Zoology Series ==" wy THE NATURAL HISTORY MUSEUM VOLUME 68 NUMBER1 27 JUNE 2002 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology Prof P.S. Rainbow Editor of Bulletin: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review before acceptance. 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Copyright © 2002 The Natural History Museum ELECTRONIC ACCESS _ This journal is included in the Cambridge Journals Online service which can be found at: http://journals.cambridge.org For further information on other Press titles access http://uk.cambridge.org or http://us.cambridge.org World list abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) ISSN 0968-0470 The Natural History Museum Zoology Series Cromwell Road Vol. 68, No. 1, pp. 1-50 London SW7 5BD Issued 27 June 2002 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Lond. (Zool.) 68(1): 1-11 Issued 27 June 2002 Another variation on the gymnure theme: description of a new species of Hylomys (Lipotyphla, Erinaceidae, Galericinae). PAULINA D. JENKINS Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD MARK F. ROBINSON Waterway Conservation & Regeneration, British Waterways, Llanthony Warehouse, Gloucester Docks, Gloucester, GLI 2EJ SYNOPSIS. A new species of Hylomys from Lao Peoples Democratic Republic is described, based on morphological comparisons with other members of the subfamily Galericinae. The relationships revealed by a phylogenetic analysis are discussed and compared with those of a previous published analysis. INTRODUCTION The Family Erinaceidae is divided into two subfamilies: the wide- spread Erinaceinae (hedgehogs) occuring in Africa, Europe and Asia, and the Galericinae (moonrats and gymnures), which is con- fined to southeast Asia, Indonesia and the Philippines. There has been considerable disagreement over the correct name to apply to the subfamily of moonrats and gymnures, summarised by Frost er al. (1991), who favoured the use of Hylomyinae. McKenna & Bell (1997) pointed out however, that the use of Galericini as a tribal name by Butler (1948) had been accepted by many subsequent writers, particularly palaeontologists, and was therefore the appro- priate name to use. In this paper we follow McKenna & Bell (1997) in using the name Galericinae. Most authors up to and including Corbet (1988), considered that the Galericinae includes five genera: Echinosorex Blainville, 1838, Hylomys Miiller, 1840, Neotetracus Trouessart, 1909, Neohylomys Shaw & Wong, 1959 and Podogymnura Mearns, 1905, all but the latter being monotypic. In their revision of the family Erinaceidae, Frost et al.(1991) con- cluded that there are only three valid genera within the Galericinae: Echinosorex, Podogymnura and Hylomys. They accepted Hylomys as a rather variable but nevertheless monophyletic genus, although they conceded that there was evidence to support the retention of Neotetracus and Neohylomys as subgenera. The genus Hylomys is widely distributed in southeast Asia and Indonesia. Hylomys suillus Miiller, 1840 occurs in Malaysia, Indo- nesia, Thailand, Vietnam, Cambodia, Lao Peoples Democratic Republic (PDR), Myanmar and southern PDR China; in Lao PDR it has been recorded from Phongsali, Xiangkhouang, Vientiane and Dong Hua Sao National Biodiversity Conservation Area (NBCA) (Robinson, 1999). A number of different subspecies have been attributed to H. suillus and the biochemical and metrical variation within this species was examined by Ruedi ef al. (1994). They recognised that much of the high level of variation could be attrib- uted to the geographical and altitudinal isolation of the named forms but demonstrated that one of these taxa, H. parvus Robinson & Kloss, 1916, merited specific status. Hylomys sinensis (Trouessart, 1909) occurs from southern China to Myanmar and northern Viet- nam; it has not been recorded from Lao PDR but is likely to occur in those areas adjacent to northern Vietnam, whence it is recorded by Osgood (1932). Hylomys hainanensis (Shaw & Wong, 1959) is © The Natural History Museum, 2002 restricted to Hainan Island, PDR China and H. parvus is known only from Sumatra, Indonesia. Another geographically isolated undescribed gymnure has been discovered recently from a region of limestone karst in the Lao PDR. While sharing many characters with other species of Hylomys this new species also differs markedly from its congeners and, furthermore, shares some features with geographically remote species of Podogymnura. The new taxon has been compared in particular with specimens of H. suillus, although there is no indication that the two species occur sympatrically, and also with H. sinensis, H. parvus and, in the absence of specimens, with the original description of H. hainanensis and the figures of the skull of this species in Frost er al. (1991). In addition comparisons were made with the other genera of Galericinae: Echinosorex gymnura (Raffles, 1822) from Malaysia and Indonesia, and Podogymnura truei Mearns, 1905 from the Philippines. In order to assess phylogenetic relationships, both the new taxon and H. parvus were analysed using the criteria employed by Frost et al. (1991). MATERIAL AND METHODS Comparative material was examined from the collections of the Natural History Museum (BMNH), London (formerly the British Museum (Natural History)), the American Museum of Natural History, New York (AMNH), the Muséum National d’ Histoire Naturelle, Paris (MNHN) and the Thailand Institute of Scientific and Technological Research, Bangkok (TISTR), as listed in Table 1. All measurements are in millimetres and were taken using digital calipers. Cranial and dental nomenclature follows Butler (1948), Novacek (1986), Frost et al. (1991) and Gould (1995). Dental notations are indicated in the text in the following manner, with premaxillary and maxillary teeth denoted by uppercase letters and mandibular teeth by lowercase: incisor (I/1), canine (C/c), premolar (P/p), molar (M/m), thus P3 refers to the third upper premolar, i2 to the second lower incisor. PHYLOGENETIC ANALYSIS Cranial, dental, skeletal and external characters were scored for the new species and H. parvus according to the character transformation 2 series employed by Frost et al. (1991: 3-15) and added to the character matrix shown in Frost et al. (1991: appendix 2) see Table 2. Branch and bound analyses were performed using Paup 4.0ba (Swofford, 1999) set at maximum parsimony, with a maximum trees setting of 1000 and all characters treated as unordered and of equal weight. Bootstrap analyses (Felsenstein, 1985) were made to pro- vide an assessments of confidence limits of nodes, with 1000 replicates of 100 random addition sequence replicates. Bremer support indices were calculated by increasing the upper bound of the shortest tree by one step, repeating the branch and bound analysis and producing a strict consensus tree; the process was repeated, progressively increasing the length of the suboptimal cladograms by a single step until all clades of interest no longer occurred on the consensus tree; the level at which each node collapsed was recorded (Kitching et al., 1998). Both accelerated (ACCTRAN) and delayed (DELTRAN) optimizations were used to map character evolution. The trees obtained were compared with those in Frost et al. (1991) and the results of the analysis are given below. RESULTS Hylomys megalotis, sp. nov. HOLOTYPE. BMNH 1999.44 (field number 5/99) male, body in alcohol, skull extracted. Collected 15 January 1999 by M. F. Robinson. TYPE LOCALITY. Environs of Ban Muang and Ban Doy, c 18 km North of Thakhek, Thakhek district, Khammouan Limestone National Biodiversity Conservation Area, Khammouan Province, Lao Peoples Democratic Republic, 17°33'15"N 104°49'30"E. Habitat: steep slopes around the base of massive limestone karst, covered in rock and large boulders, with an underlying soil base and heavily degraded mixed deciduous forest, scrub and bamboo. Low lying areas away from the karst had been cleared for cultivation of paddy rice. PARATYPES. BMNH 1999.45 (field number 14/99) collected 16 January 1999; 1999.46 (field number 15/99) and 1999.48 (field number 17/99) collected 17 January 99, females, bodies in alcohol, skulls extracted; 1999.47 (field number 16/99) collected 17 January 1999, male, skin and internal organs in alcohol, skull and skeleton. All specimens were collected by M.F. Robinson from the same locality as the holotype. DIAGNOSIS Ears large, rhinarium elongated; first and fifth digits of forefeet long, claws long, cheiridia large and rounded; cheiridia on hindfeet large, soles naked; pre-anal gland with single opening. Skull with posterior region of nasals extending to level of antorbital rim; maxilla and parietal widely separated by frontal in supraorbital region; long grooves for palatine artery present in palate; anterior palatine fo- ramina anterior to maxillary palatine suture; antorbital fossa shallow; nasolabialis fossa shallow; posteroventral maxillary process of zygoma distinct; antero-ventral process of alisphenoid present. Den- tition robust. Third upper premolar (P3) large with well developed lingual cusp and three roots. Neural spine of axis low. DESCRIPTION Medium sized Hylomys with a long tail, approximately 75% of head and body length. Pelage grey, long, soft and very fine, lacking flattened spinous hairs; individual hairs grey for most of their length, then buff with buff or black tips. Dorsal region of rhinarium narrow, elongate posteriorly; ears prominent, very large, rounded. First and P.D. JENKINS AND M.F. ROBINSON fifth digits of forefeet lengthened, claws long and moderately stout; sole and tarsal region of hindfeet naked, cheiridia large. Pre-anal gland with single opening immediately posterior to cloaca. Two pairs of inguinal mammae present. Skull elongate, moderately slender and somewhat flattened in appearance (see Figs. 1—2); dorsal profile more or less straight, showing a gradual increase in height from anterior of rostrum to braincase. Rostrum long, slender and moderately shallow, nasals long extending posteriorly to, or slightly beyond, level of antorbital rim; posterodorsal region of premaxilla widely separated from anterodorsal region of frontal by maxilla; interorbital region moder- ately narrow; supraorbital processes of frontals scarcely evident; frontals anteriorly depressed in midline; supraorbital region of frontals broad, so that the maxillaries are widely separated from the parietals; parietals extend anteriorly in supraorbital and orbital region but do not form an anterior process; supraorbital foramen present in dorso-orbital region of frontals; orbital region of maxilla broad, forming major portion of the anterior region of the orbit; orbital region of frontal constricted anteriorly by maxilla, posteriorly by parietal; orbitosphenoid anteroposteriorly expanded, optic foramen posteromedially positioned, anterodorsal to, and moder- ately well separated from, the suboptical foramen and from the ethmoid foramen (see Fig. 3); crest present leading from anterior alisphenoid diagonally across orbitosphenoid, partially obscuring optic, suboptic and sphenorbital foramina in lateral view; alisphe- noid dorsoventrally compressed, fusiform anteroventral process of alisphenoid present, well marked alisphenoid canal present; brain- case low and scarcely domed, lambdoid crest moderately well-developed laterally, low medially; mastoid large, slightly in- flated; paraoccipital process small; infraorbital foramen dorsal to P4; antorbital or prelacrimal flange present only as a low ridge; shallow antorbital fossa on anterior surface of zygoma; nasolabialis fossa shallow; maxillary component of zygoma narrow with long, slender posteroventral process ventral to well marked long jugal, slender anterodorsal process of squamosal portion of zygoma over- lying jugal; palate with paired maxillary foramina level with P2 and anterior of P3, small paired anterior palatal foramina, lying anterior to the suture between maxilla and palatine; palatal spine absent; basioccipital narrow with ridge in midline, tympanic wing of basioc- cipital slightly inflated. Mandible with deep, moderately broad coronoid process; mental foramen below p3. Dental formula: 3/3 1/1 4/4 3/3 = 22. Dentition robust (see Figs. 1— 2). First upper incisor robust, distostyle present; I2 and [3 sub-triangular, anteroflexed, distostyle present, 13 approximately half size of 12; C with anterior basal cusp and distostyle and two roots; Pl and P2 subequal in height, P2 longer than P1, both with anterior basal cusp and distostyle, P1 with two fused roots, P2 with two roots; P3 large, subequal in height to C, lingual cusp (protocone according to Gould, 1995) well developed, three roots present; P4, M1 and M2 quadrate in shape, parastyle well developed, meta- conule present on M1 and M2; M3 subtriangular in shape, with well developed parastyle and hypocone and metacone distinct in unworn dentition. First lower incisor larger than i2, both semi-procumbent, i2 larger than 13, which is anteroflexed with hypoconulid present; c anteroflexed, greater in height than i3 and p1; p1 and p2 subequal in height and both with a single root; p3 larger with two roots; p4 with well developed paraconid and talonid; m1—m3 with well developed paraconids, m3 less than half size of m1. ETYMOLOGY The name of the new species is derived from the Greek p¢éyac (megas), large; ®t6c (otos), ear; the ears are large in comparison with those of other species of Hylomys. A NEW SPECIES OF HYLOMYS MUPUARRAROUADOOANODAUOOADOURDORRUORRNORRNORRUUORUOURUUNRUOUNONUDOURUUDRDORURUORRUNURUNURUNURUORRRUND| 10cm Fig. 1 Crania from left to right of dorsal view of mandible and skull, ventral view of skull, left lateral view of skull and mandible. Top row: Hylomys megalotis BMNH 1999.47; second row: Hylomys suillus BMNH 1962.711; third row: Hylomys sinensis BMNH 1911.2.1.20; fourth row: Hylomys parvus BMNH 1919.11.8.12. \oS) P.D. JENKINS AND M.F. ROBINSON Fig. 2 Lateral view of anterior of skull, mandible and dentition of Hylomys megalotis BMNH 1999.44. Scale = 1 mm. COMPARISON WITH OTHER TAXA The new species is readily distinguished in external appearance from all other species of Hylomys. It is similar in body size but with a considerably longer tail and larger ears (see Table 3). Tail 65-74 % of head and body length in H. megalotis, 51-63 % in H. sinensis, 27-31 % in H. hainanensis, and very short (< 25 %) in H. parvus and H. suillus. The lack of flattened spinous hairs in the pelage distinguishes H. megalotis from H. suillus and H. sinensis. The rhinarium is more extensive posteriorly than in H. sinensis but more elongate and narrower than in H. suillus. The claws and first and fifth digits of the forefeet of H. sinensis, H. suillus and H. parvus are not lengthened as in H. megalotis and the cheiridia are smaller than in H. megalotis. The sole and tarsal region of the hindfoot are naked in H. megalotis, differing from the haired soles found in H. sinensis and H. suillus. The paired pre-anal glands are midway between the cloaca and the anus in H. sinensis and close to the anus in H. suillus, so differing from that of H. megalotis, in which the single opening is positioned immediately posterior to the cloaca. The skull of H. megalotis is more elongate in appearance than any of the other species of Hylomys; it is longer, with a longer, narrower rostrum, longer upper toothrow and the braincase is shallower relative to its breadth (see (see Fig. 1 and Table 3). The posterodorsal region of the premaxilla is widely separated from the anterodorsal region of the frontal by the maxilla in H. megalotis, narrowly separated in H. hainanensis but in contact or nearly in contact in H. sinensis, H. suillus and H. parvus (see Table 4). As in H. sinensis, but unlike other species of Hylomys, the