43.10’S, 180°5.43’E, 109 m, stn KOK0505/45, 16 Apr. 2005. Remarks. The species of Liocarcinus are in urgent need of revision (Davie, 2002, p459).The detailed synonymy presented here is limited to Australasia, thereby including and verifying all the records therefrom. The only species liable to be confused with L. corrugatus is Nectocarcinus antarcticus. The main differences between them are: frontal carapace margin with 3 evenly rounded lobes (4 small sub-acute teeth in N. antarcticus), anterolateral carapace margin cut into 5 prominent teeth (margin cut into 4 small teeth), and tips of dactyli on swimming legs (P5) pointed (tips bluntly rounded). However these features refer to adults and smaller specimens have anterolateral teeth that are relatively larger (see below for ontogenetic changes). Distribution. Supposedly includes the Atlantic Ocean and the Indo-West Pacific region. Known from Australia and New Zealand. The record from the Chatham Rise extends the southern limit to 42°43’ S. Depth range is 0–137 m (being a swimming crab this does not mean intertidal).

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FIGURE 21. Liocarcinus corrugatus (Pennant, 1777), female 19.2 x 16.1 mm (stn KOK0505/45): a. dorsal view; male 33.9 x 27.2 mm (WRE507CB): b. dorsal view.

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FIGURE 22. a. Thalamita danae, male CW = 58 mm (pl. 3 fig. 1, Stephenson & Hudson, 1957).b. Thalamita macropus, male CW = 24 mm (pl. 4 fig. 1, Stephenson & Hudson, 1957).

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Thalamita Latreille, 1829

Thalamita danae Stimpson, 1858 Fig. 22 a.

Thalamita danae Stimpson, 1858: 37. — Stephenson & Hudson 1957: 335, figs 2N, 3N, pl. 3, fig. 1, pls 7G, 10D. — Stephenson & Rees 1967: 70, figs 25a–c, 26a–c. — Davie 1998: 150, colour photo; 2002: 471 (fig.), 478. — Jones & Morgan 2002: 161, 163, 2 colour photos. Thalamita stimpsoni Milne Edwards, 1861: 362, pl. 35, fig. 34.— Stephenson & Hudson 1957: 356, figs 2M, 3M, pl. 6, figs 1–3, pls 8R, 9I. — Stephenson & Rees 1967: 98, fig. 36.

Material Examined. Hutchinson Bluff, Raoul I., Kermadec Is: 1 male, 10.8 mm x 7.6 mm, 29°16’S, 177°55’W, 84–114 m, beam trawl, pumice rubble, BS296, 24 Aug. 1972. Due to renovation of the collection building at Te Papa, and transfer of specimens to temporary storage, this specimen was not available for photography. Therefore a photo of Thalamita danae (male CW=58 mm) published by Stephenson & Hudson (1957: pl. 3 fig. 1) is reproduced here. For comparison a photo of Thalamita macropus (male CW = 24 mm), the other species of this genus recorded from the Kermadec Is, is included (Fig. 2 b) (Stephenson & Hudson 1957: pl. 4, fig. 1). Remarks. The blue mottled swimming crab, Thalamita danae, has strong granular ridges on outer face of cheliped palm. Its habitat includes littoral, sub-littoral depths to 40 m, rocky shores under stones and mussel clumps, on sandy-mud flats and reefs (Davie, 2002). The specimen reported here came from around 100 m, which is somewhat deeper than recorded in Australia. Takeda & Webber (2006) recorded another species, Thalamita macropus Montgomery, 1931 from Raoul and Chanter Is, Kermadec Is, 31–135 m. The main differences between the two species of Thalamita from the Kermadec Is (see Fig. 22a, b) are: T. danae has the first anterolateral tooth without subsidiary tooth (in T. macropus the first anterolateral tooth has a subsidiary basal tooth behind it on the carapace); tips of the 5 anterolateral teeth form a convex shape with the greatest width at the third teeth and last 2 teeth receding (but in T. macropus the teeth on the anterolateral margin are relatively larger and the greatest carapace width is at the last teeth so that the anterolateral margins are divergent and not convex (see Stephenson & Hudson, 1957, pl. 3, fig. 1, pl. 4, fig. 1). The straight frontal carapace margin, with 6 truncate lobes, immediately distinguishes the 2 species of Thalamita from the other portunids, Ovalipes elongatus, Nectocarcinus antarcticus and Liocarcinus corrugatus. Distribution. Red Sea, Japan, Indo-Malayan region to Australia, New Caledonia, Marshall Is, Fiji, Samoa, French Polynesia, and now further south to Raoul I., Kermadec Is. Depth range 5–100 m.

