••-V

Bulletin of the

British Museum (Natural History

s c*»«

, LIBRARY

Zoology series Vol 45 1983

British Museum (Natural History)
London 1983

Dates of publication of the parts

No 1 28 July 1983

No 2 28 July 1983

No 3 25 August 1983

No 4 25 August 1983

No 5 29 September 1983

No 6 27 October 1983

No 7 .24 November 1983

ISSN 0007-1498

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

Contents
Zoology Volume 45

Page

No 1 Problems in catfish anatomy and phylogeny exemplified by the
Neotropical Hypophthalmide (Teleostei: Siluroidei).
By G. J. Howes 1

No 2 Miscellanea

A revision of the genus Epiclintes (Ciliophora: Hypotrichida) includ-
ing a redescription of Epiclintes felis comb. n.
By P. G. Carey & E. C. Tatchell 41

Notes on the Family Lekythoporidae (Bryozoa, Cheilostomata).

By P. L. Cook & P. J. Hayward 55

A new species of Arthrolepis (Anura: Ranidae) from the West

Usambara Mountains, Tanzania.

By A. G. C. Grandison 77

The distribution behavioural ecology and breeding strategy of the

Pygmy Toad, Mertensophyrne micranotis (Lov.).

By A. G. C. Grandison & S. Ashe 85

Additional notes on bariliine cyprinid fishes.

By G. Howes 95

No 3 Bats (Mammalia: Chiroptera) from Indo- Australia.

ByJ. E. Hill . 103

No 4 On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the
interrelationships of African cichlid species flocks.
By Peter Humphry Greenwood 209

No 5 Miscellanea

D enter ammina (Lepidodeuterammind) subgen. nov., and a re-
description of Rotalina ochracea Williamson (Protozoa: Forami-
niferida).
By P. Bronnimann & J. E. Whittaker . . . ... .233

The freeliving marine nematode genus Sabatieria (Nematoda:

Comesomatidae). I. Two new species from Stonington Island,

Antarctica.

By H. M. Platt 239

New species of marine nematodes from Qingdao, China.

By Z. N. Zhang & H. M. Platt .253

Echinoderms of the Rockall Trough and adjacent areas. I. Crinoi-

dea, Asteroidea and Ophiuroidea.

By J. D. Gage, Margaret Pearson, Ailsa M. Clark, G. L. J. Paterson

& P. A. Tyler 263

No 6 The cranial muscles of loricarioid catfishes, their homologies and
value as taxonomic characters (Teleostei: Siluroidei).
By Gordon J. Howes . . 309

No 7 Miscellanea

A lectotype for Deuterammina (Deuterammind) rotaliformis
(Heron-Allen & Earland) and new trochamminids from E. Ireland
(Protozoa: Foraminiferida).
By P. Bronnimann & J. E. Whittaker . . . . . .347

Notes on Atlantic and other Asteroidea. 3. The families Ganeriidae

and Asterinidae, with description of new asterinid genus.

By Ailsa M. Clark .... .... 359

Peniculus haemuloni, a new species of copepod (Siphonostoma-
toida: Pennellidae) parasitic on Haemulon steindachneri from
Ubatuba, Brazil.
By P. D. Alexander .381

Bulletin of the

British Museum (Natural History)

Problems in catfish anatomy and
phylogeny exemplified by the Neotropical
Hypophthalmidae (Teleostei: Siluroidei)

Gordon J. Howes

Zoology series Vol 45 No 1 28 July 1983

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in for
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World List abbreviation: Bull. Br. Mm. nat. Hist. (Zool.)

© Trustees of the British Museum (Natural History), 1983

The Zoology Series is edited in the Museum's Department of Zoology
Keeper of Zoology : Dr J. G. Sheals
Editor of Bulletin : Dr C. R. Curds
Assistant Editor : Mr C. G. Ogden

ISSN 0007-1498 Zoology Series

VoUSNo 1 pp 1-39

British Museum (Natural History)

Cromwell Road

London SW7 5BD Issued 28 July 1983

j WU«bTWH

Problems in catfish anatomy and phylogeny
exemplified by the Neotropical Hypophthalmidae
(Teleostei: Siluroidei)

Gordon J. Howes

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW75BD

Contents

Introduction 1

List of genera examined 3

Form and function in Hypophthalmus and comparison with other siluroids . 4

Body shape, sensory canals and lateral line system 4

The eye and infraorbital bones 6

Cranial musculature . . . . • 9

The snout and jaws 17

The cranium 21

The complex vertebrae, swimbladder and posterior lateral line nerve . . 25

The suspensorium 39

The hyoid arch 32

The opercular series 34

The gill arches 34

Postcranial elements 35

Relationships of the Hypophthalmidae 35

Acknowledgements 33

References 38

Introduction

Of the subgroups of otophysan fishes, the siluroids — or catfishes — are the most morpho-
logically diverse. None of the other subgroups, gymnotoids, characoids and cyprinoids, has
such a range of body shape, peculiarity of dermal covering when present, and cranial
morphology. Neither do these groups display such a range of postcranial architecture,
complex fusion patterns of the caudal fin skeleton or such diverse forms of swimbladder.
This morphological diversity is reflected in the most recent familial classification of the
siluroids (Greenwood et al., 1966), which recognises thirty-one families, a number exceeding
the combined familial total of other otophysan groups.

As presently recognised, most siluroid families are morphologically well differentiated,
suggesting that each is probably a monophyletic assemblage. Indeed, the large morphological
'gaps' separating families are emphasised by the fact that seven families are monotypic, cf
one among characoids and two among gymnotoids. That the interrelationships of siluroid
families have scarcely been studied is not surprising when one considers the paucity of
anatomical data available. Even the largest families, i.e. the Pimelodidae, Ariidae and
Bagridae are represented in the anatomical literature by only meagre information, the
anatomy of some neotropical families, Auchenipteridae, Ageneiosidae, and Aspredinidae,
has scarcely been described.

The siluroids offer a wealth of character complexes involving the bony elements of the
cranium, vertebral column and fin supports, and their associated muscles. These character

Bull. Br. Mus. nat. Hist. (Zool.) 45( 1 ) : 1-39 Issued 28 July 1983

2 G. J. HOWES

complexes lend themselves readily to the application of cladistic methodology. The task of
analysis is, however, a daunting one. Not only is the number of taxa considerable, over 2000
species, but, by their very nature, siluroids are difficult to study. Most adult specimens are
large and highly ossified, and sutures between bones are often obscured in dry skeletal
material. It is therefore necessary to work, for the most part, with cleared and stained prep-
arations of small, and frequently juvenile specimens. Thus problems can arise in the
interpretation of features which undergo considerable ontogenetic change.

One of the generalisations made about siluroids is that they are benthic fishes. In fact, the
majority of species in seven families, Siluridae, Schilbeidae, Pangasiidae, Ariidae,
Ageneiosidae, Cetopsidae, Hypophthalmidae, are mid- water to surface dwellers, and only in
one third of families, Loricariidae, Amphiliidae, Callichthyidae, Astroblepidae, Doradidae,
Chacidae, Sisoridae, Cranoglanididae, can the majority of included taxa be classed as
entirely benthic. In the past more attention has been given to siluroid inter- than to intra-
relationships. The most recent analysis (Fink & Fink, 1981) recognises the siluroids as a
suborder, the Siluroidei, coordinate with their supposed sister group, the Gymnotoidei
which together form the order Siluriformes. The Siluriformes are, in turn, thought to be the
sister group of the characoids, order Characiformes, with the cyprinoids, order
Cypriniformes, representing the plesiomorph lineage and sister group of the Silurifomes +
the Characiformes.

Only the most rudimentary analyses have been attempted on the functional anatomy of
siluroids (Eaton, 1948; Alexander, 1965; Gosline, 1973 & 1975). According to Gosline
(1973) siluroids employ a feeding mechanism functionally different from that of other
otophysans. But again, lack of comparative anatomical data prohibits any worthwhile
discussion of this issue.

One attempt at cladistic analysis involving several siluroid families has been published by
Lundberg & Baskin (1969). In order to obtain an understanding of siluroid relationships
these authors chose to focus efforts '. . . on comparative studies of a single structural complex
in obtaining an understanding of siluroid relationships'. This approach, whilst accumulating
valuable comparative data, can however, produce a misleading phylogeny. The weakness
inherent in an approach that places reliance on single character complexes is discussed
below (p. 27) with reference to patterns of vertebral fusion and swimbladder encapsulation.

Roberts (1973) notes that the '...cosmopolitan distribution of siluroids and their
pseudarchaic characters have led ichthyologists to believe catfishes to be older than other
ostariophysans'. I presume 'pseudarchaic' as used by Roberts means 'neomorph' resulting in
convergences with features found in ancient groups. But, neomorphs are no indication of the
group's phyletic age. On the other hand, the cosmopolitan distribution of siluroids may
suggest an early and extensive distribution whereas the more restricted distribution of other
otophysans could indicate their later cladogenesis from the ancestral otophysan lineage.

The subject of this paper, the family Hypophthalmidae, is chosen for anatomical investi-
gation because it represents several aspects of the problems discussed above. The family is
monotypic, apparently 'isolated' phyletically, is 'pelagic' in habits, and poses intriguing
problems of functional anatomy. The object of the study is to describe the osteological and
myological features, with particular regard to the cranium, to analyse these features in terms
of their apo-plesiomorphic status, to account for their functional relevance, and to use
these data as indicators of phyletic relationship.

Giinther (1864:3:66) first recognised the distinctiveness of Hypophthalmus by placing it in
a separate subfamily, Siluroidae Anomalopterae (Group Hypophthalmina). Helogenes was
also included on the basis of its elongate anal fin, and its short dorsal and adipose fins. Cope
(1871) established the Hypophthalmidae as one of three families of his Nematognathi. In
Cope's view Hypophthalmus was so aberrant that he doubted its relationships with other
catfishes. Eigenmann & Eigenmann (1890) continued to recognise Gunther's concept of the
family but Regan (1911) placed Helogenes in a separate family thereby establishing the
Hypophthalmidae as monotypic. Chardon (1968) chose to recognise the uniqueness of the

CATFISH ANATOMY & PHYLOGENY 3

Hypophthalmidae by placing it in its own suborder, the Hypophthalmoidei. He was,
however, unable to relate the taxon closely with another of equal rank.

As yet, the taxonomy of the genus Hypophthalmus has received no thorough treatment.
Eigenmann & Eigenmann (1890) synonymised the six nominate species with H. edentatus,
the type of the genus, stating There is little doubt but that the species have been based on the
different stages of one species'. More recently, however, Roberts (1972:139) noted there are
'. . . three very distinct species of Hypophthalmus in the Amazon'. In the most recent
literature authors have recognised a second species named as H. perporosus Cope, 1878 (see
Goulding, 1980;Carvalho, 1980a).

Although Hypophthalmus is a common commercial food fish in Amazonia (see Roberts,
1972: 140), its ecology is poorly studied (for accounts see Roberts, 1972; Meschkat, 1960;
Goulding, 1980; Carvalho, 19800 & b). Hypophthalmus species are shoalers and filter-feed,
chiefly on zooplankton. According to Carvalho (19806), the larger proportion of ingested
food of H. edentatus comprises cladocerans and copepods. Gut contents of specimens
personally examined contain insect (Hymenoptera) leg and antennal fragments, and
fragmented plant material (portions of leaves, stems, pollen cuticles and seed spikes).
Carvalho (19806) classifies Hypophthalmus as a pelagic zooplanktivore but perhaps, in view
of the above catalogue of gut contents, it should more rightly be classed as a detrital filter-
feeder. As in other detritivores, Hypophthalmus has a long, convoluted gut.

Only two papers deal with any detailed aspect of hypophthalmine anatomy. Wright (1885)
described the skull and swimbladder connections. This paper, however, omits some
important features and misinterprets others, omissions obviously due to limitations imposed
by the use of dry skeletal material in which many delicate structures are lost or obliterated.
Chardon (1968) also described the swimbladder and Weberian structure. His account and
illustrations supplement those of Wright's. Roberts (1972) gives a more general account of
Hypophthalmus, and includes a fairly detailed description of the gill arch structure and its
likely mode of function.

List of genera examined

Where availability of material has permitted, the cranial anatomy of at least two species in
each polytypic genus has been examined.

A = alizarin stained specimen, D = dry skeleton, S = alcohol preserved specimens
(dissected).

Aelurichthys D
Ageniosus A, D, S
Amphilius A
Arius A, D, S
Aspredo S

Auchenipterus A, D, S
Bagrus A, D, S
Brachyplatystoma S
Branchioica A
Bunocephalus A
CallophvsusS
Chaca D
Chaetostoma D
Chrysichthys A, D, S
Clarias A, D, S
Clarotes D, S
Cochliodon A
Cryptopterus D, S
Diplomystes D, S

Doras D
Eutropius D, S
Gagata D
Galeichthys D, S
Geneidens D
Glyptosternon D, S
Goldiella S
Helogenes A, D, S
Hemisorubim D, S
Heptapterus S
Hoplosternum D, S
Hypophthalmus A, D, S
Ictalurus A, D, S
Iheringichthys A, S
Leiocassis D
Liobagrus D
Loricaria D, S
Luciopimelodus D, S
Megalonema S

Mystus D
Nematogenys D, S
Notarius D
Osteogeniosus A
Ompok D, S
Oxydoras D
Pangasius S
Pangasianodon D
Parapimelodus S
Perugia S
Phyllonemus A
Pimelodina S
Pimelodus A, D, S
Platysto malic hthys D
Plotosus D, S
Pseudauchenipterus S
Pseudobagrus A, D
Pseudocetopsis S
Pseudopimelodus D, S

4 G. J. HOWES

Pterodoras D Silurus D, S Tandanus D, S

Rhamdia D, S Sorubim A, D, S Trichomycterus A, S

Rhinodoras D Soruhimichthys D, S Wallago D, S

Schilbe A, D, S Synodontis S

Abbreviations used in the figures are given either in the relevant sections of the text, or in the
figure captions.

Form and function in Hypophthalmus and comparison with other siluroids

The principal derived features of Hypophthalmus are listed below. The reasons for
considering their derived nature are discussed under each character complex.

1 . Body shape and pattern of the sensory canals and lateral line system.

2. The position of the eyes and arrangement of their musculature.

3. The hypertrophy of the facial, and the complexity of the hyoid musculature.

4. The particular morphology of the mouth and the articulation of the mandibular barbels
with cartilaginous plates.

5. The reduction of the neurocranium and its complete ankylosis with the anterior vertebral
complex.

6. The nature of the complex vertebrae, anterior placement of the Weberian ossicles,
reduction plus encapsulation of the swimbladder, and displacement of the vagus nerve.

7. The extent of the gill openings and elongation of the branchial arch elements.

8. The elongation and modification of the hyoid arch.

Body shape, sensory canals and lateral line system (Figs 1 & 2)

Hypophthalmus has a deep and moderately compressed body with a sloped dorsal profile;
the head is broad (30% of its length) with a subtriangular snout, the eyes are set at the same
horizontal level as the mouth. There is a single pair of long maxillary barbels and two pairs
of similarly long mandibular barbels. The gill opening extends almost as far forward as the
mandibular symphysis.

The pectoral fin is narrowly triangular, its tip reaching beyond that of the small pelvic fin.
The first ray of the pectoral fin is spinous and finely serrated along its outer margin. The anal
fin is long with c. 62 branched rays, extending from the tip of the pelvic to below the adipose
fin. The dorsal fin is short-based with 1 + 6 rays; the small adipose fin lies midway between
the dorsal fin and the origin of the forked caudal fin.

In life, Hypophthalmus edentatus has a silvery blue-green colour, the barbels are dark, the
fins hyaline but with dusky tips to the pectorals, and a dark submarginal band on the anal fin.

The skin above the orbit is permeated with a network of canals which are denser and more
ramified below the frontal margin than elsewhere (Fig. 2). The entry points to this network
are through the pterotic and frontal sensory canal openings. Similar ramified patterns occur
over the antero-dorsal and ventral portions of the operculum. Their respective connections
are with the pterotic and preopercular canals.

The lateral line has dorsal and ventral branches. The longest dorsal branches are angled
caudally at 45° to the main lateral line, some extending almost to the dorsal midline; shorter
branches run cranially at a similar angle, thus forming a network (Fig. 1 ).

The longest ventral branches extend to the base of the anal fin where they curve caudally
almost at a right angle (one or two branches curve anteriorly). Shorter, angled branches cross
the upper part of the vertical lines and form the ventral counterpart of the dorsal network.
The lateral line is bifurcated caudally, the longer arm, itself often sub-branched, extends to
the 6th or 7th ray of the upper caudal lobe. The lower branch hardly reaches the base of the
caudal fin and is absent in some specimens.

CATFISH ANATOMY & PHYLOGENY

Fig. 1 Hypophthalmus edentatus\ (upper) lateral view showing ramifying lateral line system;
(lower) ventral view of head to show inferior position of the eye. Scale = 20 mm. Drawn from
specimen BMNH 1925. 10.28:327.

The body shape of Hypophthalmus is like that described by Alexander (1965) for the
African siluroid Schilbe. The salient features noted by Alexander of the schilbeid morpho-
type are a compressed trunk; long anal fin; short body cavity; reduced dorsal and adipose
fins; deep epaxial musculature reaching well-forward on the cranium and the supraoccipital
bearing a tall median crest onto which most of the dorsal musculature attaches. All these
features apply to Hypophthalmus and occur also in Old-World siluroids (Siluridae e.g.
Ompok, Cryptopterus; Schilbeidae, Pangasiidae) and the Neotropical Auchenipteridae
(Auchenipterus) and Ageniosidae. As such it seems to be a morphotype independently
derived in several lineages. Nonetheless, such a body shape is assumed to be a derived feature
for each group in which occurs as it is always accompanied by some marked, but different
modification of the swimbladder and, or, lateralis system.

Carvalho (19800) found a high proportion of fat in Hypophthalmus (up to 26% body
weight) which varied inversely to the total water content. This high fat content doubtless
contributes to the attainment of neutral buoyancy and appears to be a correlate of a reduced
swimbladder (see p. 28).

A highly ramified lateral line system similar to that in Hypophthalmus is also found in
siluroids and pangasiids, but in no other taxon investigated do the lower branches extend so
far ventrally or curve along the anal fin base. An orbital and opercular canal system like that
of Hypophthalmus occurs similarly only in Luciopimelodus. A ramifying of canals are
present on the cranial and postcranial regions of many siluroids, but generally as separate
twiglets over the head and upper opercular region only. In the Ageneiosidae and
Pimelodidae alone is there a lattice-like pattern aproaching that of Hypophthalmus. Further-
more, only in Brachyplatystoma and Luciopimelodus does such a system of canals cross the
interoperculum. Other siluroids with a similar body shape to Hypophthalmus (e.g.
Pangasius, Schilbe, Ompok} lack an intricate epidermal network.

Alexander (1965) observed that catfishes possessing the schilbeid morphotype (see above)
were more pelagic than others, and accounted for the compressed, tapered body shape in
terms of a specialised locomotion. A principal component of schilbeid locomotion is the
ability to hover by inclining the body at a 45° angle to the surface. It has not been reported
whether Hypophthalmus adopts a similar locomotory attitude. Neither is it clear whether
Hypophthalmus should be classed as 'pelagic' for it seems that these fishes may make
vertical migrations (Meschkat, 1960).

G. J. HOWES

io4-5

io1-3

Fig. 2 Hypophthalmus edentatus lateral view of head showing sensory canal system and infra-

orbitals. Composite drawing.

The eye and infraorbital bones (Figs 2-4)

Two of the more noticeable features of Hypophthalmus are the position of the eye relative
to other parts of the cranium, and the arrangement of the orbital muscles (Fig. 3).

The eye is of moderate size (12% of head length) and is situated on the ventral margin of
the head at the same horizontal level as the mouth. When viewed ventrally the eyes appear as
almost entirely inferior (Fig. 1 ).

The eye muscles, in their origins and position lateral to the adductor muscle complex, are
unlike those of any other siluroid. The anterior muscles are long and strap-like. Both the
superior obliquus (sob) and inferior obliquus (iob) have a common origin from a broad
tendinous sheet that extends dorsally across the adductor mandibulae musculature to attach
to the lateral margin of the frontal and to the lateral ethmoid. The optic nerve and those
supplying the eye muscles all run parallel to one another within an almost vertical band of
connective tissue. As in other siluroids the superior obliquus is innervated by the trochlear
nerve (niv) and the inferior obliquus by a branch of the oculomotor (niii). The rectus internus
(ri) arises from a thin tendinous strand that crosses the face of the adductor complex and runs
between the superior obliquus and inferior obliquus muscles to attach to the anteromedial
surface of the eyeball. The muscle is unnervated by a sub-branch of that branch of the
oculomotor that feeds the inferior obliquus.

The posterior set of muscles, rectus superior, r. inferior and r. externus (Fig. 4, rs; rin; re)
originate from a tendinous band which is strongly attached to the dermis. The former two
muscles are innervated by branches of the oculomotor and the latter by the abducens (nvi).

The optic nerve (nii) is thick and exits from a slit-like foramen to loop over the adductor
muscle complex and pass across its face which is grooved to accommodate it.

The infraorbital bones (io Fig. 2) are, with the exception of the 3rd, reduced to ossifications
of the sensory canal. The 1st infraorbital is axe-shaped, the common morphology of this
element amongst siluroids, the 2nd a slightly curved, cylindrical bone, the 3rd with an

CATFISH ANATOMY & PHYLOGENY

aap

et

ao lo

nso

am

mn

Fig. 3 Hypophthalmus edentatus; (upper) superficial dissection of the head showing position of the
eye muscles in relation to the cheek musculature; (lower) dorsal view of orbital region (eye
muscles removed) with frontal cut away to display the dilatator operculi muscle (do) and
olfactory nerve (ni). Drawn from specimen BMNH 1976.6.18: 123-129. Scales = 5mm.
am = adductor mandibulae; nso = rami of supraorbital nerve trunk.

antero-ventral lamellae, its posterior, tubular portion lying beneath the orbit; the 4th, the
largest of the series, curves dorso-posteriorly to beyond the border of the eye, then, at the
upper level of the orbit, curves dorso-anteriorly to extend across the interspace between the
orbit and the cranium; the 5th a small tubular bone connected to the pterotic.

Although unusual, the ventro-lateral position of the eye in Hypophthalmus is not unique
amongst siluroids. It occurs similarly in Ageniosus, some Auchenipterus and Pangasius
species. However, in all these taxa, the arrangement of the eye muscles is of the more normal
teleost condition.

In both Auchenipterus and Pangasius the eye ball is cupped by the surrounding muscles,
the posterior set passing into the optic foramen. Ageniosus is markedly similar to
Hypophthalmus in the displacement of the optic nerve tract across the adductor muscle face.
However, the musculature is extended laterally and there is a consequent orientation of the
nerve tract, unlike the vertical course followed in Hypophthalmus. Furthermore, the eye
muscle arrangement in Ageniosus is less modified in that the posterior muscles, although
highly tendinous, join in a single cylindrical tract across the adductor mandibulae muscle.

niv

Fig. 4 Hypophthalmus edentatus; the eyeball, associated muscles and nerves (the upper skin
reverted). Drawn from specimen BMNH 1976.6.18: 123-129. Scale = 5 mm.

Only in the pimelodid Sorubimichthys is there a muscle arrangement approaching that of
Hypophthalmus. Although in the former genus the eye is superior in position and the
posterior set of muscles are horizontally aligned, the anterior muscles originate from a
tendinous band above the extensor tentaculi muscle (et Fig. 10).

The inferior orientation of the eyes in Hypophthalmus may provide a partially stereo-
scopic vision which would serve to alert the fish to an approaching predator. The degree of
eye mobility in Hypophthalmus is unknown. The eye muscles are thinner, and appear
'weaker' than in the other taxa investigated, but their complete innervation suggests some
degree of eye-ball movement.

According to Dullemeijer & Barel (1977) the eye in teleosts is a functionally dominant
feature and in their opinion one with little positional plasticity. This is obviously not the
case in Hypophthalmus where the eye has shifted ventrally from the usual siluroid supero-
lateral position. In Hypophthalmus the position of the eye has apparently been 'dictated' by
the form of the adductor muscle complex. This is in direct contrast to phytoplankton feeding
percomorphs (Dullemeijer & Barel, 1977) where the eye has retained a superior position and
consequently only a small area is available in which the adductor muscle can develop and
operate.

In general, the eyes of siluroids appear to have a less dominant role than in other
otophysans or indeed other teleosts. In many siluroids the trend has been for the eyes to have
shifted into a superior position, taken to the extreme in some loricarioids (e.g. Chaetostoma
and Astroblepus] where the orbits are separated by a minimal space.

CATFISH ANATOMY & PHYLOGENY 9

Fink & Fink (1981) claim that the infraorbital series in siluroids is reduced to the canal
bearing portion of the bone. Whilst this is true for the majority of taxa it does not hold for all;
Hypophthalmus is one exception (see above). In the clariids, the posterior infraorbitals
(includes 'supraorbital' aucct.} are greatly expanded.

The recognition of reduced infraorbitals and the absence of a supraorbital bone as
synapomorphies linking siluroids with gymnotoids (Fink & Fink, 1981) is dubious. Such
reduction and loss also occurs in cyprinoids (Cobitidae). An even more pertinent point is the
differing arrangement of the infraorbitals in gymnotoids and siluroids — clearly shown by
Fink & Fink (1981, fig. 7D-E). The anterior infraorbital bones of siluroids are always
elongate and the 3rd extends someway past the orbit so that the posterior series curve dorso-
anteriorly. In gymnotoids the 3rd infraorbital is a short bone and the posterior elements are
directed diagonally in a dorso-posterior direction.

Cranial musculature (Figs 5-6)
Jaw and suspensorial muscles

When the eye, together with its muscles and the optic tract have been removed, the entire
face of the fish is seen to be covered by an almost undifferentiated adductor muscle (Fig. 5).
The orbital socket is indicated on the face of the muscle by a circular tendinous area, the
optic tract by a vertical groove (otf, Fig. 5). Dorso-posteriorly, lateral fibres of the levator
arcus palatini muscle (lap) adhere to the adductor's surface, and a more medial section of the
levator is indicated by a longitudinal division of the adductor.

The posterior origin of the adductor mandibulae is from the lateral and antero-medial
faces of the hyomandibula. Anteriorly, at the articulation of the lower jaw, there is a
myocomma and the adductor can be differentiated into a number of segments. The upper,
outer portion curves inward, its anterior border being tightly attached to the thick
mandibular nerve tract (nvmd) which follows the muscles contour as it passes medially to

lap otf

etb

etc

eta

Fig. 5 Hvpophthalmm edentatm\ superficial head musculature. Drawn from specimen BMNH

1976.6.18: 123-9. Scale = 5 mm.

10

G. J. HOWES

aap etb nvmd

atm pal

amm

bmx

A2+A,

Fig. 6 Hypophthalmus edentatus; lower jaw musculature. The palatine and extensor muscle have
been raised. In relaxed position the tendon (atm) passes between the rami of the maxillaris nerve
(nvmx). Drawn from specimen BMNH 1976.6.18: 123. Scale = 3 mm.

attach to the inner face of the lower jaw. From its insertion this part of the muscle is regarded
as section A2 (Fig. 6). There is no recognisable Aw, A2 inserting via a broad tendon stretching
across the medial face of the adductor mandibulae mass.

The antero-ventral part of the muscle lying lateral to the mandibularis nerve trunk is
divisible into an upper bundle of fibres inserting into the lip tissue covering the coronoid
process, and a ventral section inserting partially into labial tissue and partially on the lateral
face of the dentary (A,, Fig. 5). When the superficial layer of the dorsal part of A, is removed,
a medial layer is found to be continuous with A2 (amm, Fig. 6); this division appears to be
homologous with the retractor tentaculi (see below). The posterior border of the ventral part
of A, is marked by a myocomma. A short, pinnate bundle of fibres stems from the antero-
medial face of the adductor complex and inserts into a long tendon (atm, Fig. 6). The tendon
passes between the maxillary and mandibular branches of the trigeminal nerve to insert on
the maxilla at the base of the barbel.

There is no discrete medial division of the main body of the adductor mandibulae, only
posteriorly is there a separation by a medial sheet of the levator arcus palatini muscle (see
above), but the fibres of both muscles intermesh.

The central portion of the adductor arcus palatini extends between the parasphenoid and
the dorsal border of the pterygoid bones (aap, Fig. 6). A separate, posterior portion runs from
the parasphenoid to the medial face of the hyomandibula, and is identified as the adductor
hyomandibulae. The anterior part of the adductor arcus palatini is divisible into three
sections which together comprise the extensor tentaculi. The antero-dorsal part of this
muscle covers the lateral ethmoid and attaches to the posterior half of the palatine (eta, Fig.
5). The postero-medial part originates as two separate sections from the central region of the
adductor arcus palatini. The longer, postero-medial section runs forward at an oblique angle
to insert on the postero-medial portion of the palatine (etb, Fig. 5). The shorter, medial
portion runs antero-dorsally to insert on the antero-medial part of the palatine (etc, Fig. 5).

Innervation

Two thick nerve tracts stem from the trigeminofacial ganglion and exit from a single prootic
foramen. The supraorbital trunk diverges immediately. The infraorbital trunk is of great

CATFISH ANATOMY & PHYLOGENY

nvmd ,rs aap

ct

11

Imb

am

Fig. 7 Nematogenys inermis; dorso-lateral view of cheek musculature. Drawn from specimen
BMNH 1883.li.27: 45-48. Scale=10mm. co = coronoid process; ct = connective tissue;
1mb = ligament connecting maxillary barbel with mandible.

thickness, and above the jaw articulation it divides into the buccal and maxillo-mandibular
rami which pass, respectively into the root of the maxillary barbel and the lower jaw. (Fig. 6).
The buccal ramas gives off a branch to the adductor arcus palatini before bifurcating into an
outer branch serving the area of the extensor tentaculi muscle around the palatine, and an
inner branch which enters the base of the maxillary barbel. The mandibular ramus (nvmd)
curves round the anterior border of the adductor muscle complex, sending off short branches
into the musculature (Fig. 6). The trunk passes across the inner face of the dentary to run
inside the length of the bone before terminating in a bifurcation, each branch of which enters
its respective barbel (Figs 7 & 1 4).

Jaw and suspensorial muscles in other siluroids

It is generally accepted (see Winterbottom, 1974) that in teleosts the section of the adductor
mandibulae termed A, is that which forms the outer, or dorsal muscle segment inserting on
the maxilla. Section A2 is that which lies medially and, or, ventrally to A,, and inserts on the
lower jaw; the part of it which covers the medial face of the dentary is termed Aw. Further
divisions (A2a, A3 etc) are those successively medial to A, and A2 and which insert onto the
inner face of the lower jaw.

In siluroids, this seemingly straightforward nomenclature has become somewhat confused
through the presence of a muscle, serving the base of the maxillary barbel, known as the
retractor tentaculi, often referred to as the adductor tentaculi or protractor maxillae; see
Winterbottom, 1974 for synonymy. The fact that this muscle inserts on the maxilla but stems
from the medial aspect of the adductor mandibulae has led to its being homologised with
various sections of the adductor complex.

Takahasi (1925) thought that because of its maxillary insertion it was synonymous with
division A,, a hypothesis followed by Edgeworth (1935). McMurrich (18846) and Lubosch
(1938) considered the retractor tentaculi to be an inner division of the adductor. Eaton (1948)
agreed, in referring to Ictalurus thus . . .'From the fact that a stout ligament runs from the
base of the adductor tendon back and downward to join the ligamentous sheath of the

12

G. J. HOWES

Imx

Fig. 8 Callophysus macropterus; lateral view of muscular and ligamentous connections to the
upper and lower jaws. Drawn from specimen BMNH 1913.7.30: 20-22. Scale = 5 mm.
Imx = maxillary-mandibular 'ligament'; pet = palato-pterygoid connective tissue.

quadrate-articular joint, I would infer that the muscle has simply taken hold, as it were, of
the primitive ligament which runs, in such fishes as Amia calva, from the articular to the
maxillary. Thus, the ligament has now become in large part a muscle tendon. . .'

Alexander (1965) concurs with Eaton, believing that the retractor tentaculi (the adductor
tentaculi of Alexander) is derived from '. . .A3 rather than A,'.

From the arrangement in siluroid taxa I have investigated, it appears that these latter
authors are probably correct in assuming that the retractor tentaculi is part of the inner
adductor series and that its attachment with the upper jaw is a de novo development. The
possible evolutionary development of the retractor tentaculi as suggested by Eaton (1948)
and Alexander (1 965) can be amplified by the following observations.

The presumed plesiomorph condition of the adductor mandibulae muscle in siluroids is
where divisions of the muscle are not well-differentiated and insertion is entirely, or almost
entirely, on the lower jaw. Diplomystes and Nematogenys (Fig. 7) illustrate this condition
where the lower, outer part inserts musculously on the lower jaw (anguloarticular). An
upper, inner division can be distinguished by its attachment, anteriorly, to the lower part by
a broad vertical tendon which covers the postero-dorsal rim of the lower jaw. Stemming from
the antero-medial surface of this muscle, and extending across the dorsal surface of the
mandible is a thick sheet of connective tissue (ct). This sheet bifurcates, the upper strand
attaching to the posterior face of the maxilla, the lower to the distal portion of the maxilla
where it forms a sheath around the maxillary barbel.

In Callophysus, Pimelodidae, a continuous sheet of tissue extends medially from the
dentary to the anterior pterygoid process (pet, Fig. 8). That area of tissue covering the
coronoid process is thickened and, anteriorly, is separated from the main sheet so forming a
distinct 'ligament' running to the base of the maxillary barbel (Imx, Fig. 8).

A similar arrangement of the 'ligament' is present in Tandanus and Plotosus. Alexander
(1965), following Takahasi (1925), states that the deepest part of the adductor mandibulae
inserts on the articular-maxillary ligament. This is, however, incorrect. A strong ligament
does indeed stretch from the posterior aspect of the maxilla to the coronoid process of the
dentary, and appears to be homologous with the partially differentiated 'ligament' described
above in Callophysus, but no muscle inserts on it (Imx, Fig. 9). The outer portion of the
adductor mandibulae in Tandanus and Plotosus inserts on the lower jaw medial to the
ligament. The inner sections of the adductor, be they interpreted as A2 or A3, insert on the
medial face of the anguloarticular (Fig. 9).

CATFISH ANATOMY & PHYLOGENY 13

et

mx

Imx

bmx

Fig. 9 Tandanus tandanus; dorso-lateral view of jaw muscle attachments. Drawn from specimen

BMNH 1871.2.10: 4-6. Scale= 10 mm.

Kesteven (1943) also erroneously identified a retractor tentaculi, his protractor maxillae,
in Tandanus. He states that the muscle '. . .arises from the side of the skull above the anterior
attachment of the palatal arch. . .' to insert on the maxilla '. . .enswathing the posterior end
of the bone.' I find no such muscle in Tandanus tandanus (Fig. 9) nor in Plotosus plotosus
and must presume Kesteven misidentified part of the extensor tenatculi or else did not
entirely dissect the connective tissue enswathing the maxilla.

In the pimelodids Pimelodus and Iheringichthys and the bagrid Clarotes a distinct
ligament connects the base of the maxillary barbel with the coronoid process of the dentary.
However, the ligament then courses posteriorly from the coronoid process to the fascia of the
medial section of the adductor mandibulae. A further modification of this condition occurs
in one group of pimelodids, Parapimelodus and Pimelodus maculatus, whereby there are
two distinct ligaments extending from the base of the barbel, the outer attaching to the
coronoid process and the medial one to the pterygoid process. In none of these taxa does the
musculature attach to the ligaments.

In Megalonema, Pimelodidae, the maxillo-articular ligament bifrucates posteriorly, the
inner branch becoming continuous with the tendon of the medial segment of the adductor
muscle complex. The outer ligament is, however, still connected with the lower jaw. In
Sorubimichthys, Pimelodidae, a distinct ligament connects the maxilla with the coronoid
process of the dentary. However, the ligament is tightly attached along the length of the
upper jaw (premaxilla) by connective tissue. A branch of the mandibularis nerve runs into
the tissue joining the upper and lower jaws (Fig. 10).

These examples serve to illustrate the probable pattern involved in the evolution of the
retractor tentaculi. The hypothesised stages of this process are; (1) the palato-mandibular
connective tissue becomes differentiated to form a stout, free ligament connecting the
maxilla with the posterior part of the lower jaw, (2) medial fibres of the adductor mandibulae
become associated with the maxillo-palatine part of the palato-mandibular tissues, (3) the
connective tissue becomes further differentiated and tendinously links a discrete medial
section of the adductor mandibulae with the root of the maxillary barbel.

The basic conditions for the development of the retractor tentaculi are present in several
members of the Pimelodidae, Bagridae and Plotosidae and it seems very likely that the
muscle has been derived independently in several lineages. Various apomorphic conditions
of the muscle can be recognised as for example in some pimelodids such as Rhamdia where
the anterior segment is attached to the orbitosphenoid via a bifurcated tendon.

14

do

am

nvmd

pmx

Imb

Fig. 10 Sorubimichthys planiceps; dorso-lateral view of superficial jaw musculature. Drawn from
specimen BMNH 1977.5.24: 12. Scale = 5 mm. em = eye musculature, 1mb = maxillary barbel-
mandibular ligament.

If the adductor mandibulae muscles of siluroids are compared with those of other
otophysans, their arrangement is seen to correspond most closely to that of characoids,
where section A, also inserts on the dentary. Vari (1979: 317) considered that a lower jaw
insertion for the outer part of the adductor represented the plesiomorph condition. In
characoids a strong, well-differentiated maxillary-mandibular ligament is present and often
the dorso-lateral fibres of A, attach to this ligament. Vari (1979) notes that incorporation of
the ligament into the tendon of A, is a derived condition in distichodontid characoids.

In cyprinoids the section A, never inserts on the lower jaw and there is no maxillary-
mandibular ligament.

In gymnotids there is a maxillary-mandibular ligament and in Gymnotus a few fibres of the
outer part of the adductor muscle insert on this ligament; the remainder of the muscle inserts
onto the outer face of the lower jaw, as in siluroids. However, this is not the case in
Sternopygus, Eigenmannia and Rhamphichthys the only other gymnotid taxa examined,
where the outer segment of the adductor attaches to the 1st infraorbital in Sternopygus and
Eigenmannia, and to both the 1st infraorbital and the maxilla in Rhamphichthys.

Two hypotheses are available to account for the arrangement of the outer adductor
muscles in siluroids. 1. That the insertion on the lower jaw is plesiomorphic and the
ligamentous strand connecting the maxilla with the dentary is homologous with that similar
ligament in characoids. 2. That insertion on the lower jaw is a derived feature brought about
by regression of the maxilla and the loss of its intimate connection with the lower jaw. In
effect, a regression of the cyprinoid condition.

Fink & Fink (1981) favour the second hypothesis stating . . .'The conditions in some
characiformes and in siluriformes are hypothesised to be secondary reductions from a
primitive attachment to the maxilla.' Although Fink & Fink refer to some Characiformes,
the outer element of the adductor mandibulae attaches to the lower jaw in the majority of
characiform taxa. In those siluroids where the adductor mandibulae inserts on the upper jaw,
Loricariidae, Astroblepidae, it is to the premaxilla and the muscles involved are derivatives
of the medial and not the outer parts of the adductor complex (Howes, in press). Only in the
Callichthyidae is there an insertion of A, on to the maxilla, and this appears to be a derived
condition associated with the maxillary-ethmoid joint (Howes, in press). In view of these

CATFISH ANATOMY & PHYLOGENY 15

observations I would regard a lower jaw insertion of the adductor madibulae A, in siluroids
as the plesiomorph condition. The use of this character by Fink & Fink (1981) uniting the
siluroids and the gymnotoids cannot be upheld, as in some (?the majority) of gymnotoids, the
outer part of the adductor muscle inserts on the 1st infraorbital and not the lower jaw (see
above).

The opercular muscles (Figs 3 & 5)

The dilatator operculi (do, Figs 3 & 5) is an exceptionally long muscle having its area of
origin anterior to the orbit, the fibres adhering to, and covered by, the ventral margin of the
frontal. Posteriorly, above the optic tract, the muscle is exposed before becoming tendinous
and passing between the adductor mandibulae and the medial section of the levator arcus
palatini. The long insertion tendon attaches to the antero-dorsal process of the operculum.

The levator operculi (lo Fig. 3) comprises two thin segments, an upper one arising from the
postero-dorsal face of the hyomandibula and inserting on the postero-dorsal margin of the
operculum, and a medial one originating from the pterotic and inserting on the antero-
medial face of the operculum.

The adductor operculi (ao Fig. 3) originates from a fossa indenting the pterotic,
exoccipital and 'posttemporaF. It inserts on the dorso-medial face of the operculum.

The hyoid muscles (Figs 1 1 & 12)

The ventral head muscles have a complex arrangement with much integration of the various
segments.

The ventral layer exhibits two groups of muscles; those in which the fibre direction is
toward the midline and those in which it runs antero-laterally.

The largest element (hma, Fig. 11 A), belonging to the latter group, occupies half the
lateral aspect of the lower jaw. Its fibres run at an oblique angle from the anterohyal to the
dentary; anteriorly they change direction to run almost longitudinally. The outermost fibres
insert onto the forepart of the dentary whilst the remainder turn inward as a separate bundle
and are incorporated into the medial section (hmb, Fig. 1 1 A). This medial part of the muscle
also originates from the anterior part of the anterohyal. The muscle is formed from two
sheets. The ventral, medial portion (hmbl, Fig. 1 1 A) curves antero-medially, its outer fibres
inserting on the cartilaginous plate of the inner mandibular barbel, its inner fibres meeting
those of its antimere at a midline raphe. The lateral portion (hmb2, Fig. 11 A) runs
anteriorly to insert on a tendinous pad covering the cartilaginous base of the outer
mandibular barbel (see below). A separate bundle of fibres insert into the base of the barbel.
The medial fibres ofhmb2 overlie the lateral aspect of hmb 1 , but at the midline both muscle-
layers meet at a raphe.

Covering the branchiostegal rays and extending forward as a thin, longitudinally fibred
segment is muscle hmc (Fig. 11 A). Anteriorly it joins an aponeurosis into which hmbl
inserts; medially it joins its partner at a midline raphe. Extending forward from the
aponeurosis is a discrete bundle of fibres that attaches to the base of the inner mandibular
barbel (hmd, Fig. 11 A).

The large muscle (hma) is considered, from its disposition and innervation by the ramus
hyoideus VII to be the posterior protractor hyoidei. The inner divisions (hmbl and hmb2 are
also innervated by branches of the r. hyoideus VII and appear to be anterior parts of the
protractor hyoidei. The small muscle slip attaching to the outer mandibular barbel is merely
a continuation of the dorsal portion of this muscle and can be identified with that element
named by Singh (1967) as a retractor tentaculi, not to be confused with his use of this name
also for the facial muscle. The longitudinal muscle (hmc) is the hyohyoidei adductores. It is
unclear, however, if the small bundle of fibres (hmd) running from its anterior aponeurosis to
the inner mandibular barbel is indeed part of this muscle or whether it is derived from the
protractor hyoidei. This small barbel muscle should also be identified with Singh's (1967)
retractor tentaculi, which Winterbottom (1974) considered to be homologous with the

16

G. J. HOWES

im

Fig. 11 Hypophthalmus edentatus; A, ventral view of hyoid musculature (right side); B, dorso-
medial view of mandibular barbel supporting elements (left side). Thick dashed lines = nerve
path, thin-dashes and dots = outline of barbel root; C, lateral view of mouth, upper lip cut
sagittally. Drawn from specimens BMNH 1972.7.27: 675-678 & 1976.6.18: 123. Scales = 5 mm
ul = upper lip.

retractor hyoidei. Ghiot (1978) illustrated in Pimelodus five separate muscles controlling the
mandibular barbels of which he ascribed origin to 'differentiation of the hyoid protractor'.

The intermandibularis (im, Figs 1 1 A & C) forms the anterior transverse muscular floor of
the mouth. It is divided medially and each segment is attached to its respective cartilaginous
plate which serves to support the barbel (see below). In the midline, each half of the muscle
joins in a raphe which, in turn, is attached by strong connective tissue to an overlying collar
of fatty tissue (ft, Fig. 11C). This collar encircles the leading edge of the antero- and
dorsohyals and covers the basihyal.

Only in some pimelodid taxa is there such a complex arrangement of the hyoid
musculature as in Hypophthalmus. In Pimelodus, Sorubim and Pseudoplatystoma the
protractor hyoidei is complexly aponeurotic, and for its greater part is attached to the large

CATFISH ANATOMY & PHYLOGENY

17

ib

ha

Fig. 12 Sorubim lima; ventral view of hyoid musculature (dashed lines = outlines of barbel
supporting plates. Drawn from specimen BMNH 1969.7.15.26. Scale = 5 mm. ib, ob = inner and
outer mandibular barbels.

cartilaginous plates supporting the bases of the barbels (Fig. 12). Ghiot (1978) distinguished
in Pimelodus clarias five separate sections of what he termed the superior hyoidian
protractor muscle in addition to separate lateral and medial divisions. The complexity of the
hyoid musculature with separate mandibular barbel muscles attached to basal cartilages
appears to be derived for at least one group of pimelodids and the Hypophthalmidae. The
usual condition in siluroids is for the mandibular barbels to be 'embedded' within the
protractor hyoidei, with bundles of fibres inserting on the anterior and posterior bases of the
barbels and performing the function of retractors and protractors. This arrangement of hyoid
musculature is illustrated by Singh (1967) in Bagridae and Winterbottom (1974, fig. 12) in
Ictaluridae.

Unlike other otophysans, the siluroids exhibit considerable variability in the morphology
of the hyoid musculature. A comparative study of these various arrangements appears to
offer a fruitful field of potential synapomorphies.

The snout and jaws (figs 11-15)

Ramsay Wright's (1885) description of the snout region in Hypophthalmus can hardly be
bettered and the following account is merely a supplement.

The entire ethmoid region is of papyraceous bone interspersed with cartilaginous strips
(ec). Such cartilaginous areas separate the ethmoid (eth) from the lateral ethmoids and are
bridged by the spine-like nasal bones (na, Fig. 13 A). It is not possible to determine whether
the ethmoid bone is a supra- or a mesethmoid. That it is so thin, and its lateral and posterior

18

pmx

le

mx

Fig. 13 Hypophthalmus edentatus; (upper) dorsal view of ethmoid region, (lower) palatine-
maxillary articulation. Scales = 5 mm. Drawn from specimen BMNH 1972.7.27: 675-678.

margins grade into cartilage suggest that it is a single element and is entirely mesethmoidal in
origin.

The lateral ethmoid (le, Fig. 13) has a wide anterior surface with a convex antero-lateral
margin; postero-laterally its border is produced into a long, ventrally directed spine which
almost meets the third infraorbital. The medial border of each lateral ethmoid abuts the
midline mesethmoidal cartilage. The vomer is needle-like, its head slightly broadened and
resting below the ethmoid (v, Fig. 1 3).

The nostril is bordered medially and laterally by the anterior arm of the bifurcated 1st
infraorbital.

The edentulous premaxillaries (pmx, Fig. 13) are barely identifiable as separate elements,
being fused with the ethmoid and distinguishable only by their denser ossification. They are
so firmly united at the symphysis that even under a magnification of 50 x a suture is not
visible in alizarin preparations. The maxilla is a small, well-ossified, flattened and hook-
shaped bone (mx, Fig. 13). The blade of the hook supports the barbel (bmx), its base is
rounded and articulates withe the cartilaginous tip of the palatine.

The rod-like palatine (pal) is ossified, apart from its anterior and posterior tips and a small
medial disc at about its midpoint (Fig. 1 3). This disc articulates with the lateral border of the
lateral ethmoid.

Like the upper jaw, the lower jaw is also edentulous. The dentary (d) contributes to half
the length of the mandible and is strongly sutured to the anguloarticular (aa).

The coronoid process is formed largely by a dorsal extension of the Meckelian cartilage
(com, Fig. 14). The horizontal segment of this cartilage extends anteriorly along the medial
face of the dentary and also has a slight posterior extension. The mandibularis nerve passes
across the medial face of the upright part of the cartilage and then between the horizontal
segment and the dentary wall. The nerve trunk continues along the ventral rim of the dentary
and then bifurcates, each ramus inserting into the base of a corresponding mandibular
barbel. A separate rod-like cartilaginous element lies antero-dorsally to the coronoid
cartilage (cr, Fig. 14). Its distal tip is forked, and attaches to the lateral fold of thick labial
tissue. The cartilage is mobile. When the jaws are abducted, and the labial tissue made
taught, the cartilaginous rod rises to an acute angle.

CATFISH ANATOMY & PHYLOGENY 19

nvmd

.com

Fig. 14 Hypophthalmus edentatus, lower jaw (right) in medial view. Drawn from specimen BMNH
1972.7.27: 675-678. Scale = 5cm. com = coronoid process of Meckelian cartilage; lt = labial
tissue.

The barbels are supported on three cartilaginous plates (cp, Figs 1 IB & 14). The first, i.e.
the element closest to the jaw symphysis, is the most complex. Its flat proximal tip attaches
to the dentary rim and is curved where the mandibularis branch passes between it and the
dentary. The cartilage extends dorso-posteriorly as an open cylinder, its distal tip providing
the insertion site for the inter mandibularis muscle. The root of the inner barbel (ib)
bifurcates, one part ending as a short hook on the outer part of the cartilage, the other
extending along the outer side of the cylindrical section. The two inner plates are irregularly
shaped. The 2nd, middle, plate contacts the symphysial plate dorsally; the 3rd, outer, is
notched at its articulation with the dentary thus allowing transmission of the mandibular
nerve branch to the outer barbel (ob). As with the inner barbel the outer one is also
bifurcated, its thicker leg attaching to the 3rd plate, its inner, thinner leg becoming
'cartilaginous' and extending posteriorly to join the centre of the 2nd cartilaginous plate.

Uniquely amongst siluroids, the anterior portions of the ceratobranchials enter the floor of
the mouth and are curved upward, with the result that the basihyal is raised to the level of the
roof of the mouth and the dorsohyals touch the underside of the ethmoid. The elevation of
the floor of the mouth appears to be accomplished by the upward rotation of the medial
cartilaginous plates supporting the intermandibularis muscle (see p. 16). The inter-
mandibularis thus forms a hillock against the underside of the ethmoid and the thin upper
lip tightly embraces the lower jaw when the mouth closes. The pressure created by the floor
rising against the roof of the mouth serves to force water through the mesh-like gill-rakers
which filter out particulate material. The length of the gill openings presumably allows a
high volume of water to pass rapidly over the gill-arches.

The skin covering the dentary contains 7-8 transverse tubes (mst, Fig. 14) filled with a
jelly-like substance staining intensely with the mucus-specific stain alcian blue. Of the taxa
examined Iheringichthys has a similar jelly-like material in the lower lip, but it is
concentrated in ampullae in the tissue close to the jaw symphysis and not arranged in widely
spaced rows. Gelinek (1978) has described ampullary organs in the epidermis of Sorubim.
He found them to be densely grouped in the labial tissue of the upper jaw and noted their
structural similarity to those organs in Plotosus. Gelinek observed the similarities between
these organs to those of gymnotids and mormyrids, thus hypothesising a similar function,
namely electroreception.

The snout and jaws in other siluroids

The presumed plesiomorph condition of the ethmoid in siluroids is that which occurs in

20

G. J. HOWES

mx

ethc

Fig. 15 Upper, Iheringichthys labrosus, ventral view of mandibular barbel supporting elements.
Specimen BMNH 1934.8.20: 70-72. Scale = 2 mm. Lower, Luciopimelodus pati, dorsal view of
ethmoid region. Specimen BMNH 1 878.5. 1 6: 30 (dry skeleton). Scale = 3 mm. d = dentary.

many taxa belonging to the Diplomystidae, Trichomycteridae, Bagridae, Ictaluridae,
Pimelodidae, Mochokidae, Plotosidae and Loricariidae, here the ethmoid is elongate and T-
shaped and often with a median anterior notch (see Fink & Fink, 1981). Howes (1980)
mistakenly considered this type of ethmoid in Mochokidae as apomorphic.

In its presumed plesiomorph condition, the siluroid lateral ethmoid is triangular, closely
united with the parasphenoid ventromedially and with the orbitosphenoid posteriorly.

The expanded and poorly ossified ethmoid region of Hypophthalmus is considered to be a
derived condition. Other derived states of the ethmoid are those where the bone is flattened
with divergent anterior forks (Ageniosidae) is curved ventrally (some Ariidae) or is cavitous
with a medially grooved surface (some Auchenipteridae). Another derived form occurs in
one group of Pime\oididae,Piramutana, Hemisorubim, Luciopimelodus, Brachyplatystoma,
Pseudoplatystoma and Sorubimichthys. Here the mesethmoid is greatly depressed and

CATFISH ANATOMY & PHYLOGENY

na

le

21

sbc

Fig. 16 Hypophthalmus edentatus, dorsal view of cranium. Specimen BMNH 1972.7.27: 675-678

(alizarin). Scale = 10 mm.

expanded laterally (Fig. 1 5). As in Hypophthalmus the lateral portion of the ethmoid forms a
large cavity wherein lies the olfactory organ (etch, Fig. 1 5).

Derived conditions of the lateral ethmoid are those where the bone has a posterior process
contacting an anterior process of the frontal (some Ariidae and Pangasiidae) and where the
bone is extensive, its antero-lateral portion lamellate, extensively cancellous and vacuolate.
This latter condition occurs in Pangasiidae, Amphiliidae, Auchenipteridae and
Pimelodidae, but among the latter family only one taxon, Parapimelodus, approaches
Hypophthalmus in the width and shape of its mouth.

The mandibular barbels of pimelodids are particularly well-developed and in some taxa,
as in Hypophthalmus, they articulate with large cartilaginous plates, e.g. Sorubim,
Pseudoplatystoma, Iheringichthys, Pinirampus, Pimelodus (part), Luciopimelodus. The two
former genera differ from Hypophthalmus, however, in having the barbel origin remote from
the jaw symphysis. Ghiot (1978) described the cartilaginous plates in Pimelodus 'clarias' and
commented on the elasticity of the plates and their function of supporting and moving the
barbels. In Pimelodus blochii, Iheringichthys, Pinirampus and Luciopimelodus the barbels
lie close to the symphysis, the presumed plesiomorph condition, but the cartilaginous
supporting plates have a complex form resembling that in Hypophthalmus (Fig. 15). These
taxa also share with Hypophthalmus the peculiar mandibular sesamoid cartilage which
supports the lip fold, and the coronoid extension of the Meckelian cartilage (see p. 1 8
above).

The cranium (Figs 16-18)

There is little purpose in presenting a detailed description of the cranium, already described

22 G. J. HOWES

by Wright (1885). It is necessary only to comment on those features pertinent to the present
discussion.

The neurocranium is moderately broad, transversly convex, becoming high-vaulted in the
occipital region, and has a deep supraoccipital crest (Fig. 16). All the cranial bones are of a
honeycomb texture with the frontal papyraceous. Nearly all sutures are synchondral, and
fatty tissue pervades bone-muscle interspaces.

The otic region

The prootic is pierced by two foramina (pro, Fig. 17). The hyomandibular and palatine
branches of the VII cranial nerve emerge through the posterior one (fhvii) whilst the oculo-
motor, trigeminal, abducens and facial nerves emerge through the anterior foramen in the
border of the bone (ftf).

In almost all siluroids investigated there is but a single foramen piercing or indenting the
prootic. In some ariid taxa the prootic is imperforate and the foramen for the cranial nerves
is contained within the pterosphenoid. The absence of a second, posterior, opening in the
prootic of most siluroids is due to the peculiarity of the cranial blood vascular system in these
fishes. In the majority of teleosts the posterior prootic foramen serves for the passage of the
jugular vein (lateral head vein) as well as the hyomandibularis nerve trunk. In siluroids, the
jugular lies lateral to the cranial wall and is joined by its various branches in the same
aperture as that through which the trigeminal nerve complex passes.

The absence of a complete trigemino-facialis chamber in siluroids has been commented
upon by several authors as the following quotations indicate:

Allis (1908:259) referring to Ictalurus ( = Amiurus) notes that, 'The artery (external
carotid) does not apparently traverse a trigemino-facialis chamber, for although it would
seem as if this chamber must be present in some form, there is no proper indication of but
one cranial wall in this region, and that one wall would seem to be the inner wall of the
chamber; for both the external carotid and the jugular vein lie external to it.'

De Beer (1937:137) again in referring to Ictalurus comments, 'There is no lateral
commissure or pila antotica. The side wall of the skull in the orbital region therefore presents
a continuous fontanelle — the sphenoid fontanelle — between the hind edge of the preoptic
root of the orbital cartilage and the anterior basicapsular commissure. . .'

Bamford (1948:365, 367) referring to Galeichthys, 'The normal foramina for the exit of the
various parts of V and VII are absent, and the exit of these is limited to the sphenoid fissures.'
. . .'There is no trace of a pila lateralis or any similar structure.'

Alexander (1965:95) finds that in siluroids, 'The chondrocranium lacks the lateral
commissure. The trigemino-facialis chamber of the adult skull has consequently lost its
walls.'

All these authors are agreed that the siluroid condition is one where the trigemino-facialis
chamber has been modified and the prefacial commissure suppressed. Bamford (1948)
suggested that cartilaginous growth from the anterior edge of the otic capsule had crowded
the nerves and vessels into the same foramen as the optic nerve (see below concerning the
passage of the optic nerve).

The absence or reduction of an internal carotid artery in certain siluroids has also been the
subject of comment by Allis (1908), Goodrich (1930), De Beer (1937) and Bamford (1948).
According to Bamford (1948) the internal carotid has all but disappeared, with a consequent
modification of the arterial circuitry. As yet, too few siluroids have been examined to
ascertain the extent of this specialization.

The optic foramen and the suprasphenoid

In Hypophthalmus passage of the optic nerve (II) is through a separate foramen between the
pterosphenoid (pts) and the 'suprasphenoid' (see below). In many siluroids there is no
separate optic foramen, the optic nerve sharing the same opening as the trigemino-facialis

CATFISH ANATOMY & PHYLOGENY

23

obs

fhvii

pro

ss

pts

fii

Fig. 17 Neurocranium, otic region of: (upper) Hypophthalmus edentatus ventro-lateral view.
Specimen BMNH 1972.7.27: 675 (alizarin). Scale = 3mm. (centre) Auchenipterus nigripinnis
lateral view. Specimen BMNH 1910.5.26: 12 (skeleton). Scale = 3mm. (bottom) Pterodoras
granulosus lateral view. Uncatalogued skeleton. Scale = 10 mm. ss = suprasphenoid.

complex, e.g. Diplomystidae, Callichthyidae, Siluridae, Bagridae (part), Clariidae. It is more
usual in siluroids for there to be a separate foramen for the optic nerve between the
pterosphenoid and parasphenoid or the suprasphenoid.

Wright (1885) referred to a paired element in Hypophthalmus as a basisphenoid. The bone
in question is long and shallow and contacts the pterosphenoid, orbitosphenoid and prootic
(ss, Fig. 17). Between it and the pterosphenoid is a slit-like foramen through which passes the
optic nerve (fii, Fig. 17). In many taxa I have examined a similar element is present, and in
the Auchenipteridae and Doradidae it is pierced by the optic foramen (Fig. 17), a feature
recognised as synapomorphic.

24

v6

sbc

v5 v4

Fig. 18 Hypophthalmus edentatus neurocranium, occipital region (upper) in lateral, and (lower) in
ventral views. Composite drawing from alizarin and dry preparations, tsc = 'transcapular'.

McMurrich (1884#) and Kindred (1919) have described the so-called basisphenoid in
Ictalurus ( = Amiurus). Kindred (1919: 39-40) discussed the homology of this element and
decided that the term 'suprasphenoid' rather than basisphenoid would be better applied to it.
He defined the suprasphenoid as a '. . .connective tissue ossification above the parasphenoid
and between the ventral ends of the alisphenoids ( = pterosphenoids).' Daget (1964) also
advocates caution in applying the term 'basisphenoid' to this element in siluroids. Thus,
Kindred's term, suprasphenoid is adopted here as one being free of any implied homology.

The homology of the siluroid parasphenoid itself is in doubt. Allis (1919) considered the
problem in Ictalurus, (=Amiurus) and argued that the siluroid parasphenoid was an
ossification of the ventral myodome roof, the posterior myodome being absent in siluroids.

Bamford (1948:375) remarks that in Ictalurus, '. . .there are so many exceptional features
in this region of the skull . . . myodome basicranial relations, absence of trigemino-facialis
chamber, blood systems, pilae etc — that the matter cannot be regarded as settled.'

Until more is known about the occurrence of the suprasphenoid amongst siluroids, and
until more definite information on its ontogeny and relationships with other elements is
available, the value of this bone as a taxonomic character cannot be determined.

The occipital region

There are several outstanding features in this region of the cranium in Hypophthalmus, not
least of which is the ankylosis of the skull and the vertebral column (Fig. 18).

CATFISH ANATOMY & PHYLOGENY 25

The supraoccipital (so) crest is high and posteriorly is firmly united with the succeeding
neural spine (4th). Anteriorly the supraoccipital meets the frontal (f), and laterally it is
bounded by the epioccipitals. Wright's (1885) description and figure give an incorrect shape
to the supraoccipital, and those elements bounding it are misidentified.

Posteriorly the epioccipital (epo) is separated from the parapophysis of the 4th vertebrae
by an elliptical foramen through which passes the ramus lateralis (Fig. 16). This foramen is
shown in Wright's (1885) figures (fe, figs 1 & 3), but in the wrong position, as lying between
the pterotic and posttemporal.

The 'posttemporaf is an extensive bone (ptt, Fig. 18), its anterior part meeting the pterotic
(pte) and exoccipital (exo) and its medial margin bordering the parapophyses of the 4th, 5th
and 6th vertebrae. In no other siluroid does the posttemporal extend so far posteriorly, nor
articulate with the 6th vertebra. The ventral limb of the 'posttemporal' is broad and thick,
and articulates with the basioccipital. A thick ligament runs between its upper medial border
and the 4th centrum (Ipt, Fig. 18).

There has been some considerable debate concerning the identity of the element
connecting the cleithrum with the cranium in siluroids. The most recent discussions are by
Lundberg (1975) and are reconsidered by Fink & Fink (1981). The latter conclude that the
siluroid element is a composite formed from the supracleithrum, possibly the posttemporal
and Baudelot's ligament. That part of the element contacting the basioccipital and identified
by Fink & Fink (1981) as the ossified Baudelot's ligament was referred to by Kindred (1919)
as the transcapular.

I have little to add to this discussion except to confirm Lundberg's (1975) observation that
the small plate-like bone (previously thought to be an extrascapular, but identified by
Lundberg as the posttemporal) does not occur in the Hypophthalmidae. However, such an
element is present in Luciopimelodus which I have labelled here as an extrascapular so as to
conform with Fink & Fink's nomenclature (es, Fig. 20). Notwithstanding Lundberg's
evidence (1975) from the sensory canal pattern that the posttemporal is lacking, it would
seem that the supracleithrum would be the element of the clavicular series most 'easily lost'
in phylogeny; see for example Rosen (1964) where the supracleithrum is lacking in certain
atheriniforms.

The complex vertebrae, swimbladder and posterior lateral line nerve

The complex vertebrae and swimbladder (Figs 1 9-2 1 )

Both Wright (1 885) and Chardon (1968) have described the peculiar nature of the Weberian
apparatus in Hypophthalmus. The most notable character is the complex encapsulation of
the swimbladder formed of the parapophyses of the 4th centrum. The parapophyses are
produced into dorsal and ventral laminae, the ventral lamina formed partly from superficial
ossification; see below p. 28. The cavity so formed provides a capsule housing the swim-
bladder (sbc, Fig. 18).

Internally the swimbladder (sb, Fig. 19) is divided by a longitudinal septum. Externally it
has a thick tunica externa but no tunica interna. The opening between the upper and lower
parts of the parapophysis together with the lateral extension of the 'posttemporal' form a
funnel-like aperture to the exposed outer wall of the swimbladder (Fig. 19). No transverse
duct connects the swimbladder chambers.

The tripus (tr) is connected to the tunica externa just anterior to the septum. A medial
extension of the tripus articulates with the complex vertebrae (3rd centrum). Anteriorly the
tripus contacts the scaphium (sc) via a minute intercalarium (ic). The claustrum is lacking.
The scaphia provide the walls of the cavum sinus imparis. According to Wright (1885: 1 14),
the roof of the cavum sinus imparis is an ossified plate of dura mater attached to the first
centrum, but in the preparations examined it seems as if the roof is just a continuation of the
exoccipital. The 1st centrum is so far forward that it lies between the exoccipitals and above
the basioccipital (vl +2, Fig. 19).

26

G. J. HOWES

Wright (1885) maintained that the Weberian apparatus in Hypophthalmus has been
'. . .pushed (in the process of its reduction) into the foramen magnum, instead of being
outside the skull . . .'. Although that author considered the Weberian apparatus and
swimbladder to be functionless (a theory endorsed by Bamford, 1 948) this does not appear to
be so. The swimbladder is constructed as in other siluroids and there is a mobile articulation
of the Weberian ossicles, all of which suggests that the system is functional (see below).

The complex vertebrae and swimbladder in other siluroids

In siluroids, as in other otophysans, the anterior four vertebrae are modified to form the
Weberian apparatus. However, unlike other otophysans, there is a greater degree of fusion
between the centra involved, and the united elements are usually referred to as the complex
vertebrae (Chardon, 1968). In addition to this fusion of the first four centra, sometimes the
5th, and more rarely the 6th, 7th and even 8th are also incorporated in this complex.

