ISSN 0968-0470
Bulletin of The Natural History Museum
PKfcSfc WTED gg frgflAL UBQARY Zoology Series j
VOLUME 68 NUMBER 1 27 JUNE 2002 1
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.
SUBSCRIPTIONS Bulletin of the Natural History Museum, Zoology Series (ISSN 0968-0470) is published twice a year (one volume per annum) in June and November. Volume 68 will appear in 2002. The 2002 subscription price (excluding VAT) of a volume, which includes print and electronic access, is £88.00 (US $155.00 in USA, Canada and Mexico). The electronic- only price available to institutional subscribers is £79.00 (US $140.00 in USA, Canada and Mexico).
ORDERS Orders, which must be accompanied by payment, may be sent to any bookseller, subscription agent or direct to the publisher: Cambridge University Press, The Edinburgh Building, Shaftesbury Road, Cambridge CB2 2RU, UK; or in the
USA, Canada and Mexico: Cambridge University Press, Journals Department, 40 West 20th Street, New York, NY 1011 -42 1 , USA. EU subscribers (outside the UK) who are not registered for VAT should add VAT at their country's rate. VAT registered members should provide their VAT registration number. Japanese prices for institutions (including ASP delivery) are available from Kinokuniya Company Ltd, P.O. Box 55, Chitose, Tokyo 156, Japan. Prices include delivery by air. Postmaster: send address changes in USA, Canada and Mexico to: Bulletin of the Natural History Museum. Zoology Series, Cambridge University
Press, 1 10 Midland Avenue, Port Chester, New York, NY 105783-4930. Claims for missing issues should be made immediately on receipt of the subsequent issues.
Orders and enquiries for issues prior to 2002 should be sent to: Intercept Ltd., P.O. Box 716, Andover, Hampshire SP10 1YG, Telephone: (01264) 334748, Fax: (01264) 334058, Email: [email protected], Internet: http://www.intercept.co.uk
ADVERTISING Apply to the Publisher. Address enquiries to the Advertising Promoter.
COPYRIGHT AND PERMISSIONS This journal is registered with the Copyright Clearance Center, 222 Rosewood Drive, Danvers, MA 01923, USA. Organizations in the USA who are also registered with CCC may therefore photocopy material (beyond the limits permitted by Section 107 and 108 of US Copyright law) subject to payment to CCC of the per-copy fee of $16.00. This consent does not extend to multiple copying for promotional or commercial purposes. Code 0968-0470/2002 $16.00. ISITear Sheet Service, 3501 Market Street, Philadelphia, PA 19104, USA, is authorised to supply single photocopies of separate articles for private use only. Organizations authorised by the UK Copyright Licensing Agency may also copy material subject to the usual conditions. For all other use, permission should be sought from Cambridge University Press.
No part of this publication may otherwise be reproduced, stored or distributed by any means without permission in writing from Cambridge University Press, acting for the copyright holder.
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 1 . Xx£^i:2 &c3i
Bull. not. Hist. Mu.s. Loud. (Zool.) 68( 1 ): 1-1 Issued 27 June 2002
Another variation on the gymnure theme: description of a new species of Hylomys (Lipotyphla, Erinaceidae, Galericinae).
nrMFSAt LIBRARY 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, GLJ 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 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
The Family Erinaceidae is divided into two subfamilies: the wide- limestone karst in the Lao PDR. While sharing many characters with spread Erinaceinae (hedgehogs) occuring in Africa, Europe and other species of Hylomys this new species also differs markedly
Asia, and the Galericinae (moonrats and gymnures), which is con- from its congeners and, furthermore, shares some features with fined to southeast Asia, Indonesia and the Philippines. There has geographically remote species of Podogymnura. The new taxon has been considerable disagreement over the correct name to apply to been compared in particular with specimens of H. suillus, although the subfamily of moonrats and gymnures, summarised by Frost et al. there is no indication that the two species occur sympatrically, and (1991), who favoured the use of Hylomyinae. McKenna & Bell also with H. sinensis, H. panus and. in the absence of specimens, (1997) pointed out however, that the use of Galericini as a tribal with the original description of H. hainanensis and the figures of the
name by Butler (1948) had been accepted by many subsequent skull of this species in Frost et al. ( 1991 ). In addition comparisons writers, particularly palaeontologists, and was therefore the appro- were made with the other genera of Galericinae: Echinosorex priate name to use. In this paper we follow McKenna & Bell ( 1997) gymnura (Raffles, 1822) from Malaysia and Indonesia, and in using the name Galericinae. Most authors up to and including Podogymnura truei Mearns. 1905 from the Philippines. In order to Corbet (1988), considered that the Galericinae includes five genera: assess phylogenetic relationships, both the new taxon and H. parvus
Echinosorex Blainville, 1838, Hylomys Miiller, 1840, Neotetracus were analysed using the criteria employed by Frost et al. (1991). 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 <://.( 1991) con- MATERIAL AND METHODS cluded that there are only three valid genera within the Galericinae: Echinosorex, Podogymnura and Hylomys. They accepted Hylomys Comparative material was examined from the collections of the as a rather variable but nevertheless monophyletic genus, although Natural History Museum (BMNH), London (formerly the British they conceded that there was evidence to support the retention of Museum (Natural History)), the American Museum of Natural Neotetracus and Neohylomys as subgenera. History, New York (AMNH), the Museum National d'Histoire The genus Hylomys is widely distributed in southeast Asia and Naturelle, Paris (MNHN) and the Thailand Institute of Scientific Indonesia. Hylomys suillus Miiller, 1840 occurs in Malaysia, Indo- and Technological Research, Bangkok (TISTR), as listed in Table 1 nesia, Thailand. Vietnam. Cambodia, Lao Peoples Democratic All measurements are in millimetres and were taken using digital Republic (PDR), Myanmar and southern PDR China; in Lao PDR it calipers. Cranial and dental nomenclature follows Butler (1948), has been recorded from Phongsali, Xiangkhouang, Vientiane and Novacek (1986), Frost et al. (1991) and Gould (1995). Dental Dong Hua Sao National Biodiversity Conservation Area (NBCA) notations are indicated in the text in the following manner, with (Robinson, 1999). A number of different subspecies have been premaxillary and maxillary teeth denoted by uppercase letters and attributed to H. suillus and the biochemical and metrical variation mandibular teeth by lowercase: incisor (I/i), canine (C/c), premolar within this species was examined by Ruedi et al. (1994). They (P/p), molar (M/m), thus P3 refers to the third upper premolar, i2 to recognised that much of the high level of variation could be attrib- the second lower incisor. uted to the geographical and altitudinal isolation of the named forms but demonstrated that one of these taxa, H. pan'iis Robinson & Kloss, 1916, merited specific status. Hylomys sinensis (Trouessart, 1909) occurs from southern China to Myanmar and northern Viet- PHYLOGENETIC ANALYSIS 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 Cranial, dental, skeletal and external characters were scored for the
Osgood (1932). Hylomys hainanensis (Shaw & Wong, 1959) is new species and H. pan>us according to the character transformation
© The Natural History Museum, 2002 )
P.D. JENKINS AND M.F. ROBINSON series employed by Frost et al. (1991: 3-15) and added to the fifth digits of forefeet lengthened, claws long and moderately stout: character matrix shown in Frost et al. (1991: appendix 2) see Table sole and tarsal region of hindfeet naked, cheiridia large. Pre-anal
2. Branch and bound analyses were performed using Paup 4.0ba gland with single opening immediately posterior to cloaca. Two
(Swofford, 1999) set at maximum parsimony, with a maximum trees pairs of inguinal mammae present. setting of 1000 and all characters treated as unordered and of equal Skull elongate, moderately slender and somewhat flattened in weight. Bootstrap analyses (Felsenstein, 1985) were made to pro- appearance (see Figs. 1-2); dorsal profile more or less straight, vide an assessments of confidence limits of nodes, with 1000 showing a gradual increase in height from anterior of rostrum to replicates of 100 random addition sequence replicates. Bremer braincase. Rostrum long, slender and moderately shallow, nasals support indices were calculated by increasing the upper bound of the long extending posteriorly to, or slightly beyond, level of antorbital shortest tree by one step, repeating the branch and bound analysis rim; posterodorsal region of premaxilla widely separated from and producing a strict consensus tree; the process was repeated, anterodorsal region of frontal by maxilla; interorbital region moder- progressively increasing the length of the suboptimal cladograms by ately narrow; supraorbital processes of frontals scarcely evident; a single step until all clades of interest no longer occurred on the frontals anteriorly depressed in midline; supraorbital region of consensus tree; the level at which each node collapsed was recorded frontals broad, so that the maxillaries are widely separated from the (Kitching etal., 1998). Both accelerated (ACCTRAN) and delayed parietals; parietals extend anteriorly in supraorbital and orbital (DELTRAN) optimizations were used to map character evolution. region but do not form an anterior process; supraorbital foramen
The trees obtained were compared with those in Frost et al. ( 1991 present in dorso-orbital region of frontals; orbital region of maxilla and the results of the analysis are given below. 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 RESULTS 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 Hylomys megalotis, sp. nov. optic, suboptic and sphenorbital foramina in lateral view; alisphe- HOLOTYPE. BMNH 1999.44 (field number 5/99) male, body in noid dorsoventrally compressed, fusiform anteroventral process of alcohol, skull extracted. Collected 15 January 1999 by M. F. alisphenoid present, well marked alisphenoid canal present; brain- Robinson. case low and scarcely domed, lambdoid crest moderately well-developed laterally, low medially; mastoid large, slightly in- Type LOCALITY. Environs of Ban Muang and Ban Doy, c 1 8 km flated; paraoccipital process small; infraorbital foramen dorsal to North ofThakhek, Thakhek district, Khammouan Limestone National P4; antorbital or prelacrimal flange present only as a low ridge; Biodiversity Conservation Area, Khammouan Province, Lao Peoples shallow antorbital fossa on anterior surface of zygoma; nasolabialis Democratic Republic, 1 7°33' 1 5"N 1 04°49'30"E. Habitat: steep slopes fossa shallow; maxillary component of zygoma narrow with long, around the base of massive limestone karst, covered in rock and large slender posteroventral process ventral to well marked long jugal, boulders, with an underlying soil base and heavily degraded mixed slender anterodorsal process of squamosal portion of zygoma over- deciduous forest, scrub and bamboo. Low lying areas away from the lying jugal; palate with paired maxillary foramina level with P2 and karst had been cleared for cultivation of paddy rice. anterior of P3, small paired anterior palatal foramina, lying anterior PARATYPES. BMNH 1999.45 (field number 14/99) collected 16 to the suture between maxilla and palatine; palatal spine absent; January 1999; 1999.46 (field number 15/99) and 1999.48 (field basioccipital narrow with ridge in midline, tympanic wing of basioc- number 17/99) collected 17 January 99, females, bodies in alcohol, cipital slightly inflated. Mandible with deep, moderately broad skulls extracted; 1999.47 (field number 16/99) collected 17 January coronoid process; mental foramen below p3.
1999, male, skin and internal organs in alcohol, skull and skeleton. Dental formula: 3/3 1/1 4/4 3/3 = 22. Dentition robust (see Figs. 1-
All specimens were collected by M.F. Robinson from the same 2). First upper incisor robust, distostyle present; 12 and 13 locality as the holotype. sub-triangular, anteroflexed, distostyle present, 13 approximately half size of 12; C with anterior basal cusp and distostyle and two Diagnosis roots; PI and P2 subequal in height, P2 longer than PL both with Ears large, rhinarium elongated; first and fifth digits of forefeet long, anterior basal cusp and distostyle, PI with two fused roots, P2 with claws long, cheiridia large and rounded; cheiridia on hindfeet large, two roots; P3 large, subequal in height to C, lingual cusp (protocone soles naked; pre-anal gland with single opening. Skull with posterior according to Gould, 1995) well developed, three roots present: P4, region of nasals extending to level of antorbital rim; maxilla and Ml and M2 quadrate in shape, parastyle well developed, meta- parietal widely separated by frontal in supraorbital region; long conule present on Ml and M2; M3 subtriangular in shape, with well grooves for palatine artery present in palate; anterior palatine fo- developed parastyle and hypocone and metacone distinct in unworn ramina anterior to maxillary palatine suture; antorbital fossa shallow; dentition. First lower incisor larger than i2, both semi-procumbent, nasolabialis fossa shallow; posteroventral maxillary process of i2 larger than i3, which is anteroflexed with hypoconulid present; c zygoma distinct; antero- ventral process of alisphenoid present. Den- anteroflexed, greater in height than i3 and pi; pi and p2 subequal in tition robust. Third upper premolar (P3) large with well developed height and both with a single root; p3 larger with two roots; p4 with lingual cusp and three roots. Neural spine of axis low. well developed paraconid and talonid; ml-m3 with well developed Description paraconids, m3 less than half size of ml.
Medium sized Hylomys with a long tail, approximately 75% of head and body length. Pelage grey, long, soft and very fine, lacking Etymology flattened spinous hairs; individual hairs grey for most of their length, The name of the new species is derived from the Greek ueyocc; then buff with buff or black tips. Dorsal region of rhinarium narrow, (megas), large; (hrzoq (otos), ear; the ears are large in comparison elongate posteriorly; ears prominent, very large, rounded. First and with those of other species of Hylomys. A NEW SPECIES OF HYLOMYS
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. 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 portion of the nasals in the new species extend to the level of the
The new species is readily distinguished in external appearance antorbital rim. Hylomys megalotis has a shallow antorbital fossa from all other species of Hylomys. It is similar in body size but with unlike the moderately deep fossa of H. sinensis and H. parvus, and a considerably longer tail and larger ears (see Table 3). Tail 65-74 the deep fossa in H. suillus. The zygoma of//, megalotis differs from
% of head and body length in H. megalotis, 5 1-63 % in H. sinensis, all other species of Hylomys: the maxilla is considerably narrower,
27-3 1 % in H. hainanensis, and very short (< 25 %) in H. par\>us the nasolabialis fossa shallower and the jugal more extensive. As in and H. suillus. The lack of flattened spinous hairs in the pelage Podogymnura aureospinula and Echinosorex, a distinct posteroventral distinguishes H. megalotis from H. suillus and H. sinensis. The process is present on the maxillary region of the zygoma of H. rhinarium is more extensive posteriorly than in H. sinensis but more megalotis, indistinct in H. parvus but absent in H. suillus, H. sinensis elongate and narrower than in H. suillus. The claws and first and and H. hainanensis. The supraorbital process of the frontal of //. fifth digits of the forefeet of//, sinensis, H. suillus and H. pan>us are megalotis is poorly defined and blunt, the anterior process of the not lengthened as in H. megalotis and the cheiridia are smaller than parietal absent and the parietal is widely separated from the maxilla in H. megalotis. The sole and tarsal region of the hindfoot are naked by the frontal in the supraorbital region, as in Podogymnura and in H. megalotis, differing from the haired soles found in H. sinensis Echinosorex; in H. par\>us the supraorbital process of the frontal is and H. suillus. The paired pre-anal glands are midway between the poorly defined and blunt but the anterior process of the parietal is cloaca and the anus in H. sinensis and close to the anus in H. suillus, short but distinct, narrowly separated from the maxilla by the frontal; so differing from that of H. megalotis, in which the single opening in H. suillus, H. hainanensis and H. sinensis the anterior process of is positioned immediately posterior to the cloaca. the parietal is distinct, scarcely separated from the maxilla and
The skull of H. megalotis is more elongate in appearance than contributing to the well marked supraorbital process of the frontal. any of the other species of Hylomys; it is longer, with a longer, The optic and suboptic foramina are well separated in H. megalotis narrower rostrum, longer upper toothrow and the braincase is but lie close together in H. suillus, H. sinensis and H. parvus. An shallower relative to its breadth (see (see Fig. 1 and Table 3). The anteroventral process of the alisphenoid is present in H. megalotis, posterodorsal region of the premaxilla is widely separated from the unlike all other Hylomys and Podogymnura. As in Podogymnura and anterodorsal region of the frontal by the maxilla in H. megalotis, Echinosorex, the palatine foramina are small and positioned anterior narrowly separated in H. hainanensis but in contact or nearly in to the maxillary/palatine suture in H. megalotis and long grooves for contact in H. sinensis, H. suillus and H. parvus (see Table 4). As in the palatine artery are present in the palate, whereas the elongated
H. sinensis, but unlike other species of Hylomys, the posteriormost palatine foramina in other species of Hylomys lie at the maxillary/ A NEW SPECIES OF HYLOMYS
CORF
OPTF
FOV SPALF
SOPF
Fig. 3 Lateral view of left orbital region of Hylomys megalotis BMNH 1999.44. Scale = 1 mm. Abbreviations: AL - alisphenoid. AL1C - 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- tion seen in Fig. 4a, while the remaining trees all showed the tinct. The anterior opening of infraorbital canal is dorsal to P3/P4 in alternative arrangement for this subfamily (Fig. 4b). That part of the most species of Hylomys but dorsal to P4/M 1 in H. megalotis, H. tree obtained by Frost el al. (1991: Fig. 9) for the Galericinae is parvus and Echinosorex, and yet more posteriorly positioned in illustrated as part of Fig. 4a. The strict consensus tree (see Fig. 5) Podogymnura. revealed strong bootstrap support (91%) for the Galericinae and for
The dentition of H. megalotis is considerably more robust than a clade of//, suillus. H. sinensis and H. hainanensis, and this tritomy that of any of the other species of Hylomys. The dental formula is the also had a high Bremer support index. There was moderate bootstrap
same as in H. suillits and H. parvus and these species are distin- support (83% ) for a clade of H. suillus, H. sinensis, H. hainanensis guished from H. hainanensis, which lacks pi and H. sinensis, which and H. parvus, and a clade comprising all species in the genus lacks PI and pi. In H. megalotis and H. parvus, P2 has two roots, Hylomys occurred in 77% of the replicates. The bootstrap support unlike the other species which have either one or two well fused value for a clade of Echinosorex and Podogymnura was low at only roots. Unlike all other species which lack a lingual cusp, P3 of H. 64%. Within the genus Hylomys, H. megalotis was basal to all other megalotis has a well developed lingual cusp as in Echinosorex, and species. Clades with bootstrap support values less than 50%, respec-
this tooth is large with three roots as in Podogymnura and tively ofH. sinensis and H. hainanensis (42%), and H. suillus and H.
Echinosorex; p3 is larger than p2 with two roots as in H. pan'us, hainanensis (43%) were considered to be unresolved. Podogymnura and Echinosorex, while p3 is slightly smaller than p2 The shared derived character transformations (synapomorphies) with one root in H. suillus, H. sinensis and H. hainanensis. which were revealed by the analysis are recorded below, using the
format of character number quoted from Frost et al. ( 1 99 1 ) followed by character transformation state, where (0) equals the ancestral and
( ) the derived character state. RESULTS OF THE PHYLOGENETIC 1 ANALYSIS Synapomorphies of Galericinae: [8.1] Antorbital or pre-lacrimal flange: (0) not developed, lacrimal
Forty equally most parsimonious trees were retained in the branch canal visible in lateral view; ( 1 ) developed so that the lacrimal canal
and bound analysis, 141 steps in length, with a Consistency Index of is obscured in lateral view. CI 1 .000. 0.72, Retention Index of 0.93 and Rescaled Consistency Index of [10.2] Jugal size; (0) large, reaches lacrimal; (1) small, does not
0.66. In all most parsimonious trees the two subfamilies, Galericinae reach lacrimal; (2) vestigial, confined to lateral rim of zygoma; (3)
and Erinaceinae, readily segregated and the Galericinae further absent. In ACCTRAN the transformation was from 1 —> 2, in separated into two distinct groups: a clade comprising Echinosorex DELTRAN the change was from 3 -> 2. CI 1.000. and Podogymnura, the other clade confined to Hylomys. Most of the [62.1] P4 lingual roots: (0) one; (1) two unfused; (2) two fused. CI variation found among all trees occurred within the Erinaceinae, 1.000. This state occurs in all Galericinae but was shown only in since for the Galericinae twenty of the trees showed the configura- DELTRAN. P.D. JENKINS AND M.F. ROBINSON
E. gymnura
P. aureospinula
P. truei
H. sinensis
H. suillus
H. hainanensis
H. parvus
H. megalotis
Erinaceinae
E. gymnura
P. aureospinula
P. truei
H. sinensis
H. hainanensis
H. suillus
H. parvus
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 et al. (1991: fig. 9),
restricted to show only the relationships within the Galericinae and is 1 28 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 [74.1] Tibia, lateral flange on antero-superior margin: (0) absent or
Gould, 1995): (0) absent or weak; (1) present, well developed on weakly present; (1) strongly developed. CI 1.000. buccal side. CI 1.000.
[69.1] Axis, posteroventral keel: (0) absent; (1) present. CI 1.000. Synapomorphy of Hylomys: [71.1] Scapula, metacromium process: (0) deltoid, amorphous pro- [19.1] Cranio-orbital foramen in the frontal: (0) closely associated or jection; (1) long, fusiform projection. CI 1.000. joined with the ethmoid foramen; (1) foramina widely separated.
[72.1] Sacral vertebrae, neural spines: (0) not fused into continuous The terminology for this character is confusing. Frost et al. (1991) longitudinal plate; ( 1 ) fused into continuous longitudinal plate. CI used the name ophthalmic foramen (which they attributed to Butler 1.000. (1948) although this name could not be found in this paper), but [73.1] Ischium, posterodorsal process (see Gould, 1995 for correc- Gould (1995: character 19) pointed out that this foramen had been tion of error by Frost et al. 1991): (0) not greatly elongated; (1) misidentified by Butler and is the anterior opening for the superior greatly elongated. CI 1 .000. ramus of the stapedial artery. Gould also referred to this foramen as A NEW SPECIES OF HYLOMYS
E. gymnura 3 (64 P. aureospinula 4 (73) P. true!
H. sinensis 14 (97) 7 .(97' H. hainanensis 6 (83) H. suillus
6 (77) 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. related to the shortening of the skull in erinaceids. As the skull CI 1.000. shortens, the alisphenoid overlaps the orbitosphenoid, creating a
strong alisphenoid wing [character 17 of Frost et al. (1991) and Synapomorphy of Hylomys sinensis, H. hainanensis, H. Gould ( 1995)1, the degree of overlap seems to be directly related to SUILLUS AND H. PARVUS: the visibility of the suboptic foramen from lateral view and, as [4.1] Size of palatal foramina: (0) small; (1) anterior foramina pointed out by Butler (1948) the orbitosphenoid is reduced in size. elongated posteriorly; (2) anterior foramina elongated to include While the alisphenoid is more extensive in H. megalotis than in other middle palatine foramina. CI 1.000. galericines, and the suboptic and sphenorbital foramina are partially
Synapomorphies of Hylomys sinensis, H. hainanensis, H. concealed in lateral view, the orbitotemporal region is not obviously suillus: shortened. The anterior process in H. megalotis is fully ventral in
[13.1] Supraorbital process of frontal on parietal/frontal suture: (0) location and is actually or nearly in contact with a short posteroventral
absent or poorly defined; ( 1 ) sharp, well defined. CI 1 .000. process of the maxilla, thus contributing to the ventral floor of the
[16.1] Anterior process of parietal: (0) absent or very weak; (1) orbit, however the orbitosphenoid is not reduced in size. It is
extends anteriorly along the supraorbital rim to form the base of the possible that this character state in H. megalotis is not homologous
supraorbital process. CI 1.000. with that of the Erinaceinae and that it actually represents a separate
[57. 1 ] p3: (0) two roots present, larger in size than p2; ( 1 ) one root character transformation, alternatively it is scored incorrectly and present, nearly equal in size to p2; (2) absent. CI 1.000. the plesiomorphous condition should be the presence of the anterior process.
The analysis found no autapomorphic characters to define H. [22. 1 ] Palatal shelf and spine: (0) well developed spine on posterior
megalotis but the following apomorphies for this species are recorded palatal shelf; ( 1 ) spine absent or vestigial. CI 0.200. Shown only in as follows: DELTRAN, homoplasious with Podogymnura, H. parvus, and
[1.1] Posteriormost extension of nasals: (0) anterior to the level of Atelerix.
