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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

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Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset

Bull. nat. Hist. Mus. Lond. (Zool.) 68(1): 1-11

Issued 27 June 2002

Another variation on the gymnure theme:

description of a new species of Hylomys

(Lipotyphla, Erinaceidae, Galericinae).

PAULINA D. JENKINS

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

MARK F. ROBINSON

Waterway Conservation & Regeneration, British Waterways, Llanthony Warehouse, Gloucester Docks,

Gloucester, GLI 2EJ

SYNOPSIS.

A new species of Hylomys from Lao Peoples Democratic Republic is described, based on morphological

comparisons with other members of the subfamily Galericinae. The relationships revealed by a phylogenetic analysis are

discussed and compared with those of a previous published analysis.

INTRODUCTION

The Family Erinaceidae is divided into two subfamilies: the wide-

spread Erinaceinae (hedgehogs) occuring in Africa, Europe and

Asia, and the Galericinae (moonrats and gymnures), which is con-

fined to southeast Asia, Indonesia and the Philippines. There has

been considerable disagreement over the correct name to apply to

the subfamily of moonrats and gymnures, summarised by Frost er al.

(1991), who favoured the use of Hylomyinae. McKenna & Bell

(1997) pointed out however, that the use of Galericini as a tribal

name by Butler (1948) had been accepted by many subsequent

writers, particularly palaeontologists, and was therefore the appro-

priate name to use. In this paper we follow McKenna & Bell (1997)

in using the name Galericinae. Most authors up to and including

Corbet (1988), considered that the Galericinae includes five genera:

Echinosorex Blainville, 1838, Hylomys Miiller, 1840, Neotetracus

Trouessart, 1909, Neohylomys Shaw & Wong, 1959 and

Podogymnura Mearns, 1905, all but the latter being monotypic. In

their revision of the family Erinaceidae, Frost et al.(1991) con-

cluded that there are only three valid genera within the Galericinae:

Echinosorex, Podogymnura and Hylomys. They accepted Hylomys

as a rather variable but nevertheless monophyletic genus, although

they conceded that there was evidence to support the retention of

Neotetracus and Neohylomys as subgenera.

The genus Hylomys is widely distributed in southeast Asia and

Indonesia. Hylomys suillus Miiller, 1840 occurs in Malaysia, Indo-

nesia, Thailand, Vietnam, Cambodia, Lao Peoples Democratic

Republic (PDR), Myanmar and southern PDR China; in Lao PDR it

has been recorded from Phongsali, Xiangkhouang, Vientiane and

Dong Hua Sao National Biodiversity Conservation Area (NBCA)

(Robinson, 1999). A number of different subspecies have been

attributed to H. suillus and the biochemical and metrical variation

within this species was examined by Ruedi ef al. (1994). They

recognised that much of the high level of variation could be attrib-

uted to the geographical and altitudinal isolation of the named forms

but demonstrated that one of these taxa, H. parvus Robinson &

Kloss, 1916, merited specific status. Hylomys sinensis (Trouessart,

1909) occurs from southern China to Myanmar and northern Viet-

nam; it has not been recorded from Lao PDR but is likely to occur in

those areas adjacent to northern Vietnam, whence it is recorded by

Osgood (1932). Hylomys hainanensis (Shaw & Wong, 1959) is

restricted to Hainan Island, PDR China and H. parvus is known only

from Sumatra, Indonesia. Another geographically isolated

undescribed gymnure has been discovered recently from a region of

limestone karst in the Lao PDR. While sharing many characters with

other species of Hylomys this new species also differs markedly

from its congeners and, furthermore, shares some features with

geographically remote species of Podogymnura. The new taxon has

been compared in particular with specimens of H. suillus, although

there is no indication that the two species occur sympatrically, and

also with H. sinensis, H. parvus and, in the absence of specimens,

with the original description of H. hainanensis and the figures of the

skull of this species in Frost er al. (1991). In addition comparisons

were made with the other genera of Galericinae: Echinosorex

gymnura (Raffles, 1822) from Malaysia and Indonesia, and

Podogymnura truei Mearns, 1905 from the Philippines. In order to

assess phylogenetic relationships, both the new taxon and H. parvus

were analysed using the criteria employed by Frost et al. (1991).

MATERIAL AND METHODS

Comparative material was examined from the collections of the

Natural History Museum (BMNH), London (formerly the British

Museum (Natural History)), the American Museum of Natural

History, New York (AMNH), the Muséum National d’ Histoire

Naturelle, Paris (MNHN) and the Thailand Institute of Scientific

and Technological Research, Bangkok (TISTR), as listed in Table 1.

All measurements are in millimetres and were taken using digital

calipers. Cranial and dental nomenclature follows Butler (1948),

Novacek (1986), Frost et al. (1991) and Gould (1995). Dental

notations are indicated in the text in the following manner, with

premaxillary and maxillary teeth denoted by uppercase letters and

mandibular teeth by lowercase: incisor (I/1), canine (C/c), premolar

(P/p), molar (M/m), thus P3 refers to the third upper premolar, i2 to

the second lower incisor.

PHYLOGENETIC ANALYSIS

Cranial, dental, skeletal and external characters were scored for the

new species and H. parvus according to the character transformation

series employed by Frost et al. (1991: 3-15) and added to the

character matrix shown in Frost et al. (1991: appendix 2) see Table

2. Branch and bound analyses were performed using Paup 4.0ba

(Swofford, 1999) set at maximum parsimony, with a maximum trees

setting of 1000 and all characters treated as unordered and of equal

weight. Bootstrap analyses (Felsenstein, 1985) were made to pro-

vide an assessments of confidence limits of nodes, with 1000

replicates of 100 random addition sequence replicates. Bremer

support indices were calculated by increasing the upper bound of the

shortest tree by one step, repeating the branch and bound analysis

and producing a strict consensus tree; the process was repeated,

progressively increasing the length of the suboptimal cladograms by

a single step until all clades of interest no longer occurred on the

consensus tree; the level at which each node collapsed was recorded

(Kitching et al., 1998). Both accelerated (ACCTRAN) and delayed

(DELTRAN) optimizations were used to map character evolution.

The trees obtained were compared with those in Frost et al. (1991)

and the results of the analysis are given below.

RESULTS

Hylomys megalotis, sp. nov.

HOLOTYPE. BMNH 1999.44 (field number 5/99) male, body in

alcohol, skull extracted. Collected 15 January 1999 by M. F.

Robinson.

TYPE LOCALITY. Environs of Ban Muang and Ban Doy, c 18 km

North of Thakhek, Thakhek district, Khammouan Limestone National

Biodiversity Conservation Area, Khammouan Province, Lao Peoples

Democratic Republic, 17°33'15"N 104°49'30"E. Habitat: steep slopes

around the base of massive limestone karst, covered in rock and large

boulders, with an underlying soil base and heavily degraded mixed

deciduous forest, scrub and bamboo. Low lying areas away from the

karst had been cleared for cultivation of paddy rice.

PARATYPES. BMNH 1999.45 (field number 14/99) collected 16

January 1999; 1999.46 (field number 15/99) and 1999.48 (field

number 17/99) collected 17 January 99, females, bodies in alcohol,

skulls extracted; 1999.47 (field number 16/99) collected 17 January

1999, male, skin and internal organs in alcohol, skull and skeleton.

All specimens were collected by M.F. Robinson from the same

locality as the holotype.

DIAGNOSIS

Ears large, rhinarium elongated; first and fifth digits of forefeet long,

claws long, cheiridia large and rounded; cheiridia on hindfeet large,

soles naked; pre-anal gland with single opening. Skull with posterior

region of nasals extending to level of antorbital rim; maxilla and

parietal widely separated by frontal in supraorbital region; long

grooves for palatine artery present in palate; anterior palatine fo-

ramina anterior to maxillary palatine suture; antorbital fossa shallow;

nasolabialis fossa shallow; posteroventral maxillary process of

zygoma distinct; antero-ventral process of alisphenoid present. Den-

tition robust. Third upper premolar (P3) large with well developed

lingual cusp and three roots. Neural spine of axis low.

DESCRIPTION

Medium sized Hylomys with a long tail, approximately 75% of head

and body length. Pelage grey, long, soft and very fine, lacking

flattened spinous hairs; individual hairs grey for most of their length,

then buff with buff or black tips. Dorsal region of rhinarium narrow,

elongate posteriorly; ears prominent, very large, rounded. First and

P.D. JENKINS AND M.F. ROBINSON

fifth digits of forefeet lengthened, claws long and moderately stout;

sole and tarsal region of hindfeet naked, cheiridia large. Pre-anal

gland with single opening immediately posterior to cloaca. Two

pairs of inguinal mammae present.

Skull elongate, moderately slender and somewhat flattened in

appearance (see Figs. 1—2); dorsal profile more or less straight,

showing a gradual increase in height from anterior of rostrum to

braincase. Rostrum long, slender and moderately shallow, nasals

long extending posteriorly to, or slightly beyond, level of antorbital

rim; posterodorsal region of premaxilla widely separated from

anterodorsal region of frontal by maxilla; interorbital region moder-

ately narrow; supraorbital processes of frontals scarcely evident;

frontals anteriorly depressed in midline; supraorbital region of

frontals broad, so that the maxillaries are widely separated from the

parietals; parietals extend anteriorly in supraorbital and orbital

region but do not form an anterior process; supraorbital foramen

present in dorso-orbital region of frontals; orbital region of maxilla

broad, forming major portion of the anterior region of the orbit;

orbital region of frontal constricted anteriorly by maxilla, posteriorly

by parietal; orbitosphenoid anteroposteriorly expanded, optic

foramen posteromedially positioned, anterodorsal to, and moder-

ately well separated from, the suboptical foramen and from the

ethmoid foramen (see Fig. 3); crest present leading from anterior

alisphenoid diagonally across orbitosphenoid, partially obscuring

optic, suboptic and sphenorbital foramina in lateral view; alisphe-

noid dorsoventrally compressed, fusiform anteroventral process of

alisphenoid present, well marked alisphenoid canal present; brain-

case low and scarcely domed, lambdoid crest moderately

well-developed laterally, low medially; mastoid large, slightly in-

flated; paraoccipital process small; infraorbital foramen dorsal to

P4; antorbital or prelacrimal flange present only as a low ridge;

shallow antorbital fossa on anterior surface of zygoma; nasolabialis

fossa shallow; maxillary component of zygoma narrow with long,

slender posteroventral process ventral to well marked long jugal,

slender anterodorsal process of squamosal portion of zygoma over-

lying jugal; palate with paired maxillary foramina level with P2 and

anterior of P3, small paired anterior palatal foramina, lying anterior

to the suture between maxilla and palatine; palatal spine absent;

basioccipital narrow with ridge in midline, tympanic wing of basioc-

cipital slightly inflated. Mandible with deep, moderately broad

coronoid process; mental foramen below p3.

Dental formula: 3/3 1/1 4/4 3/3 = 22. Dentition robust (see Figs. 1—

2). First upper incisor robust, distostyle present; I2 and [3

sub-triangular, anteroflexed, distostyle present, 13 approximately

half size of 12; C with anterior basal cusp and distostyle and two

roots; Pl and P2 subequal in height, P2 longer than P1, both with

anterior basal cusp and distostyle, P1 with two fused roots, P2 with

two roots; P3 large, subequal in height to C, lingual cusp (protocone

according to Gould, 1995) well developed, three roots present; P4,

M1 and M2 quadrate in shape, parastyle well developed, meta-

conule present on M1 and M2; M3 subtriangular in shape, with well

developed parastyle and hypocone and metacone distinct in unworn

dentition. First lower incisor larger than i2, both semi-procumbent,

i2 larger than 13, which is anteroflexed with hypoconulid present; c

anteroflexed, greater in height than i3 and p1; p1 and p2 subequal in

height and both with a single root; p3 larger with two roots; p4 with

well developed paraconid and talonid; m1—m3 with well developed

paraconids, m3 less than half size of m1.

ETYMOLOGY

The name of the new species is derived from the Greek p¢éyac

(megas), large; ®t6c (otos), ear; the ears are large in comparison

with those of other species of Hylomys.

A NEW SPECIES OF HYLOMYS

MUPUARRAROUADOOANODAUOOADOURDORRUORRNORRNORRUUORUOURUUNRUOUNONUDOURUUDRDORURUORRUNURUNURUNURUORRRUND|

10cm

Fig. 1 Crania from left to right of dorsal view of mandible and skull, ventral view of skull, left lateral view of skull and mandible. Top row: Hylomys

megalotis BMNH 1999.47; second row: Hylomys suillus BMNH 1962.711; third row: Hylomys sinensis BMNH 1911.2.1.20; fourth row: Hylomys

parvus BMNH 1919.11.8.12.

\oS)

P.D. JENKINS AND M.F. ROBINSON

Fig. 2 Lateral view of anterior of skull, mandible and dentition of Hylomys megalotis BMNH 1999.44. Scale = 1 mm.

COMPARISON WITH OTHER TAXA

The new species is readily distinguished in external appearance

from all other species of Hylomys. It is similar in body size but with

a considerably longer tail and larger ears (see Table 3). Tail 65-74

% of head and body length in H. megalotis, 51-63 % in H. sinensis,

27-31 % in H. hainanensis, and very short (< 25 %) in H. parvus

and H. suillus. The lack of flattened spinous hairs in the pelage

distinguishes H. megalotis from H. suillus and H. sinensis. The

rhinarium is more extensive posteriorly than in H. sinensis but more

elongate and narrower than in H. suillus. The claws and first and

fifth digits of the forefeet of H. sinensis, H. suillus and H. parvus are

not lengthened as in H. megalotis and the cheiridia are smaller than

in H. megalotis. The sole and tarsal region of the hindfoot are naked

in H. megalotis, differing from the haired soles found in H. sinensis

and H. suillus. The paired pre-anal glands are midway between the

cloaca and the anus in H. sinensis and close to the anus in H. suillus,

so differing from that of H. megalotis, in which the single opening

is positioned immediately posterior to the cloaca.

The skull of H. megalotis is more elongate in appearance than

any of the other species of Hylomys; it is longer, with a longer,

narrower rostrum, longer upper toothrow and the braincase is

shallower relative to its breadth (see (see Fig. 1 and Table 3). The

posterodorsal region of the premaxilla is widely separated from the

anterodorsal region of the frontal by the maxilla in H. megalotis,

narrowly separated in H. hainanensis but in contact or nearly in

contact in H. sinensis, H. suillus and H. parvus (see Table 4). As in

H. sinensis, but unlike other species of Hylomys, the posteriormost

portion of the nasals in the new species extend to the level of the

antorbital rim. Hylomys megalotis has a shallow antorbital fossa

unlike the moderately deep fossa of H. sinensis and H. parvus, and

the deep fossa in H. suillus. The zygoma of H. megalotis differs from

all other species of Hylomys: the maxilla is considerably narrower,

the nasolabialis fossa shallower and the jugal more extensive. As in

Podogymnura aureospinula and Echinosorex, a distinct posteroventral

process is present on the maxillary region of the zygoma of H.

megalotis, indistinct in H. parvus but absent in H. suillus, H. sinensis

and H. hainanensis. The supraorbital process of the frontal of H.

megalotis is poorly defined and blunt, the anterior process of the

parietal absent and the parietal is widely separated from the maxilla

by the frontal in the supraorbital region, as in Podogymnura and

Echinosorex; in H. parvus the supraorbital process of the frontal is

poorly defined and blunt but the anterior process of the parietal is

short but distinct, narrowly separated from the maxilla by the frontal;

in A. suillus, H. hainanensis and H. sinensis the anterior process of

the parietal is distinct, scarcely separated from the maxilla and

contributing to the well marked supraorbital process of the frontal.

The optic and suboptic foramina are well separated in H. megalotis

but lie close together in H. suillus, H. sinensis and H. parvus. An

anteroventral process of the alisphenoid is present in H. megalotis,

unlike all other Hylomys and Podogymnura. As in Podogymnura and

Echinosorex, the palatine foramina are small and positioned anterior

to the maxillary/palatine suture in H. megalotis and long grooves for

the palatine artery are present in the palate, whereas the elongated

palatine foramina in other species of Hylomys lie at the maxillary/

A NEW SPECIES OF HYLOMYS

CORF

SPALF

OPTF

FOV

SOPF

Fig.3 Lateral view of left orbital region of Hylomys megalotis BMNH 1999.44. Scale = 1 mm. Abbreviations: AL — alisphenoid, ALIC — alisphenoid

canal, CORF — cranio-orbital foramen, EF — ethmoid foramen, FOV — foramen ovale, FR — frontal, MX — maxilla, OPTF — optic foramen, OS —

orbitosphenoid, PL — palatal, PR — parietal, PT — pterygoid, SOPF — suboptic foramen, SPALF — sphenopalatine foramen, SPORF — sphenorbital

foramen, SQ — squamosal.

palatine suture and the palatine artery grooves are small or indis-

tinct. The anterior opening of infraorbital canal is dorsal to P3/P4 in

most species of Hy/lomys but dorsal to P4/M1 in H. megalotis, H.

parvus and Echinosorex, and yet more posteriorly positioned in

Podogymnura.

The dentition of H. megalotis is considerably more robust than

that of any of the other species of Hylomys. The dental formula is the

same as in H. suillus and H. parvus and these species are distin-

guished from H. hainanensis, which lacks p1 and H. sinensis, which

lacks P1 and pl. In H. megalotis and H. parvus, P2 has two roots,

unlike the other species which have either one or two well fused

roots. Unlike all other species which lack a lingual cusp, P3 of H.

megalotis has a well developed lingual cusp as in Echinosorex, and

this tooth is large with three roots as in Podogymnura and

Echinosorex, p3 is larger than p2 with two roots as in H. parvus,

Podogymnura and Echinosorex, while p3 is slightly smaller than p2

with one root in A. suillus, H. sinensis and H. hainanensis.

RESULTS OF THE PHYLOGENETIC

ANALYSIS

Forty equally most parsimonious trees were retained in the branch

and bound analysis, 141 steps in length, with a Consistency Index of

0.72, Retention Index of 0.93 and Rescaled Consistency Index of

0.66. In all most parsimonious trees the two subfamilies, Galericinae

and Erinaceinae, readily segregated and the Galericinae further

separated into two distinct groups: a clade comprising Echinosorex

and Podogymnura, the other clade confined to Hylomys. Most of the

variation found among all trees occurred within the Erinaceinae,

since for the Galericinae twenty of the trees showed the configura-

tion seen in Fig. 4a, while the remaining trees all showed the

alternative arrangement for this subfamily (Fig. 4b). That part of the

tree obtained by Frost ef al. (1991: Fig.9) for the Galericinae is

illustrated as part of Fig. 4a. The strict consensus tree (see Fig. 5)

revealed strong bootstrap support (97%) for the Galericinae and for

aclade of H. suillus, H. sinensis and H. hainanensis, and this tritomy

also had a high Bremer support index. There was moderate bootstrap

support (83%) for a clade of H. suillus, H. sinensis, H. hainanensis

and H. parvus, and a clade comprising all species in the genus

Hylomys occurred in 77% of the replicates. The bootstrap support

value for a clade of Echinosorex and Podogymnura was low at only

64%. Within the genus Hylomys, H. megalotis was basal to all other

species. Clades with bootstrap support values less than 50%, respec-

tively of H. sinensis and H. hainanensis (42%), and H. suillus and H.

hainanensis (43%) were considered to be unresolved.

The shared derived character transformations (synapomorphies)

which were revealed by the analysis are recorded below, using the

format of character number quoted from Frost er al. (1991) followed

by character transformation state, where (0) equals the ancestral and

(1) the derived character state.

SYNAPOMORPHIES OF GALERICINAE:

[8.1] Antorbital or pre-lacrimal flange: (0) not developed, lacrimal

canal visible in lateral view; (1) developed so that the lacrimal canal

is obscured in lateral view. CI 1.000.

[10.2] Jugal size; (0) large, reaches lacrimal; (1) small, does not

reach lacrimal; (2) vestigial, confined to lateral rim of zygoma; (3)

absent. In ACCTRAN the transformation was from | — 2, in

DELTRAN the change was from 3 — 2. CI 1.000.

[62.1] P4 lingual roots: (0) one; (1) two unfused; (2) two fused. CI

1.000. This state occurs in all Galericinae but was shown only in

DELTRAN.

P.D. JENKINS AND M.F. ROBINSON

gymnura

aureospinula

truei

sinensis

suillus

hainanensis

parvus

megalotis

Erinaceinae

E. gymnura

P. aureospinula

P. Enwer

H. sinensis

H. hainanensis

ishh Sibi Il Wats)

lel Sen Vea AUNS)

H. megalotis

Erinaceinae

Fig. 4 Comparison of trees obtained for the Galericinae. (a) One of twenty most parsimonious trees, all showing the same configuration for the

Galericinae. Tree length 141 steps, with a Consistency Index of 0.72, a Retention Index of 0.93, and a Rescaled Consistency index of 0.66. The

branching pattern on the left shows the results from the analysis of this study, that on the right is partially redrawn from Frost er al. (1991: fig. 9),

restricted to show only the relationships within the Galericinae and is 128 steps in length with a Consistency Index of 0.76. (b) One of the remaining

twenty most parsimonious trees, showing the alternate arrangement for the Galericinae.

[66.1] M3 hypocone (see Frost. et al. 1991) or metastylar spur (see

Gould, 1995): (0) absent or weak; (1) present, well developed on

buccal side. CI 1.000.

[69.1] Axis, posteroventral keel: (0) absent; (1) present. CI 1.000.

[71.1] Scapula, metacromium process: (0) deltoid, amorphous pro-

jection; (1) long, fusiform projection. CI 1.000.