posteriormost portion of the nasals in the new species extend to the level of the antorbital rim. Hylomys megalotis has a shallow antorbital fossa unlike the moderately deep fossa of H. sinensis and H. parvus, and the deep fossa in H. suillus. The zygoma of H. megalotis differs from all other species of Hylomys: the maxilla is considerably narrower, the nasolabialis fossa shallower and the jugal more extensive. As in Podogymnura aureospinula and Echinosorex, a distinct posteroventral process is present on the maxillary region of the zygoma of H. megalotis, indistinct in H. parvus but absent in H. suillus, H. sinensis and H. hainanensis. The supraorbital process of the frontal of H. megalotis is poorly defined and blunt, the anterior process of the parietal absent and the parietal is widely separated from the maxilla by the frontal in the supraorbital region, as in Podogymnura and Echinosorex; in H. parvus the supraorbital process of the frontal is poorly defined and blunt but the anterior process of the parietal is short but distinct, narrowly separated from the maxilla by the frontal; in A. suillus, H. hainanensis and H. sinensis the anterior process of the parietal is distinct, scarcely separated from the maxilla and contributing to the well marked supraorbital process of the frontal. The optic and suboptic foramina are well separated in H. megalotis but lie close together in H. suillus, H. sinensis and H. parvus. An anteroventral process of the alisphenoid is present in H. megalotis, unlike all other Hylomys and Podogymnura. As in Podogymnura and Echinosorex, the palatine foramina are small and positioned anterior to the maxillary/palatine suture in H. megalotis and long grooves for the palatine artery are present in the palate, whereas the elongated palatine foramina in other species of Hylomys lie at the maxillary/ A NEW SPECIES OF HYLOMYS FR MX CORF SPALF al PR SQ OPTF FOV SOPF Fig.3 Lateral view of left orbital region of Hylomys megalotis BMNH 1999.44. Scale = 1 mm. Abbreviations: AL — alisphenoid, ALIC — alisphenoid canal, CORF — cranio-orbital foramen, EF — ethmoid foramen, FOV — foramen ovale, FR — frontal, MX — maxilla, OPTF — optic foramen, OS — orbitosphenoid, PL — palatal, PR — parietal, PT — pterygoid, SOPF — suboptic foramen, SPALF — sphenopalatine foramen, SPORF — sphenorbital foramen, SQ — squamosal. palatine suture and the palatine artery grooves are small or indis- tinct. The anterior opening of infraorbital canal is dorsal to P3/P4 in most species of Hy/lomys but dorsal to P4/M1 in H. megalotis, H. parvus and Echinosorex, and yet more posteriorly positioned in Podogymnura. The dentition of H. megalotis is considerably more robust than that of any of the other species of Hylomys. The dental formula is the same as in H. suillus and H. parvus and these species are distin- guished from H. hainanensis, which lacks p1 and H. sinensis, which lacks P1 and pl. In H. megalotis and H. parvus, P2 has two roots, unlike the other species which have either one or two well fused roots. Unlike all other species which lack a lingual cusp, P3 of H. megalotis has a well developed lingual cusp as in Echinosorex, and this tooth is large with three roots as in Podogymnura and Echinosorex, p3 is larger than p2 with two roots as in H. parvus, Podogymnura and Echinosorex, while p3 is slightly smaller than p2 with one root in A. suillus, H. sinensis and H. hainanensis. RESULTS OF THE PHYLOGENETIC ANALYSIS Forty equally most parsimonious trees were retained in the branch and bound analysis, 141 steps in length, with a Consistency Index of 0.72, Retention Index of 0.93 and Rescaled Consistency Index of 0.66. In all most parsimonious trees the two subfamilies, Galericinae and Erinaceinae, readily segregated and the Galericinae further separated into two distinct groups: a clade comprising Echinosorex and Podogymnura, the other clade confined to Hylomys. Most of the variation found among all trees occurred within the Erinaceinae, since for the Galericinae twenty of the trees showed the configura- tion seen in Fig. 4a, while the remaining trees all showed the alternative arrangement for this subfamily (Fig. 4b). That part of the tree obtained by Frost ef al. (1991: Fig.9) for the Galericinae is illustrated as part of Fig. 4a. The strict consensus tree (see Fig. 5) revealed strong bootstrap support (97%) for the Galericinae and for aclade of H. suillus, H. sinensis and H. hainanensis, and this tritomy also had a high Bremer support index. There was moderate bootstrap support (83%) for a clade of H. suillus, H. sinensis, H. hainanensis and H. parvus, and a clade comprising all species in the genus Hylomys occurred in 77% of the replicates. The bootstrap support value for a clade of Echinosorex and Podogymnura was low at only 64%. Within the genus Hylomys, H. megalotis was basal to all other species. Clades with bootstrap support values less than 50%, respec- tively of H. sinensis and H. hainanensis (42%), and H. suillus and H. hainanensis (43%) were considered to be unresolved. The shared derived character transformations (synapomorphies) which were revealed by the analysis are recorded below, using the format of character number quoted from Frost er al. (1991) followed by character transformation state, where (0) equals the ancestral and (1) the derived character state. SYNAPOMORPHIES OF GALERICINAE: [8.1] Antorbital or pre-lacrimal flange: (0) not developed, lacrimal canal visible in lateral view; (1) developed so that the lacrimal canal is obscured in lateral view. CI 1.000. [10.2] Jugal size; (0) large, reaches lacrimal; (1) small, does not reach lacrimal; (2) vestigial, confined to lateral rim of zygoma; (3) absent. In ACCTRAN the transformation was from | — 2, in DELTRAN the change was from 3 — 2. CI 1.000. [62.1] P4 lingual roots: (0) one; (1) two unfused; (2) two fused. CI 1.000. This state occurs in all Galericinae but was shown only in DELTRAN. P.D. JENKINS AND M.F. ROBINSON gymnura aureospinula truei sinensis suillus hainanensis parvus megalotis Erinaceinae E. gymnura P. aureospinula P. Enwer H. sinensis H. hainanensis ishh Sibi Il Wats) lel Sen Vea AUNS) H. megalotis Erinaceinae Fig. 4 Comparison of trees obtained for the Galericinae. (a) One of twenty most parsimonious trees, all showing the same configuration for the Galericinae. Tree length 141 steps, with a Consistency Index of 0.72, a Retention Index of 0.93, and a Rescaled Consistency index of 0.66. The branching pattern on the left shows the results from the analysis of this study, that on the right is partially redrawn from Frost er al. (1991: fig. 9), restricted to show only the relationships within the Galericinae and is 128 steps in length with a Consistency Index of 0.76. (b) One of the remaining twenty most parsimonious trees, showing the alternate arrangement for the Galericinae. [66.1] M3 hypocone (see Frost. et al. 1991) or metastylar spur (see Gould, 1995): (0) absent or weak; (1) present, well developed on buccal side. CI 1.000. [69.1] Axis, posteroventral keel: (0) absent; (1) present. CI 1.000. [71.1] Scapula, metacromium process: (0) deltoid, amorphous pro- jection; (1) long, fusiform projection. CI 1.000. [72.1] Sacral vertebrae, neural spines: (0) not fused into continuous longitudinal plate; (1) fused into continuous longitudinal plate. CI 1.000. [73.1] Ischium, posterodorsal process (see Gould, 1995 for correc- tion of error by Frost et al. 1991): (0) not greatly elongated; (1) greatly elongated. CI 1.000. [74.1] Tibia, lateral flange on antero-superior margin: (0) absent or weakly present; (1) strongly developed. CI 1.000. SYNAPOMORPHY OF HYLOMYS: [19.1] Cranio-orbital foramen in the frontal: (0) closely associated or joined with the ethmoid foramen; (1) foramina widely separated. The terminology for this character is confusing. Frost er al. (1991) used the name ophthalmic foramen (which they attributed to Butler (1948) although this name could not be found in this paper), but Gould (1995: character 19) pointed out that this foramen had been misidentified by Butler and is the anterior opening for the superior ramus of the stapedial artery. Gould also referred to this foramen as A NEW SPECIES OF HYLOMYS £4 (97) >8 6 EB. gymnura P. aureospinula truei H. sinensis H. hainanensis Hee (Stes H. parvus H. megalotis Erinaceinae Fig. 5 Strict consensus of 40 most parsimonious trees. The branch lengths are shown above the branches followed by the bootstrap support values in parentheses, representing the percentage of trees containing the specified clades. The Bremer support indices are given below the branches. the sphenofrontal foramen and McDowell (1958) as the sinus canal. CI 1.000. SYNAPOMORPHY OF HYLOMYS SINENSIS, H. HAINANENSIS, H. SUILLUS AND H. PARVUS: [4.1] Size of palatal foramina: (0) small; (1) anterior foramina elongated posteriorly; (2) anterior foramina elongated to include middle palatine foramina. CI 1.000. SYNAPOMORPHIES OF HYLOMYS SINENSIS, H. HAINANENSIS, H. SUILLUS: {13.1] Supraorbital process of frontal on parietal/frontal suture: (0) absent or poorly defined; (1) sharp, well defined. CI 1.000. [16.1] Anterior process of parietal: (0) absent or very weak; (1) extends anteriorly along the supraorbital rim to form the base of the supraorbital process. CI 1.000. [57.1] p3: (0) two roots present, larger in size than p2; (1) one root present, nearly equal in size to p2; (2) absent. CI 1.000. The analysis found no autapomorphic characters to define H. megalotis but the following apomorphies for this species are recorded as follows: [1.1] Posteriormost extension of nasals: (0) anterior to the level of the antorbital rim; (1) medial or posterior to the level of the antorbital rim. CI 0.333. Homoplasious with H. sinensis but also with Erinaceinae. [5.1] Location of the anterior palatine foramina: (0) at the maxilla/ palatine suture; (1) anterior to the maxilla/palatine suture. CI 0.500. Shown only in DELTRAN, homoplasious with Echinosorex and Podogymnura. [17.1] Anterior process of alisphenoid: (0) absent; (1) present. CI 0.500. Homoplasious with Erinaceinae. This character, defined as a narrow, fusiform anterior process of the orbital wing of the alisphe- noid is, according to Frost et al. (1991), related to the location of the sphenopalatine foramen and involved with shortening of the orbitotemporal region. Gould (1995) commented that the relative position of the suboptic foramen (her character 21 scored thus: (0) anterior to the sphenorbital fissure; (1) present in the medial wall of the sphenorbital fissure; (2) present in the medial wall of the sphenorbital fissure but hidden within the fissure) seems to be related to the shortening of the skull in erinaceids. As the skull shortens, the alisphenoid overlaps the orbitosphenoid, creating a strong alisphenoid wing [character 17 of Frost et al. (1991) and Gould (1995)], the degree of overlap seems to be directly related to the visibility of the suboptic foramen from lateral view and, as pointed out by Butler (1948) the orbitosphenoid is reduced in size. While the alisphenoid is more extensive in H. megalotis than in other galericines, and the suboptic and sphenorbital foramina are partially concealed in lateral view, the orbitotemporal region is not obviously shortened. The anterior process in H. megalotis is fully ventral in location and is actually or nearly in contact with a short posteroventral process of the maxilla, thus contributing to the ventral floor of the orbit, however the orbitosphenoid is not reduced in size. It is possible that this character state in H. megalotis is not homologous with that of the Erinaceinae and that it actually represents a separate character transformation, alternatively it is scored incorrectly and the plesiomorphous condition should be the presence of the anterior process. [22.1] Palatal shelf and spine: (0) well developed spine on posterior palatal shelf; (1) spine absent or vestigial. CI 0.200. Shown only in DELTRAN, homoplasious with Podogymnura, H. parvus, and Atelerix. DISCUSSION The addition of two taxa to the analysis performed by Frost et al. (1991) provided broadly similar results in that the Galericinae divided intotwo main groups: one comprising Echinosorex and Podogymnura, the second including all five species of Hylomys. The results of the current phylogenetic analysis lend support to the taxonomy proposed by Frost et al. (1991) that the three species of Hylomys considered in their analysis (H. suillus, H. sinensis and H. hainanensis) are correctly attributed to a single genus rather than the three separate genera (respectively Hylomys, Neotetracus and Neohylomys) maintained by Corbet (1988). The additional species of Hylomys however, reduced the degree of support for the genus and, on this particular morphologi- cal data set, a considerable degree of homoplasy 1s evident within the Hylomys clade. There was only one unique synapomorphy for the 8 Hylomys clade (character 19.1: the wide separation between the cranio-orbital and ethmoid foramina), two of the other apomorphies (34.1: the inflation of the mastoid region between the exoccipital and the squamosal, and 41.1: the expanded exoccipital) showing homoplasy with P. truei, while the third character state (50.1: upper canine slightly larger than the adjacent post-canine teeth) is not shown by H. sinensis (50.2: upper canine approximately equal in size to the adjacent post-canine teeth). There are no autapomorphies defining H. megalotis, which shows more plesiomorphy than the other species of Hylomys; many of its features are homoplasious with Echinosorex, Podogymnura and Erinaceinae. Hylomys is a morphologically variable genus, containing species that are generally well segregated and show little overlap in species range. Hylomys hainanensis is a geographically isolated island form and while H. sinensis and H. parvus are each parapatric with H. suillus in a few areas, Corbet (1988) pointed out that in regions where H. sinensis and H. suillus occur, H. sinensis is found at greater altitudes than H. suillus. Similarly Ruedi et al. (1994) showed that although H. parvus is currently restricted to moss forests at the peak of Gunung Kerinci, Sumatra at greater altitudes than H. suillus, the latter occurs elsewhere at greater and lesser altitudes. Both Corbet (1988) and Ruedi et al. (1994) invoked ecological factors such as competitive exclusion to explain the altitudinal segregation of these three species, but did not provide data to support this supposition. There are few distribution records of H. suillus and H. megalotis in Lao PDR, which potentially may be sympatric or possibly parapatric if H. megalotis should prove to be ecologically restricted to the specific limestone habitat in which it was first collected. Little is known about the biology of Hylomys. Hylomys sinensis is believed to be entirely terrestrial and H. swillus mainly so, although this species has also been observed climbing in low bushes (Lekagul & McNeely, 1977). Hylomys suillus occurs in hilly or montane humid forests with dense undergrowth, H. sinensis in cool damp forests in the cover of runways and under logs and rocks. In their original description Shaw & Wong (1959) reported that H. hainanensis spends most of its time in underground burrows and that the cylindrical body, short tail and claws are adaptations to a fossorial life. Hylomys parvus is apparently restricted to high alti- tude moss forest. There is no information about the behavioural ecology of H. megalotis but the limestone karst where it has been found to date is an unusual habitat and some of the morphological features of this species, such as the moderately broad forefeet with long, fairly stout claws, the long naked hindfeet with large cheiridia, the moderately long tail and the comparatively