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. DISCUSSION
[5.1] Location of the anterior palatine foramina: (0) at the maxilla/
palatine suture; ( 1 ) anterior to the maxilla/palatine suture. CI 0.500. The addition of two taxa to the analysis performed by Frost et al.
Shown only in DELTRAN, homoplasious with Echinosorex and ( 1 99 1 ) provided broadly similar results in that the Galericinae divided Podogymnura. into two main groups: one comprising Echinosorex andPodogymnura,
[17.1] Anterior process of alisphenoid: (0) absent; (1) present. CI the second including all five species of Hylomys. The results of the 0.500. Homoplasious with Erinaceinae. This character, defined as a current phylogenetic analysis lend support to the taxonomy proposed narrow, fusiform anterior process of the orbital wing of the alisphe- by Frost etal. (1991) that the three species of Hylomys considered in
noid is, according to Frost et al. (1991), related to the location of the their analysis (H. suillus, H. sinensis and H. hainanensis) are correctly sphenopalatine foramen and involved with shortening of the attributed to a single genus rather than the three separate genera orbitotemporal region. Gould (1995) commented that the relative (respectively Hylomys, Neotetracus and Neohylomys) maintained by
position of the suboptic foramen (her character 21 scored thus: (0) Corbet (1988). The additional species of Hylomys however, reduced
anterior to the sphenorbital fissure; ( 1 ) present in the medial wall of the degree of support for the genus and, on this particular morphologi-
the sphenorbital fissure; (2) present in the medial wall of the cal data set, a considerable degree of homoplasy is evident within the sphenorbital fissure but hidden within the fissure) seems to be Hylomys clade. There was only one unique synapomorphy for the 8 P.D. JENKINS AND M.F. ROBINSON
Hylomys clade (character 19.1: the wide separation between the Field work was funded by the Global Environment Facility through the cranio-orbital and ethmoid foramina), two of the other apomorphies World Bank to the Government of Lao PDR. via a contract with Burapha Development Consultants, sub-contracted to WWF-Thailand. (34. 1 : the inflation of the mastoid region between the exoccipital and the squamosal, and 41.1: the expanded exoccipital) showing We thank Bob Randal (AMNH), Jacques Cuisin (MNHN) and Nivesh Nadee (TISTR) for making specimens in their collections available to us. We homoplasy with P. truei, while the third character state (50. 1 : upper are very grateful to Andrew Cabrinovic (NHM) for specimen preparation, to canine slightly larger than the adjacent post-canine teeth) is not
Harry Taylor, Photographic Unit ( NHM) for the excellent photographs and to shown by H. sinensis (50.2: upper canine approximately equal in Karen Fisher (volunteer, NHM) for her kind assistance with documentation. size to the adjacent post-canine teeth). There are no autapomorphies We are grateful to Dr Francois Catzeflis. Institut des Sciences de l'Evolution. defining H. megalotis, which shows more plesiomorphy than the Universite Montpellier for his constructive review of the manuscript. We other species of Hylomys; many of its features are homoplasious particularly thank Dr Darrell Siebert, NHM for his generous guidance with with Echinosorex, Podogymnura and Erinaceinae. the phylogenetic analysis and constructive criticism of the manuscript. 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. REFERENCES suillus in a few areas, Corbet (1988) pointed out that in regions Blainville, H. M. D. de 1838. Recherches sur Panciennete des Mammiferes insecti- where H. sinensis and H. suillus occur, H. sinensis is found at greater vores a la surface de la terre: precedees de l'histoire de la science a ce sujet. des altitudes than H. suillus. Similarly Ruedi et al. (1994) showed that principes de leur classification et de leur distribution geographique actuelle. is currently restricted to forests at the although H. parvus moss peak Compte Rendus Hebdomadaires de Sceances de VAcademie des Sciences 6: 738- of Gunung Kerinci, Sumatra at greater altitudes than H. suillus, the 744. latter occurs elsewhere at greater and lesser altitudes. Both Corbet 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 (1988) and Ruedi et al. (1994) invoked ecological factors such as Zoological Society. London 118 (2): 446-500. competitive exclusion to explain the altitudinal segregation of these Corbet, G. B. 1988. The family Erinaceidae: a synthesis of its taxonomy, phylogeny. three species, but did not provide data to support this supposition. ecology and zoogeography. Mammal Review 18 (3): 117-172. There are few distribution records of//, suillus and H. megalotis in Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: 783-791. Lao PDR, which potentially may be sympatric or possibly parapatric
Frost, D. R., Wozencraft, W. C. & Hoffmann, R. S. 1 99 1 . Phylogenetic relationships if H. megalotis should prove to be ecologically restricted to the of hedgehogs and gymnures (Mammalia: Insectivora: Erinaceidae). Smithsonian specific limestone habitat in which it was first collected. Contributions to Zoology (518): 1-69. 19 plates.
Little is known about the biology of Hylomys. Hylomys sinensis is Gould, G. C. 1995. Hedgehog phylogeny (Mammalia. Erinaceidae) - the reciprocal
illumination of the quick and the dead. American Museum Novitates (3131): 1 —45. believed to be entirely terrestrial and H. suillus mainly so, although Kitching, I. J., Forey, P. L„ Humphries, C. J. & Williams, D. M. 1998. Cladistics. this species has also been observed climbing in low bushes (Lekagul Second edition. The theory and practice of parsimony analysis. Oxford University & McNeely, 1977). Hylomys suillus occurs in hilly or montane Press. Oxford. New York, Tokyo. humid forests with dense undergrowth, H. sinensis in cool damp Lekagul, B. & McNeely, J. A. 1977. Mammals of Thailand. Kurusapha Ladprao Press. Bangkok. forests in the cover of runways and under logs and rocks. In their McDowell, S. B. 1958. The greater Antillean insectivores. Bulletin of the American original description Shaw & Wong (1959) reported that //. Museum of Natural Histoiy 115 (3): 117-214. hainanensis spends most of its time in underground burrows and that McKenna, M. C. & Bell, S. K. 1997. Classification of mammals above the species the cylindrical body, short tail and claws are adaptations to a level. Columbia University Press, New York. Mearns, E. A. 1905. Descriptions of new genera and species of mammals from the fossorial life. Hylomys parvus is apparently restricted to high alti- Philippine Islands. Proceedings of the United States National Museum 28: 425-460. tude moss forest. There is no information about the behavioural Miiller, S. 1840. Over de Zoogdieren van den Indischen Archipel (No. 2) pp. 9-57. In ecology of H. megalotis but the limestone karst where it has been C. J. Temminck (ed.) Verhandelingen over de Natuurlijke Geschiedenis der found to date is an unusual habitat and some of the morphological Nederlandsche overzeesche bezittingen. doorde Leden der Natuurkundige commissie features of this species, such as the moderately broad forefeet with in Indie en andere Schrijvers. A. Arnz & Co., Leiden. Novacek, M. J. 1986. The skull of lepticid insectivorans and the higher-level classi- long, fairly stout claws, the long naked hindfeet with large cheiridia, fication of eutherian mammals. Bulletin of the American Museum of Natural History the moderately long tail and the comparatively flattened braincase 183(1): 1-111. may be adaptations to life in this habitat. Osgood, W. H. 1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic expeditions. Publications of the Field Musuem of Natural Histoiy. Zoology (18): 193-339.
Raffles, T. S. Descriptive a zoological collection, made on account Acknowledgements. We would like to thank the staff of WWF-Thai- 1822. catalogue of of the Honourable East India Company, in the island of Sumatra and its vicinity, land, the Lao PDR Department of Forestry (DoF) and the Forest Management under the direction of Sir Thomas Stamford Raffles, Lieutenant-Governor of Fort and Conservation Project (FOMACOP) for permission to work in Lao PDR Marlborough; with additional notices illustrative of the Natural History of these and for organising the project; in particular ( National Bouahong Phanthanousy countries. Transactions of the Linnean Society, London 13 (1): 239-274. Project Director, FOMACOP), Bouaphanh Phantavong (Deputy National Robinson, H. C. & KIoss, C. B. 1916. Preliminary diagnoses of some new species and Project Director, FOMACOP), Robert Mather and Robert Steinmetz (WWF- subspecies of mammals and birds obtained in Korinchi, West Sumatra, Feb.-June 1914. Journal the Straits Branch the Royal Asiatic Society (73): 269-278. Thailand) and Bruce Jefferies (FOMACOP), who additionally oversaw the of of Robinson, M. F. 1999. Order Insectivora, pp. 221-223 In J. W. Duckworth, R. E. Salter export of specimens. We are very grateful to Maurice Webber for his vital & Khounboline, K. (compilers) Wildlife in Lao PDR 1999 status report. IUCN, contribution to all aspects of the fieldwork. Vientiane. Assistance in the field was grateful ly received from Thongphanh Ratanalangsy Ruedi, M., Chapusiat, M. & Iskander, D. 1994. Taxonomic status of Hylomys parvus (KhammouanLimestone and DongPhu Vieng NBCA Co-ordinator), Sinnasone and Hylomys suillus (Insectivora: Erinaceidae): biochemical and morphological Seangchanthanarong (DoF) and Nousine Latvylay (driver for FOMACOP in analyses. Journal of Mammalogy 75 (4): 965-978. Swofford, D. L. 1999. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Savannakhet). The headmen and villagers from Pontong,Muang,Mouangkhai, Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. Louang, Vieng, Kengkhot, Thamtem and Tonglom provided excellent field Shaw, T. H. & Wong Song 1959. [A new Insectivore from Hainan]. Acta Zoologica support and companionship, and without this project not have been them would Sinica 11: 422^125. [In Chinese; English summary]. possible. We would like to thank Angela L. Smith for her help in measuring Trouessart, E.-L. 1909. Neotetracus sinensis, a new insectivore of the family specimens at TISTR and for her comments on the manuscript. Erinaceidae. Annals and Magazine of Natural Histoiy (8) 4: 389-391. A NEW SPECIES OF HYLOMYS
Table 1 Comparative material
Hylomys BMNH 1912.10.22.8 Perlis, Malay Peninsula [Malaysia]
BMNH I 932.4.19.3 Chapa. Tonkin, [Vietnam] BMNH 1955.1423 Jor, Batang Pasang, Perak, Federal Malay States
BMNH 1 933.4. 1.1 17-1 34 Chapa, Tonkin, [Vietnam] [Malaysia]
BMNH 1 933.4.1.536-541 Chapa, Tonkin, [Vietnam] BMNH 1955.1424 Kedah Peak, Federal Malay States [Malaysia]
BMNH 9 1 1 .2. 1 . 1 5-23 Omi-san, Omi-Hsien, S. Szechwan [Sichuan, PDR BMNH 1962.71 la Kedah Peak. Kedah, Malaya [Malaysia] China] I BMNH 1892.9.6.4 Mt. Kina Balu [Malaysia]
BMNH 1 982.205 Omi-san, Omi-Hsien, S. Szechwan [Sichuan, PDR BMNH 1895.10.4.3^ Mt. Kina Balu [Malaysia] China] I BMNH 1955.661 Mount Kinabalu. British North Borneo [Malaysia]
1 1 - 1 1 80- 1 1 MNHN 1 9 84 Ta-Tsien-Lou, Setchouen [Sichuan, PDR China] BMNH 1971.2614-2615 Mt. Kinabalu, N Borneo [Malaysia] BMNH 909. 12.13.1 Ta-Tsien-Lou, Szechuan [Sichuan, PDR China] MNHN 1889-37 Mont Kina Balu. Borneo [Malaysia] BMNH 911.8.6.1 Yengyuek. Yunnan, [PDR China] MNHN 1893-132-133 Mont Kina Balu, Borneo [Malaysia] BMNH 912.7.15.1 Ching-tsai-Yang, Yunnan, [PDR China] BMNH 1971.2616 Dusan Dankulum, Kinabalu, N Borneo [Malaysia] BMNH 914.10.23.3 Near Yang-fsi, W. Yunnan, [PDR China] BMNH 1971.2617-2618 Tinampoh, Bundu Tuhan rest house, N Borneo [Malaysia] Hylomys suillus BMNH 1919.1 1.5.7 Korinchi, Sumatra [Sumatera, Indonesia] BMNH 1909.7.20.2-3 Sima. Burma [Myanmar] AMNH 102532 Seletan. Mocamh Doewa, Sumatra [Sumatera, Indonesia] AMNH 441 1 2 Nam-Ting, Yunnan, China [PDR China] AMNH 102533 Seletan. Mocamh Doewa. Sumatra [Sumatera, Indonesia] BMNH 1925. 1 . 1 . 1 7 Bao-Ha, Tonkin, [Vietnam] AMNH 102534 Seletan, Mocamh Doewa. Sumatra [Sumatera. Indonesia] BMNH 1926.10.4.42 Xieng-Khouang, Laos [Lao PDR] AMNH 102820 Lampung. Kalianda. Sumatra [Sumatera. Indonesia] AMNH 87313 Bologens Plateau, Laos [Lao PDR] BMNH 1954.45 Tjibodas. West Java [Jawa. Indonesia] BMNH 1926.10.4.36-41 Dak-to, Annam BMNH 1954.46-48 Sodong Jerok. Idjen Massif. East Java [Jawa. MNHN 1929-320-325 Dak-to, Annam Indonesia] BMNH 1955.1422 Tasan,Chumpawn, Peninsular Siam [Thailand] BMNH 1961.1743 Tjemorosewu. Mt. Lawu. Java [Jawa. Indonesia] TISTR 54-61 1 Huey Mae Sanam, Chiengmai, Thailand AMNH 106111 Java [Jawa. Indonesia] TISTR 54-613 Trang, Muang, Chong. Thailand TISTR 54-614-615 Khao Yai National Park, Korat, Thailand Hylomys parvus TISTR 54-616 Phu Nam Tok, Saraburi, Thailand BMNH 1919.1 1.5.8-12 Korinchi, Sumatra [Sumatera, Indonesia] TISTR 54-617 Pok Nam Tok, 21 km from Saraburi, Saraburi, Thailand TISTR 54-618 Mae Sai, B. Santon Poi, Chieng Rai, Thailand Podogymnura truei TISTR 54-1498-1500 Pak Thong Chai, Sakaerat, Nakhon Ratchasima. BMNH 1953.659-660 Baclayan. E slopes of Mount Apo, Mindanao, Thailand Philippine Islands
TISTR 54- 1 8 1 1 Pak Thong Chai, 1 7 km S of Pak Thong Chai, Nakhon Ratchasima. Thailand Echinosorex gymmiru TISTR 54-1809-1810 Khon San, Pa Phu Khieo, Chaiyaphum, Thailand BMNH 1914.12.8.101-104 Bankachon, Tenasserim [Myanmar] TISTR 54-1812 Phu Kradung National Park, Loei, Thailand BMNH 1955.1452 Changkat Mentri. Perak, Federal Malay States BMNH 1960.8.4.7 Ulu Langat Forest Reserve, Kajang, Selangor, [Malaysia] [Malaysia] BMNH 1955.1453 Damansara Road. Kuala Lumpur, Selangor, Federal BMNH 1961.1158 Ulu Langat Forest Reserve. Kajang. Selangor, Malay States [Malaysia] Malaysia BMNH 1961.1 156 Rontau Panjang. Klang. Selangor, Malaya [Malaysia] BMNH 1955.1420 Semangko Pass, Pahang, Federal Malay States BMNH 1961.1 157 Sungei Buloh, Selangor. Malaya [Malaysia]
[Malaysia] BMNH 1 96 1.1157 Sungei Buloh. Selangor. Malaya [Malaysia] BMNH 1961.1 159 Padang, Jeriau, Fraser's Hill, Pahang, Malaysia BMNH 1951.179-180 Mount Dulit, Sarawak. Borneo [Malaysia] BMNH 1962.710-711 Jandai Baik, Pahang, Malaysia BMNH 1951.181 Tinjar River, Baram District. Sarawak, Borneo BMNH 1912.10.22.7 Pelarit, Perlis, Malay [Malaysia] [Malaysia]
BMNH 1955.1421 Pelarit, Perlis, Federal Malay States [Malaysia] BMNH 1 97 1 .26 1 3 1 2 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
111111111122222222223333333333 4444444444 5555555555666666666 67777777 77788 Character 12 345 67 89012 345 67 89012 3456789012 345678901234567 8901234567890123 456789012345 67 8901
Hypothetical ancestor 00 00 00?000?00000?0000000000000000 0000000000 0000000?000?00?0000000 0???000000000 Echinosorex gymnura 0000101 1221101000002001000000 00 01000 1000000100000001000100 00111001101111110000001 Podogymnura aureospimila 000010 12 2110110000201000000100? 10 00 10000001000100010111010011100110??????0000001 Podogymn ura truei 000 0100 11211010 000020100000000001100 100010010001000 10 1110100111001 10 ??1 11 10000001 Hylomys sinensis 1001010 10211100100120100000000001100 100010000100020011111111111001101111110000000 Hylomys suillus 00 01010 1021110 010012 0000000000001100 100010000000010000011111111001101111110000000 Hylomys hainanensis 01010 1021110010012 0000000000001100 1000100000000 10 00011111111100110??11110000000 Hylomys megalotis 100 010 1221100001012 0100000000001100 100010000000010000000000111001101011110000000 Hylomys parvus 000100 1121101000012 0100000000001100 100010000000010 00000011111100110????????????? Hemiechinus aethiopicus 1112 011 02131011011011001104001110011 111101101011120001112111121110210000001130200 Hemiechinus hypomelas 1112 011 02121011011011001103001110011 111101101011120001112010121110 21000 0001130200 Hemiechinus micropus 1112 011 02121011011011001104001110011 11110110101112000111211112111021000 0001130200 Hemiechinus auritus 1112 011 02111011011011001102001110000 11110110101112000111201012111021000 00 01130300 Hemiechinus collaris 1112 011 02111011011011001102 001110000 111101101011120001112010121110210000001130300 Mesechinus dauuricus 1102 011 02110011011011001101101011000 111101101011120001112010121110210000000120310 Erinaceus amurensis 1102 011 02111011011011001101011010000 1111011010112 21001112010121110210000000011110 Erinaceus concolor 1102 011 02111011011011001101011010000 111101101011221001112010121110210000000010110 Erinaceus europaeus 1102 011 02111011011011001101011010000 11110110101122100111201012111021000 00 00017110 A telerixfrontalis 1102 011 02111011011011101111011010000 111101101011120001102010121110210000000111110 Atelerix algirus 1102 011 02111011011011101111011010000 111101101011120001102011121110210000000110110 Atelerix albiventris 1102 011 02111011011011101111011010000 111101101011120001102111121110210000000210110 Atelerix sclateri 1102 011 02111011011011101111011010000 11110110101112000110 2111121110210000000110110 Tenrecoids 00000 00300010000000000000000000000 000000000000007001000000000000000000000000000 Soricoids 000 00 00 3 00 01000002 00000 00000000000 000000000000007001000100000 000000000000000000 Lepticidae (fossil) 00000 00000000000000000000000000000 000000000000000000000000000000000???0007?0???