The lateral processes of the various centra incorporated in the complex vertebrae are
always well-developed, often forming sheets of bone which may contact one another to
provide a dorsal shield to the swimbladder. The anterior ramus of the 4th vertebral para-
pophysis (following Tavolga, 1962, referred to hereafter as the Miillerian ramus) may
contact the inferior limb of the 'posttemporal' and, in some taxa where the swimbladder is
encapsuled, it may form a substantial part of the capsule. Alternatively, the Miillerian ramus
may remain free from the 'posttemporal' and curve ventrally to contact the tunica externa of
the swimbladder.

In some families (Doradidae, Auchenipteridae, Mochokidae, Malapteruridae,
Pangasiidae, Ariidae) the Miillerian ramus is free from any attachment with the
'posttemporal', its distal tip is expanded into a plate which contacts the tunica externa of the
swimbladder. Protractor muscles run from the bony expansion to the cranium and neural
spine complex supporting the dorsal fin. Miiller (1842) named this mechanism the Spring-
federapparat, now usually referred to as the elastic spring apparatus, and below as the ESA.

The plesiomorph condition of the swimbladder in siluroids is assumed to be one in which

v5 v6

Fig. 19 Hypophthalmus edentatus swimbladder and Weberian apparatus. Dorso-lateral view of
transverse section through the anterior vertebrae and occipital region. Composite drawing from
alizarin and dry preparations. fs2 = foramen of 2nd spinal nerve; ssc = semicircular canal.

CATFISH ANATOMY & PHYLOGENY

27

V5

fx

v6

Fig. 20 Luciopimelodus pati complex vertebrae in (left) dorsal, (right) ventral, and (bottom) lateral
views. Specimen BMNH 1878.5.16: 30, dry skeleton. Scale = 5 mm.

the organ is large and free from any intimate contact with the anterior vertebral
parapophyses, the lumen partially divided by a T-shaped septum, and the pneumatic duct
entering the anterior transverse chamber. In taxa with this type of swimbladder, the anterior
vertebrae (i.e. those following the complex vertebrae) demonstrate a range of fusion patterns,
from being completely unfused, Diplomystidae, to the fusion of centra 4-6 in some
Pimelodidae. In those taxa with an ESA, the swimbladder is also large and not encapsuled,
but it is notable that in all ESA taxa studied the complex centra display the maximal
vertebral consolidation (4-6 + 4-7 in some ariids). In those taxa with encapsuled swim-
bladders, a series of progressive anterior vertebral fusions is apparent. However, only three
families have the maximal fusion of 6 centra, Loricariidae, Astroblepidae and Ageneiosidae.
If progressive fusions, namely the 4th centra with the 5th, then fusion with 6th and 7th
vertebrae, are taken as successively derived conditions the resulting cladogram indicates that
swimbladder encapsulation has occurred independently in at least two lineages (Fig. 22).
However, a more rigorous study of the individual taxa within those lineages shows incon-
gruencies in this hypothesised phylogeny. For example, the nature of swimbladder
encapsulation in the Callichthyidae, Astroblepidae and Loricariidae is virtually identical,
the 'posttemporal' contributing to the lateral wall of the capsule (Alexander, 1964; 1965;
Chardon, 1968). These taxa also share a derived rib structure and dermal ossification
patterns (see Alexander, 1965). On the basis of vertebral fusion patterns alone, however,
these families appear in different lineages. Likewise, the Ageneiosidae also appears with the
Loricariidae and Astroblepidae, but the encapsulation of the swimbladder is quite different
since it is derived from superficial ossification (see below).

28 G. J. HOWES

Further incongruencies arise in this phylogeny when the detailed structure of the ESA and
the associated swimbladder are taken into account. In the Doradidae and Auchenipteridae
the organisation of this system is more elaborate than that of other ESA taxa. In both families
the epioccipital has long posterior processes which contact the 5th and 6th parapophyses.
Such features are lacking in other ESA groups. From these observations it seems very likely
that the ESA has, as in the case of swimbladder encapsulation, been achieved independently
in several lineages.

Another incongruency arises if the superficial ventral ossification of the fused vertebral
centra is taken into account. Superficial ossifications were described by Bridge & Haddon
(1894) as a continuous sheet of bone uniting the paired parapophyses and enclosing the
cardinal veins. According to Tilak (1965: 170-1 71) the superficial ossification is derived from
the tunica externa of the swimbladder. There is much variability in development of
superficial ossification. In some taxa it forms a channel or a complete tunnel enclosing the
dorsal aorta. In others it contributes to the anterior part of the swimbladder capsule.

There appears to be a trend for the more derived siluroids to have reduced and encapsuled
swimbladders. The most extreme forms of encapsulation and swimbladder reduction are
usually encountered among those taxa with accessory respiratory devices, Clariidae,
Loricariidae and Callichthyidae. In this respect the Hypophthalmidae and Ageneiosidae are
exceptional.

Alexander (1964) makes the point that encapsulation of the swimbladder results '. . .when
the anterior parapophyses have not been correspondingly reduced'. Undoubtedly this is so,
the anterior (Miillerian) ramus of the 4th parapophysis progressively curving around the
swimbladder and meeting the lamina of the 5th appears to be an ontogenetically obvious
process of encapsulation. The swimbladder capsule may also incorporate part of the ventral
superficial ossification (see above), in Loricariidae and Callichthyidae the posterior cranial
bones (epioccipital and 'posttemporal') also contribute to its formation.

Alexander (1964) advanced the hypothesis that a reduced swimbladder volume could have
evolved to compensate for increased buoyancy resulting from gas held in the accessory
respiratory organ. Qasin & Hasan (1961) suggested that such gas could even be used in
buoyancy control. Gee (1976) does not agree, pointing out the unlikelihood of a
simultaneous reduction of swimbladder volume (SBV) and development of an accessory air
breathing organ (ARO) derived from some other part of the fish's anatomy. Gee also noted
that loricariids although possessing the smallest SBV lack an ARO. Gee's thesis is that the
overiding factor in swimbladder reduction is directed towards a demersal existence wherein a
greater variety of environments may be exploited, e.g. substrate burrowing by tricho-
mycterids; leaf litter and bank 'burrowing' by loricariids. Some of these environments,
however, tend to be hypoxic and would have demanded compensatory air breathing devices.

Reduction of SBV in Hypophthalmidae and Ageneiosidae is obviously due to different
factors than those obtaining in loricariids etc, and ones related to attaining neutral, or near
neutral buoyancy. Unlike Old World 'pelagic' siluroids, schilbeids, siluriids and pangasiids,
whose neutral buoyancy depends on a large swimbladder, the Hypophthalmidae, and
probably the Ageneiosidae, rely on a high body fat content (see p. 5). More positive asser-
tions on the functional utility of SBV reduction can only be made in the light of detailed
knowledge of swimbladder physics.

Alexander (1965) states there is no known fish in which the Weberian apparatus is a
functionless vestige. However, in loricariids, callichthyids and trichomycterids the ossicles
are reduced, the claustrum and intercalarium lacking (see Chardon, 1968). To what degree
the absence of these elements impairs or alters the function of the Weberian system is
unknown.

Bamford (1948) noted that little work had been done on the ontogeny of the siluroid
Weberian apparatus as compared with that in cyprinids. There has been no redress of this
situation. Bamford's implication was that the entire Weberian system could have been
derived independently in siluroids. This hypothesis has still to be tested.

CATFISH ANATOMY & PHYLOGENY

29

nviip

nviip

nviia

cw

Fig. 21 Pathway of vagus and posterior branch of the facial nerves in gymnotoids (upper), and
siluroids (lower). Dorsal view, semi-diagrammatic drawn from dissections of Eigenmannia
virescensand Pimelodus blochii. cw = cranial wall; op = operculum.

The posterior lateral line nerve

The anterior shift and gross re-shaping of the Miillerian ramus in Hypophthalmus has
necessitated a diversion in the route of the vagal ramus lateralis, posterior lateral line nerve.
As a result, after leaving the exoccipital (fX. Fig. 18) it takes an almost vertical course then
passes through the foramen between the Miillerian ramus and epioccipital (see above,
p. 25), finally, running posteriorly across the dorsal surface of the parapophysis.

Nelson (1960) described the course of the vagus lateralis in Malapterurus as leaving the
cranium and running laterally along the ossified Baudelot's ligament (inferior posttemporal
limb and transcapular of authors) thence passing caudally along the length of the body; the
ramus lateralis vagi of authors. Such is the usual course of the nerve in other siluroids
investigated. Nelson (1960) also describes a dorsal or recurrent branch of the VII cranial
nerve which, after leaving the cranium innervates the dorsal length of the body. This nerve
appears to correspond with that described by Mithel (1964) as the ramus lateralis accessorius
in Mystus and by Juge ( 1 899) as the 'nerf de Weber' in Silurus.

The more direct course of the vagus lateralis in Hypophthalmus also occurs in
Luciopimelodus (Fig. 20). Here too the posterior tip of the 'posttemporal' is elongated to
extend as far as the 5th parapophysis, and the Miillerian ramus is elevated and strongly
curved antero-ventrally to partially encapsulate the swimbladder. These features cause the
nerve to follow a vertical route from its exit in the exoccipital and, as in Hypophthalmus to
pass through a gap between the supraoccipital, epioccipital and Miillerian ramus (Fig. 20).

Nelson (1960) noted that in gymnotoids the '. . .nerve (vagus) emerges from the postero-
lateral aspects of the cranium associated with the opercular muscles and passes superficial to
the pectoral girdle.'; after which it penetrates the body wall musculature and follows the
horizontal myoseptum.

Szabo (1974) following Nelson (1960) considers the dorsal nerve in siluroids '. . .to be of
the type', presumably homologous, as the lateral nerve trunk in gymnotoids. Fink & Fink
(1981) accept such a homology as a character in uniting siluroids and gymnotoids as sister
groups. The very different pathways of the nerve trunk in these two groups suggests however
the possibility of independent derivation.

In siluroids the vagus lateralis passes laterally across the transcapular (tsc) (Baudelot's
ligament fide Fink & Fink), over the Miillerian ramus and then ventrally, medial to the
pectoral girdle. The posterior branch of the anterior lateral nerve (ramus recurvens nervi
facialis; ramus dorsalis or ramus lateralis accessorium auctf) emerges through the
supraoccipital to course along the dorsal aspect of the body (Fig. 2 1).

30

Diplomystidae

Bagridae
Chacidae
Schilbfeidae
Siluridae

Ictaluridae

Plotosidae

Pimelodont.(pt)
Cranoglanidae

G. J. HOWES
Clariidae
Amphiliidae
Sisoridae
Callichthyidae
Trichomycteridae
Cetopsidae
Aspredinidae
Pimelodont.(pt)

Hypophthalmidae
Helogeneidae

Ageniosidae

Astroblepidae

Loricariidae

Sb

En

Ariidae(4748)

Auchenipteridae

Doradidae

Mochokidae

Malapteruridae

Pangasiidae

Sb

.44546

ESA

U

Fig. 22 Cladogram of currently recognised siluroid families based on degree of fusion of the
anterior vertebrae and nature of the swimbladder. UFU = vertebrae unfused; FU = vertebrae
fused; 44-5+ = numbers of centra added to the complex vertebrae; SbF = swimbladder free; Sb
En = swimbladder encapsuled; ESA = elastic spring apparatus. For the purposes of the diagram
the Clariidae includes the Heteropneustidae. The Pimelodontidae is represented on two lineages,
those taxa with an encapsuled swimbladder correspond with group 1 as denned on p. 37. Ariid
taxa examined appear to have the 7th (? and 8th) centra added to the complex vertebrae. It is not
certain that all ariid taxa have this many consolidated centra. Data mostly from Chardon (1968)
and pers. obs.

In gymnotoids the vagus lateralis runs postero-ventrally into the epaxialis and does not
pass over the 4th parapophysis. The posterior branch of the anterior lateral nerve does not
pass dorsally but laterally, to pierce the hyomandibula, then to run medial to the pectoral
girdle, and into the body musculature where it joins the vagus lateralis (Fig. 21).

The suspensorium (Figs 23 & 24)

In Hypophthalmus a large pterygoid bone extends between the quadrate and hyomandibula
to form a bipartite attachment with the lateral ethmoid. This articulation takes the form of a
long triangular portion and an upright process (pty, Fig. 23). Between the two anterior forks a
ligament runs anteriorly to attach to a minute bony element and thence to the lateral

CATFISH ANATOMY & PHYLOGENY

31

syc

•op ih

Fig. 23 Hvpophthalmus edentatus suspensorial and opercular bones, medial view (cartilage
dotted). Specimen BMNH 1972.7.27: 675-678. Scale = 5 mm.

ethmoid. The pterygoid (pty, Fig. 23) contacts the quadrate synchondrally, and the
hyomandibular process via a dentate suture. The hyomandibular process (hy) extends
anteriorly along the dorsal rim of the quadrate (q) which it meets synchondrally. The stem
shank of the hyomandibula contacts the central dorsal rim of the quadrate via a
synchondrosis. Posteriorly, it is separated by a wedge of cartilage (syc), which may represent
the symplectic. The interhyal (ih) articulates between this cartilaginous element, the
posterodorsal edge of the quadrate, and the upper rim of the interoperculum.

The identity of the various bones comprising the suspensorium in siluroids is doubtful.
Regan (1911) simply referred in pimelodids to 'pterygoids' lying between the hyomandibula
and palatine. Compared with most other teleosts the pterygoid series is, for the most part,
incomplete since it lacks one of the pterygoid elements usually present. Authors have been at
variance as to whether it is the ecto- or endopterygoid which is absent (see comments of
Kindred, 1919; Alexander, 1965 and Gosline, 1975).

The problems of identifying siluroid suspensorial elements is illustrated by reference to the
pimelodid Pinirampus (Fig. 24). The pterygoid is like that of Hypophthalmus in having a
sharply demarcated ventral portion that articulates with the quadrate, whilst the dorsal
portion articulates with the hyomandibular process. An area of cartilage lies between the
branches of the pterygoid, the hyomandibular process and the quadrate. Lying close to the
anterior border of the pterygoid is a scythe-shaped bone, its elongate portion directed
ventrally. The bone is connected to the pterygoid and to the lateral ethmoid via ligaments.
Between the scythe-like bone and the pterygoid border is a splint-like bone which is attached
by connective tissue to both the scythe-like element and the palatine.

Two identification schemes of the suspensorial elements are presented. In one (Fig. 24,
left) the pterygoid is considered as the metapterygoid, the scythe-shaped bone as the
endopterygoid and the splint-like bone as the ectopterygoid. In the other (Fig. 24, right) the
hyomandibula plus metapterygoid are considered as forming a single element, or,
alternatively the metapterygoid is considered lost, and the pterygoid as comprising the endo-
plus, ectopterygoid. The two isolated bones are problematical. From its position in
connective tissue the splinter-like element could be a sesamoid ossification. The scythe-
shaped bone is characteristic of one group of pimelodids (see p. 37) and again, may be a

32

G. J. HOWES

hy+mp

ect

Fig. 24 Pinirampus pirinampu suspensorial and opercular (part) bones. Specimen BMNH
1934.8.20: 104-5. Scale = 3 mm. The upper, labelled drawings represent two interpretations of
the suspensorial elements. Shaded areas represent cartilage.

sesamoid bone. A small bone occurs in Hypophthalmus in a similar position and having the
same ligamentous attachments as the scythe-shaped bone in Pinirampus (see above). It
seems unlikely that these 'isolated' elements represent an ectopterygoid as they occur in the
wrong topographical positions. The problem of suspensorial bone homologies in siluroids
will be dealt with elsewhere (Banister & Howes, in preparation).

The hyoid arch (Fig. 25)

In Hypophthalmus the anterohyal and posterohyal are exceptionally elongate and,
unusually, are of equal length. The anterohyal (ahy) bears 8 branchiostegal rays (bsr) and the
posterohyal (phy) 6. The dorsal and ventral hyals (dh, vh) are small and separated from the
anterohyal by cartilage. The interhyal (ih) articulates with the posterior tip of the
posterohyal. The urohyal(uh) is a short oval bone lacking a medial ridge.

Gosline (1974) relates the number of branchiostegal rays to the length of the free sleeve of
the branchiostegal membrane. This certainly appears to be a valid correlation in
Hypophthalmus where the gill opening extends very far forward, and may, to a certain
extent, be applied to some pimelodid taxa as well. Examples are Sorubim and
Pseudoplatystoma which also possess long gill openings and a high number of branchiostegal
rays (11-1 6).

CATFISH ANATOMY & PHYLOGENY

vh

33

ih

Fig. 25 Hypophthalmus edentatus hyoid arch, lateral view. Specimen BMNH 1972.7.27: 675-678.

Scale = 5 mm.

According to McAllister (1968) a high number of branchiostegal rays is the primitive
condition in teleosts. On this basis he considers that amongst siluroids the Old World taxa
'. . .appear to be more primitive than the South American catfishes, having 4-20
branchiostegals as opposed to 3-17. . .'. The extent of these ranges and the degree of overlap
hardly justify such a statement. From McAllister's counts for siluroid families, the pre-
dominant number of branchiostegal rays would seem to be 9-1 1 and perhaps this, by virtue
of its widespread occurrence, should be thought of as the plesiomorph number rather than
the high number advocated by McAllister. Further comparative data, particularly on the
morphology and the placement of the branchiostegal rays on the hyoid bar are necessary
before any valid statement can be made concerning the apo- or plesiomorphy of their
number.

The relationships of the interhyal to the preoperculum shows some interesting modifi-
cations in the Pimelodidae. The usual siluroid condition, as in other ostariophysans, is for
the interhyal to articulate between the quadrate, or symplectic, and the hyomandibular
limb. In most siluroids the interhyal attaches, in the absence of a symplectic, ligamentously
to both quadrate and hyomandibula, abutting against the intervening cartilage (see
description of Hypophthalmus above). In Pseudoplatystoma the interhyal has shifted
posteriorly to such an extent that its connection with the hyomandibula and quadrate is lost,

34

G. J. HOWES

Fig. 26 Hypophthalmus edentatus upper and lower (part) branchial arches, dorsal view, left side
(cartilage stippled). Specimen BMNH 1972.7.27: 675-678. Scale = 3 mm.

and instead it articulates entirely with a preopercular fossa. Intermediate stages in the
development of this condition are to be found in Goldiella, Sorubim and Brachyplatystoma
where the interhyal has shifted even further posteriorly and is ligamentously attached to the
rim of a preopercular fossa alongside the hyomandibula.

The opercular series (Fig. 23)

The preoperculum (po) of Hypophthalmus is a thin, narrow bone covering the posterior
margins of the hyomandibula and the quadrate, virtually obscuring the interoperculum. The
interoperculum (iop) is roughly triangular but with a concave ventral border where it
overlaps the pectoral fin base. The operculum (op) is almost equilaterally triangular, its
posterior edge much fretted, and its entire structure cancellous.

The opercular bones of Hypophthalmus appear to be of a 'generalised' siluroid type, i.e.
a large operculum, almost vertical preoperculum and small interoperculum. There is
considerable variability in size and shape of these elements throughout the Siluroidei
(Howes, in press). Fink & Fink ( 1 98 1 ) maintain that the triangular shape of the operculum is
synapomorphic for the siluroids and gymnotoids. However, whereas the ventro-posterior
border of the operculum in siluroids is either straight or concave, that in gymnotoids is more
often convex, as in other otophysans.

The gill arches (Figs 26-27)

Upper elements (Fig. 26). The epibranchials of Hypophthalmus edentatus are narrow-
waisted, with expanded medial surfaces and bear 25-27 long, fine gill-rakers, As in all
siluroids examined, epibranchial (eb) 3 has an uncinate process which in Hypophthalmus is
rounded and overlies the central portion of eb4. Infrapharyngobranchial(if) 3 is well-ossified
and elongate, its posterior margin separated from eb3 and eb4 by a cartilaginous wedge. Of
the two alizarin preparations at my disposal, the larger shows two distinct chondrifications in

CATFISH ANATOMY & PHYLOGENY

uh

35

cm

Fig. 27 Hypophthalmm edentatus lower branchial arch elements; (upper) dorsal view. Specimen
BMNH 1972.7.27: 675-678. Scale = 3mm; (lower) cross section of left side gill-rakers and
ceratobranchials. cm = ceratobranchial membrane.

this wedge. These cartilages suspend a large oval toothplate bearing on its ventral surface
many minute, conical teeth (tp, Fig. 26).

Lower elements (Fig. 27). The basibranchials (bb) and hypobranchials (hb) are poorly
ossified. Basibranchial 1 is a triangularly shaped cartilage lying in the plane of the antero-
hyals; basibranchials 2 and 3 are rod-like ossified elements; basibranchial 4 is incorporated
within a broad cartilaginous cornua, the ossified lateral margins of which represent hypo-
branchials 4; basibranchial 5 is represented by a triangular cartilage whose apex is ossified.
Hypobranchials 1-3 are ossified only along their leading edges.

The ceratobranchials are the most elongate of any siluroid and carry at least 200 long
gill-rakers. The first two ceratobranchials (cb) have only a single row of gill-rakers (gr), along
the outer edge; the inner edge has a low wall of fibrous connective tissue (cm). The other
ceratobranchials have a double row of gill-rakers; the 5th is dentigerous, its anterior stem is
excessively elongate and meets its fellow along the midline. The gill-rakers on cb 5 occur
only along the inner margin; anteriorly the rakers of each ceratobranchial intermesh.

Roberts (1972) described the sieving mechanism of the gill-rakers. The arrangement of this
mechanism is shown in Fig. 27 as a transverse section of the lower gill-arch. Such an arrange-
ment does not occur in any other siluroid.

36 G. J. HOWES

Postcranial elements

As this study is concerned primarily with the cranial anatomy of Hypophthalmus, no
detailed description of the postcranial anatomy is presented. The following features,
however, are worthy of note.

The pectoral girdle is of simple construction, with a greatly elongated horizontal cleithral
limb and the coracoid reduced to a horizontal lamina. Such elongation of the cleithrum is
unknown in any other siluroid.

There are 62 vertebrae, excluding the complex vertebrae. This is an exceptionally high
number amongst siluroids and is exceeded only by members of the Clariidae. Virtually no
comparative data exist for vertebral numbers in siluroids. A superficial survey indicates that
the mean siluroid count is 40-45. High counts are found among Ariidae 49-50,
Ageneiosidae 49-50, Siluridae 50-51 and Clariidae 60 + . Among the Pimelodidae only
Luciopimelodus has an exceptionally high count 47-48.

The caudal fin skeleton of Hypophthalmus is of a plesiomorph pattern with no fusion of
any hypural elements (PH;1;2;3;4;5). Lundberg & Baskin (1969) found hypural fusion and
reduction of principal caudal fin rays to be widespread conditions. The most derived hypural
fusion patterns occur in the Plotosidae, Chacidae and Loricariidae. In the latter family
there is also an expansion of the last neural spine, which contacts the upper caudal element,
i.e. the fused hypurals 3-5, uroneural and epural. Certainly the derived caudal fusion
patterns shared by the Loricariidae, Callichthyidae and Astroblepidae are congruent with
synapomorphies involving the encapsulation of the swimbladder (see p. 27).

Other caudal fin patterns are not so easy to interpret and characters such as the absence of
an hypurapophysis in the Clariidae and Plotosidae are interpreted as parallelisms by
Lundberg & Baskin. Although both clariids and plotosiids trend toward anguilliform
morphology, it is interesting to note that the hypural fusion pattern in clariids (excluding
the Heteropneustidae) is, in contrast to plotosiids, plesiomorph. Even in such a seemingly
derived taxon as Allabenchelys all the hypurals are free.

Amongst the Pimelodidae, the predominant caudal fusion pattern is a free parhypural;
1 +2; 3 + 4; 5, which as Lundberg & Baskin (1969: 40) remark is significantly more advanced
than that of any bagrid. The hypural fusion patterns yield little support for the proposed
groupings of pimelodids presented below, except that Rhamdia and Pimelodella share a
similar fusion of hypurals, 1 +2 and 3 + 4.

Relationships of the Hypophthalmidae

Anatomically, the Hypophthalmidae is a unique group. No other siluroid has such an
arrangement of the eye musculature, hypertrophy of the jaw adductor muscles, reduction of
ethmoidal ossification, complexly developed mandibular barbel supporting elements,
elongated gill- and hyoid arch elements, and profoundly modified vertebral and swimbladder
elements.

Attention was drawn in the Introduction to the poverty of published data on siluroid
comparative anatomy. In order to apply cladistic methods in resolving interrelationships
extensive comparisons must be made thus, at this stage one cannot, with any degree of
certainty identify plesiomorph or apomorph characters amongst siluroids. However, from
this survey it is clear that the sister-group to the Hypophthalmidae lies within the
Pimelodidae. Synapomorphies are manifest in the structure of the ethmoid region, the
mandibular barbel articulation, the eye muscle arrangement, the complex vertebrae, and the
sensory canal system. The following is a synopsis of these characters:

1 . Ethmoid region with lateral cavity (p. 20) — shared with Luciopimelodus, Pinirampus and
Pimelodus (part).

2. Articulation of the mandibular barbels with large cartilaginous plates (p. 21) — shared
with Luciopimelodus, Iheringichthys, Sorubim, Pinirampus, Pseudoplatystoma,
Pimelodus (part).

CATFISH ANATOMY & PHYLOGENY 37

3. Anterior eye musculature of tendinous origin — shared with Sorubimichthys

4. Network of epidermal canals crossing the orbital and opercular regions — shared with
Luciopimelodus and Iheringichthys.

5. Posterior extension of the 'posttemporal' passing beyond the edge of the Miillerian ramus,
elevation of that ramus, and vertical path of the anterior portion of the vaugus lateralis
nerve — shared with Luciopimelodus.

In the course of this study only a limited number of pimelodid taxa was studied, but even
this preliminary investigation reveals that there are at least three major groups contained
within the Pimelodidae as presently recognised, viz:

1. Those species with the swimbladder partially encapsuled and usually with a caecal
posterior border; the 5th and 6th fused within the vertebral complex; the ventral superficial
ossification of the 4th and 5th centra forming a tunnel for the dorsal aorta; absence of a
retractor tentaculi muscle.

Included taxa: Iheringichthys, Pinirampus, Callophysus, Parapimelodus, Pimelodina,
Perugia, Megalonema, Luciopimelodus, Pimelodus (part). In addition, the first four genera
and Pimelodus maculatus all have a scythe-shaped pterygoid bone lying anterior to the main
pterygoid series and attaching to the lateral ethmoid via a ligament (p. 31).

2. Those species with the swimbladder well-developed, not encapsuled, and with attached
muscles (see below); 6 vertebrae forming the anterior complex centrum, the 4th and 5th
centra elongate and forming an aortic tunnel. A retractor tentaculi muscle present.

Included taxa: Pseudoplatystoma, Sorubim, Hemisorubim, Pimelodus (part).

The distribution of swimbladder muscles throughout the Pimelodidae as a whole has still
to be recorded. Bridge & Haddon (1894) described swimbladder compressor muscles in
Pseudoplatystoma, (their Platystomd) as originating from the occiput and inserting on the
antero-ventral wall of the swimbladder. These authors also described a tensor tripodis
muscle in Pseudoplatystoma and some other pimelodontid taxa. This muscle runs from the
exoccipital and inserts tendinously into the anterior wall of the swimbladder.

Alexander (1965: 110) was unable to trace the tensor tripodis in Pimelodus blochii,
Alexander's P. clarias. However, it appears in this species as a mere strip of tendinous fibres.
The muscle is also present in Sorubim and Hemisorubim

3. Those species with the swimbladder well-developed, not encapsuled, and without
compressor muscles; 5 fused centra, sometimes with superficial ossification but without an
aortic tunnel. The lateral ethmoid is transversly convex.

Included taxa: Rhamdia, Pimelodella, Heptapterus

The above groups include taxa presently distributed amongst four subfamilies, the
Pimelodinae, Luciopimelodinae, Callophysinae and Sorubiminae. None of the three groups
defined here corresponds with any subfamily as presently recognised. In addition, there is
another subfamily, the Pseudopimelodinae (Gomes, 1946). Representatives of this latter
group have not been thoroughly investigated in the course of this study, but in those that
have been examined the anterior part, at least, of the swimbladder is divided by an extension
of the complex vertebrae.

No meaningful phylogenetic classification is accomplished in recognising the above
named pimelodid subfamilies, based as they are on a mixture of plesio- and apomorphies.
Similarly, placing the Hypophthalmidae in a separate suborder (Chardon, 1968) simply
to recognise its morphological distinctiveness, serves no purpose in elucidating its
relationships.

From the characters analysed above synapomorphies have been identified that indicate the
sister group of the Hypophthalmidae to be in that assemblage which includes
Luciopimelodus and Iheringichthys. A more refined definition of this group must await a
thorough anatomical study of the Pimelodidae.

38 G. J. HOWES

Acknowledgements

This paper has benefited greatly from the critical and constructive comments of Drs
Humphry Greenwood, Keith Banister and Paul Skelton, to all of whom I extend my sincere
thanks.

I particularly thank Bernice Brewster, Lynne Parenti, Chris Sanford and Rob Travers for
so much helpful advice and technical assistance.

I am deeply indebted to those many ichthyological colleagues and associates who, over the
years, have provided specimens and given me so freely and patiently much information on
catfishes.

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Manuscript accepted for publication 18 October 1982

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Contents

<

1 GENERAL
29 JUL1983

A revision of the genus Epiclintes (Ciliophora: Hypotrichida) including a
redescription of Epiclintes felis comb. n. By P. G. Carey & E. C. Tatchell .

Notes on the Family Lekythoporidae (Bryozoa, Cheilostomata). By P. L.
Cook & P. J. Hayward

A new species of Arthroleptis (Anura: Ranidae) from the West Usambara
Mountains, Tanzania. By A. G. C. Grandison

The distribution behavioural ecology and breeding strategy of the Pygmy
Toad, Mertensophyrne micranotis (Lov.). By A. G. C. Grandison & S. Ashe

Additional notes on bariliine cyprinid fishes. By G. Howes

41
55
77

85
95

•

•

A revision of the genus Epidintes (Ciliophora:
Hypotrichida) including a redescription of
Epidintes felis comb. n.

Philip G. Carey

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD.

Eric C. Tatchell

Anatomy Department, Guy's Hospital Medical School, London Bridge, London SE1 9RT.

Introduction

Several hundred species of ciliated protozoa have now been documented from marine
interstitial sediments (Dragesco, 1974). However few of these organisms have been
adequately described; indeed some forms although regularly encountered in the extensive
literature on this group (Hartwig, 1980) have never been redescribed since the original
accounts given by Muller (1786) and Ehrenberg (1830). In an early study, Faure-Fremiet
(1950) put forward the hypothesis that interstitial ciliates had a cosmopolitan distribution.
This has subsequently been proved correct (Hartwig, 1977), and it has been recognised that
these organisms may play a dominant role in the benthic ecosystem (Fenchel, 1969). The
opportunity to redescribe one such truly cosmopolitan species came with the discovery of
large numbers of the highly contractile hypotrich ciliate Epidintes felis (Muller, 1 786) comb,
n. in muddy creeks near the estuary of the river Thames. A brief glance through the literature
concerning this species indicated that it had been poorly described. In addition there was
some confusion concerning synonymies and it was apparent that a revision of the genus
Epidintes was long overdue.

Materials and methods

Epidintes felis were collected from two locations on the south shore of the Thames estuary:
Oare creek, National Grid reference TR0163 and Faversham creek, National Grid reference
TR0262. Both locations are near Faversham, north Kent. The organisms were found in
small shallow (5- 10 cm) lagoons just below the high tide line along the sides of the two
creeks. They were found on the surface of the fine mud and detritus at the bottom of the
lagoons or beneath the surface film of the water. The salinity of the water was 70% sea water
(SW). Subsequently this species has been found at two sites on the north shore of the
Thames, at Shoeburyness, Essex, National Grid reference TQ9485. At this locality the
organisms were found firstly at the sand surface, 20 metres from shore, and secondly in
association with the red alga Ceramium sp. collected 10 metres from shore. All sites were
permanently covered with water at low tide.

Cells were cultured in filtered 70% SW, pH 7-8 at 18°C under constant illumination from
an 8 W fluorescent lamp. Subcultures were made every 5 days. Some difficulty in handling
the organisms was experienced due to their thigmotactic nature and inate fragility but when
quickly transferred using a wide bore micropipette, this problem was solved. Silver stains of
the cortex were prepared by the method of Tuffrau (1967). Silver stained preparations were
supplemented by observations using Nomarski interference, phase contrast and brightfield

Bull. Br. Mus. not. Hist. (Zool.) 45 (2): 41-54 Issued 28 July 1983

41

42 P. G. CAREY & E. C. TATCHELL

illumination, photographs and video recordings. It was found that good structural preser-
vation could be obtained with certain fixatives, consequently some specimens were prepared
for transmission electron microscopy. Two methods of fixation were employed:

(1) Nouzarede's Method (Nouzarede, 1977) for fragile ciliates. The specimens were first
fixed in 2% osmium tetroxide in 70% SW neutralised with solid calcium carbonate, for 10
minutes. They were then rinsed in 70% SW and transferred to 10 drops of sea water to a dish
and two drops of 25% glutaraldehyde (also neutralised with solid calcium carbonate) were
added. The organisms were fixed in this solution for 30 minutes. The ciliates were then
returned to the osmium solution for a further 20 minutes.

(2) Specimens were fixed in a solution of equal parts of 3% glutaraldehyde in phosphate
buffer and 1% osmium tetroxide in phosphate buffer for 20 minutes. These two fixatives were
prepared in Millonig's buffer (Millonig, 1961).

In both cases the ciliates were washed in 10% ethanol and then dehydrated in graded
alcohol solutions before being embedded in TAAB resin. Sections were stained in a saturated
solution of uranyl acetate in 50% ethanol for 10 minutes, followed by 0-4% lead citrate in
0-1 M NaOH (CO32~ free) for 5 minutes. The first fixative preserved the pellicle most
effectively and the second was employed to preserve the subpellicular structures.

Nomenclature

Epiclintes felis (Muller, 1 786) was first identified by Muller (1 786) under the name Trichoda
felis. Although the description was terse and the accompanying diagram gave little
information, it is still possible to recognise this very distinctive hypotrich. Muller (1786)
described the form of the body as being slightly curved and transparent. It was rather rotund
but posteriorly attenuated in the 'tail'. The ventral surface was seen to be covered by
'vibrating hairs' which covered the surface of the 'trunk' right up to the apex of the tail. Bory
St Vincent (1824) added a little more information to Muller's description and transferred
the organism to the genus Oxitricha as O. felis. He noted that the tail was frequently held
curved. Once again the diagram of the species gave few details as to the placement of the
cirri. The organism was also independently decribed by Claparede & Lachmann (1858) as a
new species in the family Oxytrichina, Oxytricha auricularis. Without knowledge of
Muller's T. felis, these authors gave a fairly detailed account of their 'new' species. They
carefully described the characteristic shape of the body and the fact that the dorsal surface
was seen to be ornamented by 'short batonnets' implanted in the tegument. Detailed infor-
mation regarding arrangement of cirri was lacking.

In his paper of 1862, Stein proposed the name Epiclintes auricularis to combine Muller's
(1786) T. felis and O. auricularis of Claparede & Lachmann (1858). Stein (1864) gave the
first detailed description of E. auricularis and supplied further details in his major work
(Stein, 1 867). However Stein ( 1 862) was in error in attributing the specific name ''auricularis''
to his new genus Epiclintes. Since the name 'felis' was presented in 1786, 70 years before O.
auricularis was published, the earlier name must take precedence. The correct name of the
taxon is Epiclintes felis (Muller, 1786) comb. n. Diesing (1866) was the third author
to independently identify E. felis and ascribe it to a new genus; Claparedia Diesing, 1866.
Although an invalid name, his useful description noted that the peristome of the organism
was situated asymmetrically and the body was extremely plastic or deformable. It was also
noted that the tail region was highly contractile and caudal 'styli' were seen to be present.

Wallengren (1900) in his description of the organism he called Epiclintes ambiguua made
a taxonomic error which has subsequently led to great confusion. He stated that the species
E. ambiguua was first named by Stein (1862) after Muller's (1786) original description of
Trichoda ambiguua. This is incorrect, Stein (1862) described E. auricularis, based on the
description of T. felis Muller (1786). However Muller (1786) did describe a T. ambiguua but
it is doubtful if this organism is a hypotrich and most certainly does not belong in the genus

REVISION OF EPICLINTES 43

Epiclintes. Kahl (1932) failed to note the error and adopted the name E. ambiguus; so that
subsequently the name E. ambiguus appears regularly in the literature. Apart from this lapse
in the nomenclature of the organism, Wallengren (1900) went on to give an excellent
description of his species, which was undoubtedly E. felis. Since these first accounts, many
workers on interstitial ciliates have included information regarding this hypotrich, but none
has clarified the important details of cirral arrangement, division and behaviour.

Morphological Description
Epiclintes felis (Muller, 1786) comb. n.

E. felis may be described as a genuinely psammophilic species, that is to say, it can be found
on, or at least temporarily associated with, sand of the intertidal zone rather than between
the interstices. From Figs 1 & 4 it can be seen that the body of this hypotrich is divided into
three distinct regions; a dorsoventrally flattened 'head' region leading via a thin neck to a
rotund 'trunk' which terminates in an attenuated 'tail'. Many authors have commented on
the fact that the organism is highly contractile, with the ability to shorten to at least 25% of
its initial length. It is quite clear that the diversity of form recorded for this species is the
result of the organism having undergone contraction to a varying degree. All observations
recorded here are based on extended cells unless otherwise stated. However in either con-
tracted or relaxed state, the three regions of the body are a permanent feature. The true
appearance of the body is best seen when the organisms have been left undisturbed for some
time. In normal locomotion over the substrate, the body is held extended and frequently
slightly curved. The head and tail regions are transparent but the central trunk region is
translucent and is frequently seen to be full of food materials. In culture E. felis was seen to
feed on bacteria but frequently, in freshly collected material, diatom frustules were seen
within the body. The hypotrich is colourless but Biitschli (1887-1 889) noted a yellowish tint.
Claparede & Lachmann (1858) were the first authors to quote an accurate size for Oxytricha
auricularis. They recorded a length of 300 um. Other reports quote a range of sizes varying
from 200-300 um, Stein (1864, 1867), Mereschkowsky (1879), Rees (1884), Kahl (1932). In
this study the species was found to vary in size between 100-350 urn.

'Head' Region

This region illustrated in Figs 2 & 5 is anteriorly rounded and dorsoventrally flattened. It is
truly auriform or 'ear-like' hence the etymology of Claparede & Lachmann (1858). In
extended, normally moving animals, the head is equal to or one and a half times as broad as
the trunk region. Many earlier authors regarded the trunk as possessing greatest breadth but
it is clear that their observations were based on at least partially contracted animals. From
Fig. 5 the asymmetric adoral zone of membranelles (AZM) extends around the anterior head
region on the ventral surface and along the left side of the oral area entering a distinctive
tubular buccal cavity. The cytopharynx is clearly visible but the cytostome is not. The AZM
itself comprises some 25-35 fine membranelles held erect. These are twice the length of the
cirri on the body, and extend as far as the buccal oveture. Here the membranelles form a
distinctive 'cage'.

'Trunk' Region

This region is ovoid tapering anteriorly to the head region and posteriorly to the elongated
tail. Its length varies with the degree of contraction; in the extended state it is at least twice
the length of the anterior portion. When contracted it may equal the head region in length. In
normal locomotion the surface of the cell adjacent to the substrate is somewhat flattened.
This would account for the thigmotactic nature of this region. On contraction the dorsal
surface of the organism becomes distinctly convex in shape, the ventral surface retaining its
flattened appearance. This central region may become distended by food material. The

44

P. G. CAREY & E. C. TATCHELL

contractile vacuole is single and as Kent (1880-1882) has correctly stated, it may be situated
centrally or may be found on the left near the termination of the peristome field. The
cytoproct is clearly visible at the dorsal surface, just at the junction of the trunk region and
tail. The number and form of the macronuclei has been poorly described; early authors
described this organelle as a single hoop-shaped body. The macronuclei are in fact small

Figs. 1-3 Epiclintes felis photographed by Nomarski interference microscopy: (1) Entire
organism, dorso-lateral view, bar represents lOjim; (2) 'Head' region, ventral view, bar
represents 7 urn; (3) Tail' region, ventral view, partly contracted, bar represents 8 )im.

REVISION OF EPICLINTES 45

and numerous and are, as Wallengren (1900) noted, spherical or ovoid. Due to difficulties in
staining this hypotrich, the micronuclei have never been observed.

'Tail' Region

The tail is relatively long, see Figs 1 & 4, at least the length of head and trunk combined, and
tapers quickly from the hind regions of the trunk, then less so to its end which is gently
rounded. As seen from Figs 1 & 3 this caudal region is very characteristic due to the arrange-
ment of cirri which give it a plaited or braided appearance, especially when contracted. Kahl
(1932), for example, noted that it resembled a fishbone. It is usually dorso ventral ly flattened
and distinctly ribbon-like. This region is highly contractile, much more so than the head or
trunk. Possibly longitudinal contractile fibres, akin to the myonemes of peritrich ciliates, are
present in the tail as shortening is so rapid. These could contribute to the braided appearance
seen so clearly on contraction. The tail serves to periodically jerk the body backwards during
normal locomotion. Also on shortening it is frequently noticed that the tail may bend or flex
at the junction with the trunk region. It has the ability to bend at least 90° from the
longitudinal axis of the cell without sustaining damage. Many of the early workers noted
correctly that this region contained no food materials.

Cirri

Dorsal Cirri

It can be seen from Fig. 6 that the dorsal surface of the organism is ornamented by sensory
bristles, which take the form of very short non-motile cilia each apparently arising from a
shallow pit. There are three distinct longitudinal rows of single, equally spaced bristles in E.
felis. The first row arises at the edge of the peristome at the extreme anterior edge of the cell,
passes over the head in the midline thence to the trunk and terminates at the posterior
extremity of the tail. This row is most easily observed from the lateral view. The two other
rows lie on the dorsal surface, at the lateral edges of the organism. They pass round the
anterior margin, down the sides of the body and meet the first row at the extremity of the tail.
Claparede & Lachmann (1858) were the first authors to identify these organelles in this
species. These were the 'batonnets of the tegument'.

Ventral Cirri

Unlike the dorsal surface, the ventral surface of E. felis is equipped with many rows of short
cirri. These take the form of thick bristles that taper distally. They are used extensively in
swimming. Biitschli (1887-1889) noted correctly that these cirri constantly oscillate. Due to
the constant movement of the organism and the translucency of the trunk region, it was very
difficult to observe the numbers and arrangement of cirri without silver impregnation
methods, and this could account for the great discrepancies reported by earlier workers. The
ventral surface of the head region is equipped with three diagonal rows of cirri, Fig. 5,
running right to left from the extreme right margin of the cell to just over two thirds the width
of the body. The first lies parallel to the long axis of the cell, the second and third becoming
rather more diagonal. The third row terminates at the end of the buccal cavity. Again, almost
impossible to observe without staining is the paroral apparatus which runs from the base of
the buccal cavity and part way up its right side. The membranes are small and may be
subdivided into two or three parts. The fourth ventral row, which may be termed the first
row of the trunk region, starts in the head, passes close to the end of the buccal cavity but
continues over the trunk region becoming less diagonally slanted as it does so. The cirri in
these rows are fairly short and stumpy and are all separated to the same degree, thus cannot
be sub-divided into frontoventrals or midventrals. Stein (1864, 1867) and Biitschli
(1887-1889) both stated that three rows were present in this area. Again the number of rows
of cirri on the trunk region has been poorly reported. Earlier authors did not state clearly
whether or not these ventral rows extended into the tail. Muller's (1786) description gave
four or five rows for this central portion while Stein (1864, 1867) claimed six or seven.

46

P. G. CAREY & E. C. TATCHELL

-azm

pa

cc

6

Figs. 4-6 Epiclintes felis: (4) Entire organism, ventral view, (azm) adoral zone of membranelles,
(be) buccal cavity, (cv) contractile vacuole, (tc) transverse cirri, (me) marginal cirri, (cc) caudal
cirri, (vc) ventral cirri, (cp) cytoproct; (5) 'Head' region, ventral view, (db) dorsal bristles, (pa)
paroral apparatus, (c) cytopharynx; (6) Lateral a, and dorsal b, views illustrating arrangement
of dorsal sensory bristles.

REVISION OF EPICLINTES

47

Claparede & Lachmann (1858) noted five short strong rows in the tail region but these
probably included rows of marginal cirri which are known to be present, no definite number
of trunk rows is mentioned in their description. Rees (1884) claimed a total of twelve rows
over the entire body, his diagram shows eight on the trunk and one on the tail.
Mereschkowsky (1879) gave a total of twenty for the entire cell, nine for the central portion
and five upon the tail. The excellent diagram of Perejaslawzewa (1886) shows a total of
seven, three on the anterior and four on the trunk, two of which continue into the tail.
Wallengren's (1900) description of E. ambiguua depicted six oblique ventral rows on the
trunk and six rows beginning in the mid-trunk region and entering the tail. Nevertheless it
can be seen that there are eight rows running diagonally across the body from right to left
(Figs, 4 & 7), the last two enter the tail region and are continued to the tip, passing near the
mid-line. These cirri are of similar length and are equally separated, but all of the eight rows
can only be visualised in stained preparations. Frequently it is seen that these rows of cirri
become interlaced and form a distinctive crest down the mid-line of the tail.

Marginal cirri are a characteristic feature of this hypotrich species, see Figs 1 & 4. There is
a continuous peripheral row of short marginals beginning just behind the termination of the
peristome field on both right and left sides, which pass down the lateral edges of the body and
join at the apex of the tail. The cirri on the right side appear slightly longer than the left.
Stein (1864) noted bristle-like marginals whilst Calkins (1902) showed that they projected
from the lateral edges and that they were elongated at the posterior of the cell. The two
marginal rows lying at the periphery of the tail and the two ventral rows lying adjacent to the
mid-line have, in earlier reports, often been seen as four separate rows in this caudal
prolongation. A band of transverse cirri, clearly distinguishable from the ventral cirri by
their separate rhythm of oscillation, run down the left side from the first trunk row to the first
beginnings of the tail (Fig. 7). They approximate in size and shape to other cirri of the ventral
surface. A very short band of caudal cirri, numbering four or five emerge from the ventral
surface of the tail. These are finer and slightly longer than the ventrals and marginals and are
the caudal 'styli' seen by earlier workers.

CC

Fig. 7 Epiclintes felis, ventral view of silver stained preparation, (tc) transverse cirri, (me)
marginal cirri, (cc) caudal cirri, (vc) ventral cirri.

48 P. G. CAREY & E. C. TATCHELL

Division

Division in E.felis was observed on a few occasions during this study but precise details are
lacking, therefore a description of stomatogenesis and associated development is not given.
Wicklow (1979; pers. comm.) clarified the events accompanying stomatogenesis in this
unusual hypotrich. In a detailed study of its morphogenesis he found that E.felis possessed a
ventral primordium from which many of its ventral rows develop. The row of transverse cirri
develop from the posteriormost ends of proter and opisthe cirral streaks. Fission is isotomic,
the opisthe retaining the elongate tail and the proter swimming away from division with a
simple cylindrical body. The new AZM was seen to develop from the anteriormost of the
transverse cirri, in the mid-trunk region.

infrastructure

When sectioned, E.felis is seen to be unique with the dorsal surface of the cell covered by a
multilamellate pellicle of about sixteen layers thick (Fig. 8). This thick pellicle extends down
either side of the organism but is modified on the ventral surface. In the vicinity of the
ventral cirri the pellicle reverts to the more normal ciliate structure (Pitelka, 1969), but in
other regions the multilamellate appearance is seen but the number of layers is reduced to
five or six. This unusual pellicle does not extend up onto the dorsal sensory bristles but
terminates in a funnel-shaped structure (Fig. 9) rather than a simple pit as seen for example
in Euplotes (Ruffolo, 1976), which surrounds the base of the organelle. The repeat distance
in this thick pellicle is about 6 nm, thus giving a total width for the pellicle of about 100 nm.
The lamellae have the appearance of a series of unit membranes in a collapsed condition.
The existence of this multilayered pellicle and its function in this highly contractile
hypotrich is certainly worthy of further investigation.

Diagnosis of the Genus EPICLINTES Stein, 1862

Trichoda Muller, 1 786 (In part)

Oxytricha Claparede & Lachmann, 1858 (In part)

Claparedia Diesing, 1 866

A free swimming genus of hypotrich, with a flexible and elastic body. The organism exhibits
characteristic cycles of contraction and relaxation. The body is divided into three regions;
anteriorly the head is dorsoventrally flattened, bounded at the apex of the cell by the
peristome. It tapers gradually to the central portion which is cylindrical. The trunk is twice
the length of the head and is usually filled with food materials and granular inclusions. The
posterior portion takes the form of an attenuate tail, the length of which may roughly equal
or extend to at least three times the length of both head and trunk combined. The trunk and
tail are flattened ventrally contributing to the organisms thigmotactic abilities. The
peristome is asymmetric, commencing right and enlarging as it traverses the anterior of the
cell. The AZM terminates on the left, within the head region, in a distinct buccal cavity.
Paroral membranes are present on the right side of this buccal cavity. The ventral surface of
this organism is equipped with several rows of short stumpy cirri passing longitudinally or
diagonally across the body. These cannot be separated into frontoventrals or midventrals.
Marginal cirri are placed in one or two rows along the sides of the trunk and tail. A row of
transverse cirri is present, running from just behind the peristome to where the trunk
narrows at the start of the tail. A short band of caudal cirri is present also at the tip of this tail.
Marine.

Species Descriptions

From the literature, nine species have been attributed to the genus Epiclintes Stein, 1862.
Clearly some of these species are not valid and some have been described on more than one

REVISION OF EPICL1NTES

49

Figs. 8-9 (8) Electron micrograph of Epiclintes felis illustrating the multilamellate pellicle of the
dorsal surface, fixed by method 1 . The pellicle is 16 layers thick with a repeat distance of 6-2 nm,
bar represents 40 nm; (9) Electron micrograph of Epiclintes felis illustrating a section through
a dorsal sensory bristle, fixed by method 2, bar represents 0-1 urn.

50 P. G. CAREY & E. C. TATCHELL

occasion. Thus study reduces the number of nominal species to three, E. felis, E. caudatus
Bullington, 1940 and E. radiosa Calkins, 1902. Corliss (1979) noted that one species of
Epiclintes resembled the genus Uncinata. However no member of the genus Epiclintes
recorded in the literature resembles this giant species with its characteristic beak-like
uncinus.

Two species that have been assigned to the genus, but differ fundamentally from the others
are Epiclintes vermis Gruber, 1884 and Epiclintes pluvialis Smith, 1900. E. vermis with its
vermiform body closely resembles other hypotrich genera such as Holosticha. E. pluvialis
superficially resembles Epiclintes but clearly is in possession of a symmetrical peristome. It
lacks ventral cirri and the dorsal cirri were said to be long and 'hispid'. These two species
differ sufficiently from the diagnosis of the genus Epiclintes to warrant exclusion.

Key to species of Epiclintes

1 Frontal 'styles' absent

Frontal 'styles' well developed, numbering four or five radiosa

2 Ventral cirri in two longitudinal rows caudatus

— Ventral cirri in eleven diagonal rows felis

Epiclintes felis (Muller, 1 786) comb. n.

Trichoda felis Muller, 1 786

Oxitricha felis Bory St Vincent, 1 824

Oxytricha auricularis Claparede & Lachmann, 1858

Epiclintes auricularis Stein, 1 862

Epiclintes ambiguua Wallengren, 1900

Epiclintes ambiguus Kahl, 1932

DIAGNOSIS A psammophilic species commonly found on or adjacent to the sand surface. The
body is characterised by its extreme flexibility and contractility. When extended, three
regions of the body are apparent. There is a typically auriform anterior, a cylindrical 'trunk'
region and an elongate 'tail'. The peristome is asymmetric running around the apex of the
cell and into a buccal cavity on the left. The paroral apparatus is small and is situated on the
right of the buccal cavity. The cytopharynx is clearly visible. The width of the anterior is
equal to, or one and a half times as broad as the trunk. This central region tapers posteriorly
to the tail. It is translucent and full of food materials. The contractile vacuole is single and
placed centrally or just behind the peristome. A cytoproct lies adjacent to the dorsal surface
at the junction of trunk and tail. Macronuclei are small, oval and numerous. The tail is
elongate, at least the length of the anterior and central portions combined. It is rounded at
the tip and often displays a plaited appearance. The dorsal surface is equipped with three
longitudinal rows of sensory bristles, running from the extreme anterior to the tip of the tail,
one in the mid-line, the other two at the lateral edges. Ventrally the head is equipped with
three rows of short cirri running diagonally from right to left. The trunk has eight rows
similarly arranged, the last two of which enter and run down the tail. These two progress
right to the tip. Two lateral rows of marginal cirri are present, beginning near the peristome
and extending to the point of the tail. A longitudinal row of transverse cirri runs down the
left side of the trunk, beginning at the first trunk row and terminating where the tail region
begins. In addition, a short row of four or five caudal cirri are present at the extremity of the
tail. A prepared slide of Epiclintes felis stained by the silver impregnation technique of
Tuffrau (1967) has been deposited in the collection of the British Museum (Natural History),
accession number 1982 : 2 : 12. Average length 100-300 um but may shorten to at least 25%
of this initial length. Epiclintes felis has been recorded from many locations. Claparede &

REVISION OF EPICLINTES

51

Lachmann (1858) found the organism on the coast of Norway and noted that it had also been
collected in the Baltic Sea. Stein (1864) similarly found this species in the Baltic but more
recent researches have shown it to be truly cosmopolitan. Hartwig (1977) gave an extensive
list of its distribution and locations will only be listed briefly here; North Sea, Baltic Sea,
White Sea, Gulf of Naples, French Adriatic and Mediterranean Coasts, Black Sea, Caspian
Sea, Sea of Japan, Bay of Bengal, American Atlantic Coast, Coast of Mauretania, Brazilian
Coast and the Coast of Great Britain.

10

Fig. 10 Epiclintes caudatus After Bullington, 1 940, ventral view.

Epiclintes caudatus Bullington, 1940

DIAGNOSIS (Fig. 10) A very long and slender hypotrich, widest centrally and dorsoventrally
flattened. It is extremely extensile and contractile. The body is divided into three regions, the
anterior portion is almost but not quite equal to the central portion in width. The 'tail' is
long and narrow, slightly enlarged at the tip. The peristome is asymmetric and traverses
around the head region terminating in a buccal cavity on the left side. Sensory bristles
emerge from the dorsal surface, at the lateral edges of the cell, beginning in the mid-
peristome region and extending to the tip of the tail. Two longitudinal rows of ventral cirri
are present, situated on either side of the midline. Two rows of marginal cirri are also
present. In the original description the colour was said to be yellowish and the length at rest
was 354 um and the width 28-6 um. The three regions of the body, an asymmetric AZM, the
presence of dorsal sensory bristles, marginal cirri and a high degree of contractility con-
tribute to the inclusion of this species in the genus Epiclintes. Only two rows of ventral cirri
were noted, running longitudinally. This, together with a very elongate head region and a
very long tail process serve to distinguish it from E.felis. A marine species isolated from the

52

P. G. CAREY & E. C. TATCHELL

11

12

13 15

Figs. 11-15 (1 1) Oxytricha retractilis After Clapareide & Lachmann, 1858, ventral view of re-
laxed and contracted organisms; (12) Oxytricha longicaudata After Strethill- Wright, 1862, ven-
tral view of relaxed and contracted organisms; (13) Mitra radiosa After Quennerstedt, 1867,
lateral and ventral views of relaxed organism; (14) Mitra retractilis After Kahl, 1932, ventral
view of relaxed organism; (15) Epiclintes radiosa After Calkins, 1902, lateral and ventral views
of relaxed organism.

REVISION OF EPICLINTES 53

West Indies. These organisms usually swim in left spirals but when locomoting over a sub-
strate or the underside of the surface film they were seen to move without spiralling, in
circles, to the left and right.

Epiclintes radiosa Calkins, 1 902

Oxytricha retractilis Claparede & Lachmann, 1858
Oxytricha longicaudata Strethill-Wright, 1862
Mitra radiosa Quennerstedt, 1 867
Epiclintes retractilis Kent, 1 88 1-1 882
Mitra rectractilis Kahl, 1932

DIAGNOSIS A marine species which like E. felis has been described by many authors in
various states of contraction. O. retractilis Claparede & Lachmann (1858) and O.
longicaudata Strethill-Wright (1862) were both described in the extended state. (Figs 1 1 &
12). They identified three distinct regions of the body, a narrow anterior with asymmetric
peristome, a central region that was distinctly ovoid and an elongate highly contractile 'tail'
which nearly disappeared on shortening. Strethill-Wright (1862) noted that this tail had a
plaited appearance. Five long anterior cirri were observed by both authors. Quennerstedt
(1867), Kahl (1932) and Calkins (1902) all described the organism in the contracted state
whereupon the tail shortens considerably and the head and central portions tend to be seen
as a single unit. Quennerstedt (1867) (Fig. 13) noted the organism was five times as long as
broad and that five long straight frontal cirri, he termed 'styles', were present. This author
also noted that a single row of marginals were present upon each side of the anterior and
central portions, and a double row bordered the tail. Kahl (1932) noted that M. retractilis
had a short row of five transverse cirri by which the organism attaches to the substrate. Again
four or five long frontal bristles were noted (Fig. 14). Calkins (1902) described the peristome
as wide on the right and narrowing as it approached the left side of the body, mention was
also made of attachment to the substrate by posterior cirri (transverse cirri) of the tail (Fig.
15). Kahl (1932) was the first to group these independently described organisms under the
name Mitra retractilis. However as Kent (1881-1882) had already pointed out, the name
Mitra had previously been assigned to a genus of mollusc and was therefore preoccupied.
Thus Kahl's designation is invalid. The additional description by Calkins (1902) erected the
name Epiclintes radiosa for the organism described by Quennerstedt (1867). The sizes
quoted for these organisms are 80 urn for O. retractilis, 75 urn for M. retractilis and 45 um for
E. retractilis. This marine species has been recorded from Norway, Scotland, Heligoland,
Kiel Sea, Finland, White Sea and the Atlantic Coast of the USA. Vital information on
numbers and arrangement of cirri are unfortunately lacking, but they have been grouped due
to the characteristic frontal cirri and the conspicious highly contractile tail. Further
information is required on this interesting species before a definite decision can be taken
regarding its validity.

References

Bory de St. Vincent, J. B. 1824. Encyclopedic Methodique II, Histoire Natur. des Zoophytes, fais suite

a I'hist. nat. des vers de Bruguiere. Paris.
Bullington, W. E. 1940. Some Ciliates from the Tortugas. Papers Carnegie Inst. Wash. Tortugas

Lab. 32: 179-221.
Butschli, O. 1887-1889. Protozoa, Abt. Ill, Infusoria und System der Radiolaria. In Bronn, H. G.

(Ed.) Klassen undOrdnungdes Thiers-Reichs 1: 1098-2035. C. F. Winter, Leipzig.
Calkins, G. N. 1902. Marine Protozoa from Woods Hole. Bull. U.S. Fish. Commn. 21: 413-468.
Claparede, E. & Lachmann, J. 1858. Etudes sur les infusoires et les rhyzopodes. Mem. inst.

Genevoise. 5: 1-482.
Corliss, J. O. 1979. The Ciliated Protozoa: Characterisation, Classification and Guide to the

Literature. 2nd ed. Oxford: Pergamon.

54 P. G. CAREY & E. C. TATCHELL

Diesing, K. M. 1866. Revision der Prothelminthen. Sber. haver. Akad. Wiss., Math. Natur. Kl.

Munch. 52: 505-580.
Dragesco, J. 1974. Ecologie des Protistes Marins. In De Puytorac, P. & Grain, J. (Eds) Actualites

Protozoologiques 1: 219-228.
Ehrenberg, C. G. 1830 (1832). Beitrage zur Kenntnis der Organisation der Infusorien und ihrer

Geographischen Verbreitung, besonders in Siberien. Abh. Akad. Wiss. DDR, Year 1832, 1-88.
Faure-Fremiet, E. 1950. Ecologie des Cilies Psammophiles Littoraux. Bull. Biol. France. Beige. 84:

35-75.
Fenchel, T. 1969. The Ecology of Marine Microbenthos. IV. Structure and Function of the Benthic

Ecosystem, its Chemical and Physical Factors and the Microfauna Communities with Special

Reference to the Ciliated Protozoa. Ophelia 6: 1-1 82.
Gruber, A. 1884. Weitere Beobachtungen an Vielkernigen Infusorien. Ber. Naturf. Gesell. Freiburg, IB.

3: 57-69.
Hartwig, E. 1977. On the Interstitial Ciliate Fauna of Bermuda. Cah. Biol. Mar. 18: 1 13-126.

1980. A Bibliography of the Interstitial Ciliates (Protozoa): 1926-1979. Arch. Protistenk. 123:

422-438.

Kahl, A. 1932. Urtiere oder Protozoa I. Wimpertiere oder Ciliata (Infusoria) III. Spirotricha. In

Dahl, F. (Ed.) Die Tierwelt Deutschlands 25: 399-650. G. Fischer, Jena Pt.
Kent, W. S. 1880-1882. A Manual of the Infusoria, 1-3. David Bogue, London.
Mereschkowsky, C. 1879. Studien iiber die Protzoen des Russlands nordlichen. Archiv. fur Mikros

Anat.16: 153-248.
Millonig, G. 1961. Advantages of a phosphate buffer for OsO4 solutions in fixation. J. appl. Phys.

32: 1637.

Muller, O. F. 1 786. Animacula Infusoria Fluviatilia et Marina. Havniae et Lipsiae.
Nouzarede, M. 1977. Cytologie fonctionelle et morphologic experimental de quelques protozoaires

cilies mesopsammiques geant de la famille des Geleiidae (Kahl). Bull. Stat. Arcachon (N.S.) 28,

(Suppl.) (year 1976), 1-315.
Perejaslawzewa, S. 1886. Protozoen des Schwarzen Meers. Zapiski Novorossiiskago Obs.

Estestvoispytatelei 10: 79-1 14 (formerly: Memoiren der Neurus. Ges. derNaturh. zu Odessa.)
Pitelka, D. R. 1969. Fibrillar Systems in Protozoa. In T. T. Chen (Ed.) Research in Protozoology

3: 437-469. Pergamon Press N.Y.
Quennerstedt, A. 1867. Bidrag till Sveriges Infusorie — fauna. Ada Univ. Lund. Vol. II, Tomes 4-5

pg.41.
Rees, E. 1884. Protozoaires de 1'escault de Test. Tijschr. d. Nederl. Dierk. Vereenig. Supp. Dl:

593-673.
Ruffulo, J. J. 1976. Fine Structure of the Dorsal Bristle Complex and Pellicle of Euplotes. J.

Morphol. 148:469-487.
Smith, J. C. 1900. Notices of some undescribed infusoria, from the infusorial fauna of Louisiana.

Proc. Am. Mic. Soc. 21: 87-94.
Stein, F. R. 1862. Neue oder nicht genugend bekannten Infusorienformen Tiere. Amtl. Ber. Dl.

Naturf. u. Aerzte. 37: 161-162.
1 864. Ueber die neue gattung Epiclintes Stein. Sber K. bohm. Ges Wiss Math. Nat. Kl. (1): 44-46.

1867. Der Organismus der Infusionsthiere nach eigenen Forschungen. In Systematischer

Reihenfolge Bearbeitet, II, Leipzig.

Strethill-Wright, T. 1862. Observations on British Protozoa. Quart. J. Mic. Sci. New Ser. 2:

2 1 7-22 1 .
Tuffrau, M. 1967. Perfectionnements et practique de la technique d'impregnation au protargol des

infusoires cilies. Protistologica 3: 9 1-98.

Wallengren, H. 1900. Studierofver Ciliata Infusorien Ada Univ. Lund. 36: 1-52.
Wicklow, B. J. 1979. Epiclintes ambiguus and Kerona polyporum: comparative ultrastructure,

cortical morphogenesis and systematics of two hypotrich ciliates. J. Protozooi 26: 1 6A.

Manuscript accepted for publication 22 March 1 982

Notes on the Family Lekythoporidae (Bryozoa,
Cheilostomata)

P. L. Cook

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

P. J. Hayward

Department of Zoology, University College of Swansea, SA2 8PP

Introduction

(a) Cheilostome Ovicells

In recent years, several kinds of cheilostome ovicell have been investigated, and the origins
and relationships of the various body wall layers involved have been traced through
ontogeny. As a result, it has become obvious that the ovicell, which may be regarded as all
the walls defining and protecting the brood chamber and the contained embryo, has several
very different origins. Ovicells may be derived wholly, or in part, from diverticula of
terminal walls or from extensions of frontal walls and, or, frontal shields. They may be
derived wholly, or in part, from one or more zooids. They may be modified, interzooidal,
frontally budded kenozooids, or may be formed from one or more kenozooids budded from
the maternal zooid, which produces the ovum, or from a sequentially distal zooid or zooids
(see Cook, 1979 for brief review). The general term 'ovicell' may therefore be defined
functionally, but not morphologically, and it appears that the protective coverings for brood
chambers have often been evolved convergently and show only superficial similarities.