flattened braincase may be adaptations to life in this habitat. ACKNOWLEDGEMENTS. We would like to thank the staff of WWEF-Thai- land, the Lao PDR Department of Forestry (DoF) and the Forest Management and Conservation Project (FOMACOP) for permission to work in Lao PDR and for organising the project; in particular Bouahong Phanthanousy (National Project Director, FOMACOP), Bouaphanh Phantavong (Deputy National Project Director, FOMACOP), Robert Mather and Robert Steinmetz (WWF- Thailand) and Bruce Jefferies (FOMACOP), who additionally oversaw the export of specimens. We are very grateful to Maurice Webber for his vital contribution to all aspects of the fieldwork. Assistance inthe field was gratefully received from Thongphanh Ratanalangsy (Khammouan Limestone and DongPhu Vieng NBCA Co-ordinator), Sinnasone Seangchanthanarong (DoF) and Nousine Latvylay (driver for FOMACOP in Savannakhet). The headmen and villagers from Pontong, Muang, Mouangkhai, Louang, Vieng, Kengkhot, Thamtem and Tonglom provided excellent field support and companionship, and without them this project would nothave been possible. We would like to thank Angela L. Smith for her help in measuring specimens at TISTR and for her comments on the manuscript. P.D. JENKINS AND M.F. ROBINSON Field work was funded by the Global Environment Facility through the World Bank to the Government of Lao PDR, via a contract with Burapha Development Consultants, sub-contracted to WWF-Thailand. We thank Bob Randal (AMNH), Jacques Cuisin (MNHN) and Nivesh Nadee (TISTR) for making specimens in their collections available to us. We are very grateful to Andrew Cabrinovic (NHM) for specimen preparation, to Harry Taylor, Photographic Unit (NHM) for the excellent photographs and to Karen Fisher (volunteer, NHM) for her kind assistance with documentation. We are grateful to Dr Francois Catzeflis, Institut des Sciences del’ Evolution, Université Montpellier for his constructive review of the manuscript. We particularly thank Dr Darrell Siebert, NHM for his generous guidance with the phylogenetic analysis and constructive criticism of the manuscript. REFERENCES Blainville, H. M. D. de 1838. Recherches sur l’ancienneté des Mammiferes insecti- vores a la surface de la terre; précédées de I’histoire de la science a ce sujet, des principes de leur classification et de leur distribution géographique actuelle. Compte Rendus Hebdomadaires de Scéances de |’ Académie des Sciences 6: 738— 744. Butler, P. M. 1948. On the evolution of the skull and teeth in the Erinaceidae, with special reference to fossil material in the British Museum. Proceedings of the Zoological Society, London 118 (2): 446-500. Corbet, G. B. 1988. The family Erinaceidae: a synthesis of its taxonomy, phylogeny, ecology and zoogeography. Mammal Review 18 (3): 117-172. Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: 783-791. Frost, D. R., Wozencraft, W. C. & Hoffmann, R. S. 1991. Phylogenetic relationships of hedgehogs and gymnures (Mammalia: Insectivora: Erinaceidae). Smithsonian Contributions to Zoology (518): 1-69, 19 plates. Gould, G. C. 1995. Hedgehog phylogeny (Mammalia, Erinaceidae) — the reciprocal illumination of the quick and the dead. American Museum Novitates (3131): 1-45. Kitching, I. J., Forey, P. L., Humphries, C. J. & Williams, D. M. 1998. Cladistics. Second edition. The theory and practice of parsimony analysis. Oxford University Press, Oxford, New York, Tokyo. Lekagul, B. & McNeely, J. A. 1977. Mammals of Thailand. Kurusapha Ladprao Press, Bangkok. McDowell, S. B. 1958. The greater Antillean insectivores. Bulletin of the American Museum of Natural History 115 (3): 117-214. McKenna, M. C. & Bell, S. K. 1997. Classification of mammals above the species level. Columbia University Press, New York. Mearns, E. A. 1905. Descriptions of new genera and species of mammals from the Philippine Islands. Proceedings of the United States National Museum 28: 425-460. Miiller, S. 1840. Over de Zoogdieren van den Indischen Archipel (No. 2) pp. 9-57. In C. J. Temminck (ed.) Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche overzeesche bezittingen, door de Leden der Natuurkundige commissie in Indié en andere Schrijvers. A. Amz & Co., Leiden. Novacek, M. J. 1986. The skull of lepticid insectivorans and the higher-level classi- fication of eutherian mammals. Bulletin of the American Museum of Natural History 183 (1): 1-111. Osgood, W. H. 1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic expeditions. Publications of the Field Musuem of Natural History, Zoology (18): 193-339. Raffles, T. S. 1822. Descriptive catalogue of a zoological collection, made on account of the Honourable East India Company, in the island of Sumatra and its vicinity, under the direction of Sir Thomas Stamford Raffles, Lieutenant-Governor of Fort Marlborough; with additional notices illustrative of the Natural History of these countries. Transactions of the Linnean Society, London 13 (1): 239-274. Robinson, H. C. & Kloss, C. B. 1916. Preliminary diagnoses of some new species and subspecies of mammals and birds obtained in Korinchi, West Sumatra, Feb.—June 1914. Journal of the Straits Branch of the Royal Asiatic Society (73): 269-278. Robinson, M. F. 1999. Order Insectivora, pp. 221-223 In J. W. Duckworth, R. E. Salter & Khounboline, K. (compilers) Wildlife in Lao PDR 1999 status report. TUCN, Vientiane. Ruedi, M., Chapusiat, M. & Iskander, D. 1994. Taxonomic status of Hylomys parvus and Hylomys suillus (Insectivora: Erinaceidae): biochemical and morphological analyses. Journal of Mammalogy 75 (4): 965-978. Swofford, D. L. 1999. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. Shaw, T. H. & Wong Song 1959. [A new Insectivore from Hainan]. Acta Zoologica Sinica 11: 422-425. [In Chinese; English summary]. Trouessart, E.-L. 1909. Neotetracus sinensis, a new insectivore of the family Erinaceidae. Annals and Magazine of Natural History (8) 4: 389-391. A NEW SPECIES OF HYLOMYS Table 1 Comparative material Hylomys sinensis BMNH 1932.4.19.3 Chapa, Tonkin, [ Vietnam] BMNH 1933.4.1.117—134 Chapa, Tonkin, [Vietnam] BMNH 1933.4.1.536—541 Chapa, Tonkin, [Vietnam] BMNH 1911.2.1.15—23 Omi-san, Omi-Hsien, S. Szechwan [Sichuan, PDR China] BMNH 1982.205 Omi-san, Omi-Hsien, S. Szechwan [Sichuan, PDR China] MNHN 1911-1180-1184 Ta-Tsien-Lou, Setchouen [Sichuan, PDR China] BMNH 1909. 12.13.1 Ta-Tsien-Lou, Szechuan [Sichuan, PDR China] BMNH 1911.8.6.1 Yengyuek, Yunnan, [PDR China] BMNH 1912.7.15.1 Ching-tsai- Yang, Yunnan, [PDR China] BMNH 1914.10.23.3 Near Yang-fsi, W. Yunnan, [PDR China] Hylomys suillus BMNH 1909.7.20.2—3 Sima, Burma [Myanmar] AMNH 44112 Nam-Ting, Yunnan, China [PDR China] BMNH 1925.1.1.17 Bao-Ha, Tonkin, [Vietnam] BMNH 1926.10.4.42 Xieng-Khouang, Laos [Lao PDR] AMNH 87313 Bologens Plateau, Laos [Lao PDR] BMNH 1926.10.4.36-41 Dak-to, Annam MNHN 1929-320-325 Dak-to, Annam BMNH 1955.1422 Tasan, Chumpawn, Peninsular Siam [Thailand] TISTR 54-611 Huey Mae Sanam, Chiengmai, Thailand TISTR 54-613 Trang, Muang, Chong, Thailand TISTR 54-614—615 Khao Yai National Park, Korat, Thailand TISTR 54-616 Phu Nam Tok, Saraburi, Thailand TISTR 54-617 Pok Nam Tok, 21 km from Saraburi, Saraburi, Thailand TISTR 54-618 Mae Sai, B. Santon Poi, Chieng Rai, Thailand TISTR 54-1498-1500 Pak Thong Chai, Sakaerat, Nakhon Ratchasima, Thailand TISTR 54-1811 Pak Thong Chai, 17 km S of Pak Thong Chai, Nakhon Ratchasima, Thailand TISTR 54-1809-1810 Khon San, Pa Phu Khieo, Chaiyaphum, Thailand TISTR 54-1812 Phu Kradung National Park, Loei, Thailand BMNH 1960.8.4.7 Ulu Langat Forest Reserve, Kajang, Selangor, [Malaysia] BMNH 1961.1158 Ulu Langat Forest Reserve, Kajang, Selangor, Malaysia BMNH 1955.1420 Semangko Pass, Pahang, Federal Malay States [Malaysia] BMNH 1961.1159 Padang, Jeriau, Fraser’s Hill, Pahang, Malaysia BMNH 1962.710-711 Jandai Baik, Pahang, Malaysia BMNH 1912.10.22.7 Pelarit, Perlis, Malay [Malaysia] BMNH 1955.1421 Pelarit, Perlis, Federal Malay States [Malaysia] BMNH 1912.10.22.8 Perlis, Malay Peninsula [Malaysia] BMNH 1955.1423 Jor, Batang Pasang, Perak, Federal Malay States [Malaysia] BMNH 1955.1424 Kedah Peak, Federal Malay States [Malaysia] BMNH 1962.711a Kedah Peak, Kedah, Malaya [Malaysia] BMNH 1892.9.6.4 Mt. Kina Balu [Malaysia] BMNH 1895.10.4.3-4 Mt. Kina Balu [Malaysia] BMNH 1955.661 Mount Kinabalu, British North Borneo [Malaysia] BMNH 1971.2614—2615 Mt. Kinabalu, N Borneo [Malaysia] MNHN 1889-37 Mont Kina Balu, Borneo [Malaysia] MNHN 1893-132—133 Mont Kina Balu, Borneo [Malaysia] BMNH 1971.2616 Dusan Dankulum, Kinabalu, N Borneo [Malaysia] BMNH 1971.2617-2618 Tinampoh, Bundu Tuhan rest house, N Borneo [Malaysia] BMNH 1919.11.5.7 Korinchi, Sumatra [Sumatera, Indonesia] AMNH 102532 Seletan, Mocamh Doewa, Sumatra [Sumatera, Indonesia] AMNH 102533 Seletan, Mocamh Doewa, Sumatra [Sumatera, Indonesia] AMNH 102534 Seletan, Mocamh Doewa, Sumatra [Sumatera, Indonesia] AMNH 102820 Lampung, Kalianda, Sumatra [Sumatera, Indonesia] BMNH 1954.45 Tyibodas, West Java [Jawa, Indonesia] BMNH 1954.46—-48 Sodong Jerok, Idjen Massif, East Java [Jawa, Indonesia] BMNH 1961.1743 Tjemorosewu, Mt. Lawu, Java [Jawa, Indonesia] AMNH 106111 Java [Jawa, Indonesia] Hylomys parvus BMNH 1919.11.5.8—12 Korinchi, Sumatra [Sumatera, Indonesia] Podogymnura truei BMNH 1953.659-660 Baclayan, E slopes of Mount Apo, Mindanao, Philippine Islands Echinosorex gymnura BMNH 1914.12.8.101—104 Bankachon, Tenasserim [Myanmar] BMNH 1955.1452 Changkat Mentri, Perak, Federal Malay States [Malaysia] BMNH 1955.1453 Damansara Road, Kuala Lumpur, Selangor, Federal Malay States [Malaysia] BMNH 1961.1156 Rontau Panjang, Klang, Selangor, Malaya [Malaysia] BMNH 1961.1157 Sungei Buloh, Selangor, Malaya [Malaysia] BMNH 1961.1157 Sungei Buloh, Selangor, Malaya [Malaysia] BMNH 1951.179-180 Mount Dulit, Sarawak, Borneo [Malaysia] BMNH 1951.181 Tinjar River, Baram District, Sarawak, Borneo [Malaysia] BMNH 1971.2613 12 miles N of Kalabakan, Tawau, N Borneo [Malaysia] 10 P.D. JENKINS AND M.F. ROBINSON Table 2 Data matrix from Frost et al. (1991: Appendix 2) with the addition of characters scored for two additional taxa, Hylomys megalotis and Hylomys parvus Character Hypothetical ancestor Echinosorex gymnura Podogymnura aureospinula Podogymnura truei Hylomys sinensis Hylomys suillus Hylomys hainanensis Hylomys megalotis Hylomys parvus Hemiechinus aethiopicus Hemiechinus hypomelas Hemiechinus micropus Hemiechinus auritus Hemiechinus collaris Mesechinus dauuricus Erinaceus amurensis Erinaceus concolor Erinaceus europaeus Atelerix frontalis Atelerix algirus Atelerix albiventris Atelerix sclateri Tenrecoids Soricoids Lepticidae (fossil) 111111111122222222223333333333444444444455555555556666666666777777777788 123456789012345678901234567890123456789012345678901234567890123456789012345678901 000000000?000?00000?000000000000000000000000000000000?7000?00?7000000007?77000000000 000010112211010000020010000000001000100000010000000100010000111001101111110000001 00001001121101000002010000000000110010001001000100010111010011100110?711110000001 100101010211100100120100000000001100100010000100020011111111111001101111110000000 000101010211100100120000000000001100100010000000010000011111111001101111110000000 000101010211100100120000000000001100100010000000010000111111111001107?11110000000 100010012211000010120100000000001100100010000000010000000000111001101011110000000 111201102131011011011001104001110011111101101011120001112111121110210000001130200 111201102121011011011001103001110011111101101011120001112010121110210000001130200 111201102121011011011001104001110011111101101011120001112111121110210000001130200 111201102111011011011001102001110000111101101011120001112010121110210000001130300 111201102111011011011001102001110000111101101011120001112010121110210000001130300 110201102110011011011001101101011000111101101011120001112010121110210000000120310 110201102111011011011001101011010000111101101011221001112010121110210000000011110 110201102111011011011001101011010000111101101011221001112010121110210000000010110 11020110211101101101100110101101000011110110101122100111201012111021000000001?110 110201102111011011011101111011010000111101101011120001102010121110210000000111110 110201102111011011011101111011010000111101101011120001102011121110210000000110110 110201102111011011011101111011010000111101101011120001102111121110210000000210110 110201102111011011011101111011010000111101101011120001102111121110210000000110110 00000000030001000000000000000000000000000000000000?001000000000000000000000000000 00000000030001000002000000000000000000000000000000?001000100000000000000000000000 000000000000000000000000000000000000000000000000000000000000000000000???000?70??? Table 3. Selected measurements of Hylomys in millimetres to show variation in size and proportions. The mean, standard deviation and range are provided, with sample size in parentheses. Character Head and body length Tail length Ratio of tail length to head and body length Hindfoot length Ear length Greatest skull length Condyloincisive length Upper toothrow length Length from I1 to anterior of P4 Ratio of 11—P4 length to upper toothrow length Rostral breadth Rostral length Ratio of rostral length to rostral breadth Braincase breadth Braincase height Ratio of braincase height to braincase breadth ' Measurements from Shaw & Wong (1959). H. parvus H. sinensis H. suillus H. hainanensis' H. megalotis 107.67 + 4.50 115.75 + 4.83 1325 5.92 136.71 + 8.28 124.23 + 7.68 104-114 (3) 111-124 (6) 120-139 (10) 120-147 (7) 115.6-134.7 (4) 24.00 + 0.82 68.5 + 3.70 2B OOS 09 39.67 + 2.13 86.33 + 3.29 23-25 (3) 63-73 (6) 19-30 (8) 36-43 (6) 82.8-91.3 (4) 0.22 + 0.01 0.57 + 0.05 0.17 + 0.35 0.29 + 0.13 0.70 + 0.04 0.21-0.24 (3) 0.51—0.63 (6) 0.14—0.24 (8) 0.27-0.31 (6) 0.65-0.74 (4) 23.17 + 0.24 25.88 + 1.02 PIES OES 25.14 + 1.62 20.83 + 0.38 23-23.5 (3) 24-27 (6) 20.5—23 (9) 24-29 (7) 20.4—21.3 (4) 18.38 + 0.45 17.71 + 1.41 WSS AAS 22.43 + 0.90 NIE VOD) 18-19 (6) 15-20 (7) 16-19 (6) 20.9-23.2 (4) 32.38 + 0.82 34.45 + 0.80 34.65 + 0.60 37.76 + 1.01 30.55 (1) 31.07-33.64 (10) 33.48-35.61 (7) 33.7-35.3 (4) 36.36-39.23 (4) 30.98 + 0.82 33/3 = O67; 36.96 + 1.04 29.09 (1) 30.05-32.36 (11) 32.88-35.01 (7) 35.66-38.56 (4) 15.21 + 0.42 NG Pas O37) 17.75 + 0.64 17.26 + 0.16 19.89 + 0.47 14.64-15.61(3) 15.64-16.82 (11) —16.70-18.75 (13) ——-17.0-17.50 (5) 19.44—20.64 (4) 8.19 + 0.47 8.94 + 0.44 9.79 + 0.46 11.65 + 0.33 7.54-8.61 (3) 8.15—9.54 (11) 9.05—10.34 (13) 11.40-12.21 (4) 0.54 + 0.02 0.56 + 0.02 0.55 + 0.01 0.59 + 0.01 0.52-0.56 (3) 0.51-0.58 (11) 0.54-0.58 (13) 0.57-0.59 (4) Slt = (0), 17/ 5.92 + 0.18 6.02 + 0.21 5729 2 0als 4.98—-5.35 (3) 5.59-6.20 (11) 5.57-6.44 (13) 5.06-5.38 (4) 9.85 + 0.49 9.67 + 0.29 10.35 + 0.41 13.57 + 0.43 9.33-10.5 (3) 9.33-10.37 (11) 9.57-10.85 (13) 13.30-14.32 (4) 0.52 + 0.03 0.61 + 0.02 0.59 + 0.02 0.39 + 0.01 0.47-0.55 (3) 0.57-0.65 (11) 0.56—0.63 (10) 0.37-0.41 (4) 13.89 + 0.31 15.60 + 0.86 14.70 + 0.54 14.61 + 0.34 13.29 (1) 13.42-14.45 (11) = 12.97-15.69 (10) ~—-13.9-15.4 (4) 14.16—-15.06 (4) 9.06 + 0.34 9.47 + 0.33 8.75 + 0.19 8.63 (1) 8.55—9.67 (11) 9.33-9.85 (7) 8.49-8.95 (4) 0.65 + 0.03 0.68 + 0.02 0.60 + 0.02 0.65 (1) 0.61—-0.70 (11) 0.63—0.70 (7) 0.58-0.62 (4) 11 A NEW SPECIES OF HYLOMYS Jose] Josie] Josie] Jose] € € £ €> o01e7] ose] ose] [jes JUISAI juasqy USI juosqy POSNY ][9M Z 10 | POSNJ [JIM Z JO | c G uasaid juasqy uasaid JUASAI Wuasaid juosqy uasaid 1UdSAI Jase] Apueoiyiusis IW/td © [esioq suoy] JOLINUY [yetus JUdSd1q juasqy jusoe[pe AJaSO]9 10 poulor poqesedas [jay SPI juasqy pouljap Ajiood 10 yuasqy [Jeu JOUNSIPU] jouNsiq SLIP AWD] juasqy Jolaquy JOR]UOD UT AjIRaN XAMOSOUIYIT Jase] Apueoyiusis LW/td 0} [esi0g suo] JOWQUY [jews UdSdI1g quasqy jusoe[pe AJASO]9 10 paulor J9YI980} aso[D 2PIM juasqy juasqy é MO]]PYS JOUNSIG JOUNSIP / JOUNSIPUy peolg Ajayesapoyy MOT[EYS JOLaUy ByyIxeu Aq payesedas Ajapi Ay DANUUUKSOPOg Jase] ApYysi[s LW/td 0} [esiog suo] JOU [jews quasqy Uasalg poqesedas Ajapi Ay poqesedas [ja OPI juasqy qunyq ‘pauljap Aj100q SAISUD}XA MOT[PYS JOuNsIG jouNsiq MOLIRN MOTTRYS JoAo7] ByyIxeu Aq payeiedas Ajapi Ay SHO]D3au sdwojA yy Josie] Apysils IW/td ©} [esto JOUNSIPUL 10 [feUUS anjns uQ payesuolq juasqy juasqy poqesedas Ajapr Ay JaIAaS0} asojD MOLIRN JOUNSIP “OYs quniq ‘poulyap Aj00g [jews daaq JOUNSIpU] peoig daap Ajaqesapoy| JoWaquy yor}U00 UI SS] 10 a10/\ sawojay Jojjeus ApYystys E> [jews juasqy Pasny [JOM Z 10 | 1UdSd1q ]UdSdIq Joss ApYsils td/€d 0} jesi0g JOUNSIPUT JO |]eUIS ainjns ud paiesuoq 1Uasaid juasqy poqesedas Ajapi Ay Jayas0} aso]D MOLBU A139, ssoo0id |eyiqsovidns 0] sayngijuoS ‘padojaaap [Ja padojaaap []9,\\ [jets daoq juasqy peoig daaq JOWajUy y9e]U09 UI SSd] IO D10/\ SNIJINS samo] AY Jojjews Apystps es [jews juasqy Pasny [JOM Z 40 | juasqy juasaid Josie] Apysiys bd/€d 1 [es1og JOUNSIPUT 10 [RUS ainjns uO poiesuolq ]UASAIq juasqy payesedas Ajapi Ay G MOLBU AIDA, ssadoid |ejiqioesdns 0] sajngijUoS ‘podojaaap [Jo podojaaap [9 é é juasqy peoig daap Ajayesapoyy JOLWaUy RypIxeu Aq payesedas A[MoueN SISUAUDUIDY UO V | Jayyeuus Ayysiys €> [bug juasqy Pasny [Jam Z 10 | juasqy juasqy jenbaqng bd/Ed 1 [PS1Oq JOUNSIPU! 