[72.1] Sacral vertebrae, neural spines: (0) not fused into continuous

longitudinal plate; (1) fused into continuous longitudinal plate. CI

1.000.

[73.1] Ischium, posterodorsal process (see Gould, 1995 for correc-

tion of error by Frost et al. 1991): (0) not greatly elongated; (1)

greatly elongated. CI 1.000.

[74.1] Tibia, lateral flange on antero-superior margin: (0) absent or

weakly present; (1) strongly developed. CI 1.000.

SYNAPOMORPHY OF HYLOMYS:

[19.1] Cranio-orbital foramen in the frontal: (0) closely associated or

joined with the ethmoid foramen; (1) foramina widely separated.

The terminology for this character is confusing. Frost er al. (1991)

used the name ophthalmic foramen (which they attributed to Butler

(1948) although this name could not be found in this paper), but

Gould (1995: character 19) pointed out that this foramen had been

misidentified by Butler and is the anterior opening for the superior

ramus of the stapedial artery. Gould also referred to this foramen as

A NEW SPECIES OF HYLOMYS

£4 (97)

>8 6

EB. gymnura

P. aureospinula

truei

H. sinensis

H. hainanensis

Hee (Stes

H. parvus

H. megalotis

Erinaceinae

Fig. 5 Strict consensus of 40 most parsimonious trees. The branch lengths are shown above the branches followed by the bootstrap support values in

parentheses, representing the percentage of trees containing the specified clades. The Bremer support indices are given below the branches.

the sphenofrontal foramen and McDowell (1958) as the sinus canal.

CI 1.000.

SYNAPOMORPHY OF HYLOMYS SINENSIS, H. HAINANENSIS, H.

SUILLUS AND H. PARVUS:

[4.1] Size of palatal foramina: (0) small; (1) anterior foramina

elongated posteriorly; (2) anterior foramina elongated to include

middle palatine foramina. CI 1.000.

SYNAPOMORPHIES OF HYLOMYS SINENSIS, H. HAINANENSIS, H.

SUILLUS:

{13.1] Supraorbital process of frontal on parietal/frontal suture: (0)

absent or poorly defined; (1) sharp, well defined. CI 1.000.

[16.1] Anterior process of parietal: (0) absent or very weak; (1)

extends anteriorly along the supraorbital rim to form the base of the

supraorbital process. CI 1.000.

[57.1] p3: (0) two roots present, larger in size than p2; (1) one root

present, nearly equal in size to p2; (2) absent. CI 1.000.

The analysis found no autapomorphic characters to define H.

megalotis but the following apomorphies for this species are recorded

as follows:

[1.1] Posteriormost extension of nasals: (0) anterior to the level of

the antorbital rim; (1) medial or posterior to the level of the antorbital

rim. CI 0.333. Homoplasious with H. sinensis but also with

Erinaceinae.

[5.1] Location of the anterior palatine foramina: (0) at the maxilla/

palatine suture; (1) anterior to the maxilla/palatine suture. CI 0.500.

Shown only in DELTRAN, homoplasious with Echinosorex and

Podogymnura.

[17.1] Anterior process of alisphenoid: (0) absent; (1) present. CI

0.500. Homoplasious with Erinaceinae. This character, defined as a

narrow, fusiform anterior process of the orbital wing of the alisphe-

noid is, according to Frost et al. (1991), related to the location of the

sphenopalatine foramen and involved with shortening of the

orbitotemporal region. Gould (1995) commented that the relative

position of the suboptic foramen (her character 21 scored thus: (0)

anterior to the sphenorbital fissure; (1) present in the medial wall of

the sphenorbital fissure; (2) present in the medial wall of the

sphenorbital fissure but hidden within the fissure) seems to be

related to the shortening of the skull in erinaceids. As the skull

shortens, the alisphenoid overlaps the orbitosphenoid, creating a

strong alisphenoid wing [character 17 of Frost et al. (1991) and

Gould (1995)], the degree of overlap seems to be directly related to

the visibility of the suboptic foramen from lateral view and, as

pointed out by Butler (1948) the orbitosphenoid is reduced in size.

While the alisphenoid is more extensive in H. megalotis than in other

galericines, and the suboptic and sphenorbital foramina are partially

concealed in lateral view, the orbitotemporal region is not obviously

shortened. The anterior process in H. megalotis is fully ventral in

location and is actually or nearly in contact with a short posteroventral

process of the maxilla, thus contributing to the ventral floor of the

orbit, however the orbitosphenoid is not reduced in size. It is

possible that this character state in H. megalotis is not homologous

with that of the Erinaceinae and that it actually represents a separate

character transformation, alternatively it is scored incorrectly and

the plesiomorphous condition should be the presence of the anterior

process.

[22.1] Palatal shelf and spine: (0) well developed spine on posterior

palatal shelf; (1) spine absent or vestigial. CI 0.200. Shown only in

DELTRAN, homoplasious with Podogymnura, H. parvus, and

Atelerix.

DISCUSSION

The addition of two taxa to the analysis performed by Frost et al.

(1991) provided broadly similar results in that the Galericinae divided

intotwo main groups: one comprising Echinosorex and Podogymnura,

the second including all five species of Hylomys. The results of the

current phylogenetic analysis lend support to the taxonomy proposed

by Frost et al. (1991) that the three species of Hylomys considered in

their analysis (H. suillus, H. sinensis and H. hainanensis) are correctly

attributed to a single genus rather than the three separate genera

(respectively Hylomys, Neotetracus and Neohylomys) maintained by

Corbet (1988). The additional species of Hylomys however, reduced

the degree of support for the genus and, on this particular morphologi-

cal data set, a considerable degree of homoplasy 1s evident within the

Hylomys clade. There was only one unique synapomorphy for the

Hylomys clade (character 19.1: the wide separation between the

cranio-orbital and ethmoid foramina), two of the other apomorphies

(34.1: the inflation of the mastoid region between the exoccipital and

the squamosal, and 41.1: the expanded exoccipital) showing

homoplasy with P. truei, while the third character state (50.1: upper

canine slightly larger than the adjacent post-canine teeth) is not

shown by H. sinensis (50.2: upper canine approximately equal in

size to the adjacent post-canine teeth). There are no autapomorphies

defining H. megalotis, which shows more plesiomorphy than the

other species of Hylomys; many of its features are homoplasious

with Echinosorex, Podogymnura and Erinaceinae.

Hylomys is a morphologically variable genus, containing species

that are generally well segregated and show little overlap in species

range. Hylomys hainanensis is a geographically isolated island form

and while H. sinensis and H. parvus are each parapatric with H.

suillus in a few areas, Corbet (1988) pointed out that in regions

where H. sinensis and H. suillus occur, H. sinensis is found at greater

altitudes than H. suillus. Similarly Ruedi et al. (1994) showed that

although H. parvus is currently restricted to moss forests at the peak

of Gunung Kerinci, Sumatra at greater altitudes than H. suillus, the

latter occurs elsewhere at greater and lesser altitudes. Both Corbet

(1988) and Ruedi et al. (1994) invoked ecological factors such as

competitive exclusion to explain the altitudinal segregation of these

three species, but did not provide data to support this supposition.

There are few distribution records of H. suillus and H. megalotis in

Lao PDR, which potentially may be sympatric or possibly parapatric

if H. megalotis should prove to be ecologically restricted to the

specific limestone habitat in which it was first collected.

Little is known about the biology of Hylomys. Hylomys sinensis is

believed to be entirely terrestrial and H. swillus mainly so, although

this species has also been observed climbing in low bushes (Lekagul

& McNeely, 1977). Hylomys suillus occurs in hilly or montane

humid forests with dense undergrowth, H. sinensis in cool damp

forests in the cover of runways and under logs and rocks. In their

original description Shaw & Wong (1959) reported that H.

hainanensis spends most of its time in underground burrows and that

the cylindrical body, short tail and claws are adaptations to a

fossorial life. Hylomys parvus is apparently restricted to high alti-

tude moss forest. There is no information about the behavioural

ecology of H. megalotis but the limestone karst where it has been

found to date is an unusual habitat and some of the morphological

features of this species, such as the moderately broad forefeet with

long, fairly stout claws, the long naked hindfeet with large cheiridia,

the moderately long tail and the comparatively flattened braincase

may be adaptations to life in this habitat.

ACKNOWLEDGEMENTS. We would like to thank the staff of WWEF-Thai-

land, the Lao PDR Department of Forestry (DoF) and the Forest Management

and Conservation Project (FOMACOP) for permission to work in Lao PDR

and for organising the project; in particular Bouahong Phanthanousy (National

Project Director, FOMACOP), Bouaphanh Phantavong (Deputy National

Project Director, FOMACOP), Robert Mather and Robert Steinmetz (WWF-

Thailand) and Bruce Jefferies (FOMACOP), who additionally oversaw the

export of specimens. We are very grateful to Maurice Webber for his vital

contribution to all aspects of the fieldwork.

Assistance inthe field was gratefully received from Thongphanh Ratanalangsy

(Khammouan Limestone and DongPhu Vieng NBCA Co-ordinator), Sinnasone

Seangchanthanarong (DoF) and Nousine Latvylay (driver for FOMACOP in

Savannakhet). The headmen and villagers from Pontong, Muang, Mouangkhai,

Louang, Vieng, Kengkhot, Thamtem and Tonglom provided excellent field

support and companionship, and without them this project would nothave been

possible. We would like to thank Angela L. Smith for her help in measuring

specimens at TISTR and for her comments on the manuscript.

P.D. JENKINS AND M.F. ROBINSON

Field work was funded by the Global Environment Facility through the

World Bank to the Government of Lao PDR, via a contract with Burapha

Development Consultants, sub-contracted to WWF-Thailand.

We thank Bob Randal (AMNH), Jacques Cuisin (MNHN) and Nivesh

Nadee (TISTR) for making specimens in their collections available to us. We

are very grateful to Andrew Cabrinovic (NHM) for specimen preparation, to

Harry Taylor, Photographic Unit (NHM) for the excellent photographs and to

Karen Fisher (volunteer, NHM) for her kind assistance with documentation.

We are grateful to Dr Francois Catzeflis, Institut des Sciences del’ Evolution,

Université Montpellier for his constructive review of the manuscript. We

particularly thank Dr Darrell Siebert, NHM for his generous guidance with

the phylogenetic analysis and constructive criticism of the manuscript.

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vores a la surface de la terre; précédées de I’histoire de la science a ce sujet, des

principes de leur classification et de leur distribution géographique actuelle.

Compte Rendus Hebdomadaires de Scéances de |’ Académie des Sciences 6: 738—

744.

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special reference to fossil material in the British Museum. Proceedings of the

Zoological Society, London 118 (2): 446-500.

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ecology and zoogeography. Mammal Review 18 (3): 117-172.

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Gould, G. C. 1995. Hedgehog phylogeny (Mammalia, Erinaceidae) — the reciprocal

illumination of the quick and the dead. American Museum Novitates (3131): 1-45.

Kitching, I. J., Forey, P. L., Humphries, C. J. & Williams, D. M. 1998. Cladistics.

Second edition. The theory and practice of parsimony analysis. Oxford University

Press, Oxford, New York, Tokyo.

Lekagul, B. & McNeely, J. A. 1977. Mammals of Thailand. Kurusapha Ladprao Press,

Bangkok.

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Museum of Natural History 115 (3): 117-214.

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level. Columbia University Press, New York.

Mearns, E. A. 1905. Descriptions of new genera and species of mammals from the

Philippine Islands. Proceedings of the United States National Museum 28: 425-460.

Miiller, S. 1840. Over de Zoogdieren van den Indischen Archipel (No. 2) pp. 9-57. In

C. J. Temminck (ed.) Verhandelingen over de Natuurlijke Geschiedenis der

Nederlandsche overzeesche bezittingen, door de Leden der Natuurkundige commissie

in Indié en andere Schrijvers. A. Amz & Co., Leiden.

Novacek, M. J. 1986. The skull of lepticid insectivorans and the higher-level classi-

fication of eutherian mammals. Bulletin of the American Museum of Natural History

183 (1): 1-111.

Osgood, W. H. 1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic

expeditions. Publications of the Field Musuem of Natural History, Zoology (18):

193-339.

Raffles, T. S. 1822. Descriptive catalogue of a zoological collection, made on account

of the Honourable East India Company, in the island of Sumatra and its vicinity,

under the direction of Sir Thomas Stamford Raffles, Lieutenant-Governor of Fort

Marlborough; with additional notices illustrative of the Natural History of these

countries. Transactions of the Linnean Society, London 13 (1): 239-274.

Robinson, H. C. & Kloss, C. B. 1916. Preliminary diagnoses of some new species and

subspecies of mammals and birds obtained in Korinchi, West Sumatra, Feb.—June

1914. Journal of the Straits Branch of the Royal Asiatic Society (73): 269-278.

Robinson, M. F. 1999. Order Insectivora, pp. 221-223 In J. W. Duckworth, R. E. Salter

& Khounboline, K. (compilers) Wildlife in Lao PDR 1999 status report. TUCN,

Vientiane.

Ruedi, M., Chapusiat, M. & Iskander, D. 1994. Taxonomic status of Hylomys parvus

and Hylomys suillus (Insectivora: Erinaceidae): biochemical and morphological

analyses. Journal of Mammalogy 75 (4): 965-978.

Swofford, D. L. 1999. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other

Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts.

Shaw, T. H. & Wong Song 1959. [A new Insectivore from Hainan]. Acta Zoologica

Sinica 11: 422-425. [In Chinese; English summary].

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Erinaceidae. Annals and Magazine of Natural History (8) 4: 389-391.

A NEW SPECIES OF HYLOMYS

Table 1 Comparative material

Hylomys sinensis

BMNH 1932.4.19.3 Chapa, Tonkin, [ Vietnam]

BMNH 1933.4.1.117—134 Chapa, Tonkin, [Vietnam]

BMNH 1933.4.1.536—541 Chapa, Tonkin, [Vietnam]

BMNH 1911.2.1.15—23 Omi-san, Omi-Hsien, S. Szechwan [Sichuan, PDR

China]

BMNH 1982.205 Omi-san, Omi-Hsien, S. Szechwan [Sichuan, PDR

China]

MNHN 1911-1180-1184 Ta-Tsien-Lou, Setchouen [Sichuan, PDR China]

BMNH 1909. 12.13.1 Ta-Tsien-Lou, Szechuan [Sichuan, PDR China]

BMNH 1911.8.6.1 Yengyuek, Yunnan, [PDR China]

BMNH 1912.7.15.1 Ching-tsai- Yang, Yunnan, [PDR China]

BMNH 1914.10.23.3 Near Yang-fsi, W. Yunnan, [PDR China]

Hylomys suillus

BMNH 1909.7.20.2—3 Sima, Burma [Myanmar]

AMNH 44112 Nam-Ting, Yunnan, China [PDR China]

BMNH 1925.1.1.17 Bao-Ha, Tonkin, [Vietnam]

BMNH 1926.10.4.42 Xieng-Khouang, Laos [Lao PDR]

AMNH 87313 Bologens Plateau, Laos [Lao PDR]

BMNH 1926.10.4.36-41 Dak-to, Annam

MNHN 1929-320-325 Dak-to, Annam

BMNH 1955.1422 Tasan, Chumpawn, Peninsular Siam [Thailand]

TISTR 54-611 Huey Mae Sanam, Chiengmai, Thailand

TISTR 54-613 Trang, Muang, Chong, Thailand

TISTR 54-614—615 Khao Yai National Park, Korat, Thailand

TISTR 54-616 Phu Nam Tok, Saraburi, Thailand

TISTR 54-617 Pok Nam Tok, 21 km from Saraburi, Saraburi, Thailand

TISTR 54-618 Mae Sai, B. Santon Poi, Chieng Rai, Thailand

TISTR 54-1498-1500 Pak Thong Chai, Sakaerat, Nakhon Ratchasima,

Thailand

TISTR 54-1811 Pak Thong Chai, 17 km S of Pak Thong Chai, Nakhon

Ratchasima, Thailand

TISTR 54-1809-1810 Khon San, Pa Phu Khieo, Chaiyaphum, Thailand

TISTR 54-1812 Phu Kradung National Park, Loei, Thailand

BMNH 1960.8.4.7 Ulu Langat Forest Reserve, Kajang, Selangor,

[Malaysia]

BMNH 1961.1158 Ulu Langat Forest Reserve, Kajang, Selangor,

Malaysia

BMNH 1955.1420 Semangko Pass, Pahang, Federal Malay States

[Malaysia]

BMNH 1961.1159 Padang, Jeriau, Fraser’s Hill, Pahang, Malaysia

BMNH 1962.710-711 Jandai Baik, Pahang, Malaysia

BMNH 1912.10.22.7 Pelarit, Perlis, Malay [Malaysia]

BMNH 1955.1421 Pelarit, Perlis, Federal Malay States [Malaysia]

BMNH 1912.10.22.8 Perlis, Malay Peninsula [Malaysia]

BMNH 1955.1423 Jor, Batang Pasang, Perak, Federal Malay States

[Malaysia]

BMNH 1955.1424 Kedah Peak, Federal Malay States [Malaysia]

BMNH 1962.711a Kedah Peak, Kedah, Malaya [Malaysia]

BMNH 1892.9.6.4 Mt. Kina Balu [Malaysia]

BMNH 1895.10.4.3-4 Mt. Kina Balu [Malaysia]

BMNH 1955.661 Mount Kinabalu, British North Borneo [Malaysia]

BMNH 1971.2614—2615 Mt. Kinabalu, N Borneo [Malaysia]

MNHN 1889-37 Mont Kina Balu, Borneo [Malaysia]

MNHN 1893-132—133 Mont Kina Balu, Borneo [Malaysia]

BMNH 1971.2616 Dusan Dankulum, Kinabalu, N Borneo [Malaysia]

BMNH 1971.2617-2618 Tinampoh, Bundu Tuhan rest house, N Borneo

[Malaysia]

BMNH 1919.11.5.7 Korinchi, Sumatra [Sumatera, Indonesia]

AMNH 102532 Seletan, Mocamh Doewa, Sumatra [Sumatera, Indonesia]

AMNH 102533 Seletan, Mocamh Doewa, Sumatra [Sumatera, Indonesia]

AMNH 102534 Seletan, Mocamh Doewa, Sumatra [Sumatera, Indonesia]

AMNH 102820 Lampung, Kalianda, Sumatra [Sumatera, Indonesia]

BMNH 1954.45 Tyibodas, West Java [Jawa, Indonesia]

BMNH 1954.46—-48 Sodong Jerok, Idjen Massif, East Java [Jawa,

Indonesia]

BMNH 1961.1743 Tjemorosewu, Mt. Lawu, Java [Jawa, Indonesia]

AMNH 106111 Java [Jawa, Indonesia]

Hylomys parvus

BMNH 1919.11.5.8—12 Korinchi, Sumatra [Sumatera, Indonesia]

Podogymnura truei

BMNH 1953.659-660 Baclayan, E slopes of Mount Apo, Mindanao,

Philippine Islands

Echinosorex gymnura

BMNH 1914.12.8.101—104 Bankachon, Tenasserim [Myanmar]

BMNH 1955.1452 Changkat Mentri, Perak, Federal Malay States

[Malaysia]

BMNH 1955.1453 Damansara Road, Kuala Lumpur, Selangor, Federal

Malay States [Malaysia]

BMNH 1961.1156 Rontau Panjang, Klang, Selangor, Malaya [Malaysia]

BMNH 1961.1157 Sungei Buloh, Selangor, Malaya [Malaysia]

BMNH 1951.179-180 Mount Dulit, Sarawak, Borneo [Malaysia]

BMNH 1951.181 Tinjar River, Baram District, Sarawak, Borneo

[Malaysia]

BMNH 1971.2613 12 miles N of Kalabakan, Tawau, N Borneo [Malaysia]

P.D. JENKINS AND M.F. ROBINSON

Table 2 Data matrix from Frost et al. (1991: Appendix 2) with the addition of characters scored for two additional taxa, Hylomys megalotis and Hylomys

parvus

Character

Hypothetical ancestor

Echinosorex gymnura

Podogymnura aureospinula

Podogymnura truei

Hylomys sinensis

Hylomys suillus

Hylomys hainanensis

Hylomys megalotis

Hylomys parvus

Hemiechinus aethiopicus

Hemiechinus hypomelas

Hemiechinus micropus

Hemiechinus auritus

Hemiechinus collaris

Mesechinus dauuricus

Erinaceus amurensis

Erinaceus concolor

Erinaceus europaeus

Atelerix frontalis

Atelerix algirus

Atelerix albiventris

Atelerix sclateri

Tenrecoids

Soricoids

Lepticidae (fossil)

111111111122222222223333333333444444444455555555556666666666777777777788

123456789012345678901234567890123456789012345678901234567890123456789012345678901

000000000?000?00000?000000000000000000000000000000000?7000?00?7000000007?77000000000

000010112211010000020010000000001000100000010000000100010000111001101111110000001

00001001121101000002010000000000110010001001000100010111010011100110?711110000001

100101010211100100120100000000001100100010000100020011111111111001101111110000000

000101010211100100120000000000001100100010000000010000011111111001101111110000000

000101010211100100120000000000001100100010000000010000111111111001107?11110000000

100010012211000010120100000000001100100010000000010000000000111001101011110000000

111201102131011011011001104001110011111101101011120001112111121110210000001130200

111201102121011011011001103001110011111101101011120001112010121110210000001130200

111201102121011011011001104001110011111101101011120001112111121110210000001130200

111201102111011011011001102001110000111101101011120001112010121110210000001130300

110201102110011011011001101101011000111101101011120001112010121110210000000120310

110201102111011011011001101011010000111101101011221001112010121110210000000011110

110201102111011011011001101011010000111101101011221001112010121110210000000010110

11020110211101101101100110101101000011110110101122100111201012111021000000001?110

110201102111011011011101111011010000111101101011120001102010121110210000000111110

110201102111011011011101111011010000111101101011120001102011121110210000000110110

110201102111011011011101111011010000111101101011120001102111121110210000000210110

110201102111011011011101111011010000111101101011120001102111121110210000000110110

00000000030001000000000000000000000000000000000000?001000000000000000000000000000

00000000030001000002000000000000000000000000000000?001000100000000000000000000000

000000000000000000000000000000000000000000000000000000000000000000000???000?70???