All ovicells which have been investigated in detail, and all those which may be inferred by
inspection to have analogous ontogeny, are 'distal' in position to the opening edge of the
operculum of the maternal zooid. Generally, this 'distal' orientation is also the same as the
direction of budding of zooids 'away from' the ancestrula as the colony develops. Even in
colonies with 'reversed frontal budding' (where the distal part of the orifice is directed
towards the ancestrular region, see below), the opening of the ovicell is placed close to the
orifice of the maternal zooid, on that side of the operculum which opens to allow protrusion
of the lophophore (see Cook & Lagaaij, 1976). This position is correlated with the position of
the coelomopore through which the ovum passes to the exterior. In all cheilostomes which
have been investigated, the coelomopore is placed at the base of the 'distal' pair of tentacles.
Passage of ova into ovicells is achieved by protrusion of the lophophore and apposition of
the coelomopore to the ovicell orifice (see Silen, 1 945).

Proximally placed ovisacs, which are principally uncalcified have been reported in the
anascan genus Aetea, but their ontogeny may be intussusceptive and quite unlike that of
other known ovicells (Cook, 1977). Proximal calcified ovicells have also been described in
several cheilostome ascophoran genera, which were grouped together in the family
Lekythoporidae by Levinsen (1909), and further reviewed by Canu & Bassler (1929). The
reported occurrence of 'proximal' ovicells infers that the position of the coelomopore, or at
least the behaviour of the maternal lophophore, is radically different from all other
cheilostomes in only one group, the Lekythoporidae. Such a difference would be
fundamental, although it could be postulated for the genus Inversiula (Microporellidae),
which is apparently equally aberrant, and where the operculum is hinged distally (see

Bull. Br. Mus. nat. Hist. (Zool.) 45 (2): 55-76 Issued 28 July 1 983

55

56 P. L. COOK & P. J. HAYWARD

Moyano, 1972). The brood sac in Inversiula is an interior diverticulum of part of the
cuticular outer body wall (tentacle sheath) housed in specially enlarged brooding zooids. The
operculum is orientated almost vertically, which would apparently allow passage of ova
from a partially protruded lophophore into the ovisac in the 'normal' manner. This has
never been observed in living specimens, but Nielsen (1981) has described passage of ova
into ovicells closed by the operculum, in which the lophophore was not protruded, but was
dimorphic, with very short tentacles.

The hingeing of the operculum in the Lekythoporidae is not like that of Inversiula and
investigation of specimens of the five genera attributed to the family shows that the ovicell is
placed in the 'normal' distal position with respect to the orifice, but that the pattern of
astogeny is one of 'reversed frontal budding'. The modifications in zooidal morphology
resulting from this type of growth are described below.

(b) Frontal budding and colonial and zooidal morphology

Frontal budding is a common phenomenon only in cryptocystidean ascophorans, although it
occurs sporadically, in a modified form, initiating overgrowths in umbonuloid ascophorans
and some anascans. In some cryptocystidean ascophorans, columns of successive, frontally
budded zooids may occur (see Banta, 1972, fig. 1 (1), 3). Each zooidal element arises as an
intussusceptive expansion of frontal cuticle, with underlying epidermis, of the primary zooid
of the series. These together form the boundary wall of an expansion of hypostegal coelom,
which is presumed to be provided with nutrients from the visceral coelom by means of the
marginal frontal septulae in the frontal calcified shield. As the cuticle-covered bud reaches
its full size, a secondary cryptocystidean frontal shield, also with marginal septulae, grows
into, and partitions, its coelom into visceral and hypostegal elements. Cryptocystidean
frontal shields are interior walls (see Boardman & Cheetham, 1973), and because there is
communication between hypostegal and visceral coeloms, these zooids have the potential to
form successive series of frontally budded members vertically to the horizontal plane of the
primary zooids. Functionally, frontally budded zooids may be regarded as being identical to
primary zooids budded through septulae in distal or lateral walls; but morphologically, they
may also be regarded as extensions only of the originating primary zooid, because they have
no basal walls. An even more integrated kind of frontal budding is found where more than
one primary zooid contributes to the development of more than one frontal bud. These
interzooidal frontal buds may form nodular masses or erect branches, and are sometimes
composed of groups of integrated polymorphic zooids with differing functions (e.g.
Hippoporidra, see Cook, 1983). Frontally budded zooids frequently have morphologies and
orientations different from those of the primary series (see Boardman et al, 1969, fig. 5), and
apart from not possessing basal walls, often have no recognisable lateral walls.

In some minute, globular colonies, all budding is frontal, and the position and orientation
of the zooids is rigidly patterned and 'reversed' with respect to the direction of growth. In
these forms, the distal side of the orifice, the side on which the operculum opens, is directed
towards, not away from the ancestrular area, as it is in 'normal' colony growth. In addition,
all the calcified walls may be regarded as frontal shields, which consist of a 'concealed'
portion, from which new frontal buds arise, and an 'exposed' portion which surrounds the
orifice (see Cook & Lagaaij, 1976). Combinations of 'normal' and 'reversed' orientations of
frontal buds occur in the genus Sphaeropora (see Cook & Chimonides, 198 1 ).

In the Lekythoporidae all colonies are known or inferred to have minute, encrusting bases.
In Lekythopora and Poecilopora zooids are budded distally from the ancestrula for only one
or two astogenetic generations. The rest of the colony is composed of erect branches of
frontally budded zooids, each of which is orientated with the distal edge of the orifice towards
the outer face of the branch. Each zooid bud arises interzooidal ly from more than one zooid
of the earlier astogenetic generation and develops a cryptocystidean frontal shield with
numerous frontal septulae. Fully formed zooids are vase-shaped or columnar and all the
vertical calcified walls (whether axial or peripheral) form one continuous 'frontal' shield.

FAMILY LEKYTHOPORIDAE

57

m

m

Fig. 1 Diagrammatic, idealised longitudinal section through axis of an erect branch of
Lekythoporidae (based on Lekythopora and Poecilopord). Direction of colony growth
(astogenetically distal), arrow at right. External cuticles only shown (thin lines); Primary
calcification (thick lines); secondary and extrazooidal calcification stippled; as ascus; ax axial
frontal septulae; b topographically distal, frontal bud; ec extrazooidal coelom (derived from
amalgamation of hypostegal coelom of several zooids); he hypostegal coelom; ia interzooidal,
frontally budded avicularium; m mandible; o brood chamber within peristomial ovicell; op
operculum (direction of opening arrowed); p peristome; pa peristomial avicularium; ps
peristomial frontal septula; t tube connecting peristomial avicularium with frontal septula; ts
tentacle sheath.

58

P. L. COOK & P. J. HAYWARD

FAMILY LEKYTHOPORIDAE 59

Septulae are present in the axially directed 'concealed' part of the shield, and in the out-
wardly directed part. This, together with the circumoral region forms the equivalent of the
'exposed' part of the frontal shield in conescharelliniform colonies with 'reversed frontal'
budding (see Cook & Lagaaij, 1976, and Fig. 1). One or more circles of septulae surround the
primary orifice. The septulae allow the expansion of hypostegal coelom axially to form new
zooid buds, and peripherally to allow thickening of the exposed frontal shield calcification
and, or, the budding of peristomial or interzooidal avicularia and kenozooids. The primary
orifice is terminal, and is orientated with the proximal edge of the operculum directed axially
and in a more 'distal' position, with respect to the direction of growth of the branch, than the
outwardly directed (morphologically) distal edge (see Fig. 1). This is because the longitudinal
axes of the zooids are curved towards the outside of the branch. The peristome arises as a
ridge between the circum-oral septulae and other frontal septulae. It becomes tubular and
often very elongated, continuing the outward curve of the zooid axis. The ovicell is an
expanded diverticulum on the peripheral, morphologically distal, side of the peristome, but
the curvature of the zooid and peristome is such that it appears to be topographically
'proximal' in position (see Fig. 1). This illusion is completed in several species when
the peristome develops a small secondary 'sinus' on its astogenetically distal, but
topographically proximal side (Fig. 18).

The problem of interpreting the relationships of the primary orifice and the ovicell is
exacerbated by the length of the peristome and by the rapid development of secondary
calcification and frontally budded kenozooids in most species. The hypostegal coeloms may
even amalgamate, with the breakdown of interzooidal cuticle, and form extrazooidal coelom
with massive extrazooidal calcification. The presence of septulae and extension of
hypostegal coelom on the outer sides of the peristome also assists this calcification. The
exposed frontal shields, peristome and even ovicells sometimes become completely obscured
within only two astogenetic generations of the growing edge or tip of the branch (Fig. 2).

Longitudinal and transverse sections reveal details of the relationships of zooid budding
and communication, but because most zooidal axes are curved in more than one plane, com-
plete axial sections cannot be obtained. In colonies with regularly cylindrical branches,
Catadysis, or irregular cylindrical extensions, Lekythopora and Orthoporidra, all zooid
orifices are orientated with their proximal edges pointing axially and nearly vertically, i.e. in
the direction of growth. In colonies which appear superficially to be composed of two
'laminae' of zooids, Poecilopora, the proximal sides of orifices oif one 'lamina' are directed
towards those of the opposing 'lamina', in the same manner as those of the erect, but rooted
genus Flabellopora (see Harmer, 1957). In colonies with rounded branches with one 'frontal'
surface, Turritigera, the zooidal axes are twisted as well as curved, but the proximal edges of
the orifices are directed towards the axis of the branch. The 'basal' side of these colonies is
entirely composed of extrazooidal calcification (Fig. 8).

Avicularia occur on the edge of the peristome in all species. They are derived from
circum-oral frontal septulae, and one proximo-lateral avicularium at least is always present,
although several may occur. A long, tubular connection between the avicularian chamber
and the originating septula passes through the peristomial calcification. This often traces a
curved path, which perhaps reflects the distortions in zooidal axes which occur during calci-
fication of the bud. Sometimes the avicularia are raised on long hollow processes, which may
bear small secondary avicularia laterally as well as terminally. The long processes become
obscured during later thickening, leaving the avicularia at the surface of the branch, where
they may appear to be frontally budded interzooids. In addition, large frontally budded,
interzooidal avicularia also develop later in astogeny, especially at the bases of branches.
The apparently 'confused' budding pattern (especially in Lekythopora and Orthoporidra),
and the effects of extrazooidal calcification therefore make it difficult to distinguish the
origins of avicularian types except near the growing tips. Similar difficulties in the genus
Turbicellepora have been described by Hayward (1978), who also discussed the descriptive
terminology applicable to the large 'spatulate' avicularian rostra which often occur in
'celleporine' forms. Measurements of zooids are also difficult to define and obtain, because

60

P. L. COOK & P. J. HAYWARD

L

Figs 6-9 Scanning electron micrographs of orifices of Lekythoporidae; arrows indicate direction
of colony growth: (6) Catadysis immersum (Busk) BMNH 1963.2.12.247, Montevideo, part of
branch showing secondary orifices and frontal pores x26; (7) C. immersum, secondary orifice
with avicularia. Note astogenetically distal (topographically proximal) edge of primary orifice at
lower level x!30; (8) Turritigera reticulata sp. nov. BMNH 1890.4. 16. 2 A, Marion Island,
broken edge of branch from distal side showing one primary orifice (centre), zooid cavities, and
thick 'basal' extrazooidal calcification of branch x57; (9) T. fenestella sp. nov. BMNH
1890.4. 16. 2B, Marion Island, detail of zooid at branch tip, showing sinuate primary orifice,
secondary orifice and large avicularium x 1 12.

the shape of zooids is often irregular, the primary orifice is completely hidden and secondary
calcification obscures all limits of structures. One measure of the relative length of zooids
may be the distance between the centres of secondary orifices in distal series, but this is very
susceptible to error if peristomes are worn or broken. Opercula and primary orifices may
often be visible in broken branches, and examination from the topographically 'proximal'
(zooid interior) side may reveal differences in shape between them.

FAMILY LEKYTHOPORIDAE 61

Systematic Section
LEKYTHOPORIDAE Levinsen
Lekythoporidae Levinsen, 1909 : 89, 383.
TYPE GENUS. Lekythopora MacGillivray.

DESCRIPTION. The characters of the family are redefined as follows. Colonies erect or
semi-erect, arising from a small encrusting base which becomes strengthened by frontally
budded kenozooids, avicularia and, or, extrazooidal calcification later in astogeny. All
zooids of the secondary zone of change and repetition (see Boardman et ai, 1969) are
frontally and interzooidally budded. Zooids columnar, with a centrally placed, terminal
primary orifice. Proximal sides of orifices and asci orientated towards the central axis of the
branch. All vertical calcified walls formed as an interior, cryptocystidean shield; axial part
concealed, peripheral part and circum-oral region exposed. Shield with a hypostegal coelom
and frontal septulae, some of which form a circum-oral ring. Primary calcified orifice
straight, curved or sinuate proximally, surrounded by a raised, tubular peristome. Avicularia
arising from circum-oral septulae, occasionally numerous, always one proximo-laterally,
which appears to be topographically distal. Subrostral chambers terminal, on the edge of the
peristome or a mucronate process arising from the peristome. Rostra acute or rounded,
mandible slung on a complete bar. Subrostral chamber connected to the originating septula
by a long tube, which passes through the calcified wall of the peristome. Interzooidal,
frontally budded avicularia sporadic, variously orientated, mandible slung on a complete
bar, which may bear a ligula. Brood chamber protected by a large, globular peristomial
ovicell, sometimes with an exposed, frontal cuticular area, or with pores and small avicularia
on its surface.

REMARKS. The family has been defined (e.g. by Bassler, 1953) to include the genera
Lekythopora MacGillivray, Poecilopora MacGillivray, Turritigera Busk, Catadysis Canu &
Bassler, Orthoporidra Canu & Bassler and Actisecos Canu & Bassler. Of these, all but
Actisecos have been described as having 'proximal' ovicells. Actisecos, a rooted, lunulitiform
genus, has little in common with the other genera and was assigned to the family Actisecidae
by Harmer (1957 : 854). The remaining genera differ from each other principally in details of
budding pattern, the shape of the primary orifice and operculum, and the characters of
avicularia and ovicells.

Most of the specimens examined are small, rarely more than 10-15 mm in height, and
type material is often worn and fragmentary. A few colonies, C. immersum, O. compacta, T.
reticulata and T. fenestella, are relatively large and exceed 50 mm in height or width. Some
species show distinct substratum preferences. Lekythopora is often found on erect hornerid
and adeonid bryozoans, Poecilopora on flexible cellariiform and cellulariiform bryozoans or
on hydroids, Turritigera is often associated with other bryozoans or with polychaete tubes,
and colonies of Orthoporidra may originate on small stones.

With one exception, Poecilopora cribritheca, all species are from the Southern
hemisphere, and perhaps apart from Lekythopora, are from fairly deep, sometimes very
deep, cold waters. Generally, bottom temperatures have been given only for Antarctic
localities in published data, and those for low latitude, 'tropical' and 'subtropical' specimens
are not known (see Table 2). Several Antarctic species have been reported as abundant, but
material examined here, from the Zoological Museum Amsterdam (ZMAC), the National
Museum of Victoria (NMV), the British Museum (BMNH), and the Manchester Museum
(MM) collections, is not generally plentiful. In view of the enormous, but unworked
collections which have been made from the Antarctic shelf, and the increasing number of
species becoming known from deep waters, it should be possible, theoretically at least, to
obtain larger quantities of some of the species described here, and to analyse their population
variation in colony form and zooidal morphology. Several species appear to belong to com-
plexes, composed of widely distributed populations, each with different character corre-
lations. Additional information on early astogeny, and internal features such as tentacle

62 P. L. COOK & P. J. HAYWARD

Table 1 Average measurements of zooids of Lekythoporidae (in mm)

Lz Iz Lpo Ipo Lov lov

L. hystrix 0-70 0-40 0-12 0-11 0-20 0-40

P.anomala 0-40 0-28 0-08 0-07 0-15 0-20

P. cribritheca 0-50 0-30 0-12 0-11 0-18 0-20

T.stellata 0-50 0-30 0-13 0-16 0-20 0-25
Stn 320

T.stellata 0-60 0-35 0-13 0-18 0-25 0-35
Stn 142

T. reticulata 1-20 0-50 0-13 0-11 0-40 0-50

T. fenestella 0-75 0-35 0-17 0-12 0-25 0-30

C. immersum 0-65 0-50 0-12 0-17 0-25 0-35

O.compacta 0-80 0-50 0-19 0-23 0-40 0-50

O.solida 1-20 0-50 0-20 0-25 0-40 0-50

O.petiolatus 0-60 0-33 0-12 0-13 0-24 0-30

Table 2 Nominal Recent records of Lekythoporidae and Orthoporidroides. Additional temperature data from
Murray, 1895 and Livingstone, 1928; estimated depths and temperatures in parentheses

Depth Temperature
Species Locality data Latitude Longitude (metres) (°C)

Reference

Lekythoporidae
L. hystrix Port Phillip Heads 38°30'S 144°30'E (60) (+17)
L.hvstrix Port Western 38°30'S 145°30'E
L. hystrix Port Jackson 34°S 151°E (60) (+17)

BMNH
BMNH
BMNH

L.avicularis Port Jackson 34°S 151°E 146 (+17)

Maplestone 1909

P.anomala Port Phillip Heads 38°30'S 144°30'E (60) (+17)
P.anomala Twofold Bay 36°59'S 150°20'S 275 (+12-8)
P.anomala Maria Island 42°37'S 148°E 73

BMNH
BMNH

Thornely 1924

P. cribritheca Sulu Archipelago 6°8'N 121°19'E 275 (+12)

Harmerl957

T.stellata Montevideo 37°17'S 53°52'W 1100 +2-8
T.stellata Cape of Good Hope 35°04'S 18°37'E 275 +8-3
T.stellata Cape Horn 53°S 68°W
T.stellata Is. Topar 50°8-5'S 74°41'W 360
T.stellata Caleta Hale 47°57'S 74°41'W 40-50
T.stellata BellinghausenSea 70°S 80°48'W 7500 +0-9
T.stellata BellinghausenSea 70°20'S 83°23'W 459 +0-8
T.stellata BellinghausenSea 70°15'S 84°6'W 659 +0-8
T.stellata BellinghausenSea 70°23'S 82°47'W 480 +0-8
T.stellata BellinghausenSea 71°14'S 89°14'W 460 -0-3
T.stellata BellinghausenSea 71°18'S 88°2'W 435 -0-3

Busk 1884
Busk 1884
Waters 1905
Moyano 1974
Moyano 1974
Waters 1904
Waters 1904
Waters 1904
Waters 1904
Waters 1904
Waters 1904

T. reticulata Marion Island 38°S 46°40'W

BMNH

T. fenestella Marion Island 38°S 46°40'W

BMNH

Continues

FAMILY LEKYTHOPORIDAE

63

Species

Locality data

Latitude

Depth
Longitude (metres)

Temperature
(°C)

Reference

T. spectabilis

Uruguay

36°49'S

53°15-4'W

1661-1679

d'Hondt!981

38°16-9'S

51°56-1'W

4382-4402

d'Hondt 1981

36°55-7'S

53°01-4'W

2707

d'Hondt!981

C. immersum

Montevideo

37°17'S

53°52'W

1100

+ 2-8

Busk 18 84

C. immersum

Burdwood Bank

54°25'S

57°32'W

103

Hay ward 1980

O. compacta

BellinghausenSea

70°S

80°48'W

7500

+ 0-9

Waters 1904

O. compacta

Bellinghausen Sea

71°18'S

88°02'W

435

-0-3

Waters 1904

O. compacta

Bellinghausen Sea

71°19'S

87°37'W

436

-0-2

Waters 1904

O. compacta

Halley Bay

75°31'S

26°36'W

BMNH

O. compacta

Scotia Bay

61°S

45°W

BMNH

O. compacta

McMurdo Sound

77°05'S

164°17'E

256

BMNH

O. compacta?

South Chile

62°28-5'S

59°41-5'W

119

BMNH

O. compacta?

Queen Maude Land

70°19-4'S

24°12-6'E

Redier 1965

to

to

70°20-9'S

24°13-4'E

O. setosa

Commonwealth Bay

64°32'S

97°20'E

201

Thornely 1924

O. setosa

Commonwealth Bay

66°08'S

94°17'E

220

Thornely 1924

O. setosa

Adelie Land

66°55'S

145°21'E

782

+ 1-8

Thornely 1924

O. setosa

Adelie Land

66°32'S

141°39'E

287

+ 1-62

Thornely 1924

O. setosa

Graham Land

65°S

64°W

750

Vigeland 1952

O. setosa

Graham Land

65°S

64°W

90

Vigeland 1952

O. setosa

Graham Land

65°S

64°W

140

Vigeland 1952

O. petiolata

Cape Horn

53°13'S

68°31'W

97

+ 6-6

Waters 1905

O. petiolata

Burdwood Bank

54°25'S

57°32'W

103

Hayward 1980

O. petiolata

Houtjes Bay

34°10'S

18°10'E

BMNH

O. solida

S. W. Australia

42°42'S

134°10'E

4758

+ 1-1

Busk 1884

Orthoporidroides

O. erectus

Magellan Straits

55°S

72°W

320

( + 7-8)

BMNH

O. erectus

Montevideo

37°17'S

53°54'W

1100

+ 2-8

BMNH

O. erectus

Is. Innocentes

50°33'S

74°53'W

150

Moyano 1974

O. erectus

Is. Topar

50°8-5'S

74°41'W

360

Moyano 1974

O. erectus

Canal Zenteno

52°49'W

73°40'S

30-40

Moyano 1974

O. aff. erectus

E. Falkland Is.

52°09'S

58°14'W

79

+ 8-3

BMNH

O. aff. erectus

E. Falkland Is.

50°29'S

58°52'W

140

+ 4-78

BMNH

O. aff. erectus

E. Falkland Is.

50°30'S

58°19'W

141

+ 5-4

BMNH

(9. aff. erectus

E. Falkland Is.

52°31'S

58°29'W

146

+ 5-45

BMNH

O. aff. erectus

E. Falkland Is.

50°50'S

57°13'W

144

+ 5-61

BMNH

O. robusla

Chile

37°37'S

73°40'W

600

Moyano 1981

number, opercular variation etc., should result in a better understanding of the systematic
relationships within the Lekythoporidae, and the relationships of the family with other
ascophoran groups.

LEKYTHOPORA MacGillivray
Lekythopora MacGillivray, 1883 : 194.
TYPE SPECIES. L. hystrix MacGillivray.

64 P. L. COOK & P. J. HAYWARD

DESCRIPTION. Colonies forming short, irregularly cylindrical branches. Zooids with several
series of frontal septulae. Primary orifice sinuate, operculum extended proximally but not
distinctly sinuate. Ovicells with a frontal, marginally porous area covered by cuticle.

REMARKS. L. hystrix appears to be the only species referable to the genus. Harmer (1957 :
884) referred Phylactella lucida Hincks to Lekythopora', this species has been discussed by
Cook (1968 : 220) and referred to Celleporina. L. perplexa Harmer (1957 : 884, pi. 59, figs.
5-7, 10) is also a 'celleporine' species with 'normal' frontal budding and tabulate ovicells and
may be provisionally referred to Celleporina. Harmer (1957 : 885) also mentioned an
unnamed species of Lekythopora from Japan. Examination of his specimens shows that these
colonies, too, are referable to Celleporina. L. laciniosa Calvet (1907 : 445, pi. 29, figs. 13, 14)
is attributable to Celleporina.

Lekythopora hystrix MacGillivray

Lekythopora hystrix MacGillivray, 1883 : 194, pi. 2, figs. 6, 6a-d; 1885 : 113, pi. 2, fig. 6: 1888 : 201,

pi. 156, figs. 4-10; 1895: 106, pi. 14, figs. 1-2. Waters, 1885 : 308. Brown, 1958 : 83.
Lekythopora avicularis Maplestone, 1909 : 273, pi. 78, fig. 12.

SPECIMENS EXAMINED. BMNH, Port Phillip Heads, Victoria, 1897.5.1.934,937,
1899.5.1.1328, 1934.10.20.103. Port Western, S. Australia, 1934.2.20.23. Australia, 1883.
10.15.125-145. Port Jackson, Sydney, 1981.4.1.1. NMV, Port Phillip Heads 646 II, 64612
(fig'd MacGillivray, 1888) and 64602.

DESCRIPTION (Figs. 14a, 15). Primary zooids with short peristomes, frontally budded zooids
with long, curved peristomes, directed peripherally, sometimes with a secondary, topo-
graphically proximal, sinus. Peristomal avicularia with curved, acute rostra, placed on the
edge of the peristome. Interzooidal avicularia elongated, only slightly expanded and rounded
terminally, orientated proximally to branch growth. Ovicell with frontal area surrounded by
minute pores.

REMARKS. MacGillivray (1888 : 21 1) noted that the primary orifice was difficult to see, and
described it variously as 'rounded' (1883) or 'with a notch in the lower lip' (1888). In one
illustration (1 885, pi. 2, fig. 6), he actually figured the relationship of the proximal sinus with
a distally placed ovicell, but apparently did not notice that this orientation was in contra-
diction to his description of the ovicell as 'proximal'. However, he was aware of the reversal
of orifice orientation with respect to direction of growth in P. anomala, and compared it with
that of L. hystrix. The relationships of L. mooraboolensis Maplestone (1902 : 25, pi. 2, fig.
18) and L. kitsoni Maplestone (1902 : 25, pi. 2, fig. 19), from the Australian Tertiary, are
difficult to evaluate without specimens. L. mooraboolensis was described from a single,
small, globular specimen of 3-4 zooids and may have been the young astogenetic stage of a
colony of Conescharellina. L. kitsoni was described as ligulate and bilaminar and appears to
have had 'proximal' ovicells with a frontal area. It may be assignable to either L. hystrix or P.
anomala (see below).

Apart from L. kitsoni, L. hystrix is apparently the only species of the family which has a
fossil record. Both Waters (1885) and MacGillivray (1895) noted its occurrence from several
Tertiary Australian localities (see also Brown, 1958).

POECILOPORA MacGillivray
Poecilopora MacGillivray, 1886 : 136.
TYPE SPECIES. P. anomala MacGillivray.

DESCRIPTION. Colonies with compressed, cylindrical, or 'bilaminar' branches, bifurcating in
one plane. Zooids in opposing interdigitating series. Primary orifice sinuate, operculum
extended proximally but not sinuate. Ovicells with a frontal area covered by cuticle, visible
late in ontogeny.

FAMILY LEKYTHOPORIDAE

65

66

P. L. COOK & P. J. HAYWARD

Fig. 14 Silhouettes of opercula of Lekythoporidae and Orthoporidroides, Scale bar = 0-50 mm; (a)
Lekythopora hystrix; (b) Poecilopora cribritheca; (c) Turritigera stellata; (d) T. reticulata; (e) T.
fenestella; (f) Catadysis immersum; (g) Orthoporidra compacta; (h) 0. petiolatus; (i) 0. solida; (j)
Othoporidroides erectus.

Poecilopora anomala MacGillivray

Poecilopora anomala MacGillivray, 1886 : 136, p. 1, fig. 9; 1888 : 21 1 (as P. anomola), pi. 156, figs.

11-13.
^Turritigera stellata Thornely (not Busk), 1924 : 18.

SPECIMENS EXAMINED. BMNH, Port Phillip Heads, 1887.12.10.67, 1888.11.14.103,134,291,
1897.5.1.947.948,949, 1889.7.1.3613. Port Western, South Australia, 1899.7.1.5129.
ChallengerStn. 163 A, Twofold Bay, New South Wales, 220 m, 1899.7.1.4081, 1963.2.12.15.

DESCRIPTION (Figs. 3, 12, 13) The small colonies rarely exceed a height of 10 mm, and are
branched in one plane. The 'bilaminar' growth resembles that of Flabellopora. The primary
orifice has an indistinct sinus proximally. Zooids small, with porcellanous frontal shields
with scattered septulae. Peristomes long at lateral margins of colony, flaring terminally; short
elsewhere, with one acute avicularium. Interzooidal avicularia short, slightly expanded, and
rounded terminally, randomly orientated. Ovicells prominent even late in ontogeny,
protruding beyond the secondary thickening, frontal area surrounded by minute pores.

REMARKS. The early stages of the colony may consist of the ancestrula and one encrusting
zooid generation only ( 1 899.7. 1 .361 3).

MacGillivray (1886 : 137) noted that the reversal of orifice orientation was the explanation
for the apparently 'proximal' ovicells, and postulated that L. hystrix might have a similar
structure.

Thornely's (1924) record of 'Turritigera stellata' was not from an Antarctic locality.
Although most of the stations mentioned in her paper were from Wilkes Land, Antarctica,

FAMILY LEKYTHOPORIDAE 67

the locality 'Maria Island' (42°37'S, 148°E, 73 m) is Tasmanian (see Livingstone, 1928 : 8).
Thornely noted that her colonies were 'bilaminar', that the zooids had tubular peristomes
with only one marginal avicularium, and that the ovicells had a perforated frontal area. The
locality of her specimens indicates that they were either L. hystrix or P. anomala, and the
colony form suggests strongly that they were the latter.

The specimens from Challenger Stn 163 A were not described by Busk (1884). The other
bryozoans from this Station include a large number of cellariiform and cellulariiform species,
which provide substrata for the colonies of P. anomala.

Poecilopora cribritheca (Harmer)

Catadysis cribritheca Harmer, 1957 : 886, pi. 59, figs 9, 12-15, 22, 23.

SPECIMENS EXAMINED. ZMA, LECTOTYPE, Siboga Stn 105, Sulu Archipelago, 275 m.
BMNH, Paralectotype, as above, 1 98 1 .5.6. 1 .

DESCRIPTION (Fig. 14b). Colonies with compressed cylindrical branches. Primary orifice
with a wide sinus. Zooids with porcellanous calcification and scattered frontal septulae.
Peristomial avicularia not protuberant, rounded or subtriangular. Ovicell immersed, but the
frontal area, which has minute, slit-like pores, remains visible.

REMARKS. Harmer (1957) assigned P. cribritheca to Catadysis, which has cylindrical
branches of zooids without long peristomes but with numerous frontal pores. The
porcellanous calcification, compressed branches, long peristomes and exposed frontal area of
the ovicells of P. cribritheca are all characters typical of Poecilopora.

P. cribritheca differs from P. anomala in its slightly larger zooids, and the shape and
distribution of the avicularia. Interzooidal avicularia seem to be absent, but the material is
fragmentary, and incomplete. P. cribritheca is the only species of Lekythoporidae known to
occur in the tropical waters of the Northern hemisphere, but is from fairly deep water
(275m) near the equator.

TURRITIGERA Busk
Turritigera Busk, 1884 : 129.
TYPE SPECIES. T. stellata Busk.

DESCRIPTION. Colonies with cylindrical branches, sometimes becoming reticulate with
anastomoses. Most zooids curved so that their peristomes open on one side of the branch.
Branches dichotomous or formed by frontal ly budded zooids arising in groups at right angles
to the primary branch, in more than one plane. Primary orifice sinuate, but operculum
curved or sinuate proximal ly. Peristomial avicularia sometimes numerous, occurring on the
edge of peristomes, often on raised processes. Ovicell rapidly occluded by secondary
calcification, but often remaining prominent.

Turritigera stellata Busk

Turritigera stellata Busk, 1884 : 130, pi. 24, fig. 1. Waters, 1888 : 22, pi. 1, figs. 22, 25; 1904 : 76, pi. 5,

fig. 3, pi. 8, fig. 13; 1905 : 242, pi. 29, figs. 19, 20. Moyano, 1974 : 1 8, figs. 4, 8, 3 1-34.
not Turritigera stellata Thornely 1924, see P. anomala.

SPECIMENS EXAMINED. BMNH, Challenger Stn. 320, 1100m, 1887.12.9.517.520B,

1899.7.1.3166,3167, 1944.1.8.240,241. Challenger Stn. 142, 275m, 1887.12.9.518,519,

1899.7.1.498,499,500, 1934.2.16.13, 1944.1.8.239,242. Locality? New Zealand,

1936.12.30.155.

MM, Challenger Stn. 320, T38-40; Expedition Antarctique Beige, T 17-37.

DESCRIPTION (Figs. 4, 10, 14c). Colonies reaching a height of 20-30 mm, branched
irregularly. Primary orifice with a small, rounded proximal sinus, operculum with a distal
flange, curved proximally. Zooids with numerous frontal septulae, which become slit-like as

68 P. L- COOK & P. J. HAYWARD

calcification increases. Peristomes long, flaring terminally, with 6ne to seven terminal
avicularia, which are sometimes raised on short processes. Mandibles and rostra acute.
Interzooidal avicularia elongated, slightly expanded and rounded terminally, randomly
orientated, sometimes present on non-zooidal 'basal' side of colony. Ovicell with a small
frontal area which is rapidly obscured by frontal thickening. 18 tentacles (Waters). 'Basal'
thickening extrazooidal, with slit-like pores and minute avicularia.

REMARKS. The South American (Stn. 320), South African (Stn. 142) and Antarctic popu-
lations show some differences in characters. Colonies from Stn. 320 are rather delicate, with
numerous small peristomial avicularia, and interzooidal avicularia on the 'frontal' and 'basal'
sides. There is relatively little extrazooidal thickening (see also Moyano, 1974). Colonies
from Stn. 142 are at a later stage of ontogenetic development and consist of astogenetically
earlier fragments. They appear more robust, with considerable 'basal' and 'frontal'
thickening. The axially directed, proximal, peristomial avicularium is large and the
remainder are reduced in number, usually to two or three. Interzooidal avicularia are absent.
Colonies from Waters's (1904) Antarctic Stations are very robust, and the zooids tend to be
arranged biserially (c.f. T. reticulatd). Waters noted that some colonies possessed
anastomoses (1904, pi. 5, fig. 3a). The zooids have numerous peristomial avicularia, but
interzooidal avicularia are very rare. In contrast, specimens from ?New Zealand (locality
doubtful) are delicate but have one enlarged peristomial avicularium, like those from South
Africa. T. stellata would therefore appear to be a very variable species. The extent and
significance of this variation can only be assessed when complete colonies, showing all
astogenetic and ontogenetic stages can be compared from a wide range of populations.
Recently, for example, d'Hondt( 1981 : 41, pi. 5, figs. 1-3) has described a new, abyssal form,
T. spectabilis, from a depth range of 1661-4402 m, off Uruguay. T. spectabilis resembles
some forms of T. stellata in having only one, large peristomial avicularium. Ovicells were
absent.

The operculum of T. stellata figured by Waters (1904, pi. 5, fig. 3b) was distinctly sinuate,
like the primary orifice. Waters's preparation of opercula (MM T33) is badly preserved and
no sinus is visible. None of the numerous opercula in the Busk preparations from South
African material (BMNH 1 899.7. 1 .498,499) shows any sinus.

Turritigem reticulata sp. nov.

SPECIMENS EXAMINED. BMNH, HOLOTYPE and Paratypes, all parts of one colony, Marion
Island, 1890.4.16.2A.

ETYMOLOGY. Reticulatus (L) — net-like, referring to the colony form.

DESCRIPTION (Figs. 8, 11, 14d, 17, 18). Colonies large, 50mm in height, reticulate, with
anastomosing branches. Zooids very large, alternating and biserial, peristomes short.
Primary orifice very small, with a fairly wide sinus, operculum distinctly sinuate. Secondary
orifice often forming a topographically proximal sinus. Frontal pores numerous, becoming
slit-like. Peristomial avicularia small, one to three are bourne on a stout suboral, topo-
graphically distal process, mandibles short, triangular. Interzooidal avicularia absent.
Ovicells very large, with marginal pores; prominent at first, becoming partially obscured by
extrazooidal thickening, often orientated medially.

REMARKS. The reticulate growth form of T. reticulata is shared by T. fenestella (see below)
and, according to Waters (1904), by some colonies of T. stellata. The large zooidal
dimensions of T. reticulata, together with its very small primary orifice distinguish it from T.
fenestella.

Turritigera fenestella sp. nov.

SPECIMENS EXAMINED. BMNH, HOLOTYPE and Paratypes, all parts of one colony, Marion
Island, 1890.4. 16.2B.

FAMILY LEKYTHOPORIDAE

69

70 P. L. COOK & P. J. HAYWARD

ETYMOLOGY. Fenestella (L) — a little window, referring to the colony form.

DESCRIPTION (Figs. 9, 14e, 16). Colony large, 50mm in height, reticulate, with
anastomosing branches. Zooids irregularly arranged, fairly small, with a primary orifice with
a proximal plate, in which there is a distinct round sinus; operculum also with a distinct
sinus. Frontal pores fairly numerous, rounded, becoming obscured. Peristomial avicularia
one to three, one bourne on a very long, curved suboral process. Mandibles triangular, the
suboral mandible large, rostrum hooked terminally. Ovicells small, smooth, remaining
prominent, although obscured by extrazooidal thickening, often orientated medially.

REMARKS. The similarities in colony form of T.fenestella and T. reticulata, which were from
the same locality, tend to obscure the distinct differences in budding pattern, zooidal size and
orificial characters which exist between them. The zooids of T. fenestella are similar in size
to those of T. stellata, but differ completely in the form of the avicularian processes and
in the shape of the operculum. The three species described above form a complex, which
requires more plentiful material for elucidation.

CATADYSISCanu & Bassler
Catadysis Canu & Bassler, 1927 : 12, 23, 25.
TYPE SPECIES. Myriozoum immersum Busk ( = Schizoporella challengeria Waters).

DESCRIPTION. Colonies with regularly cylindrical branches, zooids opening on all sides.
Zooids with numerous frontal septulae, peristomes not prominent. Primary orifice with a
rounded sinus, operculum curved proximally. Peristomial avicularia small, interzooidal
avicularia absent. Ovicells imperforate.

Catadysis immersum (Busk)

Myriozoum immersum Busk, 1884 : 170, p. 25, fig. 4.

Schizoporella challengeria Waters, 1888 : 30, pi. 2, figs. 25-28.

Catadysis challengeria (Waters) Canu & Bassler, 1927 : 11, Bassler, 1953 : G233.

Catadysis immersum (Busk) Harmer, 1957 : 885. Hayward, 1980 : 712, figs. 5A-D.

SPECIMENS EXAMINED. BMNH, Challenger Stn 320, 1100m, 1887.12.9.683,684,6856,
1899.12.12.18, 1899.7.1.2295, 1934.11.12.57, 1944.1.8.333,334, 1963.2.12.221,247.
'Bruce' Stn 346 (Scottish National Antarctic Expedition) Burdwood Bank, 1936.12.30.297.

DESCRIPTION (Figs. 2, 6, 7, 140- Colonies large, 50x30 mm, branching in all planes. Zooids
with numerous frontal septulae which become immersed by extrazooidal calcification as a
series of slit-like pores. Branches cylindrical, smooth, composed of four series of zooids.
Peristomes never prominent, orifices rapidly obscured by secondary calcification, leaving
rounded pits on the branch surface. Peristomial avicularia small, proximal and distal,
mandibles triangular. Ovicells obscured very early in ontogeny.

REMARKS. Waters (1888) introduced the name 'Schizoporella challengeria' for Busk's species
because he considered that it belonged to the genus Schizoporella and that the combination
was therefore preoccupied by Onchopora immersa Haswell (1 880), which he also assigned to
Schizoporella. O. immersa is referable to the genus Tetraplaria (see Harmer, 1957 : 1055),
and as Catadysis has little in common with Schizoporella, Waters's name 'challengeria' is
unnecessary (see also Harmer, 1957 : 885, and Hayward, 1980).

Most specimens are small and fragmentary, but one from Challenger Stn 320 exceeds
50 mm in width, with numerous branches.

ORTHOPORIDRA Canu & Bassler

Orthopora Waters, 1 904 (preoccupied).
Onhoporidra Canu & Bassler, 1927 : 12, 23, 34.

TYPE SPECIES. Orthopora compacta Waters.

FAMILY LEKYTHOPORIDAE 71

DESCRIPTION. Colony with irregularly cylindrical branches. Primary orifice usually straight
proximally, but opercula may be curved. Zooids with numerous frontal septulae, calcifi-
cation smooth and very thick. Peristomial avicularia often raised on long, hollow mucronate
prominences. Ovicells large, usually imperforate, but with small frontal avicularia.

REMARKS. A large number of specimens has been recorded from many Antarctic and sub-
Antarctic localities. These appear to have the general character of O. compacta as described
below. Small differences in the primary orifice and operculum, however, indicate that
population studies are necessary, and it is possible that several distinct species will be found
to exist, once plentiful material has been examined.

Orthoporidra compacta (Waters)

Orthopora compacta Waters, 1904 : 75, pi. 5, figs. 4a-i. ?Redier, 1965 : 4, 32.

Orthoporidra compacta: Canu & Bassler, 1927 : 12; 1929 : 515, fig. 214. Rogick, 1965 : 406. Moyano,

1978:44.
ICellepora setosa Thornely, 1924 : 17, fig. 5. Livingstone, 1928 : 76, pi. 3, fig. 8, Figs. 18-20. Moyano,

1978:44.
?not Cellepora setosa: Redier, 1965 : 30; 1966 : 2.

SPECIMENS EXAMINED. MM, T10-15, 343, Waters's( 1904) preparations including operculum
(T14) and tentacles (T12), Expedition Antarctique Beige. BMNH, Halley Bay, 1966.3.4.4.
Bahia, Chile, 1971.3.26.29. Scotia Bay, S. Orkneys, 1920.12.1 1.3. McMurdo Sound, 1981.
3.1.1.

DESCRIPTION (Figs. 14g, 21, 22). Colonies large, branched in several planes. Primary orifice
and operculum slightly curved proximally. Zooids large, frontal septulae distinct, marginal
and circum-oral, in two rows across exposed frontal shield. Suboral, topographically distal,
avicularium raised on a long tubular process, mandible rounded or triangular, rostrum bar
with a ligula. Interzooidal avicularia oval or expanded terminally, rostrum bar with a ligula,
usually orientated proximally. Ovicell large with small avicularia on the surface. 24 tentacles
(Waters).

REMARKS. The only type material consists of decalcified sections and preparations etc. The
operculum (T14) differs from that figured by Waters (1904) and resembles that of 0. solida in
being slightly curved proximally, with lateral proximal lacunae which may be muscle
insertions (cf. Waters, 1904).

Thornely (1924) introduced C. setosa for specimens from several Antarctic localities. Her
specimens were redescribed by Livingstone (1928), who noted that neither the primary
orifice nor operculum were sinuate. Both authors figured large ovicells with small pores, or
avicularia?, on their surfaces.

The only available specimen named as O. setosa, which is part of the material described or
listed by Redier (1965, 1966), is a bifurcated branch from Antarctica identified by Dr A. B.
Hastings (Stn 136, Expedition Antarctique Beige, BMNH 1964.8.2.5.). Although similar
to O. compacta in most characters including those of the large, wide, ligulate interzooidal
avicularia, this specimen differs in the dimensions of the orifice and in possessing a sinuate
operculum. The primary orifice is straight proximally but has a small central notch, is longer
than wide and generally far smaller than that of O. compacta. 'C. setosa Redier' is therefore,
in part at least, a separate taxon from C. setosa Thornely and O. compacta.

Additional material is required before all the Antarctic records of these forms can be
reassessed. Redier's (1965) material of O. compacta also requires re-examination. His
record of the Philippine Islands as a locality for the species is a mistaken reference to the
illustrations included by Canu & Bassler (1929) in their paper on Philippine bryozoans.

Orthoporidra solida (Busk)
Cellepora solida Busk, 1884 : 200, pi. 29, fig. 12. Waters, 1904 : 76, pi. 5, fig. 5.

72

P. L. COOK & P. J. HAYWARD

FAMILY LEKYTHOPORIDAE 73

SPECIMENS EXAMINED. BMNH, LECTOTYPE (chosen here), Challenger Stn 160, 4758m.
1899.7.1.3525, the specimen figured by Busk, 1884. Paralectotypes 1887.12.9.775-777;
1899.7.1.3526.

DESCRIPTION (Figs. 5, 14i, 19, 20). Orthoporidra with very large zooids, with numerous,
scattered frontal septulae. Primary orifice and operculum curved proximally. Peristome not
very prominent. Peristomial avicularia proximal and paired, lateral, raised on blunt mucros,
mandibles rounded, rostra subtriangular. Interzooidal avicularia oval, large. Ovicells large,
with small avicularia and scattered pores.

REMARKS. The colonies form branched cylindrical masses 5-10 mm in height. Secondary
thickening is less developed than in O. compacta, and consists of numerous kenozooids as
well as extrazooidal tissue. The avicularian processes are short and blunt, quite unlike those
of O. compacta.

Busk (1884: 200, pi. 29, fig. 12a) described the large interzooidal avicularia (as
'operculum' in the explanation of Plate 29), and Waters (1904, pi. 5, Fig. 5) also illustrated
an operculum. The preparations in the BMNH show that the opercula are far larger than the
dimensions indicated by Waters's figure.

Orthoporidra petiolata (Waters)

Celleporapetiolata Waters 1904 : 76 (nom. nud.); 1905 : 241, pi. 29, figs. 19, 20.
Catadysis petiolata: Hayward, 1980 : 714, Figs 5 E-H.

SPECIMENS EXAMINED. MM, T2-T6, Expedition Antarctique Beige. BMNH, Houtjes Bay, S.
Africa, 1936.12.30.300c. Stn 346, Scottish National Antarctic Expedition, Burdwood Bank,
1981.3.1.12.

DESCRIPTION (Fig. 14h). Colony arising from a minute thickened base of kenozooids,
encrusting erect bryozoa, 10-15 mm in height. Zooids with marginal and circum-oral
septulae. Primary orifice and operculum with a small sinus. Suboral peristomial avicularium
raised on long processes, mandible small, rounded or elongated and triangular. Ovicells
small, prominent at first, with frontal pores and small avicularia. 14 tentacles (Waters).

REMARKS. Waters (1904) mentioned that a species which he proposed to name 'C. petiolata'
was similar in character to O. compacta. In his formal introduction (1905) he gave no further
details, and figured only the operculum and mandibles. Waters's preparation (T5) shows an
operculum with a small rounded sinus.

The generic assignment of O. petiolata is arbitrary. The orificial characters are similar to
those of some species assigned here to Turritigera (e.g. T. fenestella), and to Catadysis
immersum (see Hayward, 1980). The budding pattern of the zooids, and the distribution of
the frontal septulae resemble those of Orthoporidra compacta.

Comparison of the Lekythoporidae with the genus Orthoporidroides Moyano

Frontally budded zooids with cryptocystidean frontal shields, which form massive or
cylindrical, erect branches are typical of the Family Celleporinidae Harmer. Genera such as
Turbicellepora (see Hayward, 1978) and Celleporina (see Harmer, 1957) also have oral
avicularia derived from circum-oral frontal septulae, and often have elongated peristomes
with terminal avicularia. Large, interzooidal (vicarious) frontally budded avicularia, and
ovicells with porous frontal areas (tabulae) are also typical of these genera. The genera
included in the Lekythoporidae are therefore distinguished from those in the Celleporinidae
principally by their 'reversed' frontal budding pattern. The apparent close similarity
between the Families is illustrated by Orthoporidroides erectus (Waters).

CELLEPORINIDAE Harmer

Celleporinidae Harmer, 1957 : 899.

74 P. L. COOK & P. J. HAYWARD

ORTHOPORIDROIDES Moyano
Orthoporidroides Moyano, 1 974 : 20 1 . 1 98 1 .
TYPE SPECIES. Cellepora armata var. erecta Waters.

Orthoporidroides erectus (Waters)

Cellepora armata var. erecta Waters, 1888: 36. pi. 3, figs 4, 4 1 , 43.
Orthoporidroides erectus : Moyano, 1 974 : 2 1 , Figs. 6, 35^40.

SPECIMENS EXAMINED. BMNH, Challenger Stn 308, 320m, 1889.12.12.7. Challenger Stn
320. 1100m. 1888.3.14.3, 1899.7.1.3584, 3587, 3592, 3602, 3606; 1934.11.12.5.
Shackleton-Rowett Exped., ?S. Orkney, 1923.12.1.39,48.

DESCRIPTION (Fig. 14j). Colonies erect, branching, arising from a small, encrusting base.
Zooids budded frontally and distally, primary orifice with a shallow sinus directed towards
the outside of the branch, operculum produced into a triangular sinus. Avicularia suboral,
terminal on a long, proximally produced peristome, mandible acute. Ovicell hyperstomial,
with a small, frontal slit.

REMARKS. Moyano (1974) has illustrated the morphology of O. erectus fully and compared it
with that of Turritigera stellata. Moyano also noted similarities between O. erectus and
Cellepora petiolata Waters (1905), see above. The opercula of O. erectus figured by Moyano
resemble those illustrated by Waters, but differ slightly from those of the specimens from
Challenger Stn 320, which are very variable (Fig. 14j). These specimens were not described
by Busk ( 1 884) or Waters (1888).

Discussion

Study of the relationships of the ovicell in the Lekythoporidae illustrates the importance of
analysis of the astogenetic structure of colonies, as well as the ontogeny of their member
zooids. It is interesting that since the ovicells of these genera were first described as
'proximal' in position, this monothetic 'character' has been accepted almost without
question, and has even been modified (e.g. Harmer, 1957 : 884) in an attempt to include
other taxa in the family, a frequent occurrence in bryozoan taxonomy (see Boardman et al,
1969). The explanation of the unusual relationships of the ovicell has awaited first, the
demonstration of frontal budding by Banta (1972) and second, the realisation that in both
erect and rooted colonies, this type of astogeny may include reversal of zooidal orientation
with respect to the direction of colony growth (see Cook & Lagaaij, 1976, and Cook &
Chimonides, 1981).

The similarities in several characters among the genera assigned to the Lekythoporidae
and Celleporinidae suggest a close relationship between the families.

It is possible that parallel complexes exist, each containing groups of species with totally
different budding patterns, but with similar zooidal characters and even with similar
resultant colony forms. Orthoporidroides robusta Moyano (1981 : 182, Figs 1-7), for
example, resembles some other, unnamed Recent specimens from the Falkland Islands
(BMNH, 1981.3.1.6,9,10) in illustrating how the type of budding pattern found in O. erectus
may have been transformed into the type found in the Lekythoporidae. Colonies of both
species are large, maximum height 50-60 mm, and profusely branched, with a range of zooid
orientations. In Orthoporidroides sp. there are several sequential series of outwardly directed
interzooidal, frontally budded zooids. The more centrally placed zooids of branches, which
are those budded earlier in the astogenetic sequence, tend to have 'reversed' orientation of
orifices, while the more numerous peripheral zooids have a 'normal' orientation. Some
intermediate orientations, and ovicells in a 'proximal', lateral and distal topographical
position occur in both O. robusta and the unnamed species. This differs from O. robusta in

FAMILY LEKYTHOPORIDAE 75

its orifice shape, which has a very long, narrow sinus, and the number of peripheral zooids in
its branches.

At present it is difficult to determine whether or not astogeny and structure reflect closer
genetic links than those of zooidal morphology. In order to trace possible changes in, for
example, zooidal orientation and astogeny with time, a range of fossil specimens is necessary.
Colonies of Lekythoporidae are robust enough to be preserved in fossil sediments;
unfortunately only L. hystrix is unequivocally known to have a fossil record. Lack of records
of other species, both in Recent and fossil deposits, is probably caused by difficulties of
recognition, due to the small size of the colonies, and rapid thickening of extrazooidal
calcification, which obscures the zooidal features. Recent species tend to inhabit the deeper
shelf waters, and until more collections from these regions, particularly from high latitudes
in the Southern hemisphere become available, detailed analysis of colonies is not possible.

Although the characters of the type-species of the five genera of Lekythoporidae are
distinct, several of the other species described here show intermediate sets of correlations
which make generic assignment somewhat arbitrary.

The 'easily recognised' character of the 'proximal ovicell' has to a great extent obscured
analysis of other structures in the past. The fragmentary nature of most of the type material
requires amplification by well preserved, complete colonies before the range of astogenetic,
ontogenetic, polymorphic and microenvironmental variation of the taxa can be analysed and
described.

Acknowledgements

We are grateful to Dr M. V. Hounsome (Manchester Museum), Dr B. Smith (National
Museum of Victoria) and Dr S. van der Spoel (Zoological Museum, Amsterdam) for the
opportunity to examine specimens, and to Mr P. J. Chimonides (British Museum, Natural
History) for discussions and scanning electron microscopy.

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Manuscript accepted for publication 10 October 1982

A new species of Arthroleptis (Anura: Ranidae)
from the West Usambara Mountains, Tanzania

Alice G. C. Grandison

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

The herpetology of the Usambara Mountains, Tanga Province, Tanzania is primarily based
on the extensive collections made by Loveridge (Barbour & Loveridge, 1928). He concen-
trated his efforts on the more accessible eastern half of the Usambaras, especially around
Amani (5°06'S: 38°37'E) at an elevation of around 923 m and appears to have been the
only person who collected amphibians in the Mt. Lutindi outlier (Hindu of Moreau, 1935),
the only part of East Usambaras that rises above 1219 m and has a patch of highland and
cedar forest on its peak (1 506 m) similar to that which, until comparatively recently, covered
a vast area of West Usambaras (Moreau, 1935). Both the East and West Usambaras consist of
high mountains with dramatically steep cliffs and ridges but the western peaks and plateaux
are all above 1219m, their highest ridges reach 2286 m and the rainfall is considerably less.
The eastern and western divisions of the mountains are divided by the hot, dry trench-like
Lwengera Valley which is nearly 914m deep. The forests of the Highland Zone of the West
Usambaras, as defined by Moreau (1935) in his excellent account of the topography, ecology
and climate of these ancient mountains, remained virtually untouched by herpetologists
until the last decade when collecting has been carried out by the writer and other zoologists,
notably at ca 1 530 m, 37-8 km NE Soni above Mombo at the Mazumbai Forest Reserve and
tea plantation, 4°48'S: 38°29'E. This Forest Reserve consists of over 600 acres given to the
University of Dar es Salaam by the owner of the tea plantation, Mr John Tanner. The most
profitable collecting by the writer was done at 1530 m where an irrigation channel about
1-25 m wide follows the contour from a waterfall and river in forest passing through the
forest to emerge in part of the plantation. The stream of approximately 140 mm depth had a
slack to moderate current with a fine gravel and mud base. On its right bank it was overhung
by forest trees while its left bank had a muddy track of sodden leaves plus debris backed by
balsam and wild banana. The forest trees included Ocotea usambarensis and Newtonia.
Average annual rainfall measured over a seven year period (1954/60), was 1041 mm. From a
stone and debris in the stream and from the stream margin five Arthroleptis were obtained.
These together with specimens found later by other collectors, at Mazumbai and additional
localities in the West Usambaras between 1450m and 2134m, surpass in adult size all
known species of Arthroleptis. In colour pattern and some proportions they resemble A.
affinis Ahl, and to a much lesser extent A. adolfifriederici Nieden. A comparison of them
with long series of affinis obtained mainly at the type locality, Amani in the East Usambaras,
and with the type and paratype of Ruandan adolfifriederici suggests that an undescribed
species of Arthroleptis exists in the West Usambara Mts.

The new species is named after Mr John Tanner in recognition of the generosity and
hospitality afforded the collectors and other zoologists and his keen interest in the natural
history of Mazumbai.

Abbreviations

BM(NH)British Museum (Natural History), London

FMNH Field Museum of Natural History, Chicago

R Universitetets Zoologiske Museum, Copenhagen

ZMB Zoologisches Museum der Humboldt-Universitat Berlin

Bull. Br. Mm. nat. Hist. (Zool.) 45 (2): 77-84 Issued 28 July 1 983

77

78 A. G. C. GRANDISON

Arthroleptis tanneri sp. nov.
Arthroleptis adolftfriederici: Barbour& Loveridge 1928 (part)

HOLOTYPE. Adult female BM(NH) 1974.59, collected by A. G. C. Grandison and F. V.
Slade at Mazumbai 4°48'S 38°29'E West Usambara Mountains, Tanzania, ca. 1530m
elevation, 1000 hrs, 8 October, 1973.

PARATYPES. Same locality data as for holotype: BM(NH) 1974.60 (stained and cleared
preparation), BM(NH) 1974.61-63, 2245-2300 hrs., 8th October, 1973, BM(NH) 1974.200
16.30 hrs. 1 1th October 1973,m BM(NH) 1982.525-36, R 773 1-35, R 7736-68, R 7797-98.
Phillipshof 4°45'S 38°18'E (4 mis N of Lushoto), W. Usambara Mts: MCZ 13166. Shume-
Magamba Forest Reserve, 4°40'S 38°15'E, W. Usambara Mts, 2134m elevation, 11-16
February 1981: BM(NH) 1982.537-40. Balangai Reserve, 4°56'S 38°37'E, W. Usambara
Mts, 1450 m elevation, May 1981: BM(NH) 1982.541.

DIAGNOSIS. A large Arthroleptis related to A. adolfifriederici, separable from that species
and all other species of the genus by exceptional size (males 33-7 ±3-3: females
50-7 mm ±4-9) and stout build. Further distinguished from the Usambara A. affinis by its
shorter tibia, more widely set nares, slimmer, less pointed terminal phalanges, absence of
supernumerary tubercles under the metatarsals, chin spinules but no denticulations along
the second and third fingers in the adult male and by minor differences in the temporal
colour pattern.

DESCRIPTION OF HOLOTYPE. Gravid female 54-0 mm SVL. Head slightly broader than long,
its width 44% SVL. Length of snout from anterior border of eye to snout tip 35% of head
width. Distance between the nares 11% of SVL. Canthus rostralis sharp, loreal region
oblique, lips moderately flared. Interorbital region wider than upper eyelid, subequal to
internarial distance. Tympanum clearly visible, the diameter of the tympanic annulus
slightly greater than half the internarial distance. Snout rounded in dorsal view. Body stout.
Terminal phalanges of fingers bluntly rounded, not or barely wider than distal joint, without
circummarginal grooves. Toes long and slender, their terminal phalanges bluntly rounded
with circummarginal grooves, the maximum width of the terminal phalanx of the 3rd toe
about half the tympanic diameter and only 1/3 wider than the midpoint of the penultimate
phalanx. No outer metatarsal tubercle or tarsal fold. Inner metatarsal tubercle broadly oval,
its length 1/2 the distance from the tip of the 1st toe to the distal margin of the tubercle.
Tibial length less than length of foot (measured from tip of 4th toe to proximal margin of
inner metatarsal tubercle) 49% of SVL. No supernumerary tubercles proximal to basal
subarticular tubercles of the toes. No toe web. Skin on the head and back smooth with a
slight ridge extending along the canthus and lateral edge of the upper eyelid to form a curved
dorsolateral line on the back.

COLOUR IN ALCOHOL. Dorsum mid brown with a darker rather obscure cruciform area
extending from the interorbital area at the level of the middle of the eyelids to the middle of
the shoulders, and bordered posteriorly by a broad pale chevron-shaped zone that is of the
same intensity of brown as the top of the snout and upper eyelids. Behind the somewhat
indistinct chevron is a darker area that reaches to the vent. The tip of the snout has a short
light vertical streak. The lores are darker than the top of the snout and a prominent dark
brown band passes from the posterior corner of the upper eyelid through the upper third of
the tympanum and curves round the posterior rim of the tympanum ending at a point above
the arm insertion. The under surfaces of the tarsus and metatarsus are dark brown, except for
their outer edges which have a pale line from the heel along the external margin of the 5th
toe to its tip. The legs and forelimbs are cross banded dark and lighter brown. The back of
the thighs are mottled with pale grey. The middle of the belly is immaculate dirty white, the
gular skin and chest pale fawn with creamy white mottling. The lower lip is medium brown,
spotted or barred with white. Undersurfaces of the hind limbs dirty white with a faint greyish
network. Undersurfaces of toes rather pale, the subarticular and metatarsal tubercles cream.
Undersurfaces of hand uniform dirty cream.

NEW SPECIES OF ARTHROLEPTIS 79

HABITAT. The holotype and some of the British Museum paratypes were associated with
Rana angolensis Bocage and Phrynobatrachus kreffti Boulenger. One adult female was on a
stone jutting out of the stream, others in holes in rotten logs or under logs or on leaf litter,
while the holotype and two other paratypes were among floating plant debris and twigs at the
stream margin. The Copenhagen paratypes were reported as being on the forest floor about
1-6 km from water near to where the forest is replaced by heather on Mt. Sagara above
Mazumbai.

COLOUR IN LIFE. The general dorsal colouration of the British Museum series was
claret-brown with the chain of vertebral markings from the interorbital region to the lumbar
region of a darker shade. The hands and feet were pinkish, especially their undersurfaces.
Upper half of iris was pale gold.

VARIATION IN THE PARATYPES. There is a limited amount of variation in the distinctness of
the dorsal pattern, the dark area over the occiput and the pale chevron immediately posterior
to it being more clearly defined in younger individuals. Furthermore in the long series of
smaller specimens (R 7736-68) some tend to have dark specks at the posterior margin of the
chevron and small dark brown spots on flanks and temporal region. The cruciform area is
replaced in some of the paratypes by a dark inverted triangular zone from the posterior limit
of which two dark oblique bands extend towards the flanks. The undersurface of the head
varies from grey mottled white to an almost uniform brown which in a sexually mature adult
male has an even darker band laterally. The dorsal skin of the juvenile BM(NH) 1974.200
has small scattered warts.

SECONDARY SEXUAL CHARACTERS. As already indicated in the species diagnosis, male tanneri
are smaller than females. Only four males are available and all have unpigmented testes. The
smallest R7735 (30-0 mm SVL), is the only one that has spinules on the chin, and a dark
brown almost black area along the lower jaw medial to the barred lip. No other secondary
sexual differences are apparent. All the gravid females have large unpigmented eggs with a
maximum diameter of 3-5 mm.

COMPARISON. Although A. tanneri attains a larger size than A. affinis Ahl, the taxon
described from Amani, East Usambaras and reinstated by Skelton-Bougeois (1961), there is
a strong superficial resemblance. However, statistically significant differences at the level
P< 0-001 using two sample t-test exist in the following percentage ratios of samples of the
two species.

A. tanneri A. affinis

gravid 9 9 ^99, subadults gravid 9 9 cf^99, subadults

Head width n6 n!9 nlO n32

SVL x 44-17 x 43-34 x 42-0 x 42-01

S 1-17 S 1-86 S 1-56 S 1-29

Head width n 6 n 19 n 11 n 32

Tibia length x 88-5 x 84-31 x 77-2 x 75-36

S 5-0 S 6-09 S 2-40 S 3-38

Tibia n 6 n 19 n 13 n 32

SVL x 49-8 x 51-83 x 54-8 x 55-78

S 3-06 S 3-36 S 3-10 S 2-60

Internarial distance n 6 nil

SVL x 11-15 x 12-62

S 0-52 S 0-51

80

A. G. C. GRANDISON

No statistically significant differences in the percentage ratios of the internarial distance to
SVL were found in samples of tanneri and affinis that included halfgrown individuals as well
as adults of both sexes. Breeding males of tanneri and affinis may be distinguished by their
different secondary sexual characters. None of the sexually mature male affinis has chin
spinules but all have a row of denticulations along the inner border of the 2nd and 3rd fingers
and the 3rd finger is elongated. Of the two males obtained by Loveridge at Phillipshof West
Usambaras and identified by Barbour & Loveridge (1928) at adolfifriederici Nieden, the
larger individual (37-4 mm SVL) MCZ 13166 has neither chin spinules nor denticulated
fingers but agrees in proportions and in other characteristics with tanneri while the smaller
specimen (29- 1 mm SVL ) MCZ 1 3 1 67 is a typical affinis and has finger denticulations. Like
all the males and females of affinis from the East Usambaras, both those from the Highland
Zone (Mt. Lutindi) and from the Intermediate Forest of Amani, the Phillipshof example
MCZ 13167 bears a prominent supernumerary tubercle under the distal portion of the 2nd
and 3rd metatarsals and additional but smaller tubercles scattered over the sole of the foot. In
some individuals of affinis a small tubercle is also present at the base of the first toe between
the inner metatarsal tubercle and the subarticular tubercle (Fig. 2). Ahl (1939) in his type
descriptions of affinis and of the junior synonym A. schoenbecki, both of which were
obtained at Amani, mentions the supernumerary tubercles which bedeck the soles of the feet
in the type specimens but makes no reference to the more distal tubercles under the 2nd and
3rd metatarsals. Examination of the type material in the present study has confirmed their
presence.

Supernumerary tubercles under the metatarsals are absent from every individual of
tanneri, as well as from the holotype and paratype of A. adolfifriederici from Rugege and
Bugoya Forests Ruanda (Fig. 2). A third example also collected at Rugege and referred to
adolfifriederici by Nieden (1912) also lacks these tubercles. However adolfifriederici differs
from A. tanneri in its smaller size, the adult female holotype and paratype being 41-1 mm

Fig. 1 The Usambara Mountains, NE Tanzania. Land over 1000m is indicated by cross

hatching.

NEW SPECIES OF ARTHROLEPTIS

81

Dorsal view and undersurface of the foot of: Arthroleptis tanneri sp. nov. (A & C);
Arthroleptis af finis (B & D).

and 43-8 mm SVL, as well as in its marbled dorsal pattern (Nieden, 1912) and further differs
from both tanneri and affinis in its much narrower head and shorter internarial distance and
tibia. Percentage ratios in the holotype and paratype of adolfifriederici of head
width/SVL are 35-0 and 33-1, of internarial distance/SVL 10-70 and 10-27, of head
width/tibia length 61-0 and 55-34. A. tanneri seems to be further distinguished from either

82 A. G. C. GRANDISON

adolfifriederici or affinis by its terminal phalanges being bluntly rounded and barely wider
than the distal joint. In the Ruanda adolfifriederici and in the East Usambara series of affinis
they tend to be sharply pointed and are decidedly wider than the proximal phalanges and
joints.

In colour pattern tanneri and affinis are rather similar but whereas in tanneri the ventral
surfaces of the toes are cream, in affinis the phalanges and the soles of the feet are dark brown
and the cream subarticular tubercles are in sharp contrast. Also in affinis the supratympanic
stripe generally curves round the top of the tympanic annulus and does not pass across the
tympanic membrane.