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 H. pan'us H. sinensis H. suillus H. hainanensis 1 H. megalotis
Head and body length 107.67 ±4.50 115.75 ±4.83 132.7 ±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) Tail length 24.00 ± 0.82 68.5 ±3.70 23.19 ±3.69 39.67 ±2.13 86.33 ±3.29 23-25 (3) 63-73 (6) 19-30(8) 36-^3 (6) 82.8-91.3(4) Ratio of tail length to head and body length 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) Hindfoot length 23.17 ±0.24 25.88 ± 1.02 21.89 ±1.31 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) Ear length 18.38 ±0.45 17.71 ± 1.41 17.5 ± 1.26 22.43 ± 0.90 17. 19(2) 18-19(6) 15-20(7) 16-19(6) 20.9-23.2 (4) Greatest skull length 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) Condyloincisive length 30.98 ± 0.82 33.73 ±0.67 36.96 ±1.04
29.09 ( 1 30.05-32.36 (11) 32.88-35.01 (7) 35.66-38.56 (4) Upper toothrow length 15.21 ±0.42 16.12 ±0.37 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) Length from 11 to anterior of P4 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)
Ratio of 1 1-P4 length to upper toothrow length 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) Rostral breadth 5.11 ±0.17 5.92 ±0.18 6.02 ±0.21 5.29 ±0.13 4.98-5.35 (3) 5.59-6.20(11) 5.57-6.44(13) 5.06-5.38 (4) Rostral length 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) Ratio of rostral length to rostral breadth 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) Braincase breadth 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) Braincase height 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) Ratio of braincase height to braincase breadth 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)
' Measurements from Shaw & Wong (1959). : . j I
A NEW SPECIES OF HYLOMYS 11
-f >i c— a. = §- u ° ,so s s r"3 '— B C u 'j (N u J2 ~ .22 c <£PE q o .CJJ £> 1) o ^ < = Q W3 < 3 < £5 « .£ S !_ ~ ^ u CJ ao c 2 <= 3 — 3 1) T3 .2 -a / u u u O o -a <•" e3 — — "-» .o'< 2 =Q„. —g E c U DO < £1 C/3 < Q y? < < 3 u ~ a S3 u c_ 1 g J 8 1 j* CO _^ — —u3J 3J 11 — t^^^,^^ Tj a;' 5 k z q q u £ £ £ 5 .£? u Q K £ in: i £ 2 — u is|slll| o 12 a=' s H -5 £ ^ 3 Q- in < c/j v _) 1> *- Q. O Q. — ° lo ~ 5 ci 00 £ — B a. 5 = u u T3 u a, Z Sfl < c a 7 a. a, < iz a. o 5. 2 ~ 15 * c a. h .c Z < a < c-. £ £ 3> tn Cl — ~ w n i> > - - u. -O 2 T3 T3 -a JB ^5 ~ - - — 5j o. - a £ 22 u 5^5 = O a. £Q -E5 E < Q C/3 > rr; > 5 -= c ca «<§ -C E "a o- £ 'I 2 £ 'i — £ u g c — o C - c o o> p cd u r^ > WJM > »-— = -z: Bc o> s - fl a7^ ^<. E p P3 c E 1 U "-> 0) £ o L- C_ 15 zz la e 03 eS OS o £ B CL B s o 8 -S y .2 £ jj sssosa « „° s £ 5 >> I s 1 1 1 fi I till a. a QhP a o-a = — < n ^ 3 yy (-3>szt^- \j Bull. nur. Hist. Mus. Land. (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 NW1 4RY, UK. CONTENTS Introduction 13 Systematic Account 13 P. principe sp. nov 13 Ecological Notes 17 Acknowledgements 17 References 17 Synopsis. A new species of freshwater crab of the genus Potamonautes Macleay. 1 838 is described from Principe ( Democratic Republic of Sao Tome and Principe), an island in the Gulf of Guinea oft 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 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 The freshwater crabs reported on here were collected during a (including an adult male) becomes available. zoological expedition to the island of Principe, made recently by the Figures were prepared using a camera lucida, and the specimens Zoological Society of London. The island of Principe, together with were deposited in The Natural History Museum, London, U.K. the island of Sao Tome, constitutes a small independent country (BMNH). Abbreviations: cw, distance across the carapace at the (The Democratic Republic of SaoTome and Principe) in the Gulf of widest point; cl, carapace length measured along the median line, Guinea. Principe is the second in a chain of volcanic islands that from the anterior to the posterior margin; ch, carapace height, the make up the Atlantic Ocean Islands group, that lies off the coasts of maximum height of the cephalothorax); fw, front width measured Cameroon, Equatorial Guinea and Gabon. The other islands in this along the anterior margin; s, thoracic stemite; e, thoracic episternite; group are Bioko, Sao Tome and Annobon. Bioko is closest to the s4/s5, s4/s5, s5/s6. s6/s7, s7/s8, sternal sulci between adjacent mainland and has two species of potamonautid freshwater crabs, thoracic sternites; s4/e4, s5/e5, s6/e6, s7/e7, episternal sulci between Sudanonautes floweri (De Man, 1901) and S. granulatus (Balss, adjacent thoracic sternites and epistemites; P1-P5, pereiopods 1-5, 1929), both of which are also found in nearby Cameroon al-a6, abdominal segments 1-6, a7, telson of the abdomen. (Cumberlidge, 1993, 1995, 1999). SaoTome 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 SYSTEMATIC ACCOUNT 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 Family POTAMONAUTIDAE Bott, 1970 from streams and nearby land in cloud forest in the remote roadless Genus POTAMONAUTES MzcLeay, 1838 highland region in the southwest part of the island. The new species is compared to Potamonautes anchietae (De Brito-Capello, 1871) Potamonautes principe sp. nov. and to P. margaritarius, but differs from each in a number of (Fig. 1) important characters (Bott, 1953, 1955, 1964). Although the speci- mens from Principe are both subadult females, and ideally an adult Material examined male is needed to make a definitive identification, a description is Holotype. BMNH 2001.6907. 1 subadult female, cw 40.5, cl 27.5, ) The Natural History Museum. 2002 14 N. CUMBERLIDGE, P.F. CLARK AND J. BAILLIE ^rv 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 15 ch 1 4.8, fw 10.5 mm, summit of Pico do Principe (01° 34', 5 l''N, 07° alcohol preservation) was creamy white with a very light purplish 22', 57"E), 945 m, coll. J. E. M. Baillie, 26 Sept. 1999. tone, however one specimen was more white than the other. This Paratype. BMNH 2001.6908.1 subadult female, cw 33.2, cl 22.8, coloration was relatively uniform throughout the body including the ° 07° ch 1 1 .6, fw 1 0.0 mm, summit of Pico do Principe (0 1 34'. 5 1 "N, dorsal carapace, underside, and legs. The purplish tone was darker 22', 57"E), 945 m, coll. J. E. M. Baillie, 1 Sept. 1999. on the carapace just behind the eyes, which turned to an orange reddish color once the specimens were placed in alcohol. DIAGNOSIS. Postfrontal crest straight, smooth, spanning entire carapace, ends meeting anterolateral margins at epibranchial teeth. Distribution. This species is known only from the summit of the Exorbital tooth small, pointed; epibranchial tooth represented only Pico do Principe: (01° 34', 51'N, 07° 22', 57"E) at 945 meters, The by small granule; anterolateral margin posterior to epibranchial Democratic Republic of Sao Tome and Principe, Gulf of Guinea, tooth raised, completely smooth, continuous with posterolateral Central Africa. margin. Carapace surface completely smooth; highly arched, height ETYMOLOGY. The species is named for the island of Principe greater than front width (ch/fw 1.5). Pleural (vertical) suture on where it was collected. The species name principe is a noun in carapace sidewall Y-shaped, ends meeting exorbital and epibranchial apposition. teeth. Suborbital margin raised, completely smooth. Ischium of third maxilliped with deep vertical sulcus. Third sternal sulcus s3/s4 Taxonomic remarks. The new species is assigned to Pota- deep, v-shaped, meeting sterno-abdominal cavity. Thoracic epister- monautes because it possesses the following combination of nal sulci s4/e4, s5/e5, s6/e6 and s7/e7 distinct. Anterior inferior characters: the anterolateral margin lacks an intermediate tooth margin of merus of cheliped lined by row of small sharp teeth, with between the exorbital and epibranchial teeth; the mandibular palp is large pointed tooth near junction with carpus. two-segmented: and the third maxilliped exopod has a long flagellum. Potamonautes is a widespread genus of African freshwater crabs Description. Carapace ovoid, wide (cw/fw 3.88), highly arched found throughout Africa from Senegal to the Horn of Africa, and (ch/fw 1.45); surface completely smooth semi-circular, urogastric, from Egypt to South Africa. Bott's ( 1955) revision of the freshwater transverse branchial grooves faint. Front straight, relatively narrow, crabs of Africa recognised some 34 species in this genus. Since then, about one-quarter carapace width (fw/cw 0.26), anterior margin the number of species of Potamonautes has risen to more than 60 sharply deflexed. Postfrontal crest distinct, smooth, straight, span- (Bott, 1959, 1960, 1964, 1968, 1970; Stewart etal., 1995; Stewart, consisting epigastric, ning entire carapace, of fused postorbital 1997a,b; Daniels et al., 1998; Cumberlidge, 1999; Corace et al, crests; ends of postfrontal crest meeting anterolateral margins at 2001). Although Bott (1955) recognised 15 subgenera of epibranchial teeth. Anterolateral margin between exorbital. Potamonautes, the authors of the present study prefer to follow epibranchial teeth smooth, lacking intermediate tooth; anterolateral Cumberlidge (1999) and use Potamonautes [sensu lato \ for all margin posterior to epibranchial tooth raised, completely smooth, species, pending a revision of the entire genus (Cumberlidge, un- continuous with posterolateral margin. Exorbital tooth small, pointed; published). epibranchial tooth represented only small granule. Suborbital by It is not normally good practice to describe a new species from a margin raised, completely smooth. Suborbital, subhepatic, subadult female. However, we have decided to establish this taxon pterygostomial regions all completely smooth; sidewall divided into in the light of the distinct nature of the available morphological three parts by longitudinal (epimeral) suture (dividing suborbital, characters, and because of the isolated nature of the study area which region), subhepatic regions from pterygostomial and by Y-shaped may mean that further specimens of P. principe are unlikely to vertical (pleural) groove (dividing suborbital from subhepatic become available for some time. Characters of the gonopods, adult regions). Superior ends of Y-shaped vertical groove meeting male chelipeds. abdomen and sternum are not at present known exorbital, First epibranchial teeth. thoracic sternal sulcus si /s2 deep; because the only specimens of P. principe are subadult females. second sulcus s2/s3 deep, running horizontally across sternum; third Nevertheless, there are a number of unique characters that distinguish sulcus s3/s4 deep, v-shaped, meeting sterno-abdominal cavity. Tho- P. principe from other species in the genus. racic episternal sulci s4/e4, s5/e5, s6/e6 and s7/e7 distinct. Third COMPARISONS WITH OTHER SPECIES. Potamonautes principe is maxillipeds filling entire oral field, except for transversely ovate closest to P. anchietae, a medium-sized species of freshwater crab respiratory openings at superior lateral corners; long flagellum on from Angola (De Brito-Capello, 1871, Bott, 1953, 1955, 1964). This exopod of third maxilliped; ischium with deep vertical sulcus. species was most recently described and illustrated by Bott 1 Epistome prominent, smooth, triangular. Mandibular palp 2-seg- ( 955, p. 247-249, figs. 24, 76, 77, pi. IX, fig. la-d) as P. (Isopotamonautes) mented; terminal segment single, undivided, with hair (but no hard anchietae. Potamonautes principe and P. anchietae are similar in flap) at junction between segments. Subadult female abdomen that both species have a highly arched carapace, a prominent and subcircular, segments al-a6 of female abdomen quadrate, telson complete postfrontal crest, a pointed exorbital tooth, a small granu- (a7) a broad triangle with rounded apex; segments a5-a6 broadest. lar epibranchial tooth, a v-shaped thoracic sternal groove s3/s4, and Major cheliped of subadult female relatively slender, with elon- a similar-sized first carpal tooth on the carpus of the cheliped. gated dactylus and propodus, palm of propodus swollen; fingers of However, there are a number of characters that distinguish the digits of cheliped with small even teeth, forming long narrow specimens from Vissabenguilla, Angola (SMF 1890) described by interspace when closed. First carpal tooth of inner margin of carpus Bott (1955) as P. (I.) anchietae from the specimens from Principe of cheliped large, pointed; second carpal tooth pointed, half size of under consideration here. first tooth. Posterior inferior margin of merus of cheliped smooth, The carapace height of P. principe is greater than that of P. with few small teeth distally; medial inferior margin with row of anchietae (ch/fw P. principe 1 .45, P. anchietae 1.19), and the frontal small sharp teeth along entire length, large pointed tooth at distal margin of P. principe is narrower than that of P. anchietae (fw/cw P. end; superior surface of merus smooth. Pereiopods P2-P5 slender, principe 0.26, P. anchietae 0.39). Further, the anterolateral margins P3 longest, P5 shortest. Dactyli of P2-P5 tapering to point, each of the carapace of P. principe are completely smooth and lack teeth bearing four rows of downward-pointing short, sharp spines. of any kind, whereas these margins in P. anchietae are distinctly COLOUR. The colour of the specimens when freshly caught (before granular. The medial inferior margin of the merus of the cheliped of 16 N. CUMBERLIDGE, P.F. CLARK AND J. BAILLIE U Q tt. U i- g PQ i E 1 . A NEW FRESHWATER CRAB FROM PRINCIPE 17 P. principe has a row of distinct pointed teeth along its length, Acknowledgments. Gilles Joffroy and Tariq Stevart (Universite libre whereas this margin in P. anchietae, lacks teeth, and is either de Bruxelles) assisted in the collection of specimens. Phil Crabb (NHM granular or smooth. Finally, the suborbital and pterygostomial regions Photo. Unit) took the photographs reproduced in this paper. of P. principe are smooth, whereas these regions are distinctly granular in P. anchietae. Bott(1953, 1955, 1 964) described two subspecies of P. anchietae: REFERENCES P. (I.) a. biballensis Rathbun, 1905, and P. (I.) a. machedoi Bott, 1 964, both of which are found in Angola. However, comparison of Balss, H. 1929. Crustacea V. Potamonidae au Cameroun. In: Th. Monod, ed.. the types of these taxa with P. principe indicates that both differ Contributions a I elude de lafaune dtt Cameroun. Faune des Colonies francaises 3: substantially from the new species described here. For example, the 115-129 De Brito-Capello. F. 1871. Algumasespecies novas ou conhecidas pertencentes epibranchial tooth of P. (I.) a. biballensis is large and pointed pouco 5 dos generos Calappu e Thelphusa. Jornal de Sciencias mathematicas, physicas e whereas that of/ , principe is small and granular, and the carapace of nuluraes (Lisbon) 3: 128-134. P. (I.) a. biballensis is flattened, whereas that P. principe is high (ch/ Bott, R. 1953. Potamoniden (Crust. Decap.) von Angola. Publicacoes cuhuruis da fw P. principe 1.45, P. (I.) a. biballensis 1.25). Similarly, the Companhia de Diamantes de Angola. Lisbon 16: 133-148. 1955. epibranchial tooth of P. (I.) a. machedoi is large and pointed whereas Die SuBwasserkrabben von Afrika (Crust.. Decap.) und ihre Stammesgeschichte. Annates du Musc'e dit Congo Beige (Tervuren, Belgique) C- that of P. principe is small and granular, and the frontal margin of P. Zoohgie. Series 111. III. 3: 209-352. principe is narrower than that of P. (I.) a. machedoi (fw/cw P. 1959. Potamoniden aus West-Afrika (Crust.. Dec). Bulletin de I'lnstiiut francais principe 0.26, P. (I.) a. machedoi 0.33). d'Afrique noire 21. serie A (3): 994-1008. Potamonautes principe was also compared here with a specimen I960. Crustacea (Decapoda): Potamonidae. hi: Hansstrom. B. & others. South African Animal Life. Results of Lund University Expedition in 1950-1952: 13-18. of P. margaritarius from Sao Tome (SMF 2668), and the two taxa 1 964. Decapoden aus Angola unter besonderer Beriicksichtigung der Potamoniden can be distinguished by the following characters. The carapace of (Crust. Decap.) und einem Anhang : Die Typen von Thelphusa pelii Herklots. 1861. the latter species is distinctly flattened (ch/fw P margaritarius 0.95, Publicacoes culiurais da Companhia de Diamantes de Angola. Lisboa 69: 23-34. P. principe 1.45), the anterolateral margin of P margaritarius 1968. Decapoden aus dem Museu do Dundo (Crust. Decap.). Publicacoes culiurais da Companhia de Diamantes de Angola. Lisboa 77: 165-172. behind the epibranchial tooth is clearly toothed (whereas this margin 1970. Betrachlungen uber die Entwicklungsgeschichte und Verbreitung der is completely smooth in P principe), and the ischium of the third SUBwasser-Krabben nach der Sammlung des Naturhistorischen Museums in Gent/ maxilliped of P margaritarius is smooth and lacks a vertical suture Schweiz. Revue Suisse de Zoologie 77: 327-344. (whereas this suture is deep in P principe). Bredero. J. T., Heemskerk, VV. & Toxopeus, H. 1977. Agriculture and livestock production in Sao Tome and Principe I West Africa). Unpublished report. Foundation for Agricultural Plant Breeding. Wageningen. Corace, R. G., Cumberlidge. N. & Garms. R. 200 1 . A new species of freshwater crab from Rukwan/i. East Africa. Proceedings oj the Biological Society of Washington. ECOLOGICAL NOTES 114(1): 178-187. Cumberlidge, N. 1993. Redescription of Sudanonautes granulaius (Balss. 1929) i Pi iiamoidea, from West The island of Principe is ovoid in outline, with a total land area of Potamonautidae) Africa. Journal of Crustacean Biology 13: 805-816. about 139 sq km (Fig. 3). The highest point is Pico de Principe 1995. Redescription of Sudanonautes floweri (De Man. 1901) (Brachyura: (945 m), which is a volcanic mountain whose lower slopes are Potamoidca: Potamonautidae) from Nigeria and Central Africa. Bulletin oj the vegetated by lowland rain forest that grades into cloud forest at British Museum of Natural History (Zoology), London 61 (2): 111-119. 1999. higher elevations. The climate of Principe is tropical, hot, and The freshwater crabs of West Africa. Family Potamonautidae. Faune et Flore Tropicales 35, nstitutde recherche pour ledeveloppementdRD, ex-ORSTOM), humid, and there is a single rainy season from October to May. Paris. 1-382. Rainfall levels are high, averaging around 1,000 mm per year in Daniels, S. R., Stewart, B. A. & Gibbons, M. J. 1998. Potamonautes granulans sp. the north and 5,000 mm per year in the south, and monthly tem- ttOV, (Brachyura: Potamonautidae). a new cryptic species of river crab from the peratures range between 25° and 31° C (Bredero et al., 1977). The Olilants ri\cr system, South Africa. Crustaceana 71: 885-903. MacLeay, W. S. 1 838. Brachyurous Decapod Crustacea. Illustrations of the Zoology of two specimens were collected from the summit of the Pico do South Africa 5; being a Portion of the Objects of Natural History Chiefly Collected Principe at 945 meters above sea level on two separate occasions. during an Expedition into the Interior of South Africa, under the Direction of Dr. The terrain at the summit of the Pico do Principe is volcanic and Andrew Smith, in the Years 1834. 1 835. and 1836; Fitted Out by The Cape of Good mountainous, and the vegetation cover is cloud forest with small Hope Association for Exploring Central Africa.' In: A. Smith. Illustrations oj the Zoology South Consisting Descriptions stunted trees, and a high abundance of epiphytes and bryophytes. of Africa; Chiefly of Figures and of the Objects ofNatural History Collected During an Expedition into the Interior ofSouth The thick cloud cover and regular rainfall keep the soil and leaf Africa, in the Years 1834. 1835. and 1836: Fitted Out by The Cape of Good Hope litter very damp. On both occasions when crabs were collected the Association for Exploring Central Africa.' (Invertebrates): 1-75. weather was extremely damp and the mountain top engulfed in De Man, J. G. 1901. Description of a new fresh-water Crustacea from the Soudan: cloud cover. Both specimens were collected on land from under followed by some remarks on an allied species. Proceedings of the Zoological Society of London, 1901: 94-104. damp and decaying leaf litter. There was no river or body of water Milne-Edwards, A. 1 886. La description de quelques Crustaces du genre Thelphusa near the location where the specimens were found. The nearest recueillis par M. de Brazza dans les regions du Congo. Bulletin de la Societe stream was observed at lower altitudes (830 m), but there were Philosophiqtte Paris series 7(10): 1 48- 1 5 1 numerous temporary small pools at the summit which form after Rathbun, M. J. 1905. Les crabes d'eau douce (Potamonidae). Nouvelles Archives du Museum d'Histoire naturelle (Paris) 7(4): 159-322. heavy precipitation. It is likely that P principe can breathe air, Stewart, B. A. 1997a. Morphological and genetic differentiation between populations given its extremely terrestrial lifestyle. When crabs were disturbed of river crabs (Decapoda: Potamonautidae) from the Western Cape, South Africa, by removing the leaf litter cover, they ran rapidly across the forest with a taxonomic re-examination of Gecarcinautes brincki. Zoological Journal of floor and took cover under leaf litter or any available crevasse. the Linnean Society 199: 1—21. 1997b. Biochemical and morphological evidence for a new species of river crab Specimens were collected close to holes near the Pico do Principe, Potamonautes parvispina (Decapoda: Potamonautidae). Crustaceana 70: 737-753. but crabs were not actually observed moving in or out of these , Coke, M. & Cook, P. A. 1 995. Potamonautes dentatus. new species, a fresh-water holes, and it ramains possible that the holes may not have been dug crab (Brachyura: Potamoidea: Potamonautidae) from KwaZulu-Natal. South Africa. by the crabs. Journal of Crustacean Biology 15: 558. N. CUMBERLIDGE, P.F. CLARK AND J. BAILLIE Principe Gulf of Guinea Bombom 1° 40' N Porto Real Terriro Velho Maria Correia 1°35'N Pico do Principe Antonio Enes • ^Zj Bone do Joquei 7°20'E 7° 25' E Fig. 3 Map of Principe showing the type locality of Potamonautes principe sp.nov. Yx(-3S7-&<+-\) Bull. not. Hist. Mas. Land. (Zool.) 68( 1 ): 19-26 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 5BD, U.K. ALASDAIR J. EDWARDS Tropical and Coastal Management Studies, Department of Marine Sciences and Coastal Management, University of Newcastle, Newcastle upon Tyne NE1 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. 11,23 dorsal-fin rays. 26-28 scales in lateral series, and scales with distinct centres and radii in all fields. INTRODUCTION 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 The genus Pseudoplesiops Bleeker was recently diagnosed to membranes. Pectoral fin length was measured as the length of the included pseudoplesiopine pseudochromids with the medial lami- longest middle ray. Caudal fin length was measured as the length of nae of the pelvic bones expanded dorsally (Gill & Edwards. 1999). the lowermost ray on the dorsal hypural plate. Aside from this synapomorphy, members of the genus are distin- Counts of dorsal-, anal- and pelvic-fin spines (unsegmented rays) guished from other pseudoplesiopines in possessing the following and segmented rays are presented, respectively, as Roman and combination of external characters: scales in lateral series 26^2; Arabic numerals. If the last dorsal- or anal-fin ray was divided at its lower lip complete (uninterrupted at symphysis); and preopercular base it was counted as a single ray. Counts of branched, segmented pores usually 7 (rarely 6 or 8), with a pore present at the upper rays in the dorsal and anal fins included unbranched rays behind the terminus of the preopercle. The genus is mostly confined to the first branched ray. A value was not recorded if, due to tip damage, a eastern Indian and Pacific Oceans, but two undescribed species branched or unbranched condition could not be determined for the occur in the Maldive Islands, central Indian Ocean. Although it is segmented ray preceding the anteriormost branched ray. As in most our intention to produce a revision of the genus, we herein describe actinopterygian fishes, the upper ray in the pectoral fin is rudimen- the two Maldives species in order to make their names available for tary and rotated so that its asymmetrical medial and lateral hemitrichs a forthcoming guide to western and central Indian Ocean fishes. 