Nectocarcinus A. Milne-Edwards, 1860

Nectocarcinus antarcticus (Hombron & Jacquinot, 1846) Figs 23a, b, 24a, b

Portunus antarcticus Jacquinot, 1853: 51, figs 1–5, pl. 5. Nectocarcinus antarcticus — Milne-Edwards 1860: 220. — Miers 1874: 2, fig. 2, pl. 1; 1876: 30. — Hutton 1879: 340. — Filhol 1886: 383. — Wilson 1907: 65. — Chilton 1909: 608; 1911: 291. — Thomson 1913: 237. — Rathbun 1918: 3. — Thomson & Anderton 1921: 98, 2 figs. — Chilton & Bennett 1929: 754. — Young 1929: 151. — Powell 1937: 375, 377, 387. — Richardson 1949a: 31, fig. 1. — Ralph & Yaldwyn 1956: 74, fig. 42. — Yaldwyn 1958: 125. — Dell 1960: 5: 1963a: 44; 1963b:253; 1968: 227. — Stephenson 1962: 315; 1972: 21. — Bennett 1964: 65, fig. 130. — Inoue et al. 1968: 135, 137. — Takeda & Miyake 1969: 165, figs 4c, d. — Dell et al. 1970: 54, figs. 1b, 2b, e, 3c, d, pl. 2. — Dawson & Yaldwyn 1974: 47. — Fenwick 1978: 207. — Probert et al. 1979: 381. — Willan 1981: 230. — Rainer 1981: 37. — Probert & Wilson 1984: 390. — Wear & Fielder 1985: 49, figs 126–129. — McLay 1988: 220, fig. 49a–d. — Takeda 1990: 372, colour photograph. — Ahyong et al. 2007: 170, colour photograph.

Material Examined. Wellington Harbour 41°17’S, 174°46’E: WT046CB, 1 female, 5.5 mm x 4.7 mm

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(NIWA 48737); WLG604CB, 1 male, 10.9 mm x 8.6 mm (NIWA 48738); WLG612CB, 1 female, 7.9 mm x 6.7 mm (NIWA 48739).

FIGURE 23. Nectocarcinus antarcticus (Hombron & Jacquinot, 1846); a. male 8.9 x 7.6 mm (2NSN167CB); b. immature female 18.9 x 15.3 mm (2TRG050CB).

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FIGURE 24. Nectocarcinus antarcticus (Hombron & Jacquinot, 1846); a. mature female 28.0 x 21.9 mm (Lyttelton Harbour, 2005); b. mature male 53.8 x 40.7 mm (DUD035CB).

Tauranga Harbour 37°39’S, 176°10’E: 1 female, 11.6 mm x 9.3 mm, TRG800CB, 23 Feb. 2004; 1 female, 7.4 mm x 6.3 mm, TRG801CB, 24 Feb. 2004 (NIWA 48740); 1 female, 15.1 mm x 12.2 mm,