Osteology

The stained and cleared adult female paratype of tanneri was compared with a similarly
prepared Amani adult of affinis. Osteological differences that are interpreted as being of
taxonomic significance are limited to the skull, in particular the shape, extent of the
sphenethmoid including the degree of its dorsal exposure, the medial separation of the nasals
and the shape of the anterior borders of the frontoparietals.

In tanneri the sphenethmoid is a short squat bone, sharply concave posteriorly and with a
straight anterior border that fails to reach the palatines except for their medial posterior tips.
It has no dorsal exposure, being completely covered by the frontoparietals. The
sphenethmoid in affinis not only projects much farther forward to extend well beyond the
palatines and to the level of the anterior edges of the choanae, but its anterior and posterior
borders are of a different shape from those of tanneri, being deeply convex and straight
respectively. Furthermore the sphenethmoid has a crescentic dorsal exposure.

In both taxa the palatines are robust and contact the maxillae broadly but fail to reach the
pterygoids.

Not only are the medial borders of the nasals more convex and wider apart in affinis but
there is a greater separation between the nasals and the frontoparietals; moreover the
anterior borders of the frontoparietals project forwards instead of as in tanneri being straight.

It is considered unlikely that these differences are attributable to individual variation in
view of the author having discovered in an extensive analysis of bufonid skulls that there is
virtually no intraspecific variation in adults in the shape and size of the sphenethmoid,
palatines, nasals and frontoparietals.

Distribution of A. tanneri and its relationship with /. affinis

In the eastern half of the Usambara Mts A. affinis is known from Amani, 923 m, and from
1219 m on Mt. Lutindi (Hundu). In the neighbourhood of Amani during the dry season the
writer found affinis to be abundant among leaf litter, in cracks in the impacted earth of road
cuttings and in mounds of earth, sometimes in association with Callulina kreffti Nieden and
Nectophrynoides tornieri Roux. Loveridge obtained 192 examples from leaf strewn paths at
Amani (Barbour & Loveridge, 1928). At comparable elevations in the western half of the
Usambaras no herpetological fieldwork has been undertaken below 1219m and only one
Arthroleptis has been taken at 1219m, on the Ambangulu Estate, Korogwe District,
5°05'S, 38°26'E. The single individual, a juvenile of 20-0 mm displays features of both
tanneri and affinis. It resembles affinis in having supernumerary tubercles under its
metatarsals but its proportions correspond more closely to those of tanneri, its percentage
ratios being Head width/SVL 45-0, Head width/Tibia 90-0 and Tibia/SVL 50-0. The
existence of this aberrant individual raises the question of whether tanneri and affinis coexist
at Ambangulu and whether the 'mixed' characters of this juvenile can be attributed to
hybridisation. Further material from this and other localities at lower elevations in the West
Usambara Mts is awaited with interest.
Evidence of sympatry of A. tanneri and A. affinis rests on the two adult males collected by

NEW SPECIES OF ARTHROLEPTIS 83

Loveridge in 1926 during ten days spent at Phillipshof in the West Usambaras. Barbour &
Loveridge (1928) quote the elevation of Phillipshof as approximately 5500ft (1676m),
which accords with information given to the author by the Royal Geographical Society
based on the 1916 War Office maps of Tanganyika Territory and the Lushoto sheet
1 : 250 000 which place Phillipshof four miles north of Lushoto, 4°45'S 38°18'E, elevation
1524-1829 m. Barbour & Loveridge (1928) describe Phillipshof as 'rolling downs of grazing
land, marshy swamps and slow-flowing streams in the bottoms, scattered patches of
rain-forest on the uplands, with vast stretches of forest nearby on either side of the Malindi
road'. According to Moreau's (1935) zonation of the Usambaras Phillipshof lies in the same
Highland Zone as Mazumbai. It is not known whether the Phillipshof examples of tanneri
and affinis were found in similar niches and closely associated with each other or whether
they occupied quite different biotopes; no precise field data accompany the specimens.
Except for the Phillipshof record of sympatry and the unique characters of the Ambangulu
juvenile the available material points to tanneri and affinis being separated geographically
and by altitude with A. tanneri confined to the West Usambara Mts in the Highland forest
zone of Moreau (1935) and A. affinis occurring at the lower elevations of the Intermediate
forest zone of the East Usambara Mts.

Additional material used in study

Arthroleptis adolfifriederici Nieden

Bugoya Forest, Ruanda: ZMB 2 1 787 (Paratype).

Rugege Forest, Ruanda: ZMB 25287, FMNH 73836 (ZMB 2 1 789) (Holotype).

Arthroleptis affinis Ahl

Amani 5°06 ' S 38°37 ' E, E. Usambara Mts, Tanzania 923 m: BM(NH) 1 974. 1 77 (stained and
cleared preparation), BM(NH) 1974.170-176, BM(NH) 1974.178-185, BM(NH)
1974.186-195, MCZ 88131-278, MCZ 13153-60, FMNH 73836, ZMB 25289 (holotype of
A. schoenebecki Ahl), ZMB 23093 (holotype of A. affinis), R 7797-98, R 77 1 00.
Mt. Lutindi, E. Usambara Mts, Tanzania: MCZ 13161-65.
Phillipshof, W. Usambara Mts, Tanzania: MCZ 13167.

Acknowledgements

Most cordial thanks are extended to Dr K. M. Howell, University of Dares Salaam and to Dr
A. Schiotz, Director, Danmarks Akvarium, Copenhagen for making available collections
from the Usambaras and generously allowing me to describe the new species. Mr H. Marx
(Field Museum of Natural History, Chicago), Dr G. Peters (Zoologisches Museum
Humboldt-Univ. Berlin) and Dr E. E. Williams (Museum of Comparative Zoology, Harvard)
also kindly lent material in their care.

Dr I. Walker (Imperial College of Science, London) and Dr N. Jago (Centre for
Overseas Pest Research, London) helped in formulating plans for my Tanzanian
visit. Dr R. Laurent (Institute Miguel Lillo, Tucuman) and my colleagues Dr G.
Boxshall and Dr M. Hills gave freely of their time in discussing morphometric
analyses, Dr Hills being especially helpful with the statistical interpretation of data.

I wish also to record my thanks to members of the British Council, Staff of the
Zoology Department, University of Dar es Salaam, and the Tanzanian Government
for facilitating my research.

It is difficult to express adequately my appreciation of Mr John Tanner's kindness,
generosity and hospitality towards members of the team while staying at his home in

84 A. G. C. GRANDISON

Mazumbai. He ensured that our visit was scientifically worthwhile and enjoyable,
and I thank him most sincerely.

Dr S. N. Stuart's help with field data and literature is also gratefully acknowledged.

Field work was supported financially by the Trustees of the British Museum
(Natural History).

References

Ahl, E. 1939. Beschreibung neuer afrikanischer Frosche der Gattung Arthroleptis. Sber. Ges. naturf.

FreundeBeri 303-310.
Barbour, T. & Loveridge, A. 1928. A comparative study of the herpetological faunae of the

Uluguru and Usambara Mountains, Tanganyika Territory with descriptions of new species. Mem.

Mus. comp. Zool. Harv.. 50: 87-265.
Moreau, R. E. 1935. A synecological study of Usambara, Tanganyika Territory, with particular

reference to birds. J. Ecol. 23: 1-43.
Nieden, F. 1910. Verzeichnis der bei Amani in Deutschostafrika vorkommen den Reptilien und

Amphibien. Sber. Ges. naturf. Freunde Be rl: 44 1-4 52.
Nieden, F. 1912. Amphibia. Wiss. Ergebn. dt. Zent Afr. Exped. 4: 165-195.
Skelton-Bourgeois, M. 1961. Reptiles et batraciens d'Afrique orientale. Rev. Zool. Bot. afr. 63:

309-338.

Manuscript accepted for publication 15 July 1982

The distribution, behavioural ecology and breeding
strategy of the Pygmy Toad, Mertensophryne
micranotis (Lov.)

Alice G. C. Grandison

Zoology Department, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Sanda Ashe

P.O. Box 3, Watamu, Kenya

Introduction

Sexually mature males of the East African pygmy toad, Mertensophryne micranotis
(Loveridge) have been reported as having series of small spines around the rim of the cloaca
and at the entrance to the cloacal tube (Grandison, 1980). Development of these spines
correlates with development of the cluster of heavily keratinised thumb spines and their
function was interpreted as an adaptation for holding the vents of the male and female
securely together during mating to ensure direct transfer of sperm to the small clutch of large
eggs produced by the female. Spines in the area of an anuran's vent were first reported by
Anderson (1871). He described a large zone of curved cornified papillae in Rana gammi
Anderson, a species currently synonymised with Rana (Paa) sikkimensis Jerdon. Dubois
(1976) figured the spines in sikkimensis and suggested that cornified spinules occur also in
Rana (Paa) delacouri Angel. The life histories of neither of these Nepalese species is known.
In order to test the theory that had been postulated of internal fertilisation in
Mertensophryne micranotis fieldwork and a captive breeding programme were recently
undertaken in Kenya. New information on the species' occurrence in Kenya, its habits and
breeding behaviour is presented here.

Range and ecological preferences

M. micranotis is known to occur in Tanzania, Zanzibar and Kenya. Few examples have
been collected, perhaps because such a small-sized toad (sexual maturity is attained at
16 mm in males) is difficult to detect in leaf litter. Although the type and paratype found at
Kilosa, Tanzania and a single individual from the Uluguru Mts. were said by Loveridge
(1925) and Barbour & Loveridge (1928) respectively not to have been found in forest, else-
where the species seems to be restricted to forest or recently cut down forest. In Kenya the
species occurs both along the coast from Gede in the north (3°1 1 'S: 40° 1 'E) to Shimoni in
the south (4°38'S: 39°23'E), in the Shimba Hills National Park in the Coast Range (4°16'S:
39°22'E) and Mrima Hill (4°29'S: 39°16'E). Until comparatively recently the Coast
Province of Kenya was a near continuous forested belt overlying the approximately three
kilometre wide Pleistocene coral reef and the Pliocene and Pleistocene sandy soils adjacent
to it farther inland. Today little true forest remains in the Coast Province and very few
remnants extend to more than 50 acres. Agriculture, notably plantations of sugar, coconut,
cashew nut, maize and sisal combined with the rapid growth of Kenya's tourist industry have
taken their toll of the coastal forests. Two of the small remaining pockets are contained in the

Bull. Br. Mus. nat. Hist. (Zool.) 45 (2): 85-93 Issued 28 July 1 983

85

86

A. G. C. GRANDISON & S. ASHE

3 9

Fig. 1 Known distribution of Mertensophryne micranotis (Loveridge) in Kenya. Records

indicated by an inverted triangle.

MER TENSOPHR YNE MICRA NO TIS 87

National Monuments of Gede and Mtwana, in which are preserved the ruins of thirteenth
century Arab-African towns. Diani forest (sometimes also referred to as Jadini forest) con-
sists of 50 acres of largely untouched forest similar in composition to Gede forest; it is
privately owned. Other small forest relics encircle cleared areas that until the nineteenth
century were settlements, each housing up to 2000 of the Mijikenda peoples but which today
contain only a few houses occupied by elders of the clans and graves of their ancestors. Most
of these kayas, as these sacred villages are called, were situated along the ridge of the Coast
Range from the Shimba Hills northwards to Kilifi and at vantage points to resist incursions
from the north by the Galla farmers. By the seventeenth century the most southerly of the
Mijikenda, the Digo, vacated their Shimba kaya and established minor kayas along the
coastal plain (Spear, 1978). At one such sub-kaya, Kinondo, and at the Kombeni kaya M.
micranotis has been found. On account of their religious significance and present day use for
rituals and processions the kaya forests are provided with a measure of protection which may
safeguard their populations of micranotis.

In the southern extremity of the Coast Province two densely forested hills, Jombo and
Mrima, jut up spectacularly from the surrounding flat plains (Britton et al., 1980). No
herpetological survey has apparently ever been made of Jombo (476 m) but the senior author
made two short visits to nearby Mrima which rises to 299 m. The very deeply weathered
volcanic plug of Mrima has been prospected since the early 1950s for its rich deposits of
manganese and niobium. Its rocks weather extremely rapidly and up to 236 m of terra rosa
cover the carbonatite plug. It is the weathered material that has been mined and the
abandoned and completely unprotected mineshafts are a serious hazard to contend with
when scouring the upper reaches of the forest for the anuran denizens of the leaf litter. A
circular, flat-topped pile of moist leaves which contained duiker droppings and may have
been a communal duiker latrine on to which some animals, perhaps ants, had piled leaves
and other debris yielded juvenile M. micranotis. The pile was on a trail and was the only
damp spot found; no rain appeared to have fallen for some considerable time and no water-
filled treeholes were evident. On the bank of a path and among small pieces of weathered
carbonatite and terra rosa and harmonising perfectly in colouration another juvenile
micranotis was obtained. In these forests there are no surface ponds or streams and even after
heavy and prolonged rain the water probably rapidly disappears through the red earth and
litter that overly and form pockets in the labyrinth of holes in the coral rag of the narrow
coastal strip. On the sandy soils farther inland the rains likewise drain rapidly. The only
standing water that becomes available with the highly seasonal rains are ephemeral
reservoirs in treeholes formed by broken off limbs, in fissures in tree buttresses and in live
and fallen tree trunks, as well as in abandoned shells of the large land snail, Achatina, and in
the occasional discarded can on the forest floor. Consequently the only anurans that can
survive in such forests need to have a highly specialised breeding strategy, such as direct
development that completely omits the larval stage or one that produces a terrestrial larva,
or need to be opportunistic breeders capable of taking advantage of the seasonal water
pockets for egg laying sites. As the water holes are small and subject to rapid desiccation
small clutches capable of an accelerated rate of development would be to the species'
advantage.

The searches made of the forest litter, of trees and shrubs in these Kenyan forests have
revealed only three species of Anura, two of them arthroleptines a group known to develop
directly from eggs laid in underground chambers to fully transformed miniatures of the adult
frogs. The third, Mertensophryne micranotis, has been reported as breeding in water-filled
treeholes and land snail shells and has an unusual-shaped tadpole with an angled head that is
surrounded by a raised ring of tissue (Grandison, 1980). The anuran composition of a water-
less forest in SE Tanzania is recorded by Loveridge (1942). The area is Nchingida 10°8'S,
39°12'E on the Rondo Plateau which is the type locality of Mertensophryne micranotis
rondoensis (Loveridge), a form that Loveridge distinguished from the nominate on its less
heavily pigmented throat. Loveridge (1944) described the Rondo Plateau as waterless where
even the heaviest rainstorms drain rapidly through the sandy soils. In addition to the M. m.

88

A. G. C. GRANDISON & S. ASHE

rondoensis he recorded (1942, 1944) four other species of Anura — two arthroleptines and
two microhylids, Breviceps mossambicus Peters and Spelaeophryne methneri Ahl. While the
life history of Breviceps is known to be entirely independent of water (Wager, 1965), very
large eggs being laid in burrows often far from water and the tadpole stage passed within the
egg capsule, no aspect of the breeding cycle of Spelaeophryne has been recorded. The right
ovary removed from an adult female S. methneri in the BM collections contains 1 8 large ova,
average diameter 2 mm, and a few much smaller ova: all are unpigmented. Such a small
clutch of unpigmented eggs suggests a specialised reproductive mode that has freed the
species from an aquatic environment, although since the species lacks any obvious means of
burrowing into the litter, unlike Arthroleptis stenodactylus Pfeffer and Breviceps with their
well developed shovel-shaped metatarsal tubercles, it seems unlikely that Spelaeophryne is a
burrowing breeder. Perhaps like Nectophrynoides malcolmi Grandison (1978) it has a
terrestrial tadpole with a vascularised tail that serves as a respiratory organ, but perhaps like
Mertensophryne micranotis it depends on waterfilled holes in which to lay its small
complement of eggs.

The senior author's field study was conducted during the month of May. Mean monthly
rainfall records had suggested that this period would be the most likely to coincide with the
long rains and consequently produce water-filled breeding sites and the likelihood of witness-
ing breeding behaviour in the species. However the rains were capricious in May 198 1 and at
Diani Beach in the southern coastal belt only 19 cm fell during the entire month and for half
the period there was no rain or only a few drops (C. Harcourt, pers. comm.); as a result the
number of water-filled treeholes and land snail shells was exceedingly small and no mating
pairs were found. The standing water in all treeholes found was siphoned and checked for the
presence of anuran eggs and tadpoles but only one sample contained micranotis tadpoles; it
was in one of three holes in the buttress formation of a tree that branched near its base into
three trunks and was the only one obscured by a green leafy liana which partly shaded it from
the sun. The diameter of the tree trunk at the level of the hole was 54 cm. The other two
waterfilled holes in the same tree were at a similar height from the forest floor and although
their sizes fall within the range of variation of other oviposition sites for micranotis the holes
contained only insect larvae (Table 1). All the tadpoles, in excess of 45, had developed limbs
but still had larval mouthparts. The pH of the water was not taken and the identity of the tree
is not known.

The mammalogist, Dr G. Rathbun, reports (pers. comm.) that while studying elephant
shrews and cutting forest trails at Gede he was attracted by a faint squeak to a hole in a tree

Table 1 Records of water-filled treeholes occupied by Mertensophryne micranotis. From data
supplied by G. Rathbun (t) and L. P. Lounibos (*)

Height from

Volume of Larvae (L)

forest floor

Diameter

Depth

water

or

Location

Date

(cm)

(cm)

(cm)

(cc)

Toads (T)

t Gede

18.V.71

150-0

10-16

Full

T(cf and 9)

* Kombeni

16.V.75

700

L

16.V.75

230

L

24.iv.76

90

L

24.iv.76

30-48

3-8x5-1

5-08

55

L

27.iv.76

101-60

3-8x7-6

7-62

20

L

7.V.77

60

T (in amplexus)

* Makadara

ll.iv.76

17-78

2-5x2-5

6-35

70

L

ll.iv.76

96-52

2-5x2-5

6-35

25

L

19.iv.76

121-92

3-8x3-8

7-37

45

L

Shimoni

12.V.81

20-32

2-5x3-8

10-16

Full

L

MER TENSOPHR YNE MICRANO TIS 89

7-6 cm diameter which he tentatively identified as Lecaniodiscus fraxinifolius. The hole was
in a stump formed by a fallen limb 1-5 m from the forest floor. It contained a pair of
sexually mature M. micranotis which suggests that the hole had been selected as a potential
breeding site. Dr L. P. Lounibos (pers. comm.) provides the only other known records of the
occurrence of micranotis in water-filled treeholes. His records and details of the Gede and
Shimoni treeholes are given in Table 1 . Dr Lounibos' observations were made in the course
of his research into mosquito habitat segregation in Makadara Forest Shimba Hills and
Kombeni Forest, Rabai Location (3°55'S: 39°34'E). Samples of his Makadara Forest
tadpoles and adults were identified by A. McKay, National Museums, Kenya and the
identity of a sexually mature male obtained at Kinondo Forest was confirmed by the senior
author. It is assumed that his Kombeni tadpoles and toads were also correctly identified.
Lounibos (1981) should be consulted for rainfall records and seasonality of water in treeholes
in his study areas.

Movements, colour change and territorial behaviour

Skulking movements and exceptionally effective camouflage are striking features of M.
micranotis. Adults and subadults when disturbed on the forest floor tend to remain motion-
less or to very slowly back under a dead leaf or twig or disappear down a hole. A gravid
female unearthed from loose soil at the base of a tree flattened its body and remained inert for
some time after capture and to the extent that it resembled a dead leaf.

The disruptive dorsal colour pattern of shades of brown closely matches the leaf litter in
which micranotis is usually found and makes the animal exceedingly difficult to detect. The
darkest areas of the body are invariably the lateral band and the interocular bar. A pale hair-
like vertebral line of variable length, but usually extending from behind the eyes to the
sacrum is present in most specimens. It is sometimes flanked by an irregular dark brown
band that divides in the sacral region to form a large ring enclosing a pale zone which extends
posteriorly to a brilliant white supra-anal triangle. This white triangle is characteristic of the
species. The dark vertebral band and ring are separated from the even darker lateral band by
a broad lighter area which is particularly subject to variation in colour according to the
animal's background. For instance, in an adult male transported on white plastic foam this
dorsolateral area was a very pale grey but after a day spent on dead leaves, twigs and bark the
entire dorsum changed to dark brown, while two days later the pale grey areas changed to
khaki. Both in its dark and khaki phases the toad was barely discernible among the litter.
Recently transformed juveniles and halfgrown individuals are usually black when they
emerge from their hiding places but may acquire a greenish-brown tinge when more active.
Ventrally, both adults and young have prominent dark blotches on a white background.
Males tend to have a brighter, more contrasting colour pattern than females. Observations on
captive individuals suggest that the species is diurnal and particularly active in the morning,
also that it is territorial, with individuals having their own burrows in the leaf litter. In the
wild it is likely that members of the species occurring in forest overlying coral rag will use as
retreats the abundant holes in the coral as well as burrows in soil and leaf litter. After active
foraging the toads usually back down into their burrows. Adults are more secretive than
juveniles and were not seen to make their presence obvious by hopping or other rapid move-
ments; their movements were exceedingly slow, laborious and consisted of usually no more
than five steps at a time followed by a long pause. On the other hand, juveniles tended to
scramble and hop but such activity was interspersed with long periods of immobility.

The junior author noticed the species' propensity while in captivity to climb. Day old
toadlets climbed with apparent ease the glass sides of a casserole in which they were reared.
Generally they moved hand over hand with the body raised but when climbing clean glass
they lowered their bodies. Adults not only readily climbed the plants in the terrarium,
occasionally roosting on leaves and crawling along narrow stems but they persistently
climbed the glass sides of the rectangular tank usually by straddling a corner of the terrarium

90 A. G. C. GRANDISON & S. ASHE

and bracing their limbs. As they climbed the glass it was noticed that they left behind a trail
of liquid and that the posterior part of the toad's abdomen was flattened on the substrate. It is
believed that the liquid is expelled from the toad's cloaca and that the surface tension set up
facilitates progression over a smooth surface. By developing a technique that allows a short
and spindly legged species to climb smooth vertical surfaces the availability and variety of
treehole oviposition sites may also be increased thus enhancing the chances of survival.

Captive breeding programme

One subadult and eleven juveniles collected between 29 June and 1 1 July 1981 were reared
in captivity to sexual maturity. They were collected from an area of less than an acre in the
vicinity of the junior author's house at Watamu. The area once covered by primary forest
and an extension of the Gede Forest was cleared of all but the large trees for house building in
1980. The toads were found on and among the leaf litter overlying coral rag in or at the edge
of a surveyor's cut line as well as in a large termite-ridden rotten log in the undergrowth.

The terrarium consisted of a glass tank 40x28x22-5 cm furnished with a 2-5 cm layer
of leaf litter and sand, growing plants and logs. A coconut shell with a little rain water and
a plastic jar 12 cm deep and a 6 cm diameter, filled with rainwater and tilted at an angle
of 45° against a twisted root provided potential oviposition sites. The root acted as a climbing
frame and provided numerous hiding places. A wooden ramp in the jar of water formed an
exit.

By the 22 October, five days after the rains had started the day temperature exceeded 30°C
and night time temperature 25°C and there was high humidity, the first batch of eggs was laid
but neither mating nor egg laying was witnessed. Several toads appeared gravid.

The following accounts are condensed from the detailed notes made by the junior author
on her observations on the mating behaviour of her captive specimens.

Courtship

It remains uncertain whether females approach individual males and what signals are used to
initiate courtship because pairs were already in amplexus on each occasion when
observations were begun. However it was noticed that the soft chirp of a male in amplexus
prompted three other males that had been chasing each other around in the water container
to call and mount each other indiscriminately and their calls appeared to induce two
females to move in the direction of their calls but stop and wander off when the calling
ceased. The behaviour of the three males in the jar suggests that the normal calling-station
may be a water-filled container and that the receptive female moves in that direction when
stimulated by the advertisement call. However the mating behaviour of only six pairs has
been observed and while one pair mated in water the other pairs mated mostly out of water.
Visual as well as auditory cues may contribute to mate selection in M. micranotis for it was
noted that the three calling males in the water jar were of a brighter colour and more con-
trasting pattern than the amplectic male which was very dark coloured.

Amplexus and fertilisation

The mating of one pair, which spanned a period of over eight hours, is described in detail.
When the male first mounted the female, their eyes were in the same vertical plane while his
hind legs trailed on the ground. The amplectic position was axillary. As he drew up his legs
he stimulated the female's sacroiliacal region by drumming it with his long fourth toe then
placed his feet over her tibiotarsal joints. Half an hour later he uttered an almost inaudible
rapid ticking sound which the female answered with a soft chirp. The pair were clamped
closely to each other from snout to vent but although their white supraanal patches were
vertically in line a distance of about 2 mm separated them. After an interval of about a few

MER TENSOPHR YNE MICRA NO TIS 9 1

minutes the female alternately inflated and deflated her body, as if sighing, while his feet
were placed on her flanks. As her body deflated at 30 second intervals the male exerted a
downward squeeze with his cloacal region. Despite the female moving away and trying to
dislodge the male by scratching him first with her hind leg then with a front leg, the male
remained firmly attached, although at one point he was shifted sideways as she forced her
way under a piece of bark. Mating movements were discontinued while the female fed on
white ants but were resumed approximately two hours after observations on the amplectic
pair were begun. The male resumed tactile stimulation of the female by drumming his fourth
toe. His heels which were then placed on either side of and slightly above her white
supraanal patch seemed to channel the drop of clear liquid that trickled from his cloaca
down to hers where it appeared to be absorbed. As the female adopted a more upright stance
the male moved farther back and lower down and his mating movements became more
forceful. As he engaged in a series of thrusts his white supraanal patch was seen to curl
inwards towards that of the female. During the ensuing two hours mating movements
continued intermittently, with a rest period while the female, still with male attached, moved
to a hiding place in the litter but when the amplectic male produced a rapid, very faint
ticking sound another male called from 25 cm away in the water jar; the vocalisation elicited
a marked increase in activity by all the other males in the terrarium which chased and
mounted each other indiscriminately while their calls became louder until each call sounded
like a ten note chirp which could be heard 2-5 m away. Several females were active on the
floor of the terrarium but although one of the calling males attempted to mount a female it
did not persist when the female moved away. When the amplectic pair emerged from the
hiding place more vigorous mating movements occurred, despite the female's attempts on
two occasions to dislodge the male, and he thrust twice each time the female deflated her
body. Although both sexes were visibly vibrating no call from the female was audible but the
male emitted a ticking sound. During the next hour sporadic mating movements occurred
but his vent was above hers and although the pair entered the coconut shell partly filled with
water and moved round the slope in a clockwise direction only her foot entered the water and
the pair climbed out of the shell. The female again tried to dislodge the male while she ate
insects. Observation was discontinued for ten minutes during which time the pair
disappeared.

In five other pairs in which mating was witnessed the duration of amplexus varied from
five and a half hours to ten hours. It was noticed that the male's grip on the female's vent is so
tight that his vent drags upwards the skin surrounding her vent. One pair found in the water
jar in the 'normal' amplectic position were joined by a second male which clung to the
female's axillae in an inverted, belly to belly position. Both males engaged in mating move-
ments but two hours later when the dorsal male had disappeared the protruding cloacal
region of the underslung male was seen to curl upwards to contact that of the female and the
moisture he expelled from his vent as he thrust appeared to immediately be absorbed by the
female. Belly to belly amplexus is known to occur also in the internally fertilised Ethiopian
bufonid, Nectophrynoides malcolmi Grandison (Grandison, 1978 and Wake, 1980).

Although clutches of eggs were later found in the jar of water in the terrarium, egg laying
by mated females had not been witnessed and because males were present the possibility of
the eggs having been externally fertilised could not be ruled out. So on the next occasion
when a pair began to mate they were removed to another terrarium containing a water bowl
of similar capacity and quantity of water. When mating was completed and the male had
dismounted he was removed and returned to his original terrarium. Mating had taken place
in water. Six hours after mating the female, still in the water, began laying eggs, two strings
emerging together. Three and a half hours from the start of egg laying and at 2300 hours
oviposition had not been completed, but the following morning two eggs strings containing a
total of 22 eggs were found attached to the rim of the jar. The female had returned to the floor
of the terrarium. The eggs proved to be fertile and they developed at a similar rate to those in
previous clutches. Each of the six eggs in one string that was severed and preserved within
forty minutes of being laid was found to be at the late cleavage stage of development, but eggs

92 A. G. C. GRANDISON & S. ASHE

from another clutch where the time of laying was unknown were at early to late gastrula. The
number of eggs in five clutches were 17, 19,22,32, 32.

The aforegoing description emphasises the close contact of the cloacae during mating in
Mertensophryne micranotis and the tenacious grip that the male exerts on the female's vent
while he engages in downward thrusts and his feet are placed on each side of the female's
vent. While it can be assumed that the male's cloacal spines play a significant part in the
coupling and that internal fertilisation of the eggs takes place it is still not known whether the
spines interlock in the furrows of the female's vent.

Function of the tadpole's head 'crown'

Broadley (in Channing, 1978) suggested that the function of the head 'crown' in tadpoles of
Stephopaedes anotis (Boulenger) might be to exclude from the eyes and nostrils scum
accumulating on the water surface. The angled head and ring of raised tissue ('crown') of the
M. micranotis tadpole is closely similar to that of S. anotis (Grandison, 1980) but
observations on tadpoles of M. micranotis in the wild and in captivity suggest that the
angled head with the raised ring of tissue surrounding a saucer-like depression in which lie
the eyes and nostrils is a simple adaptation for suspending the tadpole at the meniscus, where
the highest concentration of oxygenated water is available in the small pocket of stagnant
water that is selected as a breeding site. With the 'crown' breaking the water surface the
tadpole has access to oxygen, both during its gill breathing stages and later when it has
acquired lungs, at the same time maintaining itself in a tail-down position with the minimum
expenditure of energy. In the field study it was noticed that each time the sun struck the
water surface of the treehole the tadpoles tended to rise quickly and suspend themselves at
the meniscus, particularly around the rim of the hole so that their angled heads were parallel
to the contour of the meniscus whilst the body and tail of the tadpole hung down vertically.
In such a position the ventral mouthparts had access to the algal growth lining the rim of the
cavity. When the head 'crown', which is at an angle of 45° to the body and tail, breaks the
surface tension at or near the centre of the hole where the meniscus is virtually horizontal the
body and tail of the tadpole are suspended parallel to the angle of the head.

Diet

The contents of the stomachs of three subadults that were known to have been preserved
immediately after capture were analysed. Ants constituted the major food element but mites
were found to have been almost equally favoured, particularly Linopodes (Eupodidae).
Although nine other genera of mites were present only one or two examples of each were
identified (Eupelops, Trachygalumna, Pilizetes, Liodes, Eremaezetes, Scapheremaeus,
Bdella and Spinibdelld). Beetles and thrips of kinds typical of the litter/subcortical layer were
also represented in the analysis, as well as Collembola (Symphypleona) and the macerated
remains of spiders and fly larvae. The only trace of termites was a crumpled set of wings of
Termitidae. In captivity the toads thrived on a diet of small white ants, termites and aphids.

Acknowledgements

This study was supported by the Fauna and Flora Preservation Society and the British
Museum (Natural History). Field work was made possible through the assistance of the
Office of the President of Kenya and the cooperation of Mr A. D. McKay, National
Museums of Kenya. The senior author thanks Drs L. P. Lounibos and G. Rathbun for
making available their observations on treehole occupancy by M. micranotis; her colleagues
Dr W. Sands and Miss A. Baker for analyses of stomach contents; Miss C. Harcourt for field

MER TENSOPHR YNE M1CRA NO TfS 93

assistance and rainfall records; and Mr and Mrs M. McKay and Mrs D. Webb for kind
hospitality. The assistance given by Dr R. C. Drewes, Mr S. Reilly and Mr E. Wederkinch in
making material and field data available is also gratefully acknowledged, as is Professor Ruth
Bellairs' cooperation.

References

Anderson, J. 1871. A list of the reptilian accession to the Indian Museum, from 1865 to 1870, with a

description of some new species. J. Asiat. Soc. Beng. 40: 12-39.
Barbour, T. & Loveridge, A. 1928. A comparative study of the herpetological faunae of the

Uluguru and Usambara Mountains, Tanganyika Territory with descriptions of new species. Mem.

Mm. comp. Zool. Harv. 50: 87-265.
Britton, P. L., Britton, H. A. & Coverdale, M. A. C. 1980. The avifauna of Mrima Hill, South

Kenya Coast. Scopus 4: 73-78.
Channing, A. 1978. A new bufonid genus (Amphibia: Anura) from Rhodesia. Herpetologica 34:

394-397.
Dubois, A. 1976. Les grenouilles du sous-genre Paa du Nepal (famille Ranidae genre Rand).

Cahiers nepalais. Documents no 6. C.N.R.S. vi + 275 pp. Paris.
Grandison, A. G. C. 1978. The occurrence of Nectophrynoides (Anura Bufonidae) in Ethiopia. A

new concept of the genus with a description of a new species. Monit. Zool. ital. N.S. Suppl. XI:

119-172.

1980. Aspects of breeding morphology in Mertensophryne micranotis (Anura: Bufonidae):

secondary sexual characters, eggs and tadpole. Bull. Br. Mus. nat. Hist. (Zool.) 39: 299-304.

Lounibos, L. P. 1981. Habitat segregation among African treehole mosquitoes. Ecol. Entom. 6:

129-154.
Loveridge, A. 1925. Notes on East African Batrachians, collected 1920-1923, with the description

of four new species. Proc. zool. Soc. Land.: 763-79 1 .

1942. Scientific results of a fourth expedition to forested areas in East and Central Africa. V.

Amphibians. Bull. Mus. comp. Zool. Harv. 91: 377-436.

1944. Scientific results of a fourth expedition to forested areas in East and Central Africa. VI.

Itinerary and comments. Bull. Mus. Comp. Zool. Harv. 94: 191-214.
Spear, T. T. 1978. The Kava Complex. A history of the Mijikenda peoples of the Kenya Coast to

1900. xxiv+ 172 pp. Nairobi.

Wager, V. A. 1965. The frogs of South Africa. 242 pp. Cape Town and Johannesburg.
Wake, M. 1980. The reproductive biology of Nectophrynoides malcolmi (Amphibia: Bufonidae),

with comments on the evolution of reproductive modes in the genus Nectophrynoides. Copeia no. 2:

193-200.

Manuscript accepted for publication 3 August 1982

Additional notes on bariliine cyprinid fishes

Gordon Howes

Department of Zoology, British Museum (Natural History), Cromwell Road, London

SW75BD

This addendum serves to correct and amplify statements and observations published in an
earlier study on the anatomy, phylogeny and classification of bariliine cyprinid fishes
(Howes, 1980).

The generic status of Barilius lujae Boulenger, 1909

In the synoptic account of Barilius species given in Howes (1980), Barilius lujae Blgr, 1909
was omitted. Examination of syntypes (BMNH 1908.11.26:8; 1909.4.26: 18), together with
a series of more recently collected specimens (BMNH 1975.6.20:429^438;
1976.6.20 : 407^413, including alizarin preparations), makes it clear that the species lujae
should be assigned to the genus Leptocypris.

Synapomorphies characterising Leptocypris (modified from Howes, 1980) are: shallow
lower jaw; truncated lateral ethmoid; absence of, or reduced intermandibularis muscle;
absence of, or few gill-rakers; elongated pectoral and pelvic axial scales, and overlap of the
antero-dorsal portion of the 2nd infraorbital by the posterior border of the 1st. The species
lujae has all these characters and, in addition, a type of ethmoid architecture which supports
the hypothesis that Leptocypris and Engraulicypris are closely related genera (Howes,
1980: 182).

In Leptocypris lujae the medial ethmoid notch is horseshoe-shaped as in Engraulicypris
sardella (cf. Fig. 1 here with fig. 9 in Howes, 1980). In Engraulicypris the ethmoid
indentation coincides with a foramen in the underlying vomer, whereas in L. lujae the
indentation is floored partly by the ethmoid cartilage and partly by the vomer.

Compared with other species of Leptocypris, L. lujae is longer-jawed, the posterior tip of
the maxilla extending to, or beyond, the posterior rim of the orbit; there is also a lower jaw
symphysial process. As in L. niloticus and L. weynsii, L. lujae has a non-papillate maxillary
valve. There is some variability in the length of the pelvic axial scale, it varies between
42%-50% of the pelvic fin length. However, this appears to be positively correlated with the
length of the fish and it is noted that in L. niloticus the pelvic scale is not developed to its
maximum length (25% of the pelvic fin length) until the fish is over 35 mm SL.

In the length of the jaws and the morphology of the ethmoid region, L. lujae appears to be
the most derived species. The species of Leptocypris may be identified by the following key:

1. Gill-rakers on first ceratobranchial absent; branched anal fin rays 8-9 . . . L.modestus
Gillrakers on first ceratobranchial 2-3; branched anal fin rays 1 1-15 ....

2. Branched anal fin rays 1 1-12 L. niloticus

Branched anal fin rays 14-15 3

3 Posterior tip of maxilla extending to centre of eye; lateral line scales 44-45 . . . L. weynsii
Posterior tip of maxilla extending to, or beyond posterior border of the eye; lateral line scales
38-40 L. lujae

Attention is drawn to 'Barilius' guineensis Daget, 1962, which appears, from its gross
morphology, to belong to Leptocypris (see Howes, 1980 : 191). It differs from other species,
however, in (according to Daget, 1962) having a total of 8 gill-rakers on the first arch.

Bull. Br. Mm. not. Hist. (Zool.)45 (2): 95-101 Issued 28 July 1983

95

96

G. J. HOWES

Mec

Fig.

1mm

1 Leptocypris lujae\ dorsal view of ethmovomerine region of the neurocranium.
Mec = mesethmoid cartilage, Pe = preethmoid, Se = supraethmoid, Vo = vomer.

Comments on south-east Asian Barilius species

When offering a revised concept of Barilius (Howes, 1980), based on the type species of the
genus B. barila, few of the south-east Asian species had been examined and their generic
attribution remained doubtful. Of those species in this category (Howes, 1980 : 190) the
following have now been examined: B. bernatziki Koumans, 1937 (Holotype, Basle
Museum, NHMB 5155, 77 mm SL); B. huahinensis Fowler, 1934 (USNM 103104, 52 &
66 mm SL); B. infrafasciatus Fowler, 1934 (USNM 107910, 65-75 mm SL); B. koratensis
Smith, 1931 (Holotype, USMN 90298 47-3 mm SL); B. nanensis Smith, 1945 (Paratypes
USNM 107939, 107940, 119474-119476, 53-64 mm SL); B. pulchellus Smith, 1931
(Paratypes USNM 90299, 51 & 53 mm SL; USNM 107967,39-5-69-5 mm SL).

Osteologically and meristically, these species are intermediate between the generic
subgroups (i) and (ii) previously postulated (Howes, 1980: 180), possessing characters of
both groups. This suggests that those characters used in defining the two groups are a mixture
of plesio- and apomorphies.

In the south-east Asian species, and noticeably in B. pulchellus, the ethmoid bloc is
somewhat flattened with the vomer extending anteriorly to the mesethmoid. The lateral
edges of the supraethmoid are slightly raised to produce a shallow channel. The maxilla has a
tall mid-lateral (palatine) ascending process and the palatine is broad anteriorly, slightly
overlapping the maxilla and supporting the base of the anterior barbel. Rows of tubercles
appear on the lower jaw and infraorbitals. These characters are shared with the Indian
species, B. gatensis and B. bendelisis, and those species referred to group (ii) by Howes
(1980: 190). The morphology of the ethmo-vomerine region, lateral extension of the
palatine, and the pattern of tubercle development on the lower jaw appear synapomorphic
for this group of species. As such, the concept of Barilius presented previously (Howes,
1980) is shown to be a paraphyletic one; see below, p. 99.

The taxonomy of the south-east Asian species is in an unsatisfactory state. Smith (1945)
separates B. nanensis and B. huahinensis on the differences in lateral line scale counts, dorsal
fin position, and length of the anterior (rostral) barbel. However, there appears to be

BARILIINECYPRINID FISHES 97

variability and overlap in all these characters, although the differences in body markings
between the two species quoted by Smith (1945 : 156-157) are consistent. In B. nanensis
there are 7-8 gill-rakers on the 1st ceratobranchial cf. 9-10 in B. huahinensis.

Barilius pulchellus Smith, 1931 is a distinctive species characterised by a large pectoral
axial fin lobe, 3-6 minute, spiny gill-rakers on the 1st ceratobranchial, and a black dorsal fin
membrane. Specimens identified as B. infrafasciatus Fowler, 1934 (type not seen) are similar
in virtually every respect to B. pulchellus apart from the length of the pelvic axial scale which
is 50% of the pelvic fin length cf. 25% in B. pulchellus.

Smith (1931) described Barilius koratensis as without barbels. My examination of the
unique holotype reveals both anterior and posterior barbels. Smith described only two
vertical bars, above the pectoral and below the dorsal fin, but there is also a dark patch above
the last ray of the anal fin. In other meristic and morphometric characters B. koratensis
resembles B. nanensis; these taxa may well be only representatives of different populations of
one species.

Barilius bernatziki Koumans, 1937 has 30-31 lateral line scales, the least number of all
the south-east Asian species. In its markings (7 dark vertical bands and basal caudal spot),
large pectoral axial lobe, and narrow cranium, B. bernatziki most closely resembles the
Indian species, B. gatensis.

The phyletic position of Zacco

Zacco was excluded from the bariliine group by Howes (1980) on the grounds that it lacked a
posttemporal-subtemporal connection. Fink & Fink (1981) claim this feature is present, and
an examination of additional material leads me to agree with them. If Zacco is included
within the bariliine assemblage, characters which were previously regarded as examples of
homoplasy (peculiar morphology of the anal fin, ventral prolongation of the caudal peduncle
and colour pattern), must now be viewed as synapomorphies.

Zacco must be regarded as the sister taxon to Opsariichthys and because of its less derived
cranial architecture and relatively unmodified lower jaw, it is the plesiomorph partner.
Inclusion of Zacco within the bariliine group does not alter the group's distribution mapped
in Howes (1980, fig. 47).

Monophyly of the bariliine group

According to Fink & Fink (198 1) a number of characters used by Howes (1980) to define the
bariliine group are plesiomorphic. Of these they list the ventrally open posterior myodome;
trigeminal foramen entirely within the prootic; lateral hyomandibular flange; frontal fossa;
lateral temporal foramen, and the metapterygoid postero-dorsal process. They also believe
that fusion of the 2nd and 3rd centra, only present in some bariliines, may be synapomorphic
for those taxa and other Cyprinidae.

Posterior myodome. Whilst it is recognised that a ventrally open posterior myodome often
occurs in what are regarded as plesiomorphic teleosts (Patterson, 1975) it does not follow
that the feature is itself plesiomorphic. Amongst otophysans, the posterior myodome in
characoids is developed to various degrees. In some taxa currently recognised as mono-
phyletic groups (Alestinae) there are some species with an open and others with a closed
myodome. Also, in those taxa usually regarded as plesiomorphic (e.g. Hepsetus) the
myodome is closed, whereas in those considered derived (Serrasalmidae) it is open.
Ontogenetic evidence for polarity is contradictory; in a cyprinid investigated the myodome
closes during ontogeny (Opsariichthys; see Howes, 1980), whereas in salmonids it opens
(Verraes, 1974). Functionally, an open myodome serves to extend the length of the posterior
eye muscles which then originate either from the rim of the basioccipital or from the anterior
vertebrae. From its mosaic distribution and varying morphology, it is more parsimonious to

98 G. J. HOWES

suppose that an open posterior myodome has developed independently in several lineages,
and as such is an unreliable indicator of relationship.

Hyomandibular flange, frontal fossa and temporal foramen. For the most part the same
argument may be applied to these features as to the posterior myodome, namely, mosaic
distribution through independent derivation, and thus I agree with Fink & Fink (1981) in
disregarding them. However, some mention must be made of the lateral temporal foramen.
Wu, Chen, Chen & Chen (1981) considered that the lateral temporal fossa in the Cobitidae
and Gyrinocheilidae, and the supratemporal fossa in Catostomidae are homologues of the
post-temporal fossa in other cyprinoids. This certainly appears to be so, as in all cases the
fossa is bounded by the epioccipital, pterotic and, usually, the parietal. The lateral temporal
foramen described in Opsaridium (Howes, 1980) is not to be confused with the above as it
occurs between the pterotic and sphenotic and is thus homologous with that aperture in some
characoids (e.g. Salminus).

Metapterygoid process. Although a metapterygoid dorso-posterior process occurs in other
otophysans, it differs markedly from the condition considered derived for some bariliine taxa
(Howes, 1980). In Raiamas the process is directed anterodorsally at an angle of 45° and is the
same length as the upper part of the hyomandibula. Furthermore, the process has a gutter
along its posterior border into which the levator arcus palatini muscle inserts. In other
bariliines the process is not so well developed, but in Engraulicypris and Leptocypris it is tall
and lamellate, occupying a mid-dorsal position on the metapterygoid. In these taxa it also
serves to support the LAP muscle. Whilst it is accepted that a metapterygoid process is a
common otophysan feature, in none does it possess the morphology of the process in
bariliines, nor does it provide the site of attachment of almost the entire LAP muscle.

Vertebral fusion. Fink & Fink's (1981) claim that all other members of the bariliine group are
'. . . more specialised than Opsariichthys in having the second and third centra fused' must
be treated with caution.

Fusion of centra 2 and 3 is a variable character throughout the Cyprinidae. There may be
only partial (dorsal) fusion, as in Zacco and Engraulicypris (Howes, 1980) and Pseudo-
laubuca (Howes, 1978). Monophyletic groups contain taxa exhibiting both fused and
unfused centra (e.g. aspinines and squaliobarbines) and in such cases it is not always the
more derived members of the groups which display the fusion pattern. Thus, vertebral fusion
must be regarded as having been derived independently in several lineages. Supporting Fink
& Fink's argument for the derived nature of vertebral fusion simply leads to an unresolved
reticulate pattern of relationships (Fig. 2).

Interrelationships of bariliines

Disregarding the posterior open myodome and those characters considered by Fink & Fink
(1981) as plesiomorphic, the bariliine group can be defined on only a single synapomorphy,
ie a connection between the posttemporal and subtemporal fossae. That this character has
now been found in Salmostoma (placed in the cheline group by Howes, 1979) revises the
concept of the bariliine group.

Accepting the fossae connection as synapomorphic, then two lineages can be recognised,
(1) those species with elongate pectoral and pelvic axial scales, long and expanded processes
of the 1st vertebrae, with development of a condylar joint with the basioccipital, and a
modified ethmo-vomerine region (see caption to Fig. 3), and (2) those with the pectoral and
pelvic axial scales lobate or fleshy, the rostral barbel attached to the lateral border of the
palatine, parallel rows of tubercles on the lower jaw, and distinctive body and fin patterning.

Lineage (1) includes Barilius barila, B. evezardi, B. modestus and B. vagra, Salmostoma,
Engraulicypris, Raiamas and Leptocypris. Salmostoma was regarded by Howes (1979) as the
plesiomorph sister group of 'cheline' genera. The 'chelines' share with Barilius and
Salmostoma the derived form of anterior vertebrae and ethmovomerine region, but lack the

BARILIINECYPRINID FISHES 99

Other Some Zacco.Opsariichthys

'bariliines' cyprinids & some cyprinids

Centra 2 3-^ -¥• — Centra

fused; \ / unfused

'derived'

Fig. 2 A cladogram of cyprinid relationships based on the pattern of vertebral fusion proposed by

Fink & Fink (1981); see text.

posttemporal-subtemporal connection. Salmostoma, however, shares with all other chelines
a deep, posteriorly sloped neural complex. Only one 'cheline' genus, Securicula shares the
elongate pectoral and pelvic axial scales with Salmostoma and Barilius. In the absence of
synapomorphies linking the 'cheline' genera with any other cyprinid taxon, the absence of
these features is considered as a phylogenetic loss. Current observations indicate that the
genera Rastrineobola, Neobola and Chelaethiops also belong to this lineage, and work in
progress attempts to define more precisely the relationships of these three genera.

Lineage (2) contains the remaining Barilius species — recognised as group (ii) by Howes
(1980: 1 80)— together with Opsaridium, Opsariichthys and Zacco, plus the luciosomine
genera (sensu Howes, 1980).

In summary, it would seem that the bariliine and cheline groups (sensu Howes, 1979;
1980) are paraphyletic assemblages; a revised hypothesis of the interrelationships of the
relevant taxa is presented in Fig. 3.

The dates of Opsaridium Peters and Rastrineobola Fowler

Jordan (1919) mistakenly gives the publication date of Opsaridium Peters as 1855.
Howes (1980) followed this error. The correct date is 1853 (see citation in references). The
specific name is correctly zambezensis.

The publication date of Rastrineobola Fowler is 1936, not 1934 as misprinted in Howes
(1980: 195).

Bengala Gray, 1832, a senior synonym of Megarasbora (, tint her, 1868

Giinther (1868) established the genus Megarasbora to contain Cyprinus elanga Hamilton,
1822. Megarasbora was recognised by Brittan (1954) and Howes (1980) in their respective
revisions and reviews of rasborine cyprinids. However, all three authors had overlooked the
fact that Gray (1832) had already established the genus Bengala for Hamilton's species.
Jordan (1919) recorded Megarasbora as a junior synonym of Bengala.

100

G. J. HOWKS

Fig. 3 Revised cladogram of 'bariliine' and 'cheline' relationships (dashed lines indicate
uncertain affinities). 1, posttemporal-subtemporal fossa connection; 2, pectoral and pelvic axial
scales elongate (lost in 'chelines' and Engraulicypris); 2a, 1st vertebra with expanded lateral
processes; 2b, omega-shaped ethmoid notch, vomerine arms extending straight forward; 2c,
reduction of dilatator fossa, derived condition of jaw adductor muscles; 2d, loss of
intermandibularis muscle, jaw elongation, hypertrophy of metapterygoid spine; 3, pectoral and
pelvic axial scales lobate or fleshy; 3a, palatine extended laterally, supports rostral barbel (when
present); 3b, parallel rows of tubercles on dentary; 3c, dentary with anterior notch, sexual
dimorphism exhibited by extension and expansion of anal fin rays in males; 3d, bowl-shaped
depression in supraethmoid. Characters described in Howes (1980), except 2b and 2c, taken
from work in preparation.

Acknowledgements

My sincere thanks are due to Dr Richard Vari for facilitating my study of Smith's specimens
in the United States National Museum and for his warm hospitality. I am most grateful to
Maurice Kottelat for providing information on south-east Asian Barilius and arranging loans
of specimens, and to Margaret Clarke for her assistance with nomenclatural problems. Drs
Humphry Greenwood and Keith Banister have kindly read, and critically commented on the
manuscript.

References

Brit tan, M. R. 1954. A revision of the Indo-Malayan fresh-water fish genus Rasbora. Institute of

Science & Technology, Manila. 224 pp.
Daget, J. 1962. Les poissons du Fouta Dialon et de las basse Guinee. Mem. Inst. fr. Afr. noire 65:

1-210.

BARILIINECYPRINID FISHES 101

Fink, S. V. & Fink, W. L. 1981. Interrelationships of the ostariophysan fishes (Teleostei). Zool. J.

Linn. Soc. 72 (4): 297-353.

Gray, J. E. 1 832. Illustrations of Indian Zoology. Part 2, pi. 96. London
Giinther, A. 1 868. Catalogue of fishes in the British Museum 7: 1-5 1 2.
Howes, G. J. 1978. The anatomy and relationships of the cyprinid fish Luciobrama macrocephalus

(Lacepede). Bull. Br. Mus. nat. Hist. (Zool.) 34: 1-64.
1979. Notes on the anatomy of the cyprinid fish Macrochirichthvs macrochirus with a review

of the subfamily Cultrinae. Bull. Br. Mus. nat. Hist. (Zool.) 36 (3): 147-200.

1980. The anatomy, phylogeny and classification of bariliine cyprinid fishes. Bull. Br. Mus.

nat. Hist. (Zool.) 37 (3): 129-198.
Jordan, D. S. 1919. The genera of fishes Part 3. Leland Stanford Junior Univ. Pubs. University

Series:285^MO + i-xv.
Peters, W. C. H. 1853. Uebersicht der in Mossambique beobachteten Seefische. Monats. Ber. Acad.

Berlin: 783.

Smith, H. W. 1945. The fresh-water fishes of Siam, or Thailand. Bull. U.S. natn. Mus. 188: 1-622.
Verraes, W. 1974. Discussion on some functional-morphological relations between some parts of

the chondrocramium and the osteocranium in the skull base and the skull roof, and of some soft head

parts during postembryonic development of Salmo gairdneri Richardson 1836 (Teleostei:

Salmonidae). Forma Functio 7: 28 1-292.
Wu, X., Chen, Y., Chen, X. & Chen, J. 1981. A taxonomical system and phylogenetic relationship

of the families of the suborder Cyprinoidei (Pisces). Scientia Sinica 24 (4): 563-572.

Manuscript accepted for publication 1 7 June 1982

British Museum (Natural History)

An Atlas of Freshwater Testate Amoebae

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1980, Hardcovers, 222pp, £1 7.50 (£1 8.00 by post). Co-published by British Museum
(Natural History) and Oxford University Press.

This book illustrates, using scanning electron micrographs, most of the common species of
testate amoebae that are found in freshwater habitats. Information on the biology, ecology,
geographical distribution and a classification are followed by descriptions of ninety-five
species. Each of these is illustrated by several views of the shell.

The text is designed not only to enable biologists to identify species of testate amoebae, but
to serve as an introduction to students interested in the taxonomy and biology of these fresh-
water protozoa. It will be of special interest to prptozoologists, ecologists, limnologists, water
treatment specialists and micropalaeontologists interested in recent sediments.

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Bats (Mammalia: Chiroptera) from Indo- Australia. By J. E. Hill.

On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the
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Bulletin of the

British Museum (Natural History)

Bats (Mammalia: Chiroptera) from
Indo- Australia

J. E. Hill

Zoology series Vol 45 No 3 25 August 1983

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
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© Trustees of the British Museum (Natural History), 1983

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ISSN 0007- 1 498 Zoology series

Vol 45 No. 3 pp 103-208
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 25 August 1983

Bats (Mammalia: Chiroptera) from

J.E.HH1,

Department of Zoology, British Museum (Natural History), Cromwell Road,
SW7 5BD

Contents

Synopsis . . . .
Introduction
Systematic section .
Megachiroptera .
Pteropodidae .
Pteropodinae.
Harpyionycterinae
Nyctimeninae
Macroglossinae .
Microchiroptera .
Emballonuridae
Megadermatidae
Rhinolophidae .
Hipposideridae .
Vespertilionidae
Vespertilioninae .
Miniopterinae
Murininae
Kerivoulinae .
Molossidae

Summary ....
Acknowledgements .
References.
Addendum

103
103
104
104
104
104
127
130
132
140
140
142
143
148
153
153
171
190
194
195
198
199
199
208

Synopsis

Accessions of Indo-Australian bats to the collections of the British Museum (Natural History) during
the past fifteen years are reviewed in detail, with a particular reference to the collections made in Papua
New Guinea and Sulawesi by 'Operation Drake'. Numerous species and species groups from the region
are examined and in some cases revised; a new species of Hesperoptenus from Sulawesi and a new
subspecies of Myotis adversus from the New Hebrides are described. The more important taxonomic
studies and notes made in this study and the new range records that it reports are listed in a terminal
summary.

Introduction

A variety of collections of bats from the region bounded by India in the west and the New
Hebrides, New Caledonia and Fiji to the east has been received at the British Museum
(Natural History) during the years 1967-1982. Some have come from individuals who have
obtained small numbers of bats while visiting the region, others from large, organised
expeditions that have carried out biological studies, or by donation from workers connected
with organisations and institutions in the area. Individually much of this newly accessed
material scarcely justified separate publication and record, although often unusual or of
taxonomic or faunal interest.

Towards the end of this period, however, extensive collections of bats were made in Papua
New Guinea and in Sulawesi by Mr Ben Gaskell, an ecologist and collector with 'Operation

Bull. Br. A/MS. nat. Hist. (Zool.) 45(3): 103-208

Issued 25 August 1983

104 J. E. HILL

Drake', the commemorative round-the-world voyage of the brigantine Eye of the Wind in
1978-1980 that marked the 400th anniversary of the circumnavigation of the world by Sir
Francis Drake in the Golden Hind. The voyage was interrupted from time to time so that the
scientists, technicians and young people aboard could undertake adventurous, scientific or
voluntary aid projects in various parts of the world. Bats were collected in Indo-Australia
during two of these periods, first at and around Buso and Wau in Morobe Province, Papua
New Guinea, and later in central Sulawesi, chiefly around Morowali where the objective was
ultimately to prepare a Management Plan for the Morowali Nature Reserve.

The study of the specimens obtained by 'Operation Drake', especially of those from
Sulawesi, has entailed a further and more wide-ranging examination of much of the
Indo-Australian bat collection already in London, and has prompted a further examination
of the more recently accessed material that has not before been reported in the literature. In
particular it has led to further study of a number of long standing taxonomic problems in the
area and to some taxonomic changes. Although basically this paper is concerned with
specimens obtained through the explorations initiated and carried out through 'Operation
Drake' it has thus been possible to add a variety of other studies and notes drawn from
material from other localities and sources, much of it directly relevant to the 'Drake'
collections.

Place names in many parts of the Indo-Australian region can present difficulties, there
being sometimes a choice of as many as three variants of any one designation. In general the
traditional and conventional European spelling or usage (i.e. Amboina, Ceram) has been
adopted but Sulawesi has been used throughout for Celebes, this name having come into
general use: however, the New Hebrides remain so called, although recently renamed
Vanuatu. The Indonesian part of New Guinea appears as West Irian, while Papua New
Guinea is used for the rest of the island: here, 'Province' has been omitted from locational
data so that localities appear as 'Wau, Morobe' i.e. Wau, Morobe Province. Further west,
Borneo is used as a general term for the entire island, divided into Sarawak, Sabah, Brunei
and Kalimantan.

Measurements of specimens are in millimetres and with the exception of those of
individual teeth have been made with a dial-reading micrometer. Teeth have been measured
with a mechanical stage fitted to a stereoscopic microscope. Although no standard suite of
external and cranial measurements has been used, conventional measurements are given
where required, amplified in particular cases to conform with those employed by the
describer or by previous workers on the species concerned. To avoid repetition, a notation
has been adopted for the wing elements whereby IIIm for example indicates the metacarpal of
the third digit, III1 its first phalange, IIP its second: when length measurements of these
elements are given in the text these designations appear without preamble.

Systematic Section

MEGACHIROPTERA
PTEROPODIDAE
PTEROPODINAE

Rousettus amplexicaudatus stresemanni Stein, 1 933
RousettusstresemanniSlein, 1935: 91 . Japen ( = Jobi) I, Geelvinck Bay, NE West Irian.

SPECIMENS EXAMINED. Papua New Guinea: cf BM(NH) 69.1416 Madang, 5° 14'S, 145°45'E (skin,
skull; coll. J. I. Menzies); <*BM(NH)73.1967 Rauit, West Sepik, 525 m,3°36'S, 142° 1 5 'E (in alcohol;

INDO-AUSTRALIAN BATS 105

coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973); 3dd BM(NH)
78.850-852 Baku Forest Station, Gogol Valley, Madang (78.850, 851 in alcohol, skull of 78.850
extracted, 78.852 skin only; coll. P. A. Morris); 9 BM(NH) 78.853 Baiyer R, c. 50 km NW of Mt.
Hagen, Western Highlands (in alcohol, skull extracted; coll. P. A. Morris).

REMARKS. Until recently there have been few records of stresemanni and for many years
(Laurie & Hill, 1954) it has been considered a distinct species co-existing in New Guinea
with R. amplexicaudatus brachyotis (Dobson, 1877). Latterly, however, a number of reports
from Papua New Guinea has appeared, McKean (1972) recording a series of stresemanni
from Ihu, Greig-Smith (1975) a specimen (listed above) from Rauit, with a record of an
example (also listed above) from Madang first noted (in litt.) by Menzies, and with Koopman
(1979) reporting material from Bagabag I, off the northeastern coast near Madang and later
(1982) from Dabora, Tapio and Mornuna, Milne Bay, east Papua New Guinea. Menzies
(1977) tentatively referred sub-fossil remains from Kiowa in Ghimbu Province to
stresemanni. Finally, Rookmaaker & Bergmans (1981) have listed numerous records from
localities in much of New Guinea, some evidently based on specimens previously identified
as R. a. brachyotis. Specimens in the American Museum of Natural History suggested to
Koopman (1979) that stresemanni might be properly regarded as a subspecies of R.
amplexicaudatus while Rookmaaker & Bergmans (1981) in their comprehensive review of
this species have synonymised stresemanni with R. a. amplexicaudatus (Geoffrey, 1 8 100).

Specimens recorded here from Papua New Guinea are large, corresponding closely with
the original description of stresemanni by Stein. In size they also agree with specimens from
Ihu reported by McKean (1972) and with those from New Guinea measured by Rookmaaker
& Bergmans (1981). They are generally a little larger than six from Timor examined by
Goodwin (1979) and are similarly near or exceed the upper size limits of Timorese speci-
mens (including some of those seen by Goodwin) studied by Rookmaaker & Bergmans.
Specimens from Timor may be assumed to be topotypical or nearly so of R. a.
amplexicaudatus, described originally from that island. The dimensions given by
Rookmaaker & Bergmans for examples from New Guinea are likewise generally a little
greater than those for specimens from Timor and its associated islands.

External measurements of 4dd and 1 9, with cranial measurements of 2dd and 1 9: length
of forearm 85-8-904, 85-5; greatest length of skull 38-6, 394, 38-8; condylobasal length
36-8, 37-0, 37-5; condylocanine length 35-6, 35-8, 36-0; length front of orbit to tip of nasals
13-1,13-0, 13-1; length palation to incisive foramina — , 17-5, — ; least interorbital width 8-6,
8-3, 8-3; least postorbital width 7-5, 7-6, 8-3; zygomatic width 24-7, 24-0, 22-3; width of
braincase 154, 15-6, 154; mastoid width 14-6, 14-1, 144; greatest width c'-c1 7-1, 74, 7-6,
c'-c1 (alveoli) 6-7, 6-9, 7-0; m^m2 (alveoli) 10-2, 10-3, iO-8; c-m2 13-5, 13-7, 14-0; length
complete mandible from condyles 28-6, 28-3, — ; length right ramus from condyle 30-1,
29-9, 30-0; c-m3 15-1, 14-7, 15-6.

DISCUSSION. Rookmaaker & Bergmans (1981) have reviewed R. amplexicaudatus in detail,
with a summary of earlier classification, synonymies, and many measurements derived from
a relatively large number of specimens. These authors considered size to be the only major
subspecific character in the species and recognised three size groups from the Solomon
Islands, New Guinea, the Philippine Islands, Timor and Java. Of these, the smallest
examples were found to occur on the Solomon Islands, the largest in New Guinea and the
Philippines (these latter being mutually indistinguishable); specimens from Java, like those
from the Solomons, proved significantly smaller than specimens from New Guinea and the
Philippines. Examples from Timor were found to be generally a little larger than those from
Java, but not significantly so, and a little smaller on the whole than those from New Guinea
and the Philippines, although again the differences lacked significance. However, such speci-
mens were thought by Rookmaaker & Bergmans to have a greater similarity to examples
from New Guinea and the Philippines than to those from Java. On this basis they recognised
three subspecies, R. a. brachyotis (Dobson, 18770) from the Solomon and Bismarck Islands,
R. a. infumatus (Gray, 1 870) from Sumatra and Java east to Flores and possibly Alor Island,

106 J. E. HILL

and R. a. amplexicaudatus from Sumba Island, Timor and some smaller associated islands,
New Guinea, possibly from the Molucca Islands, and from the Philippines. Specimens from
Sulawesi were not allocated to subspecies: others from Borneo, Mentawei and Engano
Islands, Malaya, Thailand and Burma were referred to R. a. amplexicaudatus, but few
examples are available from the western part of the range. At the eastern limit, however,
Smith & Hood (1981) have suggested that the population on the Bismarck Islands (R. a.
brachyotis) is subspecifically separable from that on the Solomon Islands (R. a. hedigeri
Pohle, 1953), although the distinction is less marked than the divisions between these island
subspecies and R. a. stresemanni from New Guinea.

Considerations of relative size led Rookmaaker & Bergmans (198 1 ) to associate specimens
from Timor with those from New Guinea and thus to synonymise stresemanni with R. a.
amplexicaudatus. However, their account of specimens from Sulawesi and some of its
associated islands indicates that in some ways these unallocated examples are intermediate
between R. a. infumatus and Timorese specimens ofR. a. amplexicaudatus. A study of their
detailed tabulated measurements for these populations confirms this view: also of two
specimens (BM(NH) 98.11.3.20-21) from Alor Island provisionally allocated to R. a.
infumatus by Rookmaaker & Bergmans the male is similar in size to males from Timor, but
the female to females from Java. Rookmaaker & Bergmans also made a detailed
examination of the relation between condylobasal length and zygomatic width in some of the
populations of R. amplexicaudatus. Their diagrams show that while in these dimensions
Timorese specimens lie in the lower part of the range of variation of those from New Guinea
and the Philippines and are separated more distinctly from the Javanese population the
limited Sulawesian sample tends to bridge this interval. It is also evident that the majority of
specimens from New Guinea and the Philippines exceed Timorese and Sulawesian
specimens in one or both of these dimensions. There seems, therefore, at least as much to
justify the association of Timorese specimens (R. a. amplexicaudatus) with those from
Sumatra, Java and the Lesser Sunda Islands (R. a. infumatus) as with those from New
Guinea.