Table 3. Selected measurements of Hylomys in millimetres to show variation in size and proportions. The mean, standard deviation and range are

provided, with sample size in parentheses.

Character

Head and body length

Tail length

Ratio of tail length to head and body length

Hindfoot length

Ear length

Greatest skull length

Condyloincisive length

Upper toothrow length

Length from I1 to anterior of P4

Ratio of 11—P4 length to upper toothrow length

Rostral breadth

Rostral length

Ratio of rostral length to rostral breadth

Braincase breadth

Braincase height

Ratio of braincase height to braincase breadth

' Measurements from Shaw & Wong (1959).

H. parvus H. sinensis H. suillus H. hainanensis' H. megalotis

107.67 + 4.50 115.75 + 4.83 1325 5.92 136.71 + 8.28 124.23 + 7.68

104-114 (3) 111-124 (6) 120-139 (10) 120-147 (7) 115.6-134.7 (4)

24.00 + 0.82 68.5 + 3.70 2B OOS 09 39.67 + 2.13 86.33 + 3.29

23-25 (3) 63-73 (6) 19-30 (8) 36-43 (6) 82.8-91.3 (4)

0.22 + 0.01 0.57 + 0.05 0.17 + 0.35 0.29 + 0.13 0.70 + 0.04

0.21-0.24 (3) 0.51—0.63 (6) 0.14—0.24 (8) 0.27-0.31 (6) 0.65-0.74 (4)

23.17 + 0.24 25.88 + 1.02 PIES OES 25.14 + 1.62 20.83 + 0.38

23-23.5 (3) 24-27 (6) 20.5—23 (9) 24-29 (7) 20.4—21.3 (4)

18.38 + 0.45 17.71 + 1.41 WSS AAS 22.43 + 0.90

NIE VOD) 18-19 (6) 15-20 (7) 16-19 (6) 20.9-23.2 (4)

32.38 + 0.82 34.45 + 0.80 34.65 + 0.60 37.76 + 1.01

30.55 (1) 31.07-33.64 (10) 33.48-35.61 (7) 33.7-35.3 (4) 36.36-39.23 (4)

30.98 + 0.82 33/3 = O67; 36.96 + 1.04

29.09 (1) 30.05-32.36 (11) 32.88-35.01 (7) 35.66-38.56 (4)

15.21 + 0.42 NG Pas O37) 17.75 + 0.64 17.26 + 0.16 19.89 + 0.47

14.64-15.61(3) 15.64-16.82 (11) —16.70-18.75 (13) ——-17.0-17.50 (5) 19.44—20.64 (4)

8.19 + 0.47 8.94 + 0.44 9.79 + 0.46 11.65 + 0.33

7.54-8.61 (3) 8.15—9.54 (11) 9.05—10.34 (13) 11.40-12.21 (4)

0.54 + 0.02 0.56 + 0.02 0.55 + 0.01 0.59 + 0.01

0.52-0.56 (3) 0.51-0.58 (11) 0.54-0.58 (13) 0.57-0.59 (4)

Slt = (0), 17/ 5.92 + 0.18 6.02 + 0.21 5729 2 0als

4.98—-5.35 (3) 5.59-6.20 (11) 5.57-6.44 (13) 5.06-5.38 (4)

9.85 + 0.49 9.67 + 0.29 10.35 + 0.41 13.57 + 0.43

9.33-10.5 (3) 9.33-10.37 (11) 9.57-10.85 (13) 13.30-14.32 (4)

0.52 + 0.03 0.61 + 0.02 0.59 + 0.02 0.39 + 0.01

0.47-0.55 (3) 0.57-0.65 (11) 0.56—0.63 (10) 0.37-0.41 (4)

13.89 + 0.31 15.60 + 0.86 14.70 + 0.54 14.61 + 0.34

13.29 (1) 13.42-14.45 (11) = 12.97-15.69 (10) ~—-13.9-15.4 (4) 14.16—-15.06 (4)

9.06 + 0.34 9.47 + 0.33 8.75 + 0.19

8.63 (1) 8.55—9.67 (11) 9.33-9.85 (7) 8.49-8.95 (4)

0.65 + 0.03 0.68 + 0.02 0.60 + 0.02

0.65 (1) 0.61—-0.70 (11) 0.63—0.70 (7) 0.58-0.62 (4)

A NEW SPECIES OF HYLOMYS

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Bull. nat. Hist. Mus. Lond. (Zool.) 68(1): 13-18

Issued 27 June 2002

A new species of freshwater crab (Brachyura,

Potamoidea, Potamonautidae) from Principe,

Gulf of Guinea, Central Africa

NEIL CUMBERLIDGE

Department of Biology, Northern Michigan University, Marquette, Michigan 49855, U.S.A.

PAUL F. CLARK

Department of Zoology, The Natural History Museum, Cromwell Road, London, SW7 5BD, U.K.

JONATHAN BAILLIE

Institute of Zoology, Zoological Society of London, Regents Park, London NWI 4RY, UK.

CONTENTS

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SYNOPSIS.

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A new species of freshwater crab of the genus Potamonautes Macleay, 1838 is described from Principe (Democratic

Republic of Sao Tomé and Principe), an island in the Gulf of Guinea off the coast of Central Africa. The specimens were collected

during a recent zoological expedition by the Zoological Society of London. This is also the first record of a freshwater crab on

the island of Principe.

INTRODUCTION

The freshwater crabs reported on here were collected during a

zoological expedition to the island of Principe, made recently by the

Zoological Society of London. The island of Principe, together with

the island of Sao Tomé, constitutes a small independent country

(The Democratic Republic of Sao Tomé and Principe) in the Gulf of

Guinea. Principe is the second in a chain of volcanic islands that

make up the Atlantic Ocean Islands group, that lies off the coasts of

Cameroon, Equatorial Guinea and Gabon. The other islands in this

group are Bioko, SAo Tomé and Annobon. Bioko is closest to the

mainland and has two species of potamonautid freshwater crabs,

Sudanonautes floweri (De Man, 1901) and S. granulatus (Balss,

1929), both of which are also found in nearby Cameroon

(Cumberlidge, 1993, 1995, 1999). Sao Tomé is the third island in the

group, and has one endemic species of freshwater crab, Potamonautes

margaritarius (A. Milne-Edwards, 1886). There are no records of

freshwater crabs occurring on Annobon, the fourth island in the

chain, and the furthest from the mainland.

Until the present report, freshwater crabs were not known to be

present on Principe. The new species from Principe was collected

from streams and nearby land in cloud forest in the remote roadless

highland region in the southwest part of the island. The new species

is compared to Potamonautes anchietae (De Brito-Capello, 1871)

and to P. margaritarius, but differs from each in a number of

important characters (Bott, 1953, 1955, 1964). Although the speci-

mens from Principe are both subadult females, and ideally an adult

male is needed to make a definitive identification, a description is

nevertheless provided here, based on several unique somatic

characters of the specimen. Characters of the gonopods, male abdo-

men, and male chelipeds will be described when more material

(including an adult male) becomes available.

Figures were prepared using a camera lucida, and the specimens

were deposited in The Natural History Museum, London, U.K.

(BMNH). Abbreviations: cw, distance across the carapace at the

widest point; cl, carapace length measured along the median line,

from the anterior to the posterior margin; ch, carapace height, the

maximum height of the cephalothorax); fw, front width measured

along the anterior margin; s, thoracic sternite; e, thoracic episternite;

s4/s5, s4/s5, s5/s6, s6/s7, s7/s8, sternal sulci between adjacent

thoracic sternites; s4/e4, s5/e5, s6/e6, s7/e7, episternal sulci between

adjacent thoracic sternites and episternites; P1—P5, pereiopods 1-5,

al—a6, abdominal segments 1-6, a7, telson of the abdomen.

SYSTEMATIC ACCOUNT

Family POTAMONAUTIDAE Bott, 1970

Genus POTAMONAUTES MacLeay, 1838

Potamonautes principe sp. nov.

(Fig. 1)

MATERIAL EXAMINED

Holotype. BMNH 2001.6907. 1 subadult female, cw 40.5, cl 27.5,

14 N. CUMBERLIDGE, P.F. CLARK AND J. BAILLIE

Fig. 1 | Potamonautes principe sp.nov. Holotype subadult female, cw 40.5 mm, BMNH 2001.6907. A, carapace, dorsal view; B, cephalothorax, frontal

view; C, left third maxilliped; D, abdomen; E, anterior sternum; F, right cheliped, frontal view; G, left cheliped, frontal view; H, carpus and merus of

right cheliped, dorsal view; I, carpus and merus of right cheliped, inferior view. Scale = A, H, I, 13.1 mm; B, D, C, E, 10.5 mm; F, G, 8.3 mm.

A NEW FRESHWATER CRAB FROM PRINCIPE

ch 14.8, fw 10.5 mm, summit of Pico do Principe (01° 34’, 51"N, 07°

22', 57"E), 945 m, coll. J. E. M. Baillie, 26 Sept. 1999.

Paratype. BMNH 2001.6908.1 subadult female, cw 33.2, cl 22.8,

ch 11.6, fw 10.0 mm, summit of Pico do Principe (01° 34’, 51"N, 07°

22', 57"E), 945 m, coll. J. E. M. Baillie, 1 Sept. 1999.

DIAGNOSIS. Postfrontal crest straight, smooth, spanning entire

carapace, ends meeting anterolateral margins at epibranchial teeth.

Exorbital tooth small, pointed; epibranchial tooth represented only

by small granule; anterolateral margin posterior to epibranchial

tooth raised, completely smooth, continuous with posterolateral

margin. Carapace surface completely smooth; highly arched, height

greater than front width (ch/fw 1.5). Pleural (vertical) suture on

carapace sidewall Y-shaped, ends meeting exorbital and epibranchial

teeth. Suborbital margin raised, completely smooth. Ischium of third

maxilliped with deep vertical sulcus. Third sternal sulcus s3/s4

deep, v-shaped, meeting sterno-abdominal cavity. Thoracic epister-

nal sulci s4/e4, s5/e5, s6/e6 and s7/e7 distinct. Anterior inferior

margin of merus of cheliped lined by row of small sharp teeth, with

large pointed tooth near junction with carpus.

DESCRIPTION. Carapace ovoid, wide (cw/fw 3.88), highly arched

(ch/fw 1.45); surface completely smooth semi-circular, urogastric,

transverse branchial grooves faint. Front straight, relatively narrow,

about one-quarter carapace width (fw/cw 0.26), anterior margin

sharply deflexed. Postfrontal crest distinct, smooth, straight, span-

ning entire carapace, consisting of fused epigastric, postorbital

crests; ends of postfrontal crest meeting anterolateral margins at

epibranchial teeth. Anterolateral margin between exorbital,

epibranchial teeth smooth, lacking intermediate tooth; anterolateral

margin posterior to epibranchial tooth raised, completely smooth,

continuous with posterolateral margin. Exorbital tooth small, pointed;

epibranchial tooth represented only by small granule. Suborbital

margin raised, completely smooth. Suborbital, subhepatic,

pterygostomial regions all completely smooth; sidewall divided into

three parts by longitudinal (epimeral) suture (dividing suborbital,

subhepatic regions from pterygostomial region), and by Y-shaped

vertical (pleural) groove (dividing suborbital from subhepatic

regions). Superior ends of Y-shaped vertical groove meeting

exorbital, epibranchial teeth. First thoracic sternal sulcus s1/s2 deep;

second sulcus s2/s3 deep, running horizontally across sternum; third

sulcus s3/s4 deep, v-shaped, meeting sterno-abdominal cavity. Tho-

racic episternal sulci s4/e4, s5/e5, s6/e6 and s7/e7 distinct. Third

maxillipeds filling entire oral field, except for transversely ovate

respiratory openings at superior lateral corners; long flagellum on

exopod of third maxilliped; ischium with deep vertical sulcus.

Epistome prominent, smooth, triangular. Mandibular palp 2-seg-

mented; terminal segment single, undivided, with hair (but no hard

flap) at junction between segments. Subadult female abdomen

subcircular, segments al—a6 of female abdomen quadrate, telson

(a7) a broad triangle with rounded apex; segments a5—a6 broadest.

Major cheliped of subadult female relatively slender, with elon-

gated dactylus and propodus, palm of propodus swollen; fingers of

digits of cheliped with small even teeth, forming long narrow

interspace when closed. First carpal tooth of inner margin of carpus

of cheliped large, pointed; second carpal tooth pointed, half size of

first tooth. Posterior inferior margin of merus of cheliped smooth,

with few small teeth distally; medial inferior margin with row of

small sharp teeth along entire length, large pointed tooth at distal

end; superior surface of merus smooth. Pereiopods P2—P5 slender,

P3 longest, PS shortest. Dactyli of P2—P5 tapering to point, each

bearing four rows of downward-pointing short, sharp spines.

CoLour. Thecolour of the specimens when freshly caught (before

alcohol preservation) was creamy white with a very light purplish

tone, however one specimen was more white than the other. This

coloration was relatively uniform throughout the body including the

dorsal carapace, underside, and legs. The purplish tone was darker

on the carapace just behind the eyes, which turned to an orange

reddish color once the specimens were placed in alcohol.

DISTRIBUTION. This species is known only from the summit of the

Pico do Principe: (01° 34", 51'N, 07° 22', 57"E) at 945 meters, The

Democratic Republic of Sao Tomé and Principe, Gulf of Guinea,

Central Africa.

ETYMOLOGY. The species is named for the island of Principe

where it was collected. The species name principe is a noun in

apposition.

TAXONOMIC REMARKS. The new species is assigned to Pota-

monautes because it possesses the following combination of

characters: the anterolateral margin lacks an intermediate tooth

between the exorbital and epibranchial teeth; the mandibular palp is

two-segmented; and the third maxilliped exopod has a long flagellum.

Potamonautes is a widespread genus of African freshwater crabs

found throughout Africa from Senegal to the Horn of Africa, and

from Egypt to South Africa. Bott’s (1955) revision of the freshwater

crabs of Africa recognised some 34 species in this genus. Since then,

the number of species of Potamonautes has risen to more than 60

(Bott, 1959, 1960, 1964, 1968, 1970; Stewart et al., 1995; Stewart,

1997a,b; Daniels et al., 1998; Cumberlidge, 1999; Corace et al.,

2001). Although Bott (1955) recognised 15 subgenera of

Potamonautes, the authors of the present study prefer to follow

Cumberlidge (1999) and use Potamonautes [sensu lato | for all

species, pending a revision of the entire genus (Cumberlidge, un-

published).

It is not normally good practice to describe a new species from a

subadult female. However, we have decided to establish this taxon

in the light of the distinct nature of the available morphological

characters, and because of the isolated nature of the study area which

may mean that further specimens of P. principe are unlikely to

become available for some time. Characters of the gonopods, adult

male chelipeds, abdomen and sternum are not at present known

because the only specimens of P. principe are subadult females.

Nevertheless, there are a number of unique characters that distinguish

P. principe from other species in the genus.

COMPARISONS WITH OTHER SPECIES. Potamonautes principe is

closest to P. anchietae, a medium-sized species of freshwater crab

from Angola (De Brito-Capello, 1871, Bott, 1953, 1955, 1964). This

species was most recently described and illustrated by Bott (1955, p.

247-249, figs. 24, 76, 77, pl. IX, fig. la—d) as P. (Isopotamonautes)

anchietae. Potamonautes principe and P. anchietae are similar in

that both species have a highly arched carapace, a prominent and

complete postfrontal crest, a pointed exorbital tooth, a small granu-

lar epibranchial tooth, a v-shaped thoracic sternal groove s3/s4, and

a similar-sized first carpal tooth on the carpus of the cheliped.

However, there are a number of characters that distinguish the

specimens from Vissabenguilla, Angola (SMF 1890) described by

Bott (1955) as P. (/.) anchietae from the specimens from Principe

under consideration here.

The carapace height of P. principe is greater than that of P.

anchietae (ch/fw P. principe 1.45, P. anchietae 1.19), and the frontal

margin of P. principe is narrower than that of P. anchietae (fw/cw P.

principe 0.26, P. anchietae 0.39). Further, the anterolateral margins

of the carapace of P. principe are completely smooth and lack teeth

of any kind, whereas these margins in P. anchietae are distinctly

granular. The medial inferior margin of the merus of the cheliped of

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A NEW FRESHWATER CRAB FROM PRINCIPE

P. principe has a row of distinct pointed teeth along its length,

whereas this margin in P. anchietae, lacks teeth, and is either

granular or smooth. Finally, the suborbital and pterygostomial regions

of P. principe are smooth, whereas these regions are distinctly

granular in P. anchietae.

Bott (1953, 1955, 1964) described two subspecies of P. anchietae:

P. (1.) a. biballensis Rathbun, 1905, and P. (I.) a. machedoi Bott,

1964, both of which are found in Angola. However, comparison of

the types of these taxa with P. principe indicates that both differ

substantially from the new species described here. For example, the

epibranchial tooth of P. (I.) a. biballensis is large and pointed

whereas that of P. principe is small and granular, and the carapace of

P. (1.) a. biballensis is flattened, whereas that P. principe is high (ch/

fw P. principe 1.45, P. (1.) a. biballensis 1.25). Similarly, the

epibranchial tooth of P. (/.) a. machedoi is large and pointed whereas

that of P. principe is small and granular, and the frontal margin of P.

principe is narrower than that of P. (/.) a. machedoi (fw/cw P.

principe 0.26, P. (I.) a. machedoi 0.33).

Potamonautes principe was also compared here with a specimen

of P. margaritarius from Sao Tomé (SMF 2668), and the two taxa

can be distinguished by the following characters. The carapace of

the latter species is distinctly flattened (ch/fw P. margaritarius 0.95,

P. principe 1.45), the anterolateral margin of P. margaritarius

behind the epibranchial tooth is clearly toothed (whereas this margin

is completely smooth in P. principe), and the ischium of the third

maxilliped of P. margaritarius is smooth and lacks a vertical suture

(whereas this suture is deep in P. principe).

ECOLOGICAL NOTES

The island of Principe is ovoid in outline, with a total land area of

about 139 sq km (Fig. 3). The highest point is Pico de Principe

(945 m), which is a volcanic mountain whose lower slopes are

vegetated by lowland rain forest that grades into cloud forest at

higher elevations. The climate of Principe is tropical, hot, and

humid, and there is a single rainy season from October to May.

Rainfall levels are high, averaging around 1,000 mm per year in

the north and 5,000 mm per year in the south, and monthly tem-

peratures range between 25° and 31° C (Bredero et al., 1977). The

two specimens were collected from the summit of the Pico do

Principe at 945 meters above sea level on two separate occasions.

The terrain at the summit of the Pico do Principe is volcanic and

mountainous, and the vegetation cover is cloud forest with small

stunted trees, and a high abundance of epiphytes and bryophytes.

The thick cloud cover and regular rainfall keep the soil and leaf

litter very damp. On both occasions when crabs were collected the

weather was extremely damp and the mountain top engulfed in

cloud cover. Both specimens were collected on land from under

damp and decaying leaf litter. There was no river or body of water

near the location where the specimens were found. The nearest

stream was observed at lower altitudes (830 m), but there were

numerous temporary small pools at the summit which form after

heavy precipitation. It is likely that P. principe can breathe air,

given its extremely terrestrial lifestyle. When crabs were disturbed

by removing the leaf litter cover, they ran rapidly across the forest

floor and took cover under leaf litter or any available crevasse.

Specimens were collected close to holes near the Pico do Principe,

but crabs were not actually observed moving in or out of these

holes, and it ramains possible that the holes may not have been dug

by the crabs.

ACKNOWLEDGMENTS. Gilles Joffroy and Tariq Stevart (Université libre

de Bruxelles) assisted in the collection of specimens. Phil Crabb (NHM

Photo. Unit) took the photographs reproduced in this paper.

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1995. Redescription of Sudanonautes floweri (De Man, 1901) (Brachyura:

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Paris, 1-382.

Daniels, S. R., Stewart, B. A. & Gibbons, M. J. 1998. Potamonautes granularis sp.

noy. (Brachyura: Potamonautidae), a new cryptic species of river crab from the

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South Africa 5; being a Portion of the Objects of Natural History Chiefly Collected

during an Expedition into the Interior of South Africa, under the Direction of Dr.

Andrew Smith, in the Years 1834, 1835, and 1836; Fitted Out by ‘The Cape of Good

Hope Association for Exploring Central Africa.’ In: A. Smith, Illustrations of the

Zoology of South Africa; Consisting Chiefly of Figures and Descriptions of the

Objects of Natural History Collected During an Expedition into the Interior of South

Africa, in the Years 1834, 1835, and 1836; Fitted Out by ‘The Cape of Good Hope

Association for Exploring Central Africa.’ (Invertebrates): 1-75.

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followed by some remarks on an allied species. Proceedings of the Zoological

Society of London, 1901: 94-104.

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recueillis par M. de Brazza dans les régions du Congo. Bulletin de la Société

Philosophique Paris séries 7(10): 148-151.

Rathbun, M. J. 1905. Les crabes d’eau douce (Potamonidae). Nouvelles Archives du

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Stewart, B. A. 1997a. Morphological and genetic differentiation between populations

of river crabs (Decapoda: Potamonautidae) from the Western Cape, South Africa,

with a taxonomic re-examination of Gecarcinautes brincki. Zoological Journal of

the Linnean Society 199: 1-21.