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" ‘ed al —* Pus 2 " i ~~ : | ~~ Yin 45 ey J ? 1 — ee yf . . “eer! : a if Coed Thay 4 agent roa =~ pie af tia 4 a | } 7 , ay }: : ; 7 see " We j x ves) ‘ Ny a" ae . Kx (382 Git: Sy, Bull. nat. Hist. Mus. Lond. (Zool.) 68(1): 13-18 Issued 27 June 2002 A new species of freshwater crab (Brachyura, Potamoidea, Potamonautidae) from Principe, Gulf of Guinea, Central Africa NEIL CUMBERLIDGE Department of Biology, Northern Michigan University, Marquette, Michigan 49855, U.S.A. PAUL F. CLARK Department of Zoology, The Natural History Museum, Cromwell Road, London, SW7 5BD, U.K. JONATHAN BAILLIE Institute of Zoology, Zoological Society of London, Regents Park, London NWI 4RY, UK. CONTENTS INTER CNHI ECVE aes «once pise ccupeentaanesiscate tee scvsts'sSudeunesctesauess 17 Bee BR ea ese n a NER EE a atiae See es ace cane RE ERS «wee nance os Read aher kone Manone 17 waatt as haps ae ska ac Wtanhtane sales en des spt (oases wide cas av tras eee ay 17 A new species of freshwater crab of the genus Potamonautes Macleay, 1838 is described from Principe (Democratic Republic of Sao Tomé and Principe), an island in the Gulf of Guinea off the coast of Central Africa. The specimens were collected during a recent zoological expedition by the Zoological Society of London. This is also the first record of a freshwater crab on the island of Principe. INTRODUCTION The freshwater crabs reported on here were collected during a zoological expedition to the island of Principe, made recently by the Zoological Society of London. The island of Principe, together with the island of Sao Tomé, constitutes a small independent country (The Democratic Republic of Sao Tomé and Principe) in the Gulf of Guinea. Principe is the second in a chain of volcanic islands that make up the Atlantic Ocean Islands group, that lies off the coasts of Cameroon, Equatorial Guinea and Gabon. The other islands in this group are Bioko, SAo Tomé and Annobon. Bioko is closest to the mainland and has two species of potamonautid freshwater crabs, Sudanonautes floweri (De Man, 1901) and S. granulatus (Balss, 1929), both of which are also found in nearby Cameroon (Cumberlidge, 1993, 1995, 1999). Sao Tomé is the third island in the group, and has one endemic species of freshwater crab, Potamonautes margaritarius (A. Milne-Edwards, 1886). There are no records of freshwater crabs occurring on Annobon, the fourth island in the chain, and the furthest from the mainland. Until the present report, freshwater crabs were not known to be present on Principe. The new species from Principe was collected from streams and nearby land in cloud forest in the remote roadless highland region in the southwest part of the island. The new species is compared to Potamonautes anchietae (De Brito-Capello, 1871) and to P. margaritarius, but differs from each in a number of important characters (Bott, 1953, 1955, 1964). Although the speci- mens from Principe are both subadult females, and ideally an adult male is needed to make a definitive identification, a description is © The Natural History Museum, 2002 nevertheless provided here, based on several unique somatic characters of the specimen. Characters of the gonopods, male abdo- men, and male chelipeds will be described when more material (including an adult male) becomes available. Figures were prepared using a camera lucida, and the specimens were deposited in The Natural History Museum, London, U.K. (BMNH). Abbreviations: cw, distance across the carapace at the widest point; cl, carapace length measured along the median line, from the anterior to the posterior margin; ch, carapace height, the maximum height of the cephalothorax); fw, front width measured along the anterior margin; s, thoracic sternite; e, thoracic episternite; s4/s5, s4/s5, s5/s6, s6/s7, s7/s8, sternal sulci between adjacent thoracic sternites; s4/e4, s5/e5, s6/e6, s7/e7, episternal sulci between adjacent thoracic sternites and episternites; P1—P5, pereiopods 1-5, al—a6, abdominal segments 1-6, a7, telson of the abdomen. SYSTEMATIC ACCOUNT Family POTAMONAUTIDAE Bott, 1970 Genus POTAMONAUTES MacLeay, 1838 Potamonautes principe sp. nov. (Fig. 1) MATERIAL EXAMINED Holotype. BMNH 2001.6907. 1 subadult female, cw 40.5, cl 27.5, 14 N. CUMBERLIDGE, P.F. CLARK AND J. BAILLIE Fig. 1 | Potamonautes principe sp.nov. Holotype subadult female, cw 40.5 mm, BMNH 2001.6907. A, carapace, dorsal view; B, cephalothorax, frontal view; C, left third maxilliped; D, abdomen; E, anterior sternum; F, right cheliped, frontal view; G, left cheliped, frontal view; H, carpus and merus of right cheliped, dorsal view; I, carpus and merus of right cheliped, inferior view. Scale = A, H, I, 13.1 mm; B, D, C, E, 10.5 mm; F, G, 8.3 mm. A NEW FRESHWATER CRAB FROM PRINCIPE ch 14.8, fw 10.5 mm, summit of Pico do Principe (01° 34’, 51"N, 07° 22', 57"E), 945 m, coll. J. E. M. Baillie, 26 Sept. 1999. Paratype. BMNH 2001.6908.1 subadult female, cw 33.2, cl 22.8, ch 11.6, fw 10.0 mm, summit of Pico do Principe (01° 34’, 51"N, 07° 22', 57"E), 945 m, coll. J. E. M. Baillie, 1 Sept. 1999. DIAGNOSIS. Postfrontal crest straight, smooth, spanning entire carapace, ends meeting anterolateral margins at epibranchial teeth. Exorbital tooth small, pointed; epibranchial tooth represented only by small granule; anterolateral margin posterior to epibranchial tooth raised, completely smooth, continuous with posterolateral margin. Carapace surface completely smooth; highly arched, height greater than front width (ch/fw 1.5). Pleural (vertical) suture on carapace sidewall Y-shaped, ends meeting exorbital and epibranchial teeth. Suborbital margin raised, completely smooth. Ischium of third maxilliped with deep vertical sulcus. Third sternal sulcus s3/s4 deep, v-shaped, meeting sterno-abdominal cavity. Thoracic epister- nal sulci s4/e4, s5/e5, s6/e6 and s7/e7 distinct. Anterior inferior margin of merus of cheliped lined by row of small sharp teeth, with large pointed tooth near junction with carpus. DESCRIPTION. Carapace ovoid, wide (cw/fw 3.88), highly arched (ch/fw 1.45); surface completely smooth semi-circular, urogastric, transverse branchial grooves faint. Front straight, relatively narrow, about one-quarter carapace width (fw/cw 0.26), anterior margin sharply deflexed. Postfrontal crest distinct, smooth, straight, span- ning entire carapace, consisting of fused epigastric, postorbital crests; ends of postfrontal crest meeting anterolateral margins at epibranchial teeth. Anterolateral margin between exorbital, epibranchial teeth smooth, lacking intermediate tooth; anterolateral margin posterior to epibranchial tooth raised, completely smooth, continuous with posterolateral margin. Exorbital tooth small, pointed; epibranchial tooth represented only by small granule. Suborbital margin raised, completely smooth. Suborbital, subhepatic, pterygostomial regions all completely smooth; sidewall divided into three parts by longitudinal (epimeral) suture (dividing suborbital, subhepatic regions from pterygostomial region), and by Y-shaped vertical (pleural) groove (dividing suborbital from subhepatic regions). Superior ends of Y-shaped vertical groove meeting exorbital, epibranchial teeth. First thoracic sternal sulcus s1/s2 deep; second sulcus s2/s3 deep, running horizontally across sternum; third sulcus s3/s4 deep, v-shaped, meeting sterno-abdominal cavity. Tho- racic episternal sulci s4/e4, s5/e5, s6/e6 and s7/e7 distinct. Third maxillipeds filling entire oral field, except for transversely ovate respiratory openings at superior lateral corners; long flagellum on exopod of third maxilliped; ischium with deep vertical sulcus. Epistome prominent, smooth, triangular. Mandibular palp 2-seg- mented; terminal segment single, undivided, with hair (but no hard flap) at junction between segments. Subadult female abdomen subcircular, segments al—a6 of female abdomen quadrate, telson (a7) a broad triangle with rounded apex; segments a5—a6 broadest. Major cheliped of subadult female relatively slender, with elon- gated dactylus and propodus, palm of propodus swollen; fingers of digits of cheliped with small even teeth, forming long narrow interspace when closed. First carpal tooth of inner margin of carpus of cheliped large, pointed; second carpal tooth pointed, half size of first tooth. Posterior inferior margin of merus of cheliped smooth, with few small teeth distally; medial inferior margin with row of small sharp teeth along entire length, large pointed tooth at distal end; superior surface of merus smooth. Pereiopods P2—P5 slender, P3 longest, PS shortest. Dactyli of P2—P5 tapering to point, each bearing four rows of downward-pointing short, sharp spines. CoLour. Thecolour of the specimens when freshly caught (before 15 alcohol preservation) was creamy white with a very light purplish tone, however one specimen was more white than the other. This coloration was relatively uniform throughout the body including the dorsal carapace, underside, and legs. The purplish tone was darker on the carapace just behind the eyes, which turned to an orange reddish color once the specimens were placed in alcohol. DISTRIBUTION. This species is known only from the summit of the Pico do Principe: (01° 34", 51'N, 07° 22', 57"E) at 945 meters, The Democratic Republic of Sao Tomé and Principe, Gulf of Guinea, Central Africa. ETYMOLOGY. The species is named for the island of Principe where it was collected. The species name principe is a noun in apposition. TAXONOMIC REMARKS. The new species is assigned to Pota- monautes because it possesses the following combination of characters: the anterolateral margin lacks an intermediate tooth between the exorbital and epibranchial teeth; the mandibular palp is two-segmented; and the third maxilliped exopod has a long flagellum. Potamonautes is a widespread genus of African freshwater crabs found throughout Africa from Senegal to the Horn of Africa, and from Egypt to South Africa. Bott’s (1955) revision of the freshwater crabs of Africa recognised some 34 species in this genus. Since then, the number of species of Potamonautes has risen to more than 60 (Bott, 1959, 1960, 1964, 1968, 1970; Stewart et al., 1995; Stewart, 1997a,b; Daniels et al., 1998; Cumberlidge, 1999; Corace et al., 2001). Although Bott (1955) recognised 15 subgenera of Potamonautes, the authors of the present study prefer to follow Cumberlidge (1999) and use Potamonautes [sensu lato | for all species, pending a revision of the entire genus (Cumberlidge, un- published). It is not normally good practice to describe a new species from a subadult female. However, we have decided to establish this taxon in the light of the distinct nature of the available morphological characters, and because of the isolated nature of the study area which may mean that further specimens of P. principe are unlikely to become available for some time. Characters of the gonopods, adult male chelipeds, abdomen and sternum are not at present known because the only specimens of P. principe are subadult females. Nevertheless, there are a number of unique characters that distinguish P. principe from other species in the genus. COMPARISONS WITH OTHER SPECIES. Potamonautes principe is closest to P. anchietae, a medium-sized species of freshwater crab from Angola (De Brito-Capello, 1871, Bott, 1953, 1955, 1964). This species was most recently described and illustrated by Bott (1955, p. 247-249, figs. 24, 76, 77, pl. IX, fig. la—d) as P. (Isopotamonautes) anchietae. Potamonautes principe and P. anchietae are similar in that both species have a highly arched carapace, a prominent and complete postfrontal crest, a pointed exorbital tooth, a small granu- lar epibranchial tooth, a v-shaped thoracic sternal groove s3/s4, and a similar-sized first carpal tooth on the carpus of the cheliped. However, there are a number of characters that distinguish the specimens from Vissabenguilla, Angola (SMF 1890) described by Bott (1955) as P. (/.) anchietae from the specimens from Principe under consideration here. The carapace height of P. principe is greater than that of P. anchietae (ch/fw P. principe 1.45, P. anchietae 1.19), and the frontal margin of P. principe is narrower than that of P. anchietae (fw/cw P. principe 0.26, P. anchietae 0.39). Further, the anterolateral margins of the carapace of P. principe are completely smooth and lack teeth of any kind, whereas these margins in P. anchietae are distinctly granular. The medial inferior margin of the merus of the cheliped of ‘spodtpayD “q :MoIA [RIUOL “D ‘MIA [eQUAA “g SMOIA [PSIOG ‘Y “L069' 1007 HNIN “Wi COP Mo ‘ajeuay y[npeqns adAjofoH ‘Aou'ds adioutd sajnvuowmjog 7 “Bi = | el < er pa Q zZ < w ~ < = Oo a iw eal ©) al = ~ za oO - = Oo Za A NEW FRESHWATER CRAB FROM PRINCIPE P. principe has a row of distinct pointed teeth along its length, whereas this margin in P. anchietae, lacks teeth, and is either granular or smooth. Finally, the suborbital and pterygostomial regions of P. principe are smooth, whereas these regions are distinctly granular in P. anchietae. Bott (1953, 1955, 1964) described two subspecies of P. anchietae: P. (1.) a. biballensis Rathbun, 1905, and P. (I.) a. machedoi Bott, 1964, both of which are found in Angola. However, comparison of the types of these taxa with P. principe indicates that both differ substantially from the new species described here. For example, the epibranchial tooth of P. (I.) a. biballensis is large and pointed whereas that of P. principe is small and granular, and the carapace of P. (1.) a. biballensis is flattened, whereas that P. principe is high (ch/ fw P. principe 1.45, P. (1.) a. biballensis 1.25). Similarly, the epibranchial tooth of P. (/.) a. machedoi is large and pointed whereas that of P. principe is small and granular, and the frontal margin of P. principe is narrower than that of P. (/.) a. machedoi (fw/cw P. principe 0.26, P. (I.) a. machedoi 0.33). Potamonautes principe was also compared here with a specimen of P. margaritarius from Sao Tomé (SMF 2668), and the two taxa can be distinguished by the following characters. The carapace of the latter species is distinctly flattened (ch/fw P. margaritarius 0.95, P. principe 1.45), the anterolateral margin of P. margaritarius behind the epibranchial tooth is clearly toothed (whereas this margin is completely smooth in P. principe), and the ischium of the third maxilliped of P. margaritarius is smooth and lacks a vertical suture (whereas this suture is deep in P. principe). ECOLOGICAL NOTES The island of Principe is ovoid in outline, with a total land area of about 139 sq km (Fig. 3). The highest point is Pico de Principe (945 m), which is a volcanic mountain whose lower slopes are vegetated by lowland rain forest that grades into cloud forest at higher elevations. The climate of Principe is tropical, hot, and humid, and there is a single rainy season from October to May. Rainfall levels are high, averaging around 1,000 mm per year in the north and 5,000 mm per year in the south, and monthly tem- peratures range between 25° and 31° C (Bredero et al., 1977). The two specimens were collected from the summit of the Pico do Principe at 945 meters above sea level on two separate occasions. The terrain at the summit of the Pico do Principe is volcanic and mountainous, and the vegetation cover is cloud forest with small stunted trees, and a high abundance of epiphytes and bryophytes. The thick cloud cover and regular rainfall keep the soil and leaf litter very damp. On both occasions when crabs were collected the weather was extremely damp and the mountain top engulfed in cloud cover. Both specimens were collected on land from under damp and decaying leaf litter. There was no river or body of water near the location where the specimens were found. The nearest stream was observed at lower altitudes (830 m), but there were numerous temporary small pools at the summit which form after heavy precipitation. It is likely that P. principe can breathe air, given its extremely terrestrial lifestyle. When crabs were disturbed by removing the leaf litter cover, they ran rapidly across the forest floor and took cover under leaf litter or any available crevasse. Specimens were collected close to holes near the Pico do Principe, but crabs were not actually observed moving in or out of these holes, and it ramains possible that the holes may not have been dug by the crabs. 