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TRG838CB, 25 Feb. 2004; 1 female, 18.9 mm x 15.3 mm, 37°39’S, 176°10’E, 2TRG050CB, 5 Apr. 2005. Nelson Harbour 41°15’S, 173°18’E: 1 male, 8.9 mm x 7.6 mm, 2NSN167CB, 17 Dec. 2004 (NIWA 48742). Lyttelton Harbour 43°37’S, 172°44’E: 1 female, 28 mm x 21.9 mm, 3 m, personal collection. Dunedin Harbour45°53’S, 170°31’E: 1 male, 53.8 mm x 40.7 mm, 14 m, DUD035CB, 26 Feb. 2003. Remarks. Initially the small specimens shown in Fig. 22a, b were considered to possibly be a new species of Nectocarcinus, but after examining a whole range of sizes it became apparent that these were juvenile stages of N. antarcticus because there were no adult stages that matched the juvenile features. The most parsimonious hypothesis is that they all belong to the same species. The most detailed description of N. antarcticus is to be found in Dell et al. (1970), but this is based on male and female crabs with CW>70 mm and so does not encompass juvenile differences. The main changes to morphology as size increases are: 1) in small crabs the anterolateral teeth (and post-orbital tooth) are prominent, acute, with last tooth directed laterally whereas in large crabs the anterolateral teeth are relatively inconspicuous and are directed anteriorly; 2) curvature of the anterolateral margin becomes much stronger with increasing size; 3) while the pereopods of all specimens are fringed with long setae, body tomentum only becomes evident with increasing size and maturation. Figs 22, 23 are a series of specimens increasing in size to show changes in the proportions of the carapace. In both males and females the ratio of CW/CL increases from around 1.17 (CW = 5.5mm) to 1.32 (72.7 mm). In females the ratio suddenly changes to around 1.27 upon reaching sexual maturity. Dell et al. (1970) noted that N. antarcticus females achieve maturity over a range of sizes that seem to decrease northwards. Conversely maximum size attained by both sexes seems to increase southwards to the southern off-shore islands. Ontogenetic changes in morphology are also seen in the two Australian species N. integrifrons (Latreille, 1825) and N. tuberculosus A. Milne-Edwards, 1860 (see Dell et al. 1970). Dell et al. (1970) provide a key to the six species of Nectocarcinus, but the two New Zealand species are easily distinguished because N. bennetti Takeda & Miyake, 1969 is dark red-purple and iridescent, lacks body tomentum and the male G1 is strongly curved, while N. antarcticus is red, non-iridescent, tomentose and the male G1 is not curved. Distribution. Endemic to New Zealand. The specific name “antarcticus” is probably based on the fact that the type locality was the Auckland Is, a remote locality south of New Zealand. However, the name is not a good indicator of the distribution of this species which occurs along the whole length of New Zealand between 34°S – 51°S, and a lot further north than N. bennetti which is restricted to 44°S – 53°S. The northern limit of the N. bennetti is Stewart I. and the two species overlap from 44°S – 51°S. Depth range 0–550 m.

Family Plagusiidae Dana, 1851

Percnon Gistel, 1848

Percnon planissimum (Herbst, 1804) Figs 25a, b, 26a, b.

Cancer planissimus Herbst, 1804: 3, pl. 59, fig. 3. Percnon planissimum — Chilton & Bennett 1929: 774. — Richardson 1949a: 32 (key), pl. 1, fig. 12. — Barnard 1950: 138, fig. i, j. — Tweedie 1950: 136, fig. 4d. — Edmondson 1959: 197, fig. 25c, 27a–c. — Forest & Guinot 1961: 163. — Davie 2002: 439. Leiolophus planissimus — Miers 1876: 46. Percnon pilimanus — Chilton 1911: 559. [not Acanthopus pilimanus A. Milne-Edwards, 1873 = P. affine (H. Milne Edwards, 1853)] Percnon affine — McLay 1989: 106, fig. 1A–D. [not Acanthopus affinis H. Milne Edwards, 1853]