1997b. Biochemical and morphological evidence for a new species of river crab

Potamonautes parvispina (Decapoda: Potamonautidae). Crustaceana 70: 737-753.

. Coke, M. & Cook, P. A. 1995. Potamonautes dentatus, new species, a fresh-water

crab (Brachyura: Potamoidea: Potamonautidae) from KwaZulu-Natal, South Africa.

Journal of Crustacean Biology 15: 558.

18 N. CUMBERLIDGE, P.F. CLARK AND J. BAILLIE

Principe

Gulf of Guinea

Bonbon

1° 40'N

Santo Antonio

Porto R

Bahia do Oeste ono Rest

@ Terriro Velho

(os

Maria Correia

( 1°35'N

Pico do Principe

Antonio Enes @

ay Boné do Joquei

7° 20'E TPI5'E

Fig. 3 Map of Principe showing the type locality of Potamonautes principe sp.nov.

\ x (252 644 eo

Bull. nat. Hist. Mus. Lond. (Zool.) 68(1); 19-26

A Issued 27 June 2002

Two new species of the Indo-Pacific fish genus

Pseudoplesiops (Perciformes,

Pseudochromidae, Pseudoplesiopinae)

ANTHONY C. GILL

Fish Research Group, Department of Zoology, The Natural History Museum, Cromwell Road, London SW7

SBD, U.K.

ALASDAIR J. EDWARDS

Tropical and Coastal Management Studies, Department of Marine Sciences and Coastal Management,

University of Newcastle, Newcastle upon Tyne NEI 7RU, U.K.

SYNOPSIS. Pseudoplesiops immaculatus is described from 72 specimens from throughout the West Pacific and the eastern and

central Indian Ocean. It is distinguished from congeners in having, in combination, a prominent intermandibular flap and an

unspotted operculum. Pseudoplesiops occidentalis is described from five specimens from the Maldive Islands, central Indian

Ocean. It is distinguished from congeners in having, in combination, II,23 dorsal-fin rays, 26—28 scales in lateral series, and scales

with distinct centres and radii in all fields.

INTRODUCTION

The genus Pseudoplesiops Bleeker was recently diagnosed to

included pseudoplesiopine pseudochromids with the medial lami-

nae of the pelvic bones expanded dorsally (Gill & Edwards, 1999).

Aside from this synapomorphy, members of the genus are distin-

guished from other pseudoplesiopines in possessing the following

combination of external characters: scales in lateral series 26—42;

lower lip complete (uninterrupted at symphysis); and preopercular

pores usually 7 (rarely 6 or 8), with a pore present at the upper

terminus of the preopercle. The genus is mostly confined to the

eastern Indian and Pacific Oceans, but two undescribed species

occur in the Maldive Islands, central Indian Ocean. Although it is

our intention to produce a revision of the genus, we herein describe

the two Maldives species in order to make their names available for

a forthcoming guide to western and central Indian Ocean fishes.

MATERIALS AND METHODS

Institutional abbreviations follow Leviton et al. (1985). All meas-

urements to the snout tip were made to the midanterior tip of the

upper lip. Length of specimens are given in mm standard length

(SL), which was measured from the snout tip to the middle of the

caudal peduncle at the vertical through the posterior edge of the

dorsal hypural plate. Head length was measured from the snout tip to

the posteriormost edge of the opercular membrane. Snout length

was measured over the shortest distance from the snout tip to the

orbital rim, without constricting the fleshy rim of the latter. Orbit

diameter was measured as its fleshy horizontal length. Interorbital

width was measured as the least fleshy width. Upper jaw length was

measured from the snout tip to the posterior edge of the maxilla.

Predorsal, preanal and prepelvic lengths were measured from the

snout tip to the base of the first spine of the relevant fin. Body width

was measured between the posttemporal pores. Caudal peduncle

length was measured from the base of the last anal-fin ray to the

ventral edge of the caudal fin at the vertical through the posterior

edge of the ventral hypural plate. Caudal peduncle depth was

measured obliquely between the bases of the last dorsal- and last

anal-fin rays. Measurements of fin rays excluded any filamentous

membranes. Pectoral fin length was measured as the length of the

longest middle ray. Caudal fin length was measured as the length of

the lowermost ray on the dorsal hypural plate.

Counts of dorsal-, anal- and pelvic-fin spines (unsegmented rays)

and segmented rays are presented, respectively, as Roman and

Arabic numerals. If the last dorsal- or anal-fin ray was divided at its

base it was counted as a single ray. Counts of branched, segmented

rays in the dorsal and anal fins included unbranched rays behind the

first branched ray. A value was not recorded if, due to tip damage, a

branched or unbranched condition could not be determined for the

segmented ray preceding the anteriormost branched ray. As in most

actinopterygian fishes, the upper ray in the pectoral fin is rudimen-

tary and rotated so that its asymmetrical medial and lateral hemitrichs

appear to represent two separate rays; these were counted as a single

ray. Procurrent caudal-fin ray counts were of the rays above (“up-

per’) and below (‘lower) the principal caudal-fin rays. The uppermost

principal caudal-fin ray was defined as the ray articulating with

hypural 5, and the lowermost principal caudal-fin ray was the ray

articulating with the cartilage nubbin between the distal tips of the

parhypural and the haemal spine of preural centrum 2. All

pseudoplesiopine species normally have 17 (9 + 8) principal caudal-

fin rays.

Counts of ‘scales in lateral series’ were of the posteroventrally

oriented transverse scale rows on the midside, beginning with the

row through the tubed scale at the branchial opening and ending with

the row through the scale at the midposterior edge of the hypural

plate. “Scales in transverse series’ were counted anterodorsally from

the anal-fin origin to the dorsal-fin base. Circumpeduncular scales

were counted in a zig-zag fashion around the middle of the caudal

peduncle. Gill-raker counts were of the outer rakers on the first arch,

including rudiments; the angle raker is included in the lower-limb

(second) count. Counts of pseudobranch filaments included all

rudiments. Nomenclature of head pores follows Winterbottom

(1986), as modified by Gill et al. (2000).

Counts of vertebrae are presented in the form precaudal + caudal

= total. Caudal vertebrae are defined as those with a haemal spine,

and include the terminal urostylar complex (which was counted as a

single vertebra). The pattern of insertion of supraneural (predorsal)

bones and anterior dorsal-fin pterygiophores within interneural

spaces is given as an ‘anterior dorsal-fin pterygiophore formula’

modified from the “predorsal formula’ of Ahlstrom et al. (1976).

Each supraneural is represented by an ‘S,’ neural spines are repres-

ented by slashes, and pterygiophores are represented by °3’

(indicating a pterygiophore that bears two supernumerary rays and

a serially associated ray), ‘2’ (indicating a pterygiophore that bears

a supernumerary ray and a serially associated ray) or “1” (indicating

a pterygiophore that bears only a serially associated ray). An

‘anterior anal-fin pterygiophore formula’ is also presented, and is

similar to the anterior dorsal-fin pterygiophore formula, except that

the slashes represent haemal spines. Osteological features were

determined from x-radiographs and from cleared-and-stained speci-

mens, which were prepared following the methods of Taylor & Van

Dyke (1985).

Counts and measurements are given as values or value ranges for

all type specimens, followed, where different, by values for the

holotype in parentheses. Where counts were recorded bilaterally,

both counts are presented for the holotype, separated by a slash; the

first count given is the left count.

Pseudoplesiops immaculatus sp. nov.

Bearded Dottyback

Figures 1, 2

Pseudoplesiops typus [non Bleeker, 1858]; Bleeker, 1875: 31 (in

part, specimen from Amboina).

Chilidichthys [sic] sp. 1; Allen & Steene, 1979: 26 (Christmas Island

and Cocos Keeling Atoll, Indian Ocean).

Pseudoplesiops revellei [non Schultz, 1953]; Kailola, 1987: 244

(Papua New Guinea); Allen & Steene, 1988: 180, fig. 145 (Christ-

mas Island, Indian Ocean); Paxton et al., 1989: 521 (list; Great

Barrier Reef).

Pseudoplesiops sp.; Gill in Randall et al., 1990: 131 (description;

distribution; col. fig.); Randall & Anderson, 1993: 15 (Maldive

A.C. GILL AND A.J. EDWARDS

Islands); Gill in Randall et al., 1997: 131 (description; distri-

bution; col. fig.); Kulbicki & Williams, 1997: 14 (Ouvéa Atoll,

New Caledonia).

Pseudoplesiops n. sp.; Allen & Smith-Vaniz, 1994: 10 (Cocos

(Keeling) Islands).

Pseudoplesiops sp. 1.; Gill & Edwards, 1999: 144 (list of osteologi-

cal materials); Gill, 2000: 2560 (key).

HOLOTYPE. AMS I.20757-069, 29.8 mm SL, Great Barrier Reef,

W end of Raine Island (11°36'S 144°O1'E), coral gutter, 2-20 m,

AMS and Australian Institute of Marine Science team, 13 February

1979.

PARATYPES. AMS I.17090-043, 1: 26.5 mm SL, Papua New

Guinea, Madang Harbour, S edge of Massas Island, coral reef, 1.5—

14 m, B.B. Collette et al., 31 May 1970; AMS I.17094-013, 1: 28.5

mm SL, Papua New Guinea, Trobriand Islands, N end of Kiriwinna

Island, 0-5 m, B.B. Collette and B. Goldman, 7 June 1970; AMS

I.17094-014, 1: 30.4 mm SL, collected with AMS I.17094-013;

AMS 1I.17096-008, 1: 27.8 mm SL, Papua New Guinea, Trobriand

Islands, Kiriwinna Island, off N coast of Tawai Point, coral with

sand patches, 7.5 m, B.B. Collette, 8 June 1970; AMS 1.20756-014,

1: 25.7 mm SL (subsequently cleared and stained), Great Barrier

Reef, Great Detached Reef, over coral and sand, 2-8 m, AMS and

Australian Institute of Marine Science team, 11 February 1979;

AMS 1I.20756-124, 2: 15-30 mm SL, collected with AMS I.20756-

014; AMS 1.20757-039, 1: 29.4 mm SL, collected with holotype;

AMS 1.20757-091, 3: 27.6-30.5 mm SL, collected with holotype;

AMS 1.20784-040, 2: 17.0-21.9 mm SL, Great Barrier Reef, Yonge

Reef, 1 mile N of platform, back reef knoll, 1-15 m, D.F. Hoese et

al., | December 1978; AMS I.21972-004, 1: 31.2 mm SL, Solomon

Islands, Guadalcanal, 12 km W of Honiara, over wreck, J.E. Randall,

12 July 1975; AMS 1.22616-044, 1: 20.9 mm SL, Great Barrier

Reef, Escape Reef, coral reef, 5-18 m, J.R. Paxton et al., 2 Nov-

ember 1981; AMS I.22619-025, 1: 26.0 mm SL, Great Barrier Reef,

Escape Reef North (15°49'S 45°50'E), sand slope, coral and sand in

cave under bommie, 19-22 m, J. Paxton et al., 3 November 1981;

ANSP 131727, 2: 27.2-27.5 mm SL, Cocos Keeling Islands, West

Island, off N end outside breakers (12°07'50"S 96°48'55"E),

Fig. 1 Pseudoplesiops immaculatus, AMS 1.20757-069, 29.8 mm SL, holotype, Raine Island, Great Barrier Reef. (Photo by P. Crabb.)

TWO NEW SPECIES OF PSEUDOPLESIOPS

eer ie

Fig. 2 Cephalic laterosensory pores of Pseudoplesiops immaculatus, AMS 1.20757-069, 29.8 mm SL, holotype, Raine Island, Great Barrier Reef. AIOP,

anterior interorbital pore; AN, anterior nostril; DENP, dentary pores; IMF, intermandibular flap; ITP, intertemporal pore; LSCP, pore in tubed lateral-line

scale (other details of scale omitted); NASP, nasal pores; PARP, parietal pores; PN, posterior nostril; POPP, preopercular pores; PTP, posttemporal pore;

SOBP, suborbital pores; SOTP, supraotic pores. Scaled areas shown in stipple. Scale bar = 2 mm.

0.5—1.3 m relief with sand channels, soft and stony corals, W.F.

Smith-Vaniz et al., 24 February 1974; ANSP 131728, 1: 30.6 mm

SL, Cocos Keeling Islands, West Island, ca. 1 km NNW of N end of

island (12°07'20"S 96°49'05"E), rocky patch fronting on sand, W.F.

Smith-Vaniz et al., 1 March 1974; ANSP 178041, 1: 30.8 mm SL,

Papua New Guinea, Bougainville Island, Tautsina Island, E of Kieta

Peninsula, coral reef off stack at N end of island, 0.9-7.5 m, 11

March 1965 (Te Vega Expedition Cruise no. 6, Station 247); ASIZT

57016, 1: 25.8 mm SL, Taiwan, J.P. Chen, 22 April 1994; ASIZT

56991, 1: 19.7 mm SL, Taiwan, 9 m, J.P. Chen, 21 April 1994;

BMNH 1974.5.25.973, 1: 22.8 mm SL, Papua New Guinea,

Trobriand Islands, Kiriwinna Island, E shore NW of Gusaweta,

exposed coral platform, 0—3.6 m, B.B. Collette, 17 June 1970; CAS

58686, 2: 26.3—-27.9 mm SL, Maldive Islands, Malé Atoll, reef off

SE side of Funidu Islet, inside atoll lagoon (04°11'00"N

073°30'30"E), M.G. Bradbury, 6 November 1964; MNHN 1994-45,

1: 22.6 mm SL, New Caledonia, Loyalty Islands, Ouvéa Atoll,

Bagaat Islet (20°37'18"S 166°16'08"E), vertical reef wall with small

cave and base of wall with coral rubble platform, 15—21 m, J.T.

Williams, J.-L. Menou and P. Tirard, 16 November 1991; QM

1.15527, 2: 26.8-33.8 mm SL, collected with holotype; RMNH

31188, 1: 28.7 mm SL, Ambon (Amboina); RUSI 35701, 1: 30.9

mm SL, Taiwan, off Houpihu, P.C. Heemstra, 20 January 1988;

USNM 209591, 3: 25.3-27.0 mm SL, Indonesia, point E of Tandjung

Naira, Haruka Island, surge channel, 4.5 m, V.G. Springer and M.F.

Gomon, 15 January 1973; USNM 209965, 1: 23.7 mm SL, Indone-

sia, Saparua, two stations mixed (isolated coral head surrounded by

crinkly calcareous matrix at 9 m and coral patch in 3.6 m), V.G.

Springer and M.F. Gomon, 18 January 1973; USNM 290118, 6:

17.4-28.9 mm SL, Papua New Guinea, Hermit Islands, Amot

Island, ocean side of reef at drop off (01°33'S 144°59'E), 0-15.2 m,

V.G. Springer et al., 30 October 1978; USNM 290327, 5: 27.7-29.5

mm SL, Papua New Guinea, S tip of Massas Island (05°10'18"S

145°51'24"E), 0-24 m, V.G. Springer et al., 6 November 1978;

USNM 290437, 1: 31.1 mm SL, Papua New Guinea, Ninigo Islands,

just SE of Ami Island (01°14'S 144°22'B), patch reef behind reef, 0—

4.5 m, V.G. Springer et al., 22 October 1978; USNM 290749, 3:

23.3-29.9 mm SL, Indonesia, Banda Islands, just W of N tip of

Great Banda Island (04°30'30"S 129°56'10"), 0-18 m, V.G. Springer

and M.F. Gomon, 9 March 1974; USNM 290791, 1: 30.3 mm SL,

Indonesia, Ambon, Latuhalat, Namalatu, about 150 m offshore

(03°47'S 128°06'E), 10.5-18 m, V.G. Springer et al., 14 March

1974; USNM 290808, 1: 22.0 mm SL, collected with USNM

290749; USNM 291609, 1: 25.7 mm SL, Philippine Islands, Batanes,

Batan Island, past Mahate, White Beach (20°24'45"N 121°55'00"E),

coral and encrusted boulders, 9-12 m, G.D. Johnson and W.F.

Smith-Vaniz, 1 May 1987; USNM 292031, 1: 27.8 mm SL, Papua

New Guinea, Hermit Islands, N side of W entrance (01°30'30"S

144°59°15"E), 0-12 m, V.G. Springer et al., 4 November 1978;

USNM 322986, 1: 31.8 mm SL, New Caledonia, Loyalty Islands,

Ouvéa Atoll, Motu Veiloa Islet (20°26'06"S 166°28'30"E), reef

crest, small patch reef of mostly dead coral surrounded by rubble,

0.6-3.3 m, J.T. Williams and M. Kulbicki, 17 November 1991;

USNM 322994, 1: 32.1 mm SL, New Caledonia, Loyalty Islands,

Ouvéa Atoll, Récif Draule (20°34'12"S 166°14'12"E), large depres-

sion on submerged reef crest, coral and rubble, 5 m, J.T. Williams, P.

Tirard and J.L. Menou, 16 November 1991; USNM 328198, 1: 24.5

mm SL, collected with MNHN 1994-45; USNM 356587, 1: 20.9

mm SL, Vanuatu, Shepherd Islands, Judy Reef off NW tip of Tongoa

Island (16°52'30"S 168°31'30"E), coral reef with extensive coral

development, J.T. Williams et al., 9 June 1996; USNM 357981, 1:

26.8 mm SL, Solomon Islands, Santa Cruz Islands, Reef Islands,

Fenualoa Island, just W of Nota Point (ca. 10°16'30"S 166°16'30"E),

coral reef face and outer slope, rich coral growth with sand and

rubble, 0-13 m, J.T. Williams et al., 18 September 1998; USNM

358382, 1: 20.4 mm SL, Solomon Islands, Santa Cruz Islands, Reef

Islands, Lomlom Island, Nialo Point on E side of Forrest Passage

(10°16'S 166°18'30"E), vertical reef wall and rocky surge channels

at surface, 0-35 m, J.T. Williams er al., 18 September 1998; WAM

P.26083-041, 1: 31.2 mm SL, Indian Ocean, Christmas Island, Ethel

Beach, 3-6 m, G.R. Allen and R.C. Steene, 19 May 1978; WAM

P.26085-030, 1: 25.9 mm SL, Indian Ocean, Christmas Island, Ethel

Beach, 15-20 m, G.R. Allen and R.C. Steene, 20 May 1978; WAM

P.26093-015, 1: 27.3 mm SL, Indian Ocean, Christmas Island, 1 km

W of Margaret Beach, 10 m, G.R. Allen and R.C. Steene, 25 May

1978; WAM P.26104-005, 1: 27.3 mm SL, Indian Ocean, Christmas

Island, Flying Fish Cove (10°29'S 105°40'E), 6—8 m, G.R. Allen and

R.C. Steene, | June 1978; WAM P.26107-004, 1: 24.0 mm SL,

Indian Ocean, Christmas Island, Rhonda Beach (10°29'S 105°40'E),

6-7 m, G.R. Allen and R.C. Steene, 2 June 1978; WAM P.26113-

003, 2: 25.6-28.4 mm SL, Indian Ocean, Christmas Island, Winifred

Beach (10°29'S 105°40'E), 12-14 m, G.R. Allen and R.C. Steene, 6

June 1978; WAM P.27825-034, 3: 17.6-25.4 mm SL, Papua New

Guinea, Manus Island, Los Negros Island, SE point at aerodrome,

10-40 m, G.R. Allen and R. Knight, 5 October 1982; WAM P.29626-

001, 1: 31.2 mm SL, Papua New Guinea, Port Moresby (09°30'S

147°10'E), 5—6 m, P. Colin, 24 February 1987; WAM P.29927-004,

1: 23.9 mm SL, Indian Ocean, Cocos-Keeling Islands, Direction

Island (12°05'S 096°53'E), 0.1-2.0 m, G.R. Allen, 24 February

1989.

DIAGNOSIS

A species of Pseudoplesiops with the following combination of

characters: prominent intermandibular flap present; and operculum

immaculate, without large dark spot.

DESCRIPTION. (Based on 72 specimens, 15.0—33.8 mm SL) Dor-

sal-fin rays 1,26—28 (1,27), last 3-11 (8) segmented rays branched;

anal-fin rays I-II,16—18 (1,17), last 3-9 (5) segmented rays branched;

pectoral-fin rays 15—18 (16/15), upper 1—5 (2/2) and lower 1-3 (1/1)

rays simple; pelvic-fin rays I,3-4 (1,4), all segmented rays simple;

principal caudal-fin rays 9-10 + 8 (9 + 8), the uppermost 1—2 (1) and

lowermost 1—2 (1) rays unbranched; upper procurrent caudal-fin

rays 3—5 (4); lower procurrent caudal-fin rays 2-4 (3); total caudal-

fin rays 23—25 (24); scales in lateral series 32—39 (36/35); predorsal

scales 7—12 (10); transverse scales 14—17 (16/16); scales behind eye

1-2 (2); scales to preopercular angle 3-4 (3); circumpeduncular

scales 16-17 (16); ctenoid scales beginning at 10-16 (14/14) trans-

A.C. GILL AND A.J. EDWARDS

verse scale rows behind branchial opening; gill rakers 14 + 6-10 =

7-12 (2 + 8); pseudobranch filaments 5-8 (6).

Head pores (all bilaterally paired; Fig. 2): nasal pores 2—3 (2/2);

anterior interorbital pores 1; posterior interorbital pores 0; supraotic

pores 2; suborbital pores 7—9 (8/8); posterior otic pores 0;

preopercular pores 7—8 (7/7); dentary pores 4; intertemporal pores 1;

anterior temporal pores 0; posttemporal pores 1; parietal pores 2.