17 ACKNOWLEDGMENTS. Gilles Joffroy and Tariq Stevart (Université libre de Bruxelles) assisted in the collection of specimens. Phil Crabb (NHM Photo. Unit) took the photographs reproduced in this paper. REFERENCES Balss, H. 1929. Crustacea V. Potamonidae au Cameroun. Jn: Th. Monod, ed., Contributions a l'étude de la faune du Cameroun. Faune des Colonies francaises 3: 115-129 De Brito-Capello, F. 1871. Algumas especies novas ou pouco conhecidas pertencentes dos generos Calappa e Thelphusa. Jornal de Sciencias mathematicas, physicas e naturaes (Lisboa) 3: 128-134. Bott, R. 1953. Potamoniden (Crust. Decap.) von Angola. Publicagoes culturais da Companhia de Diamantes de Angola, Lisboa 16: 133-148. —— 1955. Die Sii®wasserkrabben von Afrika (Crust., Decap.) und ihre Stammesgeschichte. Annales du Musée du Congo Belge (Tervuren, Belgique) C- Zoologie, Séries Il, Il, 3: 209-352. — 1959. Potamoniden aus West-Afrika (Crust., Dec.). Bulletin de l'Institut francais d'Afrique noire 21, série A (3): 994-1008. 1960. Crustacea (Decapoda): Potamonidae. Jn: Hansstrém, B. & others, South African Animal Life. Results of Lund University Expedition in 1950-1952: 13-18. 1964. Decapoden aus Angola unter besonderer Beriicksichtigung der Potamoniden (Crust. Decap.) und einem Anhang : Die Typen von Thelphusa pelii Herklots, 1861. Publicagoes culturais da Companhia de Diamantes de Angola, Lisboa 69: 23-34. — 1968. Decapoden aus dem Museu do Dundo (Crust. Decap.). Publicagoes culturais da Companhia de Diamantes de Angola, Lisboa 77: 165-172. 1970. Betrachtungen tiber die Entwicklungsgeschichte und Verbreitung der StiBwasser-Krabben nach der Sammlung des Naturhistorischen Museums in Genf/ Schweiz. Revue Suisse de Zoologie 77: 327-344. Bredero, J. T., Heemskerk, W. & Toxopeus, H. 1977. Agriculture and livestock production in Sao Tomé and Principe (West Africa). Unpublished report. Foundation for Agricultural Plant Breeding, Wageningen. Corace, R. G., Cumberlidge, N. & Garms, R. 2001. A new species of freshwater crab from Rukwanzi, East Africa. Proceedings of the Biological Society of Washington. 114 (1): 178-187. Cumberlidge, N. 1993. Redescription of Sudanonautes granulatus (Balss, 1929) (Potamoidea, Potamonautidae) from West Africa. Journal of Crustacean Biology 13: 805-8 16. 1995. Redescription of Sudanonautes floweri (De Man, 1901) (Brachyura: Potamoidea: Potamonautidae) from Nigeria and Central Africa. Bulletin of the British Museum of Natural History (Zoology), London 61 (2): 111-119. 1999. The freshwater crabs of West Africa. Family Potamonautidae. Faune et Flore Tropicales 35, Institut de recherche pour le développement (IRD, ex-ORSTOM), Paris, 1-382. Daniels, S. R., Stewart, B. A. & Gibbons, M. J. 1998. Potamonautes granularis sp. noy. (Brachyura: Potamonautidae), a new cryptic species of river crab from the Olifants river system, South Africa. Crustaceana 71: 885-903. MacLeay, W. S. 1838. Brachyurous Decapod Crustacea. Illustrations of the Zoology of South Africa 5; being a Portion of the Objects of Natural History Chiefly Collected during an Expedition into the Interior of South Africa, under the Direction of Dr. Andrew Smith, in the Years 1834, 1835, and 1836; Fitted Out by ‘The Cape of Good Hope Association for Exploring Central Africa.’ In: A. Smith, Illustrations of the Zoology of South Africa; Consisting Chiefly of Figures and Descriptions of the Objects of Natural History Collected During an Expedition into the Interior of South Africa, in the Years 1834, 1835, and 1836; Fitted Out by ‘The Cape of Good Hope Association for Exploring Central Africa.’ (Invertebrates): 1-75. De Man, J. G. 1901. Description of a new fresh-water Crustacea from the Soudan; followed by some remarks on an allied species. Proceedings of the Zoological Society of London, 1901: 94-104. Milne-Edwards, A. 1886. La description de quelques Crustacés du genre Thelphusa recueillis par M. de Brazza dans les régions du Congo. Bulletin de la Société Philosophique Paris séries 7(10): 148-151. Rathbun, M. J. 1905. Les crabes d’eau douce (Potamonidae). Nouvelles Archives du Muséum d’ Histoire naturelle (Paris) 7(4): 159-322. Stewart, B. A. 1997a. Morphological and genetic differentiation between populations of river crabs (Decapoda: Potamonautidae) from the Western Cape, South Africa, with a taxonomic re-examination of Gecarcinautes brincki. Zoological Journal of the Linnean Society 199: 1-21. 1997b. Biochemical and morphological evidence for a new species of river crab Potamonautes parvispina (Decapoda: Potamonautidae). Crustaceana 70: 737-753. . Coke, M. & Cook, P. A. 1995. Potamonautes dentatus, new species, a fresh-water crab (Brachyura: Potamoidea: Potamonautidae) from KwaZulu-Natal, South Africa. Journal of Crustacean Biology 15: 558. 18 N. CUMBERLIDGE, P.F. CLARK AND J. BAILLIE Principe Gulf of Guinea Bonbon 1° 40'N Santo Antonio ® Porto R Bahia do Oeste ono Rest @ Terriro Velho (os Maria Correia ( 1°35'N & Pico do Principe Antonio Enes @ b ay Boné do Joquei 9 7° 20'E TPI5'E Fig. 3 Map of Principe showing the type locality of Potamonautes principe sp.nov. \ x (252 644 eo Bull. nat. Hist. Mus. Lond. (Zool.) 68(1); 19-26 A Issued 27 June 2002 Two new species of the Indo-Pacific fish genus Pseudoplesiops (Perciformes, Pseudochromidae, Pseudoplesiopinae) ANTHONY C. GILL Fish Research Group, Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 SBD, U.K. ALASDAIR J. EDWARDS Tropical and Coastal Management Studies, Department of Marine Sciences and Coastal Management, University of Newcastle, Newcastle upon Tyne NEI 7RU, U.K. SYNOPSIS. Pseudoplesiops immaculatus is described from 72 specimens from throughout the West Pacific and the eastern and central Indian Ocean. It is distinguished from congeners in having, in combination, a prominent intermandibular flap and an unspotted operculum. Pseudoplesiops occidentalis is described from five specimens from the Maldive Islands, central Indian Ocean. It is distinguished from congeners in having, in combination, II,23 dorsal-fin rays, 26—28 scales in lateral series, and scales with distinct centres and radii in all fields. INTRODUCTION The genus Pseudoplesiops Bleeker was recently diagnosed to included pseudoplesiopine pseudochromids with the medial lami- nae of the pelvic bones expanded dorsally (Gill & Edwards, 1999). Aside from this synapomorphy, members of the genus are distin- guished from other pseudoplesiopines in possessing the following combination of external characters: scales in lateral series 26—42; lower lip complete (uninterrupted at symphysis); and preopercular pores usually 7 (rarely 6 or 8), with a pore present at the upper terminus of the preopercle. The genus is mostly confined to the eastern Indian and Pacific Oceans, but two undescribed species occur in the Maldive Islands, central Indian Ocean. Although it is our intention to produce a revision of the genus, we herein describe the two Maldives species in order to make their names available for a forthcoming guide to western and central Indian Ocean fishes. MATERIALS AND METHODS Institutional abbreviations follow Leviton et al. (1985). All meas- urements to the snout tip were made to the midanterior tip of the upper lip. Length of specimens are given in mm standard length (SL), which was measured from the snout tip to the middle of the caudal peduncle at the vertical through the posterior edge of the dorsal hypural plate. Head length was measured from the snout tip to the posteriormost edge of the opercular membrane. Snout length was measured over the shortest distance from the snout tip to the orbital rim, without constricting the fleshy rim of the latter. Orbit diameter was measured as its fleshy horizontal length. Interorbital width was measured as the least fleshy width. Upper jaw length was measured from the snout tip to the posterior edge of the maxilla. Predorsal, preanal and prepelvic lengths were measured from the snout tip to the base of the first spine of the relevant fin. Body width was measured between the posttemporal pores. Caudal peduncle length was measured from the base of the last anal-fin ray to the ventral edge of the caudal fin at the vertical through the posterior © The Natural History Museum, 2002 edge of the ventral hypural plate. Caudal peduncle depth was measured obliquely between the bases of the last dorsal- and last anal-fin rays. Measurements of fin rays excluded any filamentous membranes. Pectoral fin length was measured as the length of the longest middle ray. Caudal fin length was measured as the length of the lowermost ray on the dorsal hypural plate. Counts of dorsal-, anal- and pelvic-fin spines (unsegmented rays) and segmented rays are presented, respectively, as Roman and Arabic numerals. If the last dorsal- or anal-fin ray was divided at its base it was counted as a single ray. Counts of branched, segmented rays in the dorsal and anal fins included unbranched rays behind the first branched ray. A value was not recorded if, due to tip damage, a branched or unbranched condition could not be determined for the segmented ray preceding the anteriormost branched ray. As in most actinopterygian fishes, the upper ray in the pectoral fin is rudimen- tary and rotated so that its asymmetrical medial and lateral hemitrichs appear to represent two separate rays; these were counted as a single ray. Procurrent caudal-fin ray counts were of the rays above (“up- per’) and below (‘lower) the principal caudal-fin rays. The uppermost principal caudal-fin ray was defined as the ray articulating with hypural 5, and the lowermost principal caudal-fin ray was the ray articulating with the cartilage nubbin between the distal tips of the parhypural and the haemal spine of preural centrum 2. All pseudoplesiopine species normally have 17 (9 + 8) principal caudal- fin rays. Counts of ‘scales in lateral series’ were of the posteroventrally oriented transverse scale rows on the midside, beginning with the row through the tubed scale at the branchial opening and ending with the row through the scale at the midposterior edge of the hypural plate. “Scales in transverse series’ were counted anterodorsally from the anal-fin origin to the dorsal-fin base. Circumpeduncular scales were counted in a zig-zag fashion around the middle of the caudal peduncle. Gill-raker counts were of the outer rakers on the first arch, including rudiments; the angle raker is included in the lower-limb (second) count. Counts of pseudobranch filaments included all rudiments. Nomenclature of head pores follows Winterbottom (1986), as modified by Gill et al. (2000). Counts of vertebrae are presented in the form precaudal + caudal 20 = total. Caudal vertebrae are defined as those with a haemal spine, and include the terminal urostylar complex (which was counted as a single vertebra). The pattern of insertion of supraneural (predorsal) bones and anterior dorsal-fin pterygiophores within interneural spaces is given as an ‘anterior dorsal-fin pterygiophore formula’ modified from the “predorsal formula’ of Ahlstrom et al. (1976). Each supraneural is represented by an ‘S,’ neural spines are repres- ented by slashes, and pterygiophores are represented by °3’ (indicating a pterygiophore that bears two supernumerary rays and a serially associated ray), ‘2’ (indicating a pterygiophore that bears a supernumerary ray and a serially associated ray) or “1” (indicating a pterygiophore that bears only a serially associated ray). An ‘anterior anal-fin pterygiophore formula’ is also presented, and is similar to the anterior dorsal-fin pterygiophore formula, except that the slashes represent haemal spines. Osteological features were determined from x-radiographs and from cleared-and-stained speci- mens, which were prepared following the methods of Taylor & Van Dyke (1985). Counts and measurements are given as values or value ranges for all type specimens, followed, where different, by values for the holotype in parentheses. Where counts were recorded bilaterally, both counts are presented for the holotype, separated by a slash; the first count given is the left count. Pseudoplesiops immaculatus sp. nov. Bearded Dottyback Figures 1, 2 Pseudoplesiops typus [non Bleeker, 1858]; Bleeker, 1875: 31 (in part, specimen from Amboina). Chilidichthys [sic] sp. 1; Allen & Steene, 1979: 26 (Christmas Island and Cocos Keeling Atoll, Indian Ocean). Pseudoplesiops revellei [non Schultz, 1953]; Kailola, 1987: 244 (Papua New Guinea); Allen & Steene, 1988: 180, fig. 145 (Christ- mas Island, Indian Ocean); Paxton et al., 1989: 521 (list; Great Barrier Reef). Pseudoplesiops sp.; Gill in Randall et al., 1990: 131 (description; distribution; col. fig.); Randall & Anderson, 1993: 15 (Maldive A.C. GILL AND A.J. EDWARDS Islands); Gill in Randall et al., 1997: 131 (description; distri- bution; col. fig.); Kulbicki & Williams, 1997: 14 (Ouvéa Atoll, New Caledonia). Pseudoplesiops n. sp.; Allen & Smith-Vaniz, 1994: 10 (Cocos (Keeling) Islands). Pseudoplesiops sp. 1.; Gill & Edwards, 1999: 144 (list of osteologi- cal materials); Gill, 2000: 2560 (key). HOLOTYPE. AMS I.20757-069, 29.8 mm SL, Great Barrier Reef, W end of Raine Island (11°36'S 144°O1'E), coral gutter, 2-20 m, AMS and Australian Institute of Marine Science team, 13 February 1979. PARATYPES. AMS I.17090-043, 1: 26.5 mm SL, Papua New Guinea, Madang Harbour, S edge of Massas Island, coral reef, 1.5— 14 m, B.B. Collette et al., 31 May 1970; AMS I.17094-013, 1: 28.5 mm SL, Papua New Guinea, Trobriand Islands, N end of Kiriwinna Island, 0-5 m, B.B. Collette and B. Goldman, 7 June 1970; AMS I.17094-014, 1: 30.4 mm SL, collected with AMS I.17094-013; AMS 1I.17096-008, 1: 27.8 mm SL, Papua New Guinea, Trobriand Islands, Kiriwinna Island, off N coast of Tawai Point, coral with sand patches, 7.5 m, B.B. Collette, 8 June 1970; AMS 1.20756-014, 1: 25.7 mm SL (subsequently cleared and stained), Great Barrier Reef, Great Detached Reef, over coral and sand, 2-8 m, AMS and Australian Institute of