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Material Examined. Kermadec Is: AQ2789, 2 males, 15.2 mm x 16.6 mm, 18.4 mm x 19.9 mm, 2 females, 18.3 mm x 19.7 mm, 22.2 mm x 23.2 mm, coll. W.R.B. Oliver, 1908 (reported by Chilton, 1911 as P. pilimanus and labelled as such); AQ2397, 1 male, 29.2 mm x 30.8 mm, 3 females, 22.8 mm x 23.6 mm, 24.2 mm x 25.0 mm, 29.1 mm x 30.0 mm, coll. by W.R.B. Oliver, 1908 (reported by Chilton, 1911 as P. pilimanus and labelled as such); AQ2445, 1 female, 15.2 mm x 16.8 mm, coll. J.H. Sorensen Denham Bay, Raoul I., May, 1944; 3 males, 13.0 mm x 14.2 mm, 18.9 mm x 20.3 mm, 24.8 mm x 26.5 mm, 4 ovigerous females, 17.4 mm x 19.0 mm, 20.5 mm x 22.2 mm, 21.8 mm x 22.4 mm, 21.9 mm x 23.8 mm, 1 female, 12.3 mm x 13.7 mm, Oneraki Beach, Raoul I., coll. by DOC staff, 4 Oct. 2003. New Zealand: Tutukaka, at Poor Knights I: 2 photos taken by dive guide, Dave Abbott, CW about 19.5 mm, (cheliped size suggests a female), 35° 28′S, 174° 44′E, 4 m, among boulders, 24 Aug. 2001 (see Fig. 26a, b). Remarks. Chilton (1911) reported “two males and several females from Sunday I.” as Percnon pilimanus (A Milne-Edwards, 1873), but he debated at some length whether they should not be called P. planissimus. I think that these specimens (held as two lots in the Canterbury Museum, AQ2397, AQ2798) should be referred to P. planissimum because they have similar CW/CL ratio and exhibit the same patterns on the dorsal surface of the carapace (see Crosnier 1965: fig. 135). It should be noted that there is some variation in the length of the two spines that terminate the rostrum in P. planissimum with much longer spines in some specimens than is figured by Crosnier (1965: fig. 135). The occurrence of Percnon planissimum around the New Zealand mainland has always been uncertain. Chilton & Bennett (1929) included it on the authority of Hutton (1882) who had originally listed it as being one which should be deleted from the list of species to be excluded from the New Zealand fauna, but in his personal copy of the list the name was struck off. A double negative, making a positive, is hardly strong evidence for its presence on the mainland! The only substantial evidence seems to be from Miers (1876) who stated that there was a specimen from New Zealand in the Paris Museum collection. It has not been possible to confirm or not whether the specimen is in Paris or whether it belongs to this species. The only evidence that P. planissimum is present are some photos (see Fig. 25a, b) taken by divers in 2001 at the Poor Knights Is off the east coast of the North Is. They were seen again several times between 2001 and 2006. It is not possible to compare the CW/CL ratio with the Kermadec specimens, because of the angle from which the dorsal view photo (Fig. 25a) has been taken, but the length/width ratio of the merus of the second pereopod is the same (about 4.0) as P. planissimum. The pattern of colours on the carapace and banding on the legs are similar (see Fig. 24) even though the actual colours are not exactly the same. The photos (Fig. 25a, b) were taken in situ using colour film at a depth of around 4–5 m so that would’ve influenced their apparent colour. Takeda & Webber (2006) included Percnon affine in their list of Kermadec crabs on the assumption that Chilton (1911) had correctly identified his specimens as P. pilimanus (A. Milne-Edwards, 1873), a species that after comparing the types, Crosnier (1965) argued was synonymous with P. affine (H. Milne Edwards, 1853). However, P. affine is not part of the fauna of the Kermadec Is because Chilton’s specimens belong to P. planissimum. Chilton (1911: 559–560) reported “P. pilimanus” as being very common among rocks near the low tide mark. As to the habits of these crabs, it is worthwhile quoting the comments of the collector: “Mr. Oliver makes the following observations on the habits of this species: ‘Fairly common among rocks near low-tide mark. Very quick in its habits. Its colour somewhat resembles the rock, on which it stays perfectly still, but when any one approaches too near it darts into the water. When, after continued westerly winds, sand was driven ashore so as to bury the boulders on the north coast of Sunday I. to about half-tide mark, thousands of these crabs, retreating before the encroaching sand, congregated in heaps amongst the rocks near shore until the sand was washed away again.’ DOC staff who collected specimens from Raoul I. in 2003 found them “washed up dead and dying on sand/rocks near hot pools – many were cooked in hot pools”. It seems that these crabs were victims of volcanic activity. An ovigerous P. planissimum from Raoul I. reported above (21.9 mm x 23.8 mm) carried around 2000 very small eggs (diameter = 0.30–0.35 mm). As might be expected for a species with such a wide distribution,