As percentage of standard length (based on 34 specimens, 22.8—

33.8 mm SL): body depth at dorsal-fin origin 22.8-26.3 (23.8);

greatest body depth 23.1—28.1 (26.5); body width 11.5—13.3 (12.4);

head length 28.8-32.6 (29.9); snout length 5.3-6.2 (6.0); orbit

diameter 7.7—9.9 (8.1); interorbital width 2.2—3.6 (3.0); upper jaw

length 10.0—11.2 (10.1); depth of caudal peduncle 13.2—15.3 (14.8);

caudal peduncle length 7.3—9.4 (8.1); predorsal length 29.5-32.6

(29.5); preanal length 54.7—57.6 (56.7); prepelvic length 26.4—29.2

(28.2); length of first segmented dorsal-fin ray 8.7—10.8 (9.7);

length of third from last segmented dorsal-fin ray 16.4—20.8 (17.4);

dorsal-fin base length 60.5—67.8 (63.1); length of first segmented

anal-fin ray 9.3—12.3 (11.1); length of third from last anal-fin ray

13.6—20.4 (17.8); anal-fin base length 33.3-37.7 (35.6); caudal fin

length 20.4—26.9 (24.8); pectoral fin length 18.4—21.8 (18.5); pelvic

fin length 25.7—32.0 (27.8).

Lower lip complete; prominent intermandibular flap present

(Fig. 2); fin spines weak and flexible; anterior dorsal-fin

pterygiophore formula S/S/S + 2/1 + 1, S/S/SY + 2/1 + 1 or S/S/2/

1 + 1 (S/S/SY + 2/1 + 1); 20-22 (21) consecutive dorsal-fin

pterygiophores inserting in 1:1 relationship directly behind neural

spine 4; anterior anal-fin pterygiophore formula 2 + 1/1, 3 + 1/1 or

2+ 1 +4 1/1 (2 + I/1); 11-12 (11) consecutive anal-fin

pterygiophores inserting in 1:1 relationship directly behind hae-

mal spine 2; second segmented pelvic-fin ray longest; caudal fin

rounded to truncate or slightly emarginate; scales without distinct

centres, and with radii confined to anterior field; dorsal and anal

fins without distinct scale sheaths, although sometimes with inter-

mittent scales overlapping fin bases; anterior lateral line

represented by single tubed scale at branchial opening, followed

by intermittent series of centrally pitted scales, which terminate at

vertical through base of segmented dorsal-fin ray 21-27, or extend

slightly past end of dorsal fin (extending slightly beyond end of

dorsal fin/to vertical through base of segmented dorsal-fin ray 26

in holotype); second intermittent series of centrally pitted scales

originating on midside above anterior part of anal fin, extending

on to middle of caudal-fin base; additional centrally pitted scales

present on caudal-fin base, pits usually aligned vertically on pos-

terior part of scale sheath; scales present on cheeks (not extending

posteriorly over upper part of preopercle) and operculum (Fig. 2);

predorsal scales extending anteriorly to supratemporal commis-

sure (Fig. 2); vertebrae 12 + 20-22 (12 + 21); epurals 2; epineurals

present on vertebrae | through 17—21 (1 through 19); ribs present

on vertebrae 3 through 11-12 (3 through 12), rib on ultimate

precaudal vertebra very small to moderately small or absent.

Upper jaw with 2-6 pairs of curved, enlarged caniniform teeth

anteriorly, the medial pair smallest, and 3-4 (at symphysis) to 1—2

(on sides of jaw) irregular inner rows of small conical teeth, the teeth

of outer row of conical teeth largest; lower jaw with 2-4 pairs of

curved, enlarged caniniform teeth, the medial pair smallest, and 2—

4 (at symphysis) to 1 (on sides of jaw) irregular inner rows of small

conical teeth, the conical teeth gradually increasing in size and

becoming more curved on middle part of jaw, then becoming

abruptly smaller on posterior part of jaw; vomer with | row of small,

stout conical teeth arranged in a chevron; palatines edentate or with

small irregular patch of small conical teeth; tongue acutely pointed,

edentate.

TWO NEW SPECIES OF PSEUDOPLESIOPS

LIVE COLORATION. (Based on colour photographs of the holotype

from the Great Barrier Reef, and of paratypes and other specimens

from the Great Barrier Reef, Loyalty Islands, Solomon Islands,

Vanuatu, Christmas Island and Taiwan)

Head and body pinkish or yellowish brown to olive or bright

green, sometimes becoming pinkish to orangish brown on snout, lips

and intermandibular flap; iris pale yellow to green or brown, some-

times with reddish grey to pink area around pupil; dorsal and anal

fins yellowish, pinkish or orangish brown to olive or bright green,

becoming paler distally, with bluish grey to bright blue distal

margin, sometimes with broad pale orange to yellow stripe

submarginally; bluish grey to pale blue spot or streak at base of

alternate dorsal- and anal-fin rays; caudal fin yellowish or pinkish

brown to olive or bright green; pectoral fins pinkish, greenish or

yellowish hyaline; pelvic fins pale pink or olive to bright green,

usually bluish grey to pale blue anteriorly and distally.

PRESERVED COLORATION

Head and body pale brown, slightly darker on dorsal part of head and

body, and on lips and intermandibular flap; fins pale brown to

brownish hyaline; bluish grey to blue spots and distal margins of

dorsal and anal fins described above become greyish brown.

HABITAT AND DISTRIBUTION

This species is distributed from the Maldive Islands, east and south

to Vanuatu, and north to Taiwan. It has been collected from a variety

of reef habitats, from shallow patch reefs to reef walls at depths

ranging to at least 20 m (with some collections perhaps from as deep

as 40 m).

COMPARISONS WITH OTHER SPECIES

Pseudoplesiops immaculatus has been confused with P. revellei

Schultz, a Pacific Plate endemic (sensu Springer, 1982). The two

species closely resemble each other in morphometric and meristic

values, and both possess a prominent intermandibular flap (Fig. 2),

although this may be weakly developed or absent in small specimens

(smaller than about 18 mm SL). A low intermandibular ridge or

weak flap may be present in certain other pseudochromid species

(e.g., Chlidichthys cacatuoides, see Gill & Randall, 1994;

Pseudoplesiops occidentalis, Fig. 4), but only in P. immaculatus and

P. revellei is it well-developed. On the basis of this synapomorphy,

we suggest that the two species are sister taxa. They are distin-

guished from each other by a single coloration character: P. revellei

has a large, dark brown to black spot on the operculum, which is

absent in P. immaculatus.

REMARKS

Colour photographs of the species have been published by Allen &

Steene (1988; as P. revellei), and Randall et al., (1990, 1997 - as P.

sp.).

ETYMOLOGY

The specific epithet is from the Latin, meaning without a spot, and

alludes to the sole character distinguishing P. immaculatus from its

sister species.

Pseudoplesiops occidentalis sp. nov.

Maldives Dottyback

Figures 3, 4

Clinus sp.; Regan, 1902: 276 (description; Haddumati, Maldive

Islands).

Pseudoplesiops sp.; Randall & Anderson, 1993: 15, pl. 3e (Maldive

Islands; col. fig.).

Pseudoplesiops sp. 1; Kuiter, 1998: 77 (habitat notes; colour photo).

HOLOTYPE. BPBM 32926, 24.9 mm SL, Maldive Islands, South

Malé Atoll, Maaniyafushi Island, reef, 25-30 m, J.E. Randall, C.

Anderson and M.S. Adam, 17 March 1988.

PARATYPES. AMS 1.41004-001, 1: 21.3 mm SL (subsequently

cleared and stained), collected with holotype; BMNH 1901.12.31.77,

1: 26.2 mm SL, Maldive Islands, Haddummati Atoll (= Haddumati),

72 m, J.S. Gardiner; BPBM 32871, 2: 16.7—26.5 mm SL, Maldive

Islands, Ari Atoll, E side of reef N of Bathala Island, rubble bottom,

35 m, J.E. Randall and M.S. Adam, 6 March 1988.

DIAGNOSIS

Pseudoplesiops occidentalis is distinguished from all other pseudo-

plesiopines in having the following combination of characters:

dorsal-fin rays II,23; scales in lateral series 26—28; and scales with

distinct centres and radii in all fields.

Fig. 3 Pseudoplesiops occidentalis, BPBM 32926, 24.9 mm SL, holotype, South Malé Atoll, Maldive Islands. (Photo by P. Crabb.)

A.C. GILL AND A.J. EDWARDS

LSCP

—S

Fig.4 Cephalic sensory pores of Pseudoplesiops occidentalis, BPBM 32926, 24.9 mm SL, holotype, South Malé Atoll, Maldive Islands. Arrow indicates

atypical doubled suborbital pore (present unilaterally only in holotype). IMR, intermandibular ridge. Other abbreviations and methods of presentation

follow Fig. 2.

DESCRIPTION. (Based on five specimens, 16.7—26.5 mm SL)

Dorsal-fin rays II,23, last 6-7 (6) segmented rays branched; anal-fin

rays II,14, last 5-6 (5) segmented rays branched; pectoral-fin rays

15-16 (16/15), upper 2-4 (2/2) and lower 1-3 (1/2) rays simple;

pelvic-fin rays [,3, all segmented rays simple; principal caudal-fin

rays 9 + 8, the uppermost 1-2 (1) and lowermost 1-2 (2) rays

unbranched; upper procurrent caudal-fin rays 3—4 (4); lower

procurrent caudal-fin rays 3-4 (4); total caudal-fin rays 23-25 (25);

scales in lateral series 26-28 (27/27); predorsal scales 6-8 (8);

transverse scales 11-12 (11/11); scales behind eye 2; scales to

preopercular angle 3; circumpeduncular scales 16; ctenoid scales

beginning at 7—9 (7/9) transverse scale rows behind branchial

opening; gill rakers 2-4 + 8-11 = 10-15 (2 + 8); pseudobranch

filaments 6-7 (7).

Head pores (all bilaterally paired; Fig. 4): nasal pores 2; anterior

interorbital pores 1; posterior interorbital pores 0; supraotic pores 2;

suborbital pores 8—9 (9/8); posterior otic pores 0; preopercular pores

7; dentary pores 4; intertemporal pores 1; anterior temporal pores 0;

posttemporal pores 1; parietal pores 2.

As percentage of standard length: body depth at dorsal-fin origin

24.0—28.2 (26.5); greatest body depth 24.0-32.4 (27.7); body width

13.3-14.5 (13.3); head length 30.5—33.5 (30.5); snout length 5.0—

6.4 (5.6); orbit diameter 8.4—10.2 (8.4): interorbital width 3.64.8

(3.6); upper jaw length 11.3—-12.0 (11.6); depth of caudal peduncle

16.5-16.9 (16.5); caudal peduncle length 7.2-8.5 (7.2); predorsal

length 29.0-32.9 (31.3); preanal length 55.0-61.1 (60.2); prepelvic

length 30.1-32.5 (30.1); length of first segmented dorsal-fin ray

10.4-11.7 (10.4); length of third from last segmented dorsal-fin ray

17.7-19.7 (17.7); dorsal-fin base length 61.7—71.4 (62.7); length of

first segmented anal-fin ray 12.2—14.5 (12.9); length of third from

last anal-fin ray 17.8—18.7 (18.5); anal-fin base length 32.8-35.9

(33.7); caudal fin length 25.3—27.9 (25.3); pectoral fin length 22.5—

24.0 (23.3); pelvic fin length 27.7—35.3 (27.7).

Lower lip complete; prominent intermandibular flap absent, al-

though low ridge present in some specimens (Fig. 4); fin spines

weak and flexible; anterior dorsal-fin pterygiophore formula S/S/3/

1 + 1; 15-16 (15) consecutive dorsal-fin pterygiophores inserting in

1:1 relationship directly behind neural spine 4; anterior anal-fin

pterygiophore formula 2/1 + 1; 7—8 (8) consecutive anal-fin

pterygiophores inserting in 1:1 relationship directly behind haemal

spine 2; second segmented pelvic-fin ray longest; caudal fin weakly

rounded to rounded or slightly emarginate; scales with distinct

centres and radii in all fields; dorsal and anal fins without distinct

scale sheaths, although with basal row of body scales overlapping

fin base, particularly on posterior part of fins; anterior lateral line

represented by single tubed scale at branchial opening, followed by

intermittent series of centrally pitted scales, which terminate at

vertical through base of segmented dorsal-fin ray 17—22 (2/22):

TWO NEW SPECIES OF PSEUDOPLESIOPS

second intermittent series of centrally pitted scales originating on

midside above anterior part of anal fin, extending on to middle part

of caudal-fin base; additional centrally pitted scales present above

and below pitted scale(s) on middle part of caudal-fin base, some-

times extending on to posterior part of caudal peduncle; scales

present on cheeks (extending posteriorly over upper part of

preopercle) and operculum (Fig. 4); predorsal scales extending

anteriorly to point between anterior interorbital pores and vertical

through anterior supraotic pores (Fig. 4); vertebrae 10 + 17; epurals

2; epineurals present on vertebrae 1 through 16—18 (1 through 16);

ribs present on vertebrae 3 through 10, rib on ultimate precaudal

vertebra relatively long.

Upper jaw with 2-5 pairs of curved, enlarged caniniform teeth

anteriorly, the medial pair smallest, and 34 (at symphysis) to 1-2

(on sides of jaw) irregular inner rows of small conical teeth, the teeth

of outer row of conical teeth largest; lower jaw with 2-3 pairs of

curved, enlarged caniniform teeth, the medial pair smallest, and 2—

3 (at symphysis) to | (on sides of jaw) irregular inner rows of small

conical teeth, the conical teeth gradually increasing in size and

becoming more curved on middle part of jaw, then becoming

abruptly smaller on posterior part of jaw; vomer with | row of small,

stout conical teeth arranged in a chevron; palatines edentate or with

small irregular patch of small conical teeth; tongue pointed, eden-

tate.

LIVE COLORATION. (Based on colour photograph of the holotype,

photograph in Kuiter, 1998, and field notes taken from paratypes in

BPBM 32871 when freshly dead)

Head and body bright pinkish red to orange-red, becoming pink

ventrally and olive-red to orangish brown posteriorly; margin of

orbit orange posteriorly, becoming pale blue ventrally; posttemporal,

intertemporal, upper preopercular and upper suborbital pores indis-

tinctly edged with grey; iris bright orange-red; pectoral-fin base pale

pink to pinkish red; dorsal and anal fins bright orange-red basally,

reddish hyaline distally, with pale blue distal margin; caudal fin

greyish yellow to pale greyish red basally, remainder of fin reddish

to yellowish hyaline with pale blue distal margin; pectoral fin

pinkish to yellowish hyaline; pelvic fin pale pink basally, becoming

pale blue distally.

PRESERVED COLORATION

Head and body brown, paler ventrally; grey edging on posttemporal,

intertemporal, upper preopercular and upper suborbital pores re-

mains, becoming greyish brown; dorsal and anal fins brownish

hyaline, becoming greyish brown distally; other fins pale brown to

dusky hyaline.

HABITAT AND DISTRIBUTION

Pseudoplesiops occidentalis is known only from the Maldive Is-

lands. It has been collected from and observed on reefs in 20 to 72 m.

COMPARISONS WITH OTHER SPECIES

Pseudoplesiops occidentalis forms a monophyletic group with P.

typus Bleeker, P. rosae Schultz and several undescribed species.

With the exception of P. occidentalis, this clade is confined to the

eastern Indian Ocean and the West Pacific. It is diagnosed by

scales with distinct centres and radii in all fields. Species limits

within the clade are poorly resolved and are currently under study

by us. For the purposes of comparison with P. occidentalis, we

divide the clade into three subgroups, each of which we believe to

be monophyletic: P. occidentalis, P. rosae-complex (autapomorphy:

plate-like, median expansion of median ethmoid, pterosphenoid

and basisphenoid into orbital space); and P. typus-complex

(autapomorphy: all scales cycloid in adult specimens). Aside from

the various autapomorphies listed above, the three taxa are distin-

guished from each other by the following: number of vertebrae (10

+ 17 in P. occidentalis, 11-12 + 16-18 = 27-29, usually 11 + 17-

18 in the P. rosae-complex and 11 + 17-18 in the P.

typus-complex); number of dorsal-fin rays (II,23 in P. occidentalis,

1,22—24 in the P. rosae-complex and II,24—25 in the P. typus-

complex); number of anal-fin rays (II,14 in P. occidentalis,

I-II,12—14, usually I,13-14 in the P. rosae-complex and II-III, 14—

16, usually II,15 in the P. typus-complex), number of scales in

lateral series (26—28 in P. occidentalis, 26-29 in the P. rosae-

complex and 32-40 in the P. typus-complex); number of

circumpeduncular scales (16 in P. occidentalis, 16 in the P. rosae-

complex and 20-22 in the P. typus-complex); predorsal scalation

(6-8 scales, extending anteriorly to a point between the anterior

interorbital pores and the vertical through the anterior supraotic

pores in P. occidentalis, 5-10 scales, extending anteriorly to a

point between the anterior interorbital pores and the vertical

through the anterior supraotic pores in the P. rosae-complex, and

10-16 scales, extending anteriorly to the supratemporal commissure

in the P. typus-complex); and cheek scalation (broadly overlapping

the upper part of the preopercle in P. occidentalis and the P. rosae-

complex, versus not overlapping the upper part of the preopercle in

the P. typus-complex). Members of the P. typus-complex also

attain a much larger body size than members of the other clades

(largest examined specimen 53.0 mm SL versus 26.5 mm SL in P.

occidentalis and 26.9 mm SL in the P. rosae-complex).

Of the characters noted above that are shared by P. occidentalis

and the P. rosae-complex, three are unique within Pseudoplesiops

and suggest a sister-relationship between the two taxa: low number

of scales in lateral series; predorsal scales extending anteriorly

beyond the supratemporal commissure; and cheek scales broadly

overlapping upper part of preopercle. This relationship will be tested

in a study of the phylogeny of the Pseudochromidae currently in

progress by the first author.

REMARKS

Colour photographs of the species are provided by Randall &

Anderson (1993) and Kuiter (1998). Randall & Anderson indicate

that their photograph is of a 26 mm SL specimen in BPBM 32871,

but it is actually of the holotype (BPBM 32926).

Regan’s (1902) specimen of the species (BMNH 1901.12.31.77)

was initially identified and reported on as “Clinus sp.” A label on the

jar and catalogue entry indicates that it was subsequently

redetermined as the plesiopid Belonepterygion fasciolatum Ogilby

(apparently by M.L. Penrith). Although we were unable to locate

any references to this identification, they possibly exist.

ETYMOLOGY

The specific epithet is from the Latin, meaning of the west; P. occi-

dentalis is known only from the western-most part of the range of the

genus.

ACKNOWLEDGEMENTS. We thank the following for allowing access to

specimens in their care: G.R. Allen, A. Bentley, D. Catania, J.-P. Chen, O.A.

Crimmen, W.N. Eschmeyer, P.C. Heemstra, D.F. Hoese, S. Jewett, J. Johnson,

M. McGrouther, R.J. McKay, V. Mthombeni, L. Palmer, K. Parkinson, J.E.

Randall, S.E. Reader, T. Trnski, M. Sabaj, W. Saul, K.-T. Shao, W.F. Smith-

Vaniz, A. Y. Suzumoto, M.J.P. van Oijen and J.T. Williams. Colour photographs

were kindly provided by J.-P. Chen, D.F. Hoese, R.H. Kuiter, J.E. Randall and

J.T. Williams; P. Crabb took the black and white photographs of the two

holotypes (Figs 1 and 3). O.A. Crimmen, S. Davidson, J.P. Garcia, A.-M.

Hine, K. Parkinson, S. Raredon and S.E. Reader radiographed specimens.

R.D. Mooi and J.T. Williams critically reviewed the manuscript and provided

helpful comments.

REFERENCES

Ahlstrom, E.H., Butler, J.L. & Sumida, B.Y. 1976. Pelagic stromateoid fishes

(Pisces, Perciformes) of the eastern Pacific: kinds, distributions, and early life

histories and observations of five of these from the northwest Atlantic. Bulletin of

Marine Science 26(3): 285-402.

Allen, G.R. & Steene, R.C. 1979. The fishes of Christmas Island, Indian Ocean.

Australian National Parks and Wildlife Service, Special Publication 2: 1-81.

—-— & 1988. Fishes of Christmas Island, Indian Ocean. 197 p. Christmas Island

Natural History Association.

& Smith-Vaniz, W.F. 1994. Fishes of the Cocos (Keeling) Islands. Atoll Research

Bulletin 412: 1-21.

Bleeker, P. 1858. Bijdrage tot de kennis der vischfauna van den Goram Archipel.

Naturrkundig Tijdschrift Nederlandsch Indié 15: 197-218.

1875. Sur la famille des Pseudochromoides et révision de ses espéces

insulindiniennes. Verhandelingen der Koninklijke Akademie van Wetenschappen,

Amsterdam 15: 1-32, pls 1-3.

Gill, A.C. 2000. Pseudochromidae, pp 2557-2577. In: Carpenter, K.E. & Niem, V.H.

(eds) FAO Species Identification Guide for Fisheries Purposes. The Living Marine

Resources of the Western Central Pacific. Vol. 4. FAO, Rome.

& Edwards, A.J. 1999. Monophyly, interrelationships and description of three

new genera in the dottyback fish subfamily Pseudoplesiopinae (Teleostei: Perciformes:

Pseudochromidae). Records of the Australian Museum 52(1): 141-160.

, Mooi, R.D. & Hutchins, J.B. 2000. Description of a new subgenus and species

of the fish genus Congrogadus Giinther from Western Australia (Perciformes:

Pseudochromidae). Records of the Western Australian Museum 20(1): 69-79.