For these reasons stresemanni has been retained as a distinct subspecies in New Guinea
and possibly also in the Philippine Islands. Measurements by Rookmaaker & Bergmans
(1981) indicate that only small overall differences exist between these populations.
Philippine females have longer wing elements than the very limited New Guinea sample
examined and on the basis of a larger representation of females from New Guinea are
cranially rather smaller on the whole but with longer toothrows. Until more specimens are
available from the Molucca Islands, Sulawesi and the western part of the range of
R. amplexicaudatus from Borneo and Sumatra to Burma it seems appropriate to regard R. a.
amplexicaudatus as a valid link between the smaller R. a. infumatus and the larger R. a.
stresemanni.

Rousettus celebensis Andersen, 1 907
Rousettus celebensis Andersen, 1907a: 509. Mount Masarang, N Sulawesi, 3500 ft.

SPECIMENS EXAMINED. N Sulawesi: 99BM(NH) 78.964-967 Tangkopo, Batuangus, near Bitung (in
alcohol, BM(NH) 78.965 head, skin, others heads only; coll. A. M. Jones).

C Sulawesi: rfrf BM(NH) 81.1066-1068 Ganda Ganda, 1° 57' S, 121° 21' E; <f BM(NH) 81.1069
Sampalawa,21 km from Kolono Dale, c. 2° 00 ' S, 121° 20' E: 5 rfrf, 11 99 BM(NH) 81.1070-1085 R
Ranu, 1°51' 121° 30' E;2rfrf (yg.), 299 BM(NH)81.1086-1089 Songinbau, 1°46' S, 121°43' E;2 9$
BM(NH)81.1090-1091 Taronggo, 1° 44' S, 121° 40' E (all in alcohol; coll. B. H. Gaskell, 'Operation
Drake').

REMARKS. Andersen (1907, 1912) pointed out that this species had before then been
confused with R. brachyotis ( = R. amplexicaudatus brachyotis) and indeed earlier authors
also referred specimens since recognised as celebensis to R. amplexicaudicaudatus or to R.

INDO-AUSTRALIAN BATS 107

minor Andersen, 1907 (=R. amplexicaudatus infumatus, after Rookmaaker & Bergmans,
1981). Andersen's view (1912) that the alleged occurrence of R. amplexicaudatus (as R.
brachyotis) in Sulawesi probably rested on confusion with R. celebensis may well have led to
all Rousettus from that island being referred to the latter species, as Rookmaaker &
Bergmans (1981) suggest. These authors record specimens of R. amplexicaudatus from
Gorantalo and Talassa in Sulawesi and from the associated islands of Muna, Peleng and
Talisai: those from Peleng were referred originally to R. amplexicaudatus by Tate (1942c).
The two species may be distinguished by a number of features: celebensis has longer fur, a
furred notopatagium and much more densely pilose tibiae while the rostrum is relatively
longer with longer, more nearly parallel rather than convergent upper toothrows. The
dentition also provides good diagnostic features: the last upper premolar (pm4) is longer than
in amplexicaudatus and the molars both above and below are distinctly narrow, the crowns
of m{ and m2 clearly elongate, m^ in particular being longer than in amplexicaudatus.

Rookmaaker & Bergmans (1981) reported and examined a total of 28 Sulawesian speci-
mens (including BM(NH) 78.964-967) of R. celebensis, with 10 further examples from the
Sanghir ( = Sangihe) Islands and one from either the Talaud or Sanghir Islands: they do not
include the holotype or two others listed by Andersen (1912). Among this material there is
one extensive series, of 18 examples collected by G. Stein at Makassar. Specimens from
'Operation Drake' and others reported here therefore considerably extend the known
representation of the species. They agree closely with the accounts by Andersen (1907,
1912): one, BM(NH) 81.1068 is unusual in lacking the right upper molar (M2) and in having
only four instead of six cheekteeth in the left lower jaw, there being no trace of the last two
molars (m2, m3), while BM(NH) 8 1 . 1 075 and BM(NH) 8 1 . 1 09 1 are young adults with the last
molars (mf yet to erupt. Length of forearm in 8 adult rfc? 71-8-76-9 (74-9); in 14 adult
9972-5-80-2 (76-5). Males fall within the range of forearm lengths given by Rookmaaker &
Bergmans (198 1) for a slightly larger sample of male examples, but some females exceed the
upper limit of the female specimens measured by these authors. Andersen (1912), who may
have examined only dry skins, thought that the species might have a long tail, its length
probably about 20 mm. The length of the tail in 8 rfcf is 23-5-27-5 (25-3) and in 14 99
23-1-30-0(27-1).

DISCUSSION. I am unable to agree with Tate (1942c), who had seen only Andersen's descrip-
tion, that celebensis appears to be a member of the amplexicaudatus group, with narrower
molars, and who in fact suggested that celebensis might be a subspecies of R.
amplexicaudatus although listing it formally as a valid species. Koopman (1979) in discuss-
ing the status of stresemanni which he considers a subspecies of R. amplexicaudatus has
pointed out that celebensis is distinct, an opinion fully endorsed by Rookmaaker &
Bergmans (1981) who record both amplexicaudatus and celebensis from Gorontalo in N
Sulawesi.

Andersen (1912) recognised three subgenera in Rousettus, placing both amplexicaudatus
and celebensis in the nominate subgenus, distinguished by moderate deflection of the
braincase, the last upper premolar (pm4) not especially narrowed, its width about one third of
the palatal width between the fronts of pm4~4, the wing originating from the back of the first
toe, and by a distinct antitragal lobe. In contrast, the subgenus Stenonycteris was defined by
Andersen chiefly on the basis of a strongly deflected braincase, excessively narrow cheek-
teeth with the width of pm4 about one fifth of the palatal width between the fronts of pm4~4,
wings originating from the back of the second toe and obsolete antitragal lobe.

In many respects R. celebensis approaches the African R. lanosus Thomas, \906a
apparently the sole representative of Andersen's subgenus Stenonycteris. It has similarly
long, rather dense pelage extending on to the notopatagium, with long hairs on the forearms,
tibiae, interfemoral membrane and the underside of the lateral membrane; long thumbs and
long wings that tend towards those of the African species (wing indices of celebensis appear
in Table 1); the wing is inserted at or near the rear of the second toe; a small, reduced,
rounded antitragal lobe; the basicranial axis is somewhat deflected; the cheekteeth are gener-

108

J. E. HILL

Table 1 Wing indices (length of forearm = 1000) of 7 dd and
14 99 ofRousettuscelebensis.

Range

Mean

Thumb

dd

393-139

423

99

388-458

409

IIm

3d

444^80

458

99

427^71

446

II>

dd

106-125

117

99

93-128

108

IP-3

dd

116-142

133

(c.u.)

99

110-142

128

IIP

dd

651-669

658

99

636-694

660

III1

dd

455^69

462

99

438^74

455

IIP

dd

548-621

585

99

554-615

583

IVm

dd

631-652

637

99

617-659

635

IV

dd

330-356

344

99

321-359

340

IV2

dd

368-401

378

99

343-385

367

ym

dd •

623-651

639

99

615-645

637

V

dd

291-306

299

99

275-304

292

V2

dd

333-355

348

99

325-352

321

ally long and narrow, and, as in lanosus, the molars have little cuspidation. Andersen (1912)
considered that celebensis probably represented a modification of the R. amplexicaudatus
type, but it seems more appropriate to include this Sulawesian species with R. lanosus in
the subgenus Stenonycteris if this is to be recognised rather than in the nominate subgenus
where Andersen originally placed it.

Pteropus hypomelanus macassaricus Heude, 1 896

Pteropus macassaricus Heude, 1 896 : 1 77, footnote, pi. 5, fig. 4. Makassar, S Sulawesi.

SPECIMENS EXAMINED. C. Sulawesi: d BM(NH) 81.1092 Ganda Ganda, 1° 57' S, 121° 21' E- 99
BM(NH) 81.1093-1096 1 km NE of Tandiondo, 1° 45' S, 121° 17' E (all in alcohol, all except
BM(NH) 81.1092 heads only; coll. B. H. Gaskell, 'Operation Drake').

Pteropus alecto alecto Temminck, 1837

Pteropus alecto Temminck, 1837, 2 : 75. Menado, N Sulawesi.

SPECIMENS EXAMINED. C Sulawesi: rf, 4 99 BM(NH) 81.1097-1 101 1 km NE of Tandiondo, 1°45' S,
121° 17' E (in alcohol, heads only; coll. B. H. Gaskell, 'Operation Drake').

INDO-AUSTRALIAN BATS 109

REMARKS. Musser et #/.(1982) have demonstrated that among the several forms of Pteropus
described or reported from Sulawesi P. arquatus Miller & Hollister, 192 1 is in fact Acerodon
celebensis, as are the specimens from the northern part of the island recorded originally by
Tate (1942c) as P. argentatus. Others known from Sulawesi besides P. hypomelanus
macassaricus and P. alecto alecto are P. griseus mimus Andersen, 1908, P. caniceps dobsoni
Andersen, 1908 and P. personatusTemminck, 1825.

Pteropus conspicillatus Gould, 1 850

Pteropus conspicillatus Gould, 1 850 : 109. Fitzroy I, Queensland, Australia.

SPECIMENS EXAMINED. Papua New Guinea: rf, 9, 9 juv. BM(NH) 80.525-527 Lababia Cave, Lababia I,
Morobe, 7° 15 ' S, 147° 09 ' E (in alcohol, coll; B. H. Gaskell, 'Operation Drake').

REMARKS. Laurie & Hill (1954) listed P. c. conspicillatus from Papua New Guinea after
Matschie ( 1 899) who reported this subspecies from Bongu (5° 30 ' S, 1 45° 50 ' E) and Madang
(5° 13' S, 145° 48 'E): a second subspecies, P. c. chrysauchen Peters, 1862 was listed from
northwestern New Guinea after Tate (1942c) who recorded it from Geelvinck Bay.

These examples from Lababia Cave cannot be allocated positively to either of these
subspecies. The adult c? BM(NH) 80.525 has a uniformly dark head, the forehead, crown and
sides of the muzzle blackish, mixed especially on the forehead and on the muzzle anterior to
the eyes with buffy hairs. Beyond a slight band of buffy hairs there is no indication of the
paler eye rings characteristic of P. c. conspicillatus and the sides of the muzzle are not pale as
in this subspecies. The adult 9 BM(NH) 80.526, however, although predominantly black on
the forehead, crown and cheeks has relatively distinct paler buffy eye rings and the sides of
the muzzle above the mouth and the corresponding areas along the lower jaw are distinctly
ochraceous buff. Moreover, the black of the crown extends forward as an acutely triangular
patch whose apex intrudes between the pale superciliaries to the base of the rostrum. The
adult male agrees therefore with P. c. chrysauchen, the adult female with P. c. conspicillatus.
The collections of the British Museum (Natural History) include two further specimens (c?, 9
22.1.22.1-2) from Simbang, on the east coast of Papua New Guinea (probably at 3° 35 S,
147° 43' E) that are referable to P. c. conspicillatus although even in these the facial
markings of the male are a little less pronounced than in the female example. Length of fore-
arm in specimens from Lababia Cave (c?, 9) 1 8 1 , 1 79.

Styloctenium wallacei (Gray ', 1866)

Pteropus wallacei Gray, 1 866a : 65, fig. 1 . Makassar, S Sulawesi.

SPECIMENS EXAMINED. C Sulawesi: 9 (yg. ad.), cf BM(NH)81.1 102-1103 R Ranu, 1°51 ' S, 121° 30' E
(in alcohol): cf (yg. ad.) BM(NH) 81.1 104 (in alcohol), 9 BM(NH) 81.1 105 (skin, skull, skeleton), 9
(yg.) BM(NH) 81.1 106 (in alcohol) Tambusisi Damar, Mt. Tambusisi, c. 4000 ft, 1° 39 ' S, 121° 22 '
E (all coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Although this distinctive bat is by no means common in collections there is a
number of records (Jentink, 1883; Matschie, 1899; Andersen, 1912) from N Sulawesi and
Tate (1942c) has reported a long series from Malenge in the Togian Islands in the Gulf of
Gorantalo. There is also a hitherto unreported specimen (cf MZB 12671) from Titaeli,
Minahassa, in the collections of the Museum Zoologicum Bogoriense, Bogor.

HO J. E. HILL

Adults obtained by Operation Drake display very clearly the badger-like white facial
markings characteristic of the species: the white shoulder patches although always present
are rather small in the two male examples, one not quite fully adult.

External measurements of an adult d and 9 (BM(NH) 81.1103, 81.1105): length of
forearm 96-6, 95-5; thumb (c. u.) 43-2, 42-3; Im 12-2, 12-0; I1 23-7, 234; IIm 50-7, 51-5; II1
12-4, 12-3; II2-3 (c. u.) 12-0, 11-3; IIIm 69-0, 67-2; III1 51-3, 51-4; III2 69-0, 63-7; IVm 69-4,
68-6; IV 39-7, 38-5; IV2 40-6, 40-4; Vm 7 1 -7, 7 1 -8; V 3 1 -4, 3 1 -5; V2 33-8, 33-9; tibia 42-4,—.
Cranial measurements of an adult 9 (BM(NH) 81.1105): total length of skull to gnathion
51-8; condylobasal length 48-5; condylocanine length 444; length front of orbit-tip of nasals
16-8; palatal length 28-0; length palation-incisive foramina 23-8; length palation-basion
18-3; lachrymal width 10-5; least interorbital width 6-8; least postorbital width 5-9;
zygomatic width 28-3; width of braincase 19-7; mastoid width 17-9; orbital diameter 114;
c'-c1 (crowns) 9-5, (alveoli) 8-8; m'-m1 (crowns) 13-9, (alveoli) 12-7; c'-c1 (internally,
cingula) 5- 1 ; pm4-pm4 (internally) 6-9; width of mesopterygoid fossa 74; c-m2 19- 1 ; length of
complete mandible from condyles 364; length right ramus from condyle 38-1; coronoid
height 19-8; c-m2 19-9.

Dobsonia viridis (?) viridis (Heude, 1896)

Cephalotes viridis Heude, 1 896 : 1 76, footnote, pi. 5, fig. 1 . Kei Is.
Dobsonia viridis umbrosa Thomas, 1910a : 384. Ceram I.

SPECIMENS EXAMINED. C Sulawesi: 9, cT BM(NH) 81.1107-1108 R Ranu, 1° 51' S, 121° 30' E (in
alcohol, skulls extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These are apparently the first of viridis to be reported from Sulawesi: it occurs
otherwise on the Kei Islands and on the islands of Amboina, Buru, Ceram and Banda.
Thomas (1910#) separated specimens from Ceram as D. v. umbrosa to which Andersen
(1912) subsequently referred others from Buru and Amboina: this author (p. 825) remarked
that the reference by Thomas to a number of specimens from 'Aru' is a misprint for Buru but
in his personal copy of his paper, now in the Library of the British Museum (Natural
History), Thomas has corrected Aru to Kei [Islands]. The collections in London include
several specimens collected by W. Stalker on the Kei Islands but only a single immature
from Buru, obtained by this collector. A small average difference in colour separates
umbrosa from viridis of the Kei Islands and the two are synonymised in the listing by Laurie
& Hill (1954). A closely related form, D. crenulata Andersen, 1909 occurs on the Halmahera
group of islands to the northwest of New Guinea and has been reported recently from the
Sanghir ( = Sangihe) Islands and on the Togian Islands. It is separated from D. viridis chiefly
on account of its greater size and larger teeth.

Specimens from Sulawesi have the characteristic dentition of the viridis group of Andersen
(1909, 1912) in which m1 has a well-developed antero-internal ledge and pm| and m} have
cuspidate labial and lingual ridges. The surface cusps or ridges of mj and m2 are also strongly
developed and prominent, of pm4 much less so. The collector, B. H. Gaskell remarks of the
coloration of the adult that when freshly obtained the dorsal pelage was a light dull green in
hue, yellower towards the rear and flanks, with the upper surface of the head yellowy green
grey, sharply divided on the neck from the colour of the body. The ventral surface was
generally similar in colour to the lower back but medially light orange. The indefinite
greenish tinge in the pelage appears characteristic of D. viridis: Goodwin (1979) also
commented upon the unusual olive green colour of adults of D. peronii peronii (Geoffroy,
1810#) when living. The colour faded rapidly in specimens preserved as dry skins, in
alcohol, or in formalin.

INDO-AUSTRALIAN BATS 1 1 1

In size both Sulawesian examples are similar to or exceed the largest of Moluccan speci-
mens of viridis and their cheekteeth, especially of the male, are generally longer and slightly
larger, particularly pm4, mj and m2. To some extent, therefore, they approach crenulata but
their canines are smaller and their cheekteeth generally narrower, like those of viridis from
the Moluccas.

Measurements of an adult 9 and c? (BM(NH) 81.1107-1108): length of forearm 113-5,
125-4; IIIm 71-6, 77-1; IVm 65-4, 70-0; Vm 66-2, 71-0; total length of skull to gnathion 48-2,
50-1; condylobasal length 46-2, 47-8; condylocanine length 45-7, 47-6; rostral length 15-6,
15-3; length front of orbit-tip of nasals 12-2, 1 1-9; palatal length 24-1, 25-0; length palation-
incisive foramina 21-1, 22-1; length palation-basion 19-1, 19-6; lachrymal width 11-8, 11-6;
least interorbital width 8-4, 8-2; least postorbital width 7-5, 6-4; zygomatic width 29-4, 31-1;
width of braincase 19-7, 19-7; mastoid width 18-4, 18-3; orbital diameter 9-9, 9-9; c'-c1
(crowns) 9-3, 9-5, (alveoli) 8-7, 8-7; m'-m1 (crowns) 14-6, 15-1, (alveoli) 13-8, 14-4; m2-m2
(crowns) 12-8, 12-9, (alveoli) 12-5, 12-6; c'-c1 (internally, cingula) 3-4, 3-7; pm4-pm4
(internally, 7-6. 7-7; width of mesopterygoid fossa 6-0, 5-8; c-m2 (crowns) 18-8, 19-8,
(cingula) 18-4, 19-4; length complete mandible from condyles 36-5, 37-4; length right ramus
from condyle 37-7, 38-9; coronoid height 20-3, 21-8; c-m3 (crowns) 20-0, 20-7, (cingula) 19-8,
20-4. Length/width of: c1 3-86/2-48, 3-82/2-54; pm3 3-94/2-88, 4-02/3-22; pm4 4-02/2-72,
4-32/3-14; m1 5-15/2-44, 5-17/2-83; m2 2-34/1-56, 2-37/1-61; c, 2-64/2-17, 2-65/2-22; pm2
1-40/1-34, 1-30/1-45; pm3 3-72/2-31, 3-90/2-52; pm4 4-02/2-53, 4-22/2-62; m1 3-99/2-15,
4-21/2-35; m2 3-30/2-08, 3-31/2-16; m3 1-90/1-31, 1-85/1-43.

DISCUSSION. The viridis group of Dobsonia as proposed by Andersen (1909, 1912) includes
besides D. viridis and D. crenulata the further species D. praedatrix Andersen, 1909 from the
Bismarck Archipelago and D. inermis Andersen, 1909 and D. nesea Andersen, 1909 from
the Solomon Islands. Since Andersen wrote a number of changes have been made to this
classification. Troughton (1936) regarded nesea as a subspecies of D. inermis while Rabor
(1952) in describing D. viridis chapmani from Negros Island in the Philippine Islands treated
crenulata as a further subspecies of viridis. Pohle (1953) considered crenulata and praedatrix
to be subspecies of D. viridis and inermis (including nesea) probably so. Laurie & Hill (1954)
retained the arrangement of Andersen but united nesea with inermis as a subspecies.

More recently, Bergmans (1975) discussed the viridis group in some detail, describing a
further species, D. beauforti from Waigeo Island in the northern Moluccas, and rejecting the
views of Rabor and Pohle. Since then he has (1978) examined the group yet further and has
established that chapmani does not fulfil the appropriate diagnostic criteria but seems more
likely to belong to the moluccensis group. Moreover, Bergmans is now of the opinion that
praedatrix and inermis (including nesea} differ sufficiently from the two (sic) other species
that they should form one or two species groups by themselves. Dobsonia beauforti is said to
be morphologically allied to D. viridis but is appreciably smaller in forearm and skull
measurements, in size much like D. inermis from the Solomon Islands. No specimens of D.
beauforti have been examined.

Only the holotype (BM(NH) 60.8.26.2) of crenulata is available for study in London.
However, De Jong & Bergmans (1981) have reviewed this taxon, recording it for the first time
from the Sanghir ( = Sangihe) Islands and from the Togian Islands, and have provided
measurements of a number of specimens. These authors refer to earlier views (Rabor, 1952;
Pohle, 1953) that crenulata is a large subspecies of D. viridis, but regard the union of the two
into the same species as premature, since specimens of each sex from any one population
are as yet insufficient to establish its range of size variation. The specimens from Sulawesi
approach and in some respects equal crenulata in external and cranial dimensions, but on
the whole have slightly narrower cheek teeth similar in width to those of viridis. They have
smaller canines than the subadult holotype of crenulata, corresponding closely to those of
viridis. This limited sample therefore suggests strongly that viridis and crenulata are likely
to prove conspecific, a view provisionally adopted here.

112 J. E. HILL

Dobsonia praedatrix Andersen, 1 909

Dobsonia praedatrix Andersen, 1909 : 532. Duke of York I, Bismarck Archipelago.

SPECIMENS EXAMINED. Bismarck Archipelago: 9, rf, 9 BM(NH) 69.304-306 Kareeba Plantation,
Keravat, New Britain, 4° 1 8 ' S, 1 52° 01' E (skins, skulls; coll. J. I. Menzies).

REMARKS. These relatively recently collected specimens confirm the diagnostic features used
by Andersen ( 1 909, 1 9 1 2) in separating praedatrix from viridis and crenulata. As pointed out
by Andersen, the rostrum is heavily built and is broader and more massive, with a consider-
ably wider interorbital region. The teeth of the male example are similar in size to those of
the holotype but in the female specimens are generally slightly smaller, overall much as the
teeth of viridis. Andersen (1912) observed correctly that pm| in the holotype of praedatrix,
however, are practically as large as in crenulata but in each of the specimens reported here
these teeth are smaller than those of crenulata and are nearer in size to pm| of viridis. The
labial and lingual longitudinal ridges of pmf and m{ are faintly cuspidate in BM(NH) 69.305
but scarcely if at all cuspidate in BM(NH) 69.304 and 69.306. The longitudinal ridges of
these teeth in the holotype are slightly more cuspidate, but the teeth are little worn. As in
viridis, crenulata and in the holotype of praedatrix mj and m2 have well developed surfacial
cusps or ridges: the surface cusp of pm4 is low and very undeveloped.

The three specimens closely resemble the holotype in colour. Dorsally, the shoulders and
neck are brownish, strongly tinged with black in the male, the head blackish brown, this
darker colour extending medially down the nape and neck in a narrow line; the ventral
surface of the body is brownish drab, medially with a faint wash of brighter tawny olive, the
underside of the neck and chin with a much sparser covering of longer, brownish hairs.

The male example is larger cranially than the two female specimens, adding support to
Bergmans (1975) who suggested that sexual dimorphism in size might well occur in some at
least of Dobsonia. Measurements of d1 BM(NH) 69.305, 9969.304, 69.306 in that order:
length of forearm 116-3, 117-3, 112-2; total length of skull to gnathion 50-6, 48-3, 48-5;
condylobasal length 47-6, 45-6, 45-5; condylocanine length 47-3, 45-2, 45-0; length front of
orbit-tip of nasals 14-5, 13-3, 13-1; palatal length 25-5, 23-7, 24-2; length palation-incisive
foramina 22-6, 21-7, 21-7; length palation-basion 19-1, 18-8, 18-5; lachrymal width 13-8,
12-8, 12-9; least interorbital width 10-1, 9-7, 10-0; least postorbital width 8-2, 7-7, 7-9;
zygomatic width 31-4, 30-0, 30-4; width of braincase 20-4, 19-4, 19-6; mastoid width 19-0,
17-8, 17-8; orbital diameter 10-8, 10-8, 10-6; c'-c1 (crowns) 9-7, 9-4, 9-4, (alveoli) 9-2, 8-8,
8-8; m'-m1 (crowns) 15-5, 14-7, 14-1, (alveoli) 14-7, 14-0, 1 3-5; c'-c1 (internally, cingula) 4-0,
4-1, 3-9; pm4-pm4 (internally) 8-0, 7-8, 7-5; width of mesopterygoid fossa 5-8, 5-9, 5-5; c-m2
20-0, 18-4, 18-5; length complete mandible from condyles 37-8, 36-0, 36-0; length right
ramus from condyle 39-2, 37-2, 37-5; coronoid height 21-6,21-7,21 -6; c-m3 21-1,19-2,19-3.
Length/width of cheekteeth: pm3 4-41/3-08, 4-04/2-81, 4-06/2-84; pm4 4-22/3-14, 3-97/2-82,
3-95/2-83; m1 5-31/2-74, 4-73/2-61, 4-70/2-72; m2 2-45/1-64, 2-06/1-49, 2-15/1-42; pm2
1-39/1-55, 1-13/1-43, 1-24/1-44; pm3 4-09/2-61, 3-87/2-39, 3-84/2-32; pm4 4-42/2-78,
3-89/2-48, 4-01/2-46; m, 4-14/2-46, 3-60/2-21, 3-91/2-20; m2 3-31/2-37, 2-98/2-02,
2-96/2-01 ;m3 1-95/1-55, 1-79/1-46, 1-60/1-33.

DISCUSSION. Examination of these adult examples confirms that Bergmans (1975) correctly
rejected the suggestion by Pohle (1953) that praedatrix should be allied subspecifically to D.
viridis, but I am less convinced that it should be removed from the viridis group as the
former author (1978) has since suggested. If dental characters are to remain the chief criteria
by which the species of Dobsonia are classified, then praedatrix must be included in the
viridis group as Andersen (1909, 1912) envisaged it, except that the degree of cuspidation of
the longitudinal ridges of pm| and m{ is less than is general in the group or is sometimes
virtually absent. In this respect praedatrix approaches the peronii group of Andersen (loc.
cit.) in which these ridges are simple, but m1 reputedly lacks the well marked antero-internal
basal ledge characteristic of the viridis group. The limited material of D. peronii in London

INDO-AUSTRALIAN BATS 1 1 3

indicates that in this species m1 has at least a small antero-internal ledge and that the
character may not be as emphatic as Andersen implied, although the ledge is much more
developed in D. viridis and its allies. As Bergmans (1978) suggested, D. peronii, by current
classification the sole member of the peronii group, may be more closely related to the viridis
group than to any other but as this author pointed out, D. peronii differs from D. viridis and
from its allies in the outline of the rostrum, which is lower, relatively longer and curves
downward less abruptly from the braincase. Bergmans also remarked that within the viridis
group peronii comes closest to the viridis-crenulata-beauforti (sub)group (sic) but dentally it
seems to approach more closely to praedatrix. The greenish coloration of D. viridis when
alive also appears in D. peronii peronii according to Goodwin (1979).

Bergmans (1978) advanced the opinion that the inclusion of praedatrix from the Bismarck
Archipelago and inermis (including nesea) from the Solomon Islands in the viridis group
indicated a disregard of zoogeographical considerations. I find no conviction in this assertion
since the group as presently constituted occupies the arc of islands to the west, northwest,
north and northeast of New Guinea, perhaps without interruption since some remain
zoologically poorly known. A biogeographic model of this nature was suggested by Smith &
Hood (1981). Indeed, if D. peronii is regarded as a member of the viridis group it then
includes a succession of island species and subspecies extending from Nusa Penida (near
Bali) in the west to the Solomon Islands in the east, but apparently excluding New Guinea.

These considerations suggest that the peronii and viridis groups of Andersen (1909, 1912)
might be merged to form a peronii group that includes D. peronii, D. viridis and its close
allies, D. praedatrix and D. inermis. It can be summarized:

D. peronii grandis Bergmans, 1978 Nusa Penida I,

Sumbawa I.

D. peronii sumbana Andersen, 1 909 Sumba I.

D. peronii subsp. W. Flores I, Alor I,

(see Bergmans, 1 978) Wetar I, Babar I.

D. peronii peronii (Geoffrey, 1810#) Timor I.

D. viridis (?) viridis (Heude, 1 896) Sulawesi, Amboina I,

Buru I, Ceram I,
Banda Is, Kei Is.

D. viridis (?) crenulata Rau I, Morotai I,

Andersen, 1909 Halmahera I, Ternate I,

Batchian I, Togian Is,
Sanghir ( = Sangihe) Is.

D. beauforti Bergmans, 1 975 Waigeo I.

D. praedatrix Andersen, 1 909 New Britain, New Ireland,

Duke of York I.

D. inermis nesea Andersen, 1 909 N and WC Solomon Is:

Shortland, Alu, Ghizo,
Rubiana, Bougainville,
New Georgia.

D. inermis inermis Andersen, 1 909 S and EC Solomon Is:

San Christoval, Ugi,
Ysabel, Rennell.

Phillips (1968) considered nesea a synonym of inermis, but McKean (1972) retained it as a valid
subspecies on the basis of Troughton's (1936) assertion that inermis is darker in colour.

The occurrence of D. viridis in Sulawesi and (as crenulata) on the Togian and Sanghir
Islands also has some bearing on Bergman's (1978) view of possible former distribution
routes for Dobsonia towards the Lesser Sunda Islands. He suggested that D. peronii and D.
moluccensis or their ancestors in these islands (there is only one local record of the latter,
from Semau ( = Samoa) Island, near Timor) might have originated from New Guinea
(moluccensis} or from the Aru, Kei or Timorlaut Islands (peronii), possibly moving along

114 J. E. HILL

former land bridges, or alternatively along a more northerly route through the southern
Moluccas and Sulawesi. A species of the moluccensis group, D. exoleta, has been known
from Sulawesi for many years and Bergmans pointed out that the concept of the more
northerly route implied that it is the closest living relative of D. peronii, although he
considered exoleta to be less closely related to peronii than to the members of the viridis
group. If Bergman's views are accepted the presence of D. viridis in Sulawesi resolves this
apparent paradox.

Dobsonia moluccensis moluccensis (Quoy & Gaimard, 1830)

Hypoderma moluccensis Quoy & Gaimard. 1830.1 : 86, Atlas, pi. 11. Amboina.

SPECIMENS EXAMINED. Molucca Is: BM(NH) 75.2140 Lihura limestone caves, near Ruhuwa, SC Ceram
I (crania, mandible, fragments, teeth; coll. R. F. Ellen).

REMARKS. This cave material compares favourably with D. m. moluccensis from Buru,
Amboina and Ceram. Length of c-m2 (alveoli) in two examples 23- 1 , 23-5.

Dobsonia moluccensis magna Thomas, 1905

Dobsonia magna Thomas, 1905# : 423. Tamata, Mambare R, Papua New Guinea, 100 ft.

SPECIMENS EXAMINED. Papua New Guinea: 9 BM(NH) 69.307 Brown R, near Port Moresby, c. 9° 27 ' S,
147° 08 ' E; d BM(NH) 69.308 Tupuselaia ( = Tupuselei), 9° 33 ' S, 147° 19 ' E (both skins, skulls; coll.
J. I. Menzies): d1 BM(NH) 73.1969 Kairiru Ridge, centre of Kairiru I, near Wewak, East Sepik,
c. 2000ft; <5 BM(NH) 73.1970 Victoria Bay, NW end of Kairiru I (both in alcohol; coll. Aberdeen
University Exploration Society Expedition to New Guinea, 1973); BM(NH) 78.202-209 ? Near Mt.
Hagen, Western Highlands (crania, damaged); BM(NH) 78.210-211 Baiyer R, Western Highlands
(mandibles, damaged); BM(NH) 78.212-249 Tuman, Kubor Range, Western Highlands (crania,
damaged); BM(NH) 78.250-255 Upper Lai Valley, Southern Highlands (crania, mandibles, damaged);
9 BM(NH) 78.854 About 10 km S of Madang(in alcohol); d BM(NH) 78.855 About 10 km S of Madang,
c. 40 m (skin); rf BM(NH) 78.856 Baiyer R, c. 50 km NW of Mt. Hagen, Western Highlands, 1300 m (in
alcohol) (all coll. P. A. Morris); BM(NH) 79.2015 (skin), BM(NH) 78.3004 (skull) Haelaelinga
Settlement, on Was ( = Wage) R, Nipa, Southern Highlands (coll. P. Sillitoe); rf, 9 BM(NH)
80.528-529 Buso, Morobe, 7° 17 ' S, 147° 08 ' E (in alcohol; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. The cranial and mandibular material BM(NH) 79.202-255 was obtained from
kitchen middens: most show signs of heating or burning. The rear of the cranium in such
specimens has been broken open to allow the extraction of the brain, a circumstance noted
by Menzies (1977) of material from similar accumulations at the Kiowa and Yaku rock
shelters.

Dobsonia exoleta Andersen, 1909

Dobsonia exoleta Andersen, 1909 : 531, 533. Tomohon, Minahassa, Sulawesi.

SPECIMENS EXAMINED. C Sulawesi: cf BM(NH) 8 1 . 1 1 09 Ganda Ganda, 1 ° 57 ' S, 1 2 1 ° 2 1' E (in alcohol,
skull extracted); <5 BM(NH) 81.1238 Tapu Waru, 1° 51' S, 121° 22' E (in alcohol) (both coll. B. H.
Gaskell, 'Operation Drake').

REMARKS. The relatively unworn dentition of BM(NH) 81.1109 agrees closely with the
holotype (BM(NH) 99. 1 0. 1 .4) and with the account by Andersen (1912). The antero-internal
corner of pm4 is developed into a conspicuous ledge with elevated rim; centrally the ledge is
raised into a small cusp; pm3 has a similar, narrower ledge; the antero-internal corner of m1 is

INDO-AUSTRALIAN BATS 1 1 5

low and platform-like but little differentiated except by a shallow antero-internal notch in
the longitudinal ridge; a slight antero-internal basal ledge in pm3, rather more developed in
pm4; m, with simple lingual ridge, no trace of an antero-internal ledge or cusp. There is a
well developed posterior basal ledge on pm3, a similar but slightly narrower ledge on pm4;
pm3 has likewise a slightly wider posterior basal ledge than pm4. Median surface cusps or
ridges are well developed in mj and m2, in the latter the ridge extending uninterruptedly
through the length of the tooth; there is a low surface cusp on the posterior face of pm4 that is
absent from the more eroded dentition of BM(NH) 81.1238.

Measurements (rf BM(NH) 81.1 109, rf 81.1238, skull of 81.1 109): length of forearm 1 16-8,
1 16-7; total length of skull to gnathion 5 1 -7; condylobasal length 49-9; condylocanine length
49-8; length front of orbit-tip of nasals 14-4; palatal length 25-8; length palation-incisive
foramina 23-4; length palation-basion 20-8; lachrymal width 12-4; least interorbital width
8-4; least postorbital width 7-0; zygomatic width 31-8; width of braincase 20-3; mastoid
width 19-5; orbital diameter 11-1; c'-c1 (crowns) 10-5, (alveoli) 9-7; m'-m1 (crowns) 16-9,
(alveoli) 15-9; c^c1 (internally, cingula) 4-1; pm4-pm4 (internally) 8-2; width of
mesopterygoid fossa 6-1; c-m2 22-0; length complete mandible from condyles 39-7;
length right ramus from condyle 4 1 -0; coronoid height 21-2; c-m3 23-3.

DISCUSSION. Dentally D. exoleta agrees with D. moluccensis from the Molucca Islands, New
Guinea and from some of its associated islands rather than with D. peronii from the Lesser
Sunda Islands. The Sulawesian species differs most conspicuously from peronii in its lack of
differentiation of the antero-internal corner of m, into a distinct cusp or small ledge, the
inner or lingual ridge of the tooth being perfectly simple and lacking any notch or division.
Andersen (1912) drew attention to a number of similarities between exoleta and D.
pannietensis (De Vis, 1905) from the Trobriand Islands, which he thought differed from
exoleta only in smaller size and in the lack of surface ridging on mp so considering the two
taxa closely related. Later, Thomas (1914^) in describing D. anderseni remarked that this
form from the islands of Manus and Ruk in the Bismarck Archipelago was intermediate in
size between exoleta and moluccensis, an opinion confirmed to some extent by Bergmans
(1979) who has examined specimens from a number of other islands in the Archipelago. As
in pannietensis the dentition of anderseni is like that of moluccensis. Laurie & Hill (1954)
recognised exoleta as a distinct species but listed both pannietensis and anderseni as sub-
species of D. moluccensis. More recently, De Jong & Bergmans (1981) reviewed known
specimens of exoleta in some detail and considered it to be a distinct species related in dental
morphology to chapmani Rabor, 1952 from the Philippines, moluccensis, anderseni and
pannietensis.

Bergmans (1975, 1978, 1979) has discussed and re-examined pannietensis and anderseni.
He concluded (1975, 1979) that both should be considered specifically distinct on account of
their smaller size when compared with D. moluccensis moluccensis and D. m. magna but it
is not clear to what extent they differ from each other. This author (1975) suggested that to
consider all of these conspecific involved the acceptance of an exceptional size range within
one species, although (1979) he has himself accepted a considerable range of size among
specimens from the islands (Louisiades, D'Entrecasteaux, Trobriand, Woodlark) immedi-
ately to the east of New Guinea that he considers all referable to pannietensis. Examination
of the measurements provided by Bergmans (1979) suggests that some at least of the speci-
mens that he refers to pannietensis are similar in size to anderseni: indeed, they overlap the
measurements given by this author (1975, 1979) for the latter, although admittedly sexual
differences in size may be involved. Moreover, Koopman (1979) in recording specimens
from some of the small islands (Karkar, Bagabag, Umboi) off northeastern New Guinea and
from the Bismarck Archipelago suggested that populations intermediate between magna and
anderseni are to be found on Karkar and Umboi. This author has since (1982) discussed the
question in some detail and while retaining anderseni as a subspecies of D. moluccensis
considered pannietensis specifically valid, at least until the genus is fully revised. Insufficient
specimens are available in London to determine the matter definitively, but for the present I

116 J. E. HILL

am unconvinced that anderseni and pannietensis should be considered species distinct from
moluccensis: Bergmans (1978) himself admits that both are more closely related to
moluccensis than to other species of the genus.

Bergmans (1979) provided a brief discussion of exoleta and compared it with D.
moluccensis pannietensis, drawing attention to a number of differences. Of these, the
relatively narrower interorbital and postorbital regions of exoleta also separate it (in the
limited sample available) from D. m. anderseni, D. m. magna and D. m. moluccensis. For
the present, therefore, I retain it as a distinct species and agree with Andersen (1912) in
regarding it as the Sulawesian representative of the moluccensis group. The species is known
to occur (De Jong & Bergmans, 1981) on Sulawesi, on the Togian Islands (Malenge) in the
Gulf of Gorantalo, and on Muna Island.

Dobsonia minor (Dobson, 1879)

Cephalotes minor Dobson (1878), 1879 : 875. Amberbaki, NW New Guinea.

SPECIMENS EXAMINED. Papua New Guinea: 9 9 BM(NH) 73.1971-1973 Rauit, 3° 36' S, 142° 15' E (in
alcohol); BM(NH) 74.337-338 Near Rauit (skulls) (all coll. Aberdeen University Exploration Society
Expedition to New Guinea, 1973).

Cynopterus brachyotis brachyotis (Muller, 1838)

Pachysoma brachyotis Muller, 1838 : 146. R Dewei, Borneo.

SPECIMENS EXAMINED. Burma: 3 cTrf, 4 99 BM(NH) 78.64-70, 99 BM(NH) 78.151-153 British
Embassy Residence Compound, Rangoon (in alcohol; coll. D. W. & G. Walton).

N Sumatra: dd, 9 BM(NH) 81.675-677 Bohorok R, near Bukit Lawang, Langkat Reserve, Gunung
Leuser Reserve complex (in alcohol; coll. R. Aveling).

S Sumatra: dd BM(NH) 78.1114-1115 Banda hurip, Pulas District, S Lampong; (?), 9 BM(NH)
78.1116-1117 Asahan, Jabung District, Lampong; 9 BM(NH) 78.1118 Lebung Dadup, Asahan,
Jabung District, Lampong; 9 BM(NH) 78.1 1 19 Sukaraja tiga, Sukadana District, Lampong; d BM(NH)
78.1120 Pring Kumpul, Pring Sewn District, Lampong; 3d BM(NH) 78.1122-1123 Air Nanigan,
Pulau Penggung District (all skins, skulls; presented by Museum Zoologicum Bogoriense).

N Sulawesi; d (?), (?) BM(NH) 78.968-970 Tangkopo Batuangus, near Bitung (in alcohol; coll.
A. M. Jones).

C Sulawesi: (?) BM(NH) 79.2333-2337 Shore of Lake Matano, c. 5 km W of Soroako (in alcohol,
very bad condition; coll. P. Holmes); 21 dd (2 yg.), 1 0 9 9 , neonate BM(NH) 81.1 006- 1 036, R Ranu, 1 °
51' S, 121° 30' E:2 dd, 5 99 (2 yg.) BM(NH) 8 1.1037-1044 GandaGanda, 1° 57' S, 121° 21' E (all in
alcohol; coll. B. H. Gaskell, 'Operation Drake').

Banggui Archipelago: cTBM(NH) 81.1045 Potil Besar 1; rfdBM(NH) 81.1046-1048 Kelara, Besar 1
(all in alcohol; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Although C. brachyotis is a relatively commonly collected bat in southeastern
Asia these specimens are of especial interest since those from Rangoon appear to be the first
recorded from Burma and the furthest west the species has been reported on the Asian
mainland while there have been few previous records from Sumatra and Sulawesi.
Specimens of C. brachyotis from southern Sumatra are from an area that proves to be of
particular taxonomic interest when the status of its congeners sphinx and titthaecheileus is
considered. In size all of these specimens agree well with the extensive representation of C. b.
brachyotis in the British Museum (Natural History) which suggests a range of forearm length
of 57-66 for this subspecies over its range from Burma and Indochina to Sumatra and
Sulawesi. Specimens from southern Sumatra with length of forearm 61-0-65-7 and m'-m1
(crowns) 8-2-9-0 approach the weakly defined subspecies C. b. javanicus Andersen, 1910
from Java but fall within the limits given by this author for C. b. brachyotis.

INDO-AUSTRALIAN BATS 1 1 7

DISCUSSION. Cynopterus minor Revilliod, 1911. The status of this nominal taxon must
remain uncertain for the present. It is known so far apparently only from the holotype from
Lambuja, SE Sulawesi, a small individual with a forearm length of 53 that may be a young
adult or unusually small example of C. brachyotis. It seems unlikely, however, that Revilliod
should fail to recognise it as such.

In addition the name minor poses an involved nomenclatorial problem. It was proposed
by Lyon in 1 908 as Niadius minor for the taxon subsequently recognised as the Sumatran
representative of Cynopterus horsfieldi by Andersen (1912) who considered Niadius Miller,
1906 a synonym of Cynopterus. As a result, Andersen (1912 : 827) regarded Cynopterus
horsfieldi minor (Lyon, 1908) to be preoccupied by the combination Cynopterus
(Cynonycteris) minor used by Trouessart (1878) for the species otherwise known as
Rousettus minor (Dobson, 1873<2,6), as it is today. In rejecting minor of Lyon, 1908 on this
account Andersen proposed Cynopterus horsfieldi lyoni in its place, a substitute that
subsequently (Robinson & Kloss, 1918, 1919; Chasen, 1940) came into use. Hill (196 la)
reviewed the circumstances surrounding this situation and on the basis of verbal advice in
1960 from the International Commission for Zoological Nomenclature that secondary
homonyms are not to be permanently rejected concluded that since Cynopterus
(Cynonycteris) minor as used by Trouessart referred to a Rousettus the name minor as
proposed by Lyon in 1908 should become again the valid subspecific epithet for the
Horsfield fruit bat of Sumatra and Malaya.

However, the current International Code of Zoological Nomenclature provides (Art. 59,
(b), (i)) that a junior secondary homonym rejected before 1961 is permanently rejected and
cannot be restored unless the employment of the replacement name is contrary to existing
usage. Thus Cynopterus horsfieldi lyoni Andersen, 1912 must become the valid name for the
Sumatran and Malayan representatives of C. horsfieldi. At the same time the Code (Art. 59,
(b), (ii)) states that if the secondary homonymy has been overlooked or the junior name not
replaced, and the taxa in question are no longer congeneric, the junior name is not to be
rejected, even though one name was originally proposed in the current genus of the other.
Therefore Cynopterus minor Revilliod, 1911, having by these rulings no competitor within
Cynopterus (minor of Lyon, 1908 having been rejected and replaced by lyoni of Andersen,
1912) remains a valid name in this genus, and is unaffected by the combination Cynopterus
(Cynonycteris) minor used by Trouessart (1 878).

Cynopterus sphinx angulatus Miller, 1 898

Cynopterus angulatus Miller, 1 898 : 3 1 6. Trang, S Thailand.

SPECIMENS EXAMINED. S Sumatra: 99 BM(NH) 78.1112-1113 Wai Miring, Kota Agung District,
Lampong; rf BM(NH) 78.1121 Pring Kumpul, Pring Sewn District, Lampong; 5<fd, 2 99 BM(NH)
78.1124-1130 Air Naningan, Pulau Panggung District (all skins, skulls; presented by Museum
Zoologicum Bogoriense).
Krakatau 1, 6° 10' S, 105° 26 ' E: cf BM(NH) 74.240 (in alcohol, skull extracted; coll. G. Lincoln).

REMARKS. Hill & Thonglongya (1972) discussed the affinities of angulatus and considered it
to be a subspecies of C. sphinx (Vahl, 1797) rather than of C. brachyotis with which (after
Andersen, 1912) it had been previously associated by the majority of authors. According to
Andersen (loc. cit.) angulatus occurs as far to the northwest as northern Burma and Assam:
however, a much wider representation of C. sphinx from Burma and Thailand than was
available to Andersen suggests that Burmese specimens (length of forearm (24) 68-3-74-5)
should be referred to the larger C. s. sphinx and that those from Thailand (length of forearm
(12) 64-9-73-7) represent the slightly smaller C. s. angulatus. Dammerman (1938) records
specimens from Krakatau Island and from the nearby island of Verlaten as angulatus, with
which their size (length of forearm 69-72, greatest length of skull 30-3-33-2, c-m1 10-1 1-2)
clearly associates them. These islands apparently lie at the known limit of the southeastern
extension of C. sphinx as it is understood here. A single specimen (BM(NH) 23.10.7.22) from

118 J. E. HILL

the island of Simalur, off the northwest coast of Sumatra is not angulatus as Thomas (19236)
recorded it but is an example of C. horsfieldi (Gray, 1843).

(?) Cynopterus sphinx pagensis Miller, 1906

Cynopterus pagensis Miller, 19066: 62 North Pagi I, Mentawei Is.

SECIMENS EXAMINED. Siberut I, Mentawei Is: 2 dd (1 yg.) 299 BM(NH) 78.2914-2918 Base of Teitei
Bulak, Sabeuleleu, Paitan R, off Saibi R; cf BM(NH) 78.2919 Near Tolailai, off Paitan R; d BM(NH)
78.2920 Near Sibosua R, off Paitan R (all in alcohol, skulls of BM(NH) 78.29 1 5-2920 extracted; coll. J.
J. Whitten).

REMARKS. Specimens from Siberut Island average slightly larger in some respects (Table 2)
than those from Sumatra. They may represent pagensis, which was synonymised with
angulatus by Andersen (1912) but listed as a valid subspecies (of C. brachyotis) by Chasen
(1940) (who included Siberut and Sipora I in the range of pagensis) on account of its
apparently short ears when compared with angulatus and similarly retained by Tate (1942c)
on grounds of darker colour. Two (BM(NH) 95. 1 .9.3-4) from Sipora were thought represent-
ative of pagensis by Andersen (loc. cit.) and are similar to those from Siberut. The Siberut
specimens have relatively short ears (16-1-17-8) while the ears of the two from Sipora are
rather longer (18-2-19-0). If this material in fact represents pagensis then as Andersen
suggested this taxon is very close to angulatus. Specimens of C. sphinx from Krakatau and
Verlaten Island also seem slightly larger than many Sumatran examples, but the sample size
is small.

Cynopterus titthaecheileus titthaecheileus (Temminck, 1825)

Pteropus titthaecheileus Temminck, 1825 : 198, pi. 15, figs. 17, 19,20. Buitenzorg, Java.

SPECIMENS EXAMINED. S Sumatra: cf, 9 BM(NH) 78.1106-1107 Bandan harip, Palas District, S
Lampong; cf BM(NH) 78.1 108 Relung Lelok, Natar District, S Lampong; cf BM(NH) 78.1 109 Pring
kumpul, Pring Sewn District, S Lampong; 9 BM(NH)78.1 1 10 Air Naningan, Pulau Panggung District,
Lampong; cf BM(NH) 78.1 1 1 1 Gisting, Talang Padang District, Lampong (all skins, skulls; presented
by Museum Zoologicum Bogoriense).

Krakatau I, 6° 10' S, 105° 26' E: 9, 2 cfcf (1 imm.) BM(NH) 74.237-239 (in alcohol, skulls of
BM(NH) 74.237-238 extracted; coll. G. Lincoln).

REMARKS. Specimens of Cynopterus here reported from S Sumatra and from Krakatoa are
of considerable taxonomic interest. They fall clearly (in part from Krakatoa) into separate
sympatric or near sympatric groups, corresponding to the taxa identified by Andersen (1912)
as brachyotis, angulatus and titthaecheileus. Those of titthaecheileus from S Sumatra and
Krakatoa average slightly smaller in some respects (Table 2) than titthaecheileus from Java
but the differences are slight. They can be readily distinguished from C. sphinx angulatus
(with which in three instances recorded here they occur) by their considerably larger size,
relatively longer rostrum which is less markedly concave in dorsal profile, curving down less
abruptly from the braincase, and their heavier canines. In turn angulatus although less char-
acteristically larger than brachyotis (also recorded here sympatrically or nearly so with the
other two taxa) has when compared with this smallest of the three forms a more rectangular,
less tapered and more robust rostrum, with a longer palate that is generally wider, especially
anteriorly, and larger, heavier cheekteeth. These three taxa I recognise as distinct species,
namely C. brachyotis (brachyotis), C. sphinx (angulatus) and C. titthaecheileus (titthaechei-
leus).

Andersen (1912) recorded both C. brachyotis brachyotis and C. sphinx angulatus from
Krapoh, Deli-Bedagei in northeastern Sumatra but although this author also recorded C.
titthaecheileus from Sumatra he had no sympatric occurrence of this large bat with either of

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120 J. E. HILL

the other two species. Dammerman (1938) while reporting C. sphinx angulatus from
Krakatoa and Verlaten I also recorded a specimen of C. titthaecheileus from the nearby
island of Sebesi.

DISCUSSION. Andersen (1912) associated titthaecheileus with C. sphinx as a subspecies,
notwithstanding that his arrangement of this part of the genus involved a wide geographical
hiatus between C. s. sphinx in India, Burma and northern Thailand and 'C. s. titthaecheileus'
in Sumatra and Java. This author considered angulatus, which fills this gap, to be a
subspecies of C. brachyotis, a circumstance leading him to the conclusion that sympatric
subspecies of C. brachyotis of different geographic origins, namely C. b. brachyotis and C. b.
angulatus were to be found in the Malay Peninsula and in Sumatra. Although initially
challenged by other authors, notably by Kloss and by his colleague Robinson (Kloss, 1911,
1916, 1917, 1919, Andersen & Kloss, 1915, Robinson & Kloss, 1915a, 19156, 1918) who
thought that angulatus might more properly be considered a subspecies of C. sphinx, this
concept prevailed until quite recently.

Chasen (1940) attempted to resolve the distributional dilemma created by Andersen by
limiting C. b. brachyotis to Malaya, Sumatra, Borneo and some of the associated islands, C.
b. angulatus to southern Thailand and its coastal islands, and to the Natuna and Anamba
Islands. Thus this author envisaged a large form in the north of the Malay Peninsula, a
smaller form occupying the rest of the area from northern Malaya to Borneo, Sumatra, and,
as C. b.javanicus Andersen, 1910, to Java. Hill (1961<a) followed the arrangement proposed
by Chasen but pointed out that specimens in the British Museum (Natural History) indicated
a much wider zone of intergradation in northern Malaya than Chasen envisaged. However,
both views prove unacceptable since larger specimens with forearm 67-5-73 are found also
in Sumatra and agree with the size range that Chasen gives for 'C. b. angulatus', while small
animals corresponding to C. b. brachyotis occur in Thailand (Hill & Thonglongya, 1972) and
are here recorded as far west as Rangoon in Burma.

Hill & Thonglongya (1972) reviewed the history of angulatus and concluded from a study
of the collections of the British Museum (Natural History) that little taxonomic significance
could be given to the features used by Andersen (1912) to distinguish C. sphinx from 'C.
brachyotis angulatus', which he thought sympatric in Burma and northern Thailand.
Consequently these authors regarded angulatus a subspecies of C. sphinx: indeed, specimens
in London demonstrate that the large Indian C. s. gangeticus Andersen, 1910 and the slightly
smaller C. s. sphinx from northeastern India and Burma merge in Burma and Thailand into
the characteristically smaller C. s. angulatus which extends southward into the Malay
Peninsula to Sumatra and possibly east to Borneo. Sody (1930) refers a specimen from
Moerah Teweh, Montellat River, southeast central Borneo to Cynopterus brachyotis
angulatus: his measurements, given in some detail because clearly he recognised the
importance of the specimen agree closely with those of Sumatran examples ascribed to C.
sphinx angulatus, of which there is as yet no published record from Java or from Sulawesi.
The Sumatran and Javan C. titthaecheileus differs from both C. sphinx sphinx and C. s.
gangeticus not only in its rather larger size, although large individuals of gangeticus are
similar in size to the smallest of titthaecheileus, but also in having a much heavier, more
robust skull with broader, higher and more substantial rostrum that is relatively longer. It
differs similarly from C. s. angulatus but these distinguishing features are more pronounced
when it is compared with this smaller form of C. sphinx.

Lack of material prevented Hill & Thonglongya (1972) from any detailed consideration of
Sumatran brachyotis, angulatus and titthaecheileus, these authors following Andersen
(1912) in treating the latter as a subspecies of C. sphinx and, by implication, regarding
Sumatran specimens attributed to it as large examples of C. s. angulatus. Until recently, the
collections in London contained only an immature skeleton from Sumatra referred to
titthaecheileus: other early Sumatran records of this species apparently rest on Andersen
(loc. cit.) who examined most, if not all of the specimens before then reported in the
literature.

INDO-AUSTRALIAN BATS

121

Cynopterus major Miller, 19606, from Nias Island, off west Sumatra is listed by Chasen
(1940) as a subspecies of C. sphinx. Its very large size (length of forearm 75-5-82-0) however,
suggests that more probably it is a subspecies of C. titthaecheileus. Similarly, C. sphinx
terminus Sody, 19406 from Timor is also a large form (length of forearm 75-2-83) but has
a relatively compact skull (condylobasal length 30-2-32-2), with a stout but short rostrum
(Sody, 19406, Goodwin, 1979). On geographical grounds, however, it is more likely to rep-
resent C. titthaecheileus: its relatively small skull might well justify recognition as a distinct
species.

The relevant species and subspecies of Cynopterus may be summarised:

Cynopterus brachyotis (Muller, 1 838)
C. b. ceylonensisGray, 1870
C. b. brachysoma Dobson, 1871a,c
(andamanensis Dobson, 18736)
C. 6. brachyotis (Mullet, 1838)

Cb.altitudinis Hill, \96la

C. b. minutus Miller, 19066
C. 6. concolor Sody, 1 940<2
C. b. javanicus Andersen, 1910
C. b. insularum Andersen, 1910
Cynopterus minor Revilliod, 1911
Cynopterus archipelagus Taylor, 1934
Cynopterus sphinx (Vahl, 1 797)
C. s. gangeticus Andersen, 1910
C.s. sphinx (Vahl, 1797)

C. s. angulatus Miller, 1898

C. s. scherzeri Zelebor, 1869
C. s. serasani Paradise, 1971
C.s. babiLyon, 1916

(?) C. s. pagensis Miller, 19066
Cynopterus titthaecheileus (Temminck, 1 825)
C. t. titthaecheileus (Temminck, 1825)

(?) C. t. major Miller, 19066
(?) Cynopterus terminus Sody, 1 9406

Sri Lanka
Andaman Is.

S Burma to Vietnam, Malaya,
Sumatra, Borneo and many
small associated islands,
Bawean I, Philippine Is,
Talaud I, Sulawesi,
Peleng I.

Malayan highlands.

Nias I, off W Sumatra.

Engano I, off W Sumatra

Java, Madura I, Bali

Kangean I, Mata Siri I.

SE Sulawesi

Polillo I, Philippine Is.

C, NW India.

Sri Lanka, peninsular
and NE India, Burma

N Burma to S China,
Hainan I, Vietnam,
Langkawi I, N Malaya,
Sumatra, Krakatoa I.
Verlaten I, (?) Borneo.

Car Nicobar I.

Serasan I, Natuna Is.

Babi I,nearSimalurI,
off W Sumatra.

Mentawei Is, off W Sumatra.

Sumatra, Krakatoa I,

Sebesi I, Java, Lombok I.
Nias I, off W Sumatra.
Timor I.

Megaerops (?) ecaudatus (Temminck, 1837)

Pachysoma ecaudatum Temminck, 1837 : 94. Padang, W Sumatra.

SPECIMEN EXAMINED. N India: 9 BM(NH) 16.3.25.1 Pashok, Darjeeling, 3000 ft (skin, skull; coll. N. A.
Baptista, Bombay Natural History Society's Mammal Survey of India, Burma and Ceylon).

REMARKS. This specimen is one of two originally reported from Pashok by Wroughton
(1916) as Cynopterus sphinx, to which he referred two others from Tong Song, also in the

122 J. E. HILL

vicinity of Darjeeling. There is no record of these three remaining specimens in the British
Museum (Natural History) and they are presumably in the collections of the Bombay
Natural History Society. A further specimen, BM(NH) 16.3.25.2 from Gopaldhara, 17 miles
W of Sonada Railway Station, SW of Darjeeling is unreported by Wroughton and without
doubt represents Cynopterus sphinx.

There is no previous published record of Megaerops from further west than northern
Thailand, whence M. ecaudatus was reported by Hill & Thonglongya (1972), the species
extending otherwise to Malaya, Sumatra and Borneo. This Indian example is similar in
many features to specimens from Chiangmai, northern Thailand (<5 BM(NH) 70.1440) and
Vietnam (BM(NH) 26.10.14.13) but its upper incisors, upper canines and anterior lower
premolars (pm2) are a little more massive. The narial branch of the premaxilla is less
attenuated in its upper part but is equalled in this respect by specimens from Pahang,
Malaya. Specimens from India, northern Thailand and Vietnam are slightly larger in some
respects than those from Malaya but the differences are small and the number of examples
limited.

Hill & Boeadi (1978) drew attention to variation in rostral profile and outline in M.
ecaudatus, indicating that in specimens from Thailand and Vietnam the rostrum is slightly
shorter and less elevated anteriorly than in Peninsular and Bornean examples, the rostral
profile sloping more gradually to the tip. These authors also found variation in the degree of
lateral compression of the rostrum. The specimens from northern India, northern Thailand
and Vietnam have a short, broad rostrum that is not markedly concave frontally: others from
southern Thailand, Malaya and Borneo have a slightly longer rostrum with a rather more
abruptly concave profile, the rostrum sharply compressed laterally to give its upper part
distinctly and strongly concave lateral margins. A series from Pahang, however, tends to
some extent to bridge this difference, some specimens (for example 99 BM(NH) 60.734,
60.736) having rostra that are a little less abruptly concave in profile and less laterally
compressed than in more extreme Peninsular or Bornean examples. For these reasons
northern specimens are referred to M. ecaudatus with some hesitation.

Measurements (9 BM(NH) 16.3.25.1, rf BM(NH) 70.1440 from Doi Pahompok, Fang
District, Chiangmai, N. Thailand, c. 18° 43 ' N, 98° 59' E and 9 BM(NH) 26.10.4.13 from
Dak-to, Annam, Vietnam, in that order); length of forearm 60-5, 55-7, 59-9; greatest length of
skull to gnathion 27-4, 26-9, 28-2; condylobasal length 26-7, 25-6, 27-4; condylocanine
length 26-6, 25-7, 27-2; length front of orbit-tip of nasals 6-9, 6-5, 7-1; palatal length 13-9,
13-3, 14-1; length palation-basion 10-8, 10-6, 11-1; median depth of premaxillae 1-4, 1-5,
1-4; rostral height at front of c1 5-0, 4-7, 4-7; rostral height at centre of pm3 6-5, 6-3, 6-0;
lachrymal width 9-6, — , 9-8; least interorbital width 5-2, 5-5, 5-7; least postorbital width 5-3,
6-3, 6-0; zygomatic width 17-8, 18-5, — ; width of braincase 12-1, 12-2, 12-3; mastoid width
11-6, 11-7, 11-8; orbital diameter 8-1,8-0, 7-9; c'-c1 (crowns) 6-0, 54, 5-5, (alveoli), 5-5, 5-1,
5-3; m'-m1 (crowns) — , 8-6, 8-3, (alveoli) 8-3, 8-5, 8-4; c'-c1 (cingula, internally) 2-7, 2-8, 2-6;
pm4-pm4 (crowns, externally) — , 8-8, 8-4, (internally) — , 5-9, 5-3; width of mesopterygoid
fossa 4-2, 4-0, 4-2; c-m1 (crowns) 8-6, 8-5, 8-9, (alveoli) 8-3, 8-1, 8-5; length complete
mandible from condyles 19-9, 18-9, 19-7; length right ramus from condyle 20-6, 20-0, 20-7;
c-m2 (crowns)—, 9-4, 9-8, (alveoli) 9-5, 9-3, 9-6.

Measurements of selected teeth (BM(NH) 16.3.25.1, 70.1440, 26.10.4.13): length i2 0-56,
0-48, 0-5 1 ; width i2 0-58, 0-4 1 , 0-44; length i3 0-53, 0-46, 0-46; width i3 0-45, 0-35, 0-43; length
c1 (from front face) 1-98, 1-71, 1-82; width c1 1-67, 1-39, 1-52; length pm2 0-71, 0-70, 0-61;
width pm2 1-02, 0-87, 0-80.

Chironax melanocephalus (Temminck, 1825)
Pteropus melanocephalus Temminck, 1825 : 190, pis. 12, 16, figs. 3, 4. Bantam, Java.

INDO-AUSTRALIAN BATS 123

SPECIMENS EXAMINED. N Sumatra: 99 BM(NH) 81.678-679 Near Bukit Lawang, Bohorok R (Bohorok
orang utan rehabilitation centre), Langkat Reserve, Gunung Leuser Reserve complex (in alcohol; coll.
R. Aveling).

W Java: 3 drf, 3 99 BM(NH) 78.1085-1088 (skins, skulls), BM(NH) 78.1089-1090 (in alcohol)
Cibodas, Mt. Gede (presented by Museum Zoologicum Bogoriense).

SPECIMENS REPORTED BUT NOT EXAMINED. N Sulawesi: 99 Zoologisch Museum, Amsterdam ZMA
2 1 .638-639 Over an upper tributary of Sungei Mauk, Cagar Alam Dumoga, 960 m (in alcohol; coll. K.
D. Bishop, F. G. Rozendaal and W. F. Rodenburg).

REMARKS. There are relatively few references to this species in the literature and for many
years it was known only from the original material from Java, in the Rijksmuseum van
Naturrlijke Historic, Leiden (Andersen, 1912). Thomas (1923#) reported a specimen
(BM(NH) 23.1.2.2, a juvenile) from Nias Island, off west Sumatra, Chasen (1940) sub-
sequently recording specimens from northern Sumatra and from Selangor, Malaya. More
recently further specimens have been reported from Selangor by Hill (196 la) and Hill &
Thonglongya (1972) have recorded others from Pahang, Malaya and from Nakon Sri
Thamrat in southern Thailand. Finally, Hill (1974) provisionally referred a young adult from
Sulawesi to this species, which does not appear to have been reported from Java since orig-
inally described by Temminck. Moreover, none of the specimens subsequently recorded
from other parts of the Malaysian region appear to have been compared directly with Javan
material. The additional specimens reported here are of some interest since those from Java
enable such a comparison to be made, not only with Malayan but also with Sumatran
examples. Additionally, Dr. W. Bergmans has provided details and measurements of two
specimens from Sulawesi in the Zoologisch Museum, Amsterdam that he has identified as
C. melanocephalus.

Andersen (1912) found the two 'cotypes' in the Rijkmuseum van Natuurlijke Historic to
be so faded as to be unsuitable for description. According to Temminck (1825) 'The hairs
on the back are of two colours, a yellowish white at the base and an ashy black at the tip;
nape, top of the head and muzzle black; the hairs diverging from a common centre on the
sides of the neck, serving probably to conceal an apparatus that secretes an odorous fluid.
All of the other parts beneath are a yellowish white and dull; membranes a dark brown.'