Marine Science team, 11 February 1979; AMS 1I.20756-124, 2: 15-30 mm SL, collected with AMS I.20756- 014; AMS 1.20757-039, 1: 29.4 mm SL, collected with holotype; AMS 1.20757-091, 3: 27.6-30.5 mm SL, collected with holotype; AMS 1.20784-040, 2: 17.0-21.9 mm SL, Great Barrier Reef, Yonge Reef, 1 mile N of platform, back reef knoll, 1-15 m, D.F. Hoese et al., | December 1978; AMS I.21972-004, 1: 31.2 mm SL, Solomon Islands, Guadalcanal, 12 km W of Honiara, over wreck, J.E. Randall, 12 July 1975; AMS 1.22616-044, 1: 20.9 mm SL, Great Barrier Reef, Escape Reef, coral reef, 5-18 m, J.R. Paxton et al., 2 Nov- ember 1981; AMS I.22619-025, 1: 26.0 mm SL, Great Barrier Reef, Escape Reef North (15°49'S 45°50'E), sand slope, coral and sand in cave under bommie, 19-22 m, J. Paxton et al., 3 November 1981; ANSP 131727, 2: 27.2-27.5 mm SL, Cocos Keeling Islands, West Island, off N end outside breakers (12°07'50"S 96°48'55"E), Fig. 1 Pseudoplesiops immaculatus, AMS 1.20757-069, 29.8 mm SL, holotype, Raine Island, Great Barrier Reef. (Photo by P. Crabb.) TWO NEW SPECIES OF PSEUDOPLESIOPS eer ie Fig. 2 Cephalic laterosensory pores of Pseudoplesiops immaculatus, AMS 1.20757-069, 29.8 mm SL, holotype, Raine Island, Great Barrier Reef. AIOP, anterior interorbital pore; AN, anterior nostril; DENP, dentary pores; IMF, intermandibular flap; ITP, intertemporal pore; LSCP, pore in tubed lateral-line scale (other details of scale omitted); NASP, nasal pores; PARP, parietal pores; PN, posterior nostril; POPP, preopercular pores; PTP, posttemporal pore; SOBP, suborbital pores; SOTP, supraotic pores. Scaled areas shown in stipple. Scale bar = 2 mm. 0.5—1.3 m relief with sand channels, soft and stony corals, W.F. Smith-Vaniz et al., 24 February 1974; ANSP 131728, 1: 30.6 mm SL, Cocos Keeling Islands, West Island, ca. 1 km NNW of N end of island (12°07'20"S 96°49'05"E), rocky patch fronting on sand, W.F. Smith-Vaniz et al., 1 March 1974; ANSP 178041, 1: 30.8 mm SL, Papua New Guinea, Bougainville Island, Tautsina Island, E of Kieta Peninsula, coral reef off stack at N end of island, 0.9-7.5 m, 11 March 1965 (Te Vega Expedition Cruise no. 6, Station 247); ASIZT 57016, 1: 25.8 mm SL, Taiwan, J.P. Chen, 22 April 1994; ASIZT 56991, 1: 19.7 mm SL, Taiwan, 9 m, J.P. Chen, 21 April 1994; BMNH 1974.5.25.973, 1: 22.8 mm SL, Papua New Guinea, Trobriand Islands, Kiriwinna Island, E shore NW of Gusaweta, exposed coral platform, 0—3.6 m, B.B. Collette, 17 June 1970; CAS 58686, 2: 26.3—-27.9 mm SL, Maldive Islands, Malé Atoll, reef off SE side of Funidu Islet, inside atoll lagoon (04°11'00"N 073°30'30"E), M.G. Bradbury, 6 November 1964; MNHN 1994-45, 1: 22.6 mm SL, New Caledonia, Loyalty Islands, Ouvéa Atoll, Bagaat Islet (20°37'18"S 166°16'08"E), vertical reef wall with small cave and base of wall with coral rubble platform, 15—21 m, J.T. Williams, J.-L. Menou and P. Tirard, 16 November 1991; QM 1.15527, 2: 26.8-33.8 mm SL, collected with holotype; RMNH 31188, 1: 28.7 mm SL, Ambon (Amboina); RUSI 35701, 1: 30.9 mm SL, Taiwan, off Houpihu, P.C. Heemstra, 20 January 1988; USNM 209591, 3: 25.3-27.0 mm SL, Indonesia, point E of Tandjung Naira, Haruka Island, surge channel, 4.5 m, V.G. Springer and M.F. Gomon, 15 January 1973; USNM 209965, 1: 23.7 mm SL, Indone- sia, Saparua, two stations mixed (isolated coral head surrounded by crinkly calcareous matrix at 9 m and coral patch in 3.6 m), V.G. Springer and M.F. Gomon, 18 January 1973; USNM 290118, 6: 17.4-28.9 mm SL, Papua New Guinea, Hermit Islands, Amot Island, ocean side of reef at drop off (01°33'S 144°59'E), 0-15.2 m, V.G. Springer et al., 30 October 1978; USNM 290327, 5: 27.7-29.5 mm SL, Papua New Guinea, S tip of Massas Island (05°10'18"S 145°51'24"E), 0-24 m, V.G. Springer et al., 6 November 1978; USNM 290437, 1: 31.1 mm SL, Papua New Guinea, Ninigo Islands, just SE of Ami Island (01°14'S 144°22'B), patch reef behind reef, 0— 4.5 m, V.G. Springer et al., 22 October 1978; USNM 290749, 3: 23.3-29.9 mm SL, Indonesia, Banda Islands, just W of N tip of Great Banda Island (04°30'30"S 129°56'10"), 0-18 m, V.G. Springer and M.F. Gomon, 9 March 1974; USNM 290791, 1: 30.3 mm SL, Indonesia, Ambon, Latuhalat, Namalatu, about 150 m offshore (03°47'S 128°06'E), 10.5-18 m, V.G. Springer et al., 14 March 1974; USNM 290808, 1: 22.0 mm SL, collected with USNM 290749; USNM 291609, 1: 25.7 mm SL, Philippine Islands, Batanes, 22 Batan Island, past Mahate, White Beach (20°24'45"N 121°55'00"E), coral and encrusted boulders, 9-12 m, G.D. Johnson and W.F. Smith-Vaniz, 1 May 1987; USNM 292031, 1: 27.8 mm SL, Papua New Guinea, Hermit Islands, N side of W entrance (01°30'30"S 144°59°15"E), 0-12 m, V.G. Springer et al., 4 November 1978; USNM 322986, 1: 31.8 mm SL, New Caledonia, Loyalty Islands, Ouvéa Atoll, Motu Veiloa Islet (20°26'06"S 166°28'30"E), reef crest, small patch reef of mostly dead coral surrounded by rubble, 0.6-3.3 m, J.T. Williams and M. Kulbicki, 17 November 1991; USNM 322994, 1: 32.1 mm SL, New Caledonia, Loyalty Islands, Ouvéa Atoll, Récif Draule (20°34'12"S 166°14'12"E), large depres- sion on submerged reef crest, coral and rubble, 5 m, J.T. Williams, P. Tirard and J.L. Menou, 16 November 1991; USNM 328198, 1: 24.5 mm SL, collected with MNHN 1994-45; USNM 356587, 1: 20.9 mm SL, Vanuatu, Shepherd Islands, Judy Reef off NW tip of Tongoa Island (16°52'30"S 168°31'30"E), coral reef with extensive coral development, J.T. Williams et al., 9 June 1996; USNM 357981, 1: 26.8 mm SL, Solomon Islands, Santa Cruz Islands, Reef Islands, Fenualoa Island, just W of Nota Point (ca. 10°16'30"S 166°16'30"E), coral reef face and outer slope, rich coral growth with sand and rubble, 0-13 m, J.T. Williams et al., 18 September 1998; USNM 358382, 1: 20.4 mm SL, Solomon Islands, Santa Cruz Islands, Reef Islands, Lomlom Island, Nialo Point on E side of Forrest Passage (10°16'S 166°18'30"E), vertical reef wall and rocky surge channels at surface, 0-35 m, J.T. Williams er al., 18 September 1998; WAM P.26083-041, 1: 31.2 mm SL, Indian Ocean, Christmas Island, Ethel Beach, 3-6 m, G.R. Allen and R.C. Steene, 19 May 1978; WAM P.26085-030, 1: 25.9 mm SL, Indian Ocean, Christmas Island, Ethel Beach, 15-20 m, G.R. Allen and R.C. Steene, 20 May 1978; WAM P.26093-015, 1: 27.3 mm SL, Indian Ocean, Christmas Island, 1 km W of Margaret Beach, 10 m, G.R. Allen and R.C. Steene, 25 May 1978; WAM P.26104-005, 1: 27.3 mm SL, Indian Ocean, Christmas Island, Flying Fish Cove (10°29'S 105°40'E), 6—8 m, G.R. Allen and R.C. Steene, | June 1978; WAM P.26107-004, 1: 24.0 mm SL, Indian Ocean, Christmas Island, Rhonda Beach (10°29'S 105°40'E), 6-7 m, G.R. Allen and R.C. Steene, 2 June 1978; WAM P.26113- 003, 2: 25.6-28.4 mm SL, Indian Ocean, Christmas Island, Winifred Beach (10°29'S 105°40'E), 12-14 m, G.R. Allen and R.C. Steene, 6 June 1978; WAM P.27825-034, 3: 17.6-25.4 mm SL, Papua New Guinea, Manus Island, Los Negros Island, SE point at aerodrome, 10-40 m, G.R. Allen and R. Knight, 5 October 1982; WAM P.29626- 001, 1: 31.2 mm SL, Papua New Guinea, Port Moresby (09°30'S 147°10'E), 5—6 m, P. Colin, 24 February 1987; WAM P.29927-004, 1: 23.9 mm SL, Indian Ocean, Cocos-Keeling Islands, Direction Island (12°05'S 096°53'E), 0.1-2.0 m, G.R. Allen, 24 February 1989. DIAGNOSIS A species of Pseudoplesiops with the following combination of characters: prominent intermandibular flap present; and operculum immaculate, without large dark spot. DESCRIPTION. (Based on 72 specimens, 15.0—33.8 mm SL) Dor- sal-fin rays 1,26—28 (1,27), last 3-11 (8) segmented rays branched; anal-fin rays I-II,16—18 (1,17), last 3-9 (5) segmented rays branched; pectoral-fin rays 15—18 (16/15), upper 1—5 (2/2) and lower 1-3 (1/1) rays simple; pelvic-fin rays I,3-4 (1,4), all segmented rays simple; principal caudal-fin rays 9-10 + 8 (9 + 8), the uppermost 1—2 (1) and lowermost 1—2 (1) rays unbranched; upper procurrent caudal-fin rays 3—5 (4); lower procurrent caudal-fin rays 2-4 (3); total caudal- fin rays 23—25 (24); scales in lateral series 32—39 (36/35); predorsal scales 7—12 (10); transverse scales 14—17 (16/16); scales behind eye 1-2 (2); scales to preopercular angle 3-4 (3); circumpeduncular scales 16-17 (16); ctenoid scales beginning at 10-16 (14/14) trans- A.C. GILL AND A.J. EDWARDS verse scale rows behind branchial opening; gill rakers 14 + 6-10 = 7-12 (2 + 8); pseudobranch filaments 5-8 (6). Head pores (all bilaterally paired; Fig. 2): nasal pores 2—3 (2/2); anterior interorbital pores 1; posterior interorbital pores 0; supraotic pores 2; suborbital pores 7—9 (8/8); posterior otic pores 0; preopercular pores 7—8 (7/7); dentary pores 4; intertemporal pores 1; anterior temporal pores 0; posttemporal pores 1; parietal pores 2. As percentage of standard length (based on 34 specimens, 22.8— 33.8 mm SL): body depth at dorsal-fin origin 22.8-26.3 (23.8); greatest body depth 23.1—28.1 (26.5); body width 11.5—13.3 (12.4); head length 28.8-32.6 (29.9); snout length 5.3-6.2 (6.0); orbit diameter 7.7—9.9 (8.1); interorbital width 2.2—3.6 (3.0); upper jaw length 10.0—11.2 (10.1); depth of caudal peduncle 13.2—15.3 (14.8); caudal peduncle length 7.3—9.4 (8.1); predorsal length 29.5-32.6 (29.5); preanal length 54.7—57.6 (56.7); prepelvic length 26.4—29.2 (28.2); length of first segmented dorsal-fin ray 8.7—10.8 (9.7); length of third from last segmented dorsal-fin ray 16.4—20.8 (17.4); dorsal-fin base length 60.5—67.8 (63.1); length of first segmented anal-fin ray 9.3—12.3 (11.1); length of third from last anal-fin ray 13.6—20.4 (17.8); anal-fin base length 33.3-37.7 (35.6); caudal fin length 20.4—26.9 (24.8); pectoral fin length 18.4—21.8 (18.5); pelvic fin length 25.7—32.0 (27.8). Lower lip complete; prominent intermandibular flap present (Fig. 2); fin spines weak and flexible; anterior dorsal-fin pterygiophore formula S/S/S + 2/1 + 1, S/S/SY + 2/1 + 1 or S/S/2/ 1 + 1 (S/S/SY + 2/1 + 1); 20-22 (21) consecutive dorsal-fin pterygiophores inserting in 1:1 relationship directly behind neural spine 4; anterior anal-fin pterygiophore formula 2 + 1/1, 3 + 1/1 or 2+ 1 +4 1/1 (2 + I/1); 11-12 (11) consecutive anal-fin pterygiophores inserting in 1:1 relationship directly behind hae- mal spine 2; second segmented pelvic-fin ray longest; caudal fin rounded to truncate or slightly emarginate; scales without distinct centres, and with radii confined to anterior field; dorsal and anal fins without distinct scale sheaths, although sometimes with inter- mittent scales overlapping fin bases; anterior lateral line represented by single tubed scale at branchial opening, followed by intermittent series of centrally pitted scales, which terminate at vertical through base of segmented dorsal-fin ray 21-27, or extend slightly past end of dorsal fin (extending slightly beyond end of dorsal fin/to vertical through base of segmented dorsal-fin ray 26 in holotype); second intermittent series of centrally pitted scales originating on midside above anterior part of anal fin, extending on to middle of caudal-fin base; additional centrally pitted scales present on caudal-fin base, pits usually aligned vertically on pos- terior part of scale sheath; scales present on cheeks (not extending posteriorly over upper part of preopercle) and operculum (Fig. 2); predorsal scales extending anteriorly to supratemporal commis- sure (Fig. 2); vertebrae 12 + 20-22 (12 + 21); epurals 2; epineurals present on vertebrae | through 17—21 (1 through 19); ribs present on vertebrae 3 through 11-12 (3 through 12), rib on ultimate precaudal vertebra very small to moderately small or absent. Upper jaw with 2-6 pairs of curved, enlarged caniniform teeth anteriorly, the medial pair smallest, and 3-4 (at symphysis) to 1—2 (on sides of jaw) irregular inner rows of small conical teeth, the teeth of outer row of conical teeth largest; lower jaw with 2-4 pairs of curved, enlarged caniniform teeth, the medial pair smallest, and 2— 4 (at symphysis) to 1 (on sides of jaw) irregular inner rows of small conical teeth, the conical teeth gradually increasing in size and becoming more curved on middle part of jaw, then becoming abruptly smaller on posterior part of jaw; vomer with | row of small, stout conical teeth arranged in a chevron; palatines edentate or with small irregular patch of small conical teeth; tongue acutely pointed, edentate. TWO NEW SPECIES OF PSEUDOPLESIOPS LIVE COLORATION. (Based on colour photographs of the holotype from the Great Barrier Reef, and of paratypes and other specimens from the Great Barrier Reef, Loyalty Islands, Solomon Islands, Vanuatu, Christmas Island and Taiwan) Head and body pinkish or yellowish brown to olive or bright green, sometimes becoming pinkish to orangish brown on snout, lips and intermandibular flap; iris pale yellow to green or brown, some- times with reddish grey to pink area around pupil; dorsal and anal fins yellowish, pinkish or orangish brown to olive or bright green, becoming paler distally, with bluish grey to bright blue distal margin, sometimes with broad pale orange to yellow stripe submarginally; bluish grey to pale blue spot or streak at base of alternate dorsal- and anal-fin rays; caudal fin yellowish or pinkish brown to olive or bright green; pectoral fins pinkish, greenish or yellowish hyaline; pelvic fins pale pink or olive to bright green, usually bluish grey to pale blue anteriorly and distally. PRESERVED COLORATION Head and body pale brown, slightly darker on dorsal part of head and body, and on lips and intermandibular flap; fins pale brown to brownish hyaline; bluish grey to blue spots and distal margins of dorsal and anal fins described above become greyish brown. HABITAT AND DISTRIBUTION This species is distributed from the Maldive Islands, east and south to Vanuatu, and north to Taiwan. It has been collected from a variety of reef habitats, from shallow patch reefs to reef walls at depths ranging to at least 20 m (with some collections perhaps from as deep as 40 m). COMPARISONS WITH OTHER SPECIES Pseudoplesiops immaculatus has been confused with P. revellei Schultz, a Pacific Plate endemic (sensu Springer, 1982). The two species closely resemble each other in morphometric and meristic values, and both possess a prominent intermandibular flap (Fig. 2), although this may be weakly developed or absent in small specimens (smaller than about 18 mm SL). A low intermandibular ridge or weak flap may be present in certain other pseudochromid species (e.g., Chlidichthys cacatuoides, see Gill & Randall, 1994; Pseudoplesiops occidentalis, Fig. 4), but only in P. immaculatus and 23 P. revellei is it well-developed. On the basis of this synapomorphy, we suggest that the two species are sister taxa. They are distin- guished from each other by a single coloration character: P. revellei has a large, dark brown to black spot on the operculum, which is absent in P. immaculatus. REMARKS Colour photographs of the species have been published by Allen & Steene (1988; as P. revellei), and Randall et al., (1990, 1997 - as P. sp.). ETYMOLOGY The specific epithet is from the Latin, meaning without a spot, and alludes to the sole character distinguishing P. immaculatus from its sister species. Pseudoplesiops occidentalis sp. nov. Maldives Dottyback Figures 3, 4 Clinus sp.; Regan, 1902: 276 (description; Haddumati, Maldive Islands). Pseudoplesiops sp.; Randall & Anderson, 1993: 15, pl. 3e (Maldive Islands; col. fig.). Pseudoplesiops sp. 1; Kuiter, 1998: 77 (habitat notes; colour photo). HOLOTYPE. BPBM 32926, 24.9 mm SL, Maldive Islands, South Malé Atoll, Maaniyafushi Island, reef, 25-30 m, J.E. Randall, C. Anderson and M.S. Adam, 17 March 1988. PARATYPES. AMS 1.41004-001, 1: 21.3 mm SL (subsequently cleared and stained), collected with holotype; BMNH 1901.12.31.77, 1: 26.2 mm SL, Maldive Islands, Haddummati Atoll (= Haddumati), 72 m, J.S. Gardiner; BPBM 32871, 2: 16.7—26.5 mm SL, Maldive Islands, Ari Atoll, E side of reef N of Bathala Island, rubble bottom, 35 m, J.E. Randall and M.S. Adam, 6 March 1988. DIAGNOSIS Pseudoplesiops occidentalis is distinguished from all other pseudo- plesiopines in having the following combination of characters: dorsal-fin rays II,23; scales in lateral series 26—28; and scales with distinct centres and radii in all fields. Fig. 3 Pseudoplesiops occidentalis, BPBM 32926, 24.9 mm SL, holotype, South Malé Atoll, Maldive Islands. (Photo by P. Crabb.) 