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 (Mower) the principal caudal-fin rays. The uppermost MATERIALS AND METHODS principal caudal-fin ray was defined as the ray articulating with hypural 5, and the lowermost principal caudal-fin ray was the ray Institutional abbreviations follow Leviton et al. (1985). All meas- articulating with the cartilage nubbin between the distal tips of the urements to the snout tip were made to the midanterior tip of the parhypural and the haemal spine of preural centrum 2. All upper lip. Length of specimens are given in mm standard length pseudoplesiopine species normally have 1 7 (9 + 8) principal caudal- (SL), which was measured from the snout tip to the middle of the fin rays. caudal peduncle at the vertical through the posterior edge of the Counts of 'scales in lateral series' were of the posteroventrally dorsal hypural plate. Head length was measured from the snout tip to oriented transverse scale rows on the midside, beginning with the the posteriormost edge of the opercular membrane. Snout length row through the tubed scale at the branchial opening and ending with was measured over the shortest distance from the snout tip to the the row through the scale at the midposterior edge of the hypural orbital rim, without constricting the fleshy rim of the latter. Orbit plate. 'Scales in transverse series' were counted anterodorsally from diameter was measured as its fleshy horizontal length. Interorbital the anal-fin origin to the dorsal-fin base. Circumpeduncular scales width was measured as the least fleshy width. Upper jaw length was were counted in a zig-zag fashion around the middle of the caudal measured from the snout tip to the posterior edge of the maxilla. peduncle. Gill-raker counts were of the outer rakers on the first arch, Predorsal, preanal and prepelvic lengths were measured from the including rudiments; the angle raker is included in the lower-limb snout tip to the base of the first spine of the relevant fin. Body width (second) count. Counts of pseudobranch filaments included all was measured between the posttemporal pores. Caudal peduncle rudiments. Nomenclature of head pores follows Winterbottom length was measured from the base of the last anal-fin ray to the ( 1986), as modified by Gill et al. (2000). ventral edge of the caudal fin at the vertical through the posterior Counts of vertebrae are presented in the form precaudal + caudal © The Natural History Museum. 2002 20 A.C. GILL AND A.J. EDWARDS = total. Caudal vertebrae are defined as those with a haemal spine, Islands); Gill in Randall et al., 1997: 131 (description; distri- and include the terminal urostylar complex (which was counted as a bution; col. fig.); Kulbicki & Williams, 1997: 14 (Ouvea Atoll, single vertebra). The pattern of insertion of supraneural (predorsal) New Caledonia). bones and anterior dorsal-fin pterygiophores within interneural Pseudoplesiops n. sp.; Allen & Smith-Vaniz, 1994: 10 (Cocos spaces is given as an 'anterior dorsal-fin pterygiophore formula' (Keeling) Islands). modified from the 'predorsal formula' of Ahlstrom et al. (1976). Pseudoplesiops sp. 1.; Gill & Edwards, 1999: 144 (list of osteologi- Each supraneural is represented by an 'S,' neural spines are repres- cal materials); Gill, 2000: 2560 (key). ented by slashes, and pterygiophores are represented by '3' HOLOTYPE. AMS 1.20757-069, 29.8 mm SL, Great Barrier Reef, (indicating a pterygiophore that bears two supernumerary rays and W end of Raine Island (11°36'S 144°01'E), coral gutter, 2-20 m, a serially associated ray), '2' (indicating a pterygiophore that bears AMS and Australian Institute of Marine Science team, 13 February a supernumerary ray and a serially associated ray) or ' 1' (indicating 1979. a pterygiophore that bears only a serially associated ray). An 'anterior anal-fin pterygiophore formula' is also presented, and is PARATYPES. AMS 1.17090-043, 1: 26.5 mm SL, Papua New similar to the anterior dorsal-fin pterygiophore formula, except that Guinea, Madang Harbour, S edge of Massas Island, coral reef, 1.5- the slashes represent haemal spines. Osteological features were 14 m, B.B. Collette etal., 31 May 1970; AMS 1.17094-013, 1: 28.5 determined from x-radiographs and from cleared-and-stained speci- mm SL, Papua New Guinea, Trobriand Islands, N end of Kiriwinna mens, which were prepared following the methods of Taylor & Van Island, 0-5 m, B.B. Collette and B. Goldman, 7 June 1970; AMS Dyke (1985). 1.17094-014, 1: 30.4 mm SL, collected with AMS 1.17094-013; Counts and measurements are given as values or value ranges for AMS 1.17096-008, 1: 27.8 mm SL, Papua New Guinea, Trobriand all type specimens, followed, where different, by values for the Islands, Kiriwinna Island, off N coast of Tawai Point, coral with holotype in parentheses. Where counts were recorded bilaterally, sand patches, 7.5 m, B.B. Collette, 8 June 1970; AMS 1.20756-014, both counts are presented for the holotype, separated by a slash; the 1: 25.7 mm SL (subsequently cleared and stained). Great Barrier first count given is the left count. Reef, Great Detached Reef, over coral and sand, 2-8 m, AMS and Australian Institute of Marine Science team, 11 February 1979; Pseudoplesiops immaculatus sp. nov. AMS 1.20756-124, 2: 15-30 mm SL, collected with AMS 1.20756- 014; AMS 1.20757-039, 1: 29.4 mm SL, collected with holotype; Bearded Dottyback AMS 1.20757-091, 3: 27.6-30.5 mm SL, collected with holotype; Figures 1 , 2 AMS 1.20784-040, 2: 17.0-21.9 mm SL, Great Barrier Reef, Yonge Pseudoplesiops typus [non Bleeker, 1858]; Bleeker, 1875: 31 (in Reef, 1 mile N of platform, back reef knoll, 1-15 m, D.F Hoese et part, specimen from Amboina). al, 1 December 1978; AMS 1.21972-004, 1: 31.2 mm SL, Solomon Chilidichthys [sic] sp. 1 ; Allen & Steene, 1979: 26 (Christmas Island Islands, Guadalcanal, 1 2 km W of Honiara, over wreck, J.E. Randall. and Cocos Keeling Atoll, Indian Ocean). 12 July 1975; AMS 1.22616-044, 1: 20.9 mm SL, Great Barrier Pseudoplesiops revellei [non Schultz, 1953]; Kailola, 1987: 244 Reef, Escape Reef, coral reef, 5-18 m, J.R. Paxton et al., 2 Nov- (Papua New Guinea); Allen & Steene, 1988: 180, fig. 145 (Christ- ember 1981; AMS 1.226 1 9-025, 1 : 26.0 mm SL, Great Barrier Reef, mas Island, Indian Ocean); Paxton et al., 1989: 521 (list; Great Escape Reef North ( 15°49'S 45°50'E), sand slope, coral and sand in Barrier Reef). cave under bommie, 19-22 m, J. Paxton et al, 3 November 1981; Pseudoplesiops sp.; Gill in Randall et al., 1990: 131 (description; ANSP 131727, 2: 27.2-27.5 mm SL, Cocos Keeling Islands, West distribution; col. fig.); Randall & Anderson, 1993: 15 (Maldive 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 R Crabb.) 1 TWO NEW SPECIES OF PSEUDOPLESIOPS 21 NASP DENP POPP SOBP Fig. 2 Cephalic laterosensory pores of Pseudoplesiops immaculatus, AMS 1.20757-069. 29.8 mm SL. holotype. Raine Island. Great Barrier Reef. AIOR anterior interorbital pore: AN, anterior nostril; DENR dentary pores; IMF. intermandibular Hap; ITR intertemporal pore; LSCR pore in tubed lateral-line scale (other details of scale omitted); NASR nasal pores; PARR parietal pores; PN. posterior nostril: POPP. preopercular pores; PTR 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. mm SL, Taiwan, off Houpihu, P.C. Heemstra, 20 January 1988; Smith-Vaniz et ai, 24 February 1974; ANSP 131728, 1: 30.6 mm USNM 20959 1,3: 25.3-27.0 mm SL, Indonesia, point E of Tandjung SL, Cocos Keeling Islands, West Island, ca. 1 km NNW of N end of Naira, Haruka Island, surge channel, 4.5 m, VG. Springer and M.F. island (12°07'20"S 96°49'05"E), rocky patch fronting on sand, W.F. Gomon, 15 January 1973; USNM 209965, 1: 23.7 mm SL, Indone- Smith-Vaniz et al, 1 March 1974; ANSP 178041, 1: 30.8 mm SL, sia, Saparua, two stations mixed (isolated coral head surrounded by Papua New Guinea, Bougainville Island, Tautsina Island, E of Kieta crinkly calcareous matrix at 9 m and coral patch in 3.6 m), VG. Peninsula, coral reef off stack at N end of island, 0.9-7.5 m. 1 Springer and M.F. Gomon, 18 January 1973; USNM 290118, 6: March 1965 (Te Vega Expedition Cruise no. 6, Station 247); ASIZT 17.4-28.9 mm SL, Papua New Guinea, Hermit Islands, Amot 57016, 1: 25.8 mm SL, Taiwan, J.P. Chen, 22 April 1994; ASIZT Island, ocean side of reef at drop off (01°33'S 144°59'E), 0-15.2 m, 56991, 1: 19.7 mm SL, Taiwan, 9 m, J.P. Chen, 21 April 1994; VG. Springer et al, 30 October 1978; USNM 290327. 5: 27.7-29.5 BMNH 1974.5.25.973, 1: 22.8 mm SL, Papua New Guinea, mm SL, Papua New Guinea, S tip of Massas Island (05°10'18"S Trobriand Islands, Kiriwinna Island, E shore NW of Gusaweta, 145°51'24"E), 0-24 m, VG. Springer et al., 6 November 1978; exposed coral platform, 0-3.6 m, B.B. Collette, 17 June 1970; CAS USNM 290437, 1 : 3 1 . 1 mm SL, Papua New Guinea, Ninigo Islands, 58686, 2: 26.3-27.9 mm SL, Maldive Islands, Male Atoll, reef off just SE of Ami Island (01 °14'S 144°22'E), patch reef behind reef, 0- SE side of Funidu Islet, inside atoll lagoon (04°H'00"N 4.5 m. VG. Springer et al, 22 October 1978; USNM 290749, 3: 073°30'30"E), M.G.Bradbury, 6 November 1964; MNHN 1994-45, 23.3-29.9 mm SL, Indonesia, Banda Islands, just W of N tip of 1: 22.6 mm SL, New Caledonia, Loyalty Islands, Ouvea Atoll, Great Banda Island (04°30'30"S 129°56'10"), 0-18 m,VG. Springer Bagaat Islet (20°37'18"S 166°16'08"E), vertical reef wall with small and M.F. Gomon, 9 March 1974; USNM 290791, 1: 30.3 mm SL, cave and base of wall with coral rubble platform, 15-21 m, J.T. Indonesia, Ambon, Latuhalat, Namalatu, about 150 m offshore Williams, J.-L. Menou and P. Tirard, 16 November 1991; QM (03°47'S 128°06'E), 10.5-18 m, VG. Springer et al., 14 March 1.15527, 2: 26.8-33.8 mm SL, collected with holotype; RMNH 1974; USNM 290808, 1: 22.0 mm SL, collected with USNM 31188, 1: 28.7 mm SL, Ambon (Amboina); RUSI 35701, 1: 30.9 290749; USNM 29 1 609, 1 : 25.7 mm SL, Philippine Islands, Batanes, ; 22 A.C. GILL AND A.J. EDWARDS Batan Island, past Mahate, White Beach (20°24'45"N 12 1°55'00"E), verse scale rows behind branchial opening; gill rakers 1-4 + 6-10 = coral and encrusted boulders, 9-12 m, G.D. Johnson and W.F. 7-12 (2 + 8); pseudobranch filaments 5-8 (6). Smith- Vaniz, 1 May 1987; USNM 292031, 1: 27.8 mm SL, Papua Head pores (all bilaterally paired; Fig. 2): nasal pores 2-3 (2/2); New Guinea, Hermit Islands, N side of W entrance (01°30'30"S anterior interorbital pores 1 ; posterior interorbital pores 0; supraotic 144°59'15"E), 0-12 m, V.G. Springer et ai, 4 November 1978; pores 2; suborbital pores 7-9 (8/8); posterior otic pores 0; USNM 322986, 1: 31.8 mm SL, New Caledonia, Loyalty Islands, preopercular pores 7-8 (7/7); dentary pores 4; intertemporal pores 1 Ouvea Atoll, Motu Veiloa Islet (20°26'06"S 166°28'30"E), reef anterior temporal pores 0; posttemporal pores 1; parietal pores 2. crest, small patch reef of mostly dead coral surrounded by rubble, As percentage of standard length (based on 34 specimens, 22.8- 0.6-3.3 m, J.T. Williams and M. Kulbicki, 17 November 1991; 33.8 mm SL): body depth at dorsal-fin origin 22.8-26.3 (23.8); USNM 322994, 1: 32.1 mm SL, New Caledonia, Loyalty Islands, greatest body depth 23.1-28.1 (26.5); body width 11.5-13.3(12.4); 4' Ouvea Atoll, Recif Draule (20°34' 1 2"S 1 66° 1 1 2"E), large depres- head length 28.8-32.6 (29.9); snout length 5.3-6.2 (6.0); orbit sion on submerged reef crest, coral and rubble, 5 m, J.T. Williams, P. diameter 7.7-9.9 (8.1); interorbital width 2.2-3.6 (3.0); upper jaw Tirard and J.L. Menou, 16 November 1991; USNM 328198, 1:24.5 length 10.0-11.2(10.1); depth of caudal peduncle 13.2-15.3(14.8); mm SL, collected with MNHN 1994-45; USNM 356587, 1: 20.9 caudal peduncle length 7.3-9.4 (8.1); predorsal length 29.5-32.6 mm SL, Vanuatu, Shepherd Islands, Judy Reef off NW tip of Tongoa (29.5); preanal length 54.7-57.6 (56.7); prepelvic length 26.4-29.2 Island (16°52'30"S 168°31'30"E), coral reef with extensive coral (28.2); length of first segmented dorsal-fin ray 8.7-10.8 (9.7); development, J.T. Williams et ai, 9 June 1996; USNM 357981, 1: length of third from last segmented dorsal-fin ray 16.4-20.8 (17.4); 26.8 mm SL, Solomon Islands, Santa Cruz Islands, Reef Islands, dorsal-fin base length 60.5-67.8 (63.1); length of first segmented Fenualoa Island, just W of Nota Point (ca. 10°16'30"S 166°16'30"E), anal-fin ray 9.3-12.3 (11.1); length of third from last anal-fin ray coral reef face and outer slope, rich coral growth with sand and 13.6-20.4 (17.8); anal-fin base length 33.3-37.7 (35.6); caudal fin rubble, 0-13 m, J.T. Williams et ai, 18 September 1998; USNM length 20.4-26.9 (24.8); pectoral fin length 18.4-21.8 (18.5); pelvic 358382, 1: 20.4 mm SL, Solomon Islands, Santa Cruz Islands, Reef fin length 25.7-32.0 (27.8). Islands, Lomlom Island, Nialo Point on E side of Forrest Passage Lower lip complete; prominent intermandibular flap present (10°16'S 166°18'30"E), vertical reef wall and rocky surge channels (Fig. 2); fin spines weak and flexible; anterior dorsal-fin v at surface, 0-35 m, J.T. Williams et ai, 18 September 1998; WAM pterygiophore formula S/S/S + 2/1 + 1, S/S/S + 2/1 + 1 or S/S/2/ v P.26083-04 1 , 1 : 3 1 .2 mm SL, Indian Ocean, Christmas Island, Ethel 1 + 1 (S/S/S + 2/1 + 1); 20-22 (21) consecutive dorsal-fin Beach, 3-6 m, G.R. Allen and R.C. Steene, 19 May 1978; WAM pterygiophores inserting in 1 : 1 relationship directly behind neural P.26085-030, 1 : 25.9 mm SL, Indian Ocean, Christmas Island, Ethel spine 4; anterior anal-fin pterygiophore formula 2 + 1/1,3 + 1/1 or Beach, 15-20 m, G.R. Allen and R.C. Steene, 20 May 1978; WAM 2 + 1 + 1/1 (2 + 1/1); 11-12 (11) consecutive anal-fin P.26093-015, 1 : 27.3 mm SL, Indian Ocean, Christmas Island, 1 km pterygiophores inserting in 1:1 relationship directly behind hae- W of Margaret Beach, 10 m, G.R. Allen and R.C. Steene, 25 May mal spine 2; second segmented pelvic-fin ray longest; caudal fin 1978; WAM P.26104-005, 1 : 27.3 mm SL, Indian Ocean, Christmas rounded to truncate or slightly emarginate; scales without distinct Island, Flying Fish Cove (10°29'S 105°40'E), 6-8 m, G.R. Allen and centres, and with radii confined to anterior field; dorsal and anal R.C. Steene, 1 June 1978; WAM P.26107-004, 1: 24.0 mm SL, fins without distinct scale sheaths, although sometimes with inter- Indian Ocean, Christmas Island, Rhonda Beach (10°29'S 105°40E), mittent scales overlapping fin bases; anterior lateral line 6-7 m, G.R. Allen and R.C. Steene, 2 June 1978; WAM P.26113- represented by single tubed scale at branchial opening, followed 003, 2: 25.6-28.4 mm SL, Indian Ocean, Christmas Island, Winifred by intermittent series of centrally pitted scales, which terminate at Beach (10°29'S 105°40'E), 12-14 m, G.R. Allen and R.C. Steene, 6 vertical through base of segmented dorsal-fin ray 21-27, or extend June 1978; WAM P.27825-034, 3: 17.6-25.4 mm SL, Papua New slightly past end of dorsal fin (extending slightly beyond end of Guinea, Manus Island, Los Negros Island, SE point at aerodrome, dorsal fin/to vertical through base of segmented dorsal-fin ray 26 10^0 m, G.R. Allen and R. Knight, 5 October 1982; WAM P.29626- in holotype); second intermittent series of centrally pitted scales 001, 1: 31.2 mm SL, Papua New Guinea, Port Moresby (09°30'S originating on midside above anterior part of anal fin, extending 147°10'E), 5-6 m, P. Colin, 24 February 1987; WAM P.29927-004, on to middle of caudal-fin base; additional centrally pitted scales 1: 23.9 mm SL, Indian Ocean, Cocos-Keeling Islands, Direction present on caudal-fin base, pits usually aligned vertically on pos- Island (12°05'S 096°53'E), 0.1-2.0 m, G.R. Allen, 24 February terior part of scale sheath; scales present on cheeks (not extending 1989. posteriorly over upper part of preopercle) and operculum (Fig. 2); predorsal scales extending anteriorly to supratemporal commis- Diagnosis sure (Fig. 2); vertebrae 12 + 20-22 (12 + 21); epurals 2; epineurals A species of Pseudoplesiops with the following combination of present on vertebrae 1 through 17-21 (1 through 19); ribs present characters: prominent intermandibular flap present; and operculum on vertebrae 3 through 11-12 (3 through 12), rib on ultimate immaculate, without large dark spot. precaudal vertebra very small to moderately small or absent. Description. (Based on 72 specimens, 15.0-33.8 mm SL) Dor- Upper jaw with 2-6 pairs of curved, enlarged caniniform teeth sal-fin rays 1,26-28 (1,27), last 3-11 (8) segmented rays branched; anteriorly, the medial pair smallest, and 3-4 (at symphysis) to 1-2 anal-fin rays 1-11,16-1 8 (1, 17), last 3-9 (5) segmented rays branched; (on sides ofjaw) irregular inner rows of small conical teeth, the teeth pectoral-fin rays 15-18(16/15), upper 1-5 (2/2) and lower 1-3 (1/1) of outer row of conical teeth largest; lower jaw with 2-4 pairs of rays simple; pelvic-fin rays 1,3—4 (1,4), all segmented rays simple; curved, enlarged caniniform teeth, the medial pair smallest, and 2- principal caudal-fin rays 9-10 + 8 (9 + 8), the uppermost 1-2 (1) and 4 (at symphysis) to 1 (on sides of jaw) irregular inner rows of small lowermost 1-2 (1) rays unbranched; upper procurrent caudal-fin conical teeth, the conical teeth gradually increasing in size and rays 3-5 (4); lower procurrent caudal-fin rays 2-\ (3); total caudal- becoming more curved on middle part of jaw, then becoming fin rays 23-25 (24); scales in lateral series 32-39 (36/35); predorsal abruptly smaller on posterior part ofjaw; vomer with 1 row of small, scales 7-12 (10); transverse scales 14-17 (16/16); scales behind eye stout conical teeth arranged in a chevron; palatines edentate or with 1-2 (2); scales to preopercular angle 3-4 (3); circumpeduncular small irregular patch of small conical teeth; tongue acutely pointed, scales 16-17 (16); ctenoid scales beginning at 10-16 (14/14) trans- edentate. TWO NEW SPECIES OF PSEUDOPLESIOPS 23 Live coloration. (Based on colour photographs of the holotype P. revellei is it well-developed. On the basis of this synapomorphy, from the Great Barrier Reef, and of paratypes and other specimens we suggest that the two species are sister taxa. They are distin- from the Great Barrier Reef, Loyalty Islands, Solomon Islands, guished from each other by a single coloration character: P. revellei Vanuatu, Christmas Island and Taiwan) has a large, dark brown to black spot on the operculum, which is Head and body pinkish or yellowish brown to olive or bright absent in P. immaculatus. to brown snout, lips green, sometimes becoming pinkish orangish on Remarks and intermandibular flap; iris pale yellow to green or brown, some- Colour photographs of the species have been published by Allen & times with reddish grey to pink area around pupil; dorsal and anal Steene (1988; as P. revellei), and Randall etal., (1990, 1997 - as P. fins yellowish, pinkish or orangish brown to olive or bright green, sp.). becoming paler distally, with bluish grey to bright blue distal margin, sometimes with broad pale orange to yellow stripe Etymology submarginally; bluish grey to pale blue spot or streak at base of The specific epithet is from the Latin, meaning without a spot, and alternate dorsal- and anal-fin rays; caudal fin yellowish or pinkish alludes to the sole character distinguishing P. immaculatus from its brown to olive or bright green; pectoral fins pinkish, greenish or sister species. yellowish hyaline; pelvic fins pale pink or olive to bright green, Pseudoplesiops occidentalis sp. nov. usually bluish grey to pale blue anteriorly and distally. Maldives Dottyback Preserved coloration Figures 3, 4 Head and body pale brown, slightly darker on dorsal part of head and body, and on lips and intermandibular flap; fins pale brown to Clinus sp.; Regan, 1902: 276 (description; Haddumati, Maldive brownish hyaline; bluish grey to blue spots and distal margins of Islands). dorsal and anal fins described above become greyish brown. Pseudoplesiops sp.; Randall & Anderson, 1993: 15, pi. 3e (Maldive Islands; col. fig.). Habitat and Distribution Pseudoplesiops sp. 1 ; Kuiter, 1998: 77 (habitat notes; colour photo). This species is distributed from the Maldive Islands, east and south HOLOTYPE. 32926, 24.9 SL, Maldive Islands, South to Vanuatu, and north to Taiwan. It has been collected from a variety BPBM mm of reef habitats, from shallow patch reefs to reef walls at depths Male Atoll, Maaniyafushi Island, reef, 25-30 m, J.E. Randall, C. Anderson and M.S. 17 1988. ranging to at least 20 m (with some collections perhaps from as deep Adam, March as 40 m). PARATYPES. AMS 1.41004-001, I: 21.3 mm SL (subsequently cleared and stained), collected with holotype; BMNH 1 901 . 1 2.3 1 .77, Comparisons with other species 1 : 26.2 mm SL, Maldive Islands, Haddummati Atoll (= Haddumati), Pseudoplesiops immaculatus has been confused with P. revellei 72 m, J.S. Gardiner; BPBM 32871, 2: 16.7-26.5 mm SL, Maldive Schultz, a Pacific Plate endemic {sensu Springer, 1982). The two Islands, Ari Atoll, E side of reef N of Bathala Island, rubble bottom, species closely resemble each other in morphometric and meristic 35 m, J.E. Randall and M.S. Adam, 6 March 1988. values, and both possess a prominent intermandibular flap (Fig. 2), although this may be weakly developed or absent in small specimens Diagnosis (smaller than about 18 mm SL). A low intermandibular ridge or Pseudoplesiops occidentalis is distinguished from all other pseudo- weak flap may be present in certain other pseudochromid species plesiopines in having the following combination of characters: (e.g., Chlidichthys cacatuoides, see Gill & Randall, 1994; dorsal-fin rays 11,23; scales in lateral series 26-28; and scales with Pseudoplesiops occidentalism Fig. 4), but only in P. immaculatus and distinct centres and radii in all fields. Fig. 3 Pseudoplesiops occidentalis, BPBM 32926, 24.9 mm SL, holotype. South Male Atoll, Maldive Islands. (Photo by P. Crabb.) 24 A.C. GILL AND A.J. EDWARDS PTP LSCP DENP IMR POPP i i Fig. 4 Cephalic sensory pores of Pseudoplesiops occidentalis, BPBM 32926. 24.9 mm SL, holotype. South Male 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) 16.5-16.9 (16.5); caudal peduncle length 7.2-8.5 (7.2); predorsal Dorsal-fin rays 11,23, last 6-7 (6) segmented rays branched; anal-fin length 29.0-32.9 (31.3); preanal length 55.0-61.1 (60.2); prepelvic rays 11,14, last 5-6 (5) segmented rays branched; pectoral-fin rays length 30.1-32.5 (30.1); length of first segmented dorsal-fin ray 15-16 (16/15), upper 2-4 (2/2) and lower 1-3 (1/2) rays simple; 10.4-11.7 (10.4); length of third from last segmented dorsal-fin ray pelvic-fin rays 1,3, all segmented rays simple; principal caudal-fin 17.7-19.7 (17.7); dorsal-fin base length 61. 7-71.4 (62.7); length of rays 9 + 8, the uppermost 1-2 (1) and lowermost 1-2 (2) rays first segmented anal-fin ray 12.2-14.5 (12.9); length of third from unbranched; upper procurrent caudal-fin rays 3-4 (4); lower last anal-fin ray 17.8-18.7 (18.5); anal-fin base length 32.8-35.9 procurrent caudal-fin rays 3~4 (4); total caudal-fin rays 23-25 (25); (33.7); caudal fin length 25.3-27.9 (25.3): pectoral fin length 22.5- scales in lateral series 26-28 (27/27); predorsal scales 6-8 (8); 24.0 (23.3); pelvic fin length 27.7-35.3 (27.7). transverse scales 11-12 (11/11); scales behind eye 2; scales to Lower lip complete; prominent intermandibular flap absent, al- preopercular angle 3; circumpeduncular scales 16; ctenoid scales though low ridge present in some specimens (Fig. 4); fin spines beginning at 7-9 (7/9) transverse scale rows behind branchial weak and flexible; anterior dorsal-fin pterygiophore formula S/S/3/ opening; gill rakers 2-\ + 8-11 = 10-15 (2 + 8); pseudobranch 1 + 1; 15-16 (15) consecutive dorsal-fin pterygiophores inserting in filaments 6-7 (7). 1:1 relationship directly behind neural spine 4; anterior anal-fin Head pores (all bilaterally paired; Fig. 4): nasal pores 2; anterior pterygiophore formula 2/1 + 1; 7-8 (8) consecutive anal-fin interorbital pores 1 ; posterior interorbital pores 0; supraotic pores 2; pterygiophores inserting in 1:1 relationship directly behind haemal suborbital pores 8-9 (9/8); posterior otic pores 0; preopercular pores spine 2; second segmented pelvic-fin ray longest; caudal fin weakly 7; dentary pores 4; intertemporal pores 1 ; anterior temporal pores 0; rounded to rounded or slightly emarginate; scales with distinct posttemporal pores 1 ; parietal pores 2. centres and radii in all fields; dorsal and anal fins without distinct As percentage of standard length: body depth at dorsal-fin origin scale sheaths, although with basal row of body scales overlapping 24.0-28.2 (26.5); greatest body depth 24.0-32.4 (27.7); body width fin base, particularly on posterior part of fins; anterior lateral line 13.3-14.5 (13.3); head length 30.5-33.5 (30.5); snout length 5.0- represented by single tubed scale at branchial opening, followed by 6.4 (5.6); orbit diameter 8.4-10.2 (8.4); interorbital width 3.6-4.8 intermittent series of centrally pitted scales, which terminate at (3.6); upper jaw length 11.3-12.0 (11.6); depth of caudal peduncle vertical through base of segmented dorsal-fin ray 17-22 (?