Specimens from Java agree closely in pelage colour with the description by Temminck.
Dorsally the hairs are whitish or slightly yellowish white at the base and for much of their
length, and are tipped with dark brown or blackish brown; the nape and the head are a
darker, more blackish tint, while the underparts are dull greyish white, tinged with yellow.
As Temminck remarked, there is a radial tuft of hairs at the side of the neck: these hairs are
greyish white throughout their length and are a little coarser in texture than the surrounding
pelage. The tuft itself is usually slightly more obvious in male than in female specimens. A
male example (BM(NH) 67. 1488) in dry preservation from Pahang, Malaya is very similar in
overall coloration to Javan specimens but the neck tufts are orange, while a female (BM(NH)
70.1441) from southern Thailand has a slightly less blackish head and is browner ventrally,
the neck tufts less prominent than in the male from Pahang (Hill & Thonglongya, 1972) and
less distinctly orange. One of the two females (BM(NH) 81.679) recorded here from Sumatra
has quite well marked orange neck tufts but the other example has no obvious neck patches.
In both the throat is white or creamy white. Other specimens from Pahang (BM(NH)
67.1484-1486), Trengganu (BM(NH) 75.1233) and Selangor (BM(NH) 60.865-878) in
Malaya, also preserved in alcohol, confirm that the neck tufts or patches are generally less
developed in females and indicate that they are not invariably orange or tinged with orange
although inevitably some changes in colour have occurred through prolonged immersion.
The young adult female (BM(NH) 73.1802) from southern Sulawesi provisionally referred to
C. melanocephalus by Hill (1974) is a little paler dorsally than Javan specimens and is tinged
with grey over the shoulders: the nape and head are dark greyish brown rather than black,
and it has prominent white neck patches, the white extending to the throat. According to Dr.
Bergmans one of the Sulawesian specimens (ZMA 21.639) in Amsterdam agrees in colour

124 J. E. HILL

with the description (Hill, 1974) of this specimen: the other (ZMA 21.638) is somewhat
lighter on the head and back but may be subadult.

Wing indices (Table 3) for the Javan specimens are in broad agreement with those given by
Andersen (1912) for the two 'cotypes' of C. melanocephalus, although not unexpectedly
there is a greater variation in the lengths of the digital components than was found by this
author in so limited a sample. The metacarpals of the third, fourth and fifth digits, the first
and second phalanges of the third digit and the second phalanges of the fourth and fifth digits
of examples from Java are generally relatively shorter than are those of specimens from
Sumatra and from the Malayan mainland, the third metacarpal especially so. On the basis of
the two 'cotypes' Andersen (loc. cit.) noted of the wing of Chironax when compared with
that of the closely related genus Balionycteris that the metacarpals of the third, fourth and
fifth digits were not lengthened (lengthened in Balionycteris) the second phalange of the third
digit was subequal in length to its metacarpal (much shorter in Balionycteris) and that the
second phalanges of the fourth and fifth digits were slightly but distinctly longer than the
associated first phalanges of the same digits (shorter in Balionycteris). The Javan specimens
tend to support these contentions: in Sumatran and Malayan specimens the metacarpals of
the third, fourth and fifth digits are lengthened to approach the condition found in
Balionycteris but the second phalange in each of these digits is also correspondingly
lengthened to retain more or less the relative proportions of Chironax. Specimens from
Sulawesi have similarly lengthened third, fourth and fifth metacarpals but the second
phalange of the third, fourth digits and on occasion of the fifth digit is relatively shortened as
in Balionycteris.

Cranially the Javan specimens are similar in structure and size to those from Malaya, with
similar teeth, as have the specimens from Sumatra: the second upper premolar (pm3) has a
well-marked antero-external supplementary cusp and the third upper premolar (pm4) is
more or less rectangular as in Malayan specimens. The young adult (BM(NH) 73.1802 from
Sulawesi thus differs from Javan examples in much the same ways as it was found to differ
(Hill, 1974) from those from Malaya: in some respects the skull is a little smaller, its supra-
orbital region is slightly more swollen and the postorbital processes a little more massively
developed, while pm3 lacks a well developed antero-external supplementary cusp and pm4 is
more rounded, dental features in which it also differs from the Sumatran specimens. Two
points not mentioned in the account of BM(NH) 73.1802 are that the rear edge of the bony
post-palate is slightly concave rather than straight or nearly so as in Javan and Malayan
examples, and that one of the inner lower incisors (i2) is missing. Dr Bergmans remarks that
in both Sulawesian examples in Amsterdam pm3 also lacks an antero-external supplemen-
tary cusp: possibly this feature will be found to characterise the Sulawesian population.

External measurements of six specimens (except where indicated in parentheses) from
Java (BM(NH) 78.1085-1090), with those of two (BM(NH) 81.678-679) from Sumatra:
length of forearm 42-8^5-7, 4 1 -9, 42-3; IIIm 27-7-29-8, 28-9, 29-2; III1 20-9-22-0, 2 1 -0, 2 1 -4;
IIP 25-8-28-2, 25-2, 27-5; IVm 26-5-28-2, 26-9, 27-3; IV1 16-1-17-2, 16-4, 16-7; IV2
17-1-18-9, 16-8, 18-2; Vm 27-8-29-5, 28-0, 28-1; V1 14-0-14-9, 15-0, 14-5; V2 15-1-16-0,
14-8, 15-9; length of ear from opening (2) 9-9, 10-5, 10-4, 10-9; greatest width of ear (2) 7-6,
7-8, 7-8, 7-3; length of tibia (2) 16-1, 16-2, 16-0, 15-9; length of foot (c.u.) (2) 10-5, 10-9,
9-5,9-4.

Wing measurements (by Dr W. Bergmans) of two specimens (ZMA 21.638-639) from
Sulawesi: length of forearm 45-6, 45-6; III"1 32-0, 32-4; III1 23-1, 23-7; IIP 23-3, 26-8; IVm
30-8, 31-0; IV1 18-2, 17-9; IV2 15-2, 17-1; Vm 32-6, 32-4; V1 14-6, 15-3; V2 13-8, 15-5.

Cranial measurements of four specimens (except where indicated in parentheses) from
Java (BM(NH) 78.1085-1088): greatest length of skull 21-9-23-3; condylobasal length (3)
21-5-22-3; condylocanine length 21-0-22-0; length front of orbit-tip of nasals 5-0-5-5; length
orbit-nares 5-1-5-6; palatal length (3) 10-4-11-7; length palation-incisive foramina (3)
8-6-10-0; length palation-basion (3) 8-1-8-7; lachrymal width 6-0-6-3; least interorbital
width 4-3^-4; least postorbital width 4-6-5-3; zygomatic width 14-5-14-8; width of
braincase 9-8-10-2; mastoid width 10-0-10-5; orbital diameter 5-6-6-0; c'-c1 (crowns) (3)

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126 J. E. HILL

4-4-^-5, (alveoli) (3) 4-1-4-2, (cingula, internally) (3) 1-9-2-2; pm4-pm4 (crowns) (3)
6-4-6-8, (alveoli) 6-1-6-6, (internally) (3) 3-8-4-1; m'-m' (crowns) 6-1,6-5, (alveoli) 6-1-6-4;
width mesopterygoid fossa 3-1-3-3; c-m1 (crowns) (3) 6-6-7-3; length complete mandible
from condyles 15-1-16-3; length right ramus from condyle 16-1-17-2; coronoid height
7-6-8-7; c-m2 (crowns) 7-5-7-9.

Thoopterus nigrescens (Gray, 1870)

Cynopterus marginatus var. nigrescens Gray, 1 870 : 123. Morty ( = Morotai) Island.
Cynopterus latidens Dobson, 1878 : 86. Morty ( = Morotai) Island.

SPECIMENS EXAMINED. C Sulawesi: 9, 2 dd BM(NH) 81.1049-1051 R Ranu, 1° 51 ' S, 122° 22' E;
8 rfrf, 6 99 BM(NH)81.1052-1065 Tambusisi Damar, Mt. 4000 ft. 1° 39' S, 121° 22 ' E (in alcohol.
BM(NH) 8 1.1054, 81.1056, 81.1058-1061, 81.1063-1064 heads only; coll. B. H. Gaskell, 'Operation
Drake').

REMARKS. There are few records of this long-haired cynopterine fruit bat which is known so
far with certainty from Morotai Island and from Menado and Minahassa in northern
Sulawesi. Its reported occurrence on Luzon Island in the Philippines is thought doubtful by
Taylor (1934) although it may extend to the southernmost of these islands. Length of forearm
in four examples 76-5-78-9.

DISCUSSION. This large bat is similar in many respects to the relatively recently described
genus Latidens Thonglongya, 1972 from southern India, which is almost identical in
external form and colour and has a similarly long, strong rostrum. Dentally, however,
Latidens approaches the Malaysian genus Penthetor, differing in the outline of m1, which is
more or less square and not wedge-shaped, and in the wider, rather more square outline of
pm4 and m,. The latter teeth, however, also have a low surface cusp and in this respect
approach Thoopterus. These and other features suggest that Thoopterus is represented in
Malaysia by Penthetor as was thought by Andersen (1912) and in southern India by
Latidens. The three genera can be readily distinguished from each other by the incisive
dentition, Thoopterus having incisors \ — \, Penthetor \ — \, and Latidens \ — \.

Aethalops alecto (Thomas, 1923)

Aethalodes alecto Thomas, 1923a : 251. Indrapura Peak, Sumatra, 7300 ft.

SPECIMENS EXAMINED. W Java: <t, 9 BM(NH) 75.590-591 Cibodas, Mt. Cede, 1450 m, c. 6° 50' S, 106°
44 ' E (skins, skulls; presented Museum Zoologicum Bogoriense).

REMARKS. Two subspecies of the small fruit bat A. alecto are recognised, A. a. alecto from the
Malayan mainland and from Sumatra, and A. a. aequalis Allen, 19380 from Sabah and
Sarawak. Their diagnostic characters are reviewed in part by Hill (196 la, 1966a), who also
gave measurements. The species has not been formally recorded hitherto from Java but that
island is included within its distribution by Honacki et al. (1982). These specimens cannot be
referred definitively to either of the described subspecies. They are similar in most respects to
A. a. alecto but have longer forearms and longer and more massive incisors, with the anterior
lower premolar (pm2) larger (about twice the crown area of the lower incisor (i2) against one
and one half times the crown area of i2 in A. a. alecto) and more substantial. They differ more
noticeably from A. a. aequalis in rather longer forearms, longer skull and toothrows, in
slightly heavier incisors, with the inner upper incisors (i2) distinctly shorter than the outer
tooth (i3), in having the lower canine rounded antero-internally, not especially flattened, in
larger anterior premolars (pm^) with pm2 in particular more massive (in A. a. aequalis it is
about equal in crown area to i2), the second premolars (pm^) less closely approximated to the

INDO-AUSTRALIAN BATS 127

canines, pm3 having a well developed anterior secondary basal cusp while pm3 is rounded
anteriorly, lacking any projecting vertical anterior ridge.

Measurements (BM(NH) 75.590, 75.591): length of forearm 50-3, 51-0; greatest length of
skull — , 254; condylobasal length — , 24-8; condylocanine length — , 24-5; palatal length
13-3, 13-2; lachrymal width 6-8, 6-3; least interorbital width 5-4, 5-1; least postorbital width
5-6, 5-3; zygomatic width 16-1, 15-5; width of braincase 11-0, 10-2; mastoid width 11-6, 10-5;
c'-c1 (crowns) 5-4, 5-1, (alveoli) 4-9, 4-5; pm4-pm4 (crowns) 8-0, 7-4, (alveoli) 7-5, 7-0; c-m1
(crowns) 8-5, 8-1, (alveoli) 8-0, 7-7; length complete mandible from condyles 18-6, 18-2;
length right ramus from condyle 19-4, 18-8; c-m2 (crowns) 9-3, 8-8, (alveoli) 9-0, 8-6.

Javan specimens of A. alecto are clearly closely related to A. a. alecto from Malaya and
Sumatra, (whence only the holotype appears to have been recorded) differing only in slightly
greater size in some respects, in the greater length and size of the incisive dentition and in
larger pm2. The Bornean A. a. aequalis differs more from either A. a. alecto or from the Javan
examples than these populations do from each other. It has long upper incisors (similar to
those of the Javan specimens although slightly smaller) but i2 is equal or nearly equal in
length to i3, the inner face of the lower canine opposite the incisor is flattened or even broadly
but shallowly grooved, pm3 lacks any well developed anterior secondary basal cusp and the
anterior face of pm3 is narrowly ridged from the cingulum to the tip of the tooth. For the
present the subspecific designation of the Javanese specimens is left undecided, pending
more material to establish clearly the limits of their variation.

let halo ps alecto aequalis Allen, 1938

Aethalops aequalis Allen, 1938 : 497. Lumu Lumu, Mt. Kinabalu, Sabah, 5500 ft.

SPECIMENS EXAMINED. Sarawak, Borneo: 9 9 BM(NH) 78.590-59 1 Camp 2, Gunung Mulu, Melinau, 3°
49' N, 115° 50' E, 550m (in alcohol; obtained Earl of Cranbrook, Royal Geographical Society
Expedition to Mulu, 1978).

REMARKS. These specimens supplement earlier records of A. a. aequalis (summarised by
Medway, 1977) all from the highlands of north and northeastern Borneo. They are, however,
from a somewhat lower altitude than any previously obtained, the subspecies having not
before been collected below some 3000 ft.

HARPYIONYCTERINAE

Harpyionycteris celebensis Miller & Hollister, 192 1

Harpyionycteris celebensis Miller & Hollister, 1921 : 99. Gimpoe, C Sulawesi.

SPECIMENS EXAMINED. C Sulawesi: 2 dd, 5 99 (2 yg.) BM(NH) 81.1150-1156 R Ranu, 1° 51' S, 121°
30 ' E (in alcohol, skulls of BM(NH) 8 1 . 1 1 50-1 1 5 1 , 8 1 . 1 1 53, 8 1 . 1 1 55 extracted); 2 rfrf( 1 yg.) BM(NH)
81.1157-1158 Tambusisi Damar, Mt. Tambusisi, c. 4000 ft, 1° 39 ' S, 1 2 1° 23 ' E (BM(NH) 8 1 . 1 1 57 in
alcohol, 81.1 158 skull only) (all coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Harpyionycteris celebensis was said by Miller & Hollister (1921) to be like H.
whiteheadi from the Philippine Islands but to have a conspicuous secondary cusp on each
side (i.e. on the anterior and posterior faces) of the second upper premolar (pm3) and to have
molars with lower crowns and relatively higher cusps. Tate (195 1) in a detailed evaluation of
Harpyionycteris suggested that celebensis was probably no more than subspecifically related
to whiteheadi but made no close comparison of their distinguishing features. This suggestion

128 J. E. HILL

was adopted by Laurie & Hill (1954) who considered Harpyionycteris monotypic, celebensis
being listed as a subspecies of//, whiteheadi.

More recently, Peterson & Fenton (1970) have reviewed the specimens then known of
Harpyionycteris, considering celebensis to be a distinct species on account of its shorter,
broader upper canine which is less proclivous than in whiteheadi and has a much wider
posterior cusp when viewed laterally, and also to some extent in the presence in celebensis of
a well developed secondary cusp on the posterior flange of pm3. These authors apparently
had available two adult examples from Sulawesi, one the holotype of celebensis. The series
reported here from Sulawesi suggests that there is a considerable degree of variation in these
features: variation in dental patterns in this genus can be quite extensive, as Peterson &
Fenton pointed out in relation to the upper molars of the series from Negros Island,
Philippine Islands that they described as H. whiteheadi negrosensis.

The series from Sulawesi consists of three subadult or young specimens and six adults. As
in the series from Negros Island examined by Peterson & Fenton, immature specimens have
a rather more strongly deflected braincase than do mature examples, and have a wider
postorbital constriction. These authors also found the lower incisors generally absent from
specimens of H. whiteheadi that they examined, three of twelve examples having only one
incisor (on one side only) all of the others lacking lower incisors. Two immatures and two
adults of the series from Sulawesi have two small, slightly spatulate lower incisors tightly
sandwiched between the anterior part of the canine bases: one immature and two adults have
one lower incisor, in one example (BM(NH) 81.1155) placed to one side with some
indication that the corresponding tooth on the other side has been lost, in the others more
centrally situated; and in two adults lower incisors are lacking.

The upper canines of specimens from Sulawesi are less proclivous and are generally rather
more massive but shorter than those of the holotype of//, whiteheadi although in some the
difference in size is small. The posterior canine cusp is variable in size: in a minority of
specimens the cusp is large and massive, lacking any definite point, like that of the specimen
(AMNH 153590) illustrated by Peterson & Fenton (1970), but more frequently the cusp is
more or less triangular with a definite point, like that of the specimen photographed by Tate
(1951, figs 1-3). Although generally larger than the corresponding cusp in the holotype of//.
whiteheadi it may be on occasion similar in size to its counterpart in that species or even
slightly smaller.

The cuspidation of pm3 varies considerably in this series. The tooth in lateral profile
ranges from a relatively high crowned structure with small but well developed
supplementary cusps on its anterior and posterior margins extending along about three
quarters the length of the tooth to just beneath the tip of the main cusp through stages in
which the cusps are lower, about halfway along the length of the tooth, or present only
anteriorly, the posterior cusp represented only by a flexure of the margin of the tooth, absent
anteriorly but present posteriorly, or both effectively absent, their presence indicated only by
a faint curvature of the edge of the main cusp. The tooth as a whole is lower than in the
holotype of//, whiteheadi, although similar in overall bulk.

The molar teeth do not seem to be especially low crowned or high cusped in relation to
those of//, whiteheadi, and there is no great degree of variation in their cuspidation such as
that noted by Peterson & Fenton in the series that they named H. whiteheadi negrosensis
from Negros Island. Cranially, specimens from Sulawesi have the median part of the palate
more deeply excavated and transversely arched than does the holotype of H. whiteheadi, a
feature not mentioned by Peterson & Fenton for the Sulawesian examples that they
examined. Tate (1951) remarked that the yellowish markings on the wings of//, whiteheadi
might be significant but similar yellowish spots or blotches are to be found on the wings of
specimens from Sulawesi.

The palate ridges of Sulawesian specimens are similar to those of Philippine examples as
illustrated by Sanborn (1952) and illustrated and described by Peterson & Fenton (1970).
There are five anterior undivided ridges and three divided posterior ridges: these latter are
greatly variable and the central ridge of the three is sometimes reduced to small paired raised

INDO-AUSTRALIAN BATS 129

swellings on each side of the median division, or tends to blend and merge with the first of the
divided ridges. There is also an undivided ridge much more posteriorly situated near the rear
edge of the palate.

External measurements of six adults: length of forearm 84-2-88-3; length of thumb (c. u.)
35-1-37-3; IIm 39-2-42-3; II1 11-2-12-0; II2-3 (c. u.) 11-3-12-8; IIIm 57-6-62-4; III1 45-6-47-8;
III2 57-8-63-3; IVm 54-0-58-5; IV1 36- 1-37-8; IV2 35-3-37-5; Vm 55-9-58-3; V1 28-9-3 1 -3; V2
32-7-35-7; length of tibia 29-2-29-5; length of foot (c.u.) 22-2-23-6.

Cranial measurements of four adults: greatest length of skull 41-2-42-0; condylobasal
length 39-2-39-7; condylocanine length 39-0-39-6; length front of orbit-tip of nasals
10-4-11-5; palatal length 21-3-22-1; length of palation-incisive foramina 17-0-18-1; length
palation-basion 15-7-16-4; lachrymal width 10-0-11-0; least interorbital width 6-4-7-0; least
postorbital width 5-7-6-4; zygomatic width 23-4-24-1; width of braincase 15-7-16-7; height
of braincase 13-3-14-3; mastoid width 14-6-15-6; orbital diameter 8-9-9-6; c'-c1 (crowns)
7-4-8-2, (alveoli) 6-8-7-4; (cingula, internally) 2-8-3-1; pm3-pm3 (crowns) 8-8-9-5; (alveoli)
8-3-9-1, (internally) 5-0-5-7; pm4-pm4 (crowns) 9-5-10-7, (alveoli) 9-1-10-3, (internally)
5-4-6-3; m2-m2 (crowns) 10-7-11-7, (alveoli) 10-6-11-6, (internally) 7-3-8-1; width
mesopterygoid fossa 4-6-5-2; c-m2 16-6-16-8; length complete mandible from condyles
31-5-32-3; length right ramus from condyle 32-4—33-9; depth mandible between pm4 and m,
4-0-4-5; depth mandible behind m3 5-3-5-7; coronoid height 15-1-15-8; c-m, 1 7-7-18-4.

DISCUSSION. Peterson & Fenton (1970) present a persuasive argument for the recognition of
celebensis as a distinct species, supported by morphological features drawn from the skull
and dentition, by zoogeographical argument based on the considerable distance over open
water separating Sulawesi from the nearest small islands that might serve as stepping stones
from the Philippines and the separation of these islands from Sulawesi by Wallace's Line.
They also base their view on the fact that the genus has not been recorded from Borneo
which is less widely separated from Sulawesi at its point of closest approach (in fact Borneo
lies on the other side of Wallace's Line and is connected to the Philippine Islands by two
chains of relatively narrowly separated islands) and on a presumed dietary specialisation
limiting Harpyionycteris to high altitude species of fruit.

Certain of the morphological characters, however, are clearly valueless or perhaps are
unreliable for diagnostic purposes. The presence or absence of an accessory posterior cusp
on the second upper premolar (pm3) evidently lacks taxonomic reliability and the
dimensions and form of the accessory canine cusp do not seem firm characters on which
diagnosis can be based. The degree of proclivity of the upper canine may also prove
unreliable: although in celebensis the upper canines are generally less proclivous than in
whiteheadi the figure by Peterson & Fenton (1970, fig. 6) indicates that the upper canines of
the subspecies H. whiteheadi negrosensis are nearer in proclivity to those of celebensis than
to those of//, w. whiteheadi. According to Peterson & Fenton (1970) no more than three
examples of this latter subspecies are known and of these only the holotype has been avail-
able for comparison: this suggests that celebensis differs from H. w. whiteheadi in its
generally less forwardly projecting upper canines which are usually shorter and antero-
posteriorly a little broader, and in a rather more deeply excavated median palate.

The zoogeographical case put forward by Peterson & Fenton (1970) also deserves further
consideration. Apart from Harpyionycteris only Acerodon, Dobsonia and Nyctimene among
other fruit bats are known so far to bridge the water gap between Sulawesi and the Philippine
Islands yet apparently do not occur in Borneo. The large Acerodon occurs also on the Talaud
Islands, and Dobsonia has one form, chapmani, on Negros Island (where Harpyionycteris
is also found) apparently related to a predominantly Moluccan and New Guinea species,
D. moluccensis. The genus Nyctimene has also been reported from Negros Island (Rabor
et al., 1970). The distribution pattern is clearly an unusual one for fruit bats and possibly
as Peterson & Fenton suggest the Sulawesian population of Harpyionycteris may have been
long isolated. On the other hand, Sulawesian specimens from R Ranu show that
Harpyionycteris is not exclusively a high altitude bat, although in Sulawesi it was also

130 J. E. HILL

obtained at a considerable altitude on Mount Tambusisi, and not necessarily dependent on
high altitude species of fruit that might not be available on small islands. It is probably
correct to say as Peterson & Fenton (1970) do that it is a specialised feeder: its multicuspid
teeth suggest a diet of hard fruit.

In view of these various considerations, for the present I also regard celebensis as
specifically distinct from H. whiteheadi, but with strong reservations that this view may well
be modified when fully adequate comparative material is available. In particular, further
specimens of//, whiteheadi whiteheadi are required to establish the extent of dental variation
in this subspecies, if any. It also seems possible that the distribution of Harpyionycteris is
imperfectly known as yet, and that it may be found to occur at least on some of the other
islands that abound in the Philippine and Sulawesian regions. Such additional material
might prove highly relevant to any further consideration of the relative status of the two taxa.

NYCTIMENINAE

Nyctimene albiventer papuanus Andersen, 1910

Nyctimene papuanus Andersen, 1910 : 621. Milne Bay, Papua New Guinea.

SPECIMENS EXAMINED. Papua New Guinea: cf BM(NH) 69.1417 Olsobip, Upper Fly R, 1500-2000 ft
(skin, skull; coll. J. I. Menzies); 99BM(NH) 73.2106-2025 Kairiru ridge, centre of Kairiru I, near
Wewak, East Sepik, c. 2000 ft (in alcohol; coll. Aberdeen University Exploration Society Expedition to
New Guinea, 1973); 9 BM(NH) 78.867 Sapi Creek Forest Reserve, near Baku, Gogol Valley, Madang,
c. 40 m (in alcohol), 9 BM(NH) 78.868 Sapi Creek, c. 8 km E of Baku (skin, skull) (both coll. P. A.
Morris); d BM(NH) 80.565 Lobota Cave, Morobe, 7° 15' S, 147° 09' E, 18 d-rf, 11 99 BM(NH)
80.566-594 Buso, Morobe, 7° 17' S, 147° 08' E (all in alcohol; coll. B. H. Gaskell, 'Operation
Drake').

Nyctimene cephalotes cephalotes (Pallas, 1767)

Vespertilio cephalotes Pallas, 1767: 10, pis. 1, 2. 'Moluccas'. Type locality fixed on Amboina
I by Andersen (19 12).

SPECIMENS EXAMINED. Molucca Is: d1, 9, d BM(NH) 81.1128-1130 Kaiteto, Amboina I (in alcohol;
coll. B. H. Gaskell, 'Operation Drake').

C Sulawesi: 14cfc?,499 BM(NH) 81.1 131-1 148 R Ranu, 1° 51' S, 121° 30' E (in alcohol; coll. B. H.
Gaskell, 'Operation Drake').

Nyctimene major lullulae Thomas, 1 904

Nyctimene major lullulae Thomas, 1904 : 197. Woodlark I, Trobriand Is.

SPECIMENS EXAMINED. Papua New Guinea: d d BM(NH) 75. 1 86 1-1 862 Kadovar I, Schouten Is, 3° 38
S, 144° 36 ' E (in alcohol; coll. A. M. Jones, Aberdeen University Exploration Society).

REMARKS. In overall size (length of forearm 69-9, 67-5; m'-m1 (crowns) 9-8, 9-9; (alveoli) 9-6,
9-7; c-m1 12-0, 1 1-9) and in the size of their teeth these specimens generally approach more
closely to N. m. lullulae than to either of the other subspecies known from the islands
surrounding the northeastern and eastern coasts of New Guinea, namely N. m. major
(Dobson, 18770) from the Bismarck Archipelago and N. m. geminus Andersen, 1910 from
eastern New Guinea and its associated islands.

INDO-AUSTRALI AN BATS 1 3 1

DISCUSSION. Koopman (1979) has discussed the distribution of N. major on the islands off
northeastern New Guinea in some detail. This author found that while the larger subspecies
N. m. major occurred on the more outlying islands of the Bismarck Archipelago, smaller
specimens from the inshore islands of Karkar and Bagabag are similar in size to N. m.
lullulae from the more easterly island of Woodlark. Kadovar Island lies considerably to the
northwest of Karkar and Bagabag and is evidently occupied by a population of similar
smaller overall size. More recently, Koopman (1982) has examined variation in this species
on the islands off the eastern tip of New Guinea.

Nyctimene aello (Thomas, 1 900)

Cephalotes aello Thomas, 1 900 : 2 1 6. Milne Bay, Papua New Guinea.

SPECIMENS EXAMINED. Papua New Guinea: d, (?) BM(NH) 68.1 109-1 1 10 Laloki, 9° 26' S, 147° 20' E
(skins, skulls; coll. J. I. Menzies); d, 9 BM(NH) 73.203 1-2032 Victoria Bay, NW end of Kairiru I, near
Wewak, East Sepik; d, 9 BM(NH) 73.2033-2034 Sabor Kunai, E end of Kairiru I (all in alcohol; coll.
Aberdeen University Exploration Society Expedition to New Guinea, 1973); 6 cfcf, 9 BM(NH)
80.595-60 1 Buso, Morobe, 7° 1 7 ' S, 1 47° 08 ' E (in alcohol; coll. B. H. Gaskell, 'Operation Drake')-

REMARKS. Specimens in dry preservation from Laloki are pale brown dorsally and have a
broad black dorsal stripe extending almost to the head: one is fulvous ventrally, the other
more buffy brown, the flanks in both rather more brownish. Those from Kairiru and Buso
are brownish dorsally, with a similar generally broad dorsal stripe: the throat, chest and
centre of the belly is dull buffy white or buffy brown shading into brownish on the flanks. All,
however, have been preserved in alcohol.

DISCUSSION. Laurie & Hill (1954) list N. aello with two subspecies, N. a. aello from eastern
New Guinea and N. a. celaeano Thomas, 1922a & b from the western and northwestern
parts of the island. According to Thomas (\922a & b) celaeano is slightly smaller than aello
and is browner and less yellowish or fulvous, the dorsal surface less yellow and much of the
ventral surface dull buffy white, the sides brown, not fulvous as in aello. These differences in
colour between the holotypes of celaeno (BM(NH) 22.2.2.2) and aello (BM(NH) 99.12.3.1)
are not entirely supported by the newly acquired material reported here or by other
specimens in dry preservation in the collections of the British Museum (Natural History).
The holotype of 'celaeno has the head, both above and below, the nape and the shoulders pale
yellowish white, in contrast to the more fulvous coloration of the holotype of aello: the
remainder of the dorsal surface is slightly paler than in aello and the ventral surface is dull
buffy white, lacking the orange tinge characteristic of this specimen. A second specimen
(BM(NH) 29.5.27.3) referred to celaeno, from Wasjor, West Irian almost exactly resembles
the holotype of aello in coloration, corresponding closely in ventral colour to the more
brightly coloured (BM(NH) 68.1110) of the two from Laloki. BM(NH) 1939.1351 from
Tamata, Mambare River and BM(NH) 1939.1352 from Takar are a slightly paler brown
dorsally than the holotype of aello and ventrally are less strongly tinged with fulvous or
orange, corresponding to but a little brighter ventrally than the duller (BM(NH) 68.1 109) of
the two Laloki examples.

The coloration of the holotype of celaeno is characteristic of immersion in alcohol:
Thomas (\922b) remarked 'Adult male, skinned out of spirit' and its original labels show
that in fact it was collected in 1910 (not 1912 as stated by Thomas, \922b). It may not have
been prepared as a dry skin until 1 92 1 or 1 922. Careful examination shows that it does retain
ventrally some trace of a brighter colour along the flanks, although much bleached.
Moreover, there is little of this brighter ventral coloration in specimens of aello that although
relatively recently collected have been preserved in alcohol. There is every indication,
therefore that the colour of the holotype of celaeno is of limited value as a comparative
feature.

132 J. E. HILL

However, there is a number of cranial and dental features by which the holotype of celaeno
differs from aello. It has a much shorter, rather broader rostrum, wider interorbital and
intertemporal areas, the frontal region is more elevated in contrast to the rather depressed
frontal region of aello (commented upon by Thomas, 1922£), the supraorbital ridges are
more swollen and inflated, and on the whole the teeth are generally smaller, the upper
canines less obviously proclivous than in aello. For the present it seems more appropriate to
consider it a distinct species allied to N. aello, at least until more specimens can be examined
to establish its status: BM(NH) 29.5.27.3 from Wasjor, West Irian, hitherto referred to
celaeno, agrees externally and cranially with N. aello to which it should be allocated. Tate
(1942c) thought celaeno apparently closely allied to N. major from New Guinea and its
associated islands but this species differs quite widely

Paranyctimene raptor Tate, 1 942

Paranyctimene raptor Tate, 1942a : 1 . Oroville Camp, Fly R, about 4 miles below mouth of Elavala R,
Papua New Guinea.

SPECIMENS EXAMINED. (?) BM(NH) 68.1111, cf BM(NH) 69.317 Laloki, about 12 miles N of Port
Moresby, 9° 26' S, 147° 20' E (skins, skulls; coll. J. I. Menzies); 2 dd, 3 99 (1 yg.) BM(NH)
73.2024-2028 Rauit, West Sepik, 1750 ft, 3° 36' S, 142° 15' E; rf BM(NH) 73.2029 Biip, J mile SE of
Rauit, 1650ft; d, 9 BM(NH) 73.2030, 73.2035 Rauit R, W of Rauit, 900ft (all in alcohol; coll.
Aberdeen University Exploration Society Expedition to New Guinea, 1973); 9 BM(NH) 75.1860
Rauit, West Sepik (in alcohol; coll. A. M. Jones, Aberdeen University Exploration Society); 9 BM(NH)
78.869 Baiyer R, c. 50 km NW of Mt. Hagen, Western Highlands; 9 BM(NH) 78.870 Sapi Creek Forest
Reserve, near Baku, Gogol Valley, Madang; cf BM(NH) 78.871 Baiyer R area, Western Highlands; d
BM(NH) 78.872 Mt Hagen (town), Western Highlands (all in alcohol; coll. P. A. Morris); 2 dd, 2 99
BM(NH) 80.602-605 Buso, Morobe; 9 BM(NH) 81.1 149 Avi, Mt. Hagen, Western Highlands (all in
alcohol; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Specimens BM(NH) 73.2024-2030, 73.2035 are those reported by Grieg-Smith
(1975) as Nyctimene draconilla Thomas, \922b, which is similar in size and coloration to
Paranyctimene raptor. This author commented upon the wide range extension that they
indicated for draconilla (considered a subspecies ofN. albiventer (Gray, 1863) by Laurie &
Hill, 1954) and on their sympatric occurrence with N. albiventer, suggesting therefore that
draconilla should be considered a distinct species. However, Koopman (1979) has reported
the sympatric occurrence of N. albiventer papuanus with N. draconilla on the Upper Fly
River in Papua New Guinea, later (1982) confirming his identification of draconilla from
this locality.

MACROGLOSSINAE

Eonycteris spelaea (?) glandifera Lawrence, 1939

Eonycteris spelaea glandifera Lawrence, 1939: 38. Montalban Caves, Rizal Province, Luzon,
Philippine Is.

SPECIMENS EXAMINED. S Sulawesi: 2 99(1 yg.), d BM(NH) 81.11 10-1 1 12 Lalonggasu Meeto (Tomba
Watu Cave), 18 km from Kendari Central (in alcohol, skull of BM(NH) 81.1 1 12 extracted; coll. B. H.
Gaskell, 'Operation Drake').

REMARKS. These specimens have longer forearms and cranially are a little larger in some
respects than examples from Burma, Thailand, Malaya, Sumatra and Borneo. Moreover, the
rostrum is larger, more massive and less tapered than in specimens from more westerly
localities. The teeth of the male example, especially the canines, are a little larger than those
of the adult female. Measurements: length of forearm (rf BM(NH) 81.1112, 9 BM(NH)
81.1111) 76-2, 74-2; cranial dimensions (rf BM(NH) 81.1 1 12): greatest length of skull 35-8;

INDO-AUSTRALIAN BATS 133

greatest length of skull to canine 34-1; condylobasal length 34-0; condylocanine length 32-1;
basal length to canine 30-1; palatal length 18-5; least interorbital width 6-9; least postorbital
width 7-3; zygomatic width 22-4; width of braincase 14-3; mastoid width 13-8; width across
occipital crests 13-1; c'-c1 (alveoli) 7-5; m'-m1 (crowns) 9-2; (alveoli) 9-2; c-m2 13-0; length
complete mandible from condyles 26- 1 ; length right ramus from condyle 27-0; c-m3 14-2.

DISCUSSION. As Goodwin (1979) has pointed out, lack of material has led to some
uncertainty in the classification of this genus. Detailed studies of Eonycteris may be said to
begin with Andersen (1912), who recognised three species, E. spelaea (Dobson, 187 la & c),
widely distributed from Burma to Sumatra, Java, the larger E. major Andersen, 1910 from
northern Borneo, and, more doubtfully, E. rosenbergi Jentink, 1889 from Sulawesi,
considering the last to be perhaps an aberrant spelaea. Subsequently Miller (1913) described
E. robusta from Luzon Island, Philippine Islands. Chasen (1931) reported spelaea from
Borneo, while Taylor (1934) reported further specimens from Luzon as robusta and
described a second species, longicauda, from that island. However, Lawrence (1939) pointed
out that Taylor had applied robusta to a hitherto undescribed subspecies of E. spelaea for
which she proposed the name glandifera, and that in describing longicauda Taylor had in
fact renamed robusta of Miller. Lawrence also indicated that specimens from Sumatra had
no consistent characters separating them from E. s. glandifera, although Andersen (1912)
had earlier thought Sumatran Eonycteris to be the same as those from Burma.

Tate (1942c) discussed the genus in some detail, recognising (excluding rosenbergi from
Sulawesi) spelaea, major and robusta as distinct species. He concluded that two subspecies of
E. spelaea could be recognised: one, E. s. spelaea he thought to extend from Burma and
Vietnam through Malaya and Sumatra to east (sic) and central Java and possibly to Borneo
(cf. Chasen, 1931), the other, E. s. glandifera occurring in the Philippines, in Bali, perhaps in
eastern (sic) Java and probably in Borneo. Tate tentatively referred specimens from Bali to
E. s. glandifera, others from southern Sumatra emphatically to E. s. spelaea rather than to
glandifera as was done by Lawrence. More recently, Goodwin (1979) has provided an
account of a female specimen from Timor Island which on size and coloration also appears
to represent E. s. glandifera. Like Andersen (1912), Tate (loc. cit.) thought rosenbergi from
Sulawesi most probably based on an anomalous specimens of spelaea. He also suggested that
robusta ( — longicauda) represented the Bornean major in the Philippine Islands and greatly
extended the range of the,latter species by referring to it a specimen from North Pagi Island,
in the Mentawei Islands off west Sumatra.

The larger size and strong rostrum of specimens from Sulawesi suggests that they too
should be referred to E. s. glandifera. Moreover, the Sulawesian male has the distinct throat
band of long, tawny hairs described by Lawrence in glandifera and noted also in specimens
from Bali by Tate (1942c). However, this feature seems less exclusive than implied by Tate
who found no indication of this throat colouring in males of E. s. spelaea. Male specimens
from Sarawak in the collections of the British Museum (Natural History) have an ochraceous
tawny or even russet throat band although in other respects they agree closely with E. s.
spelaea from the Asian mainland that like those seen by Tate lack any such coloration.

The status of Callinycteris rosenbergi Jentink, 1889 ( — Eonycteris rosenbergi} remains
uncertain. It is known only from the immature holotype in the Rijksmuseum van
Natuurlijke Historic, Leiden, re-described in some detail by Andersen (1912). According to
this author rosenbergi differs from E. spelaea in the absence of the last lower molar (m3) on
each side of the jaw; as he points out, m3 is occasionally missing at least from one side of the
jaw in E. spelaea. Miller (1907) who had seen a photograph of the holotype remarked on its
heavy dentition. It is possible therefore that it is a young adult example of E. s. glandifera:
certainly the palate ridges as illustrated by Jentink are characteristically the same as those of
E. s. glandifera from Sulawesi. If this proves to be so, then rosenbergi must replace
glandifera as the earlier name. In discussing this question Tate (1942c) erroneously refers to
rosenbergi as bernsteini (a name actually used in the hipposiderid genus Coelops), thus
creating a nomen nudum.

134 J. E. HILL

Macroglossus minimus lagochilus Matschie, 1 899

Macroglossus lagochilus Matschie, 1899 : 96. Buru I, Molucca Is.

SPECIMENS EXAMINED. C Sulawesi: <f BM(NH) 81.1113 Ganda Ganda, 1°57'S, 121°21 ' E; Scfcf, 5 99
BM(NH) 81.1 1 14-1 123 R Ranu, 1° 51 ' S, 121° 21 ' E; rf, 9 BM(NH)81.1 124-1 125 Tambusisi Damar,
Mt. Tambusisi, c. 4000 ft, 1 ° 39 ' S, 1 2 1 ° 22 ' E (in alcohol; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Specimens from Sulawesi agree closely with those from the Molucca Islands and
from Borneo. Length of forearm in thirteen examples 38-4-41-9.

DISCUSSION. There is considerable taxonomic confusion in the genus Macroglossus, not least
resulting from its treatment by Andersen (1911, 1912), and identification difficulties have
until recently been common with specimens from the western part of its range in Malaysia,
Sumatra and Java. Mr A. N. Start (in litt.) has examined and studied Macroglossus from this
region, but his findings have yet to be published. In the meantime, however, his views have
influenced Lekagul & McNeely (1977) and Medway (1978) (or indirectly Ziegler, 1982) but
these authors give no justification for their adoption of combinations of names differing from
the generally used classification of Andersen (1912).

Thomas (1889) first recognised the existence of two species of Macroglossus when
examining specimens from Aola, Guadalcanal Island in the Solomon Islands. These he
distinguished readily from others from Java by the presence of a deep internarial groove
extending to the upper lip, much shorter face, i.e. rostrum, and smaller size, their forearm
lengths ranging from 38-43 mm, while in the Javan specimens the length of the forearm
ranged from 45-48 mm. These specimens from the Solomon Islands he wrongly
called M. australis (actually Syconycteris australis): in fact, they represent the form called M.
lagochilus by Andersen (1912). Those from Java Thomas called M. minimus: their size
refers them undoubtedly to the form called M. minimus sobrinus by Andersen (1911,
1912).

Andersen (1911, 1912) reviewed the genus in considerable detail. Basing his conclusions
on the directionality of the nares and the extent and depth of the internarial groove, this
author recognised two species, M. minimus, Indo-Chinese and Indo-Malayan, and M.
lagochilus, Austro-Malayan, but with no distributional overlap. According to Andersen,
minimus extended eastward through Java to Madura and Kangean Islands, perhaps to
Timor, while lagochilus extended westward to Borneo and the Philippine Islands. He also
recognised the existence of two forms in Java, one smaller, with shorter rostrum, which he
called M. minimus minimus, the other larger, with relatively longer rostrum, which he called
M. minimus sobrinus, this latter subspecies ranging to Sumatra, the Malay Peninsula,
Thailand, Burma and northeastern India. Specimens examined by Thomas (1889) clearly
represent this form, against which he compared lagochilus from the Solomon Islands. It is
also worth noting that many years earlier Temminck (1837) had remarked upon differences
in muzzle length in Macroglossus, recording that specimens from Sumatra had excessively
long muzzles; the muzzles of those from Java proved a little shorter and little different from
the muzzles of specimens from Timor, while others from Amboina Island had remarkably
short muzzles compared with those of Sumatran examples. Andersen (loc. cit.) regarded M.
minimus minimus as Javan, with M. m. sobrinus extending westward from Java through
Sumatra to the Asian mainland: he also thought that M. m. minimus might be found to occur
on Sumatra and in the Malay peninsula. Since Andersen wrote the Malaysian range of
lagochilus has been much extended, Chasen (1940) for example recording it from as far west
as Peninsular Thailand, Malaya and Nias Island, off west Sumatra, while Lekagul &
McNeely (1977) record specimens (as M. minimus) from southeastern Thailand and
Vietnam. It has not, however, been recorded from Java.

The directionality of the nares and the extent and depth of the internarial groove are
characters that are sometimes variable or subjective, distorted in preserved specimens and
sometimes difficult to observe. Goodwin (1979) for example was unable to discern any

INDO-AUSTRALIAN BATS 135

difference in the direction in which the nostrils face between Malayan sobrinus and
lagochilus from Malaya, Sulawesi and the Solomon Islands, although this author pointed out
that in lagochilus the margins of the nares are raised to show a tendency towards the develop-
ment of incipient tubes.

Specimens in the British Museum (Natural History) establish quite clearly the presence of
two forms in Java. The larger of these (length of forearm (4) 45-5^48-1, condylobasal length
(4) 26-4-26-9) has a long rostrum (length orbit-nares (5) 10-4-11-0) and the lower jaw
projects forward beneath the incisors to form a distinct sub-square chin. The nostrils are not
raised marginally to form an incipient tubular structure and the internarial groove is
represented at most by a narrow linear depression extending about halfway to the upper lip,
or is obsolete. These correspond to the form called sobrinus by Andersen (1911, 1912). The
smaller (length of forearm (19) 39-5-44-2, condylobasal length (9) 22-8-25-3) has a short
rostrum (length orbit-nares (10) 8-1-9-2) and the mandible slopes posteriorly beneath the
incisors, with rarely any suggestion of a squarish chin. The nostrils show no suggestion of a
tubular form, but the internarial groove extends as a narrow linear depression to the upper
lip, which is not divided. These correspond to the form called minimus Geoffrey, 1810a by
Andersen (loc. cit.). It is not certain to which of the two Javan forms Geoffrey's name refers
and the syntypes apparently are lost (Andersen, 1912) but for the present it seems both
convenient and sensible to retain Andersen's fixation of this appropriate name on to the
smaller form. Similar smaller specimens come from Madura and Kangean Islands. Tate
(1942c) clearly appreciated the differences between minimus and sobrinus, recording the
former from Cheribon, Java and from Bali, and the latter from North Pagi Island, Mentawei
Islands, off west Sumatra. Although direct sympatry has not been demonstrated the
conclusion that these represent distinct species seems unavoidable.

The collections also contain specimens of the larger form from Sumatra, Nias Island,
Simalur Island, Malaya, Thailand and the island of Koh Samui. These agree in every
essential point with the limited Javan representation: following Andersen (1912) they should
be referred to sobrinus which is considered to be a distinct species, ranging certainly from
Burma and Thailand south through the Malay Peninsula and Sumatra to Java. Length of
forearm (16) 44-2-49-8, condylobasal length (6) 25-4-28-6, and length orbit-nares (6)
10-1-11-7.

Medway (1978) differentiated the two forms in Malaya by the features of the head and
rhinarium: in the smaller, (hitherto called lagochilus, cf. Chasen, 1940) the head is shorter,
the nostrils forward pointing, with a distinct median internarial groove extending to the
upper lip and flanked by two smaller grooves while in the larger (sobrinus) he noted that the
head is longer, the nostrils more upwardly directed, and the median groove vestigial. He
called the smaller form M. minimus minimus, thereby implying the synonymy of lagochilus
with the Javan minimus. However, specimens from Malaya, Borneo, the Philippine Islands
and Sulawesi have the margins of the nares raised to form a slightly tubular structure as was
noted by Goodwin (1979) of specimens from Timor, and they have generally a more
prominent internarial groove than do specimens from Java. In these respects and in size they
agree with lagochilus from the Molucca Islands and for this reason I maintain lagochilus as a
distinct subspecies, extending from southern Thailand and southern Vietnam through the
Malay Peninsula to Borneo, the Philippines, Sulawesi, the Molucca Islands and Timor.
Apparently neither M. minimus minimus nor M. minimus lagochilus have been recorded
from Sumatra but Chasen (1940) reported the latter (with sobrinus) from the island of Nias,
off the west coast.

The species and subspecies of Macroglossus may be briefly summarised:

Macroglossus minimus (Geoffrey,
M. m. lagochilus Matschie, 1 899 S. Thailand, S Vietnam,

Malaya, Nias I,
Sirhassen I, (?) Sri
Buat I; (?) Bunguran I,

136 J.E.HILL

North Natuna Is; Borneo to
Philippine Is, Sulawesi,
Peleng I, Sanghir Is
and Molucca Is.

M. m. minimus (Geoffroy, 1 8 1 Oa) Java, Bali, Madura I,

Kangean Is.

M. m. nanus Matschie, 1 899 Aru Is, Kei Is, Mysol I,

New Guinea, Bismarck
Archipelago, Admiralty Is;
Queensland, Australia.

M. m. pygmaeus Andersen, 1911 Murray I, Torres Straits

M. m. microtus Andersen, 1911 Bougainville, San

Christobal, Guadalcanal
and Florida Is,
Solomon Is.

McKean (1972) considered M. m. pygmaeus and M. m. microtis to be synonyms of
M. m. nanus. His measurements, which do not include any of topotypes either of
pygmaeus or of microtis, certainly suggest that this may be so.

Macroglossus sobrinus Andersen, 1911

M. s. sobrinus Andersen, 1911 Burma and Thailand to

Sumatra, Nias I,
Krakatoa I and Java.

M. s.fraternus Chasen & Kloss, 1927 Sipora and Siberut Is,

Mentawei Is.

Macroglossus minimus nanus Matschie, 1899

Macroglossus nanus Matschie, 1899 : 98. Lamellana, New Britain, Bismarck Archipelago.

SPECIMENS EXAMINED. Papua New Guinea: 9 BM(NH) 73.1974 Victoria Bay, Kairiru ridge, NW end of
Kairiru I, near Wewak, East Sepik (in alcohol; coll. Aberdeen University Exploration Society
Expedition to New Guinea, 1973); $$BM(NH) 75.1840 Sepik, BM(NH) 75.1841 Rauit, West Sepik, 3°
36 ' S, 142° 1 5 ' E (both in alcohol; coll. A. M. Jones, Aberdeen University Exploration Society); (?) 9 9
BM(NH) 78.858-859 Sapi Creek Forest Reserve, near Baku, Gogol Valley, c. 40 m (in alcohol; coll. P.
A.Morris); 9 BM(NH) 80.530 Kuni, Morobe, 7° 22' S, 147° 1 1' E; 3 drf (1 yg.), 4 99(1 yg.) BM(NH)
80.53 1-537 Buso, Morobe, 7° 1 7 ' S, 147° 08 ' E (all in alcohol; coll. B. H. Gaskell, 'Operation Drake').
New Britain, Bismarck Archipelago: 9 cf BM(NH) 69.309-310 Kareeba Plantation, Keravat. 4° 18 '
S, 1 52° 01' E (skins, skulls; coll. J. I. Menzies).

REMARKS. Apart from the cranial and dental characters to which Andersen (1912) drew
attention, M. m. nanus is generally a little smaller than M. m. lagochilus, the length of the
forearm in nanus (13) 36-2-42-8, in lagochilus (32) 38-0-44-0. Specimens from the Aru
Islands and the Bismarck Archipelago are similar to those from New Guinea with length of
forearm (6) 36-3-38-7. Hitherto unrecorded examples in London include BM(NH) 37.2.16.1
from north of Eilanden R, West Irian, and BM(NH) 29.5.27.7 from Wasjor, also in West
Irian. A single specimen, BM(NH) 15.3.5.5 from Piara, Cape York, Queensland, Australia
agrees closely with nanus in size and length of rostrum rather than with M. m. pygmaeus
from Murray Island, Torres Straits. Tate (1952) also recorded M. m. nanus from Cape York:
McKean (1972) reports specimens from Australia.

Macroglossus sobrinus sobrinus Andersen, 1 9 1

Macroglossus sobrinus Andersen, 1911 : 642. Gunong Igari, Perak, Malaya.

INDO-AUSTRALIAN BATS 137

SPECIMENS EXAMINED. Krakatoa I, 06° 10' S, 105° 26' E: 9 BM(NH) 74.241 (in alcohol; coll. G.
Lincoln).

REMARKS. There appears to be no previous record of Macroglossus from Krakatoa: among
bats, Dammerman (1938) reported only Rousettus ample xicaudatus, Cynopterus brachyotis
angulatus ( = C. sphinx angulatus), Cynopterus horsfieldi minor ( — C. h. lyoni) and
Hipposideros diadema from the island. This specimen has slightly forwardly directed
nostrils and a moderate internarial groove that extends towards the upper lip as a narrow
linear depression, but does not reach it. The keel beneath the mandibular symphysis is
prominently developed into an anteriorly projecting, square chin. Length of forearm 45-7.

Macroglossus sobrinus fmternus Chasen & Kloss, 1927

Macroglossus minimus fraternus Chasen & Kloss 1927: 836, Sipora I. Mentawei Is, off west coast
Sumatra.

SPECIMENS EXAMINED. Siberut I, Mentawei Is, off west coast Sumatra: 9, 4cTcf BM(NH) 78.2921-2925
Base of Tetei Bulak, Sabeuleleu, Paitan R, offSaibi R (in alcohol; coll. J. J. Whitten).

REMARKS. Chasen & Kloss (1927) referred an adult male from Siberut to fraternus. These
specimens with length of forearm 49-3-50-9 agree in size with the original material examined
by these authors, differing from the nominate subspecies in their greater dimensions. In all
the nostrils open laterally and face to the side, and the internarial groove is obsolete or at best
is represented by a shallow linear depression that does not extend anteriorly beyond the
immediate internarial area. Possibly specimens reported by Tate (1942c) from North Pagi
Island, Mentawei Islands as M. minimus sobrinus ( = M. sobrinus sobrinus), of which he
notes some are very large, also represent M. s. fraternus

Syconycteris australis papuana (Matschie, 1899)

Macroglossus (Syconycteris) papuanus Matschie, 1899: 95, 99. Andai, NW West Irian.

SPECIMENS EXAMINED. Papu^ New Guinea: d, 9 BM(NH) 69.31 1-312 Schrader Mts., above Kaironk,
8500ft c. 5° 10' S, 144° 26 E; 99 BM(NH) 69.313-314 Efogi, Owen Stanley Mts., c. 40 miles E of
Port Moresby, 3000 ft, 9° 07 ' S, 147° 42 ' E (all skins, skulls; coll. J. I. Menzies); 6 dd, 4 99 BM(NH)
73.1975-1984 Biip, \ mile SE of Rauit, West Sepik, c. 1650ft, 3° 36' S, 142° 15' E: 4dd BM(NH)
73. 1985-1988 Rauit, 1750ft, 3° 36' S, 142° 15' E: 13 d <?, 8 99(1 yg.) BM(NH) 73.1989-2009 Kairiru
ridge, centre of Kairiru I, near Wewak, East Sepik, c. 2000 ft; 3rfd, 9 BM(NH) 73.2010-2013 Victoria
Bay, NW end of Kairiru I: d BM(NH) 73.2014 Sabor, E end of Kairiru I (all in alcohol); BM(NH)
74.339-374 Near Rauit, 3° 36' S, 142° 15' E (skulls only) (all coll. Aberdeen University Exploration
Society Expedition to New Guinea, 1973); d BM(NH) 75.1848 Wageo I. Schouten Is. East Sepik; 3 dd
BM(NH) 75.1 849-1 85 1 St. Xaviers, Kairiru I, East Sepik, c. 3° 2 1' S, 1 43° 36 ' E; 4 dd, 4 9 9 BM(NH)
75.1852-1859 Sepik (all in alcohol; coll. A. M. Jones, Aberdeen University Exploration Society); 3 dd,
2 99 BM(NH) 76.381-385 Okapa, Eastern Highlands, 6500 ft (in alcohol, coll. H. King); 9 BM(NH)
78.860 Baku, Gogol Valley, Madang (skin only); 99 BM(NH) 78.861-862 Baiyer R, c. 50 km NW of
Mt. Hagen, Western Highlands, 1300 m; 9 BM(NH) 78.863 Sapi Creek Forest Reserve, near Baku,
Gogol Valley, Madang, c. 40m; 99 BM(NH) 78.864-865 Baiyer R area, Western Highlands, c.
1300 m; d BM(NH) Mt. Hagen (town), Western Highlands (all in alcohol) (all coll. P. A. Morris); d
BM(NH) 80.538 7 km WSW of Buso, Morobe; d BM(NH) 80.539 Rasange, Morobe; 9 BM(NH) 80.540
Mt. Misson, Wau, Morobe; 15'dd, 7 99, 2 (?) BM(NH) 80.541-564 Buso, Morobe (all in alcohol; coll.
B. H. Gaskell, 'Operation Drake').

DISCUSSION. Andersen (1912) recognised three species of Syconycteris, separated only by
the features of the post-canine dentition, the last premolars (pm|) and first molar (mj) in S.
crassa (Thomas, 1895) being elongate and decidedly more than half as wide as long, those of
S. australis (Peters, 18670) linear, their width only half their length, while S. naias

138 J. E. HILL

Andersen, 1911 was distinguished from the latter by the absence of the last upper (m2) and
lower (m3) molars. At the time that Andersen wrote none had been found to be sympatric, S.
crassa extending from Amboina in the Molucca Islands to New Guinea and some associated
islands, S. australis occurring only in Queensland, and S. naias only on Woodlark Island in
the Trobriand Islands. Tate (1942c) reported both S. crassa and S. australis from New
Guinea, the two being collected together in the Central Division of Papua behind Port
Moresby: Laurie & Hill (1954) followed the arrangement proposed by Andersen and
implicitly adopted by Tate. Since then a highland species, S. hobbit Ziegler, 1982 has been
described.

Lidicker and Ziegler (1968) reported a high degree of variability in the number of
post-canine teeth in specimens of the australis type from the islands off southeastern New
Guinea, including Woodlark, with the post-canine formula varying from 5 to | and
exceptionally to f, as a result considering naias to be no more than subspecifically distinct
from australis. A similar incidence of dental anomalies, the absence of one to four molars, or
the presence of one or two supernumerary molars, was found by McKean (1972) in a large
number of specimens referred to australis or to crassa. This author considered naias and
australis synonymous, and pointed out that Brass (1959) had already recorded the latter,
apparently with normal dentition, from Woodlark Island. Moreover, McKean found that in
a large series of specimens from Papua New Guinea that he referred to crassa some had teeth
nearly as narrow as australis, while a few juveniles without fully developed teeth could easily
have been mistaken for that form. He concluded that but for the account by Tate (1942c) he
would have considered crassa and australis only subspecifically related.

When first examined at the British Museum (Natural History) in 1974 the series of skulls
(BM(NH) 74.339-374) led Hill (in Grieg-Smith, 1975) to the conclusion that crassa as
understood by Andersen (1912) and Tate (1942c) could not be separated from australis
since the dimensions of the relevant cheekteeth extended over most of the range of both
alleged species. This variation does not conform to the definitions of Andersen or of Lidicker
& Ziegler who redefined Andersen's dental criteria, pm| and mj according to these authors
being less than one and one half times as long as broad in crassa, but in australis being one
and one half times as long as broad, or more. Within the series now available pm4 varies in
length from approximately 1 10-200% of its width, pm4 from 140-230%, m1 from 100-180%
and m, from 150-200%. Moreover, on occasion in a single specimen pm4 and sometimes m1
conform to the broader, elongate outline considered characteristic of crassa while pm4 and
m, are narrower and more linear, their proportions more nearly those ascribed to australis.
The variation in tooth proportions is such that there is no point at which a division into two
species on the basis of this character can be made.

Lidicker & Zeigler (1968) add that in crassa m2 and m3 are normally present although
sometimes absent in australis, that the inner upper incisors tend to be slightly separated in
crassa but in contact in its congener, and that the latter has shorter fur. McKean (vide supra)
has demonstrated a wide degree of variation in the number of post-canine teeth in specimens
from eastern New Guinea and this topic is also discussed in some detail by Koopman (1982)
who concludes that molar number is not a good species character: among BM(NH)
74.339-374 only one specimen (74.344) has an obviously anomalous dental formula, lacking
m2 and m3 on both sides. In this specimen, pm4 and m1 are sub-circular, pm4 and nij rather
more linear in outline, thus in the upper jaw conforming to the definition of crassa, in the
mandible tending towards australis.

The series includes specimens which have the inner upper incisors convergent, in contact,
very slightly separated, or not at all convergent and more distantly separated by a narrow but
obvious interspace. However, this variation does not correlate with the features of pm| and
mj. At the extremes, BM(NH) 74.339 for example has inner upper incisors that stand
separately but has strongly linear pm| and mj, while BM(NH) 74.343 has convergent incisors
that touch but sub-circular pm4 and m1, their mandibular counterparts only slightly elongate
and not at all linear. Koopman (1979) could find no dichotomy into broad and narrow
toothed forms, while Ziegler (1982) in describing the montane species S. hobbit examined six

INDO-AUSTRALIAN BATS 139

of the specimens referred to australis or to crassa by Tate (1942c) and was unable to find any
consistent differences in tooth dimensions that would differentiate the two putative species.
Moreover, Ziegler also could not establish any external, cranial or dental characters to
suggest such a distinction among the extensive representation of relevant Syconycteris in the
Bernice P. Bishop Museum, Hawaii.

The recognition that australis is conspecific with crassa means that as the prior name
australis must become the specific epithet and possibly the subspecific name for the
population in New Guinea usually called papuana. However, Koopman (1979) remarked
that Australian specimens of which he had seen a limited sample tend to be smaller than
those from New Guinea and tentatively retains australis and papuana as distinct subspecies.
The remaining subspecies of S. australis also seem only slightly differentiated. They include
crassa (Thomas, 1895) from the islands southeast of New Guinea, keyensis Andersen, 1911
from the Kei Islands, major Andersen, 1911 from Amboina and Ceram Islands, in the
Molucca Islands, and finschi (Matschie, 1 899) from the Bismarck Archipelago. McKean
(1972) synonymised keyensis and finschi with papuana, and regarded naias as a synonym
of australis ( = S. australis australis), Lidicker & Ziegler (vide supra) having considered the
latter two to be subspecifically related as does Koopman (1982), who carried out a detailed
examination of specimens from eastern Papua and its associated islands. However, Koopman
(1979) retained finschi on account of its slightly smaller size when compared with papuana:
measurements of BM(NH) 74.339-374 (condylobasal length 23-2-24-8, width ofbraincase
10-2-1 1-4) agree with those of New Guinea specimens measured by this author and support
his contention. Possibly naias from Woodlark Island is synonymous with australis but may
prove to be another weakly separable subspecies.

According to Zeigler (1982) the dorsal pelage of S. australis in any area is apparently
slightly less dense and less woolly, inter alia, than that of S. hobbit, the two so far being found
to be sympatric only at Mount Kaindi, Morobe, Papua New Guinea. The two specimens
(BM(NH) 69.31 1-312) of S. australis reported here from the Schrader Mountains at 8500 ft
have longer, slightly denser fur than do those from lower locations, and are also a little darker
in overall coloration, thus confirming the observation by Ziegler that high level examples
were the darkest to be found in the species. It seems possible that a high altitude subspecies of
S. australis, which according to Ziegler occurs as high as at least 3000 m, might be recognised
in due course.

m

Syconycteris australis major Andersen 1911

Syconycteris crassa major Andersen, 1911: 643, Amboina I. Molucca Is.

SPECIMENS EXAMINED. Molucca Is: dd BM(NH) 81.1 126-1 127 Kaiteto, Amboina I (in alcohol; coll.
B. H. Gaskell, 'Operation Drake').

REMARKS. In length of forearm (46-0, 47-5) these specimens fall within the range given by
Andersen ( 1 9 1 1 , 1 9 1 2) for this larger subspecies.

Notopteris macdonaldi macdonaldi Gray, 1859

Notopteris macdonaldi Gray, 1 859 : 38, pi. 67. Viti Levu I, Fiji Is.

SPECIMENS EXAMINED. Fiji Is: 6 dd, 6 99 (1 yg.) BM(NH) 78.2394-2405 Saweni Navosi/Koro Bulia,
Sigatoka Valley, Viti Levu I, 1 7° 39 ' S, 1 77° 30 ' E (skins, skulls; coll. J. C. Pernetta).

New Hebrides: dd BM(NH) 73.1319-1320 (1 yg.) Analghaut Village, Aneityum I, near sea level; 3
dd, 9 BM(NH) 73.1321-1324 About 2 km E of 'Bethel', Tanna I, 120m; 3 dd (2 nurselings), 4 99,
1 (?) BM(NH) 73.1325-1332 Watantup, about 4 miles N of Ipota, east coast Erromanga I, at sea level;
d BM(NH) 73.1333 Nokowoula, Espiritu Santo I, 3700ft (all in alcohol except BM(NH)
73.1331-1332, skins; coll. or obtained by the Earl of Cranbrook, Royal Society Expedition to the New
Hebrides, 1971).

140 J. E. HILL

REMARKS. Specimens obtained by the Royal Society Expedition in 1971 extend N.
macdonaldi to most of the major islands of the New Hebrides, whence before it has been
known in the literature apparently only from Aneityum (Andersen, 1912). The collections of
the British Museum (Natural History) also include two females (BM(NH) 26.6.4.7-8) from
Efate Island.

DISCUSSION. The two subspecies of N. macdonaldi, N. m. macdonaldi (Fiji Is and New
Hebrides) and N. m. neocaledonica Trouessart, 1908 (New Caledonia) differ chiefly in the
smaller size of the latter (Andersen, 1912, Revilliod, 1914, Sanborn & Nicholson, 1950):
specimens in the collections of the British Museum (Natural History) suggest also that
generally neocaledonica has a lower, rather less massive rostrum than macdonaldi.
Measurements given by Revilliod (1914) and by Sanborn & Nicholson (1950) when
comparing Fijian macdonaldi with neocaldonica indicate a considerable difference in
forearm length that is not entirely supported by specimens in London. Sanborn & Nicholson
in fact suggested that the forearm length of Fijian macdonaldi is always longer than in
neocaledonica, although they had seen no adult male of this latter subspecies. These authors
indicated that in Fijian macdonaldi the length of the forearm did not fall below 63-9 and that
in neocaledonica it did not exceed 61-5. However, a series of specimens (BM(NH)
19.10.8.5-15) from Mount Tambignon, New Caledonia, at 2500 ft have forearms ranging in
length from 60-2-63-4 in males and in females from 60-3-64-3.

Specimens from the Fiji Islands and from the New Hebrides tend to further bridge the
alleged size difference. Six Fijian males range in forearm length from 63-5-68-8 (but Sanborn
& Nicholson (1950) report a male of 71-3), five females from 63-0-66-0. In the New Hebrides
a male from Aneityum has a forearm length of 69-9 and a female (BM(NH) 25.12.14.1 1) of
65-1. Three males from Tanna have forearms varying in length from 65-0-69-0, a female a
forearm length of 62-8. The adult male from Erromanga has a forearm length of 61-5, four
females from the same island having forearms ranging in length from 61-6-63-6, while the
adult but not old male from Espiritu Santo has a forearm length of 64-8.

Sanborn & Nicholson (1950) reported the presence of a strong sagittal crest extending from
the frontals to well-developed lambdoidal crests in males of N. m. macdonaldi, females
lacking a sagittal crest but possessing similar well-developed lambdoidal crests. A slightly
less developed sagittal crest is also present in the older males of N. m. neocaledonica.

MICROCHIROPTERA
EMBALLONURIDAE

Emballonura alecto alecto (Eydoux & Gervais, 1836)

Vespertilio (Nycticeius) alecto Eydoux & Gervais, 1 836 : 7. Manila, Luzon I, Philippine Is.