24 A.C. GILL AND A.J. EDWARDS LSCP —S Fig.4 Cephalic sensory pores of Pseudoplesiops occidentalis, BPBM 32926, 24.9 mm SL, holotype, South Malé Atoll, Maldive Islands. Arrow indicates atypical doubled suborbital pore (present unilaterally only in holotype). IMR, intermandibular ridge. Other abbreviations and methods of presentation follow Fig. 2. DESCRIPTION. (Based on five specimens, 16.7—26.5 mm SL) Dorsal-fin rays II,23, last 6-7 (6) segmented rays branched; anal-fin rays II,14, last 5-6 (5) segmented rays branched; pectoral-fin rays 15-16 (16/15), upper 2-4 (2/2) and lower 1-3 (1/2) rays simple; pelvic-fin rays [,3, all segmented rays simple; principal caudal-fin rays 9 + 8, the uppermost 1-2 (1) and lowermost 1-2 (2) rays unbranched; upper procurrent caudal-fin rays 3—4 (4); lower procurrent caudal-fin rays 3-4 (4); total caudal-fin rays 23-25 (25); scales in lateral series 26-28 (27/27); predorsal scales 6-8 (8); transverse scales 11-12 (11/11); scales behind eye 2; scales to preopercular angle 3; circumpeduncular scales 16; ctenoid scales beginning at 7—9 (7/9) transverse scale rows behind branchial opening; gill rakers 2-4 + 8-11 = 10-15 (2 + 8); pseudobranch filaments 6-7 (7). Head pores (all bilaterally paired; Fig. 4): nasal pores 2; anterior interorbital pores 1; posterior interorbital pores 0; supraotic pores 2; suborbital pores 8—9 (9/8); posterior otic pores 0; preopercular pores 7; dentary pores 4; intertemporal pores 1; anterior temporal pores 0; posttemporal pores 1; parietal pores 2. As percentage of standard length: body depth at dorsal-fin origin 24.0—28.2 (26.5); greatest body depth 24.0-32.4 (27.7); body width 13.3-14.5 (13.3); head length 30.5—33.5 (30.5); snout length 5.0— 6.4 (5.6); orbit diameter 8.4—10.2 (8.4): interorbital width 3.64.8 (3.6); upper jaw length 11.3—-12.0 (11.6); depth of caudal peduncle 16.5-16.9 (16.5); caudal peduncle length 7.2-8.5 (7.2); predorsal length 29.0-32.9 (31.3); preanal length 55.0-61.1 (60.2); prepelvic length 30.1-32.5 (30.1); length of first segmented dorsal-fin ray 10.4-11.7 (10.4); length of third from last segmented dorsal-fin ray 17.7-19.7 (17.7); dorsal-fin base length 61.7—71.4 (62.7); length of first segmented anal-fin ray 12.2—14.5 (12.9); length of third from last anal-fin ray 17.8—18.7 (18.5); anal-fin base length 32.8-35.9 (33.7); caudal fin length 25.3—27.9 (25.3); pectoral fin length 22.5— 24.0 (23.3); pelvic fin length 27.7—35.3 (27.7). Lower lip complete; prominent intermandibular flap absent, al- though low ridge present in some specimens (Fig. 4); fin spines weak and flexible; anterior dorsal-fin pterygiophore formula S/S/3/ 1 + 1; 15-16 (15) consecutive dorsal-fin pterygiophores inserting in 1:1 relationship directly behind neural spine 4; anterior anal-fin pterygiophore formula 2/1 + 1; 7—8 (8) consecutive anal-fin pterygiophores inserting in 1:1 relationship directly behind haemal spine 2; second segmented pelvic-fin ray longest; caudal fin weakly rounded to rounded or slightly emarginate; scales with distinct centres and radii in all fields; dorsal and anal fins without distinct scale sheaths, although with basal row of body scales overlapping fin base, particularly on posterior part of fins; anterior lateral line represented by single tubed scale at branchial opening, followed by intermittent series of centrally pitted scales, which terminate at vertical through base of segmented dorsal-fin ray 17—22 (2/22): TWO NEW SPECIES OF PSEUDOPLESIOPS second intermittent series of centrally pitted scales originating on midside above anterior part of anal fin, extending on to middle part of caudal-fin base; additional centrally pitted scales present above and below pitted scale(s) on middle part of caudal-fin base, some- times extending on to posterior part of caudal peduncle; scales present on cheeks (extending posteriorly over upper part of preopercle) and operculum (Fig. 4); predorsal scales extending anteriorly to point between anterior interorbital pores and vertical through anterior supraotic pores (Fig. 4); vertebrae 10 + 17; epurals 2; epineurals present on vertebrae 1 through 16—18 (1 through 16); ribs present on vertebrae 3 through 10, rib on ultimate precaudal vertebra relatively long. Upper jaw with 2-5 pairs of curved, enlarged caniniform teeth anteriorly, the medial pair smallest, and 34 (at symphysis) to 1-2 (on sides of jaw) irregular inner rows of small conical teeth, the teeth of outer row of conical teeth largest; lower jaw with 2-3 pairs of curved, enlarged caniniform teeth, the medial pair smallest, and 2— 3 (at symphysis) to | (on sides of jaw) irregular inner rows of small conical teeth, the conical teeth gradually increasing in size and becoming more curved on middle part of jaw, then becoming abruptly smaller on posterior part of jaw; vomer with | row of small, stout conical teeth arranged in a chevron; palatines edentate or with small irregular patch of small conical teeth; tongue pointed, eden- tate. LIVE COLORATION. (Based on colour photograph of the holotype, photograph in Kuiter, 1998, and field notes taken from paratypes in BPBM 32871 when freshly dead) Head and body bright pinkish red to orange-red, becoming pink ventrally and olive-red to orangish brown posteriorly; margin of orbit orange posteriorly, becoming pale blue ventrally; posttemporal, intertemporal, upper preopercular and upper suborbital pores indis- tinctly edged with grey; iris bright orange-red; pectoral-fin base pale pink to pinkish red; dorsal and anal fins bright orange-red basally, reddish hyaline distally, with pale blue distal margin; caudal fin greyish yellow to pale greyish red basally, remainder of fin reddish to yellowish hyaline with pale blue distal margin; pectoral fin pinkish to yellowish hyaline; pelvic fin pale pink basally, becoming pale blue distally. PRESERVED COLORATION Head and body brown, paler ventrally; grey edging on posttemporal, intertemporal, upper preopercular and upper suborbital pores re- mains, becoming greyish brown; dorsal and anal fins brownish hyaline, becoming greyish brown distally; other fins pale brown to dusky hyaline. HABITAT AND DISTRIBUTION Pseudoplesiops occidentalis is known only from the Maldive Is- lands. It has been collected from and observed on reefs in 20 to 72 m. COMPARISONS WITH OTHER SPECIES Pseudoplesiops occidentalis forms a monophyletic group with P. typus Bleeker, P. rosae Schultz and several undescribed species. With the exception of P. occidentalis, this clade is confined to the eastern Indian Ocean and the West Pacific. It is diagnosed by scales with distinct centres and radii in all fields. Species limits within the clade are poorly resolved and are currently under study by us. For the purposes of comparison with P. occidentalis, we divide the clade into three subgroups, each of which we believe to be monophyletic: P. occidentalis, P. rosae-complex (autapomorphy: plate-like, median expansion of median ethmoid, pterosphenoid and basisphenoid into orbital space); and P. typus-complex (autapomorphy: all scales cycloid in adult specimens). Aside from the various autapomorphies listed above, the three taxa are distin- 25 guished from each other by the following: number of vertebrae (10 + 17 in P. occidentalis, 11-12 + 16-18 = 27-29, usually 11 + 17- 18 in the P. rosae-complex and 11 + 17-18 in the P. typus-complex); number of dorsal-fin rays (II,23 in P. occidentalis, 1,22—24 in the P. rosae-complex and II,24—25 in the P. typus- complex); number of anal-fin rays (II,14 in P. occidentalis, I-II,12—14, usually I,13-14 in the P. rosae-complex and II-III, 14— 16, usually II,15 in the P. typus-complex), number of scales in lateral series (26—28 in P. occidentalis, 26-29 in the P. rosae- complex and 32-40 in the P. typus-complex); number of circumpeduncular scales (16 in P. occidentalis, 16 in the P. rosae- complex and 20-22 in the P. typus-complex); predorsal scalation (6-8 scales, extending anteriorly to a point between the anterior interorbital pores and the vertical through the anterior supraotic pores in P. occidentalis, 5-10 scales, extending anteriorly to a point between the anterior interorbital pores and the vertical through the anterior supraotic pores in the P. rosae-complex, and 10-16 scales, extending anteriorly to the supratemporal commissure in the P. typus-complex); and cheek scalation (broadly overlapping the upper part of the preopercle in P. occidentalis and the P. rosae- complex, versus not overlapping the upper part of the preopercle in the P. typus-complex). Members of the P. typus-complex also attain a much larger body size than members of the other clades (largest examined specimen 53.0 mm SL versus 26.5 mm SL in P. occidentalis and 26.9 mm SL in the P. rosae-complex). Of the characters noted above that are shared by P. occidentalis and the P. rosae-complex, three are unique within Pseudoplesiops and suggest a sister-relationship between the two taxa: low number of scales in lateral series; predorsal scales extending anteriorly beyond the supratemporal commissure; and cheek scales broadly overlapping upper part of preopercle. This relationship will be tested in a study of the phylogeny of the Pseudochromidae currently in progress by the first author. REMARKS Colour photographs of the species are provided by Randall & Anderson (1993) and Kuiter (1998). Randall & Anderson indicate that their photograph is of a 26 mm SL specimen in BPBM 32871, but it is actually of the holotype (BPBM 32926). Regan’s (1902) specimen of the species (BMNH 1901.12.31.77) was initially identified and reported on as “Clinus sp.” A label on the jar and catalogue entry indicates that it was subsequently redetermined as the plesiopid Belonepterygion fasciolatum Ogilby (apparently by M.L. Penrith). Although we were unable to locate any references to this identification, they possibly exist. ETYMOLOGY The specific epithet is from the Latin, meaning of the west; P. occi- dentalis is known only from the western-most part of the range of the genus. ACKNOWLEDGEMENTS. We thank the following for allowing access to specimens in their care: G.R. Allen, A. Bentley, D. Catania, J.-P. Chen, O.A. Crimmen, W.N. Eschmeyer, P.C. Heemstra, D.F. Hoese, S. Jewett, J. Johnson, M. McGrouther, R.J. McKay, V. Mthombeni, L. Palmer, K. Parkinson, J.E. Randall, S.E. Reader, T. Trnski, M. Sabaj, W. Saul, K.-T. Shao, W.F. Smith- Vaniz, A. Y. Suzumoto, M.J.P. van Oijen and J.T. Williams. Colour photographs were kindly provided by J.-P. Chen, D.F. Hoese, R.H. Kuiter, J.E. Randall and J.T. Williams; P. Crabb took the black and white photographs of the two holotypes (Figs 1 and 3). O.A. Crimmen, S. Davidson, J.P. Garcia, A.-M. Hine, K. Parkinson, S. Raredon and S.E. Reader radiographed specimens. R.D. Mooi and J.T. Williams critically reviewed the manuscript and provided helpful comments. 26 REFERENCES Ahlstrom, E.H., Butler, J.L. & Sumida, B.Y. 1976. Pelagic stromateoid fishes (Pisces, Perciformes) of the eastern Pacific: kinds, distributions, and early life histories and observations of five of these from the northwest Atlantic. Bulletin of Marine Science 26(3): 285-402. Allen, G.R. & Steene, R.C. 1979. The fishes of Christmas Island, Indian Ocean. Australian National Parks and Wildlife Service, Special Publication 2: 1-81. —-— & 1988. Fishes of Christmas Island, Indian Ocean. 197 p. Christmas Island Natural History Association. & Smith-Vaniz, W.F. 1994. Fishes of the Cocos (Keeling) Islands. Atoll Research Bulletin 412: 1-21. Bleeker, P. 1858. Bijdrage tot de kennis der vischfauna van den Goram Archipel. Naturrkundig Tijdschrift Nederlandsch Indié 15: 197-218. 1875. Sur la famille des Pseudochromoides et révision de ses espéces insulindiniennes. Verhandelingen der Koninklijke Akademie van Wetenschappen, Amsterdam 15: 1-32, pls 1-3. Gill, A.C. 2000. Pseudochromidae, pp 2557-2577. In: Carpenter, K.E. & Niem, V.H. (eds) FAO Species Identification Guide for Fisheries Purposes. The Living Marine Resources of the Western Central Pacific. Vol. 4. FAO, Rome. & Edwards, A.J. 1999. Monophyly, interrelationships and description of three new genera in the dottyback fish subfamily Pseudoplesiopinae (Teleostei: Perciformes: Pseudochromidae). Records of the Australian Museum 52(1): 141-160. , Mooi, R.D. & Hutchins, J.B. 2000. Description of a new subgenus and species of the fish genus Congrogadus Giinther from Western Australia (Perciformes: Pseudochromidae). Records of the Western Australian Museum 20(1): 69-79. & Randall, J.E. 1994. Chlidichthys cacatuoides, anew species of pseudoplesiopine dottyback from Oman, with a diagnosis of the genus Chlidichthys Smith, and new record of Pseudochromis punctatus Kotthaus from Oman (Teleostei: Perciformes: A.C. GILL AND A.J. EDWARDS Pseudochromidae). Revue francaise d’Aquariologie Herpétologie 21(1-—2): 11—18. Kailola, P.J. 1987. The fishes of Papua New Guinea: a revised and annotated checklist. Volume two. Scorpaenidae to Callionymidae. Papua New Guinea Department of Fisheries and Marine Resources Research Bulletin 41: 195-418. Kuiter, R.H. 1998. Photo Guide to Fishes of the Maldives. 257 p. Atoll Editions, Apollo Bay. Kulbicki, M. & Williams, J.T. 1997. Checklist of the shorefishes of Ouvéa Atoll, New Caledonia. Atoll Research Bulletin 444: 1-26. Leviton, A.E., Gibbs, R.H., Jr., Heal, E. & Dawson, C.E. 1985. Standards in her- petology and ichthyology: part 1. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985(3): 802-832. Paxton, J.R., Hoese, D.F., Allen, G.R. & Hanley, J.F. 1989. Zoological Catalogue of Australia. Volume 7. Pisces. Petromyzontidae to Carangidae. 665 p. Australian Government Publishing Service, Canberra. Randall, J.E., Allen, G.R. & Steene, R.C. 1990. Fishes of the Great Barrier Reef and Coral Sea. 507 p. Crawford House Press, Bathurst. - & . 1997. Fishes of the Great Barrier Reef and Coral Sea. Revised and expanded edition. 557 p. Crawford House Press, Bathurst. & Anderson, R.C. 1993. Annotated checklist of the epipelagic and shore fishes of the Maldive Islands. Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology 59: 1-47, pls 1-8. Regan, C.T. 1902. On the fishes from the Maldive Islands. The Fauna and Geography of the Maldive and Laccadive Archipelagos 1(3): 272-281. Springer, V.G. 1982. Pacific Plate biogeography, with special reference to shorefishes. Smithsonian Contributions to Zoology 367: 1-182. Taylor, W.R. & Van Dyke, G.C. 1985. Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium 9(2): 107-119. Winterbottom, R. 1986. Revision and vicariance biogeography of the subfamily Congrogadinae (Pisces: Perciformes: Pseudochromidae). Indo-Pacific Fishes 9: 1— 34, pl. I. [dated 1985, but actually published 1986] Me (as2et5. 