50 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. it is likely that the larval stages spend a long period in the plankton during which they are transported considerable distances. The eggs are carried in two distinct clutches (left and right) in the narrow brood chamber beneath the flattened abdomen. The gonopores are not operculate.

FIGURE 25. Percnon planissimum (Herbst, 1804) Oneraki Beach, Raoul I., Kermadec Is: a. male 24.8 x 26.5 mm; b. ovigerous female 21.9 x 23.8 mm.

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FIGURE 26. Percnon planissimum (Herbst, 1804): ?female, CW about 19.5 mm (Poor Knights Island): a. dorsal view, b. frontal view (scanned photos).

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Distribution. Red Sea, East Africa, Cocos-Keeling Is, Japan, southeast Australia, Middleton and Elizabeth Reefs (Tasman Sea), Niue, Tahiti (French Polynesia), Hawaiian Is, and now the Kermadec Is and New Zealand. P. planissimum is a very widespread Indo-West Pacific species. Inhabits rocky substrates in the depth range 0–5 m.

Family Ocypodidae Rafinesque, 1815

Ocypode Weber, 1795

Ocypode pallidula Jacquinot, in Hombron & Jacquinot, 1846 Figs 27a, b, 28a, b, 29a, b.

Ocypode pallidula Jacquinot, in Hombron & Jacquinot, 1846: pl. 6, fig. 1 — Sakai & Turkay 1976: 87, figs 14, 15. — Jones 1988: 36 (Table 1). — Takeda & Webber 2006: 223, fig. 17A–C. — Davie 2002: 358. Ocypode laevis Dana, 1852: 325: 1855: 325, pl. 20, fig. 2. Ocypode cordimana — Jacquinot & Lucas 1853: 65. [Not Ocypode cordimanus Latreille, 1818.] Ocypoda kuhlii — Chilton 1911: 561. [Not Ocypode kuhlii De Haan, 1835.]