& Randall, J.E. 1994. Chlidichthys cacatuoides, anew species of pseudoplesiopine

dottyback from Oman, with a diagnosis of the genus Chlidichthys Smith, and new

record of Pseudochromis punctatus Kotthaus from Oman (Teleostei: Perciformes:

A.C. GILL AND A.J. EDWARDS

Pseudochromidae). Revue francaise d’Aquariologie Herpétologie 21(1-—2): 11—18.

Kailola, P.J. 1987. The fishes of Papua New Guinea: a revised and annotated checklist.

Volume two. Scorpaenidae to Callionymidae. Papua New Guinea Department of

Fisheries and Marine Resources Research Bulletin 41: 195-418.

Kuiter, R.H. 1998. Photo Guide to Fishes of the Maldives. 257 p. Atoll Editions,

Apollo Bay.

Kulbicki, M. & Williams, J.T. 1997. Checklist of the shorefishes of Ouvéa Atoll, New

Caledonia. Atoll Research Bulletin 444: 1-26.

Leviton, A.E., Gibbs, R.H., Jr., Heal, E. & Dawson, C.E. 1985. Standards in her-

petology and ichthyology: part 1. Standard symbolic codes for institutional resource

collections in herpetology and ichthyology. Copeia 1985(3): 802-832.

Paxton, J.R., Hoese, D.F., Allen, G.R. & Hanley, J.F. 1989. Zoological Catalogue of

Australia. Volume 7. Pisces. Petromyzontidae to Carangidae. 665 p. Australian

Government Publishing Service, Canberra.

Randall, J.E., Allen, G.R. & Steene, R.C. 1990. Fishes of the Great Barrier Reef and

Coral Sea. 507 p. Crawford House Press, Bathurst.

- & . 1997. Fishes of the Great Barrier Reef and Coral Sea. Revised and

expanded edition. 557 p. Crawford House Press, Bathurst.

& Anderson, R.C. 1993. Annotated checklist of the epipelagic and shore fishes

of the Maldive Islands. Ichthyological Bulletin of the J.L.B. Smith Institute of

Ichthyology 59: 1-47, pls 1-8.

Regan, C.T. 1902. On the fishes from the Maldive Islands. The Fauna and Geography

of the Maldive and Laccadive Archipelagos 1(3): 272-281.

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Smithsonian Contributions to Zoology 367: 1-182.

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small fishes and other vertebrates for bone and cartilage study. Cybium 9(2): 107-119.

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Congrogadinae (Pisces: Perciformes: Pseudochromidae). Indo-Pacific Fishes 9: 1—

34, pl. I. [dated 1985, but actually published 1986]

Me (as2et5. 1)

Bull. nat. Hist. Mus. Lond. (Zool.) 68(1): 27-37

Issued 27 June 2002

A redescription of Sousa chinensis (Osbeck,

1765) (Mammalia, Delphinidae) and

designation of a neotype

L.J. PORTER

The Swire Institute of Marine Science, The University of Hong Kong, Cape d’Aguilar, Shek O, Hong Kong

Special Administrative Region, China.

Synopsis. The holotype of the delphinid Sousa chinensis, held in the collection of the Royal College of Surgeons, London, was

destroyed during the Second World War. The neotype is described herein from a male specimen obtained from the waters of Hong

Kong (the Hong Kong Special Administrative Region of China since | July 1997). This dolphin was found newly dead in May

1996 and death determined as heart failure brought about by entanglement with fishing gear. There is currently debate over

divisions within the genus Sousa although recent reviews suggest that the five nominal species currently designated likely

comprise but one. A description of the neotype’s external appearance and skeletal elements is given and compared with the

information that remains for the holotype and for other specimens obtained from Hong Kong waters.

INTRODUCTION

The species Sousa chinensis (formerly Delphinus sinensis) (Osbeck,

1765) was designated after observations were made of a population

of delphinids within the Pearl River Estuary, Guangdong Province,

Southern China, i.e. close to Hong Kong. It was over one hundred

years after the species was named, however, that the first skeleton

was described from an almost entire individual collected from

Xiamen (formerly Amoy), Fujian Province, approximately 700km

northeast of Hong Kong (Flower, 1870). This specimen and, thus,

the holotype of the species was held in the collections of the Royal

College of Surgeons, London, but was subsequently destroyed

during the Second World War.

Osbeck’s original account noted ‘Snow-white Dolphins tumbled

about the ship; but at a distance they seemed nothing different from

the common species, except in their white colour’. The holotype was

physically mature and although the external appearance of this

particular individual was not described, the typical coloration of

members of the Xiamen population was noted to be ‘milky white,

with pinkish fins and black eyes’ (Flower, 1870). The individuals

that remain in the Pearl River and Jiulong Jiang River (Xiamen)

Estuaries today, are similar in external appearances (Porter, 1998).

Four other species are included in the genus Sousa (Table 1),

although there has been debate as to where the taxonomic divisions

within it, if any, actually occur (Pilleri & Gihr, 1972; Mitchell, 1975;

Zhou et al., 1980; Wang & Sun, 1982; Ross et al., 1994). The most

recent classification of the genus as comprising one species was

based upon a morphological review (Ross ef al., 1996) and a

molecular investigation of the genus concurred with this definition

(Cockcroft et al., 1997). Both of these publications, however, noted

that the limited availability of data from parts of the species’ range,

particularly Asia, did not allow definitive segregation at a sub-

specific level and, as such, the taxonomy of the genus is not

resolved.

There have been many strandings of Sousa chinensis in Hong

Kong waters, which comprise the eastern portion of the Pearl River

Estuary, since 1990 (Porter et al., 1998) although the cause of death

in many cases could not be determined due to either the decomposi-

tion of, or damage to, the remains. In May 1996, however, a fresh

specimen identified as S. chinensis was retrieved and on which a full

necropsy was performed. The cause of death was determined as

heart failure, most likely due to entanglement with fishing gear, as

deduced by the presence of fresh net abrasions on the rostrum and

neck area. This specimen has been chosen for designation as the

neotype of S. chinensis because: (a), it was obtained from the area

from which the species was originally identified; (b), it was neither

significantly different from the description of the extant holotype

nor other individuals measured from the Hong Kong population of

S. chinensis; (c), its death was perceived to be accidental rather than

a consequence of any ailment or abnormality and (d), its remains

were intact.

MATERIALS AND METHODS

A modification of the criteria described by Perrin (1975) was used to

measure the skeleton obtained from Hong Kong and herein desig-

nated as the neotype. The tympanoperiotic bones were measured in

accordance with Kasuya (1973). Other skeletal remains of the same

species were also examined and measured to obtain a better range of

values for the species in this component of its range. All measure-

ments were made with EPI precision calipers and recorded to the

nearest millimetre or, in the case of smaller bones, 0.1 mm. Each

measurement was taken several times until a confident value was

established. An inventory of the measurements recorded for the

neotype is included as Appendix I.

The neotype and other Hong Kong specimens were aged from

three teeth, selected from the upper left jaw, which were sliced to a

thickness of 110um. The sections were etched for one hour by

emersion in 5% formic acid and the dentinal growth layer groups

then counted under a light microscope. One pair of dark and light

stained layers is equivalent to one dentinal growth layer group

which is assumed to be gained annually (Kasuya, 1976).

The equivalent holotype measurements of Sousa chinensis

(Flower, 1870) were transformed into metric units and compared

with corresponding values for the neotype. The original drawings of

Flower’s (1870) and photographs of the neotype were also com-

pared. The skeletal characters of both the holotype and neotype were

28 L.J. PORTER

pes aeesse

; 5%

gurere®

Fig. 1 The skull and jaw bones of the neotype of Sousa chinensis seen from a, lateral; b, ventral; c, ramus dorsal and d, dorsal aspects

REDESCRIPTION OF THE CHINESE WHITE DOLPHIN

Table 1 Five species classification of the genus Sousa according to

morphological differences and geographical distribution (Ross et al.,

1994).

Nominal species Geographical range

S. chinensis (Osbeck, 1765)

S. plumbea (Cuvier, 1829)

S. lentigenosa (Owen, 1866)

S. teuzii (Kukenthal, 1892)

S. borneensis (Lydekker, 1901)

China to Australia

South Africa to India

India to Southeast Asia

West Africa

Rare form in Borneo, overlapping

with ‘chinensis’

then compared with the range of measurements obtained from other

dead adult specimens of S. chinensis stranded on Hong Kong beaches.

RE-DESCRIPTION OF SOUSA CHINENSIS

(OSBECK, 1765)

Neotype: ZD 1999.360, sub-adult male, N 022° 20' 00; E 113° 55’

30, The Brothers Islands, Hong Kong’s territorial waters, collected

by staff of the Swire Institute of Marine Science, The University of

Hong Kong. Donated to the Natural History Museum, London,

under CITES permit APO/EL 3227/99.

The gently-tapering rostrum of this specimen comprised 60.7% of

the total skull length and was positioned low near the base of the

cranium (Fig. la). The upper and lower tooth rows comprised 89%

and 63%, respectively, of the rostrum and ramus lengths, with 3 1—35

Fig. 2 The a, foramen magnum; b, atlas and c, axis of the neotype of Sousa chinensis

teeth in each row (Fig. 1b & c). The teeth were conical and pointed

with little wear. Five dentinal layers were counted from sectioned

teeth and the pulp cavity was open. The premaxillaries were in

contact for almost the entirety of the rostrum length, with only a

small section of the vomer exposed. Storage of the specimen in a dry

environment, however, has subsequently caused the premaxillaries

to gape (Fig. 1d). The premaxillaries were roughened dorsally

where they met the less dense maxillaries, at the base of the nasal

aperture. The posterior extremity of the rounded, right, premaxillary

extended to the anterior of the nasal aperture, whereas the pointed

left premaxillary was truncated at the mid-margin of the opening.

The premaxillaries, therefore, encircled two thirds of the aperture, in

L.J. PORTER

which each nasal passage was deep-set and elliptical. The antorbital

notch was not developed. The two occipital condyle margins were

sharply defined and the anterior margin pointed. The channel between

the condyles was long and triangular. The foramen magnum was

ovoid with a slightly extended dorsal margin (Fig. 2a). Both the

temporal and post-temporal fossae were ovoid with sharply-defined

margins. The temporal fossae protruded where they met which, in

ventral view, gave the cranium a relatively broad appearance. The

orbits were rounded and the lacrimals, positioned at right angles to

the orbit, were blunt with prominent dorsal ridges. The cranial

articulating surface of the atlas was broad with short, transverse

processes and a robust, relative to the rest of the vertebrae, neural

Fig. 3 The a, vertebral column; b, vertebral ribs and c, sternal and free floating ribs of the neotype of Sousa chinensis

REDESCRIPTION OF THE CHINESE WHITE DOLPHIN

spine (Fig. 2b). The axis was not fused to the atlas and had short

processes but a well-developed neural spine (Fig. 2c).

There were seven cervical, 12 thoracic, 23 lumbar and 18 caudal

vertebrae. No vertebrae were fused (Fig. 3a). The first vertical

perforating foramen appeared on the 25th vertebra which was also

the first to have greatly reduced metapophyses and the last with

distinct transverse and neural processes. There were 12 vertebral

ribs on both the left and right sides, six of which on each side were

double-headed (Fig. 3b). There were five sternal ribs on each side

(Fig. 3c).

The sternum was notched with a perforation on the left side and

two lateral processes on each side (Fig. 4a; note that the right side of

the sternum was damaged during preparation). There was one

Sil

mesosternal element which equalled two thirds of the manubrium.

The radius and ulna were longer than the humerus, although the

latter was wider and more robust. The transverse breadth of the

carpals was short and the pectoral fin elongated (Fig. 4b). The

scapula had a sharply pointed coracovertebral angle, a broad

metacromion process and a smaller, pointed, coracoid process (Fig.

4c).

The bullae of the tympanoperiotic bones were rounded and

bulbous and the dorsal plane of the periotic convex; the fundus of the

internal auditory meatus and the aquaductus fallopii were ovoid; the

aquaductus cochleae had a distinct ridge; the join between the

cochlear portion and the rest of the periotic was rounded and

smooth.

Fig.4 The a, sternum and b, pectoral fin skeletal elements of the neotype of Sousa chinensis

HOLOTYPE

(a) occipital condyles

NEOTYPE

HOLOTYPE

NEOTYPE

L.J. PORTER

Fig.5 A copy of the original illustrations of Flower’s (1870) holotype and the corresponding views of the neotype showing differences in the a, occipital

condyles; b, lacrimal flares and c, temporal fossae

REDESCRIPTION OF THE CHINESE WHITE DOLPHIN

COLORATION. Mainly pink with light grey around the head and

darker pigmentation along the length of the back, including the

leading and trailing edge of the dorsal fin and the ventral fringe of

the fluke. There was dark, fragmented pigmentation on the melon,

around the eyes and across the dorsal cape. Although both the dorsal

and ventral surfaces of the fluke were pink, the dorsal surface of the

pectoral fins were still grey and had only just begun to lighten.

EXTERNAL APPEARANCE. The body was slender but with a promi-

nent melon. Neither the ‘robustness’ nor prominent peduncle, typical

of other, adult individuals observed and examined from Hong Kong

waters were present (Porter, 1998). The dorsal fin was slightly

falcate and the flukes were small with a distinct notch. The pectoral

fins were broad in the middle but tapered both distally and at the

base. The ‘hump-back’, reportedly typical of populations west of

Indonesia, has not been recorded from any individual in the South

China Sea (Reeves & Leatherwood, 1994).

COMMENTS. During the post-mortem it was noted that the indi-

vidual had deep scarring around the pectoral fins and torso. This

healed injury was consistent with abrasion from either a rope or a

similar object. It cannot be determined if this injury caused any

major deformity to the pectoral fin bones although some pathology

of them was observed. These injuries were distinct from those

associated with the animal’s death.

ETYMOLOGY. The species name is derived from its geographical

location, i.e. both sinensis and chinensis referring to ‘China’. The

meaning of the current genus name Sousa has become obscure.

Although the taxonomy of the genus Sousa has yet to be resolved, in

accordance with the International Code of Zoological Nomencla-

ture, this neotype must bear the name of the holotype.

COMPARATIVE ANALYSIS OF SPECIMENS

The age of nine Hong Kong specimens of Sousa chinensis used for

comparative skeletal purposes in this study, ranged between 4 and

14 growth layer groups, the former number having been established

as the age where rapid juvenile growth rate decreases and adult

dimensions are gained (Porter, 1998). No Hong Kong specimen had

reached physical maturity as determined by fusion of post-cranial

elements and closure of the tooth pulp cavity. There was, therefore,

no physically mature specimen to compare with the holotype. The

most obvious difference, therefore, between the skeletal measure-

ments available for the holotype and those for the neotype is that of

size (Table 2). The holotype had two less lumbar vertebrae, four

more caudal vertebrae and six more chevron bones than the neotype.

The holotype had slightly fewer teeth than the neotype but both

counts are within the range identified for other Hong Kong speci-

mens (Table 3). A comparison of the remaining diagrams of the

holotype with the neotype illustrates clearly the similarities in

general shape and skull configuration. Both have large ovoid crani-

ums with robust and elongated rostrums, the eliptical nares are deep

set and the vomer is exposed (Fig. 5). Differences are apparent,

however, in the occipital condyles (a) and the lacrimal flares (b),

which were more prominent in the holotype. The shape of the

holotype temporal fossae also differed from those of the neotype (c):

in the former they were swept back towards the posterior of the skull

resulting in a smaller width between them. There are differences in

the degree of ossification of the scapulae, the holotype being more

robust with larger coracoid and metacromion processes (Fig. 6).

The measurements obtained from the holotype, the neotype and

Table 2 Sousa chinensis principal skeletal dimensions and vertebral

counts of the Hong Kong neotype and the holotype from Fujian

Province (Flower, 1870).

Measurement Neotype Holotype

1 Condylobasal length 481 526

2 Length of rostrum. 292 325

3 Width of rostrum 110.1 I)

5 Width of rostrum at midlength 44.8 47

16 Greatest parietal width 140.9 163

32 Length upper tooth row to tip rostrum 260 284

33(UL) Number of teeth (UL) 35 33

34(UR) Number of teeth (UR) 35 32

35(LL) Number of teeth (LL) 31 32

36(LR) Number of teeth (LR) 32 31

a7 Length of lower tooth row. 259 279

38 Greatest length of left ramus. 407 457

39 Greatest height of left ramus. 82.5 91

40a Length of symphysis 113.3 140

48a Number of cervical vertebrae i 7

48 Number of thoracic vertebrae 12 12

49 Number of lumbar vertebrae 12 10

50 Number of caudal vertebrae 18 22

51 Total number of vertebrae 49 51

78 Number of vertebral ribs (left) 12 12

79 Number of vertebral ribs (right) 12 12

96 Number of chevron bones 8 14

105 Greatest length of coracoid process. 21.5 37

106 Greatest width of coracoid process. 11.6 29

107 Greatest width of metacromion process. 54.5 30

108 Greatest length of humerus. 63.9 73

109 Greatest width of humerus distally. 39:5 57

110 Greatest length of radius. 74 79

111 Greatest width of radius distally. 38.2 47

112 Greatest length of ulna. 65.9 67

HOLOTYPE

coracoid process

metacromion process

NEOTYPE

Fig. 6 A copy of the original illustrations of Flower’s (1870) holotype

and the corresponding views of the neotype demonstrating differences

in the form of the coracoid and metacromion processes of the scapula.

550 5

500

450

400

350

3 300

(3)

2 250

uo}

200

150

100

320 -

270 -

= 220 -

—

S |

is 170

i“)

<b)

2 120 -

L.J. PORTER

rT]

| e

| a

| °

rT]

e | @ Neotype |

K w Holotype |

+ a oe

Condylobasal length Ramus length Rostrum length Upper tooth row length Symphysis length

5 ° Neotype

A m Holotype

i‘

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

Table 3 The range (and average number) of teeth counted from Hong

Kong Sousa chinensis

Left Right

Upper 32-36 (33.6) 32-35 (33.6)

Lower 28-34 (31.5) 27-33 (31.1)

the other eight Hong Kong specimens lie within a discrete range.

The greatest variation is seen between length measurements of the

skull, i.e. condylobasal (450-526 mm), rostral (257-325 mm),

upper tooth row (205.5—284 mm), ramus (397-457 mm) and sym-

physis (82.5-258 mm). The physical maturity of the holotype is

reflected in its location at the top of each of these length values (Fig.

7a). Least variation is seen between the width dimensions of the

skulls, i.e., parietal (137.4-171 mm), rostral (99.2-119 mm), mid-

rostral (47.8—62 mm), mandibular fossae (119.4—-130.9 mm) and the

lower tooth row length (247-279 mm) (Fig. 7b).

DISCUSSION

The ages of nine stranded specimens of Sousa chinensis examined

from Hong Kong waters were determined to lie between 4 and 14

years. It is clear from the original description that the holotype was

physically mature whereas no specimen obtained from Hong Kong

was. This is reflected in the comparison of skeletal characters where

the holotype is consistently the largest in terms of skull length

measurements. The greater ossification of the scapula also is indica-

tive of the holotype’s greater age. The measurements obtained for

the neotype appear to be typical for Hong Kong waters, although the

sample size is currently limited. Those measurements available for

the holotype are, however, within the ranges obtained for Hong

Kong specimens. The difference between vertebral counts of the

holotype and neotype is small and such variation in other delphinids

appears typical (Perrin, 1975). It is not known if the holotype was

typical of the population present in Xiamen waters at the time it was

collected (1870) and, indeed, as both Hong Kong and Xiamen

coastal areas have been extensively developed, any adaptive changes

that both populations of Sousa chinensis may have undergone since

the 17" century cannot be gauged. In addition, the latest review of

the genus (Ross et al., 1996) observes that osteological form is

similar across its global range, with any slight differences being

environmental rather than genetic.

The uncertainty that prevails over the taxonomy of the genus

Sousa can only be dispelled through the continued investigation and

comparison of specimens from throughout its range. As the holotype

has been destroyed, the designation of a neotype is required so that

such comparative studies can be conducted in relation to a defined

type and, thus, the taxonomy and life history traits of the genus

resolved. In addition, the herein designated neotype was obtained

from the area in which the species was originally identified (Osbeck,

1765) and the same geographic region that the holotype was col-

lected from (Flower, 1870). The meristic data of the neotype are

neither significantly different from what is known of the original

description of a Sousa chinensis skeleton (Flower 1870) nor the

sample of other stranded specimens measured from Hong Kong

waters. The specimen described herein, therefore, fulfils the criteria

as laid out by the International Code of Zoological Nomenclature for

designation as a neotype and, as such, has now been lodged in the

vertebrate collection of the Natural History Museum, London

(Accession No. ZD 1999.360).

ACKNOWLEDGEMENTS. The study of Hong Kong stranded marine mam-

mals was funded by the Agriculture, Fisheries and Conservation Department

(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 staff of the licensing division of AFCD are gratefully acknowledged for

their assistance in processing the CITES export permit. Many thanks also to

Dr. G. Ross, Environment Australia, who freely gave of his expertise and

greatly assisted in the taxonomic procedures and interpretations presented in

this paper. The author also wishes to thank Prof. Brian Morton, Director, The

Swire Institute of Marine Science, The University of Hong Kong, for his

support and encouragement for this and other studies of Sousa chinensis

conducted at the institute.

REFERENCES

Cockcroft, V.G., Leatherwood, S., Goodwin J. & Porter, L. 1997. The phylogeny of

humpback dolphins genus Sousa: insights through mtDNA analyses. Working paper

SC/49/SM25 14pp. In: Proceedings of the 49" International Whaling Commission

Meeting, Scientific Committee Annual Meeting, Bournemouth, United Kingdom,

1997. International Whaling Commission, Cambridge.

Flower, W. 1870. Description of the skeleton of the Chinese white dolphin (Delphinus

sinensis Osbeck ). Transactions of the Zoological Society of London 7: 151-160.