SPECIMENS EXAMINED. C Sulawesi: <td BM(NH) 82.1-16 Tapu Waru, 1° 55 ' S, 121° 22 ' E; 9 BM(NH)
82.17GandaGanda, 1°57' S, 121° 21' E; 9, <J BM(NH) 82.18-19 Tarongga, 1°44' S, 121° 40' E (all
in alcohol, skulls of BM(NH) 82. 1-5, 82. 1 3 extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These specimens are referred to E. alecto rather than to the closely similar species
E. monticola Temminck, 1838 by virtue of their rather large skulls which have a relatively
long antemolar region, with a marked diastema between the first upper premolar (pm2) and
the second upper premolar (pm4). Tate & Archbold (1939) recorded E. alecto from
Likeopang, north Sulawesi and from Peleng Island on the basis of specimens in the United
States National Museum of Natural History, Washington. These are evidently those reported
among others (in this case from Limpoeang) by Shamel (1940) as E. monticola rivalis
Thomas, 1915c ( = E. alecto rivalis). Although comparative material is limited, specimens
from Sulawesi and the Molucca Islands agree most closely in several points of size with
examples from the Philippine islands and are referred to the nominate subspecies.

INDO-AUSTRALIAN BATS 141

Emballonura monticola also occurs in Sulawesi whence it has been recorded from the
southern part of the island by Tate & Archbold (1939).

Measurements of specimens from Sulawesi: length of forearm (19) 43-1-48-3; greatest
length of skull (6) 14-7-15-1; condylobasal length (6) 13-5-14-0; condylocanine length (6)
12-9-13-2; width of rostral swellings (6) 5-9-6-2; least interorbital width (4) 4-7-5-0; least
postorbital width (6) 2-7-3-0; zygomatic width (5) 8-7-9-1; width of braincase (6) 6-8-7-1;
mastoid width (6) 7-5-8-2; cl-cl (alveoli) (6) 3-6^-0; m3-m3 (6) 6-1-6-4; c-m3 (6) 5-5-5-7;
length complete mandible from condyles (4) 9-8-10-0; length right ramus from condyle (6)
10-1-10-6; c-m3 (6) 5-6-5-9.

Emballonura nigrescens papuanus Thomas, 1914

Emballonura papuanus Thomas, 19146 : 443. Wakatimi, Mimika R, SW west Irian.

SPECIMENS EXAMINED. Papua New Guinea: d BM(NH) 80.606 Buso, Morobe, 7° 17' S, 1 47° 08 ' E (in
alcohol; coll. B. H. Gaskell, 'Operation Drake').

C Sulawesi: rf BM(NH) 82.20 R Ranu, 1° 51' S, 121° 30' E (in alcohol, skull extracted; coll. B. H.
Gaskell, 'Operation Drake').

REMARKS. The short, blunt muzzle, widely separated elliptical nostrils and long, narrow
tragus with backwardly directed tip quite clearly refer these specimens to E. nigrescens. The
example from Sulawesi has a slightly larger and more inflated braincase than E. n. nigrescens
(Gray, 1843a from the Moluccan islands of Amboina, Buru and Ceram and on this account
is referred to E. n. papuanus, otherwise known from Ternate Island, New Guinea and the
Schouten and Kei Islands. According to Thomas (1914&) the rostrum of E. n. papuanus,
is also short and stumpy but this feature is not at all obvious in the series in the British
Museum (Natural History). Tate & Archbold (1939) and Shamel (1940) have also recorded
the subspecies from Sulawesi. Measurements of the Sulawesian example BM(NH) 82.20:
length of forearm 32-2; greatest length of skull 1 1 -0; condylobasal length 10-4; condylocanine
length 10-2; width rostral swellings 4-0; least interorbital width 3-0; least postorbital width
2-3; zygomatic width — ; width of braincase 6-0; mastoid width 6-2; ^-c1 (alveoli) 3-0; m3-m3
5-1; c-m3 4-2; length complete mandible from condyles — ; length right ramus from condyle
7-9; c-m3 4-4.

Taphozous melanopogon Temminck, 1 84 1

Taphozous melanopogon Temminck, 1841 : 287, pi. 60, figs. 8, 9. Bantam, W Java.

SPECIMENS EXAMINED. S. Sulawesi: 99 BM(NH) 82.12-22 Lalonggasu Meeto, (Tomba Watu Cave),
18 km from Kendari Central (in alcohol, skulls (of BM(NH) 81.22 fragmentary) extracted; coll. B. H.
Gaskell, 'Operation Drake').

REMARKS. Although these specimens have relatively long forearms, the distribution of the
fur at the sides of the body, unlengthened rostrum, rather narrow, elongate braincase and
smaller teeth, especially the canines, suggests that they should be referred to T. melanopogon
rather than to the closely similar species T. theobaldi Dobson, 1872a. The latter was
reported from Java by Thomas & Wroughton (1909) on the basis of specimens now in the
collections of the British Museum (Natural History): these, although smaller than a very
limited sample of T. theobaldi from India, Burma and Thailand have longer forearms
(71-1-72-5 in ten examples) than the specimens from Sulawesi here referred to T.
melanopogon, and their skulls are generally larger. Unfortunately, few specimens of T.
melanopogon are known from Java: the species occurs otherwise from India and southern
China to the Philippine Islands and the islands of Sumbawa, Savu and Timor in the Lesser

142 J. E. HILL

Sunda chain. These from Sulawesi are the first of T. melanopogon to be recorded from that
island: in size they are similar to those reported by Goodwin (1979) from Timor, or to T. m.
achates Thomas, 1 9 1 5b from Savu Island.

Measurements (BM(NH) 82.21, 82.22): length of forearm 68-5, 67-7; greatest length of
skull — , — ; condylobasal length — , — ; condylocanine length 21-6, — ; least interorbital
width 64, 6-4; least postorbital width 5-3; — ; zygomatic width 13-1, — ; width of braincase
10-3, — ; mastoid width 11-6, — ; c'-c1 (alveoli) 4-3, — ; m3-m3 9-3; — ; c-m3 9-8, 9-5; length
complete mandible from condyles 16-8, 16-4; length right ramus from condyle 17-4, 16-9;
c-m3 10-7, 10-4.

DISCUSSION. The genus Taphozous has remained unrecorded from Sulawesi until
comparatively recently, although its overall Oriental distribution from the Indian sub-
continent to the Philippine Islands and Australia clearly indicates that it might be expected
to occur on that island. Apart from T. (Taphozous) melanopogon, the pouch-bearing bat T.
(Saccolaimus) saccolaimus Temminck, 1841 has been reported lately from Sulawesi, Fieler
(1980) having identified an old specimen dating from 1883 in the Staatliches Museum fur
Tierkunde, Dresden as an example of this species. It has been known to occur hitherto from
India to Borneo, Java and Timor, and may extend to New Guinea, the Solomon Islands, the
Northern Territory of Australia and northeastern Queensland if nudicluniatus De Vis, 1905
is conspecific with saccolaimus as Goodwin (1979), McKean et al. (1980) and Koopman
(1982) suggest.

MEGADERMATIDAE

Megadema spas ma niasense Lyon, 1916

Megaderma niasense Lyon, 1916 : 440. Nias I, off W Sumatra.

SPECIMENS EXAMINED. Siberut 1, Mentawei Is: dd BM(NH) 78.2926-2928 Near Sibosua R, Paitan R, off
Saibi R; 99 BM(NH) 78.2929-2931 Ridge to the E of Sibosua R (all in alcohol, skulls of BM(NH)
78.2929-2930 extracted; coll. J. J. Whitten).

REMARKS. There is no previously published record of M. spasma from Siberut although the
species has been recorded widely (Lyon, 1916, Chasen, 1940) from the island chain of which
Siberut forms a part. These specimens agree in many respects with the description of
naisense from the island of Nias to the northwest of Siberut: they are a little smaller on the
whole than M. s. trifolium Geoffrey, 18106 from Java and southern Sumatra, with the parts
of the maxillae over the canines less enlarged and the tympanic bullae and teeth generally
smaller. External measurements: length of forearm (6) 55-9-58-8; length of ear (6) 32-0-33-4;
length of tibia (6) 31-1-32-2. Cranial measurements (BM(NH) 78.2929, 78.2930): greatest
length of skull 24-8, 25-5; condylocanine length 21-5, 22-8; least interorbital width 3-9, 4-0;
zygomatic width 13-6, 14-1; width of braincase 10-5, 10-3; mastoid width 11-0, 11-5; c'-c1
(alveoli) 4-8, 5-1; m3-m3 8-0, 8-3; c-m3 9-2, 9-9; length complete mandible from condyles
16-1, 17-3; length right ramus from condyle 16-7, 17-9;c-m3 10-1, 10-9.

Megaderma spasma celebensis Shamel, 1940

Megaderma spasma celebensis Shamel, 1940 : 352. Likoepang, Sulawesi.

SPECIMENS EXAMINED. C Sulawesi: 3 9 9, 2 dd BM(NH) 82.23-27 GandaGanda, 1° 57' S, 121° 21' E; 5
cfrf,3 99 Songinbau, 1°46' S, 121°43' E (all in alcohol, skulls of BM(NH) 82.24-27, 82.29-31, 82.33,
82.35 extracted; coll. B. H. Gaskell, 'Operation Drake').

INDO-AUSTRALIAN BATS 143

REMARKS. Andersen & Wroughton (1910) and Andersen (1918) referred specimens of M.
spasma (Linnaeus, 1758) from Sulawesi and from the Philippine Islands to the nominate
subspecies, also known from its type locality, Ternate Island in the northern Moluccas.
Later, Shamel (1940) separated Sulawesian material as M. s. celebensis on account of its
generally smaller skull and the absence of any distinct contrast in colour between the dorsal
and ventral surfaces of the body. Tate (1941&) who evidently had not seen the account by
Shamel considered specimens from Sulawesi inseparable from those from Java, presumably
M. s. trifolium.

This series of Sulawesian specimens from 'Operation Drake' confirms the small cranial
size noted by Shamel (1940) when describing celebensis: all agree in size with those from
Sulawesi measured by this author and are rather smaller than the Philippine specimens that
he examined. They are also somewhat smaller than a series of M. s. trifolium from Java.

Measurements: length of forearm (13) 52-2-57-1; length of ear (13) 35-3-37-4; length of
tibia (13) 29-2-30-5; greatest length of skull (9) 23-9-24-4; condylocanine length (9)
21-1-21-6; least interorbital width (9) 3-7-4-0; zygomatic width (9) 13-3-14-0; width of
braincase (9) 9-9-10-7; mastoid width (8) 10-5-11-0; c'-c1 (alveoli) (9) 4-6-5-0; m3-m3 (9)
7-4-8-1; c-m3 (9) 8-8-9-3; length complete mandible from condyles (9) 15-4-16-1; length
right ramus from condyle (9) 16-2-16-6; c-m3 (9) 9-9-10-3. Fourteen examples of M 5.
trifolium from Java have a condylocanine length of 21-8-23-4 and a length c-m3 of 9-3-9-8:
three from Kangean Island are of similar size with corresponding measurements of 2 1 -8-22-5
and 9-2-9-5.

RHINOLOPHIDAE

Rhinolophus celebensis celebensis Andersen, 1 905

Rhinolophus celebensis Andersen, 1905a : 83, pi. 3, figs. 4a, 4b, Makassar, S Sulawesi

SPECIMENS EXAMINED. C Sulawesi: 9 BM(NH)82.36 R Ranu, 1°51 ' S, 121° 30' E; 5 rfd, 5 99 BM(NH)
82.37-46 Songinbau, 1°46' S, 121° 43' E; 99 BM(NH) 82.47-48 Taronggo, 1°44' S, 121° 40' E: d,
9 BM(NH) 82.49-50 Kabalo, Mt. Tambusisi, c. 300 m, 1° 40' S, 121° 20' E (all in alcohol, skulls of
BM(NH) 82.36-37, 82.39-40, 82.48, 82.50 extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These specimens agree favourably with the holotype ofR. c. celebensis and with
the limited sample of this taxon in the collections of the British Museum (Natural History).
They supplement the records of celebensis from various localities in Sulawesi by Andersen
(19050), Tate & Archbold (1939) and Bergmans & Rozendaal (1982). Measurements: length
of forearm (15) 39-7-43-4; condylocanine length (6) 15-5-16-0; supraorbital length (junction
of supraorbital crests to nares) (6) 5-0-5-1; palatal length (6) 2-0-2-3; length rostral inflations
(6) 1-9-2-1; width rostral inflations (6) 3-1-3-6; width of rostrum (6)4-6-5-0; least interorbital
width (6) 2-1-2-4; zygomatic width (6) 8-7-9-0; width of braincase (6) 7-4-7-7; mastoid width
(6) 8-4-8-7; c'-c1 (alveoli) (6) 4-2-4-6; m3-m3 (6) 6-3-6-5; c-m, (6) 6-8-7-1; length
complete mandible from condyles (5) 11-6-11-7; length right ramus from condyle (5)
1 1-9-1 2-0; c-m3 (6) 6-8-7-1.

DISCUSSION. Goodwin (1979) in describing parvus from the island of Timor discussed the
relationships of celebensis to borneensis Peters, 1861 from Borneo and to javanicus
Andersen, 1918 from Java, concluding that all belonged to the same species, R. borneensis.
This author remarked that the most useful taxonomic features of the skull in the borneensis
group (sic) are the size and shape of the anterior median nasal inflations, the width of the
interorbital constriction, and the position of the junction of the supraorbital crests,
characters also used for the most part by Andersen ( 1 905# ) in his classic study of the complex
to which these forms belong, the simplex group of Andersen, \9Q5a or megaphyllus group of
Andersen, 1918, later renamed the ferrumequinum group by Tate & Archbold, 1939. Apart
from those already mentioned, named taxa immediately relevant to this question also

144 J. E. HILL

include importunus Chasen, 1939 from eastern Java, madurenesis Andersen, 1918 from the
nearby island of Madura, spadix Miller, 1901 from the South Natuna and Karimata Islands
and chaseni Sanborn, 1939 from Con Son Island (Pulo Condore), off the southeastern coast
oflndochina.

The small series ofcelebensis reported here confirms the general similarity to borneensis
suggested by Andersen (1905a) and formalised by Goodwin (1979) in considering the two
conspecific. There is some variability in the size and shape of the anterior median narial
inflations: the holotype (BM(NH) 97. 1.3.19) ofcelebensis has rather short inflations which in
other examples are longer antero-posteriorly. On the whole the anterior narial inflations are
a little smaller than those of a limited sample of borneensis but the smallest of borneensis are
closely similar to the largest ofcelebensis. The narial inflations ofcelebensis are less inflated
than those of R. keyensis Peters, 1871 from the Molucca Islands to the east, although
generally similarly narrow, and in this species the facial part of the skull tends to be longer
than in celebensis, with the junction of the supraorbital and the sagittal crests lying at a point
more or less behind the middle of the orbit rather than more or less in front of it. As Goodwin
(1979) pointed out, borneensis differs more from celebensis and its immediate allies
javanicus and parvus than these do from each other in its more strongly arched fronto-
parietal area and more bulbous occipital region, although these characters are less clear when
series are examined. In addition to the generally larger and wider narial inflations and more
inflated braincase, borneensis and its close allies spadix and chaseni are usually a little larger
cranially than any of the more easterly forms. Goodwin (1979) omitted any mention of
madurensis in his account of parvus, to which in fact it is similar in size and structure.
Comparison suggests that details of coloration apart (the sole parvus available is in alcohol)
the two may be separated only by the slightly less globular narial inflations of madurensis.
It should be noted that the 'condylocanine length' given by Goodwin (1979 : 107, tab. 1)
for parvus, javanicus, celebensis and borneensis is in fact the greatest length of the skull.

Through the courtesy of Dr C. J. Smeenk of the Rijksmuseum van Naturlijke Historic,
Leiden it has been possible to examine one (RMNH 15320) of the original specimens of
importunus Chasen, 1939 from Tjiawitali, near Wijnkoops Bay, east Java. Further
comparison confirms the view expressed by Chasen in the original account that cranially it is
very like borneensis, with which it agrees in size, inflated braincase and bulbous occipital
region, differing in these features from javanicus, known so far from west Java. As Chasen
pointed out, it is clearly very closely related to borneensis to which it is very similar.
Although admitting this close relationship and also indicating clearly that it could not be
placed with javanicus he was reluctant to unite importunus with borneensis largely because
the known occurrences of importunus in eastern Java and of javanicus in western Java might
imply that they replaced each other geographically. However, further examination leaves
little doubt that importunus must be allied with borneensis.

For these reasons and in view of the limited material as yet available for some of the forms
involved, I prefer to retain borneensis and celebensis as distinct species, the latter to include
javanicus, madurensis and parvus as subspecies, although the very small size of madurensis
and parvus might justify their specific distinction (as madurensis) from celebensis.
Rhinolophus borneensis includes spadix, chaseni and importunus as subspecies, the last
possibly occurring sympatrically with R. celebensis javanicus in Java. Possibly the very
small virgo Andersen, 1905<2 from Luzon Island, Philippine Islands should also be associated
with R. celebensis, while nereis Andersen, 1905a from Pulo Siantan, Anamba Islands
evidently represents R. borneensis, but is considerably larger.

Rhinolophus pusillus pusillus Temminck, 1834

Rhinolophus pusillus Temminck, 1 834 : 28, pi. 1 , fig. 9. Java.

SPKCIMENS EXAMINED. Borneo: d BM(NH) 78.2491 About 25 km inland from Sangkulirang, a small
port at the mouth of the R Baa, East Kalimantan, c. 0° 59 ' N, 1 17° 55 ' E (in alcohol, skull extracted;
coll. S.M.Jeffrey).

INDO-AUSTRALIAN BATS 145

REMARKS. Although known from India and southern China to the Anamba Islands, Java and
Madura Island, R. pusillus has not been recorded hitherto from Borneo. In size this specimen
agrees closely with those from Java and the smaller islands of Madura, Tioman and Penang
recorded by Hill (1974) and also with the skin from Aur ( = Aor) Island reported earlier by
this author (1960) as of R. minutillus Miller, \906a ( = R. pusillus minutillus, originally
described by Miller (1900) as R. minutus) from Siantan Island, Anamba Islands. Specimens
from the Malayan offshore islands, Madura Island and Borneo all seem referrable to the
nominate subspecies R. p. pusillus.

Measurements of BM(NH) 78.2491: length of forearm 37-0; greatest length of skull to
canine 15-6; condylocanine length 13-8; width across rostral swellings 4-1; least interorbital
width 2-1; zygomatic width — ; width of braincase 7-0; mastoid width 7-9; c'-c1 (alveoli) 3-8;
m3-m3 5-8; length complete mandible from condyles 9-9; length right ramus from condyle
10-3; c-m3 6-2.

Rhinolopus euryotis tatar Bergmans & Rozendaal, 1982

Rhinolophus tatar Bergmans & Rozendaal, 1982 : 170. Moinakom River, Dumoga Nature Reserve,
North Sulawesi, 525 m, 0° 41' M. 124° 03 ' E.

SPECIMENS EXAMINED. C Sulawesi: 9 drf, 17 99 BM(NH) 82.51-76 Songinbau, 1°46' N, 121° 43 ' E; 12
d-d1, 7 99 BM(NH) 82.77-95 Taronggo, 1°44' N, 121° 40' E; 9 BM(NH) 82.96 R Ranu, P51' S, 121°
30' E (all in alcohol, skulls of BM(NH) 82.51, 82.53-55, 82.78-82, 92.84-88, 82.90 extracted; coll. B.
H. Gaskell, 'Operation Drake').

S Sulawesi: dd BM(NH) 82.97-98 Lalonggasu Meeto (Tomba Watu Cave), 18 km from Kendari
Central (in alcohol, skull of BM(NH) 82.98 extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. This subspecies (length of forearm 48-4-53-2) includes the smallest of/?, euryotis,
with broad noseleaf that covers the muzzle, the anterior leaf with a very shallow
emargination, the margins of this notch thickened, the thickening extending posteriorly as
narrow ridges over the leaf to delimit a shallow longitudinal median groove terminating at
the edge of the narial depression in a small, thickened, posteriorly directed projection. The
internarial region is expanded laterally into a wide, angular cup and the sella is wide, slightly
ovate-pyriform in frontal outline. Its connecting process is high, arcuate and inserted
anteriorly at or near the upper margin of the sella, with a lower insertion posteriorly on the
intermediate part of the leaf. The lancet is high, cuneate, with an acute point, and is densely
pilose. The ears are large, extending to the tip of the muzzle when laid forward; their anterior
or medial margin is convex beneath a small, acutely pointed tip, their posterior margin
concave just beneath tip, otherwise convex with a well developed more or less rectangular
antitragal lobe.

The skull is relatively small with short, broad rostrum and elongate braincase, the anterior
median rostral inflations relatively large, swollen, projecting, high and rounded and there is a
shallow triangular frontal depression bounded laterally by moderate suprorbital ridges. The
median cranial crest is well developed, more evident anteriorly than posteriorly and the
zygomata are not greatly expanded, the zygomatic width subequal to or only slightly exceed-
ing the mastoid width. The palate is short, its length one third or less the length of the
maxillary toothrow and the cochleae are expanded, only narrowly separated basioccipitally.
The anterior upper premolar (pm2) is small, in the toothrow, not compressed and the second
lower premolar (pm3) is very small, extruded from the row.

Externally this subspecies may be distinguished from the other described forms of R.
euryotis by its shorter forearm, generally smaller size, and, for the most part, by its narrower
sella which is usually less expanded laterally and often less ovate-pyriform than in the other
members of the species. In the majority of its features it is most similar to R. e. aruensis
Andersen, 1907/7 (Aru Islands) or to R. e. burius Hinton, 1925 (Buru Island, Molucca
Islands) but has a less massive and narrower rostrum, slightly less inflated braincase and
slightly smaller and less massive teeth, especially the canines. Its shorter forearm and

146 J. E. HILL

narrower sella provide obvious features distinguishing tatar from R. e. euryotis Temminck,
1835 (Amboina and Ceram Islands, Molucca Islands), R. e. timidus Andersen, 19056
(Batchian Island and New Guinea) and from R. e. praestans Andersen, 19056 (Kei Islands):
in these, especially in R. e. euryotis, the noseleaf is more fleshy and the features of the
antenarial region more greatly developed, with a more emphatic longitudinal groove across
the anterior leaf terminating in a larger posterior projection.

DISCUSSION. Tate & Archbold (1939) provided the first reports of/?, euryotis from Sulawesi,
recording specimens from the south and southeast of the island. These authors thought
Sulawesian specimens most probably referable to R. e. euryotis, although apparently they
had not examined specimens from Amboina, its type locality, or any of the other associated
islands in the Moluccas. They relied instead on the description of euryotis by Temminck
(1835) and in particular upon his statement that the forearm in Amboinese specimens
measured '2 pouces' which they converted as 50-8 mm. They remarked that the treatment by
Andersen (19056) of euryotis was somewhat invalidated by the assumption that the forearm
of the nominate subspecies was 56 mm in length, against this lower figure given by its
describer, and postulated larger subspecies (praestans, timidus and burius) in the Kei Islands,
Batchian and New Guinea, and on Buru, with smaller subspecies (aruensis, euryotis) in the
Aru Islands and in Amboina and Sulawesi. Specimens reported by these authors from
Sulawesi agree closely in size with tatar as described by Bergmans & Rozendaal (1982) and
with the specimens reported here.

A small series of R. euryotis in the collections of the British Museum (Natural History)
from Amboina and the nearby island of Ceram has forearm lengths ranging from 55-1-584,
considerably larger than the value assumed by Tate & Archbold (1939) for Amboinese
specimens. In fact, these authors have converted Temminck's measurements as English
pouces (one pouce = 254 mm), but Temminck used the French equivalent (one
pouce = 27-07 mm) which when converted gives a value of 54-1 mm, much closer to the
forearm lengths of Amboinese specimens available in London. This deduction is confirmed
by Bergmans & Rozendaal (1982) who quote approximate forearm lengths (measured by Dr
C. J. Smeenk) of 54-9, 55-1 and 54-5 for the three syntypes of euryotis in the Rijksmuseum
van Natuurlijke Historic, Leiden.

The specimens reported here agree exactly with the description of tatar by Bergmans &
Rozendaal, who considered it a distinct species, morphologically nearest to R. arcuatus
Peters, 1871. This latter species is widely distributed from Luzon Island (arcuatus) and
Mindanao Island (exiguus Andersen, 19056) in the Philippine Islands to Borneo
(proconsulis Hill, 1959), Sumatra (beccarii Andersen, 1907c), Buru Island in the Molucca
Islands (toxopeusi Hinton, 1925), Wetter Island in the Flores Sea (angustifolius Sanborn,
1939) (thought probably a distinct species by Bergmans & Rozendaal, 1982) and New
Guinea (mcintyrei Hill & Schlitter, 1982). Although very similar both externally and
cranially to R. euryotis, its anterior noseleaf lacks the characteristic broad median groove of
this species flanked by swollen longitudinal ridges and extending across the leaf to the inter-
narial region, where it terminates at a small projection. Instead, the narrow anterior
emargination of the leaf is prolonged posteriorly as a narrow linear groove onto the face of
the leaf, this groove extending less than halfway to the internarial region. Moreover,
although the largest of R. arcuatus (proconsulis, mcintyrei) have forearm lengths that reach
or overlap those tatar the skull is slightly smaller, with smaller teeth.

When first examined the specimens reported here were referred to R. euryotis: although
they differ sharply from R. e. euryotis in size and in the lesser degree of development of the
antenarial structures they are closely approached in the former respect by R. e. aruensis and
in the latter by this subspecies and to some extent by R. e. burius. Specimens of R. e.
praestans and R. e. timidus are nearer to R. e. euryotis in these features. As Bergmans &
Rozendaal (1982) pointed out, in comparison with R. euryotis the teeth of tatar are not
small, the zygomatic width is not narrowed and the sagittal crest is not low. Consequently,
they considered the skull of tatar to display a tendency towards R. euryotis although on its

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external features they considered tatar to be nearest to R. arcuatus. However, Bergmans &
Rozendaal (1982) appear to have made no direct comparison with any representative of R.
arcuatus, and of R. euryotis only with examples of the nominate subspecies from Amboina.
Comparison with all but angustifolius among R. arcuatus, and with all of the described
form of R. euryotis leads me to the conclusion that tatar is best considered a subspecies of
this latter, a finding that confirms the allocation of similar specimens from Sulawesi to R.
euryotis by Tate & Archbold (1939).

In view of the relatively small size of R. e. tatar it is of some interest to remark that
Koopman (1982) has reported three specimens of R. euryotis from Kiriwina island, off
eastern Papua New Guinea that are of similar size (Table 4), and one that is little larger
(approaching R. e. timidus) from New Britain, but Smith & Hood (198 1 ) have also recorded a
much larger example from the same island. Koopman (1982) was inclined to doubt the
validity of some at least of the subspecies ofR. euryotis then recognised.

HIPPOSIDERIDAE

Hipposideros bicolor bicolor (Temminck, 1834)

Rhinolophus bicolor Temminck, 1834: 19, pi. 1, fig. 3; 1835: 18 (further description). Lectotype

designated and type locality restricted to Anjer coast, northwestern Java by Tate ( 1 94 1 a).
Hipposideros javanicus Sody, 1937« : 215. Babakan, Kroja, Tjilatjap, central Java.

SPECIMEN EXAMINED. W Java: 9 Rijksmuseum van Natuurlijke Historic 29304 Tjilatjap (skin, skull;
coll. 25 October 1929, apparently by H. J. V. Sody).

REMARKS. This specimen appears to be one of those recorded by Sody (1930) as H. galeritus
longicauda (Peters, 1861). However, the noseleaf lacks the lateral supplementary leaflets
characteristic of//, galeritus and its allies and the specimen proves on further examination to
represent H. bicolor bicolor. Dorsally, it is pale brown, the hairs with creamy or whitish bases
that show through the darker tipping: the throat and chest are whitish, the remainder of the
ventral surface brownish buff. The skull is elongate in outline, the rostrum and palate
sharply tapered anteriorly; the narial swellings are only slightly inflated and there is a low
sagittal crest; the zygomatic width is less than the mastoid width, the zygoma robust with low
jugal eminence; the interparietal region is swollen and rather pronounced; the palation is
shallowly V-shaped with wide mesopterygoid fossa; the sphenoidal bridge is wide, partially
concealing elongate lateral apertures and there is a shallow oval sphenoidal depression; the
cochlea are a little wider than their distance apart. The anterior upper premolar (pm2) is very
small, slightly extruded into a recess between the canine and the second upper premolar
(pm4); the posterior ridge of m3 is about one half the length of the anterior ridge; the crown
area of the outer lower incisors is very slightly greater than the crown area of the inner pair;
anterior lower premolar (pm2) threequarters the length and two thirds the height of the
second lower premolar (pm4) and two thirds its crown area.

Measurements: length of forearm 45-8; ear not measurable; length of tail c. 39; length of
tibia c. 21; length of foot (c. u.) c. 8; greatest length of skull 19-0; condylobasal length 17-0;
condylocanine length 16-8; basal length 14-6; palatal length 6-5; width across rostral
swellings 4-8; least interorbital width 2-9; zygomatic width 9-3; width of braincase 8-7;
mastoid width 9-6; c'-c1 (alveoli) 4-1; m3-m3 6-1; c-m3 6-5; m1"3 4-0; length complete
mandible from condyles 1 1 -7; length right ramus from condyle 12-0; c-m3 7-0.

DISCUSSION. This example agrees closely with the account by Tate (194 la) of the lectotype
of H. bicolor (Temminck, 1834) in the Rijksmuseum van Natuurlijke Historic and with the
description of H. javanicus by Sody (1937«), thought by Tate (loc. cit.) and by Hill (1963) to
be a synonym of bicolor. Sody almost concurrently (1937&) recorded javanicus from the
island of Banka: the specimen that he reports is slightly smaller in some respects either than
the holotype of javanicus or the specimen of bicolor from Java reported here. There seems
little doubt \hatjavanicus must be considered a synonym of//, b. bicolor.

INDO-AUSTRALIAN BATS 149

Hipposideros ater aruensis Gray, 1858

HipposiderosaruensisGray, 1858 : 107. Aru Islands.

SPECIMENS EXAMINED. Papua New Guinea: 99 BM(NH) 73.2036-2037 (both yg.) Kairiru Cave, near St
Xavier's Mission, Kairiru I, near Wewak, East Sepik, c. 3° 2 1 ' S, 1 43° 36 ' E (in alcohol; coll. Aberdeen
University Exploration Society Expedition to New Guinea, 1973); d BM(NH) 80.612 S Tunnel, Bulolu
Gorge, Wau, Morobe (in alcohol; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Little material is available from the more eastern part of the range of//, ater and
for the present, following Hill (1963), specimens from New Guinea are referred to H. a.
aruensis. McKean (1972) records specimens from Ambunti, Papua New Guinea. Length of
forearm (BM(NH) 80.6 1 2) 40-9.

Hipposideros maggietaylome erroris Smith & Hill, 198 1

Hipposideros maggietaylorae erroris Smith & Hill, 1981 : 14. Yaguam Sulfur Cave, 5 miles
S and 3 miles W of Madang, Madang Province, Papua New Guinea, 5° 1 7 ' S, 145° 45 ' E.

SPECIMENS EXAMINED. Papua New Guinea: 4 dd (2 yg.), 499 BM(NH) 73.2038-2045 Kairiru
Cave, near St Xavier's Mission, Kairiru I, near Wewak, East Sepik, c. 3° 21' S, 143° 36' E (in
alcohol; coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973);
99 (1 yg.) BM(NH) 75.1863-1865 St Xaviers, Kairiru I (in alcohol, skull of BM(NH) 75.1864
extracted; coll. A. M. Jones, Aberdeen University Exploration Society); d, 9 BM(NH) 78.875-876
About 10 km S of Madang, 5° 1 5 ' S, 145° 45 ' E (in alcohol; coll. P. A. Morris); d1, 9 9 (1 yg.) BM(NH)
80.638-643 War Tunnel, Salamana, Morobe, 7° 03' S, 147° 03' E (in alcohol; coll. B. H. Gaskell,
'Operation Drake').

REMARKS. These specimens are listed by Smith & Hill (198 1) in their account of//, maggie-
taylorae: those obtained by 'Operation Drake' are recorded erroneously as BM(NH)
80.516-522 (the suffixes in fact their collection numbers).

Hipposideros ridleyi Robinson & Kloss, 1911

Hipposideros ridleyi Robinson & Kloss, 1911 : 241 Botanic Gardens, Singapore.

SPECIMEN EXAMINED. Borneo: d BM(NH) 82.160. Sepilok Forest Reserve, Sabah (in alcohol; coll. P.
Zborowski).

REMARKS. This specimen is the first of H. ridleyi to be recorded from Borneo, the species
being otherwise known from the type locality and from Selangor on the Malayan mainland
(Medway, 1978). It may be easily recognised by its very large sub-triangular ears (length from
meatus c. 21-23) broad noseleaf that lacks lateral supplementary leaflets, the expansion of
the internarial septum to form a concave, saucer-like disc between and in front of the nostrils
and by its high posterior leaf, the upper part supported by three septa defining four deep
pockets. Length of forearm in the Bornean example 48-1, in the holotype (BM(NH) 61.329)
47-3 and in a specimen (BM(NH) 75.2000) from a locality between Kuala Kangsar and
Rawang, Selangor, 48-5.

Hipposideros cervinus cervinus (Gould, 1863)

Rhinolophus cervinus Gould, 1863 : pi. 34, letterpress. 'Caves on Albany Island' (label on skin of
holotype). Cape York, Queensland, Australia.

150 J. E. HILL

SPECIMENS EXAMINED. C Sulawesi: 8 dd, 5 99 BM(NH) 82.99-1 1 1 Taronggo, 1° 44' S, 121° 40' E; 99
BM(NH) 82.112-113 R Ranu, 1° 51' S, 121° 30' E (all in alcohol, skulls of BM(NH) 82.100,
82.109-1 10, 82. 1 12 extracted; coll. B. H. Gaskell, 'Operation Drake').

S Sulawesi: 99 BM(NH) 82.1 14-1 15 Lalonggasu Meeto (Tomba Watu Cave), 18 km from Kendari
Central (in alcohol, skull of BM(NH) 82. 1 1 5 extracted; coll. B. H. Gaskell, 'Operation Drake').

Papua New Guinea: dd BM(NH) 75.1866-1867 Wageo I, Schouten Is, East Sepik; 9 BM(NH)
75. 1 868 Kadovar I, Schouten Is; d , 3 9 9 BM(NH) 75. 1 869-1 872 East Sepik (all in alcohol; coll. A. M.
Jones, Aberdeen University Exploration Society); d BM(NH) 76.386 Karkar I, Madang (in alcohol;
coll. H. King); 9 BM(NH) 78.877 About 10 km N of Baku, Gogol Valley, Madang, c 40 m (flat skin); 4
dd, 1 99 BM(NH) 78.878-888 8 km W of Baku, c. 50m; (in alcohol) (all coll. P. A. Morris); 99
BM(NH) 80.607-6 1 1 N Tunnel, Bulolu Gorge, Wau, Morobe, 7° 1 9 ' S, 1 46° 44 ' E; d BM(NH) 80.6 1 2
S Tunnel, Bulolu Gorge; d BM(NH) Lae, Morobe, 6° 49 ' S, 1 47° 03 ' E; 9 9 BM(NH) 80.6 14-617 Buso,
Morobe, 7° 1 7 ' S, 1 47° 08 ' E; 6 dd, 8 9 9 BM(NH) 80.6 1 8-63 1 Siboma, Morobe; 2 dd , 4 9 9 BM(NH)
80.632-637 Coastal cave W of Kui Village, Morobe, 7° 22' S, 147° 1 1' E (all in alcohol; coll. B. H.
Gaskell, 'Operation Drake').

New Hebrides: 2 dd, 2 99 BM(NH) 73.1334-1337 Grotte Montmartre, Port Vila, Efate I, 20 m; 6
dd, 3 99 BM(NH) 73.1338-1346 Mission Montmatre, Port Vila; 4 dd, 2 99 BM(NH) 73.1347-1352
'Pig Cave', Harris Plantation, N coast Efate I, 40 m; c?, 9 BM(NH) 73.1353-1354 'Pig Cave', Narabut
Camp, Efate I; 3 dd, 3 99 BM(NH) 73.1355-1360 Lomboh Cave, Litzlitz, Port Stanley Bay, Malekula
I, 5m; cf, 3 99 BM(NH) 73.1361-1364 Lipelip Cave, Amok, Malekula I, 440m; 2 dd, 9 BM(NH)
73.1365-1367 Aouta Plantation, Aore I; dd BM(NH) 73.1368-1369 New Hebrides (all in alcohol,
obtained by Earl of Cranbrook, Royal Society Expedition to the New Hebrides, 1973).

REMARKS. Specimens from Sulawesi appear to be the first of//, c. cervinus to be recorded as
such from that island, although earlier records of//, galeritus galeritus from Gimpoe or Bada
and Peleng Island by Shamel (1940) and of E. galeritus celebensis from Talassa and
Banti-moerang by Tate ( 1 94 1 a) may refer in fact to cervinus. A recent study at the British
Museum (Natural History) by Jenkins & Hill (1981) has indicated that two species, namely
//. galeritus Cantor, 1846 (including insolens Lyon, 1911 as a subspecies) and H. cervinus
(including labuanensis Tomes, 18590 as a subspecies) occur in Borneo. Hipposideros
celebensis Sody, 1936 based on specimens initially referred by this author (1930) to H.
galeritus galeritus, has from the measurements given by Sody a broader braincase and
shorter toothrow than the Sulawesian specimens here referred to H. cervinus, but Jenkins &
Hill found two paratypes of celebensis to have longer toothrows and narrower zygomatic and
braincase widths than Sody's measurements indicated. The paratypes unquestionably
represent H. cervinus and these authors suggested from the available evidence that only H. c.
cervinus occurs in Sulawesi, specimens from that island approaching the more easterly
nominate subspecies rather than the geographically nearer Bornean subspecies //. c.
labuanensis.

Measurements of specimens from Sulawesi: length of forearm (17) 45-0-49-1; condy-
locanine length (5) 14-3-14-9; width of rostrum (5) 4-9-5-1; least interorbital width (5)
2-5-2-8; zygomatic width (5) 8-8-9-1; width of braincase (5) 7-7-8-0; mastoid width (5)
8-6-8-9; c'-c1 (alveoli) (5) 3-7-3-9; m3-m3 (5) 6-0-6-1; c-m3 (5) 5-8-6-0; length complete
mandible from condyles (5) 10-3-10-8; length right ramus from condyle (5) 10-7-1 1-1; c-m3
(5)6-3-6-4.

Hipposideros diadema pullatus Andersen, 1 905

Hipposideros diadema pullatus Andersen, 1905c: 498. Haveri, Papua New Guinea, 700 m.

SPECIMENS EXAMINED. Papua New Guinea: d BM(NH) 78.889 (flat skin), 9,rf 78.890-891 (in alcohol)
About 10 km S of Madang (coll. P. A. Morris).

Hipposideros dinops pelingensis Shamel, 1940
Hipposideros pelingensis Shamel, 1 940 : 353. Peling ( = Peleng) I. E of Sulawesi.

INDO-AUSTRALIAN BATS 1 5 1

SPECIMENS OBTAINED. S Sulawesi: d, 2 99 BM(NH) 82.116-118 Lalonggasu Meeto (Tomba Watu
Cave), 18 km from Kendari Central (in alcohol, skulls of BM(NH) 82.1 17-1 18 extracted; coll. B. H.
Gaskell, 'Operation Drake').

REMARKS. Shamel (1940) diagnosed pelingensis solely on the basis of its shorter tibia (length
38-2-41-0) when compared with H. dinops Andersen, 1905c from the Solomon Islands. Tate
(1941<2 : 376) recorded pelingensis from Talassa, (Maros), south Sulawesi, listing (p. 391) the
specimens as H. diadema pelingensis although (p. 376) comparing and associating
pelingensis with dinops. Hill (1963) considered pelingensis a subspecies of//, dinops but had
seen no examples of the Sulawesian form.

These specimens agree favourably in size and structure with the nominate subspecies from
the Solomon Islands but the rostrum is very slightly narrower, the upper canines a little
smaller at the base and the toothrows a little shorter. However, in most of these features they
closely resemble the smallest of the available specimens from the Solomons, an example
(BM(NH) 67.21 18) from Malaita Island reported by Hill (197 la) who gave measurements of
H. d. dinops and discussed variation in this subspecies.

External measurements: length of forearm (3) 93-4-96-9; length of tibia (3) 39-9-40-9.
Cranial measurements (d1 BM(NH) 82.117, 9 82.118): greatest length of skull 36-5, 36-0;
condylobasal length 32-4, 32-4; condylocanine length 31-5, 31-4; palatal length 13-2, 13-0;
rostral width 10-2, 9-9; anteorbital width 10-0, 9-0; length of anterorbital foramen 2-8, 2-7;
width of anteorbital foramen 0-7, 0-8; least interorbital width 3-6, 3-9; zygomatic width 20-8,
20-2; width of braincase 13-9, 13-7; mastoid width 16-7, 16-1; d-c1 (alveoli) 9-3, 8-9; m3-m3
12-7, 12-7; c-m3 13-8, 13-4; length c1 4-12, 3-97; width c1 2-83, 2-85; length complete
mandible from condyles 24-7, 24-5; length right ramus from condyle 25-5, 25-4; top of
condyle-tip of angular process 5-8, 6-1; bottom of condyle-tip of coronoid process 7-2, 7-6;
tip of angular process-tip of coronoid process 10-9, Il-7;c-m3 15-6, 14-9.

DISCUSSION. Specimens of//, d. pelingensis are of particular interest in connection with the
very large //. inexpectatus Laurie & Hill, 1954, no material of pelingensis being available in
London when this was described. Direct comparison shows pelingensis to be considerably
smaller, with lower, narrower rostrum, much less developed sagittal and lambdoidal crests,
less expanded zygomata and much smaller teeth. There is especially a great contrast in the
large and heavy mandible of inexpectatus and the relatively lighter and by comparison
apparently delicate mandible of pelingensis. Until recently H. inexpectatus was known in
the literature solely from its holotype (BM(NH) 25.6.5.19) from Poso, north Sulawesi but
Fieler (1981) has reported two further examples, from Gorontalo and Minahassa. They are
specimens that have been for many years in the collections of the Staatliches Museum fur
Tierkunde in Dresden, formerly identified as //. diadema (Geoffroy, 1813).

Aselliscus tricuspidatus novehebridensis Sanborn & Nicholson, 1950

Aselliscus tricuspidatus novehebridensis Sanborn & Nicholson, 1950 : 461. Cave on Segond Channel,
Espiritu Santo I, New Hebrides.

SPECIMENS EXAMINED. New Hebrides: 15 <J<J, 8 99 BM(NH) 73.1370-1392 Aouta, Acre I; 2 <?<J, 2 99
BM(NH) 73.1393-1396 Aouta Plantation, Acre I; <?, 9 BM(NH) 73.1397-1398 Hog Harbour, Espiritu
Santo I, 150 ft; 3 dd, 2 99 BM(NH) 73.1399-1403 Senwar Cave, Tenmial, NW coast of Malekula I,
40 m (BM(NH) 73.1 393, 73. 1 396 skins, skulls, all others in alcohol; obtained Earl of Cranbrook, Royal
Society Expedition to the New Hebrides, 1971).

REMARKS. In size (length of forearm 39-8-43-5) these specimens agree with A. t.
novehebridensis as described by Sanborn & Nicholson (1950) and, indeed, extend the range
of forearm length given by these authors. They are generally larger than specimens from the
Solomon Islands referred to A. t. tricuspidatus (Temminck, 1834) by Hill (1956, 197 la) or
others from islands in the same group reported by Sanborn & Beecher ( 1 947).

152 J. E. HILL

External measurements: length of forearm (20 d-d1) 39-8-42-2 (41-2), (14 99) 40-3-43-5
(42-0). Cranial measurements (rf BM(NH) 73.1396, 9 73.1393): greatest length of skull 15-5,
15-7; condylobasal length 13-6, 14-0; condylocanine length 13-2, 13-6; palatal length 2-6,
2-7; rostral width 4-9, 5-0; least interorbital width 1-7, 1-7; zygomatic width 7-9, 7-8; width of
braincase 6-1,6-0; mastoid width 7-1,7-1; c'-c1 (alveoli) 4-0, 3-9; m3-m3 5-5, 5-5; c-m3 5-7,
5-7; length complete mandible from condyles 9-8, 10-0; length right ramus from condyle
10-0, 10-2; c-m3 6-0, 6-1.

Aselliscus tricuspidatus (Temminck, 1 834)

Rhinolophus tricuspidatus Temminck, 1 834 : 20, pi. 1 , fig. 4. Amboina I, Molucca Is.

SPECIMEN EXAMINED. Papua New Guinea: rf BM(NH) 78.892 (yg.) 8 km E of Baku, Gogol Valley,
Madang, 50 m (in alcohol; coll. P. A. Morris).

REMARKS. No attempt has been made to allocate this young specimen to subspecies: in any
event little comparative material from New Guinea is available in the collection of the
British Museum (Natural History). There is however a substantial representation from the
Moluccan islands of Bum and Ceram and from Batchian Island, the Kei Islands, and the
Solomon Islands. These are smaller (length of forearm (99) 47-2-42-0; condylocanine length
(37) 12-1-13-0; palatal length (43) 2-1-2-5; zygomatic width (41) 6-6-7-3; width of braincase
(44) 5-4-5-9; mastoid width (44) 6-3-6-9; c'-c1 (alveoli) (45) 3-1-3-7; m3-m3 (48) 4-6-5-2;
c-m3 (52) 5-0-5-4) than A. t. novehebridensis and are referred to A. t. tricuspidatus. According
to Koopman (1982), similarly small specimens are found on the east Papuan islands.
Specimens reported from Papua New Guinea by McKean (1972) and from West Irian by
Koopman (1982), however, are larger and tend to bridge the size differences between
tricuspidatus and novehebridensis or to approach the latter.

Coelops robinsoni Bonhote, 1908

Coelops robinsoni Bonhote, 1908 : 4. Foot of Mt. Tahan, Pahang, Malaya.

SPECIMEN EXAMINED. Borneo: rf BM(NH) 79.1398 Deer Cave, Gunung Mulu National Park, Sarawak
(partially mummified, now in alcohol, skull extracted; found S. Proctor, Royal Geographical Society
Expedition to Gunung Mulu, 1977-78).

REMARKS. The small leaf-nosed bat C. robinsoni is known from two specimens from Malaya,
one the holotype (BM(NH) 6.10.4.9) in the British Museum (Natural History), the other
(USNM 175000) from Port Swettenham, in the United States National Museum of Natural
History, and from a specimen from Teratau Island, off southern Thailand, reported by
Chasen (1940). The species was reviewed in some detail by Hill (1972) who pointed out
that two specimens referred to C. robinsoni by Robinson & Kloss (\9\5b) are in fact C.
frithii. This Bornean specimen agrees closely with the holotype of C. robinsoni but has a very
slightly wider skull. The discovery of the species in Borneo perhaps adds some weight to the
suggestion by Hill (1972) that C. hirsuta (Miller, 1910) from Mindoro Island in the
Philippines is closely allied to C. robinsoni, which indeed it may represent. Unfortunately, it
is so far known only from the holotype skin.

Measurements of BM(NH) 79.1398: length of forearm 36-5; greatest length of skull to
canine 14-6; condylocanine length 12-7; rostral width 3-9; least interorbital width 1-9;
zygomatic width — ; width of braincase — ; mastoid width — ; c'-c1 (alveoli) 3-2; m3-m3 5-0;
c-m3 5-0; length complete mandible from condyles 8-6; length right ramus from condyle 8-8;
c-m3 5-5.

INDO-AUSTRALIAN BATS 153

VESPERTILIONIDAE
VESPERTILIONINAE

Myotis muricola (?) browni Taylor, 1 934

Myotis browni Taylor, 1 934 : 288. Near Saub, Cotobato, Mindanao I, Philippine Is.

SPECIMENS EXAMINED. C Sulawesi: rf, 4 99 BM(NH) 82.119-123 R Ranu, 1° 51 ' S, 121° 30' E (in
alcohol, all skulls extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These small examples are referred provisionally to browni, a small representative
of M. muricola similar in size to M. m. caliginosus (Tomes, 1859) from the southern
Himalaya or to M. m. niasensis Lyon, 1916 from Nias Island, off west Sumatra. Further
material from the Philippines and Sulawesi might lead to the recognition of a Sulawesian
subspecies: certainly as represented by these specimens from the Ranu River members of the
Sulawesian population are considerably smaller than others of M. muricola from Thailand,
Malaya, Borneo, Java, the Lesser Sunda Islands, and Amboina. There appears to be no
previous record of M muricola as such from Sulawesi.

Measurements of five specimens, except where indicated: length of forearm: 31-4-32-7;
greatest length of skull 12-3-13-0; condylobasal length 11-8-12-6; condylocanine length
11-2-12-0; least interorbital width 2-8-3-0; zygomatic width (3) 7-8-8-0; width of braincase
5-9-6-0; c'-c1 (alveoli) (5) 3-2; m3-m3 5-1-5-4; i^m3 5-4-5-8; c-m3 4-9-5-3; length complete
mandible from condyles 8-8-9-5; length right ramus from condyle 9-0-9-6; c-m3 5-4-5-8.

DISCUSSION. There is some confusion over the correct name for small Myotis with small feet
(subgenus Selysius) occurring in Indo-Australia, the issue apparently turning on the species
to be found in the northern part of the Indian sub-continent. Numerous names have been
applied to these small bats with forearm 30-36 in length and with pmf reduced but not
intruded or only slightly intruded from the toothrows: the major question is whether or not
these should be associated with the otherwise Palaearctic species M. mystacinus (Kuhl,
1819) or should be considered a distinct species M. muricola (Gray, 1846). Indo-Australian
names involved are:

Vespertilio muricola Hodgson, 1841 (nom. nud.)

Vespertilio muricola Gray, 1 846 Nepal

Vespertilio siligorensis Horsfield, 1855 Nepal
Vespertilio darjelingensis Horsfield, 1855

Vespertilio (Pternopterus) lobipes Peters, 1 867 b Burma

Vespertilio caliginosus Tomes,, 18596 India

Vespertilio nipalensis Dobson, 18716 Nepal

Vespertilio blanfordi Dobson, 18716 Himalayas

Vespertilio moupinensis Milne Edwards, 1 872 Szechuan

Myotis niasensis Lyon, 1916 Nias I

Myotis meinertzhageni Thomas, 1926 Kashmir

(?) Myotis latirostris Kishida, 1 932 Taiwan

Myotis browni Taylor, 1 934 Mindanao I,

Philippine Is.

Myotis herrei Taylor, 1 934 Luzon I,

Philippine Is.
Myotis muricola orii Kuroda, 1 935 Taiwan

Thomas ( 1 9 1 5 a) divided M. muricola from a mystacinus group on account of its allegedly
broader, more solidly built skull, heavier teeth, especially the canines, and the greater
intrusion of the posterior of the two small premolars (pm|). According to this author, M.
muricola included lobipes, while his mystacinus group included caliginosus, blanfordi,

154 J. E. HILL

nipalensis, siligorensis and darjelingensis. Thomas recognised two 'forms' within the
mystacinus group, one with low braincase and canines of normal size to include caliginosus,
with synonyms blanfordi and perhaps nipalensis, the other with a high crown and reduced
canines to include siligorensis and its synonym darjelingensis. Myotis siligorensis is today
recognised as a distinct and valid species.

The entire question was reviewed in some detail by Tate (194 Id) who was unable to make
any certain application of the names that he discussed beyond recognising (p. 543) muricola
( = moupinensis), caliginosus ( = blanfordi) and nipalensis ( = meinertzhageni) as subspecies
of M. mystacinus, with siligorensis a valid species in its own right. Elsewhere (p. 545) Tate
considered herrei and browni probably related to niasensis which he thought to be a
subspecies of M. mystacinus, with the suggestion that lobipes may also represent this species.
Finally, Tate (p. 564) remarked that he did not believe muricola to be the correct subspecific
name for any East Indian representative and suggested that all should be referred to M.
mystacinus: specimens from Sumatra, Java and Borneo he allocated to niasensis with the
proviso that this might be a synonym of lobipes, presumably intending that they should be
called M. mystacinus niasensis.

Chasen (1940) called most Sundanesian specimens M. muricola muricola but employed
niasensis as a valid subspecies of M. muricola for those from Nias Island, while Laurie & Hill
(1954) followed the lead given by Tate (194 Id) in employing M. mystacinus but used
muricola as the subspecific name for specimens from the Lesser Sunda Islands, and, by
implication, from much of Sundanesia. Findley (1972) recognised mystacinus and muricola
groups within Selysius and considered these forms unlikely to be conspecific as once was
thought since Tate (194 Id) had recorded them sympatrically in Nepal, apparently through a
reference to Scully (1887) who reported 'mystacinus' from that country (later described as
siligorensis) although in fact Tate listed muricola as a subspecies of mystacinus. Findley
included meinertzhageni in mystacinus and caliginosus, blanfordi and moupinensis in
muricola, with the suggestion that muricola also includes the various populations of tropical
Asia, having associated browni, latirostris and niasensis (inter alia) with his muricola group.
More recently, Corbet (1978) has excluded all but meinertzhageni among the Indian forms
from any discussion of M mystacinus, considering muricola specifically distinct.

The collections of the British Museum (Natural History) indicate that two distinctively
coloured Myotis (excluding siligorensis} occur in the northern part of the Indian sub-
continent, although cranially they are very similar. One of these is exemplified by a small
series of specimens (BM(NH) 16.7.29. 37^1, 16.7.28.90, part of those reported as muricola
by Wroughton, 1917) from Hasimara Tea Estate, E bank of Toorsa River, c. 8 miles from
Bhutan Hills, Jalpaiguri District, Bhutan Duars, 500-600 ft, c. 26° 50' N, 89° 20' E. These
are brownish dorsally, the hairs black or blackish brown at the base and for most of their
length, tipped with shiny, paler ochraceous brown. Ventrally the specimens are whitish, the
hairs black based, tipped with white or creamy white, the black undercolour showing
through to a limited extent. The collections also include a similar specimen (BM(NH)
9.7.27.3) from Sirguffara, Kashmir, 6000 ft in which the white ventral tipping is a little less
evident, and a further example (BM(NH) 23.9.1.1 2), reported by Lindsay ( 1 926) as muricola,
from Chirot, in the Rattan Valley, Lahul, near the junction of Chandra and Bhaga Rivers,
9800 ft with the dorsal pelage more heavily tipped with burnished ochraceous. A specimen
(BM(NH) 71.13) in alcohol, from Kokernag, Kashmir evidently also has brownish dorsal
pelage and whitish underparts. These specimens clearly represent M. mystacinus and agree
closely with the description of M. mystacinus nipalensis by Dobson (187 \b) to which they
are referred.

Specimens of this type appear to have been the basis of the remark by Allen & Coolidge
(1940) that muricola of the Nepal-Himalaya foothills has the dorsal pelage tipped with
shiny-ochraceous rather than duller brown, with a whitish rather than yellowish ventral
surface. Thus in contrasting such specimens with East Indian examples they apparently
compared their more eastern specimens with M. mystacinus rather than with muricola as
they assumed. Tate (194 Id) quotes these authors in support of the distinctiveness of

INDO-AUSTRALIAN BATS 155

supposed Himalayan muricola: possibly their comment influenced his opinion that muricola
should not be applied as a subspecific name to any East Indian form.

Tate (1941 d) and Corbet (1978) suggested that meinertzhageni might be synonymous
with nipalensis, Ellerman & Morrison-Scott (195 1) in fact listing it as a provisional synonym
of nipalensis. However, the holotype BM(NH) 26.3.1.1 of meinertzhageni is considerably
paler dorsally than the specimens here referred to nipalensis, the pelage heavily tipped with
creamy white, faintly tinged with ochraceous, the ventral surface with pelage that is white
tipped over a black base.

The collections also include further specimens (BM(NH) 23.9.1.13-14, identified as
caliginosus by Lindsay, 1926) from Samayala, in the Kangra Valley, that evidently represent
a smaller, darker form than M. mystacinus, its dorsal pelage darker and blacker, less
profusely tipped with shiny brown, its ventral pelage tipped with ochraceous brown rather
than with whitish, although both dorsally and ventrally the hairs are blackish at the base.
Similar dark-bellied specimens in the collections come from other localities in the Punjab
(BM(NH) 7.11.21.4, 10.1.18.17-18), Sikkim (BM(NH) 91.10.7.57, 15.9.1.21), Kashmir
(BM(NH) 8.7.6.10) and Pakistan (from Murree Hills) (BM(NH) 71.1570-1573) at altitudes
from 6500-7600 ft: the highest at which these are represented is by a specimen (BM(NH)
14.7.10.55) from the Pindar Valley at 10 700 ft. All of these specimens agree closely with
caliginosus or with blanfordi which is evidently a synonym of caliginosus: in turn this last is
similar to but slightly larger than muricola. There seems little doubt that these are
conspecific and that M. muricola caliginosus is the westernmost representative of M.
muricola, overlapping in parts of its range with M. mystacinus nipalensis.

Further east, specimens of muricola from Nepal, in wet preservation, have the character-
istically dark ventral surface of caliginosus, lacking any white tipping. Dry examples from
Thailand, Vietnam, Borneo and Java have a brownish dorsal surface similar to that of
caliginosus, the hairs blackish brown at the base and tipped with ochraceous brown. The
ventral surface is buffy brown to greyish black, the hairs having dark bases with buffy brown
or greyish buff tips, but often with much of the darker undercolour showing through the
paler tipping. Lack of mainland material prevents any proper assessment of possible sub-
specific variation: as Chasen ( 1 940) pointed out the pattern of size variation in Malaya,
Sumatra, Borneo and Java throws doubt on the necessity of recognising more than one
Sundanesian subspecies. For the present, therefore, I refer specimens from this area and its
many islands except Nias to one subspecies, M. muricola muricola. Myotis niasensis Lyon,
1916 is very small and may well justify distinction as M. muricola niasensis, while the
Philippine forms herrei and browni also seem separable on similar grounds, together with
small examples from Sulawesi.

No attempt has been made in this brief survey to discuss named forms occurring outside
the Indo-Australian region, or to examine the relationship of M. muricola to any of the other
Palaearctic forms now recognised. This question is briefly reviewed by Corbet (1978).

Myotis ater (Peters, 1866)

Vespertilio ater Peters, 1 866a : 1 8. Ternate I.

Vespertilio adversus var. amboinensis Peters, 1 8666 : 400. Amboina I, Molucca Is.

(?) Vespertilio australis Dobson, 1 878 : 3 1 7. New South Wales, Australia.

SPECIMENS EXAMINED. C Sulawesi: d BM(NH) 82.124 R Ranu, 1° 51' S, 121° 30' E (in alcohol, skull
extracted; coll. B. H. Gaskell, 'Operation Drake').

Siberut I. Mentawei Is: 2 dd (1 yg.). 899 BM(NH) 78.2932-2936 Headquarters of Sinmuri R, off
Saibi R (in alcohol, skulls of BM(NH) 78.2933-2934 extracted; coll. J. J. Whitten).
REMARKS. Although similar to M. muricola, M. ater is larger, with a heavier, wider rostrum
and more inflated braincase; pm3 is small, intruded from the toothrow while pm, is slightly
intruded, squashed between pm2 and pm4. As Tate (1941 d) correctly pointed out, pm2 is less

156 J. E. HILL

reduced in ater (amboinensis) than in muricola and its allies. This specimen complements
others (BM(NH) 97.1.12.48-51, 7.1.1.504, 7.1.1.507) in the collections in London from
Sulawesi: Shamel (1940) and Tate ( 1 94 1 d) recorded (as amboinensis) specimens from several
Sulawesian localities to which Tate adds Vagian and Peleng Islands, and Papua New Guinea.

Measurements of BM(NH) 82.124, with those of other Sulawesian specimens in
parentheses: length of forearm 36-5 ((6) 36-1-38-4); greatest length of skull 13-9 (14-2, 14-4);
condylobasal length 13-3, (13-5, 13-7); condylocanine length 12-6 (12-8, 13-1); least inter-
orbital width 3-5 (3-4, 3-5); zygomatic width 8-7 ( — ); width of braincase 6-4 (6-9); mastoid
width 7-0 (7-6); c'-c1 (alveoli) 3-6 ((6) 3-8-4-0); m3-m3 5-6 ((5) 5-9-6-2); i^m3 6-5 ((6)
6-6-6-8); c-m3 5-4 ((6) 5-6-5-7); length complete mandible from condyles 10-4 ((3)
10-4-10-7); length right ramus from condyle 10-7 ((6) 20-7-10-9); c-m, 5-8 ((6) 5-9-6-1.

Specimens from Siberut Island in the Mentawei Islands whence M. ater has not before
been recorded agree in structure and size with M. ater from Sulawesi and the Molucca
Islands, except that prn^ are a little less reduced and less sharply intruded from the toothrows.
Measurements of Siberut specimens: length of forearm (4) 34-8-36-2; greatest length of skull
14-1, 14-3; condylobasal length 13-5, 13 -6; condylocanine length 12-9, 12-9; least interorbital
width 3-5, 3-6; zygomatic width 9-3, 9-2; width of braincase 6-6, 6-5; mastoid width 7-5, 7-5;
c'-c1 (alveoli) 3-9, 3-9; m3-m3 6-0, 6-2; i^m3 6-6, 6-6; c-m3 5-6, 5-6; length complete
mandible from condyles 10-4, 10-5; length right ramus from condyle 10-9, 10-9; c-m3 6-0,
5-9.

DISCUSSION. Thomas (19150) considered M. ater to be a larger ally of M. muricola from
Sulawesi and the Moluccan Islands of Amboina, Buru and Ceram. Tate ( 1 94 1 d) retained ater
in a distinct section of the subgenus Selysius (p. 545) although he also indicated (p. 564) that
ater, as amboinensis with which Tate thought it possibly synonymous, might be a subspecies
of M. mystacinus ( = M. muricola in the present sense). Laurie & Hill (1954) listed ater as a
subspecies of M. mystacinus ( = M. muricola}: however, specimens from the Ranu River
show the two forms to occur sympatrically in Sulawesi. Moreover, a small specimen
(BM(NH) 80.1.17.3) from Amboina (length of forearm 34-3, condylobasal length 12-9,
condylocanine length 12-3, c-m3 5-2) with small pm2 and pm^ slightly intruded does not
represent ambionensis of Peters (length of forearm 37-5) but instead is referable to M.
muricola. Although the collections in London contain no example of ater (amboinensis)
from Amboina, specimens from Buru (BM(NH) 10.3.3.28-30, 23.1.1.14) and Ceram
(BM(NH) 10.3.4.32-43, 23.1.2.15) are large (length of forearm (16) 36-8-39-2, condylobasal
length (11) 13-7-14-1, condylocanine length (11) 13-0-13-5, c-m3 (11)5-6-5-9) and represent
this species.

Hill (1962) and Medway (1965, 1977) regarded ater and muricola as conspecific, treating
both as subspecies of M. mystacinus (=M. muricola in the present sense). The former of
these authors provisionally associated nugax Allen & Coolidge, 1940 from northern Borneo
and Culion Island in the Philippines (Sanborn, 1952) with M. mystacinus ( = M. muricola) as
a possible subspecies, while the latter considered it to be the subspecies of M. mystacinus
(=M. muricola) in the highlands of northwestern Borneo, with M. m. muricola in the
lowlands. However, on description and size it appears to represent M. ater and it is possible
that the two may be conspecific as Tate (194 Id) suggested, but the length of the toothrow of
nugax given by Allen & Coolidge is rather large for ater. Tate (194 Id) considered that these
authors apparently had included the incisors rather than measuring from the canine as is
customary and gave for MCZ 36075 (the holotype is MCZ 36076) in the Museum of
Comparative Zoology, Harvard, a value for i^m3 of 6-5 and for c-m3 of 5-5, closely corres-
ponding to M. ater.

Although similar in numerous points of size to M. abbotti Lyon, 1916 from North Pagi
Island in the Mentawei Islands, specimens of M. ater from Siberut quite clearly show no
relation to that form. Comparison with the account by Lyon indicates that the interorbital
region and braincase of abbotti are considerably wider: moreover, Tate (194 Id) considered
abbotti to belong to the subgenus Leuconoe rather than to Selysius, although Lyon originally

INDO-AUSTRALIAN BATS 157

described it as a subspecies of M. muricola. Tate in fact thought it belonged with M. adversus
Horsfield, 1824, possibly as a synonym of M a. carimatae Miller, \9Q6d. Diagnostic features
not mentioned by Lyon can be gleaned from Shamel (1942) who noted that the wing
membrane is attached to the ankle, thus differing from M. ater and from M. horsfieldii in
which it is attached to the metatarsus, that pm3 is much smaller than pm2, its tip only slightly
above the cingulum of the latter tooth, crowded completely from the toothrow with pm2 and
pm4 almost in contact, while pm3 is in the toothrow but crowded slightly to the inside. These
features suggest alliance to M. hasseltii (Temminck, 1840) of which the nominate subspecies
occurs in Java, Sumatra and Malaya, M. h. continentis Shamel, 1942 in Thailand and Burma
and M. h. macellus (Temminck, 1840) in Borneo. Specimens apparently referable to M.
hasseltii have also been obtained in Sri Lanka (Hill, \916a).