/22); TWO NEW SPECIES OF PSEUDOPLESIOPS 25 second intermittent series of centrally pitted scales originating on guished from each other by the following: number of vertebrae (10 midside above anterior part of anal fin, extending on to middle part + 17 in P. occidentalis, 11-12 + 16-18 = 27-29, usually 11 + 17- of caudal-fin base; additional centrally pitted scales present above 18 in the P. rosae-complex and 11 + 17-18 in the P. and below pitted scale(s) on middle part of caudal-fin base, some- fy/?ws-complex); number of dorsal-fin rays (11,23 in P. occidentalis, times extending on to posterior part of caudal peduncle; scales 1,22-24 in the P. rosae-complex and 11,24-25 in the P. typus- present on cheeks (extending posteriorly over upper part of complex); number of anal-fin rays (11,14 in P. occidentalis, preopercle) and operculum (Fig. 4); predorsal scales extending 1-11,12-14, usually 1,13-14 in the P. rosae-complex and 11-111,14- anteriorly to point between anterior interorbital pores and vertical 16, usually 11,15 in the P. ry/n/s-complex), number of scales in through anterior supraotic pores (Fig. 4); vertebrae 10 + 17; epurals lateral series (26-28 in P. occidentalis, 26-29 in the P. rosae- 2; epineurals present on vertebrae 1 through 16-18 (1 through 16); complex and 32-40 in the P. typus-complex); number of ribs present on vertebrae 3 through 10, rib on ultimate precaudal circumpeduncular scales (16 in P. occidentalis, 16 in the P. rosae- vertebra relatively long. complex and 20-22 in the P. npiis-comp\ex); predorsal scalation Upper jaw with 2-5 pairs of curved, enlarged caniniform teeth (6-8 scales, extending anteriorly to a point between the anterior anteriorly, the medial pair smallest, and 3-4 (at symphysis) to 1-2 interorbital pores and the vertical through the anterior supraotic (on sides ofjaw) irregular inner rows of small conical teeth, the teeth pores in P. occidentalis, 5-10 scales, extending anteriorly to a of outer row of conical teeth largest; lower jaw with 2-3 pairs of point between the anterior interorbital pores and the vertical curved, enlarged caniniform teeth, the medial pair smallest, and 2- through the anterior supraotic pores in the P. rosae-complex, and 3 (at symphysis) to 1 (on sides of jaw) irregular inner rows of small 10-16 scales, extending anteriorly to the supratemporal commissure conical teeth, the conical teeth gradually increasing in size and in the P. ryp«.v-complex); and cheek scalation (broadly overlapping becoming more curved on middle part of jaw, then becoming the upper part of the preopercle in P. occidentalis and the P. rosae- abruptly smaller on posterior part ofjaw; vomer with 1 row of small, complex, versus not overlapping the upper part of the preopercle in stout conical teeth arranged in a chevron; palatines edentate or with the P. rvpws-complex). Members of the P. fy/JMs-complex also small irregular patch of small conical teeth; tongue pointed, eden- attain a much larger body size than members of the other clades tate. (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). LIVE COLORATION. (Based on colour photograph of the holotype, Of the characters noted above that are shared by P. occidentalis photograph in Kuiter, 1998, and field notes taken from paratypes in and the P. rosae-complex, three are unique within Pseudoplesiops BPBM 32871 when freshly dead) and suggest a sister-relationship between the two taxa: low number Head and body bright pinkish red to orange-red, becoming pink of scales in lateral series; predorsal scales extending anteriorly ventrally and olive-red to orangish brown posteriorly; margin of beyond the supratemporal commissure; and cheek scales broadly orbit orange posteriorly, becoming pale blue ventrally; posttemporal, overlapping upper part of preopercle. This relationship will be tested intertemporal, upper preopercular and upper suborbital pores indis- in a study of the phylogeny of the Pseudochromidae currently in tinctly edged with grey; iris bright orange-red; pectoral-fin base pale progress by the first author. pink to pinkish red; dorsal and anal fins bright orange-red basally. reddish hyaline distally, with pale blue distal margin; caudal fin Remarks greyish yellow to pale greyish red basally, remainder of fin reddish Colour photographs of the species are provided by Randall & to yellowish hyaline with pale blue distal margin; pectoral fin Anderson (1993) and Kuiter (1998). Randall & Anderson indicate pinkish to yellowish hyaline; pelvic fin pale pink basally. becoming that their photograph is of a 26 mm SL specimen in BPBM 32871, pale blue distally. but it is actually of the holotype (BPBM 32926). Regan's (1902) specimen of the species ( BMNH 190 1 . 1 2.3 1 .77) Preserved coloration was initially identified and reported on as "Clinus sp." A label on the Head and body brown, paler ventrally; grey edging on posttemporal. jar and catalogue entry indicates that it was subsequently intertemporal, upper preopercular and upper suborbital pores re- redetermined as the plesiopid Belonepterygion fasciolatum Ogilby mains, becoming greyish brown; dorsal and anal fins brownish (apparently by M.L. Penrith). Although we were unable to locate hyaline, becoming greyish brown distally; other fins pale brown to any references to this identification, they possibly exist. dusky hyaline. Etymology Habitat and distribution The specific epithet is from the Latin, meaning of the west; P. occi- Pseudoplesiops occidentalis is known only from the Maldive Is- dentalis is known only from the western-most part of the range of the lands. It has been collected from and observed on reefs in 20 to 72 m. genus. Comparisons with other species Pseudoplesiops occidentalis forms a monophyletic group with P. typus Bleeker, P. rosae Schultz and several undescribed species. Acknowledgements. We thank the following for allowing access to With the exception of P. occidentalis, this clade is confined to the specimens in their care: G.R. Allen. A. Bentley, D. Catania. J. -P. Chen, O.A. eastern Indian Ocean and the West Pacific. It is diagnosed by Crimmen, W.N. Eschmeyer, P.C. Heemstra, D.F. Hoese, S. Jewett. J. Johnson, scales with distinct centres and radii in all fields. Species limits M. McGrouther, R.J. McKay. V. Mthombeni, L. Palmer, K. Parkinson, J.E. within the clade are poorly resolved and are currently under study Randall, S.E. Reader, T. Trnski. M. Sabaj, W. Saul, K.-T. Shao, W.F. Smith- by us. For the purposes of comparison with P. occidentalis, we Vaniz, A. Y. Suzumoto, M.J. P. van Oijen and J.T. Williams. Colour photographs divide the clade into three subgroups, of which believe to each we were kindly provided by J. -P. Chen, D.F. Hoese. R.H. Kuiter, J.E. Randall and be monophyletic: P. occidentalis, P. rosae-complex (autapomorphy: J.T. Williams; P. Crabb took the black and white photographs of the two plate-like, median expansion of median ethmoid, pterosphenoid holotypes (Figs 1 and 3). O.A. Crimmen, S. Davidson, J. P. Garcia, A.-M. and basisphenoid into orbital space); and P. rvpws-complex Hine, K. Parkinson. S. Raredon and S.E. Reader radiographed specimens. (autapomorphy: all scales cycloid in adult specimens). Aside from R.D. Mooi and J.T. Williams critically reviewed the manuscript and provided the various autapomorphies listed above, the three taxa are distin- helpful comments. 26 A.C. GILL AND A.J. EDWARDS Pseudochromidae). Revue francaise d'Aquariologie Herpetologie 21( 1 —2): 11 — 18. REFERENCES 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 Ahlstrom, E.H., Butler, J.L. & Sumida, B.Y. 1976. Pelagic stromateoid fishes Fisheries and Marine Resources Research Bulletin 41: 195^418. (Pisces. Perciformes) of the eastern Pacific: kinds, distributions, and early life Kuiter, R.H. 1998. Photo Guide to Fishes of the Maldives. 257 p. Atoll Editions. histories and observations of five of these from the northwest Atlantic. Bulletin of Apollo Bay. Marine Science 26(3): 285-^102. Kulbicki, M. & Williams, J.T. 1997. Checklist of the shorefishes of Ouvea Atoll. New Allen, G.R. & Steene, R.C. 1979. The fishes of Christmas Island, Indian Ocean. Caledonia. Atoll Research Bulletin 444: 1—26. Australian National Parks and Wildlife Service, Special Publication 2: 1-81. Leviton, A.E., Gibbs, R.H., Jr., Heal, E. & Dawson, C.E. 1985. Standards in her- & 1988. Fishes of Christmas Island, Indian Ocean. 197 p. Christmas Island petology and ichthyology: part 1. Standard symbolic codes for institutional resource Natural History Association. collections in herpetology and ichthyology. Copeia 1985(3): 802-832. & Smith-Vaniz, W.F. 1994. Fishes of theCocos (Keeling) Islands. Atoll Research Paxton, J.R., Hoese, D.F., Allen, G.R. & Hanley, J.F. 1989. Zoological Catalogue of Bulletin 412: 1-21. Australia. Volume 7. Pisces. Petromyzontidae to Carangidae. 665 p. Australian Bleeker, P. 1858. Bijdrage tot de kennis der vischfauna van den Goram Archipel. Government Publishing Service. Canberra. Naturrkundig Tijdschrifl Nederlandsch Indie 15: 197-218. Randall, J.E., Allen, G.R. & Steene, R.C. 1990. Fishes of the Great Barrier Reefand 1875. Sur la famille des Pseudochromoi'des et revision de ses especes Coral Sea. 507 p. Crawford House Press. Bathurst. insulindiniennes. Verhandelingen der Koninklijke Akademie van Wetenschdppen, , & . 1997. Fishes of the Great Barrier Reefand Coral Sea. Revised and Amsterdam 15: 1-32, pis 1-3. expanded edition. 557 p. Crawford House Press. Bathurst. Gill, A.C. 2000. Pseudochromidae, pp 2557-2577. In: Carpenter, K.E. & Niem. V.H. & Anderson, R.C. 1993. Annotated checklist of the epipelagic and shore fishes (eds) FAO Species Identification Guide for Fisheries Purposes. The Living Marine of the Maldive Islands. Ichthyological Bulletin of the J.L.B. Smith Institute of Resources of the Western Central Pacific. Vol. 4. FAO. Rome. Ichthyology 59: 1-+7. pis 1-8. & Edwards, A.J. 1999. Monophyly, interrelationships and description of three Regan, C.T. 1 902. On the fishes from the Maldive Islands. The Fauna and Geography new genera in the dottyback fish subfamily Pseudoplesiopinae (Teleostei: Perciformes: of the Maldive and Laccadive Archipelagos 1(3): 272-281. Pseudochromidae). Records of the Australian Museum 52( 1): 141-160. Springer, V.G. 1982. Pacific Plate biogeography, with special reference to shorefishes. , Mooi. R.D. & Hutchins, J.B. 2000. Description of a new subgenus and species Smithsonian Contributions to Zoology 367: 1-182. of the fish genus Congrogadus Gunther from Western Australia (Perciformes: Taylor, W.R. & Van Dyke, G.C. 1 985. Revised procedures for staining and clearing Pseudochromidae). Records of the Western Australian Museum 20( 1 ): 69-79. small fishes and other vertebrates for bone and cartilage study. Cybium 9(2): 107-1 19. & Randall, J.E. 1994. Chlidichthyscacatitoides. a new species ofpseudoplesiopine Winterbottom, R. 1986. Revision and vicariance biogeography of the subfamily dottyback from Oman, with a diagnosis of the genus Chlidichthys Smith, and new Congrogadinae (Pisces: Perciformes: Pseudochromidae). Indo-Pacific Fishes 9: 1- record of Pseudochromis punctattts Kotthaus from Oman (Teleostei: Perciformes: 34. pi. I. [dated 1985. but actually published 1986] ^X(j£fZ£^. l^ Bull. not. 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 ofHong Kong. Cape d'Aguilar, Shek O, Hong Kong Special Administrative Region, China. Synopsis. The holotypeofthedelphinid 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 I 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 specimen identified as 5. 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 The species Sousa chinensis ( formerly Delphin its sinensis) (Osbeck, deduced by the presence of fresh net abrasions on the rostrum and 1765) was designated after observations were made of a population neck area. This specimen has been chosen for designation as the of delphinids within the Pearl River Estuary, Guangdong Province, neotype of S. chinensis because: (a), it was obtained from the area Southern China, i.e. close to Hong Kong. It was over one hundred from which the species was originally identified; (b), it was neither years after the species was named, however, that the first skeleton significantly different from the description of the extant holotype was described from an almost entire individual collected from nor other individuals measured from the Hong Kong population of Xiamen (formerly Amoy), Fujian Province, approximately 7()()km S. chinensis: (c), its death was perceived to be accidental rather than northeast of Hong Kong (Flower, 1870). This specimen and, thus, a consequence of any ailment or abnormality and (d). its remains the holotype of the species was held in the collections of the Royal were intact. 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 MATERIALS AND METHODS the common species, except in their white colour'. The holotype was physically mature and although the external appearance of this A modification of the criteria described by Perrin ( 1975) was used to particular individual was not described, the typical coloration of measure the skeleton obtained from Hong Kong and herein desig- members of the Xiamen population was noted to be 'milky white, nated as the neotype. The tympanoperiotic bones were measured in with pinkish fins and black eyes' (Flower, 1870). The individuals accordance with Kasuya ( 1973). Other skeletal remains of the same that remain in the Pearl River and Jiulong Jiang River (Xiamen) species were also examined and measured to obtain a better range of Estuaries today, are similar in external appearances (Porter, 1998). values for the species in this component of its range. All measure- Four other species are included in the genus Sousa (Table 1), ments were made with EPI precision calipers and recorded to the although there has been debate as to where the taxonomic divisions nearest millimetre or, in the case of smaller bones, 0.1 mm. Each within it, if any, actually occur (Pilleri&Gihr, 1972: Mitchell, 1975; measurement was taken several times until a confident value was Zhou era/., 1980; Wang & Sun, 1982; Ross etal, 1994). The most established. An inventory of the measurements recorded for the recent classification of the genus as comprising one species was neotype is included as Appendix I. based upon a morphological review (Ross et al., 1996) and a The neotype and other Hong Kong specimens were aged from molecular investigation of the genus concurred with this definition three teeth, selected from the upper left jaw, which were sliced to a (Cockcroft etal., 1997). Both of these publications, however, noted thickness of HOum. The sections were etched for one hour by that the limited availability of data from parts of the species' range, emersion in 5% formic acid and the dentinal growth layer groups particularly Asia, did not allow definitive segregation at a sub- then counted under a light microscope. One pair of dark and light specific level and, as such, the taxonomy of the genus is not stained layers is equivalent to one dentinal growth layer group resolved. which is assumed to be gained annually (Kasuya, 1976). There have been many strandings of Sousa chinensis in Hong The equivalent holotype measurements of Sousa chinensis Kong waters, which comprise the eastern portion of the Pearl River (Flower, 1870) were transformed into metric units and compared Estuary, since 1990 (Porter etal., 1998) although the cause of death with corresponding values for the neotype. The original drawings of in many cases could not be determined due to either the decomposi- Flower's (1870) and photographs of the neotype were also com- tion of, or damage to, the remains. In May 1996, however, a fresh pared. The skeletal characters of both the holotype and neotype were © The Natural History Museum. 2002 28 L.J. PORTER 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 29 Table 1 Five species classification of the genus Sousa according to morphological differences and geographical distribution (Ross ci al., RE-DESCRIPTION OF SOUSA CHINENSIS 1994). (OSBECK, 1765) Nominal species Geographical range Neotype: ZD 1999.360, sub-adult male. N 022° 20' 00; E 113° 55' S. chinensis (Osbeck. 1765) China to Australia 30, The Brothers Islands, Hong Kong's territorial waters, collected 5. plumbea (Cuvier, 1829) South Africa to India by staff of the Swire Institute of Marine Science, The University of 5. lentigenosa (Owen, 1866) India to Southeast Asia Hong Kong. Donated to the Natural History Museum, London, (Kukenthal, 1892) West Africa S. teuzii under CITES permit APO/EL 3227/99. 5. borneensis (Lydekker. 1901) Rare form in Borneo, overlapping The gently-tapering rostrum of this specimen comprised 60.7% of with "chinensis' the total skull length and was positioned low near the base of the then compared with the range of measurements obtained from other cranium (Fig. la). The upper and lower tooth rows comprised 89% dead adult specimens of 5. chinensis stranded on Hong Kong beaches. 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 L.J. PORTER teeth in each row (Fig. lb & c). The teeth were conical and pointed which each nasal passage was deep-set and elliptical. The antorbital with little wear. Five dentinal layers were counted from sectioned notch was not developed. The two occipital condyle margins were teeth and the pulp cavity was open. The premaxillaries were in sharply defined and the anterior margin pointed. The channel between contact for almost the entirety of the rostrum length, with only a the condyles was long and triangular. The foramen magnum was small section of the vomer exposed. Storage of the specimen in a dry ovoid with a slightly extended dorsal margin (Fig. 2a). Both the environment, however, has subsequently caused the premaxillaries temporal and post-temporal fossae were ovoid with sharply-defined to gape (Fig. Id). The premaxillaries were roughened dorsally margins. The temporal fossae protruded where they met which, in where they met the less dense maxillaries, at the base of the nasal ventral view, gave the cranium a relatively broad appearance. The aperture. The posterior extremity of the rounded, right, premaxillary orbits were rounded and the lacrimals, positioned at right angles to extended to the anterior of the nasal aperture, whereas the pointed the orbit, were blunt with prominent dorsal ridges. The cranial left premaxillary was truncated at the mid-margin of the opening. articulating surface of the atlas was broad with short, transverse The premaxillaries, therefore, encircled two thirds of the aperture, in 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 31 spine (Fig. 2b). The axis was not fused to the atlas and had short mesosternal element which equalled two thirds of the manubrium. processes but a well-developed neural spine (Fig. 2c). The radius and ulna were longer than the humerus, although the There were seven cervical, 12 thoracic, 23 lumbar and 18 caudal latter was wider and more robust. The transverse breadth of the vertebrae. No vertebrae were fused (Fig. 3a). The first vertical carpals was short and the pectoral fin elongated (Fig. 4b). The perforating foramen appeared on the 25th vertebra which was also scapula had a sharply pointed coracovertebral angle, a broad the first to have greatly reduced metapophyses and the last with metacromion process and a smaller, pointed, coracoid process (Fig. distinct transverse and neural processes. There were 12 vertebral 4c). ribs on both the left and right sides, six of which on each side were The bullae of the tympanoperiotic bones were rounded and double-headed (Fig. 3b). There were five sternal ribs on each side bulbous and the dorsal plane of the periotic convex; the fundus of the (Fig. 3c). internal auditory meatus and the aquaductus fallopii were ovoid; the The sternum was notched with a perforation on the left side and aquaductus cochleae had a distinct ridge; the join between the two lateral processes on each side (Fig. 4a; note that the right side of cochlear portion and the rest of the periotic was rounded and the sternum was damaged during preparation). There was one smooth. Fig. 4 The a, sternum and b, pectoral fin skeletal elements of the neotype of Sousa chinensis 32 L.J. PORTER HOLOTYPE -^.^ttry^fcCfcfcSfe&wa****.., *?&m0#^§eff#*M&am&*#*' (a) occipital condyles NEOTYPE (c) temporal tossae NEOTYPE Fig. 5 A copy of the original illustrations of Flower's ( 1 870) 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 33 COLORATION. Mainly pink with light grey around the head and Table 2 Sousa chinensis principal skeletal dimensions and vertebral darker pigmentation along the length of the back, including the counts of the Hong Kong neotype and the holotype from Fujian Province (Flower, 1870). leading and trailing edge of the dorsal fin and the ventral fringe of the fluke. There was dark, fragmented pigmentation on the melon, Measurement Neotype Holotype 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 1 Condylobasal length 481 526 2 Length of rostrum. 292 325 pectoral fins were still grey and had only just begun to lighten. 3 Width of rostrum 110.1 119 External appearance. The body was slender but with a promi- 5 Width of rostrum at midlength 44.8 47 16 Greatest parietal width 140.9 163 nent melon. Neither the 'robustness' nor prominent peduncle, typical 32 Length upper tooth row to tip rostrum 260 284 of other, adult individuals observed and examined from Hong Kong 33(UL) Number of teeth (UL) 35 33 present (Porter. 1998). dorsal fin slightly waters were The was 34(UR) Number of teeth (UR) 35 32 falcate and the flukes were small with a distinct notch. The pectoral 35(LL) Number of teeth (LL) 31 32 fins were broad in the middle but tapered both distal ly and at the 36(LR) Number of teeth (LR) 32 31 base. The 'hump-back', reportedly typical of populations west of 37 Length of lower tooth row. 259 279 38 Greatest length of left ramus. 407 457 Indonesia, has not been recorded from any individual in the South 39 Greatest height of left ramus. 82.5 91 China Sea (Reeves & Leatherwood, 1994). 40a Length of symphysis 113.3 140 48a Number of cervical vertebrae 7 7 COMMENTS. During the post-mortem it was noted that the indi- 48 Number of thoracic vertebrae 12 12 vidual had deep scarring around the pectoral fins and torso. This 49 Number of lumbar vertebrae 12 10 healed injury was consistent with abrasion from either a rope or a 50 Number of caudal vertebrae 18 22 similar object. It cannot be determined if this injury caused any 51 Total number of vertebrae 49 51 major deformity to the pectoral fin bones although some pathology 78 Number of vertebral ribs (left) 12 12 of them was observed. These injuries were distinct from those 79 Number of vertebral ribs ( right 12 12 96 Number of chevron bones 14 associated with the animal's death. 8 105 Greatest length of coracoid process. 21.5 37 Etymology. The species name is derived from its geographical 106 Greatest width of coracoid process. 11.6 29 107 Greatest w idth of metacromion process. 54.5 location, i.e. both sinensis and chinensis referring to 'China'. The 30 108 Greatest length of humerus. 63.9 73 meaning of the current genus name Sousa has become obscure. 109 Greatest width of humerus distally. 39.5 57 Although the taxonomy of the genus Sousa has yet to be resolved, in 110 Greatest length of radius. 74 79 accordance with the International Code of Zoological Nomencla- in Greatest width of radius distally. 38.2 47 ture, this neotype must bear the name of the holotype. 112 Greatest length of ulna. 65.9 67 COMPARATIVE ANALYSIS OF SPECIMENS HOLOTYPE The age of nine Hong Kong specimens of Sousa chinensis used for comparative skeletal purposes in this study, ranged between 4 and coracoid process 14 growth layer groups, the former number having been established metacromion process 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. 