Material Examined. Kermadec Is: AQ2422, 1 female, 22.3 mm x 20.6 mm, coll. W.R.B. Oliver, 1908, Canterbury Museum (reported by Chilton, 1911); AQ2376, 1 juv., 6.5 mm x 5.3 mm, 1 female, 7.3 mm x 6.2 mm, 1 male, 9.6 mm x 8.4 mm, coll. J.H. Sorensen, sandy beach, Raoul I., 2 May 1944. Oneraki Beach, Raoul I.: 2 males, 19.0 mm x 16.4 mm, 19.4 mm x 16.7 mm, coll. by Marion Rhodes, 18 Apr. 2003; 1 female, 18.4 mm x 15.8 mm, coll. by Erica Cammack, 20 June 2003; 1 male, 20.5 mm x 18.6 mm, 1 female, 22.5 mm x 20.3 mm, coll. Kala Sivaguru, DOC, 7 Aug. 2007. Norfolk I.: AQ2420, 2 males, 23.1 mm x 20.3 mm, 24.1 mm x 20.8 mm, 29° 00′S, 168° 00′E, coll. by W.R.B. Oliver, 1913. New Zealand: 1 male, 19.4 mm x 17.4 mm, Pakiri Beach, Leigh Marine Reserve, Auckland, 36° 17′S, 174° 48′E, 11 May 2001. Remarks. Ghost crabs are usually found in intertidal burrows on sandy beaches. The main features used to identify the species of Ocypode are the presence/absence of ocular stylets, presence/absence of “brushes of hairs” on the walking legs and the number, shape and arrangement of striae on the stridulatory crest found on the inside of the major cheliped palm. Chilton (1911) reported Ocypoda kuhlii De Haan from “Sunday I.” and “Thursday I.” (old names for islands in the Kermadec Group). These specimens, deposited in the Canterbury Museum, have been re-examined and found to be Ocypode pallidula and they agree very well with ghost crabs of the same species reported by Takeda & Webber (2006). The stridulatory ridge of O. kuhlii is made up of 7–17 round or ovoid tubercles whereas in O. pallidula there are 25–40 evenly spaced fine striae (Jones, 1988). The number of striae on the chelipeds of the material examined here was 20–25 for males (mean = 23) and 15–19 for females (mean = 17). Sexual dimorphism of the stridulatory ridge has not been reported before, but it is probably the result of positive allometry of the male cheliped. Takeda & Webber (2006) figure the stridulatory ridge of O. pallidula (their Fig. 17A) which has around 22 evenly spaced striae on the inside of the propodus. The range in number of striae given by Jones (1988) for O. pallidula, needs to be revised from, 25–40 to 15–40, so as to accommodate females. The number of striae is still one of the best characters to use, along with a few others, to identify ghost crabs. The majority (7/11) of crabs reported here have the major cheliped on the left instead of the right hand side. Sound is produced underground in the burrow (Hicks etal, 1984) by vertical movements of the palm rubbing the stridulatory ridge against the plectrum, which is a pair of closely spaced ridges on the ventral margin of the cheliped ischium. Whether on the left or on the right, the minor cheliped never had any striae on the inner surface of the propodus. Immature crabs lack a stridulatory ridge. There is only one species of ghost crab on the Kermadec Is and O. kuhlii should be omitted from the

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FIGURE 27. Ocypode pallidula Jacquinot, in Hombron & Jacquinot, 1846: female 22.5 mm x 20.3 mm, carapace cracked on left hand side (Oneraki Beach, Raoul I., Kermadec Is): a. dorsal view, b. ventral view.

The discovery of one adult male specimen of O. pallidula on the beach at Leigh Marine Reserve is the first reliable record of a ghost crab from the New Zealand mainland. Two explanations seem possible: either it is a natural occurrence resulting from recruitment of larvae produced at perhaps Norfolk I. or the Kermadec Is

54 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. or it is an escapee from a diver’s bag left on the beach. The single record of a mature-sized crab found in a burrow on the beach suggests that it may well be an escapee, which implies that the survived transport to New Zealand in amongst diving gear. Since ghost crabs are semi-terrestrial it is not impossible that it could have survived several days inside a damp bag. If the occurrence was natural we would have expected to find several crabs in burrows on the beach. Given that level of foot traffic along the beach and the nearby marine laboratory, it seems likely that someone would have noticed more burrows if they were there, so the one-off event suggests that there has been a breach of biosecurity and the crab has escaped while the owner was off diving in the marine reserve. A natural recruit to the beach would have had to have been there for perhaps a

FIGURE 28. Ocypode pallidula Jacquinot, in Hombron & Jacquinot, 1846: female 22.5 x 20.3 mm, carapace cracked on left hand side (Oneraki Beach, Raoul I., Kermadec Is): a. frontal view; male 20.7 x 18.6 mm (Oneraki Beach, Raoul I., Kermadec Is): b. ventral view.