1) Bull. nat. Hist. Mus. Lond. (Zool.) 68(1): 27-37 ~\ Issued 27 June 2002 A redescription of Sousa chinensis (Osbeck, 1765) (Mammalia, Delphinidae) and designation of a neotype L.J. PORTER The Swire Institute of Marine Science, The University of Hong Kong, Cape d’Aguilar, Shek O, Hong Kong Special Administrative Region, China. Synopsis. The holotype of the delphinid Sousa chinensis, held in the collection of the Royal College of Surgeons, London, was destroyed during the Second World War. The neotype is described herein from a male specimen obtained from the waters of Hong Kong (the Hong Kong Special Administrative Region of China since | July 1997). This dolphin was found newly dead in May 1996 and death determined as heart failure brought about by entanglement with fishing gear. There is currently debate over divisions within the genus Sousa although recent reviews suggest that the five nominal species currently designated likely comprise but one. A description of the neotype’s external appearance and skeletal elements is given and compared with the information that remains for the holotype and for other specimens obtained from Hong Kong waters. INTRODUCTION The species Sousa chinensis (formerly Delphinus sinensis) (Osbeck, 1765) was designated after observations were made of a population of delphinids within the Pearl River Estuary, Guangdong Province, Southern China, i.e. close to Hong Kong. It was over one hundred years after the species was named, however, that the first skeleton was described from an almost entire individual collected from Xiamen (formerly Amoy), Fujian Province, approximately 700km northeast of Hong Kong (Flower, 1870). This specimen and, thus, the holotype of the species was held in the collections of the Royal College of Surgeons, London, but was subsequently destroyed during the Second World War. Osbeck’s original account noted ‘Snow-white Dolphins tumbled about the ship; but at a distance they seemed nothing different from the common species, except in their white colour’. The holotype was physically mature and although the external appearance of this particular individual was not described, the typical coloration of members of the Xiamen population was noted to be ‘milky white, with pinkish fins and black eyes’ (Flower, 1870). The individuals that remain in the Pearl River and Jiulong Jiang River (Xiamen) Estuaries today, are similar in external appearances (Porter, 1998). Four other species are included in the genus Sousa (Table 1), although there has been debate as to where the taxonomic divisions within it, if any, actually occur (Pilleri & Gihr, 1972; Mitchell, 1975; Zhou et al., 1980; Wang & Sun, 1982; Ross et al., 1994). The most recent classification of the genus as comprising one species was based upon a morphological review (Ross ef al., 1996) and a molecular investigation of the genus concurred with this definition (Cockcroft et al., 1997). Both of these publications, however, noted that the limited availability of data from parts of the species’ range, particularly Asia, did not allow definitive segregation at a sub- specific level and, as such, the taxonomy of the genus is not resolved. There have been many strandings of Sousa chinensis in Hong Kong waters, which comprise the eastern portion of the Pearl River Estuary, since 1990 (Porter et al., 1998) although the cause of death in many cases could not be determined due to either the decomposi- tion of, or damage to, the remains. In May 1996, however, a fresh © The Natural History Museum, 2002 specimen identified as S. chinensis was retrieved and on which a full necropsy was performed. The cause of death was determined as heart failure, most likely due to entanglement with fishing gear, as deduced by the presence of fresh net abrasions on the rostrum and neck area. This specimen has been chosen for designation as the neotype of S. chinensis because: (a), it was obtained from the area from which the species was originally identified; (b), it was neither significantly different from the description of the extant holotype nor other individuals measured from the Hong Kong population of S. chinensis; (c), its death was perceived to be accidental rather than a consequence of any ailment or abnormality and (d), its remains were intact. MATERIALS AND METHODS A modification of the criteria described by Perrin (1975) was used to measure the skeleton obtained from Hong Kong and herein desig- nated as the neotype. The tympanoperiotic bones were measured in accordance with Kasuya (1973). Other skeletal remains of the same species were also examined and measured to obtain a better range of values for the species in this component of its range. All measure- ments were made with EPI precision calipers and recorded to the nearest millimetre or, in the case of smaller bones, 0.1 mm. Each measurement was taken several times until a confident value was established. An inventory of the measurements recorded for the neotype is included as Appendix I. The neotype and other Hong Kong specimens were aged from three teeth, selected from the upper left jaw, which were sliced to a thickness of 110um. The sections were etched for one hour by emersion in 5% formic acid and the dentinal growth layer groups then counted under a light microscope. One pair of dark and light stained layers is equivalent to one dentinal growth layer group which is assumed to be gained annually (Kasuya, 1976). The equivalent holotype measurements of Sousa chinensis (Flower, 1870) were transformed into metric units and compared with corresponding values for the neotype. The original drawings of Flower’s (1870) and photographs of the neotype were also com- pared. The skeletal characters of both the holotype and neotype were 28 L.J. PORTER pes aeesse ; 5% gurere® Fig. 1 The skull and jaw bones of the neotype of Sousa chinensis seen from a, lateral; b, ventral; c, ramus dorsal and d, dorsal aspects REDESCRIPTION OF THE CHINESE WHITE DOLPHIN Table 1 Five species classification of the genus Sousa according to morphological differences and geographical distribution (Ross et al., 1994). Nominal species Geographical range S. chinensis (Osbeck, 1765) S. plumbea (Cuvier, 1829) S. lentigenosa (Owen, 1866) S. teuzii (Kukenthal, 1892) S. borneensis (Lydekker, 1901) China to Australia South Africa to India India to Southeast Asia West Africa Rare form in Borneo, overlapping with ‘chinensis’ then compared with the range of measurements obtained from other dead adult specimens of S. chinensis stranded on Hong Kong beaches. 29 RE-DESCRIPTION OF SOUSA CHINENSIS (OSBECK, 1765) Neotype: ZD 1999.360, sub-adult male, N 022° 20' 00; E 113° 55’ 30, The Brothers Islands, Hong Kong’s territorial waters, collected by staff of the Swire Institute of Marine Science, The University of Hong Kong. Donated to the Natural History Museum, London, under CITES permit APO/EL 3227/99. The gently-tapering rostrum of this specimen comprised 60.7% of the total skull length and was positioned low near the base of the cranium (Fig. la). The upper and lower tooth rows comprised 89% and 63%, respectively, of the rostrum and ramus lengths, with 3 1—35 Fig. 2 The a, foramen magnum; b, atlas and c, axis of the neotype of Sousa chinensis 30 teeth in each row (Fig. 1b & c). The teeth were conical and pointed with little wear. Five dentinal layers were counted from sectioned teeth and the pulp cavity was open. The premaxillaries were in contact for almost the entirety of the rostrum length, with only a small section of the vomer exposed. Storage of the specimen in a dry environment, however, has subsequently caused the premaxillaries to gape (Fig. 1d). The premaxillaries were roughened dorsally where they met the less dense maxillaries, at the base of the nasal aperture. The posterior extremity of the rounded, right, premaxillary extended to the anterior of the nasal aperture, whereas the pointed left premaxillary was truncated at the mid-margin of the opening. The premaxillaries, therefore, encircled two thirds of the aperture, in L.J. PORTER which each nasal passage was deep-set and elliptical. The antorbital notch was not developed. The two occipital condyle margins were sharply defined and the anterior margin pointed. The channel between the condyles was long and triangular. The foramen magnum was ovoid with a slightly extended dorsal margin (Fig. 2a). Both the temporal and post-temporal fossae were ovoid with sharply-defined margins. The temporal fossae protruded where they met which, in ventral view, gave the cranium a relatively broad appearance. The orbits were rounded and the lacrimals, positioned at right angles to the orbit, were blunt with prominent dorsal ridges. The cranial articulating surface of the atlas was broad with short, transverse processes and a robust, relative to the rest of the vertebrae, neural Fig. 3 The a, vertebral column; b, vertebral ribs and c, sternal and free floating ribs of the neotype of Sousa chinensis REDESCRIPTION OF THE CHINESE WHITE DOLPHIN spine (Fig. 2b). The axis was not fused to the atlas and had short processes but a well-developed neural spine (Fig. 2c). There were seven cervical, 12 thoracic, 23 lumbar and 18 caudal vertebrae. No vertebrae were fused (Fig. 3a). The first vertical perforating foramen appeared on the 25th vertebra which was also the first to have greatly reduced metapophyses and the last with distinct transverse and neural processes. There were 12 vertebral ribs on both the left and right sides, six of which on each side were double-headed (Fig. 3b). There were five sternal ribs on each side (Fig. 3c). The sternum was notched with a perforation on the left side and two lateral processes on each side (Fig. 4a; note that the right side of the sternum was damaged during preparation). There was one Sil mesosternal element which equalled two thirds of the manubrium. The radius and ulna were longer than the humerus, although the latter was wider and more robust. The transverse breadth of the carpals was short and the pectoral fin elongated (Fig. 4b). The scapula had a sharply pointed coracovertebral angle, a broad metacromion process and a smaller, pointed, coracoid process (Fig. 4c). The bullae of the tympanoperiotic bones were rounded and bulbous and the dorsal plane of the periotic convex; the fundus of the internal auditory meatus and the aquaductus fallopii were ovoid; the aquaductus cochleae had a distinct ridge; the join between the cochlear portion and the rest of the periotic was rounded and smooth. Fig.4 The a, sternum and b, pectoral fin skeletal elements of the neotype of Sousa chinensis WW i) HOLOTYPE (a) occipital condyles NEOTYPE HOLOTYPE NEOTYPE L.J. PORTER Fig.5 A copy of the original illustrations of Flower’s (1870) holotype and the corresponding views of the neotype showing differences in the a, occipital condyles; b, lacrimal flares and c, temporal fossae REDESCRIPTION OF THE CHINESE WHITE DOLPHIN COLORATION. Mainly pink with light grey around the head and darker pigmentation along the length of the back, including the leading and trailing edge of the dorsal fin and the ventral fringe of the fluke. There was dark, fragmented pigmentation on the melon, around the eyes and across the dorsal cape. Although both the dorsal and ventral surfaces of the fluke were pink, the dorsal surface of the pectoral fins were still grey and had only just begun to lighten. EXTERNAL APPEARANCE. The body was slender but with a promi- nent melon. Neither the ‘robustness’ nor prominent peduncle, typical of other, adult individuals observed and examined from Hong Kong waters were present (Porter, 1998). The dorsal fin was slightly falcate and the flukes were small with a distinct notch. The pectoral fins were broad in the middle but tapered both distally and at the base. The ‘hump-back’, reportedly typical of populations west of Indonesia, has not been recorded from any individual in the South China Sea (Reeves & Leatherwood, 1994). COMMENTS. During the post-mortem it was noted that the indi- vidual had deep scarring around the pectoral fins and torso. This healed injury was consistent with abrasion from either a rope or a similar object. It cannot be determined if this injury caused any major deformity to the pectoral fin bones although some pathology of them was observed. These injuries were distinct from those associated with the animal’s death. ETYMOLOGY. The species name is derived from its geographical location, i.e. both sinensis and chinensis referring to ‘China’. The meaning of the current genus name Sousa has become obscure. Although the taxonomy of the genus Sousa has yet to be resolved, in accordance with the International Code of Zoological Nomencla- ture, this neotype must bear the name of the holotype. COMPARATIVE ANALYSIS OF SPECIMENS The age of nine Hong Kong specimens of Sousa chinensis used for comparative skeletal purposes in this study, ranged between 4 and 14 growth layer groups, the former number having been established as the age where rapid juvenile growth rate decreases and adult dimensions are gained (Porter, 1998). No Hong Kong specimen had reached physical maturity as determined by fusion of post-cranial elements and closure of the tooth pulp cavity. There was, therefore, no physically mature specimen to compare with the holotype. The most obvious difference, therefore, between the skeletal measure- ments available for the holotype and those for the neotype is that of size (Table 2). The holotype had two less lumbar vertebrae, four more caudal vertebrae and six more chevron bones than the neotype. The holotype had slightly fewer teeth than the neotype but both counts are within the range identified for other Hong Kong speci- mens (Table 3). A comparison of the remaining diagrams of the holotype with the neotype illustrates clearly the similarities in general shape and skull configuration. Both have large ovoid crani- ums with robust and elongated rostrums, the eliptical nares are deep set and the vomer is exposed (Fig. 5). Differences are apparent, however, in the occipital condyles (a) and the lacrimal flares (b), which were more prominent in the holotype. The shape of the holotype temporal fossae also differed from those of the neotype (c): in the former they were swept back towards the posterior of the skull resulting in a smaller width between them. There are differences in the degree of ossification of the scapulae, the holotype being more robust with larger coracoid and metacromion processes (Fig. 6). The measurements obtained from the holotype, the neotype and 85 Table 2 Sousa chinensis principal skeletal dimensions and vertebral counts of the Hong Kong neotype and the holotype from Fujian Province (Flower, 1870). Measurement Neotype Holotype 1 Condylobasal length 481 526 2 Length of rostrum. 292 325 3 Width of rostrum 110.1 I) 5 Width of rostrum at midlength 44.8 47 16 Greatest parietal width 140.9 163 32 Length upper tooth row to tip rostrum 260 284 33(UL) Number of teeth (UL) 35 33 34(UR) Number of teeth (UR) 35 32 35(LL) Number of teeth (LL) 31 32 36(LR) Number of teeth (LR) 32 31 a7 Length of lower tooth row. 259 279 38 Greatest length of left ramus. 407 457 39 Greatest height of left ramus. 82.5 91 40a Length of symphysis 113.3 140 48a Number of cervical vertebrae i 7 48 Number of thoracic vertebrae 12 12 49 Number of lumbar vertebrae 12 10 50 Number of caudal vertebrae 18 22 51 Total number of vertebrae 49 51 78 Number of vertebral ribs (left) 12 12 79 Number of vertebral ribs (right) 12 12 96 Number of chevron bones 8 14 105 Greatest length of coracoid process. 21.5 37 106 Greatest width of coracoid process. 11.6 29 107 Greatest width of metacromion process. 54.5 30 108 Greatest length of humerus. 63.9 73 109 Greatest width of humerus distally. 39:5 57 110 Greatest length of radius. 74 79 111 Greatest width of radius distally. 38.2 47 112 Greatest length of ulna. 65.9 67 HOLOTYPE coracoid process metacromion process NEOTYPE Fig. 6 A copy of the original illustrations of Flower’s (1870) holotype and the corresponding views of the neotype demonstrating differences in the form of the coracoid and metacromion processes of the scapula. 34 550 5 500 450 400 350 a. = 3 300 (3) = oO 2 250 uo} oO ni = n 200 150 100 50 320 - 270 - = 220 - = — eo = S | is 170 oe = =| i“) S Une Viet erarne 1" 1) ae = , ane cara 0) Leatelins page) i 04 ye dey ay Gl at cpa geiyeAW, OS eee th > a a — — es ~ ne