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 Fig. 6 A copy of the original illustrations of Flower's ( 1 870) holotype robust with larger coracoid and metacromion processes (Fig. 6). and the corresponding views of the neotype demonstrating differences The measurements obtained from the holotype, the neotype and in the form of the coracoid and metacromion processes of the scapula. 34 L.J. PORTER 550 500 450 400 350 g Neotype 300 Holotype 250 200 150 100 50 Condylobasal length Ramus length Rostrum length Upper tooth row length Symphysis length 320 270 220 > Neotype 170 Holotype - 3 eso 120 70 20 Lower tooth row Parietal width Mandibular fossa Rostrum width Ramus height Rostrum width at length length midlength Fig. 7 Range of length and width measurements for the skulls of Sousa chinensis obtained from Hong Kong and the holotype demonstrating the a. greatest and b, least variation between individuals. REDESCRIPTION OF THE CHINESE WHITE DOLPHIN 35 Table 3 The range (and average number) of teeth counted from Hong vertebrate collection of the Natural History Museum, London Kong Sousa chinensis (Accession No. ZD 1999.360). Left Right Acknowledgements. The study of Hong Kong stranded marine mam- Upper 32-36 (33.6) 32-35 (33.6) mals was funded by the Agriculture, Fisheries and Conservation Department Lower 28-34(31.5) 27-33(31.1) (AFCD) of the Government of the Hong Kong Special Administrative Region. A special acknowledgement is made to pathologist Dr. L. Simms ( AFCD) for his expertise and use of post mortem facilities. Mr. K.C. Choi and the other eight Hong Kong specimens lie within a discrete range. the staff of the licensing division of AFCD are gratefully acknowledged for The greatest variation is seen between length measurements of the their assistance in processing the CITES export permit. Many thanks also to skull, i.e. condylobasal (450-526 mm), rostral (257-325 mm), Dr. G. Ross, Environment Australia, who freely gave of his expertise and (205.5-284 upper tooth row mm), ramus (397^57 mm) and sym- greatly assisted in the taxonomic procedures and interpretations presented in physis (82.5-258 mm). The physical maturity of the holotype is this paper. The author also wishes to thank Prof. Brian Morton. Director. The reflected in its location at the top of each of these length values (Fig. Swire Institute of Marine Science, The University of Hong Kong, for his 7a). Least variation is seen between the width dimensions of the support and encouragement for this and other studies of Sousa chinensis skulls, i.e., parietal (137.4-171 mm), rostral (99.2-119 mm), mid- conducted at the institute. rostral (47.8-62 mm), mandibular fossae (1 19.4—130.9 mm) and the lower tooth row length (247-279 mm) (Fig. 7b). REFERENCES DISCUSSION Cockcroft, V.G., Leatherwood, S., Goodwin J. & Porter, L. 1997. The phylogeny of humpback dolphins genus Sousa: insights through mlDNA analyses. Working paper The ages of nine stranded specimens of Sousa chinensis examined SC/49/SM25 14pp. In: Proceedings of the 49"' International Whaling Commission Meeting, Scientific Committee Annual Meeting. Bournemouth. United Kingdom. from Hong Kong waters were determined to lie between 4 and 14 1997. International Whaling Commission. Cambridge. years. It is clear from the original description that the holotype was Flower, W. 1 870. Description of the skeleton of the Chinese white dolphin (Delphinus physically mature whereas no specimen obtained from Hong Kong sinensis Osbeck ). Transactions of the Zoological Society of London 7: 151-160. kasu.\a. T. 1973. toothed whales was. This is reflected in the comparison of skeletal characters where Systematic consideration of recent based on the morphology of tympanoperiotic bone. Scientific Report of the Whales Research the holotype is consistently the largest in terms of skull length Institute, Tokyo2S: 1-103. measurements. The greater ossification of the scapula also is indica- 1976. Reconsideration of life history parameters of the spotted and striped tive of the holotype's greater age. The measurements obtained for dolphins based on cemental layers. Scientific Report of the Whales Research Institute the neotype appear to be typical for Hong Kong waters, although the 28: 73-106. I, K ii ki hi ha W. 1 892. Solatia ieus:ii a. sp.. einplanzenfressender Delphin aus Kamerun. sample size is currently limited. Those measurements available for Zoologischen. Jahrbuchen 6: 442—t4d the holotype are, however, within the ranges obtained for Hong Mitchell. E. 1975. Report of the meeting on smaller cetaceans. Montreal. April l-l 1. Kong specimens. The difference between vertebral counts of the 1974. In Review of Biology and fisheries lor Smaller Cetaceans. Journal of the holotype and neotype is small and such variation in other delphinids Fisheries Research Board oj Camilla 24: 2505-2513. Osbeck, P. 1765. Reise nach Ostindien und China. (Translated from the original appears typical (Perrin, 1975). It is not known if the holotype was Swedish by J.Ci. Georgi and J.C. Koppe. Rostock. [In German] typical of the population present in Xiamen waters at the time it was Perrin, W.F. 1975. Variation of Spotted and Spinner porpoise (genus Stenella) in the collected (1870) and, indeed, as both Hong Kong and Xiamen Eastern Tropical Pacific and Hawaii. Bulletin of the Scripps Institute of Oceanogra- coastal areas have been extensively developed, any adaptive changes phy University of California Press. California, I'illei i. 1972. Contribution knowledge that both populations of Sousa chinensis may have undergone since G, & Gihr, M. to the of the cetaceans of Pakistan with particular reference to the genera Neomeris, Sousa. Delphinus and the 17"' century cannot be gauged. In addition, the latest review of Tursiops and description of a new Chinese porpoise (Neomeris asiaeorientalis). (Ross et observes that is the genus ai. 1996) osteological form Investions ofCetacea 4:107-162. similar across its global range, with any slight differences being Porter, LJ. 1 998. The taxonomy, ecology and conservation of Sousa chinensis (Osbeck, environmental rather than genetic. 1765) (Cetacea: Delphinidae) in Hong Kong waters. PhD thesis. The University of Hong Kong. The uncertainty that prevails over the taxonomy of the genus Reeves, R.R. & Leatherwood, S. 1 994. Dolphins. Porpoises and Whales: 1994-1998 Sousa can only be dispelled through the continued investigation and Action Plan lor the Conservation oj Cetaceans. IUCN/Species Survival Commis- comparison of specimens from throughout its range. As the holotype sion/Cetacean Specialist Group, Cambridge. has been destroyed, the designation of a neotype is required so that Ross, G.J.B., Heinsohn, G.E. & Cockcroft, V.G. 1994. Humpback dolphins Sousa chinensis (Osbeck. 1765). Sousa plumbea (G. Cuvier. 1829) and Sousa leuszii such comparative studies can be conducted in relation to a defined (Kukenthal. 1892). In: Handbook of Marine Mammals, Volume 5: 23-42 Ridgeway. type and, thus, the taxonomy and life history traits of the genus S.H. & Harrison, R. (Eds). Academic Press. London. resolved. In addition, the herein designated neotype was obtained Ross G.J.B., Heinsohn, G.E., Cockcroft, V.G., Parsons, E.C.M., Porter, L.J. & from the area in which the species was originally identified (Osbeck, Preen, A. 1996. Review of the taxonomic status of the humpbacked dolphins, genus Sousa. Working paper UNEP/SEA95/WPI9. In: Perrin, W.F., Dolar, M.L. & 1765) and the same geographic region that the holotype was col- 25pp. Alava, M.N.R. (eds.) Report of the Workshop on the Biology and Conservation of lected from (Flower, 1870). The meristic data of the neotype are Small Cetaceans and Dugongs of Southeast Asia, Dumaguete 1995. UNEP (W)/EAS neither significantly different from what is known of the original WG.I/2. Bangkok: United Nations Environment Programme. Bangkok. description of a Sousa chinensis skeleton (Flower 1 870) nor the Wang, P. & Sun, J. 1982. Studies on the Zhonghua white dolphin. Sousa chinensis. sample of other stranded specimens measured from Hong Kong from the South China Sea. Transactions of the Liaoning Zoological Society 3: 67-74 (In Chinese). waters. The specimen described herein, therefore, fulfils the criteria Zhou, K, Li, Y, Qian, W. & Yang, G. 1980. Notes on three species of dolphins from as laid the International for out by Code of Zoological Nomenclature the South China Sea and Jiulongjiang River. Oceanologica et Limnologia Sinica 11: designation as a neotype and, as such, has now been lodged in the 306-313 [In Chinese with an English Summary]. 2 36 L.J. PORTER articulating surface to distal end of process 24.5 APPENDIX 1 Number of cervical vertebrae with incomplete neural arches 2 Cervical vertebra on which left ventrolateral process reaches Skeletal measurements of the neotype. greatest development 5 lh Measurement Neotype Vertebra on which first vertical perforating foramen appears 25 lh First vertebra with greatly reduced metapophyses 25 th Condylobasal length 481 Last vertebra with distinct transverse processes 25 lh Length of rostrum 292 Last vertebra with distinct neural process 25 th Width of rostrum 110.1 First vertebra with unfused epiphysis 25"' Width of rostrum at 60 mm anterior 77.1 Last vertebra with unfused epiphysis 25 th Width of rostrum at midlength 44.8 rd First caudal vertebra with vertical neural spine 3 Premaxilleries width, at rostral midlength 28.9 Length of neural spine of first thoracic - from anterodorsal Width of rostrum at 3/4 length, measured from posterior end 34.1 margin of neural canal to tip of spine 36.5 Distance from tip of rostrum to external nares 323.0 Length of neural spine of second thoracic vertebra 46.5 Distance from tip of rostrum to internal nares 300.0 Length of neural spine of tenth thoracic vertebra 65 Greatest preorbital width 185 Length of neural spine of last thoracic vertebra 69.4 Greatest postorbital width 208 Height of first thoracic vertebra - from internal anterodorsal Greatest width of external nares 50.5 margin of neural canal to bottom of anterior face of body 55.0 Zygomatic width 203.0 Greatest width of first thoracic vertebra - across lateral processes 89.2 Greatest width premaxilleries 82.0 Height of the first lumbar vertebra 57.5 Greatest parietal width 140.9 Greatest width of the first lumbar vertebra 1 64.9 Least width between temporal fossa 137.0 Length of 23d centrum, exclusive of epiphyses, along vertical Greatest length of left temporal fossa 86.6 midline 40.1 Greatest width of temporal fossa (at right angles to length) 79.9 Number of vertebral ribs - left 1 Major diameter of left temporal fossa 51.1 Number of vertebral ribs - right 1 Minor diameter of left temporal fossa 45.6 Number of two-headed ribs - left 6 Length of orbit 48.6 Number of two-headed ribs - right 6 Length of antero-lateral face of lacrimal measured to exclude Number of floating ribs - left 1 the flare on the rim 29.3 Number of floating ribs - right 2 Greatest width internal nares 60.5 Number of sternal ribs - left Greatest length pterygoid 10.30 5 Number of sternal ribs - right 5 Length upper tooth row to tip rostrum 260.0 Greatest length of first left vertebral rib 123.6 Number of teeth - upper left 35 Width of first left vertebral Number of teeth - upper right 35 rib at apex of proximal curvature 19.7 Greatest length of longest left vertebral rib Number of teeth - lower left 31 141.4 Number of teeth - lower right 32 Greatest width of manubrium 88.8 Length of lower tooth row to tip mandible 259.0 Length of manubrium along midline 84 Greatest length of left ramus 407.0 Depth of anterior notch of manubrium 1 Greatest height of left ramus 82.5 Length of foramen in manubrium 5 Length of left mandibular fossa 119.4 Number of chevron bones 7 th Length of symphysis 113.3 Vertebra bearing first chevron bone 24 s1 Greatest width of left thyrohyal proximally 18.4 Vertebra bearing last chevron bone 31 Greatest length of left thyrohyal 66.2 Greatest length of first half of largest chevron bone 44.3 Greatest width of left stylohyal 13.4 Greatest length of first half of last chevron bone 22.9 Greatest length of left stylohyal 79.4 Height of scapula - from posterior margin of glenoid fossa Number of thoracic vertebrae, defined as equal to number of to coracovertebral angle 125.1 ribs on side with greatest number Length of scapula - from posterior margin of glenoid fossa Number of lumbar vertebrae to glenovertebral angle 110.3 Number of caudal vertebrae Greatest length of coracoid process - from anterior margin Total number of vertebrae 49 of glenoid fossa 21.5 Number of fused cervical vertebrae none Greatest width of coracoid process 11.6 Greatest width of articulating surface of atlas 101.9 Greatest width of metacromion process - from apex of ventral Height of atlas - from internal anterodorsal margin of neural curvature to vertebral apex 54.5 canal to bottom of anterior face of body 60.9 Greatest length of humerus, measured on ventral side of flipper 63.9 Length of lateral process of atlas - from margin of anterior Greatest width of humerus distally 39.5 articulating surface to farthest point at end of process 28.3 Greatest length of radius 74 Greatest length of neural spine of atlas 41.1 Greatest width of radius distally 38.2 Height of dorsal ridge of atlas in anterior view 22.8 Greatest length of ulna 65.9 Length of right dorsolateral spine of atlas 25.1 Transverse breadth of proximal row of carpals 73 Length of lateral process of axis - from margin of posterior Greatest length of left pelvic rudiment 108.7 REDESCRIPTION OF THE CHINESE WHITE DOLPHIN 37 Measurement of the tympanoperiotic bones of the neotype. Measurement Neotype Left Right Length of tympanic bulla, from anterior tip to posterior end of outer posterior prominence 33.7 34.3 Anterior tip to posterior end of inner prominence 32.8 33.2 Postero- ventral tip of outer posterior prominence to tip of sigmoid process 21.6 24.6 Postero-ventral tip of outer posterior prominence to tip of conical process 17.9 18.0 Width of tympanic bulla, at the level of the sigmoid process 20.7 21.3 Height of tympanic bulla, from tip of sigmoid process to ventral keel 22.6 25.0 Width across inner and outer posterior prominences 18.5 18.3 Greater depth of interprominental notch 5.3 4.7 Width of upper border of sigmoid process - 5.0 Width of the posterior branch of lower tympanic aperture 2.0 1.2 The thicker side between outer and inner posterior prominences 8.6 10.5 Length of periotic, from tip of anterior process to posterior end of posterior process, measured on a straight line parallel 32.0 31.9 with cerebral border Thickness of superior process at the level of upper tympanic aperature 11.8 13.0 Width of periotic across cochlear portion and superior process, at the level of upper tympanic aperture 21.8 22.2 Least distance between the margins of the fundus of the internal auditory meatus and of the aperature of ductus endolymphaticus 2.3 2.4 Least distance between the margins of the fundus of the internal auditory meatus and of the aperture of aquaductus cochleae 4.9 4.2 Length of articular facet of the posterior process of periotic for the posterior process of tympanic bulla 18.3 15.9 Antero-posterior diameter of cochlear portion 21.3 19.0 Maximum depth of inner part of involvucrum 9.3 9.1 )C^C^^^^'^) Bull. nar. Hist. Mus. Land. (Zool.) 68( 1 ): 39-50 Issued 27 June 2002 A new genus of groundwater Ameiridae (Copepoda, Harpacticoida) from boreholes in Western Australia and the artificial status of Stygonitocrella Petkovski, 1976 WONCHOEL LEE Department of Life Sciences, College of Natural Sciences. Hanyang University, Seoul 133-791, Korea: e-mail: wlee @ hanyang.ac. kr RONY HUYS* Department of Zoology, The Natural History Museum. Cromwell Road. London SW7 5BD, U.K.: e-mail: [email protected] SYNOPSIS. Examination of the copepod fauna inhabiting 50m deep production bores on Barrow Island (northwestern Australia), resulted in the discovery of an unusual ameirid which cannot be placed in any extant genus. Both sexes are characterized by a unisetose antennary exopod and extreme reduction in the swimming legs (particularly the endopods) and P5. Males lack a defined P6 closing off the single genital aperture and have an extraordinarily large spermatophore. Females similarly display a highly reduced genital field. The new species shows superficial similarities to both Psammonitocrella Rouch and Stygonitocrella Reid, Hunt & Stanley, however the combined presence of a sexually dimorphic inner basal spine on PI. a completely fused genital double-somite, reduced antennary exopod and vestigial P5 excludes it from either genus. Some problems in the current classification of freshwater Ameiridae are highlighted, with particular reference to the genus Stygonitocrella. A new genus Neonitocrella is proposed for Stygonitocrella insularis (Miura, 1962). INTRODUCTION found yet. The subterranean fauna contains Thermosbaenacea, Syncarida and a diverse amphipod community including bogidiellids and melitids. Our samples were taken in a long abandoned water The arid to semi-arid north-west of Western Australia has a rich production bore (47 m depth) and an abandoned anode bore (55 m stygofauna including the only vertebrate troglobites known to occur depth) that was used in the electrolytic protection of the oil wells. in Australasia, the Blind Gudgeon, Milyeringa Veritas Whitley, and The latter would have gassed chlorine and had a pH of <2 when in the Cave Eel. Ophisternon candidum (Mees), and two, supposedly service (Humphreys, pers. comm). sympatric, congeneric shrimps, Stygiocaris lancifera Holthuis and S. stylifera Holthuis with tethyan affinities (Humphreys, 1993). Recently, the freshwater copepods of the Cape Range karst area have been the subject of intensive study, resulting in the discovery MATERIALS AND METHODS and description of several cyclopoids belonging to the genera Metacyclops Kiefer, Mesocyclops Sars, Microcyclops Claus, Specimens were dissected in lactic acid and the dissected parts were Apocyclops Lindberg, Diacyclops Kiefer and Halicyclops Norman mounted on slides in lactophenol mounting medium. Preparations (Pesce et al., \996a-b\ Pesce & De Laurentiis, 1996; De Laurentiis were sealed with Glyceel or transparent nail varnish. All drawings etai, 1999). have been prepared using a camera lucida on a Zeiss Axioskop Here we report on the discovery of a remarkable harpacticoid in differential interference contrast microscope. abandoned production bore holes on Barrow Island off the Cape Males and females of Inermipes humphreysi gen et sp. nov. were Range coast. Barrow Island has a typical island Ghyben-Herzberg examined with a Philips XL30 scanning electron microscope. Speci- system with a freshwater lens overlying salt water. The hydrology of mens were prepared by dehydration through graded acetone, critical the superficial karst is little known despite being a production oil point dried, mounted on stubs and sputter-coated with gold or field since the early 1960s and being the target of 'produced water' palladium. disposal until recently. It lies on the North West Shelf of Western The descriptive terminology is adopted from Huys et al. (1996). Australia and up to about 8-10,000 years ago it would have been Abbreviations used in the text are: ae, aesthetasc; P1-P6, first to part of the mainland (and throughout most of the last few million sixth thoracopod; exp(enp)-l(2. 3) to denote the proximal (middle, years) (Humphreys, 2000). The entire island is likely to be an distal) segment of a ramus. Type material is deposited in the anchialine system but to date an entry point for divers has not been collections of the Western Australian Museum, Perth (WAM) and The Natural History Museum, London (BMNH). * Author for correspondence © The Natural History Museum, 2002 9 40 W. LEE AND R. HUYS SYSTEMATICS 1999.1 106), BES 792, depth 50-55m, 1 December 1992 and 999, 88 in 70% alcohol (BMNH 1999.1107-1124), BES 793, 1 Decem- ber 1992. Paratypes deposited in WAM are (a) 1 8 dissected on 7 slides, and 499, 6 88 in 70% alcohol, BES 792, depth 50-55m, 1 Family AMEIRIDAE Monard, 1927 December 1992, (b) 1 8 in 70% alcohol, BES 798, depth 45-50m, Genus INERMIPES gen. nov. 2 December 1992, and (c) 5 99, 15 88 in 70% alcohol, BES 810, depth 50-55m, 2 December 1992 (WAM C244 15-244 17). All Diagnosis. Ameiridae. Body elongate, cylindrical, and vermi- specimens are from the type locality and were collected by Dr W.F. form without distinct surface ornamentation except for ventral Humphreys using a plankton net with a 125 m mesh and of a size spinule patterns on abdomen. Cephalothorax and other somites with suitable for the borehole (Pesce et al., 1996a). smooth posterior margin. Genital and first abdominal somites com- pletely fused forming double-somite; original segmentation not Description discernible. Genital field with small copulatory pore located in Female. Total body length 540-667um (n=10; mean = 592 urn: median depression. Anal operculum well developed. measured from anterior margin of rostrum to posterior margin of Sexual dimorphism in antennule, PI inner basal spine, P5, P6, caudal rami). Largest width measured at posterior margin of genital genital segmentation and abdominal spinulation. double-somite: 106pm. Genital double-somite swollen. No distinct Rostrum fused to cephalothorax, not defined at base. Antennule demarcation between urosome and prosome (Fig. IB). 8-segmented in 9, 10-segmented and subchirocer in 3; segment 1 Cephalothorax with smooth posterior margin; pleural areas small without seta; aesthetascs on segments 4 and 8(9) or on 5 and 1 ( 8 ); and rounded; sensillae and few pores present as illustrated in Fig. apical acrothek consisting of minute aesthetasc and 2 setae. Antenna 1A-B. Rostrum not defined at base (Fig. IB), with pair of tiny with separate basis and endopod; exopod minute, with 1 seta. sensillae near apex. Mandible with 2-segmented palp, comprising unarmed basis and Pedigerous somites completely smooth. All prosomites without endopod with 4 setae. Maxillule with strongly developed arthrite, defined hyaline frills; hind margin smooth; separated by large with 2 naked seta on anterior surface and 8 spines/setae around distal membranous areas. Body not markedly constricted between indi- margin; coxa with 2 setae and 1 spine; palp cylindrical with 5 setae, vidual somites. P4-bearing somite distinctly narrower than preceding probably representing basis only. Maxilla with trisetose endite on ones. syncoxa; allobasis drawn out in remarkably long claw; endopod a Urosome (Fig. 1A-B) 5-segmented, comprising P5-bearing small bisetose segment. Maxilliped with unarmed syncoxa; basis somite, genital double-somite and 3 free abdominal somites. PS- elongate, with very long endopodal pinnate claw. bearing somite, and genital double-somite without surface PI with 3-segmented rami; basis without outer seta, inner spine ornamentation, except for dorsal pairs of sensillae. Free abdominal sexually dimorphic; exp- 1 elongate, exp-2 without inner seta, exp-3 somites with several rows of minute spinules laterally (Fig. 1 A) and with 4 elements; endopod with formula [1.0.020]. ventrally (Fig. 2C); hyaline frills of urosomites not present. Ovary P2-P4 with 3-segmented exopods and 1 -segmented endopods. large, about half of body length (Fig. 1A-B). Bases without outer seta. Exp- 1 markedly elongate; exp-2 with very Genital double-somite (Figs 1A-B, 2C) wider than long; com- strong inner seta; apical setae of exp-3 very long. Endopods minute, pletely fused, with weakly pronounced constriction bilaterally with very long apical seta(e). Armature formula: (possibly indicating original segmentation). Genital field (Fig. 2C) with small copulatory pore (arrowed in Fig. 2C) located in median Exopod Endopod depression; gonopores fused medially forming single genital slit covered on both sides by opercula derived from sixth legs; no P2 0.1.022 010 distinct armature discernible; seminal receptacles fused forming P3 0.1.022 020 P4 0.1.122 010 large median sac. Anal somite (Fig. 2C-D) with well developed spinulose opercu- lum flanked by spinular rows; with pair of dorsal sensillae anterior P5 rudimentary with baseoendopod fused to somite and repres- to operculum. Caudal rami (Fig. 2C-D) longer than wide; each ented only by outer basal seta. Exopod a small segment with 1 seta ramus with 7 setae; setae I—III and VI-VII bare, setae IV and V with in $ and 1 multipinnate fused spine plus 1 seta in 8. P6 rudimentary, minute spinules; setae I and II positioned dorsolaterally with seta I forming unarmed median operculum in 9; asymmetrical in 3 (with being much shorter than seta II; seta III positioned laterally; setae IV dextral or sinistral configuration), represented by opercular un- and V fused basally, each with predesigned fracture planes at base; armed plate. seta V longest (longer than whole urosomal segments combined); Caudal ramus short, with 7 setae; seta V longest. seta VI small; seta VII tri-articulate at base. Bases of setae typically Spermatophore extraordinarily large, longer than half the body surrounded by few tiny spinules. Inner margin of each ramus with length. dorsolateral concavity. gen. et sp. nov. TYPE AND ONLY SPECIES. Inermipes humphreysi Antennule (Fig. 3A) 8-segmented, slender; with well developed sclerite around base of segment 1; all setae bare. Segment 1 without Etymology. The generic name alludes to the absence of the outer seta; 3 longest; segment 4 with aesthetasc. Armature for- seta on the bases of the swimming legs. Gender: feminine. segment mula: l-[0], 2-[7], 3-[7], 4-[2 + (1 + ae)], 5-[2], 6-[2], 7-[4], 8-[3 + Inermipes humphreysi sp. nov. acrothek]. Apical acrothek consisting of slender aesthetasc fused basally to 2 setae. All segments without surface ornamentation. Type locality. Barrow Island, Western Australia (37°5046"N, Antenna (Fig. 3B) 4-segmented. Coxa small, without ornamenta- 31°31'35"W), stn BES 792, borehole, 50-55m deep, 1 December tion. Basis with spinules along proximal outer margin, and distal 1992. inner margin; abexopodal seta absent. Exopod minute, 1 -segmented, TYPE MATERIAL. Holotype 9 dissected on 9 slides (WAMC244 14). with 1 pinnate apical seta. Endopod 2-segmented; proximal segment Paratypes deposited in NHM are 1 9 dissected on 8 slides (BMNH with 2 rows of spinules along abexopodal margin; distal segment NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER 41 Fig. Inermipes 1 humphreysi gen. et sp. nov. A, habitus 9, lateral; B, habitus 2, dorsal; C, habitus 6\ dorsal. 