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FIGURE 29. Ocypode pallidula Jacquinot, in Hombron & Jacquinot, 1846: a. male 20.7 x 18.6 mm (Oneraki Beach, Raoul I., Kermadec Is), frontal view; stridulatory ridges of b. male (20.7 x 18.6 mm) and c. female 22.5 x 20.3 mm) cheliped palms. Scale bar = 2 mm. year before reaching adulthood and would have created many burrows during that time. Subsequent searches after the initial discovery have not revealed the presence of any more crabs. Perhaps Biosecurity staff should

56 · Zootaxa 2111 © 2009 Magnolia Press McLAY TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. pay more attention to dive bags and their contents brought into New Zealand by passengers. Given the very wide distribution of O. pallidula (see below) it is difficult to say exactly where the interloper may have come from. The nearest sources frequented by divers would be Norfolk I., Cook Is and Queensland, Australia. Distribution. Madagascar, Mauritius, Moluccas, Timor, Celebes, New Guinea, Australia (Queensland, Coral Sea), Norfolk I., Kermadec Is, Cook Is, French Polynesia (type locality: Gambier Is, Tuamotu Archipelago), Johnston I, Laysan I and Midway. It is clearly a widespread Indo-West Pacific species. Ocypode pallidula inhabits the intertidal zone on sandy shores. Brachyuran Fauna of the Kermadec Is The first compilation of the crab fauna of the Kermadec Is, north of mainland New Zealand, was done by Chilton (1911) who listed 25 species. Almost 100 years was to pass before this number was revised: Takeda & Webber (2006) were able to list 57 species, including 5 that were new. Seven additional species, 4 Leucosiidae, 2 and 1 Xanthidae, await future publication, so that altogether the fauna could include 64 species. However Takeda & Webber did not view all the species in their list and relied upon identifications made by Chilton (1911) for some records. I have examined Chilton’s specimens, held by the Canterbury Museum, and I am able to correct the list that he produced and thus the list provided by Takeda & Webber (2006). Three species should be omitted: “Ovalipes bipustulatus” as used by Chilton in that context refers to O. elongatus, and similarly, Percnon affine refers to P. planissimum, and Ocypode kuhlii refers to O. pallidula. McLay (2007) recorded 2 new families associated with hydro-thermal vents near the Kermadecs, represented by Gandalfus puia McLay, 2007 ( Williams, 1980) and Xenograpsus ngatama McLay, 2007 (Xenograpsidae Ng, Davie, Schubart & Ng, 2007). From the same collections Ng & McLay (2007) discovered 2 new species, Euryxanthops chiltoni Ng & McLay, 2007 (Xanthidae Macleay, 1838) and Medaeops serenei Ng & McLay, 2007 (Xanthidae Macleay, 1838) from the Macauley Caldera north of the Kermadec Is. In the present paper one new species, Rochina ahyongi sp. nov. is described from the Rumble Seamount near the Kermadec Is and Thalamita danae is added to the Kermadec fauna. Not all of Chilton’s collection in the Canterbury Museum has been identified to species and so several new records and/or species will be added. Thus at the moment the number of species known from the Kermadec Is area sits at 67.

Acknowledgements

I wish to thank Steve O’Shea, formerly at NIWA (now at AUT, Auckland), for sending me some of the specimens and the photo of Mursia australiensis, Anne-Nina Loertz for access to the NIWA collections, and Ashley Rowden, Kareen Schnabel and Andrew Hosie provided valuable help to sort out the location details for the samples. Shane Ahyong also provided valuable assistance and advice. I am grateful to Peter Castro (California State Polytechnic University) for confirming the identity of Pseudopalicus undulatus and for editorial help. I would also like to thank Anthony Saville and Simon Pollard Canterbury Museum, Christchurch for loan of specimens and access to their collections. Alain Crosnier (MNHN) was kind enough to send me the specimen of Dynomene pilumnoides from Fiji. Dave Abbott took the photos of Percnon planissimum. The help of Karla Sivaguru (Department of Conservation, Auckland) was invaluable for obtaining recent crab specimens from Raoul I, Kermadec Is.

References

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