Kasuya, T. 1973. Systematic consideration of recent toothed whales based on the

morphology of tympanoperiotic bone. Scientific Report of the Whales Research

Institute, Tokyo 25: 1-103.

1976. Reconsideration of life history parameters of the spotted and striped

dolphins based on cemental layers. Scientific Report of the Whales Research Institute

28: 73-106.

Kukenthal, W. 1892. Sotalia teusziin. sp., einplanzenfressender Delphin aus Kamerun.

Zoologischen. Jahrbuchen 6: 442446.

Mitchell, E. 1975. Report of the meeting on smaller cetaceans, Montreal, April 1-11,

1974. In Review of Biology and Fisheries for Smaller Cetaceans. Journal of the

Fisheries Research Board of Canada 24: 2505-2513.

Osbeck, P. 1765. Reise nach Ostindien und China. (Translated from the original

Swedish by J.G. Georgi and J.C. Koppe. Rostock. [In German]

Perrin, W.F. 1975. Variation of Spotted and Spinner porpoise (genus Srenel/a) in the

Eastern Tropical Pacific and Hawaii. Bulletin of the Scripps Institute of Oceanogra-

phy. University of California Press, California.

Pilleri, G, & Gihr, M. 1972. Contribution to the knowledge of the cetaceans of

Pakistan with particular reference to the genera Neomeris, Sousa, Delphinus and

Tursiops and description of a new Chinese porpoise (Neomeris asiaeorientalis).

Investions of Cetacea 4:107-162.

Porter, LJ. 1998. The taxonomy, ecology and conservation of Sousa chinensis (Osbeck,

1765) (Cetacea: Delphinidae) in Hong Kong waters. PhD thesis. The University of

Hong Kong.

Reeves, R.R. & Leatherwood, S. 1994. Dolphins, Porpoises and Whales: 1994-1998

Action Plan for the Conservation of Cetaceans. (\UCN/Species Survival Commis-

sion/Cetacean Specialist Group, Cambridge.

Ross, G.J.B., Heinsohn, G.E. & Cockcroft, V.G. 1994. Humpback dolphins Sousa

chinensis (Osbeck, 1765), Sousa plumbea (G. Cuvier, 1829) and Sousa teuszii

(Kukenthal, 1892). In: Handbook of Marine Mammals, Volume 5: 23-42 Ridgeway,

S.H. & Harrison, R. (Eds). Academic Press, London.

Ross G.J.B., Heinsohn, G.E., Cockcroft, V.G., Parsons, E.C.M., Porter, L.J. &

Preen, A. 1996. Review of the taxonomic status of the humpbacked dolphins, genus

Sousa. Working paper UNEP/SEA95/WP19. 25pp. In: Perrin, W.F., Dolar, M.L. &

Alava, M.N.R. (eds.) Report of the Workshop on the Biology and Conservation of

Small Cetaceans and Dugongs of Southeast Asia, Dumaguete 1995, UNEP (W)/EAS

WG.1/2. Bangkok: United Nations Environment Programme, Bangkok.

Wang, P. & Sun, J. 1982. Studies on the Zhonghua white dolphin, Sousa chinensis,

from the South China Sea. Transactions of the Liaoning Zoological Society 3: 67-74

(In Chinese).

Zhou, K, Li, Y, Qian, W. & Yang, G. 1980. Notes on three species of dolphins from

the South China Sea and Jiulongjiang River. Oceanologica et Limnologia Sinica 11:

306-313 [In Chinese with an English Summary].

APPENDIX 1

Skeletal measurements of the neotype.

Measurement

Condylobasal length

Length of rostrum

Width of rostrum

Width of rostrum at 60 mm anterior

Width of rostrum at midlength

Premaxilleries width, at rostral midlength

Width of rostrum at 3/4 length, measured from posterior end

Distance from tip of rostrum to external nares

Distance from tip of rostrum to internal nares

Greatest preorbital width

Greatest postorbital width

Greatest width of external nares

Zygomatic width

Greatest width premaxilleries

Greatest parietal width

Least width between temporal fossa

Greatest length of left temporal fossa

Greatest width of temporal fossa (at right angles to length)

Major diameter of left temporal fossa

Minor diameter of left temporal fossa

Length of orbit

Length of antero-lateral face of lacrimal measured to exclude

the flare on the rim

Greatest width internal nares

Greatest length pterygoid

Length upper tooth row to tip rostrum

Number of teeth — upper left

Number of teeth — upper right

Number of teeth — lower left

Number of teeth — lower right

Length of lower tooth row to tip mandible

Greatest length of left ramus

Greatest height of left ramus

Length of left mandibular fossa

Length of symphysis

Greatest width of left thyrohyal proximally

Greatest length of left thyrohyal

Greatest width of left stylohyal

Greatest length of left stylohyal

Number of thoracic vertebrae, defined as equal to number of

ribs on side with greatest number

Number of lumbar vertebrae

Number of caudal vertebrae

Total number of vertebrae

Number of fused cervical vertebrae

Greatest width of articulating surface of atlas

Height of atlas — from internal anterodorsal margin of neural

canal to bottom of anterior face of body

Length of lateral process of atlas — from margin of anterior

articulating surface to farthest point at end of process

Greatest length of neural spine of atlas

Height of dorsal ridge of atlas in anterior view

Length of right dorsolateral spine of atlas

Length of lateral process of axis — from margin of posterior

Neotype

481

292

110.1

ial

44.8

28.9

34.1

323.0

300.0

185

208

50.5

203.0

82.0

140.9

137.0

86.6

WSS)

Sle

45.6

48.6

29.3

60.5

10.30

260.0

259.0

407.0

82.5

119.4

StS

18.4

66.2

13.4

79.4

none

101.9

60.9

28.3

41.1

22.8

L.J. PORTER

articulating surface to distal end of process 24.5

Number of cervical vertebrae with incomplete neural arches 2

Cervical vertebra on which left ventrolateral process reaches

greatest development By

Vertebra on which first vertical perforating foramen appears Dye)

First vertebra with greatly reduced metapophyses 250

Last vertebra with distinct transverse processes 2s

Last vertebra with distinct neural process 25%

First vertebra with unfused epiphysis 25m

Last vertebra with unfused epiphysis 25%

First caudal vertebra with vertical neural spine ae

Length of neural spine of first thoracic — from anterodorsal

margin of neural canal to tip of spine 36.5

Length of neural spine of second thoracic vertebra 46.5

Length of neural spine of tenth thoracic vertebra 65

Length of neural spine of last thoracic vertebra 69.4

Height of first thoracic vertebra — from internal anterodorsal

margin of neural canal to bottom of anterior face of body 55.0

Greatest width of first thoracic vertebra — across lateral processes 89.2

Height of the first lumbar vertebra yes)

Greatest width of the first lumbar vertebra 164.9

Length of 23d centrum, exclusive of epiphyses, along vertical

midline 40.1

Number of vertebral ribs — left 12

Number of vertebral ribs — right 12

Number of two-headed ribs — left 6

Number of two-headed ribs — right 6

Number of floating ribs — left l

Number of floating ribs — right 2

Number of sternal ribs — left 5

Number of sternal ribs — right 5

Greatest length of first left vertebral rib 123.6

Width of first left vertebral rib at apex of proximal curvature IS) 7/

Greatest length of longest left vertebral rib 141.4

Greatest width of manubrium 88.8

Length of manubrium along midline 84

Depth of anterior notch of manubrium 12

Length of foramen in manubrium 5

Number of chevron bones 7

Vertebra bearing first chevron bone 24"

Vertebra bearing last chevron bone Sills

Greatest length of first half of largest chevron bone 44.3

Greatest length of first half of last chevron bone 22.9

Height of scapula — from posterior margin of glenoid fossa

to coracovertebral angle 1251

Length of scapula — from posterior margin of glenoid fossa

to glenovertebral angle 110.3

Greatest length of coracoid process — from anterior margin

of glenoid fossa PHS)

Greatest width of coracoid process 11.6

Greatest width of metacromion process — from apex of ventral

curvature to vertebral apex 54.5

Greatest length of humerus, measured on ventral side of flipper 63.9

Greatest width of humerus distally SEES)

Greatest length of radius 74

Greatest width of radius distally 38.2

Greatest length of ulna 65.9

Transverse breadth of proximal row of carpals We

Greatest length of left pelvic rudiment 108.7

REDESCRIPTION OF THE CHINESE WHITE DOLPHIN

Measurement of the tympanoperiotic bones of the neotype.

Measurement

Length of tympanic bulla, from anterior tip to posterior end of outer posterior prominence

Anterior tip to posterior end of inner prominence

Postero-ventral tip of outer posterior prominence to tip of sigmoid process

Postero-ventral tip of outer posterior prominence to tip of conical process

Width of tympanic bulla, at the level of the sigmoid process

Height of tympanic bulla, from tip of sigmoid process to ventral keel

Width across inner and outer posterior prominences

Greater depth of interprominental notch

Width of upper border of sigmoid process

Width of the posterior branch of lower tympanic aperture

The thicker side between outer and inner posterior prominences

Length of periotic, from tip of anterior process to posterior end of posterior process, measured on a straight line parallel

with cerebral border

Thickness of superior process at the level of upper tympanic aperature

Width of periotic across cochlear portion and superior process, at the level of upper tympanic aperture

Least distance between the margins of the fundus of the internal auditory meatus and of the aperature of ductus endolymphaticus

Least distance between the margins of the fundus of the internal auditory meatus and of the aperture of aquaductus cochleae

Length of articular facet of the posterior process of periotic for the posterior process of tympanic bulla

Antero-posterior diameter of cochlear portion

Maximum depth of inner part of involvucrum

Sil

Neotype

Right

34.3

B32

24.6

18.0

2-3

25.0

18.3

4.7

5.0

1.2

10.5

31.9

13.0

22.2

2.4

4.2

15.9

19.0

9.1

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Bull. nat. Hist. Mus. Lond. (Zool.) 68(1): 39-50

1S eS Saar

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 SBD, U.K.; e-mail:

rjh@nhm.ac.uk

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 PS.

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 P1, a completely fused genital double-somite,

reduced antennary exopod and vestigial PS 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

The arid to semi-arid north-west of Western Australia has a rich

stygofauna including the only vertebrate troglobites known to occur

in Australasia, the Blind Gudgeon, Milyeringa veritas Whitley, and

the Cave Eel, Ophisternon candidum (Mees), and two, supposedly

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

and description of several cyclopoids belonging to the genera

Metacyclops Kiefer, Mesocyclops Sars, Microcyclops Claus,

Apocyclops Lindberg, Diacyclops Kiefer and Halicyclops Norman

(Pesce et al., 1996a—b; Pesce & De Laurentiis, 1996; De Laurentiis

et al., 1999).

Here we report on the discovery of a remarkable harpacticoid in

abandoned production bore holes on Barrow Island off the Cape

Range coast. Barrow Island has a typical island Ghyben-Herzberg

system with a freshwater lens overlying salt water. The hydrology of

the superficial karst is little known despite being a production oil

field since the early 1960s and being the target of ‘produced water’

disposal until recently. It lies on the North West Shelf of Western

Australia and up to about 8—10,000 years ago it would have been

part of the mainland (and throughout most of the last few million

years) (Humphreys, 2000). The entire island is likely to be an

anchialine system but to date an entry point for divers has not been

* Author for correspondence

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

production bore (47 m depth) and an abandoned anode bore (55 m

depth) that was used in the electrolytic protection of the oil wells.

The latter would have gassed chlorine and had a pH of <2 when in

service (Humphreys, pers. comm).

MATERIALS AND METHODS

Specimens were dissected in lactic acid and the dissected parts were

mounted on slides in lactophenol mounting medium. Preparations

were sealed with Glyceel or transparent nail varnish. All drawings

have been prepared using a camera lucida on a Zeiss Axioskop

differential interference contrast microscope.

Males and females of Jnermipes humphreysi gen et sp. noy. were

examined with a Philips XL30 scanning electron microscope. Speci-

mens were prepared by dehydration through graded acetone, critical

point dried, mounted on stubs and sputter-coated with gold or

palladium.

The descriptive terminology is adopted from Huys et al. (1996).

Abbreviations used in the text are: ae, aesthetasc; P1—P6, first to

sixth thoracopod; exp(enp)-1(2, 3) to denote the proximal (middle,

distal) segment of a ramus. Type material is deposited in the

collections of the Western Australian Museum, Perth (WAM) and

The Natural History Museum, London (BMNH).

SYSTEMATICS

Family AMEIRIDAE Monard, 1927

Genus INERMIPES gen. nov.

DIAGNOSIS. Ameiridae. Body elongate, cylindrical, and vermi-

form without distinct surface ornamentation except for ventral

spinule patterns on abdomen. Cephalothorax and other somites with

smooth posterior margin. Genital and first abdominal somites com-

pletely fused forming double-somite; original segmentation not

discernible. Genital field with small copulatory pore located in

median depression. Anal operculum well developed.

Sexual dimorphism in antennule, P1 inner basal spine, P5, P6,

genital segmentation and abdominal spinulation.

Rostrum fused to cephalothorax, not defined at base. Antennule

8-segmented in, 10-segmented and subchirocer in d; segment 1

without seta; aesthetascs on segments 4 and 8() or on 5 and 10 (d);

apical acrothek consisting of minute aesthetasc and 2 setae. Antenna

with separate basis and endopod; exopod minute, with | seta.

Mandible with 2-segmented palp, comprising unarmed basis and

endopod with 4 setae. Maxillule with strongly developed arthrite,

with 2 naked seta on anterior surface and 8 spines/setae around distal

margin; coxa with 2 setae and | spine; palp cylindrical with 5 setae,

probably representing basis only. Maxilla with trisetose endite on

syncoxa; allobasis drawn out in remarkably long claw; endopod a

small bisetose segment. Maxilliped with unarmed syncoxa; basis

elongate, with very long endopodal pinnate claw.

P1 with 3-segmented rami; basis without outer seta, inner spine

sexually dimorphic; exp-1 elongate, exp-2 without inner seta, exp-3

with 4 elements; endopod with formula [1.0.020].

P2—-P4 with 3-segmented exopods and 1-segmented endopods.

Bases without outer seta. Exp-1 markedly elongate; exp-2 with very

strong inner seta; apical setae of exp-3 very long. Endopods minute,

with very long apical seta(e). Armature formula:

Exopod Endopod

b2 0.1.022 010

P3 0.1.022 020

P4 ORII22. 010

PS rudimentary with baseoendopod fused to somite and repres-

ented only by outer basal seta. Exopod a small segment with | seta

in 2 and | multipinnate fused spine plus | setain d. P6 rudimentary,

forming unarmed median operculum in 9; asymmetrical in d (with

dextral or sinistral configuration), represented by opercular un-

armed plate.

Caudal ramus short, with 7 setae; seta V longest.

Spermatophore extraordinarily large, longer than half the body

length.

TYPE AND ONLY SPECIES. Jnermipes humphreysi gen. et sp. nov.

ETYMOLOGY. The generic name alludes to the absence of the outer

seta on the bases of the swimming legs. Gender: feminine.

Inermipes humphreysi sp. nov.

TYPE LOCALITY. Barrow Island, Western Australia (37°50'46"N,

31°31'35"W), stn BES 792, borehole, 50—55m deep, | December

1992.

TYPEMATERIAL. Holotype 2 dissected on 9 slides (WAM C24414).

Paratypes deposited in NHM are 12 dissected on 8 slides (BMNH

W. LEE AND R. HUYS

1999.1106), BES 792, depth 50-55m, 1 December 1992 and 922, 9

36 in 70% alcohol (BMNH 1999.1107—1124), BES 793, 1 Decem-

ber 1992. Paratypes deposited in WAM are (a) 1 6 dissected on 7

slides, and 499, 6 dd in 70% alcohol, BES 792, depth 50—-55m, 1

December 1992, (b) 1 6d in 70% alcohol, BES 798, depth 45—50m,

2 December 1992, and (c) 522, 15 36 in 70% alcohol, BES 810,

depth 50-55m, 2 December 1992 (WAM C24415-24417). All

specimens are from the type locality and were collected by Dr W.F.

Humphreys using a plankton net with a 125 m mesh and of a size

suitable for the borehole (Pesce et al., 1996a).

DESCRIPTION

FEMALE. ‘Total body length 540-667um (n=10; mean = 592 um;

measured from anterior margin of rostrum to posterior margin of

caudal rami). Largest width measured at posterior margin of genital

double-somite: 106m. Genital double-somite swollen. No distinct

demarcation between urosome and prosome (Fig. 1B).

Cephalothorax with smooth posterior margin; pleural areas small

and rounded; sensillae and few pores present as illustrated in Fig.

1A-B. Rostrum not defined at base (Fig. 1B), with pair of tiny

sensillae near apex.

Pedigerous somites completely smooth. All prosomites without

defined hyaline frills; hind margin smooth; separated by large

membranous areas. Body not markedly constricted between indi-

vidual somites. P4-bearing somite distinctly narrower than preceding

ones.

Urosome (Fig. 1!A—B) 5-segmented, comprising P5-bearing

somite, genital double-somite and 3 free abdominal somites. P5-

bearing somite, and genital double-somite without surface

ornamentation, except for dorsal pairs of sensillae. Free abdominal

somites with several rows of minute spinules laterally (Fig. 1A) and

ventrally (Fig. 2C); hyaline frills of urosomites not present. Ovary

large, about half of body length (Fig. 1A—B).

Genital double-somite (Figs 1A-B, 2C) wider than long; com-

pletely fused, with weakly pronounced constriction bilaterally

(possibly indicating original segmentation). Genital field (Fig. 2C)

with small copulatory pore (arrowed in Fig. 2C) located in median

depression; gonopores fused medially forming single genital slit

covered on both sides by opercula derived from sixth legs; no

distinct armature discernible; seminal receptacles fused forming

large median sac.

Anal somite (Fig. 2C—D) with well developed spinulose opercu-

lum flanked by spinular rows; with pair of dorsal sensillae anterior

to operculum. Caudal rami (Fig. 2C—D) longer than wide; each

ramus with 7 setae; setae I-III and VI-VII bare, setae IV and V with

minute spinules; setae I and II positioned dorsolaterally with seta I

being much shorter than seta IJ; seta III positioned laterally; setae 1V

and V fused basally, each with predesigned fracture planes at base;

seta V longest (longer than whole urosomal segments combined);

seta VI small; seta VII tri-articulate at base. Bases of setae typically

surrounded by few tiny spinules. Inner margin of each ramus with

dorsolateral concavity.

Antennule (Fig. 3A) 8-segmented, slender; with well developed

sclerite around base of segment 1; all setae bare. Segment 1 without

seta; segment 3 longest; segment 4 with aesthetasc. Armature for-

mula: 1-[0], 2-[7], 3-[7], 4-[2 + (1 + ae)], 5-[2], 6-[2], 7-[4], 8-[3 +

acrothek]. Apical acrothek consisting of slender aesthetasc fused

basally to 2 setae. All segments without surface ornamentation.

Antenna (Fig. 3B) 4-segmented. Coxa small, without ornamenta-

tion. Basis with spinules along proximal outer margin, and distal

inner margin; abexopodal seta absent. Exopod minute, 1-segmented,

with | pinnate apical seta. Endopod 2-segmented; proximal segment

with 2 rows of spinules along abexopodal margin; distal segment

NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER

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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, PS [normal]; H, PS [aberrant].

NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER

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Fig. 3 Inermipes humphreysi gen. et sp. nov. Female: A, antennule; B, antenna; C, mandible; D, maxilliped. Male: E, antennule.

longer than proximal; lateral armature arising in proximal half,

consisting of | pinnate and 2 bare setae; apical armature consisting

of 5 geniculate setae (1 geniculate seta fused basally to 1 pinnate seta

and small tube pore; see insert); distal endopod segment with row of

spinules along abexopodal margin and 2 transverse hyaline frills

subapically.

Labrum with elaborate spinular ornamentation and pores as in

Fig. 5A-B.

Mandible (Fig. 3C) with well developed gnathobase bearing

several fine, multicuspidate teeth around distal margin and 1 pinnate

spine at dorsal corner; | row of spinules near base of palp. Palp 2-

segmented; proximal segment without ornamentation; distal segment

with 4 naked setae arranged in 2 pairs; subapical pair fused basally.

Paragnaths (Fig. 5A) strongly developed lobes with medially

directed hair-like setules, separated by medial lobe covered with

dense pattern of short setules.

Maxillule (Figs 2A, SA—B). Praecoxa without ornamentation;

arthrite elongate, strongly developed, with 2 naked setae along outer

margin, 2 short setae on anterior surface, and 6 spines/setae around

distal margin; 3 distal spines unipinnate and bearing pore (Fig. SA—

B) at base of proximal spinule. Coxa with cylindrical endite bearing

2 naked setae and | curved, pinnate spine. Palp represented by basis

and possibly incorporated rami, forming cylindrical segment with 5

naked setae around distal margin.

Maxilla (Fig. 2B) without surface ornamentation on syncoxa;

medial margin membranous and bearing | coxal endite with 1

pinnate spine and 2 naked setae. Allobasis drawn out into very long,

slightly curved, unipinnate claw; allobasal claw with spinules along

distal half of inner margin and transverse row of posterior spinules

halfway the length; accessory armature consisting of | pinnate spine

on posterior surface, and | tube pore along outer margin. Endopod 1-

segmented, and located on anterior surface of allobasis; ornamented

with 2 naked setae.

Maxilliped (Fig. 3D) without ornamentation on syncoxa. Basis

smooth without ornamentation. Endopod drawn out into very long,

unipinnate claw without accessory armature; longer than basis.