42 W. LEE AND R. HUYS Fig. 2 Inermipes humphreysi gen. et sp. nov. Female: A, maxillule; B, maxilla; C, urosome, ventral [copulatory pore indicated by arrow]; D, anal somite and caudal rami, dorsal; E, P5. Male: F, P5-bearing and genital somites, ventral [P5 aberrant on left side], ventral; G. P5 [normal]; H. P5 [aberrant]. NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER 43 Male: E, antennule. Fig. 3 Inermipes humphreysi gen. et sp. nov. Female: A, antennule; B, antenna; C, mandible; D, maxilliped. 44 W. LEE AND R. HUYS longer than proximal; lateral armature arising in proximal half, seta; endopodal lobe not expressed. Exopod forming minute seg- consisting of 1 pinnate and 2 bare setae; apical armature consisting ment with 1 bipinnate apical seta. of 5 geniculate setae ( 1 geniculate seta fused basally to 1 pinnate seta Male. Body larger than in 9. Somites bearing P2-P4 wider than and small tube pore; see insert); distal endopod segment with row of in 9. Body length 515-690 um (n=20; mean = 622 um; measured spinules along abexopodal margin and 2 transverse hyaline frills from anterior margin of rostrum to posterior margin of caudal rami). subapically. Largest width measured at P6-bearing somite: 109 urn. Labrum with elaborate spinular ornamentation and pores as in Prosome (Fig. 1C) 4-segmented, comprising cephalothorax and 3 Fig. 5A-B. free pedigerous somites. No distinct demarcation between urosome Mandible (Fig. 3C) with well developed gnathobase bearing and prosome. Cephalothorax with smooth posterior margin; pleural several fine, multicuspidate teeth around distal margin and 1 pinnate areas small and rounded; sensillae and few pores present as illustrated spine at dorsal corner; 1 row of spinules near base of palp. Palp 2- in Fig. 1C. Rostrum not defined at base (Fig. 1C), with pair of tiny segmented; proximal segment without ornamentation; distal segment sensillae near apex. with 4 naked setae arranged in 2 pairs; subapical pair fused basally. Pedigerous somites not covered with spinules. All prosomites Paragnaths (Fig. 5A) strongly developed lobes with medially without defined hyaline frills but separated by large membranous directed hair-like setules, separated by medial lobe covered with areas. Body without marked constrictions between individual dense pattern of short setules. somites; P4-bearing somite not distinctly narrower than others. Maxillule (Figs 2A, 5A-B). Praecoxa without ornamentation; Urosome (Fig. 1C) 6-segmented. comprising P5-bearing somite, arthrite elongate, strongly developed, with 2 naked setae along outer genital somite and 4 abdominal somites. P5-bearing somite and margin, 2 short setae on anterior surface, and 6 spines/setae around genital somite without surface ornamentation, except for pairs of distal margin; 3 distal spines unipinnate and bearing pore (Fig. 5A- dorsal sensillae. Free abdominal somites, including anal somite, B) at base of proximal spinule. Coxa with cylindrical endite bearing with several rows of spinules laterally and ventrally; dorsal hind 2 naked setae and 1 curved, pinnate spine. Palp represented by basis margin smooth without any ornamentation. Hyaline frills of and possibly incorporated rami, forming cylindrical segment with 5 urosomites not present. Spermatophore extraordinarily large, about naked setae around distal margin. half of body length. Maxilla (Fig. 2B) without surface ornamentation on syncoxa; Antennule (Figs 3E, 5C) 10-segmented; haplocer, with medial margin membranous and bearing 1 coxal endite with 1 geniculation between segments 7 and 8. Segment 1 longest, un- pinnate spine and 2 naked setae. Allobasis drawn out into very long, armed. Some elements on segments 4, 6 and 8 (inserts in Fig. 3E; slightly curved, unipinnate claw; allobasal claw with spinules along Fig. 5C) very small. Segment 5 not swollen. Segment 8 with 1 small distal half of inner margin and transverse row of posterior spinules fused spine in median anterior margin. Armature formula: l-[0], 2- halfway the length; accessory armature consisting of 1 pinnate spine [7], 3-[6], 4-[l + 1 small spine], 5-[4 + (1 + ae)], 6-[l], 7-[l], 8-[l + on posterior surface, and 1 tube pore along outer margin. Endopod 1- 1 fused spine], 9-[4], 10-[5 + acrothek]. 1 seta on fifth segment very segmented, and located on anterior surface of allobasis; ornamented small [arrowed in Fig. 5C). Apical acrothek consisting of slender with 2 naked setae. aesthetasc and 2 naked setae. Maxilliped (Fig. 3D) without ornamentation on syncoxa. Basis Inner basal spine of PI modified into robust barbed element (Fig. smooth without ornamentation. Endopod drawn out into very long, 4E). P2-P4 as in 9 unipinnate claw without accessory armature; longer than basis. Fifth pair of legs (Figs 2F-H, 5D) fused to supporting somite. Swimming legs P1-P4 (Fig. 4A-D) with wide intercoxal sclerites Baseoendopod with short setophore bearing outer basal seta; and well developed praecoxae, both without ornamentation. Coxae endopodal lobe not expressed. Exopod with 2 elements; 1 apical and bases with anterior rows of surface spinules as figured. Bases tripinnate spine fused to exopod, and 1 small slender bipinnate seta without outer seta. Exopods 3-segmented, endopods 3-segmented on subapical outer margin (Figs 2G, 5D); abnormal exopod (Fig. (PI), or 1-segmented (P2-P4). 2H) resembling female condition frequently observed on one or PI (Fig. 4A) with large coxa; with 1 row of spinules on anterior either side; only 1 out of 9 males showed normal P5 exopod on both surface. Basis with 1 strong, bipinnate spine and long setules along sides. inner margin; rows of spinules along outer distal margin, and near Sixth pair of legs (Figs 2F, 5D) asymmetrical ; represented on both base of exopod. Exp-1 and -2 with 1 bipinnate slender spine; exp-3 sides by small membranous plate (fused to ventral wall of support- with 1 bipinnate spine, 1 long unipinnate seta and 2 bipinnate ing somite along one side; articulating at base and covering gonopore weakly geniculate setae. Endopod slightly shorter than exopod; enp- along other side; dextral and sinistral configurations present); with- 1 with 1 bipinnate inner seta; enp-2 without seta; enp-3 with 2 out additional ornamentation. geniculate setae apically. All segments of exopod and endopod ornamented with small spinules and setules along outer and inner Etymology. The species is named in honour of Dr William F margin as figured. Humphreys (Western Australian Museum, Perth), who collected the P2-P4 (Fig. 4B-D). Coxae without ornamentation; basis with material. row of spinules along outer margin; additional spinules along inner margin, and on anterior surface of P3-P4; all segments with pattern of spinules as figured; inner and outer margins of exopod and endopod segments with setules or spinules. P2 exp-1 longer than DISCUSSION exp-2 and -3 combined; P3 exp-1 subequal to exp-2 and -3 com- bined; P4 exp-1 shorter than exp-2 and -3 combined. P2-P4 with Inermipes gen nov. is placed in the family Ameiridae on account of strong inner seta on enp-2, outer exopodal spines slender, terminal the morphology of the antennules, mouthparts and the sexual dimor- setae very long. Endopods minute, represented by small segment. phism in the inner basal spine of leg 1. Within the group of Spine and setal formulae as in generic diagnosis. freshwater ameirids it can be readily identified by the unisetose Fifth pair of legs (Fig. 2C, E) fused to supporting somite. antennary exopod, the lack of the outer basal seta on P1-P4 the Baseoendopod represented by small, outer setophore bearing basal extreme reduction in the swimming legs (particularly the endopods) 45 NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER A, PI; B, P2; C, P3; D, P4. Male: E, coxa and basis of PI. Fig. 4 Inermipes humphreysi gen. et sp. nov. Female: 46 W. LEE AND R. HUYS Fig. S Inermipes humphreysi gen. et sp. nov. SEM micrographs. A. oral area showing labrum. paragnath and part of maxillule; B. oral area (different angle); C, antennule 6 [tiny seta on segment 5 arrowed]; D. abdomen 6, showing P5 and P6. ventral. Scale bars: 5 |jm (A-B). 2 pm (C), 20 |am (D). NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER 47 and the P5. Males lack a defined P6 closing off the single genital the wrong genus. For example. Nitocrella petkovskii Pesce, 1980 aperture and have an extraordinarily large spermatophore. Females and Stygonitocrella colchica (Borutzky & Michailova-Neikova, similarly display a highly reduced genital field. The size of the 1970) are very closely related, differing essentially only in the spermatophore, occupying nearly half of the body, is remarkable. expression of the segment boundary between P4 enp-1 and -2, but Except for representatives of the genus Apodopsyllus Kunz are nevertheless placed in different genera. The close zoogeographical (Paramesochridae) where similarly sized spermatophores have been connection between N. petkovskii (NW Iran) and S. colchica (W reported, male harpacticoid copepods produce small spermatophores, Georgia) is noteworthy in this context. typically not exceeding the length of two body somites. Given the /. humphreysi shows superficial similarities to both Psammoni- highly reduced genital apertures of /. humphreysi it is difficult to tocrella and Stygonitocrella, currently the most advanced genera imagine how the spermatophore can be successfully extruded and within the Ameiridae. however the combined presence of a sexually transferred to the female. dimorphic inner basal spine on PI, a completely fused genital double-somite, reduced antennary exopod and vestigial P5 excludes it from either genus. The genus Psammonitocrella was proposed by Taxonomy of freshwater Ameiridae Rouch (1992) to accommodate two interstitial species collected in The primary taxonomic subdivisions in freshwater Ameiridae have the hyporheic zone of an intermittent desert stream in Arizona. Its traditionally been based on swimming leg segmentation (Lang, familial placement has been questioned by Martinez Arbizu & 1965; Petkovski, 1976) and have ignored other, more phylo- Moura (1994) who removed the genus from the Ameiridae and genetically informative characters such as setation patterns and regarded it as the sistergroup of the Parastenocarididae. This course mouthpart morphology (Galassi et al., 1999). This simplistic of action was primarily based on the loss of the outer spine of PI approach has led to: (1) the generally unsatisfactory practice of exp-2 and the absence of a sexually dimorphic inner basal spine on describing new species virtually exclusively on leg characters with- leg 1. The juvenile morphology of the P5 in both sexes and the out consideration of cephalic appendages, genital field morphology presence of separate genital and first abdominal somites in the adult or even female abdominal segmentation; (2) the establishment of female strongly suggest a paedomorphic origin for Psammonitocrella. unnatural genera such as Stygonitocrella, and (3) the blurring of The absence of sexual dimorphism in the inner basal spine of leg 1 generic boundaries. Central to this confusion stands the genus should be re-evaluated in this context. The modification of this spine Nitocrella which has served as a taxonomic repository for freshwa- in male Ameiridae appears not until the final moult. Delaying the ter Ameiridae since its proposal by Chappuis ( 1924). Lang ( 1965) expression of this character beyond the final moult (post-displace- removed all species displaying 3-segmented P2-P4 endopods to a ment) would result in the secondary loss of sexual dimorphism. The new genus Parapseudoleptomesochra and created a second genus, absence of a modified inner basal spine in Psammonitocrella is Pseudoleptomesochrella, to accommodate all Nitocrella species therefore to be regarded as autapomorphic and it is proposed here to characterized by 2-segmented P2-P4 endopods and the presence of remove the genus from its floating status and to reallocate it to the an inner seta on P2-P4 exp-1. Even under its revised taxonomic Ameiridae. concept the genus Nitocrella continued to accumulate a large number Similarities between lnermipes and Psammonitocrella are found of new species which prompted Petkovski (1976) to subdivide the in the presence of only two setation elements on the distal endopod 2- genus even further. He suggested to group only species with segment of leg 1 , the absence of the outer basal seta in P1-P2, and segmented P2-P4 endopods in Nitocrella s. restr. and to reallocate the reduced P4 endopod. It is conceivable that these shared characters all remaining species with alternative endopodal segmentation in are the product of convergence since both genera differ significantly two new genera, Nitocrellopsis (P2-P3 3-segmented. P4 2-seg- in the morphology of the antenna, maxilla, maxilliped and swim- mented) and Stygonitocrella (P2-P3 1 -2-segmented, P4 ming leg ornamentation. 1 -segmented). Petkovski (1976) also recognized three subgroups in The supposedly cosmopolitan genus Stygonitocrella was diag- Nitocrella based on the number of armature elements on P4 exp-3: nosed solely on the basis of the 1 -segmented P4 endopod (Petkovski, the hirta- (3-4 setae/spines), chappuisi- (5 setae/spines) and 1976). Comparison of the swimming leg setal formulae of the 13 vasconica-groups (6 setae/spines). Although these groups have met species currently included strongly indicates the presence of several with general acceptance, their monophyletic status has never been discrete lineages within this genus, each exhibiting a typical arma- challenged. Furthermore, since Petkovski ( 1976) did not designate a ture pattern (Table 1 ) and a restricted geographical distribution type species for Stygonitocrella, nor for Nitocrellopsis, both generic (Table 2). This subdivision is admittedly based on swimming leg names were unavailable until recently. Galassi etal. (1999) fixed A'. characters only but we suspect it to be at least partly reinforced by rouchi Galassi, De Laurentiis & Dole-Olivier, 1999 as the type of mouthpart and antennulary characters when they become available. Nitocrellopsis, making the name available with their authorship. A Formal recognition of these lineages as distinct genera is impossible similar course of action was taken by Reid et al. (in press) who since most descriptions are severely lacking in detail and many of designated S. montana (Noodt, 1965) as the type of Stygonitocrella. them are based on very few specimens or one sex only. For example, Leg segmentation characters should be used with caution when the description of P. djirgalanica is completely lacking in illustra- inferring relationships in derived lineages. For example, the evolu- tions (Borutzky, 1978). In addition, the type material of the great tionary instability of endopodal segmentation is illustrated by the majority of its species is no longer extant and additional records genus Psammonitocrella Rouch which contains species with 1- (P. have not been added. 5. petkovskii differs from its congeners in the longifurcata) or 2-segmented P2-P3 endopods (P. boultoni). Unique absence of the inner seta on P3 exp-2, an element which is present in derived characters such as the loss of the outer spine on PI exp-2 all other congeners. Attempts to trace the single female on which this leave no doubt that both Psammonitocrella species shared a com- description was based failed (Galassi, pers. comm.). mon ancestor, and hence the discrepancy observed in endopodal As pointed out by Reid et al. (in press) the generic placement of segmentation has to be interpreted as the result of intrageneric S. orghidani (Petkovski, 1973) is questionable. The original descrip- evolution. Utilizing endopodal segmentation patterns in defining tion is very concise, showing illustrations only of the antenna, generic boundaries is potentially misleading. Overweighing such caudal rami and the fifth legs. The exopodal armature of the swim- characters at the expense of others can result in assigning species to ming legs is largely unknown, apart from Petkovski's statement that 1 48 W. LEE AND R. HUYS -I- EL, + +P-P- + + +-+ + + +^- + + e« I+ Hi t/> &H 1+ + +^- + + + + + + + + ~- + + as 1i - SO - e>- 1+ + +^- + + + + + + + + I I u £3 e-. I \ c~ + + o- c~ + o- + + c~ + + I — c-~ O © © —< e-~ © • — t>- o. o- © d O O 3* (NmmcNmcirocirocom mmmrn O O CS — O (N e>- (N m t>- C-- O O O fN O CN O CN (N (N I © O — O c^- c> Nn^tO't^t't^t't I m © CS 0©©OIOc^-OCNCNc«-)C'->© O O —< © o- 0. — rjr^^toc^-^-Tj-Tt'S-'^-m rMOmtN oooooooooo O O I OOOOOOOOOO o © CN (N CN — rv. CNCMCNCNCNCNCN o o ©©55©o©S«©W©°. °. — OOOCN cn —P. OOOOOO: cn OOH r OOOOSpp2 M (N (N — r . (NtSMNNNOI NNOO ppOp — ppppppp^jpOpp ddddc-'ddddddd — do'o'd OO OO OO OOO 2 g 5 5 2 2 2 o 5 2 5 = ==52 000—: oooddodd — ddo NNNN^tNIMNNNMN CN CN O — poop — pppppppSpPPP ~- — — : — ' — — _ — — —,„-_ — ddddo'-ddddddd~dddd— c> — — OOOO c-. 0000000 OOOO OOOO(N cn m m OOOOOOOm cn m cn m m cn OOOOm m M (N d d d d d d d d d d— d d d d d _;_:_'_:_:od — — — '— — c-: d © CN CN CN CN c. N N N O) M (N M CN (N CN CN M M (N M f CM N N IN N (N M CN CN CM CN O O O O o OOOOOOO poop o d — —'• o -^ d d d d d — d d d d c- d d d d d d d d d d d Ot *o £ -2 1 « i- s o U a g e .u -s: £ <* is o -3 a o zsS q ** -s: t^ S -^ 3 go S 00 5 = *• 00-5 2 5 > 3 £ * 1 s S o "a oo CN.-5 a -2 t Table 2 Species groups within Stygoniwcrella showing zoogeographical connection, habitat and number of specimens known Species Distribution Habitat Material dubia (Chappuis, 1937) North Spain cavernicolous 1 3 guadalfensis Rouch, 1 985 Andalusia hyporheic 9 99,4 63 karamani (Petkovski, 1959) Macedonia phreatic 2 99,3 33 Ijovuschkini (Borutzky, 1967) Ukraine cavernicolous 3 99,2 do* colchica (Borutzky & Michailova-Neikova. 1970) West Georgia cavernicolous 3 99,4 33 petkovskii Pesce, 1 985 Lesbos, Greece phreatic 1 9 tianschanica (Borutzky, 1972) Kirghiztan phreatic 2 99,3 33 pseudotianschanica (Sterba, 1973) Afghanistan phreatic 2 99 djirgalanica (Borutzky. 1978) Kirghiztan phreatic 7 99,2 33 montuna (Noodt, 1965) Argentina phreatic 4 99,6 33 orghidani (Petkovski. 1973) Cuba cavernicolous 4 99,2 33 insularis (Miura. 1962) Ryukyu Islands phreatic 2 99, 1 d the distal exopod segment of P2-P4 possesses an inner seta. The P2-P4 with 3-segmented exopods and 1 -segmented (P2-P3) or species is unusual in having a bisetose antennary exopod and rudimentary (P4) endopods. Bases with outer seta. Apical setae of remarkably reduced fifth legs which consist of a median transverse exp-3 very long. P3 endopod 8 with inner distal seta shorter and plate retaining only the outer basal setae. Petkovski (1973) points outer distal spine longer than in 9. P4 endopod represented by minute out several similarities between S. orghidani and Nitocrella negreai knob. Armature formula: such as the large size of the antennules and antennae, the short caudal rami and the elongate PI endopod. He regarded these Exopod Endopod characters as evidence for a distinct lineage within Nitocrella sensu lato. Apart from the 1 -segmented P4 endopod there is no evidence P2 0.1.022 020 for a relationship with any of the other species currently included in P3 0.1.022 020 P4 0.1.122 Stygonitocrella and in view of its fragmentary description we pro- pose to relegate it to species incertae sedis within this genus. P5 rudimentary with baseoendopod fused to somite and repres- The only oriental species, S. insularis. was described from the ented only by outer basal seta. Exopod a small segment with 2 setae Ryukyu archipelago (Miura, 1962c/) and shows a number of unique in 9 and 1 pinnate fused spine plus 2 setae in 8. P6 rudimentary, characteristics within Stygonitocrella. It is regarded here as the type forming unarmed median operculum in 9; unconfirmed in 8. species of a new genus Neonitocrella on account of the bisetose Caudal ramus short, with 7 setae; seta V longest. antennary exopod, the loss of the P4 endopod (represented by a rudimentary knob) and the reduced P5 in both sexes. The PI TYPE AND ONLY SPECIES. Nitocrella insularis Miura. 1962 = armature pattern, the reduced antennary exopod and the character- Neonitocrella insularis, (Miura, 1962) comb. nov. istic structure of the male P5 exopod suggest a sistergroup relationship with /. humphreysi, to which it is also closest zoogeographically. Conversely, Nitocrella japonica described from Shikoku Island Acknowledgements. The authors would like to thank Dr W.F. Humphreys (Western Australian Museum. Australia) for providing us with (Japan) (Miura, 1962i>) also shows a remarkable similarity with /. the copepod material. of us (W.L.) acknowledges financial support humphreysi in the structure of the antenna (unisetose exopod). One from the Korea Research Foundation provided for the programme year 1997. maxilla (elongate allobasis) and maxilliped (slender basis and claw) but exhibits a more primitive leg formula. This suggests that the genus Inermipes could have been derived from a N. japonica-hke ancestor through swimming leg reduction. REFERENCES Borutzky, E.V.. 1967. Biospeologica Sovietica XXXII. On the genus Nitocrella gen. nov. Genus NEONITOCRELLA Chappuis (Copepoda. Harpacticoida). Byultetin' Moskovskogo Obshchestva Ispytatelei Prirody, Otdel Biolohichesky, New Series 72(3): 32-39. [In Russian]. DIAGNOSIS. Ameiridae. Body elongate, cylindrical, and vermi- . 1972. Copepoda Harpacticoida from subterranean water of the shore of Issyk-kul form without distinct surface ornamentation except for ventral and Southern Kisilkum. Trudy Zoologicheskogo Instituta, Leningrad 51: 98-119. spinule patterns on abdomen. Cephalothorax and other somites with . 1978. Copepoda Harpacticoida grunlovykh vod tyan'shanya. In: Issledovaniya faune sovelskogo soyuza (bespozvonochnye 1 ryby). Sbornik smooth posterior margin. Genital and first abdominal somites appar- po Trudov Zoologicheskogo Muzeya MGU 16: 66-91. [In Russian]. ently separated. Anal operculum well developed. & Mikhailova-Neikova, M. 1970. Harpacticoida (Copepoda) of Caves in West in Sexual dimorphism antennule, P3 endopod, P5, P6, and in Georgia. Azerbaijan and Middle Asia. Zoologicheskij Zhurnal 49: 1334-1341. genital segmentation; unconfirmed for inner basal spine of PI. Chappuis, P.A. 1924. Descriptions preliminaires de Copepodes nouveaux de Serbie. Rostrum fused to cephalothorax, not defined at base. Antennule 8- Buletinul Societdfii de Stiinje din Cluj 2: 27-45. . 