Swimming legs P1—P4 (Fig. 4A—D) with wide intercoxal sclerites

and well developed praecoxae, both without ornamentation. Coxae

and bases with anterior rows of surface spinules as figured. Bases

without outer seta. Exopods 3-segmented, endopods 3-segmented

(P1), or 1-segmented (P2—P4).

P1 (Fig. 4A) with large coxa; with | row of spinules on anterior

surface. Basis with | strong, bipinnate spine and long setules along

inner margin; rows of spinules along outer distal margin, and near

base of exopod. Exp-1 and -2 with | bipinnate slender spine; exp-3

with | bipinnate spine, 1 long unipinnate seta and 2 bipinnate

weakly geniculate setae. Endopod slightly shorter than exopod; enp-

1 with 1 bipinnate inner seta; enp-2 without seta; enp-3 with 2

geniculate setae apically. All segments of exopod and endopod

ornamented with small spinules and setules along outer and inner

margin as figured.

P2—P4 (Fig. 4B—D). Coxae without ornamentation; basis with

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

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

strong inner seta on enp-2, outer exopodal spines slender, terminal

setae very long. Endopods minute, represented by small segment.

Spine and setal formulae as in generic diagnosis.

Fifth pair of legs (Fig. 2C, E) fused to supporting somite.

Baseoendopod represented by small, outer setophore bearing basal

W. LEE AND R. HUYS

seta; endopodal lobe not expressed. Exopod forming minute seg-

ment with | bipinnate apical seta.

MALE. Body larger than in 2. Somites bearing P2—P4 wider than

in 2. Body length 515-690 um (n=20; mean = 622 um; measured

from anterior margin of rostrum to posterior margin of caudal rami).

Largest width measured at P6-bearing somite: 109 um.

Prosome (Fig. 1C) 4-segmented, comprising cephalothorax and 3

free pedigerous somites. No distinct demarcation between urosome

and prosome. Cephalothorax with smooth posterior margin; pleural

areas small and rounded; sensillae and few pores present as illustrated

in Fig. 1C. Rostrum not defined at base (Fig. 1C), with pair of tiny

sensillae near apex.

Pedigerous somites not covered with spinules. All prosomites

without defined hyaline frills but separated by large membranous

areas. Body without marked constrictions between individual

somites; P4-bearing somite not distinctly narrower than others.

Urosome (Fig. 1C) 6-segmented, comprising P5-bearing somite,

genital somite and 4 abdominal somites. P5-bearing somite and

genital somite without surface ornamentation, except for pairs of

dorsal sensillae. Free abdominal somites, including anal somite,

with several rows of spinules laterally and ventrally; dorsal hind

margin smooth without any ornamentation. Hyaline frills of

urosomites not present. Spermatophore extraordinarily large, about

half of body length.

Antennule (Figs 3E, 5C) 10-segmented; haplocer, with

geniculation between segments 7 and 8. Segment | longest, un-

armed. Some elements on segments 4, 6 and 8 (inserts in Fig. 3E;

Fig. 5C) very small. Segment 5 not swollen. Segment 8 with | small

fused spine in median anterior margin. Armature formula: 1-[0], 2-

[7], 3-[6], 4-[1 + 1 small spine], 5-[4 + (1 + ae)], 6-[1], 7-[1], 8-[1 +

1 fused spine], 9-[4], 10-[5 + acrothek]. 1 seta on fifth segment very

small [arrowed in Fig. SC). Apical acrothek consisting of slender

aesthetasc and 2 naked setae.

Inner basal spine of P1 modified into robust barbed element (Fig.

4E). P2—P4 as in &.

Fifth pair of legs (Figs 2F—H, 5D) fused to supporting somite.

Baseoendopod with short setophore bearing outer basal seta;

endopodal lobe not expressed. Exopod with 2 elements; 1 apical

tripinnate spine fused to exopod, and | small slender bipinnate seta

on subapical outer margin (Figs 2G, 5D); abnormal exopod (Fig.

2H) resembling female condition frequently observed on one or

either side; only 1 out of 9 males showed normal P5 exopod on both

sides.

Sixth pair of legs (Figs 2F, 5D) asymmetrical; represented on both

sides by small membranous plate (fused to ventral wall of support-

ing somite along one side; articulating at base and covering gonopore

along other side; dextral and sinistral configurations present); with-

out additional ornamentation.

ETYMOLOGY. The species is named in honour of Dr William F.

Humphreys (Western Australian Museum, Perth), who collected the

material.

DISCUSSION

Inermipes gen nov. is placed in the family Ameiridae on account of

the morphology of the antennules, mouthparts and the sexual dimor-

phism in the inner basal spine of leg 1. Within the group of

freshwater ameirids it can be readily identified by the unisetose

antennary exopod, the lack of the outer basal seta on P1—P4 the

extreme reduction in the swimming legs (particularly the endopods)

NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER

50u

d basis of P1.

sp. nov. Female: A, P1; B, P2; C, P3: D, P4. Male: E, coxa an

Fig. 4 Inermipes humphreysi gen. et

46 W. LEE AND R. HUYS

Fig.5 Jnermipes humphreysi gen. et sp. nov. SEM micrographs. A, oral area showing labrum, paragnath and part of maxillule; B, oral area (different

angle); C, antennule d [tiny seta on segment 5 arrowed]; D, abdomen d, showing P5 and P6, ventral. Scale bars: 5 um (A-B), 2 um (C), 20 um (D).

NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER

and the 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 size of the

spermatophore, occupying nearly half of the body, is remarkable.

Except for representatives of the genus Apodopsyllus Kunz

(Paramesochridae) where similarly sized spermatophores have been

reported, male harpacticoid copepods produce small spermatophores,

typically not exceeding the length of two body somites. Given the

highly reduced genital apertures of /. humphreysi it is difficult to

imagine how the spermatophore can be successfully extruded and

transferred to the female.

Taxonomy of freshwater Ameiridae

The primary taxonomic subdivisions in freshwater Ameiridae have

traditionally been based on swimming leg segmentation (Lang,

1965; Petkovski, 1976) and have ignored other, more phylo-

genetically informative characters such as setation patterns and

mouthpart morphology (Galassi et al., 1999). This simplistic

approach has led to: (1) the generally unsatisfactory practice of

describing new species virtually exclusively on leg characters with-

out consideration of cephalic appendages, genital field morphology

or even female abdominal segmentation; (2) the establishment of

unnatural genera such as Stygonitocrella, and (3) the blurring of

generic boundaries. Central to this confusion stands the genus

Nitocrella which has served as a taxonomic repository for freshwa-

ter Ameiridae since its proposal by Chappuis (1924). Lang (1965)

removed all species displaying 3-segmented P2-P4 endopods to a

new genus Parapseudoleptomesochra and created a second genus,

Pseudoleptomesochrella, to accommodate all Nitocrella species

characterized by 2-segmented P2—P4 endopods and the presence of

an inner seta on P2—P4 exp-1. Even under its revised taxonomic

concept the genus Nitocrella continued to accumulate a large number

of new species which prompted Petkovski (1976) to subdivide the

genus even further. He suggested to group only species with 2-

segmented P2—P4 endopods in Nitocrella s. restr. and to reallocate

all remaining species with alternative endopodal segmentation in

two new genera, Nitocrellopsis (P2—P3 3-segmented, P4 2-seg-

mented) and Stygonitocrella (P2—P3 1—2-segmented, P4

1-segmented). Petkovski (1976) also recognized three subgroups in

Nitocrella based on the number of armature elements on P4 exp-3:

the hirta- (3-4 setae/spines), chappuisi- (5 setae/spines) and

vasconica-groups (6 setae/spines). Although these groups have met

with general acceptance, their monophyletic status has never been

challenged. Furthermore, since Petkovski (1976) did not designate a

type species for Stygonitocrella, nor for Nitocrellopsis, both generic

names were unavailable until recently. Galassi et al. (1999) fixed N.

rouchi Galassi, De Laurentiis & Dole-Olivier, 1999 as the type of

Nitocrellopsis, making the name available with their authorship. A

similar course of action was taken by Reid ef al. (in press) who

designated S. montana (Noodt, 1965) as the type of Stygonitocrella.

Leg segmentation characters should be used with caution when

inferring relationships in derived lineages. For example, the evolu-

tionary instability of endopodal segmentation is illustrated by the

genus Psammonitocrella Rouch which contains species with 1- (P.

longifurcata) or 2-segmented P2—P3 endopods (P. boultoni). Unique

derived characters such as the loss of the outer spine on P| exp-2

leave no doubt that both Psammonitocrella species shared a com-

mon ancestor, and hence the discrepancy observed in endopodal

segmentation has to be interpreted as the result of intrageneric

evolution. Utilizing endopodal segmentation patterns in defining

generic boundaries is potentially misleading. Overweighing such

characters at the expense of others can result in assigning species to

the wrong genus. For example, Nitocrella petkovskii Pesce, 1980

and Stygonitocrella colchica (Borutzky & Michailova-Neikova,

1970) are very closely related, differing essentially only in the

expression of the segment boundary between P4 enp-1 and -2, but

are nevertheless placed in different genera. The close zoogeographical

connection between N. petkovskii (NW Iran) and S. colchica (W

Georgia) is noteworthy in this context.

I. humphreysi shows superficial similarities to both Psammoni-

tocrella and Stygonitocrella, currently the most advanced genera

within the Ameiridae, however the combined presence of a sexually

dimorphic inner basal spine on Pl, a completely fused genital

double-somite, reduced antennary exopod and vestigial PS excludes

it from either genus. The genus Psammonitocrella was proposed by

Rouch (1992) to accommodate two interstitial species collected in

the hyporheic zone of an intermittent desert stream in Arizona. Its

familial placement has been questioned by Martinez Arbizu &

Moura (1994) who removed the genus from the Ameiridae and

regarded it as the sistergroup of the Parastenocarididae. This course

of action was primarily based on the loss of the outer spine of P1

exp-2 and the absence of a sexually dimorphic inner basal spine on

leg 1. The juvenile morphology of the P5 in both sexes and the

presence of separate genital and first abdominal somites in the adult

female strongly suggest a paedomorphic origin for Psammonitocrella.

The absence of sexual dimorphism in the inner basal spine of leg 1

should be re-evaluated in this context. The modification of this spine

in male Ameiridae appears not until the final moult. Delaying the

expression of this character beyond the final moult (post-displace-

ment) would result in the secondary loss of sexual dimorphism. The

absence of a modified inner basal spine in Psammonitocrella is

therefore to be regarded as autapomorphic and it is proposed here to

remove the genus from its floating status and to reallocate it to the

Ameiridae.

Similarities between /nermipes and Psammonitocrella are found

in the presence of only two setation elements on the distal endopod

segment of leg 1, the absence of the outer basal seta in P1—P2, and

the reduced P4 endopod. It is conceivable that these shared characters

are the product of convergence since both genera differ significantly

in the morphology of the antenna, maxilla, maxilliped and swim-

ming leg ornamentation.

The supposedly cosmopolitan genus Stygonitocrella was diag-

nosed solely on the basis of the 1-segmented P4 endopod (Petkovski,

1976). Comparison of the swimming leg setal formulae of the 13

species currently included strongly indicates the presence of several

discrete lineages within this genus, each exhibiting a typical arma-

ture pattern (Table 1) and a restricted geographical distribution

(Table 2). This subdivision is admittedly based on swimming leg

characters only but we suspect it to be at least partly reinforced by

mouthpart and antennulary characters when they become available.

Formal recognition of these lineages as distinct genera is impossible

since most descriptions are severely lacking in detail and many of

them are based on very few specimens or one sex only. For example,

the description of P. djirgalanica is completely lacking in illustra-

tions (Borutzky, 1978). In addition, the type material of the great

majority of its species is no longer extant and additional records

have not been added. S. petkovskii differs from its congeners in the

absence of the inner seta on P3 exp-2, an element which is present in

all other congeners. Attempts to trace the single female on which this

description was based failed (Galassi, pers. comm. ).

As pointed out by Reid er al. (in press) the generic placement of

S. orghidani (Petkovski, 1973) is questionable. The original descrip-

tion is very concise, showing illustrations only of the antenna,

caudal rami and the fifth legs. The exopodal armature of the swim-

ming legs is largely unknown, apart from Petkovski’s statement that

W. LEE AND R. HUYS

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NEW AMEIRIDAE FROM W. AUSTRALIAN GROUNDWATER

Table 2 Species groups within Stygonitocrella showing zoogeographical connection, habitat and number of specimens known

Species Distribution Habitat Material

dubia (Chappuis, 1937) North Spain cavernicolous Id

guadalfensis Rouch, 1985 Andalusia hyporheic 929.466

karamani (Petkovski, 1959) Macedonia phreatic DOO ISS

ljovuschkini (Borutzky, 1967) Ukraine cavernicolous Sh LP) Chis

colchica (Borutzky & Michailova-Neikova, 1970) West Georgia cavernicolous 3 29.446

tianschanica (Borutzky, 1972) Kirghiztan phreatic PSII) (oie)

pseudotianschanica (Stérba, 1973) Afghanistan phreatic 2 ISN,

djirgalanica (Borutzky, 1978) Kirghiztan phreatic VEL POS

montana (Noodt, 1965) Argentina phreatic 422.636

orghidani (Petkovski, 1973) Cuba cavernicolous 499.246

insularis (Miura, 1962) Ryukyu Islands phreatic 2 Ped

the distal exopod segment of P2—P4 possesses an inner seta. The

species is unusual in having a bisetose antennary exopod and

remarkably reduced fifth legs which consist of a median transverse

plate retaining only the outer basal setae. Petkovski (1973) points

out several similarities between S. orghidani and Nitocrella negreai

such as the large size of the antennules and antennae, the short

caudal rami and the elongate Pl endopod. He regarded these

characters as evidence for a distinct lineage within Nitocrella sensu

lato. Apart from the 1-segmented P4 endopod there is no evidence

for a relationship with any of the other species currently included in

Stygonitocrella and in view of its fragmentary description we pro-

pose to relegate it to species incertae sedis within this genus.

The only oriental species, S. insularis, was described from the

Ryukyu archipelago (Miura, 1962a) and shows a number of unique

characteristics within Stygonitocrella. It is regarded here as the type

species of a new genus Neonitocrella on account of the bisetose

antennary exopod, the loss of the P4 endopod (represented by a

rudimentary knob) and the reduced P5 in both sexes. The Pl

armature pattern, the reduced antennary exopod and the character-

istic structure of the male PS exopod suggest a sistergroup relationship

with /. humphreysi, to which it is also closest zoogeographically.

Conversely, Nitocrella japonica described from Shikoku Island

(Japan) (Miura, 19626) also shows a remarkable similarity with /.

humphreysi in the structure of the antenna (unisetose exopod),

maxilla (elongate allobasis) and maxilliped (slender basis and claw)

but exhibits a more primitive leg formula. This suggests that the

genus /nermipes could have been derived from a N. japonica-like

ancestor through swimming leg reduction.

Genus NEONITOCRELLA gen. noy.

DIAGNOsIS. Ameiridae. Body elongate, cylindrical, and vermi-

form without distinct surface ornamentation except for ventral

spinule patterns on abdomen. Cephalothorax and other somites with

smooth posterior margin. Genital and first abdominal somites appar-

ently separated. Anal operculum well developed.

Sexual dimorphism in antennule, P3 endopod, P5, P6, and in

genital segmentation; unconfirmed for inner basal spine of P1.

Rostrum fused to cephalothorax, not defined at base. Antennule 8-

segmented in 2, unknown in d; segment | with seta; aesthetascs on

segments 4 and possibly 8 in 9, unknown in 6. Antenna with separate

basis and endopod; exopod well developed segment, with 2 setae.

Mandible with 2-segmented palp, comprising unarmed basis and

endopod with 4 setae. Maxillule, maxilla and maxilliped unconfirmed.

P1 with 3-segmented rami; basis without outer seta; exp-1 not

elongate, exp-2 without inner seta, exp-3 with 4 elements; endopod

with formula [1.0.020].

P2—P4 with 3-segmented exopods and 1-segmented (P2—P3) or

rudimentary (P4) endopods. Bases with outer seta. Apical setae of

exp-3 very long. P3 endopod ¢ with inner distal seta shorter and

outer distal spine longer than in °. P4 endopod represented by minute

knob. Armature formula:

Exopod Endopod

P2 0.1.022 020

P3 0.1.022 020

P4 0.1.122 -

P5 rudimentary with baseoendopod fused to somite and repres-

ented only by outer basal seta. Exopod a small segment with 2 setae

in and | pinnate fused spine plus 2 setae in d. P6 rudimentary,

forming unarmed median operculum in 2; unconfirmed in d.

Caudal ramus short, with 7 setae; seta V longest.

TYPE AND ONLY SPECIES. WNitocrella insularis Miura, 1962 =

Neonitocrella insularis (Miura, 1962) comb. nov.

ACKNOWLEDGEMENTS. The authors would like to thank Dr W.F.

Humphreys (Western Australian Museum, Australia) for providing us with

the copepod material. One of us (W.L.) acknowledges financial support from

the Korea Research Foundation provided for the programme year 1997.

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7. Make sure you distinguish numerals from letters, e.g. zero (0)

from O; one (1) from | (el) and I.

8. Distinguish hyphen, en rule (longer than a hyphen, used

without a space at each end to signify ‘and’ or ‘to’, e.g. the Harrison—

Nelson technique, 91-95%, and increasingly used with a space at

each end parenthetically), and em rule (longer than an en rule, used

with a space at each end parenthetically) by: hyphen, two hyphens

and three hyphens, respectively. Be consistent with rule used paren-

thetically.

9. Use two carriage returns to indicate beginnings of paragraphs.

10. Be consistent with the presentation of each grade of heading

(see Text below).

Title. The title page should be arranged with the full title; name(s)

of author(s) without academic titles; institutional address(es); sug-

gested running title; address for correspondence.

Synopsis. Each paper should have an abstract not exceeding 200

words. This should summarise the main results and conclusions of

the study, together with such other information to make it suitable

for publication in abstracting journals without change. References

must not be included in the abstract.

Text. All papers should have an Introduction, Acknowledge-

ments (where applicable) and References; Materials and Methods

should be included unless inappropriate. Other major headings are

left to the author’s discretion and the requirements of the paper,

subject to the Editors’ approval. Three levels of text headings and

sub-headings should be followed. All should be ranged left and be in

upper and lower case. Supra-generic systematic headings only

should be in capitals; generic and specific names are to be in italics,

underlined. Authorities for species names should be cited only in the

first instance. Footnotes should be avoided if at all possible.

References. References should be listed alphabetically. Authori-

ties for species names should not be included under References,

unless clarification is relevant. The author’s name, in bold and lower

case except for the initial letter, should immediately be followed by

the date after a single space. Where an author is listed more than

once, the second and subsequent entries should be denoted by a long

dash. These entries should be in date order. Joint authorship papers

follow the entries for the first author and an ‘&’ should be used

instead of ‘and’ to connect joint authors. Journal titles should be

entered in full. Examples: (i) Journals: England, K.W. 1987. Certain

Actinaria (Cnidaria, Anthozoa) from the Red Sea and tropical Indo-

Pacific Ocean. Bulletin of the British Museum (Natural History),

Zoology 53: 206-292. (ii) Books: Jeon, K.W. 1973. The Biology of

Amoeba. 628 p. Academic Press, New York & London. (iii) Articles

from books: Hartman, W.D. 1981. Form and distribution of silica in

sponges. pp. 453-493. In: Simpson, T.L. & Volcani, B.E. (eds)

Silicon and Siliceous Structures in Biological Systems. Springer-

Verlag, New York.

Tables. Each table should be typed on a separate sheet designed to

extend across a single or double column width of a Journal page. It

should have a brief specific title, be self-explanatory and be supple-

mentary to the text. Limited space in the Journal means that only

modest listing of primary data may be accepted. Lengthy material,

such as non-essential locality lists, tables of measurements or details

of mathematical derivations should be deposited in the Biological

Data Collection of the Department of Library Services, The Natural

History Museum, and reference should be made to them in the text.

Illustrations

DRAWINGS — Figures should be designed to go across single (84 mm

wide) or double (174 mm wide) column width of the Journal page,

type area 235 x 174 mm. Drawings should be in black on white stiff

card with a line weight and lettering suitable for the same reduction

throughout, ideally not more than 40%. After reduction the smallest

lettering should be not less than 10 pt (3 mm). Tracing paper should

ideally be avoided because of the possibility of shadows when

scanned. All artwork must have bulletin, author and figure number

included, outside of the image area, and must be free of pencil, glue

or tape marks.

PHOTOGRAPHS — All photographs should be prepared to the final size

of reproduction, mounted upon stiff card and labelled with press-on

lettering (eg Letraset). They can be mounted on white or black

background; a black background must be evenly black all over; any

background must be free of all pencil and glue marks within the

image area. All figures should be numbered consecutively as a

single series. Legends, brief and precise, must indicate scale and

explain symbols and letters. Photos, when components of figure-

plates should be abutted, trimmed as regular rectangles or close

trimmed up to edge of specimen. Joins etc. can be removed at the

scanning stage but at extra cost. Cropping instructions, if any, should

be indicated on an overlay or marked on a photocopy of the figure.

SIZE — Maximum size of artwork for use of flatbed scanners is A3.

Larger artwork has to be reduced photographically prior to scan-

ning, therefore adding to expense.

Symbols in text. Male and female symbols within the text should

be flagged within curly brackets to enable setter to do a swift global

search.

Reprints. 25 reprints will be provided free of charge per paper.

Orders for additional reprints can be submitted to the publisher on

the form provided with the proofs. Later orders cannot be accepted.