1937. Subterrane Harpacticoiden aus Nord-Spanien. Buletinul Societdfii de segmented in 9, unknown in 8; segment 1 with seta; aesthetascs on Stiinte din Cluj 8: 556-571. segments 4 and possibly 8 in 9, unknown in 8. Antenna with separate De Laurentiis, P., Pesce, G.L. & Humphreys, W.F. 1999. Copepods from ground basis and endopod; exopod well developed segment, with 2 setae. waters of Western Australia. IV. Cyclopids from basibs and craton aquifers (Crusta- Mandible with 2-segmented palp, comprising unarmed basis and cea: Copepoda: Cyclopidae). Records ofthe Western Australian Museum 19: 243-257. Galassi, D.M..P, De Laurentiis, P. & Dole-Olivier, M.-J. 1999. Nitocrellopsis rouchi endopod with 4 setae. Maxillule, maxilla and maxilliped unconfirmed. sp. n., a new ameirid harpacticoid from phreatic waters in France (Copepoda: PI with 3-segmented rami; basis without outer seta; exp-1 not Harpacticoida: Ameiridae). Hydrobiologia 412: 177-189. elongate, without seta, exp-2 inner exp-3 with 4 elements; endopod Humphreys, W.F. 1 993. Stygofauna in semi-arid tropical western Australia: A Tethyan with formula [1.0.020]. connection? Memoires de Biospeologie 20: 1 1 1-1 16. 50 W. LEE AND R. HUYS . 2000. The hypogean fauna of the Cape Range peninsula and Barrow Island. (Crustacea: Copepoda: Cyclopidae). Records of the Western Australian Museum 18: North-West Australia. Pp. 581-601. In. H. Wilkens, D.C. Culver & W.F. Humphreys 67-76. , . 1 (eds). Ecosystems of the World. Vol. 30. Subterranean Ecosystems. Elsevier, & 996/7. Copepods from ground waters of Western Australia, II . The Amsterdam. genus Halicyclops (Crustacea: Copepoda: Cyclopidae). Records of the Western Huys, R., Gee, J.M., Moore, C.G. & Hamond, R. 1 996. Synopses ofthe British Fauna Australian Museum 18: 77-85. (New Series): Marine and Brackish Water Harpacticoid Copepods Part /.viii + Petkovski, T.K. 1959. Neue und bemerkenswerte Harpacticoide RuderfuBkrebse 352p. Field Studies Council, Shrewsbury, United Kingdom. (Crust. Cop.) aus den Grundgewassern Jugoslaviens. Acta Musei Macedonici Lang, K. 1965. Copepoda Harpacticoida from the Califomian Pacific Coast. Kungliga scientiarum naturalium 6: 101-119. Svenska VetenskapsAkademiens Handlingar (4)10(2): 1-560. . 1973. Subterrane Siisswasser Harpacticoida von Kuba (Vorlaufige Mitteilung). Miura, Y. 1962a. Subterranean harpacticoid copepods of the Amami Group of the Resultats des Expeditions Biospeologiques Cuhano-Roumaines a Cuba 1: 125- Ryukyu Islands. Annotationes zoologicae Japonenses 35: 95-105. 141. . . 1962/;. Three new harpacticoid copepods from subterranean waters of Shikoku in 1976. Drei neue Nitocrelta-Aften von Kuba. zugleich eine Revision des Genus Japan. Japanese Journal of Zoology 13: 267-274. Nitocrella Chappuis (s. restr.) (Crustacea, Copepoda, Ameiridae). Acta Musei Noodt, W. 1965. Crustacea subterranea aus Argentinien. Beitrdge zur Neotropischen Macedonici scientiarum naturalium 15: 1-26. Fauna 4: 84-129. Reid, J.W., Hunt G.W. & Stanley E.H. (in press). A new species of Stygonitocrella Pesce, G.L. 1 985. Stygobiological researches in subterranean waters of Lesbos (Greece) (Crustacea: Copepoda: Ameiridae), the first report of the genus in North America. and description of Stygonitocrella petkovskii n. sp. (Crustacea Copepoda: Ameiridae). Proceedings of the Biological Society of Washington, in press. Acta Musei Macedonici scientiarum naturalium 12: 125—139. Rouch, R. 1985. Une nouvelle Stygonitocrella (Copepoda. Harpacticoida) des eaux & . 1996. Copepods from ground waters of Western Australia. III. Diacyclops souterraines d'Andalousie. Espagne. Stygologia 1: 118-127. humphreysi n. sp. and comments on the Diacyclops crassicaudis-comp\ex (Cope- . 1992. Une nouveau genre d' Ameiridae (Copepoda. Harpacticoida) dans le milieu poda: Cyclopidae). Crustaceana 69: 524—531. hyporheique d'un cours d'eau de l'Arizona. Stygologia 7: 149-157. , & Humphreys, W.F. 1996n. Copepods from ground waters of Western Sterba, O. 1973. Die neuen Harpacticidenarten der Gattung Nitocrella (Crustacea. Australia, I. The genera Meiacyciops. Mesocyclops, Microcyclops and Apocyclops Copepoda) aus Afghanistan. Zoologischer Anzeiger 190: 333-342. 1 ) Bulletin of The Natural History Museum Zoology Series Earlier Zoology Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. Volume 57 comments on its phylogenetic relationships. G. J. Howes A review of the serranochromine cichlid fish genera No. 1 Morphology and biometry of twelve soil testate amoebae Pharyngochromis, Sargochromis, Serranochromis and (Protozoa, Rhizopoda) from Australia. Africa and Austria. G. Chetia Luftenegger & W. Foissner (Teleostei: Labroidei). P. H. Greenwood A revision of Danielssenia Boeck and Psammis Sars with the A revision of Cothumia (Ciliophora: Peritrichida) and its establishment of two new genera A rchisenia and Bathypsammis morphological relatives. A. Warren & J. Paynter (Harpacticoida: Paranannopidae). R. Huys and J. M. Indian Ocean echinoderms collected during the Sinbad Voyage Gee. A new species of Syrticola Willems & Claeys. 1982 (Cope- (1980-81): 2. Asteroidea. L. M. Marsh & A. R. G. Price poda: Harpacticoida) from Japan with notes on the type The identity and taxonomic status of Tilapia amoldi Bilchrist species. R. Huys and S. Ohtsuka and Thompson, 1917 (Teleostei. Cichlidae). P. H. Greenwood Erratum £40.30 Anatomy, phylogeny and txonomy of the gadoid fish genus Macruronus Giinther. 1873. with a revised hypothesis of No. 2 The status of the Persian Gulf sea snake Hydrophis lapemoides gadoid phylogeny. G. J. Howes £38.50 (Gray. 1849) (Serpentes. Hydrophiidae). A. Redsted Rasmussen. No. 2 The pharyngobranchial organ of mugilid fishes; its structure, Taxonomic revision of some Recent agglutinated foraminifera variability, ontogeny, possible function and taxonomic utility. from the Malay Archipelago, in the Milieu Collection. The I. J. Harrison & G. J. Howes Natural History Museum, London. P. Bionntmann and J. E. Cranial anatomy and phylogeny of the South-East Asian Whittaker. catfish genus Belodontichthys. G. J. Howes & A. Fumihito Foregut anatomy, feeding mechanisms, relationships and A collection of seasnakes from Thailand with new records of classification of the Conoidea (=Toxoglossa)(Gastropoda). J. D. Hydrophis belcheri (Gray). C. J. McCarthy & D. A. Warrell Taylor. Y. I. Kantor and A. V. Sy soev. 1 993. Pp. 97- 1 70. £40.30 The copepod inhabitants of sponges and algae from Hong Kong. S. Malt The freshwater cyclopoid of Nigeria, with an illustrated key to Volume 60 all species. G. A. Boxshall & E. I. Braide species (Echinodermata: Asteroidea) A new of Ferdina from No. 1 A new subfamily and genus in Achatinidae (Pulmonata: the Sultanate of Oman with discussion of the relationships ol Sigmurethra). A. R. Mead. the genus within the family Ophidiasteridae. L. M. Marsh & A. On Recent species of Spiraserpula Regenhardt. 1961. a C. Campbell £38.50 serpulid polychaete genus hitherto known only from Creta- ceous and Tertiary fossils. T. Gottfried Pillai and H.A. Ten Volume 58 Hove. 1994. Pp. 1-104. £40.30 No. 1 The morphology and phylogeny of the Cerastinae (Pulmonata: No. Phylogenetic relationships between arietellid genera (Copepoda: Pupilloidea). P. B. Mordan Calanoida ). with the establishment of three new genera. S. A redescription of the uniquely polychromatic African cichlid Ohtsuka. G. A. Boxshall and H. S. J. Roe. 1994. Pp. 105-172. fish Tilapia guinasana Trewavas. 1936. P. H. Greenwood £40.30 A revision and redescription of the monotypic cichlid genus Pharyngochromis (Teleostei, Labroidei). P. H. Greenwood Description of a new species of Microgale ( Insectivora: Volume 61 Tenrecidae) from eastern Madagascar. P. D. Jenkins Studies on the deep-sea Protobranchia (Bivalvia): the family No. 1 A revised familial classification for certain cirrhitoid genera Nuculidae. P. M. Rhind and J. A. Allen £40.30 (Teleostei. Percoidei Cirrhitoidea). with comments on the group"s monophyly and taxonomic ranking. PH. Greenwood. No. 2 Notes on the anatomy and classification of ophidiiform fishes Studies on the deep-sea Protobranchia (Bivalvia): the Sub- with particular reference to the abyssal genus Acanthomis family Yoldiellinae. J. A. Allen, H.L. Sanders and F Hannah. Giinther, 1878. G. J. Howes 1995. Pp. 1-90. £40.30 Morphology and morphogenesis of the soil filiate Bakuella edaphoni nov. spec, and revision of the genus Bakuella No. 2 Primary studies on a mandibulohyoid 'ligament' and other Agamaliev & Alekperov, 1976 (Ciliophora, Hypotrichida). W. intrabucal connective tissue linkages in cirrhitid. latrid and Song, N. Wilbert and H. Berger cheilodactylid fishes (Perciformes: Cirrhitoidei). P.H. A new genus and species of freshwater crab from Cameroon, Greenwood. West Africa (Crustacea, Brachyura, Potamoidea, A new species of Crocidura (Insectivora: Soricidae) recovered Potamonautidae). N. Cumberlidge and P. F. Clark from owl pellets in Thailand. P.D. Jenkins and A.L. Smith. On the discovery of the male of Monnonilla Giesbrecht. 1 89 Redescription of Sudanonautes Floweri (De Man, 1901 (Copepoda: Mormonilloida) R. Huys, G. A. Boxshall and R. (Brachyura: Potamoidea: Potamonautidae) from Nigeria and Bottger-Schnack £40.30 Central Africa. N. Cumberlidge. Association of epaxial musculature with dorsal-fin Volume 59 pterygiophores in acanthomorph fishes, and its phylogenetic No. 1 A new snake from St Lucia, West Indies. G. Underwood significance. R.D. Mooi and A.C. Gill. 1995. Pp. 91-138. Anatomy of the Melanonidae (Teleostei: Gadiformes). with £40.30 . Volume 62 Volume 65 No. 1 Phylogenetic relationships of Toad-headed lizards No. 1 Deep-sea conolidean gastropods collected by the John Murray {Phrynocephalus, Agamidae) based on morphology. Expedfition, 1933-34. A. V. Sysoev. E.N. Arnold. Reassessment of 'Calcinus' astathes Stebbing 1924 (Crustacea: Rita sacerdotum, a valid species of catfish from Anomura: Paguridea: Diogenidae). P.A. McLaughlin. Myanmar (Pisces, Bagridae). C.J. Ferraris, Jr. On a new species of Ophidiaster (Echinodermata: Asterpodea) Indian Ocean echinoderms collected during the Sinbad Voyage from southern China. Y. Liao and A.M. Clark. (1980-81): 4. Crinoidea. J.I.M. Crossland and A.R.G. Price. The life cycle of Paracyclops fimbriatus (Fischer. 1853) On the hybrid status of Rothschild's Parakeet Psittacula (Copepoda. Cyclopoida). S. Karaytug and G.A. Boxshall. 1996. intermedia Pp 1-70. £40.30 (Aves, Psittacidae). PC. Rasmussen and N.J. Collar. A review of the genus Bargamannia Totton, 1954 No. 2 Indian Ocean echinoderms collected during the Sindbad Voyage (Siphonophorae. Physonecta. Pyrostephidae). PR. Pugh. 1999. (1980-81): 3. Ophiuroidea and Echinoidea. A.R.G. Price and Pp. 1-72. £40.30 F.W.E. Rowe. No. 2 Systematics and phylogeny oiZausodes C.B. Wilson, 1932 Rare cyclopoid copepods (Crustacea) from Mediterranean (Copepoda. Harpacticoida, Harpacticidae), including three new littoral caves. D. Jaume and G.A. Boxshall. species from the northern Gulf of Mexico. L. Bouck. D. Thistle Studies on the deep-sea Protobranchia (Bivalvia): the family and R. Huys. Neilonellidae and the family Nuculanidae. J. A. Allen and Nybelinia southwelli sp. nov. (Cestoda. Trypanorhyncha) with H.L. Sanders. 1996. Pp 71-132. £40.30 the re-description of N. perideraeus (Shipley & Hornell, 1906) and the synonymy of N. herdmani (Shipley & Hornell. 1906) Volume 63 with Kotorella pronosoma (Stossich, 1901). H.W. Palm and T. Walter. No. 1 A new species of Microgale (Insectivora: Tenrecidae), with Nybelinia Poche. 1926, Heteronybelinia gen. nov. and comments on the status of four other taxa of shrew tenrecs. Mixonybelinia gen. nov. (Cestoda. Trypanorhyncha) The PD. Jenkins, C.J. Raxworthy and R.A. Nussbaum. Natural History Museum, London. H.W. Palm. Notes on the anatomy and relationships of Sundasalanx A new species of Microgale (Lipotyphla. Tenrecidae) from Roberts (Teleostei, Clupeidae), with descriptions of four new isolated forest in southwestern Madagascar. PD. Jenkins and species from Borneo. D.J. Siebert. S.M. Goodman. Redescription of and lectotype designation for Balistes Modes of ear reduction in iguanian lizards (Reptilia, Iguania): macropepsis Boulenger, 1 887, a senior synonym of different paths to similar ends. E.N. Arnold. 1 999. Pp. 73- 1 7 1 Canthidermis longirostris Tortonese, 1954 and C. villosus £40.30 Fedoryako, 1979 (Teleostei. Tetraodontiformes, Balistidae). A.C. Gill and J.E. Randall. Volume 66 new species of crassispirine gastropod from the Houtman A No. 1 Generic concepts in the Clymnestridae (Copepoda, Abrolhos islands, Western Australia (Gastropoda, Conoidea, Harpacticoida): revision and revival. R. Huys and S. Conroy- Crassispirinae). A.V. Sysoev and J.D. Taylor. Dalton. Foregut anatomy and relationships of the Crassispirinae Basal resolution of laophontid phylogeny and the paraphyly of (Gastropoda, Conoidea). Y.I. Kantor. A. Medinskaya and Esola Edwards. R. Huys and W. Lee. 2000. Pp. 1-107. £40.30 J.D. Taylor. 1997. Pp 1-92. £40.30 No. 2 The shallow-water marine Mollusca of the Estero Elefantes and No. 2 The lucinid bivalve genus Cardiolucina (Mollusc: Bivalvia: Laguna San Rafael, southern Chile. D.G. Reid and C. Osorio. Lucinidae); systematics, anatomy and relationships. J.D. Taylor Remarks on the genera Schizocuma Bacescu, 1972 and and E.A. Glover. Styloptocuma Bacescu & Muradian, 1974 (Crustacea, A new species of water mouse, of the genus Chibchanomys Cumacea). I. Petrescu. (Rodentia: Muridae: Sigmondontinae) from Ecuador. PD. A new species of Sigambra (Polychaeta. Pilargidae) from the Jenkins and A. A. Barnett. abyssal plains of the NE Atlantic. G.L.J. Paterson and A.G. A new species in the asterinid genus Patiriella (Echinodermata: Glover. 2000. Pp. 1 09- 1 70. £40.30 Asteroidea) from Dhofar, southern Oman: a temperate taxon in Volume 67 a tropical locality. A.C. Campbell and F.W.E. Rowe. Morphological observations on Oncaea mediterranea (Claus, No. 1 Freshwater nematodes from Loch Ness. Scotland Part I. The 1863) (Copepoda: Poecilostomatoida) with a comparison of orders Tylenchida Thorne, 1949 and Rhabditida Chitwood, Red Sea and eastern Mediterranean populations. R. Bottger- 1933 (Nematoda, Secernentea). FR. Wanless and R. Hunter. Schnack and R. Huys. 1997. Pp. 93-147. £40.30 Taxonomy of Oncaeidae (Copepoda. Poecilostomatoida) from the Red Sea. II. Seven species of Oncaea s.str. R. Bottger- Schnack. Volume 64 Review of the False Smooth snake genus Macroprotodon No. 1 A revision of the cladoceran genus Simocephalus (Crustacea, (Serpentes. Colubridae) in Algeria with a description of a new Daphniidae). Marina J. Orlova-Bienkowskaja. species. E. Wade. 2001. Pp. 1-107. £40.30 Structural niche, limb morphology and locomotion in lacertid No. 2 Type material of Stegocephalidae Dana, 1855 (Crustacea, lizards (Squamata, Lacertidae); a preliminary survey. Amphipoda) in the collections of The Natural History Museum, E.N. Arnold. London, including the description of seven new species. J. Hetereleotris georgegilli, a new species of gobiid fish, with Berge, W. Vaderand A. Galan. notes on other Mauritian Hetereleotris species. Anthony C. Gill The genus Ischioscia Verhoeff, 1 928 in Venezuela, with the Revision of Schismatorhynehos Bleeker, 1855 (Teleostei, description of six new species (Crustacea, Oniscidea, Cyprinidae), with the description of two new species from Philosciidae). Andreas Leistikow. Borneo. Darrell J. Siebert and Agus H. Tjakrawidjaja. 1998. A review of the genus Erenna Bedot, 1904 (Siphonophora, Pp. 1-109. £40.30 Physonectae). PR. Pugh. No. 2 Partial revision of Paracyclops Claus, 1893 (Copepoda, A new species of loach, genus Nemacheilus (Osteichthyes, Cyclopoida, Cyclopidae) with descriptions of four new species. Balitoridae) from Aceh, Sumatra, Indonesia. R.K. Hadiaty and S. Karaytug and G.A. Boxhall. D.J. Siebert. First records and a new subspecies of Rhinolophus stheno Revision of the western Indian Ocean fish subfamily (Chiroptera, Rhinolophidae) from Vietnam. G. Csorba and P. Anisochrominae (Perciformes, Pseudochromidae). A.C. Gill Jenkins. 1998. Pp. 111-211. £40.30 and R. Fricke. 2001 . Pp. 109-207. £40.30 . GUIDE FOR AUTHORS Aims and scope. The Bulletin of the British Museum (Natural sub-headings should be followed. All should be ranged left and be in History) Zoology, was established specifically to accommodate upper and lower case. Supra-generic systematic headings only manuscripts relevant to the Collections in the Department of Zool- should be in capitals; generic and specific names are to be in italics, ogy. It provides an outlet for the publication of taxonomic papers underlined. Authorities for species names should be cited only in the which, because of their length, prove difficult to publish elsewhere. first instance. Footnotes should be avoided if at all possible. Preference is given to original contributions in English whose References. References should be listed alphabetically. Authori- contents are based on the Collections, or the description of speci- ties for species names should not be included under References, mens which are being donated to enhance them. Acceptance of unless clarification is relevant. The author's name, in bold and lower manuscripts is at the discretion of the Editor, on the understanding case except for the initial letter, should immediately be followed by that they have not been submitted or published elsewhere and the date after a single space. Where an author is listed more than become the copyright of the Trustees of the Natural History Mu- once, the second and subsequent entries should be denoted by a long seum. All submissions will be reviewed by at least two referees. dash. These entries should be in date order. Joint authorship papers Submission of manuscripts. Initially three clear, complete cop- follow the entries for the first author and an '&' should be used ies should be submitted in the style and format of the Bulletin. The instead of 'and' to connect joint authors. Journal titles should be text must be typed double-spaced throughout, including references, entered in full. Examples: (i) Journals: England, K.W. 1987. Certain tables and legends to figures, on one side of A4 paper with 2.5 cm Actinaria (Cnidaria, Anthozoa) from the Red Sea and tropical Indo- margins. All pages should be numbered consecutively, beginning Pacific Ocean. Bulletin of the British Museum (Natural History), with the title page as p. 1 . SI units should be used where appropriate. Zoology 53: 206-292. (ii) Books: Jeon, K.W. 1973. The Biology of Whenever possible a copy of the text, once the paper has been Amoeba. 628 p. Academic Press, New York & London, (iii) Articles accepted, should also be provided on floppy disc (see below). Discs from books: Hartman, W.D. 1981. Form and distribution of silica in should only be sent after final acceptance, as papers generally need sponges, pp. 453-^93. In: Simpson, T.L. & Volcani, B.E. (eds) revision after refereeing. If it is impossible to provide an appropriate Silicon and Siliceous Structures in Biological Systems. Springer- disc please ensure that the final typescript is clearly printed. Verlag, New York. Authors are requested to ensure that their manuscripts are in final Tables. Each table should be typed on a separate sheet designed to format, because corrections at proof stage may be charged to the extend across a single or double column width of a Journal page. It author. Additions at proof stage will not normally be allowed. Page should have a brief specific title, be self-explanatory and be supple- proofs only will be sent. mentary to the text. Limited space in the Journal means that only Word-processor discs. Please follow these instructions. modest listing of primary data may be accepted. Lengthy material, 1 Ensure that the disc you send contains only the final version of such as non-essential locality lists, tables of measurements or details the paper and is identical to the typescript. of mathematical derivations should be deposited in the Biological 2. Label the disc with the author's name, title of the paper and Data Collection of the Department of Library Services, The Natural the word-processor programme used. Indicate whether IBM or History Museum, and reference should be made to them in the text. Apple Mac (IBM preferred). Illustrations 3. Supply the file in the word-processor format; if there is a drawings - Figures should be designed to go across single (84 mm facility to save in ASCII please submit the file in ASCII as well. wide) or double (174 mm wide) column width of the Journal page, 4. Specify any unusual non-keyboard characters on the front type area 235 x 1 74 mm. Drawings should be in black on white stiff page of the hard copy. card with a line weight and lettering suitable for the same reduction 5. Do not right-justify the text. throughout, ideally not more than 40%. After reduction the smallest 6. Do not set a left-hand margin. lettering should be not less than 10 pt (3 mm). Tracing paper should 7. Make sure you distinguish numerals from letters, e.g. zero (0) ideally be avoided because of the possibility of shadows when from O; one ( 1 ) from 1 (el) and I. scanned. All artwork must have bulletin, author and figure number 8. Distinguish hyphen, en rule (longer than a hyphen, used included, outside of the image area, and must be free of pencil, glue without a space at each end to signify "and' or 'to', e.g. the Harrison- or tape marks. Nelson technique, 91-95%, and increasingly used with a space at PHOTOGRAPHS - All photographs should be prepared to the final size each end parenthetically), and em rule (longer than an en rule, used of reproduction, mounted upon stiff card and labelled with press-on with a space at each end parenthetically) by: hyphen, two hyphens lettering (eg Letraset). They can be mounted on white or black and three hyphens, respectively. Be consistent with rule used paren- background; a black background must be evenly black all over: any thetically. background must be free of all pencil and glue marks within the 9. Use two carriage returns to indicate beginnings of paragraphs. image area. All figures should be numbered consecutively as a 10. Be consistent with the presentation of each grade of heading single series. Legends, brief and precise, must indicate scale and (see Text below). explain symbols and letters. Photos, when components of figure- Title. The title page should be arranged with the full title; name(s) plates should be abutted, trimmed as regular rectangles or close of author(s) without academic titles; institutional address(es); sug- trimmed up to edge of specimen. Joins etc. can be removed at the gested running title; address for correspondence. scanning stage but at extra cost. Cropping instructions, if any, should be indicated on an overlay or marked on a photocopy of the figure. Synopsis. Each paper should have an abstract not exceeding 200 SIZE - Maximum size of artwork for use of flatbed scanners is A3. words. This should summarise the main results and conclusions of Larger artwork has to be reduced photographically prior to scan- the study, together with such other information to make it suitable ning, therefore adding to expense. for publication in abstracting journals without change. References must not be included in the abstract. Symbols in text. Male and female symbols within the text should be flagged within curly brackets to enable setter to do a swift global Text. All papers should have an Introduction, Acknowledge- search. ments (where applicable) and References; Materials and Methods should be included unless inappropriate. Other major headings are Reprints. 25 reprints will be provided free of charge per paper. left to the author's discretion and the requirements of the paper, Orders for additional reprints can be submitted to the publisher on subject to the Editors' approval. Three levels of text headings and the form provided with the proofs. Later orders cannot be accepted. 1 Another variation on the gymnure theme: description of a new species of Hylomys (Lipotyphla, Erinaceidae, Galericinae). P. D. Jenkins and M.F. Robinson 13 A new species of freshwater crab (Brachyura, Potamoidea, Potamonautidae) from Principe, Gulf of Guinea, Central Africa N. Cumberlidge, P. F. Clark and J. Baillie 19 Two new species of the Indo-Pacific fish genus Pseudoplesiops (Perciformes, Pseudo- chromidae, Pseudoplesiopinae) A.C. Gill and A.J. Edwards 27 A redescription of Sousa chinensis (Osbeck, 1765) (Mammalia, Delphinidae) and designa- tion of a neotype L.J. Porter 39 A new genus of groundwater Ameiridae (Copepoda, Harpacticoida) from boreholes in Western Australia and the artificial status of Stygonitocrella Petkovski, 1976 W. Lee and R. Huys ZOOLOGY SERIES Vol. 68, No. 1, June 2002