The status of Vespertilio australis Dobson, 1878, allegedly from New South Wales,
Australia has remained in doubt for many years. However, McKean (1970) has reproduced a
detailed letter from Dr A. M. Husson describing its holotype, housed in the Rijksmuseum
van Natuurlijke Historic, Leiden. The description, especially of the dentition, and the
measurements that Dr Husson has provided agree closely with M. ater. Measurements given
for the holotype of australis, with comparable values for ater from Sulawesi and the
Moluccas in parentheses: length of forearm (ex Dobson, who gives 1 ' 55 ' ') 39-5 ((23)
36-1-39-2); interorbital width 3-9 ((14) 3-3-3-5); c'-c1 3-7 ((18) 3-6-4-3); i2-m3 6-9 ((18)
6-5-7-1); c-m3 5-8 ((18) 5-4-5-9); i-m3 7-0 ((15) 6-9-7-5), c-m3 6-3 ((18) 5-8-6-3.

Dr Husson rejects any relationship between australis and M. adversus but suggests instead
that on dental features australis might be allied to M. muricola: to this McKean added that
Laurie & Hill (1954) considered this a subspecies of M. mystacinus with the further sub-
species M. mystacinus ater extending to Papua. The available evidence suggests strongly that
australis is at least conspecific with M. ater but the origins of the holotype and only known
specimen are obscure. Although it is on a mount marked 'Sydney N. Hollande' it may have
been mislabelled, collected elsewhere, or may be a ship-assisted vagrant as Allison (1980)
suggested.

Myotis horsfieldii horsfieldii (Temminck, 1 840)

Vespertilio horsfieldii Temminck, 1840 : 226. Java.

Leuconoe lepidus Thomas, 1 9 1 5d : 171. Baram, Sarawak, Borneo.

SPECIMENS EXAMINED. C Sulawesi: cf, 5 99 BM(NH) 82.125-130 Taronggo, 1° 44' S, 121° 40' E (in
alcohol, skulls extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Specimens from Sulawesi are cranially a little larger in some respects than those
from Thailand (M. h. deignani Shamel, 1942) or Malaya, Borneo and Java (M. h. horsfieldii;
the Bornean form lepidus was considered only weakly if at all separable by Hill (1974) and
Medway (1977) has since synonymised it with the nominate subspecies), but the differences
are very slight. Measurements appear in Table 5.

DISCUSSION. Myotis horsfieldii was first recorded as such from Sulawesi by Hill (1974) who
reported a specimen (BM(NH) 73.1804) from Wawondula in the southern part of the island.
Subsequently Jones & Maa (1976) reported an individual from Makassar (5° 07' S, 1 19° 24'
E) as M. horsfieldii with the comment that Jentink (1883) had listed this species from
northern but not southern Sulawesi and that Laurie & Hill (1954) had listed it from Peleng
Island. However, neither Jentink nor Laurie & Hill mention M. horsfieldii but instead report
M. adversus (the record in Laurie & Hill was drawn from Tate (194 Id) who also reported
adversus from Menado in northern Sulawesi) so that the exact identity of the specimen in
the United States National Museum of Natural History reported by Jones & Maa must
remain uncertain.

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Hill (1976#) thought peshwa Thomas, 1915a from India and dryas Andersen, \9Qlb
closely related to M. horsfieldii of which both are here considered provisional subspecies. It
is also possible that M. jeannei Taylor, 1934 from Zamboanga Island, Philippine Islands
may represent M. horsfieldii: there is reasonable agreement in size and the wing membrane
in jeannei is attached to the foot about halfway between the heel and the base of the toe, a
characteristic feature of M. horsfieldii when compared with M. adversus or M. hasseltii in
which it attached at the end of the tibia or at the ankle.

Myotis hasseltii continents Shamel, 1 942

Myotis adversus continentis Shamel, 1942 : 323. Bangkok, Thailand.

SPECIMEN EXAMINED. Burma: cf BM(NH) 78.154 British Embassy Residence compound, Rangoon (in
alcohol, skull extracted; coll. D. W. & G. Walton).

REMARKS. This specimen is the first of M hasseltii to be reported from Burma, although the
species is known to occur farther north in Thailand, at Chiangmai (BM(NH) 9.10.1 1.6-9).
It agrees closely with examples of M. h. continentis from Thailand but is slightly larger in
some respects (Table 6), approaching M. h. hasseltii from the Malay Peninsula and Java,
or the specimens from Sri Lanka discussed by Hill ( 1 91 6a). The second upper premolar (pm3)
of the Burmese specimen is very small, totally intruded from the row, with pm2 and pm4
in contact, while pm3 is small, slightly intruded but nevertheless separates pm2 and pm4.

No examples of M. hasseltii were obtained by 'Operation Drake' in Sulawesi, whence its
reported occurrence apparently rests on specimens recorded by Jentink (1887, 1888) from
Gorontalo in the northern part of the island.

Myotis adversus moluccarum (Thomas, 1915)

Leuconoe moluccarum Thomas, \9\5d : 1 70. Ara, Kei Islands.

Anamygdon solomonis Troughton, 1929 : 89. Rovianna I, New Georgia group, Solomon Is.

SPECIMENS EXAMINED. C Sulawesi: 2 rfrf, 9 BM(NH) 82.131-133 R Ranu, 1° 51' S, 121° 30' E (in
alcohol, skulls extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. Tate (\94\d) referred specimens of M. adversus from Sulawesi, Peleng Island and
West Irian to M. a. moluccarum, with which these agree closely. They are rather smaller
than M. a. adversus (Horsfield, 1 824) from Java: the Bornean and Sumatran subspecies M. a.
carimatae Miller, \9Q6d is very similar to moluccarum but only limited material is available
for comparison.

Myotis adversus orientis subsp. nov.

HOLOTYPE. New Hebrides: 9 BM(NH) 73.1412 Eastern cave at cliff base behind Aouta
Plantation, Aore I. Collected 24 August 1971 by the Earl of Cranbrook, presented by the
Royal Society Expedition to the New Hebrides, 1971. Original number 104-02. Skin and
skull.

OTHER MATERIAL. New Hebrides: 3 cfrf, 7 99 BM(NH) 73.1404-141 1, 73.1413 (in alcohol,
skulls extracted), 73.1414 (skin, skull). All from type locality (obtained Earl of Cranbrook,
Royal Society Expedition to the New Hebrides, 1971).

DIAGNOSIS. A member of the subgenus Leuconoe, generally slightly larger (Table 7) in most
respects than the geographically adjacent M. a. moluccarum from the Solomon Islands, New

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Length of forearm (1
Greatest length of skull
Condylobasal length
Condylocanine length
Least interorbital width
Zygomatic width
Width of braincase
Mastoid width
c'-c1 (alveoli)
m3-m3
c-m3
Length of complete mandible
from condyles i
Length of right ramus i
c-m, (

162 J. E. HILL

Guinea, the Moluccas and Sulawesi, approaching M. a. adversus from Java in size; differing
from these, from M. a. carimatae from Borneo and from M. a. macropus (Gould, 1855) from
Australia in a slightly greater degree of frontal elevation of the rostrum, the frontal profile
less concave, in wider, more substantial interorbital region, and in greater inflation of the
braincase, which is more globose, especially anteriorly, slightly higher, and more elevated
posteriorly.

DESCRIPTION. Externally very like M. a. adversus or M. a. moluccarum with relatively large,
rounded ears reaching almost to tip of muzzle when laid forward; tragus in length a little less
than half length of ear, slender, pointed, its anterior margin straight, its posterior margin
gently convex; wing inserted at ankle. Pelage dense and woolly, dorsally blackish brown, the
hairs dark at the base and for most of their length, tipped lightly with greyish or buffy white,
the tipping forming no more than a slight overlay, the ventral surface greyish white, the hairs
blackish at the base, heavily and densely tipped with grey white, posteriorly with little or no
black at the base. Dorsal pelage darker and blacker than brownish examples of moluccarum
from Sulawesi and the Molucca Islands but these are old specimens that have been
preserved for many years: a similarly old specimen from Choiseul Island in the Solomon
Islands approaches examples from the New Hebrides in dorsal colour.

Skull strongly constructed, with broad, high rostrum, its upper surface flattened
posteriorly with a broad, shallow median longitudinal depression; frontal profile only
slightly depressed; interorbital region broad, relatively massive, least interorbital width
28-30% of condylobasal length, 54-57% of width of braincase; 26-28% of condylobasal
length, 49-54% of width of braincase in adversus, carimatae and moluccarum; braincase
inflated, globose, anteriorly relatively full, elevated posteriorly; palate short and wide, with
strong post-palatal spine, supported by thin lateral laminae; second upper premolar (pm3)
small, its crown area about one third crown area of anterior tooth (pm2), almost in row,
slightly intruded, or intruded from row to lie in a recess between pm2 and last upper
premolar (pm4); second lower premolar (pm3) in crown area about one third crown area of
anterior tooth (pm2), almost in row but compressed between pm2 and last lower premolar
(pm4), or sometimes slightly intruded.

Measurements of M. a. orientis appear in Table 7.

ETYMOLOGY. An eastern representative of M. adversus.

DISCUSSION. Excluding very large species such as M. macrotarsus (Waterhouse, 1845) from
the Philippine Islands and northern Borneo and M. stalkeri Thomas, 19100 from the Kei
Islands leaves in Indo-Australia a group of large-footed bats of the subgenus Leuconoe with
forearms ranging in length from 36-44 and with pm2 and pm3 variably reduced and variably
situated in the toothrows. Their current classification has been established by views and
opinions scattered through several publications. These include Tate (194 Id), Medway
(1965, 1977), Findlay (1972), Hill &Thonglongya (1972) and Hill, (1972, 1974, 1976a).The
arrangement adopted here has been developed from these and may be summarized:

1 . Wing inserted on metatarsus M- horsfieldii
Wing inserted at end of tibia or at ankle

2. Fur short, velvety; post-palatal extension short, its median

spicule lacking thin bony supporting laminae; pm3
minute, intruded, pm2 and pm4 in contact or nearly so,
pm3 small, only partially in row or completely intruded M. hasseltii

Fur, dense, woolly; post-palatal extension long, its median
spicule supported by thin bony laminae; pm3, not greatly
reduced, pm3 not usually much intruded, pm, usually in
row or only slightly intruded M. adversus

Myotis horsfieldii deignani Shamel, 1 942 Thailand, Hainan I,

Hong Kong (Findley,
1972).

INDO-AUSTRALIAN BATS

Myotis horsfieldii horsfieldii (Temminck, 1 840)
(incl. lepidus Thomas, \9\5d)

Myotis horsfieldii (!)peshwa Thomas,

19 15a (see Hill, 1976a)
Myotis horsfieldii (?) dryas Andersen,

19076 (see Hill, 1976a)
Myotis horsfieldii (l)jeannei

Taylor, 1934
Myotis hasseltii continentis Shamel, 1942

Myotis hasseltii hasseltii

Temminck, 1840
Myotis hasseltii macellus

Temminck, 1840
Myotis hasseltii (?) abbotti Lyon, 1916

Myotis adversus adversus Horsfield,

1824

Myotis adversus carimatae Miller, 1 906 d
Myotis adversus moluccarum Thomas,

\9\5d

Myotis adversus orientis Hill, 1983
Myotis adversus macropus Gould, 1855

163

Malaya, Borneo,
Java, Bali, Sulawesi.

India.

South Andaman I.

Zamboanga I,

Philippine Is.
Burma, Thailand,

Kampuchea,

(?) Sri Lanka.
Malaya, Rhio Archipelago,

Sumatra, Java.
Borneo.

North Pagi I, Mentawei

Is, offW Sumatra.
Java, (?) Malaya.

Borneo, Karimata I.
Sulawesi, Molucca

Is, New Guinea,

Solomon Is.
New Hebrides.
Australia.

Pipistrellus javanicus (Gray, 1838)

Scotophilus javanicus Gray, 1838 : 498 (renaming of Vespertilion javanais F. Cuvier, 1832). Java.

SPECIMENS EXAMINED. C Sulawesi: 9 BM(NH) 82.134 R Ranu, 1° 51' S, 121° 30' E (in alcohol, skull
extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. This specimen from Sulawesi agrees in most respects with examples of P.
javanicus from Java and from the island of Madura. It differs from these only in the slightly
greater concavity of its rostral profile, in having the frontal part of the braincase a little more
inflated and in a slightly higher posterior upper canine cusp. Measurements: length of fore-
arm 32-1; greatest length of skull 13-6; condylobasal length 12-7; condylocanine length 12-3;
least interorbital width 3-5; zygomatic width 8-6; width of braincase 6-7; mastoid width 7-3;
c'-c1 (alveoli) 4-3; m3-m3 6-2; c-m3 4-9; length complete mandible from condyles 9-4; length
right ramus from condyle 9-8; c-m3 5-2.

DISCUSSION. Several species of Pipistrellus have been reported or described from Sulawesi.
These include P. javanicus (Gray, 1838) by Dobson (1878) (as abramus, a subspecies of P.
javanicus according to Laurie & Hill, 1954), Meyer (1899) (as abramus) and Shamel (1940)
(also as abramus); P. imbricatus (Horsfield, 1824) by Tate (19426); possibly P. papuanus
(Peters & Doria, 1881) by Laurie & Hill (1954); P. petersi (Meyer, 1899), described from
northern Sulawesi and reported again by Shamel (1940) and Tate (19426), and P.
minahassae (Meyer, 1899), also described from northern Sulawesi and again reported
provisionally by Tate (19426). Of specimens recorded as abramus by Dobson (1878), one
(BM(NH) 72.4.11.4) from northern Sulawesi is neither javanicus nor is it papuanus, as
Laurie & Hill (1954) thought it might be, so far as can be judged from the damaged skull. A
second specimen, a skin only, cannot be found. The cranial features of the holotypes of
petersi and minahassae have never been recorded, but Tate (19426) provides information
and measurements drawn from the referred specimens that he reported.

164 J. E. HILL

There are few records of specimens referable to P.javanicus from the area east of Java and
Sulawesi. I have examined specimens supposedly of this species in the collection of the
Zoologisches Museum (Museum fur Naturkunde), Berlin from Bali (ZMB 90465,
90468-90471) and from Sumba Island (ZMB 92152-92157), the latter those reported as P.
tralatitius ( = P. javanicus) by Pohle (1950). None in fact represent P. javanicus but instead
are examples of Myotis muricola, recorded from Sumba by Forcart (1952). Schwarz (1914)
reported P. tralatitius from Timor Island, but these records from Lelogama and Bonleo may
refer to P. tennis as Goodwin (1979) suggested. Other specimens from Timor recorded as P.
tralatitius by Pohle (1950) prove (vide infra) to be examples of P. tenuis. Dobson (1878)
recorded two specimens of Pipistrellus abramus from 'Australia', having earlier (1876)
suggested that the species extended to that country. These are BM(NH) 47.7.21.12-13, in
alcohol: their skulls have now been extracted. The two specimens agree closely with P.
javanicus which evidently they represent: a further specimen, BM(NH) 80.1.17.4 from the
Aru Islands may also represent this species but is rather smaller.

Pipistrellus tenuis papuanus (Peters & Doria, 1881)

Vesperugo papuanus Peters & Doria, 1881: 696. Salawatti, West Irian, NW New Guinea.
Vesperugo papuanus orientalis Meyer, 1899 : 14. Bongu, Astrolabe Bay, Papua New Guinea.

SPECIMENS EXAMINED. Papua New Guinea: cf, 99 BM(NH) 73.2046-2048 Kairiru I, near Wewak, East
Sepik (in alcohol; coll. Aberdeen University Exploration Society Expedition to New Guinea, 1973); 99
BM(NH) 80.644-645 Wau Gulf, Morobe, 7° 19' S, 140° 44' E; 99 BM(NH) 80.646-651 Lae, Morobe,
c. 6° 49 ' S, 1 47° 03 ' E; rf. 9 9 BM(NH) 82. 148-1 52 Cave near Avi, Mt. Hagen (all in alcohol; coll. B. H.
Gaskell, 'Operation Drake'); 9 BM(NH) 82.157 Lake Murray Wildlife Station, Boboa, Western
Province (in alcohol; coll. I. M. Redmond).

REMARKS. These specimens with forearm lengths ranging from 29-3-31-6 are in good agree-
ment with P. t. papuanus as represented in the collections of the British Museum (Natural
History).

DISCUSSION. Laurie & Hill (1954) recognised papuanus as a distinct species in New Guinea,
together with P. angulatus of which they listed three subspecies, P. a. angulatus Peters, 1880
from the Bismarck Archipelago, P. a. collinus Thomas, 1920 from mainland New Guinea
and P. a. ponceleti Troughton, 1936 from the Solomon Islands, in general following the leads
provided by Tate (1942&). Lidicker & Ziegler (1968) adopted the classification offered by
Laurie & Hill but tentatively raised ponceleti to specific rank, and added P. imbricatus
(Horsfield, 1824) to the New Guinea list by referring specimens from Balim River, West
Irian and Emirau Island in the Bismarck Archipelago to that species. More recently,
Koopman (1973) has reviewed the Indo- Australian pipistrelles associated with P. tenuis,
with particular reference to the forms known from New Guinea. His study led him to unite
papuanus, angulatus, collinus and ponceleti (and also nitidus Tomes, 1858 from Borneo
(referred to P. tenuis as a subspecies by Medway, 1965), subulidens Miller, 1900 from
Sirhassen Island, sewelanus Oey, 1960 from Lombok Island and murrayi Andrews, 1900
from Christmas Island) into a single species, P. tenuis (Temminck, 1 840). He mentioned but
did not discuss the records of P. imbricatus by Lidicker & Ziegler.

Although the collections of the British Museum (Natural History) include a substantial
number of specimens that can be referred to P. t. papuanus, there are few that can be
confidently identified with collinus, angulatus or ponceleti. However, there is some evidence
to confirm Koopman's (1973) division of Papuan examples into a small, relatively lowland
form (papuanus) and a larger highland form (collinus), separated also by the dimensions of
the palate, which is relatively short and narrow in the lowland form but longer and wider in
the montane form. Additionally, montane specimens have a concave rostral profile in con-
trast to the flatter, less depressed profile usual in lowland examples. No montane specimens

INDO-AUSTRALIAN BATS 165

are available from West Irian, but lowland examples from the western part of the island
confirm the indication by Koopman (Figs. 1 , 3) that here the lowland form has a longer skull
nearer in size to that of the montane form, with a correspondingly slightly longer and slightly
wider palate. However, Koopman examined only three variables and it is also clear from his
diagrams (Figs. 1-3) that high montane (especially above 1800 m) specimens differ consider-
ably from those from lower altitudes in the dimensions of the palate: those from altitudes of
1000-1800 m in Papua ally themselves in palatal size with the high montane examples (Fig.
2) but in West Irian (Fig. 3) with more truly lowland specimens. Similarly, in length of skull
(Fig. 1) specimens from the intermediate zone in Papua are like those from higher altitudes
but in West Irian are approached in size by lowland examples. The variation in eastern New
Guinea and its associated islands has been analysed in greater detail by Koopman (1982).
Pipistrellus tennis papuanus also occurs on the Bismarck Archipelago whence Smith &
Hood (198 1) have recorded specimens that they refer to this form, together with other larger
examples that apparently represent angulatus, the two perhaps being separated also by
roosting preference. Moreover, specimens examined by Koopman (1973) from New Britain
with forearms 28-30 in length may also represent P. t. papuanus.

It is unlikely that specimens from Emirau Island, Bismarck Archipelago and from the
Balim River, West Irian represent P. imbricatus as Lidicker & Ziegler (1968) averred.
According to these authors the specimen from Emirau (from their illustration, fig. 4) has a
concave rostral profile and has an anterior upper premolar (pm2) of cross-sectional area
about equal to two thirds of the cross-sectional area of the outer upper incisor (i3). However,
in a series of P. imbricatus from Java in the British Museum (Natural History) the rostral
profile is straight or nearly so, a point reinforced by Koopman (1973) who recorded that
imbricatus has a flat, not concave forehead, and pm2 is very small, sometimes minute, not
more and usually less than one third the cross-sectional area of i3. Tate (1942) identified the
Balim River specimens (AMNH 109979-109980) as 'P. imbricatus collinus', and Smith &
Hood (1981) suggest that for the present the Emirau specimen seems best referred to
angulatus.

Pohle (1950) recorded P. tralatitius( = P.javanicus) from Timor on the basis of specimens
in the Zoologische Museum (Museum fur Naturkunde), Berlin, collected by G. Stein in
1932. I have examined the skulls of these specimens (ZMB 92107-92111), all from
Tjamplong and they prove not to represent P. javanicus but to be closely similar to if not
identical with P. tenuis papuanus. Goodwin (1979) also reports P. tennis from Timor, and
gives measurements (taken by Dr K. F. Koopman) of further specimens from Tjamplong
collected by Stein and now in the Museum Zoologicum Bogojiense, Bogor. Two specimens
reported from Timor by Schwarz (1914) as P. tralatitius may also be P. tenuis, as Goodwin
suggests. Koopman (1973) allocates Timorese specimens to P. t. sewelanns, along with those
from Sulawesi.

Collett (1897) recorded a single specimen from Roebuck Bay, near Broome, northwestern
Australia as P. tennis. This specimen, collected by Dahl and now in the Oslo Museum, was
sent to the British Museum (Natural History) in 1966 for examination. It proved to agree
closely with P. t. papuanus but to be slightly larger cranially, with a narrower, blade-like
post-palatal spine and slightly less prominent basioccipital pits, and to have a slightly less
prominent posterior upper canine cusp. More recently, in 1976, two specimens (AMNH
216135-216136) from the collections of the American Museum of Natural History were
examined for Dr Karl F. Koopman. These agree basically with P. t. papuanus but have the
narial emargination a little more rounded and less acute posteriorly, the palate slightly more
domed and less prominent posterior cusps on the upper canines. Although for the present I
would refer these specimens to P. t. papuanus it seems probable that more material might
well show the Australian population to be subspecifically separable.

The limited material in the British Museum (Natural History) supports the association of
papuanus with tenuis as Koopman (1973) proposed, although of the latter only two so-called
'cotypes' (BM(NH) 7.1.1 .407^08) are available for the study, Tate ( 1 9426) having selected as
lectotype a specimen in the Rijksmuseum van Natuurlijke Historic, Leiden. On the basis of

166 J. E. HILL

this sample I am inclined to agree that tenuis and papuanus are conspecific: both are
characterised by small size, short premaxillae, the narial emargination quite sharply
narrowed posteriorly, the anterior upper premolar (pm2) moderate in size, situated in the
internal angle between c1 and the posterior upper premolar (pm4), which almost touch or are
in contact. Of the two forms, P. t. tenuis is apparently slightly the larger: it is similar in size to
a small series of murrayi from Christmas Island which also clearly represents P. tenuis and
may even be synonymous with the nominate subspecies. A specimen (AMNH 107827) in
the American Museum of Natural History from Bratan, Bali examined in 1970 for Dr Karl
F. Koopman also represents this complex. Then labelled 'Pipistrellus (coromandra)
imbricatus subsp' which very evidently it is not, it is apparently the example identified by
Tate ( 1 9426) as P. tenuis. It is similar to the 'cotypes' of tenuis or to murrayi but the skull is a
trifle longer and wider and the toothrows slightly longer, in these respects approaching
papuanus. Koopman (1973) refers specimens from Bali and Java to P. t. nitidus. The cranial
measurements of specimens discussed appear in Table 8.

Pipistrellus vordermanni (Jentink, 1890)

Vesperugo vordermanni Jentink, 1890 : 152. Billiton I.

SPECIMEN EXAMINED. Borneo: 9 BM(NH) 82.547 Samunsan Wildlife Sanctuary, Sarawak, (in alcohol,
skull extracted; coll. N. A. MacKenzie).

REMARKS. This is the first of P. vordermanni to be recorded from Borneo; no direct
comparison has been possible but the specimen agrees closely with the original account by
Jentink. Among Bornean pipistrelles the species may be readily recognised by its large ears,
slightly hatchet-shaped tragus with anteriorly directed point, whitish wings, short skull, full,
rounded braincase and short rostrum, small supraorbital tubercles, deep zygomata with
distinct postorbital process, deep basial pits, minute anterior upper premolar (pm2) which
is completely intruded into a recess between the canine and the posterior upper premolar
(pm4), the two latter teeth in contact, and much reduced anterior lower premolar (pm2).

DISCUSSION. Chasen (1940) synonymised vordermanni and curtatus Miller, 1911 from
Engano Island, off the west coast of Sumatra, with the Sumatran macrotis (Temminck, 1840)
which he considered a subspecies of P. imbricatus (Horsfield, 1 824) from Java. However,
all differ sharply from imbricatus in a decidedly shorter rostrum and consequently shorter
palate, a point emphasised by Miller in his description of curtatus. Tate (19426) treated
macrotis, vordermanni and curtatus as distinct species, associating them with P. cadornae
Thomas, 1916 from Burma in the savii group of Pipistrellus. Hill (1962) followed Tate in
considering macrotis and vordermanni distinct species but removed them, with cadornae,
from any close relationship with P. savii (Bonaparte, 1837). As Tate (19426) pointed out,
the four forms clearly constitute an assemblage characterised by the cranial and dental
characters noted above. The larger size of cadornae sets it rather apart from macrotis, vorder-
manni and curtatus which may well prove to be conspecific.

Measurements of the Bornean example of vordermanni with (in parentheses) those of an
example (BM(NH) 23.1.2.12) of macrotis from Sebang, Sumatra: length of forearm 30-5
(31-5); length of ear 15-1 (14-7); greatest length of skull 114 (12-2); condylobasal length 11-1
(11-8); condylocanine length 10-8 (11-5); width across supraorbital tubercles 4-6 (4-7);
zygomatic width 8-2 (84); least interorbital width 3-5 (3-5); width of braincase 64 (6-5);
mastoid width 7-0 (7-3); c'-c1 (alveoli) 4-0 (3-9); m3-m3 5-1 (5-3); c-m3 3-8 (4-0); length of
complete mandible from condyles — (7-8); length of right ramus from condyle — (8-3); c-m3
4-2 (44). Measurements of type material of macrotis, vordermanni and cadornae are given
by Tate ( 1 9426) and of the holotype and additional specimens of cadornae by Hill ( 1 962).

INDO-AUSTRALIAN BATS

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Pipistrellus societatis Hill, 1972

Pipistrellus societatis Hill, 1972 : 34. Base Camp, Gunong Benom, Pahang, Malaya, 800 ft, 3° 51' N,
102° 1 1 ' E.

SPECIMEN EXAMINED. Malaya: 9 BM(NH) 81.1802 Sungai Tekam Forest Reserve, Pahang (in alcohol,
skull extracted; coll. A. O. Johns).

REMARKS. This is the first example of P. societatis to be reported since its description, the
holotype (BM(NH) 67.1605) being then the only known example. This newly obtained
specimen is in good general agreement with the holotype but is slightly larger in some
respects and has, however, a small, low sagittal crest that is absent from the original example.
The narial emargination is very slightly narrowed posteriorly and the last upper molar (m3) is
slightly less reduced than in the holotype, with a more evident but small metacone and short
third commissure, although as in the holotype the tooth is flattened antero-posteriorly and
more platelet-like when compared with the closely related species P. circumdatus. As in the
holotype the hypoconid and entoconid of the last lower molar (m3) are also slightly reduced
in comparison with those of the related species.

Measurements: length of forearm 39-4; greatest length of skull 15-0; condylobasal length
14-3; condylocanine length 14-1; length orbit-gnathion 3-9; palatal length, excluding post-
palatal spine 6-7; length palatal bridge, excluding post-palatal spine 5-1; width across ante-
orbital foramina 3-2; lachrymal width 6-8; width across supraorbital tubercles 6-0; zygomatic
width 10-5; least interorbital width 4-3; width of braincase 7-5; mastoid width 8-4; c'-c1
(alveoli) 4-8; m3-m3 7-0; c-m3 5-5; length of complete mandible from condyles 10-6; length of
right ramus from condyle 1 1-0; c-m3 5-9.

DISCUSSION. Hill (1972, 1974) reported an example of the closely related P. circumdatus
from Eraser's Hill, Pahang. Certain features of this species and of P. societatis tend towards
the Australian genus Chalinolobus and more distantly towards the nominal genus
Glauconycteris from Africa. Both species show an approach to Chalinolobus in the develop-
ment of a slightly pendent lobule at the base of the outer margin of the ear, inserted a little
behind and below the angle of the mouth, in an elevated, rounded braincase and the inflation
of the supraorbital region, a shortening of the rostrum and a narrowing of the anterior palatal
emargination, the arrangement of the upper incisors into a straight line and the reduction of
the anterior upper premolar (pm2) almost to the point of obsolescence.

Philetor brachypterus rohui Thomas, 1 902

Philetor rohui Thomas, 1902 : 220. Albert Edward Range, eastern Papua New Guinea, 6000 ft.

SPECIMENS EXAMINED. Papua New Guinea: rf, 99BM(NH) 80.652-656, <f BM(NH) 80.694 Buso,
Morobe, 6° 4 1 ' S, 1 47° 1 3 ' E (in alcohol; coll. B. H. Gaskell; 'Operation Drake').

Philetor brachypterus verecundus (Chasen, 1940)

Eptesicus verecundus Chasen, 1940 : 53. Mt. Kledang, Perak, Malaya, 2646 ft.

SPECIMEN EXAMINED. Borneo: rf BM(NH) 78. 1 542 Gunung Mulu, Sarawak, 2600 ft (in alcohol; coll. D.
R. Wells, Royal Geographical Society Expedition to Gunung Mulu, 1977-78).

REMARKS. This species has been recorded previously from two Bornean localities, Pangkalan
Lobang, at Niah, Sarawak, and Poring National Park, at Ranau in Sabah (Lim, Chai &
Muul, 1974; Medway, 1977). Two of these specimens were examined at the British Museum
(Natural History) in 1975 and found to approach P. b. rohui from New Guinea in the width

INDO-AUSTRALIAN BATS 169

of the narial emargination rather than P. b. verecundus from Malaya. However, a specimen
(BM(NH) 73.597) from Ulu Gombok, Selangor, Malaya has a similarly narrower
emargination and as Medway (1977) suggests, this feature seems of little value in establishing
subspecies. Among specimens in the British Museum (Natural History) the forearm of those
of P. b. verecundus is a little longer on the whole than in P. b. rohui: in twenty Malayan
examples the length of the forearm ranges from 33-0-37-6, in thirty one from New Guinea
from 3 1 -3-35-6. In size the Bornean example (length of forearm 35-4) agrees with those from
Malaya (Lim, Chai & Muul, 1974, give forearm lengths of 33-36 for eight further speci-
mens from Borneo). However, Kock (1981) records specimens from New Britain in the
Bismarck Archipelago and from New Guinea that are similar in forearm length to those from
Malaya. For the present Bornean specimens are referred to P. b. verecundus, but subspecific
validities are very provisional in P. brachypterus, the features of its Sumatran representative,
(nominally P. b. brachypterus) remaining largely unknown (Hill, 19666 19716). The
collections in London include an unreported series (BM(NH) 74.415-434) of P. b.
verecundus from Paloh Forest Reserve, Paloh, Kluang, Johore, Malaya.

Hesperoptenus (Milithronycteris) gaskelli sp. nov.

HOLOTYPE. Central Sulawesi: 9 (yg. ad.) BM(NH) 82.135. River Ranu, 1° 51 ' S, 121° 30' E.
Collected 19 April 1980 by B. H. Gaskell, 'Operation Drake'. In alcohol, skull extracted.

DIAGNOSIS. Characteristically a member of the subgenus Milithronycteris similar in many
respects to H. tickelli or H. tomesi, differing principally from H. blanfordi in much larger
size, naked internarial region, the dorsal surface of the forearm lacking any dense covering of
hair, the absence of a fleshy pad at the base of the thumb, inflated braincase, narrow post-
orbital region and relatively less reduced outer upper incisors (i3~3) and last upper molars
(p|). Differing from H. tickelli and H. tomesi in smaller size and smaller teeth (a comparison
of the young adult holotype with two examples of H. tickelli of similar or even slightly
younger age shows the dentition of the latter to be substantially larger) the canine and post-
canine teeth little more than one half the basal area of their counterparts in H. tickelli, and
for the most part rather less than one half the basal area of those of//, tomesi, inner upper
incisors (i2~2) similarly smaller, the lower incisors about one third smaller than those of//.
tickelli and about half the size of those of//, tomesi. Differing further from H. tickelli in its
rich, dark, blackish rather than yellowish b^own or straw brown overall coloration, in much
shortened second phalanx of the third digit, in the development of small supraorbital
tubercles and slightly more vertical orbital margin, deeper, more sharply sculpted
basioccipital pits, a narrower cingulum on the outer upper incisor (i3), more flattened last
upper molar (m3) and a slightly greater degree of reduction of the last lower molar (m3); from
H. tomesi in a lesser degree of intrusion of the outer upper incisor (i3), in a lack of thickening
of the first and second lower incisors (i]_2) and a less thickened outer lower incisor (i3), its
cusp pattern remaining evident.

DESCRIPTION. Muzzle low, wide, laterally swollen, sparsely pilose, with medianly a very
shallow longitudinal groove behind the nostrils; narial openings nearly circular, moderately
separated; a few sparse, short hairs fringing the upper lip beneath nostrils, the
hairs laterally longer and rather denser; internarial region nearly naked; upper part of muzzle
anteriorly with a sparse cover of long hairs, a short, broad median longitudinal band of
similar hairs extending from a point about halfway between nostrils and eyes to widen
posteriorly between ears and join fur of crown; lateral parts of upper lip above canines and
premolars swollen, sparsely furred, the hairs usually stiff and bristle-like; sides of muzzle
around and beneath eyes nearly to angle of mouth with a thin cover of longer hairs; a small
wart above the anterior canthus of the eye, bearing a long vibrissa and a few shorter hairs;
lower lip with a small, undivided median pad; anteriorly ventral surface of head only
sparsely pilose with a small almost totally naked area beneath the symphysis menti,

170 J. E. HILL

preceding a low, almost imperceptible elevation that bears a few long hairs; a few longer
vibrissae at sides of muzzle.

Ear thick and fleshy, subqua.drangular, its anterior margin with well developed posteriorly
directed lobe at its base, proximally strongly convex, distally almost straight, the tip broadly
rounded, posterior margin of ear strongly convex, with prominent fleshy antitragal lobe, the
lobe thickened along its outer margin, inserted quadrately behind and a little below the angle
of the mouth; tragus rather fleshy, slightly less in length than one half the length of the ear,
more or less hatchet shaped, its anterior margin straight to a narrow, more or less anteriorly
directed tip, its upper margin horizontal, the posterior margin strongly convex to well
developed subtriangular basal lobe; outer surface of conch quite densely haired in its basal
half, distal parts of inner surface and lower part of outer face of tragus sparsely haired.

Thumb relatively long; third metacarpal slightly the longest of metacarpals two to five,
fourth very little snorter, fifth rather more so, the second the shortest; second phalange of
third digit shorter than first phalange; upper surface of thumb with a thin cover of short
hairs; upper surface of forearm naked except for a few hairs near the elbow; toes sparsely
haired dorsally; upper surface of uropatagium sparsely haired for its proximal half; base of
tail similarly furred both above and below, its tip protruding beyond uropatagium; calcar
long, extending across about two thirds of the uropatagial margin; a small linear post-
calcarial lobe, lacking any definite supporting cartilaginous spur.

Colour (from specimen in alcohol) both above and below a rich dark blackish chocolate
brown, the hairs more brownish at the base but otherwise darker and lustrous.

Skull evidently of medium size with broad, uninflated braincase; postorbital region
moderately constricted; supraorbital region abruptly expanded, with well developed
superior anteorbital tubercles; anterior orbital margin sharply defined, in frontal aspect
nearly vertical; anteorbital foramen large, closed by a narrow bar of bone; maxillary margin
of orbit heavy, flange-like, in ventral view forming a wide ledge alongside the toothrow;
zygomata slender; narial emargination more or less U-shaped, slightly tapered posteriorly,
with broadly rounded apex; post-palatal extension short, with broad median spine;
basioccipital pits prominent, sharply incised into basioccipital.

Inner upper incisor (i2) caniniform, massive, unicuspid, at its base equal in area almost to
one half the basal area of the canine, with well developed cingulum, anterior face of tooth
rounded, its postero-internal face deeply excavated, its postero-external face shallowly so,
the depressions separated by a narrow ridge; outer upper incisor (i3) more or less ovate, with
wide basin-like cingulum and central three-faced cusp, basal area of tooth a little less than
one half basal area of i2, the cusp extending a little above the cingula of i2 or of c1, its postero-
internal face excavated; tooth intruded from row to lie almost directly behind i2, its anterior
edge in contact with that tooth and on a line joining the anterior edges of c1"1, its posterior
edge on a line joining the centres of these teeth, its labial part intruded between i2 and c1, with
its postero-lateral edge in contact with the antero-lateral face of c1; i2 and c1 otherwise
separated by a narrow interspace of about one third the basal diameter of i2. Upper canine
with strong cingulum; upper premolar (pm4) slightly compressed in toothrow; last upper
molar (m3) in crown area about one half crown area of m1, with metacone and three
commissures, the third short. Lower incisors (i] _3) with tricuspid cutting edge, i2 overlapping
i, by almost one half its width, i3 thickened, with low, small posterior supporting cusp,
similarly imbricating i2; anterior lower premolar (pm2) in crown area about one third crown
area of posterior lower premolar (pm4), slightly compressed in row; last lower molar (m3)
little reduced, entoconid and hypoconid low, the tooth slightly narrowed posteriorly.

MEASUREMENTS. External measurements of holotype with (in parentheses) those of two males (BM(NH)
13.2.10.27-28) of H. tickelli from Sri Lanka, of similar or younger age: length of forearm 40-1 (47-9,
48-2); of thumb with claw 6-0 (7-9, 8-3); of IP 35-3 (38-9, 41-3); of IIP 36-3 (41-4, 43-3); of III1 16-2
(17-5, 19-0); of IIP 13-3 (16-1, 16-6); of IVm 36-0 (40-8, 41-6); of IV 13-4 (14-6, 16-1); of IV2
7-8 (9-6, 10-7); of Vm 35-1 (39-8, 41-1); of V 8-4 (10-1, 10-4); V2 4-9 (7-8, 7-7); length of tibia
16-5 (19-1, 20-4).

INDO-AUSTRALI AN BATS 1 7 1

Cranial measurements of holotype with (in parentheses) those of BM(NH) 13.2.10.27:
greatest length of skull 15-6 (17-1); condylobasal length 15-2 ( — ); condylocanine length 13-1
( — ); length orbit-gnathion 4-0 (4*7); palatal length 5-9 (7-0); width across anteorbital
foramina 5-6 (6-3); width across front of orbits 7-8 (8-4); width across supraorbital swellings
6-6 (6-9); least interorbital width 4-7 (4-7); zygomatic width 11-4 ( — ); width of braincase 8-3
(9-2); height of braincase 5-8 (6-1); mastoid width 9-0 (9-8); c'-c1 (alveoli) 5-4 (6-1); m3-m3 7-8
(8-6); c-m3 6-0 (7-0); length complete mandible from condyles 11-5 (13-0); length right ramus
from condyle 12-0(1 3-6); c-m3 6-5 (7-8).

DISCUSSION. The genus Hesperoptenus was reviewed in detail by Hill (19766): it has not been
recorded previously from Sulawesi. The nominate subgenus with the sole species H. doriae
Peters, 1869 is known from Malaya and Borneo. In the subgenus Milithronycteris the small,
very distinctive H. blanfordi (Dobson, 18776) has been reported from southern Burma,
Thailand and Malaya, H. tickelli (Blyth, 1851) from India and Sri Lanka east to the
Andaman Islands and Thailand, and H. tomesi Thomas, 1905 from Malaya and Borneo. In
coloration, shortening of the second phalanx of the third digit, the presence of well-
developed supraorbital tubercles, more nearly vertical anteorbital margin, a narrow
cingulum on the outer upper incisor (i3) and in the slightly greater degree of reduction of the
last upper molar (m3) this new species approaches H. tomesi, although clearly very much
smaller. However, its lower incisors (ii_3) are relatively less massive than in tomesi, i3 in
particular although large and with a low posterior cingulum cusp nevertheless retaining a
more obvious tricuspid edge. In these respects the new species tends towards H. tickelli.

ETYMOLOGY. It gives me much pleasure to associate this new species with Mr B. H. Gaskell,
the collector whose work with 'Operation Drake' has produced an excellent sample of the
Sulawesian bat fauna.

MINIOPTERINAE

Miniopterus australis australis Tomes, 1858

Miniopterus australis Tomes, 1 858 : 125. Loyalty Islands, South Pacific.

SPECIMENS EXAMINED. New Hebrides: 2d d, 2 99 BM(NH) 73.1415-1418 Grotte Montmartre, near Port
Vila, Efate I, 20 m; 8 dd, 7 99 BM(NH) 73.1419-1433 Mission Montmartre, near Port Vila, 40 m; 5
dd, 3 99 2undet. BM(NH) 73.1434-1443 Montmartre, near Port Vila; d BM(NH) 73. 1444 'Pig Cave',
Narabut Camp, Efate I; 7 dd, 4 99 BM(NH) 73.1445-1455 'Pig Cave', Harris Plantation, N coast of
Efate I; 2 dd, 9 BM(NH) 73.1456-1458 Ijiptho Cave, NW Aneityum I, sea level; 9, 2 dd BM(NH)
73.1459-1461 Paul's Bungalow site, near 'Bethel', Tanna I, sea level; 3 99 BM(NH) 73.1462-1464
Siwi, Tanna I; 3rfcTBM(NH) 73.1465-1467 Senwar Cave, Tenmial, NW coast of Malekula I, 40 m; dd
BM(NH) 73.1468-1473 Amok, Malekula I, 485 m; 2 dd, 2 99 BM(NH) 73.1474-1477 Abounatori
Village, Malo I; 2 dd, 4 99 BM(NH) 73.1478-1483 Aouta Plantation, Acre I; 5 cTcf, 4 99 BM(NH)
73.1484-1492 Hog Harbour, Espiritu Santo I; 9 BM(NH) 73.1493 Nokowoula, Espiritu Santo I; 2 dd
BM(NH) 73.1494-1495 Rock shelter 'Wonatsuri', near Nokowoula, Espiritu Santo I, 3550 ft; 3 dd, 3
99 BM(NH) 73.1496-1501 New Hebrides (all in alcohol except BM(NH) 73.1462, skin and skull;
obtained Earl ofCranbrook, Royal Society Expedition to New Hebrides, 1971).

TYPE MATERIAL AND TYPE LOCALITY. Tomes (1858) based Miniopterus australis on a total of
six specimens, all apparently now in the collections of the British Museum (Natural History).
The species was said originally to be from Australia but Dobson (1876) indicated that the
specimens that he considered to be the 'types' came in fact from the Loyalty Islands, their
bottle in the British Museum being so labelled. Later (1878) he listed these specimens as the
'types', again recording them from the Loyalty Islands. Revilliod (1914) drew attention to
this contradiction and noted that Oldfield Thomas had confirmed that this latter locality was
the provenance indicated on the labels of these specimens and in the relevant accession
register of the British Museum (Natural History). As a result Revilliod accepted the Loyalty
Islands as the correct type locality. Peterson (19810) in an abstract of a paper entitled The

172 J. E. HILL

systematic status of Miniopterus australis and related forms stated that the 'holotype' of M.
australis supposedly from the Loyalty Islands proves to differ from population samples from
these islands and from New Caledonia but is indistinguishable from the holotype of M.
tibialis (Tomes, 1858) from Amboina in the Molucca Islands. He proposed therefore to fix
the type locality ofM. australis as eastern Australia.

Five of Tomes' original specimens were accessed by John Edward Gray, then responsible
for vertebrates at the British Museum, within the registration group BM(NH) 54.5.19.1-22, a
mixed collection listed without locality or ascribed variously either to the Loyalty Islands or
to New Zealand, presented by Sir George Grey. Of these, BM(NH) 54.5.19.1 is recorded
simply as 'Vespertilio' without locality or further data, BM(NH) 54.5.19.4-7 being listed
again as 'Vespertilio 'but with the additional data 'In spirit' and 'Loyalty Islands'. The speci-
men BM(NH) 54.5.19.1 is a skin with skull in situ, and has been labelled 'Australia' by J. E.
Gray. Tomes' sixth specimen appears to be BM(NH) 7.1.1.555 (i.e. the Tomes collection,
accessed in 1907), a skin with rostral fragment and mandible attached and according to
Tomes (1858) specimen 'a' of Gray's (1843&) listing of Trilatitus blepotis from Timor, a
specimen received from the Leyden Museum. Its label bears the notes 'F™. B.M. Apr. 1857'
and 'Timor, from the Ley. Mus.', apparently in Tomes' own hand. Evidently it was given to
Tomes from the British Museum collections and was returned fifty years later when the
whole of his collection was received into the National Museum.

No holotype was designated by Tomes for M. australis but Dobson (1876, 1878) clearly
considered only the specimens in alcohol BM(NH) 54.5.19.4-7 to be the 'types'. Oldfield
Thomas evidently considered this question in some detail, no doubt in response to the point
raised by Revilliod, and has clearly re-bottled and in two cases labelled these specimens with
appropriate annotations. The specimen tabulated as No. 1 by Tomes he considered to be
BM(NH) 54.5.19.4, having labelled it 'No. 1 of Tomes'. It bears another label on which the
words Miniopterus', 'Loyalty Islands' and 'Sir G. Grey' in a second hand can just be
discerned: a further label is no longer decipherable. This specimen is now in an individual
bottle. Its external label has been written by Thomas and is annotated 'No. 1 of Tome's
australis (No. 2 made lectotype)'. He has also written the label attached to its skull which is
annotated 'No. 1 of Tome's M. australis. Not typical. No. 2. made lectotype'. The specimen
is in fact an example of M. robustior Revilliod, 1914, a point made evident by Revilliod in
the original account. This taxon is known so far only from the Loyalty Islands.

The remaining three alcoholic specimens are now together in one bottle with a modern
external label giving their locality as Loyalty Islands. One, a male, has been labelled by
Thomas as BM(NH) 54.5.19.5, the label annotated 'No. 2 of Tomes'. It is the only one of
the three to have its skull extracted: the skull label has been written by Thomas and is
annotated 'Lectotype'. australis. No. 2 of Tomes'. The other two specimens, a female and
male, have no labels but are apparently 3 and 4 of Tomes. This author noted that the four
specimens so far discussed were preserved in spirit in the British Museum: his 5 'a specimen
in skin in the same collection, the whole of them having been collected in Australia by Sir G.
Grey, K.C.B., and presented to the National Collection' is clearly BM(NH) 54.5.19.1. Its
label locality, not confirmed by the accessions register, of 'Australia' may well have led
Tomes to assume that all of the specimens came from that continent, whence Grey collected
numerous mammals. Tomes specimen 6 is apparently BM(NH) 7.1.1.555.

Tate (\94\e) quotes BM(NH) 54.5.19.5 as the type specimen of M. australis, evidently
accepting the unpublished designation made by Thomas. There seems no reason either to
query the validity of this selection or to doubt that the specimen is from the Loyalty Islands:
it is recorded with this provenance in the accessions register and the bottle that contained it
and the other three alcoholic examples registered at the same time was apparently labelled as
such (Dobson, 1876, 1878) by J. E. Gray: moreover, one of these proves subsequently to
belong to a taxon so far known only from these islands. Since no formal selection appears to
have been made, BM(NH) 54.5.19.5 is here designated the lectotype of Miniopterus australis
Tomes, 1858 with the Loyalty Islands as its type locality, in agreement with the archival
evidence.

INDO-AUSTRALIAN BATS 173

Maeda (1982) has provided an erroneous corollary. This author stated (p. 4) that BM(NH)
14.5.23.2 is one of Tomes' series of specimens and was re-described by Revilliod (1914) as
Miniopterus australis robustior, remarking later (p. 7) that it is the holotype of this taxon.
Both of these statements are incorrect. BM(NH) 14.5.23.2 was presented to the British
Museum (Natural History) by F. Sarasin and J. Roux in 19 14 and is one of the syntypes ofM
robustior. it has no connection at all with Tomes. Revilliod (1914) makes it clear that
robustior was based on nine examples from Lifou, Loyalty Islands and that Tomes No. 1
(now BM(NH) 54.5.19.4) must be referred to it, an allocation confirmed according to
Revilliod by a cranial comparison made by Oldfield Thomas.

REMARKS. The ears of M. australis are rounded and lack any definite point; the tragus is
more or less parallel-sided for most of its length, its upper part curving forward to a greater or
lesser extent, the upper part of the anterior margin concave and the posterior part convex.
The anterior projection of the upper part of the tragus is more marked in specimens from the
Loyalty Islands, New Caledonia and the New Hebrides than is usual in those from more
westerly localities: in the eastern specimens the tragus has a nearly horizontal upper margin,
often slightly dentate, points to which Revilliod (1914) first drew attention.

The braincase is small, slightly globose but not markedly inflated, rather low, with low
lambdoidal and sagittal crests, the occipital region not inflated, in profile lower than the
highest point of the braincase in the frontal region; the rostrum is low, relatively wide and
long, with a moderate median longitudinal sulcus; palate long, domed; there are shallow
basioccipital pits, not extending anteriorly to any great extent. The upper canines are strong,
with massive base and stout shaft; anterior upper premolar (pm2) large, triangular in section,
slightly compressed in row; lingual margins of m1 and m2 about equal, m1 with well
developed hypocone, in m2 the hypocone evident but slightly less developed; mandibular
dentition with no especial peculiarities; lower premolars (pm2_4) slightly compressed.

Externally M. australis, if paululus Hollister, 1913 from the Philippines and shortridgei
Laurie & Hill, 1957 from Java are included, is the smallest of Australasian Miniopterus with
the length of the forearm ranging from 34-4^41-5 and of the tibia from 13-5-17-0; the tail is
also relatively short, generally about 40 in length but varying from 35^5, only rarely
achieving these extremes. Size differences among specimens in the collections of the British
Museum (Natural History) are summarised in Table 9.

DISCUSSION. Although until recently (Maeda, 1982) thought to extend to Thailand (Lekagul
& McNeely, 1977) specimens from that country in the British Museum (Natural History)
represent M. pusillus Dobson, 1876 (then considered a subspecies of M. australis), as do
others from the Nicobar Islands. It appears therefore that M. australis as understood here is
distributed from the Philippine Islands, Borneo and Java east to Australia, New Caledonia
and the Loyalty Islands.

Specimens from the islands of Malaita, Guadalcanal and San Christobal in the Solomon
Islands have slightly longer forearms and are sometimes larger cranially than are those from
the Loyalty Islands, New Caledonia and the New Hebrides. There is also a slight difference
in tragal structure as already mentioned. These points in part have led Maeda (1982) to
describe the Malaita and San Christobal examples as a new species, M. solomonensis.
Specific rank seems scarcely justified for these specimens which are approached in several
respects by those from some of the surrounding populations but certainly a distinct sub-
species may be recognised. It is also likely that specimens from Queensland, New Guinea,
the Aru Islands, Batchian Island and the Moluccan Islands of Amboina and Ceram should be
separated as M. a. tibialis (Tomes, 1858) (considered a synonym of M a. australis by Laurie
& Hill, 1954) but sample sizes are small in some cases and may be misleading in the
determination of subspecific limits. In Borneo M. a. witkampi Sody, 1930 differs little from
these in contrast to the markedly smaller forms M. a. paululus Hollister, 1913 from the
Philippine Islands and M. a. shortridgei Laurie & Hill, 1957 from Java. Sanborn (1952)
synonymised paululus with australis with which he suggested that tibialis might also be
synonymous. On the other hand, Peterson (198 la) has implied that paululus and shortridgei

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INDO-AUSTRALIAN BATS 175

may be distinct from australis, while Maeda (1982) has raised paululus to specific rank and
considered shortridgei its synonym. With these bats from the Philippines and Java, however,
he associated small specimens from Rennell Island in the Solomon Islands to produce a
somewhat anomalous distribution pattern for his proposed M. paululus. Koopman (1982)
also recognised paululus as a distinct species in the islands off eastern Papua, the New
Hebrides and Timor, although strongly tempted to treat australis and paululus as
conspecific. The largest of specimens from Rennell Island and the New Hebrides, however,
approach or overlap (Table 9) the smallest of australis from the Loyalty Islands and New
Caledonia. Specimens from Timor are similar to shortridgei from Java but are sometimes
slightly larger, as Goodwin (1979) pointed out.

Miniopterus australis tibialis (Tomes, 1858)

Vespertilio tibialis Tomes, 1858 : 126. Amboina I, Molucca Is.

SPECIMENS EXAMINED. C Sulawesi: cf BM(NH) 82.136 Taronggo, 1° 44' S, 121° 40' E; 9 BM(NH)
82. 137 R Ranu, 1 ° 5 1 ' S, 1 2 1° 30 ' E (in alcohol, skull of BM(NH) 82. 1 36 lacking, of 82. 137 extracted;
coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These specimens are the first of M australis to be reported from Sulawesi, where
on distributional grounds the species might be expected to occur. For the present they are
referred to M. a. tibialis which is considered a probably distinct subspecies: they are similar
in size (Table 9) to those from the Molucca Islands, New Guinea, the Aru Islands and
Queensland discussed above.

Miniopterus pusillus pusillus Dobson, 1876

Miniopterus pusillusDobson, 1876 : 162. Nicobar Islands.

SPECIMENS EXAMINED. Thailand: d, 9 BM(NH) 78.2381-2382 Ban Huai Luang Cave, Chae Horn,
Lampang (skins, skulls); 9 BM(NH) 79.1422 Tham Hai Luang, Chag Hok, Lampang (in alcohol, skull
extracted) (all presented by the Thai National Reference Collection, Bangkok).

Sumatra: d JJW 15 Lian Penerukan, near Bohorok, Langkat (in alcohol, skull extracted; coll. J. J.
Whitten).

W Java: 9 BM(NH) 61.1781 Gua Tjibureum, Tjibodas, 2000 m; 9 BM(NH) 6 1 . 1 79 1 Klapanunggal,
Tjileungsi (skins, skulls; coll. Earl of Cranbrook & P. Jauffret).

TYPE MATERIAL AND TYPE LOCALITY. Tate (1941e) gave the type locality of Miniopterus
pusillus Dobson, 1876 as 'Madras, India' as did Ellerman & Morrison-Scott (1951) who also
said 'But Wroughton gave Nicobar Islands as type locality'. Wroughton (1918) gave the type
locality as 'Nicobars' and cited a specimen in the Indian Museum (No. 185) collected by
Stoliczka as the lectotype.

The original description by Dobson provides no precise type locality but lists measure-
ments of a specimen from the Nicobar Islands and of one from Madras. It includes a reference
to an earlier paper (1871c) by Dobson in which he discussed specimens from the Nicobar
Islands (collected by Stoliczka) that he referred to M. australis with some reservation. In his
Catalogue of the Chiroptera in the collections of the British Museum (1878) Dobson listed
the specimen from Madras and also others reputedly from the Philippine Islands under
Miniopterus schreibersii var. a (i.e. pusillus) but gave measurements only for a Nicobarese
specimen, taken evidently from his original work, and included the Andamans and Nicobars
in the distribution of this form. Significantly, he did not indicate that the Madras specimen
was the type. Clearly Dobson intended pusillus to apply to the Nicobarese specimens to

176 J. E. HILL

which (1871c) he first drew attention, so that Wroughton's (1918) designation of a lectotype
and type locality is justified. The choice of Madras as type locality and by implication of
BM(NH) 52.8.12.8 from there as lectotype appears to rest only on its citation as the first
specimen listed in Dobson's Catalogue ( 1 878).

REMARKS. Miniopterus pusillus is similar in many respects to M. australis but may be
distinguished from this species by its generally longer forearm that ranges from 39-3-45-5,
longer tibia ranging from 16-5-22-0 and longer tail, its length varying from 43-5-52-0; its
braincase is more globose, more elevated and higher, rising abruptly from the rostrum and is
more elevated posteriorly, the height at the occiput nearly equal or equal to its height at the
frontal; the rostrum is relatively short and is weak anteriorly when compared with that of M
australis, posteriorly wider and a little deeper; the upper canines are long and slender; as in
M. australis m1 has a well developed hypocone but the hypocone of m2 is small or absent, the
lingual shelf of this tooth more or less parallel-sided with no postero-internal projection.

The species is sympatric (chiefly as M. p. macrocneme} with M. australis over the eastern
part of its range: although the two can generally be distinguished by the slightly greater
external size of pusillus in locations where they occur together, such comparisons of speci-
mens of each from different, widely separated localities can result in overlap. Dimensions of
specimens in the collections of the British Museum (Natural History) appear in Table 10.

Besides specimens already noted, the collections in London include examples of M. p.
pusillus from the Nicobar Islands (BM(NH) 90.6.20.2, 6.12.1.32-33) and from the island of
Koh Lak, off the southeastern coast of Thailand (BM(NH) 17.2.6.4-7). Maeda (1982)
referred BM(NH) 67.1810 from Bukit Cheras, near Panching, Kuantan, Pahang, Malaya to
pusillus, but comparison suggests that more properly it represents M. medius with which it
was first identified by Hill (1972). The specimens reputedly from the Philippine Islands
listed by Dobson (1878) with, in parentheses, the locality 'Erumango' are BM(NH) A-C, not
having been accessed formally or registered. They were obtained from Mr Hugh Cuming, a
well known collector of Philippine mammals during the latter part of the last century. I have
been unable to locate 'Erumango' in the Philippine Islands but there is an island called
Eromango in the New Hebrides: Hoffman (1887) suggested that Dobson's designation was
an error for New Hebrides. In cranial dimensions (Table 10) these specimens agree more
closely with examples of M. p. macrocneme from New Guinea, the Solomon Islands and
the New Hebrides rather than with more western specimens of this subspecies or with M.
p. pusillus and also have the longer tibia characteristic of macrocneme rather than the shorter
tibia of pusillus. Maeda (1982) also thought that they should be referred to macrocneme.
A further specimen, BM(NH) 74.5.27.6, labelled 'Philippine Islands' and purchased from
a Mr Salmin may represent M. pusillus, tentatively recorded from Baguio, Benguet Island
by Taylor (1934).

The specimen reported here from Sumatra appears to be the first of M. pusillus to be
recorded from that island, and the examples BM(NH) 6 1 . 1 78 1 from Gua Tjibureum, Tjibo-
das and BM(NH) 61-1791 from Klapanunngal seem also to be the first of the species to have
been recorded from Java. The Javan specimens were initially reported by Maeda (1982), who
also referred others from Amboina, Ceram and the Kei Islands to pusillus, but these seem
on comparison to be nearer to macrocneme, like those from New Guinea. On the other hand,
specimens reported from Timor by Goodwin (1979) as M. pusillus have longer forearms like
macrocneme but skulls that are similar in length to those of the western subspecies.

DISCUSSION. Recently accessed specimens from Thailand have made possible a wider
assessment of M. pusillus than was hitherto feasible in London, although the available
material remains inadequate for any detailed analysis of subspecific variation. In particular
the newly acquired material has demonstrated that the limited number of Siamese specimens
hitherto in the collections should not be referred to M. australis as once they were, but
instead to M. pusillus which itself is specifically distinct.

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Miniopterus pusillus macrocneme Revilliod, 1914

Miniopterus macrocneme Revilliod, 1914 : 360. New Caledonia and Loyalty Islands.

SPECIMENS EXAMINED. C Sulawesi: rf, 9 BM(NH)83.138-139 R Ranu, 1° 51 ' S, 121° 40' E (in alcohol,
skulls extracted; coll. B. H. Gaskell, 'Operation Drake').

New Hebrides: 5 rfrf, 5 99, 1 undetermined BM(NH) 73.1502-1512 'Pig Cave', Harris Plantation, N
coast Efate I, 40 m; rf, 2 9 9 BM(NH) 73. 1 5 1 3-1 5 1 5 'Pig Cave', Narabut Camp, Efate I, 800 ft; 2 drf, 2
99 BM(NH) 73.1516-1519 'Pig Cave', Efate I; 2 drf, 2 99 BM(NH) 73.1520-1523 Paul's bungalow
site, near 'Bethel', Tanna I, sea level; 2 rfrf BM(NH) 73. 1 524-1 525 Siwi, Tanna I; 2 dcf, 2 9 9 BM(NH)
73.1526-1529 Arvat, E coast Erromanga I, sea level; rf, 9 BM(NH) 73.1530-1531 Womban Wovoula
Cave, Abouanatori village, Malo I, 50 m; rf, 9 BM(NH) 73.1532-1533 Abouanatori village; d BM(NH)
73. 1 534 Aouta, Acre I; <f, 9 BM(NH) 73. 1 535-1 536 Limboh, near Litzlitz, Port Stanley Bay, Malekula
I; 99 BM(NH) 73.1537-1541 Lomboh Cave, near Litzlitz, 5 m; 3 dd, 2 99 BM(NH) 73.1542-1546
Lipelip Cave, Amok, Malekula I, 440 m (all in alcohol; obtained Earl of Cranbrook, Royal Society
Expedition to New Hebrides, 1971).

REMARKS. The first record of M. pusillus from Sulawesi was provided by Hill (1974) who
reported specimens (as M. medius macrocneme) from Wawondula in the southern part of the
island. For the present Sulawesian specimens are referred to M. p. macrocneme but the total
of material available is quite inadequate for the proper evaluation of subspecies or their
boundaries. However, these specimens and those obtained earlier have the long tibiae
(Table 10) usual in this more eastern subspecies.

Miniopterus pusillus macrocneme occurs with M. australis throughout the range of the
subspecies but may be distinguished by the characters noted above under M. p. pusillus.
According to Revilliod (19 14) macrocneme may be differentiated from australis by the shape
of the tragus, which in the latter he stated projects anteriorly in its distal part, with a nearly
horizontal, irregularly dentate upper margin while in macrocneme the tragus is less strongly
projected forward and there is no well marked dentate upper margin. The anterior projection
of the upper part of the tragus is more pronounced in specimens of australis from New
Caledonia and the Loyalty Islands (whence Revilliod drew his observations) and the New
Hebrides than in those from more westerly locations and in such examples the tragus has
a nearly horizontal upper margin, sometimes slightly dentate, both features that disappear
from more westerly populations. Neither has therefore any extensive diagnostic value.

DISCUSSION. Hill (197 la, 1974) considered macrocneme (an eastern form extending
apparently from Sulawesi east to the New Hebrides, New Caledonia and the Loyalty Islands)
conspecific with medius Thomas & Wroughton, 1909 from Java but Peterson (198 1/?) has
suggested that macrocneme represents the pusillus species group and that medius belongs to
the fuscus group of species. Newly acquired specimens have established medius in southern
Thailand in near sympatry with pusillus, while others whose skulls have become available
recently have shown it to occur sympatrically with pusillus in Java; moreover the additional
material has suggested that medius must be considered specifically distinct, and thai pusillus
and macrocneme are most probably conspecific.

Maeda (1982) seems uncertain in his group allocation of macrocneme but finally places it
as a species in the fuscus group along with M. fuscus and M. medius. However, the
discriminatory ratios given by Maeda for macrocneme overlap considerably into those that
he gives for the pusillus group, and direct comparison of specimens reveals close similarities
of structure. For this reason I agree with Peterson (198 1/?) that macrocneme and pusillus
should be associated, and incline further to the view, used here, that they are conspecific.

Miniopterus medius Thomas & Wroughton, 1 909

Miniopterus medius Thomas & Wroughton, 1909 : 382, Kalipoetjang, Tji-Tandoei R, Java.

SPECIMENS EXAMINED. Thailand: d BM(NH) 78.1424 Ban Kok Chang, Nakhon Sri Thamrat (in
alcohol, skull extracted; presented by Thai National Reference Collection).

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180 J. E. HILL

Malaya: rf, 9 BM(NH) 69.1250-1251 Base of Gunong Ledang ( = G. Ophir), Johore, 2° 21' N, 102°
39' E; d1 BM(NH) 75.1296 Sungei Kelambang Camp, Ulu Setiu, Besut, Trengganu, c. 5° 25' N, 102°
42 ' E (all in alcohol; obtained by the Earl of Cranbrook).

Java: 3 dd , 4 9 9 BM(NH) 61.1 766-1 768 (in alcohol, skulls of BM(NH) 61.1 766, 6 1 . 1 768 extracted),
BM(NH) 61.1782-1783 (skins, skulls), BM(NH) 61.1784-1785 (in alcohol, skulls extracted)
Klapanunngal, Tjileungsi, 280 m (coll. Earl of Cranbrook & P. Jauffret).

Papua New Guinea: 9 BM(NH) 80.657 Mt Kainoi, Wau, 7° 19' S, 146° 44' E (in alcohol, skull
extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. There appear to be no definitive published records of M. medius from the area
east of Java. Baker & Bird (1936) recorded medius from Espiritu Santo Island in the New
Hebrides but two of their specimens still available (BM(NH) 36.4.8.1-2) in fact prove to be
M. pusillus macrocneme. Similarly, the specimens from Papua New Guinea reported as
medius by McKean (1972) seem likely to represent macrocneme (Koopman, 1982 thinks
MH 984 from Putei perhaps referable to M. schreibersii): certainly examples (BM(NH)
50.1802-1803) from Tomba, in the Hagen Range, Papua New Guinea, first reported by
Laurie (1952) as M. schreibersii blepotis but later thought to represent medius in fact belong
to macrocneme while another (BM(NH) 39.1403) from Nadimo Valley, The Gebroeders,
Weyland Range, West Irian although previously identified as medius is also macrocneme.
These specimens were among those leading Hill ( 1 97 1 a) to consider macrocneme conspecific
with medius. The specimen reported here from Mount Kainoi provides evidence of near
sympatry between medius and M. pusillus macrocneme in eastern New Guinea: as pointed
out above, medius also occurs near or with M. p. pusillus in Thailand and Java.

Measurements of M. medius in the collections of the British Museum (Natural History)
appear in Table 1 1 . The largest of M pusillus equal or overlap the smallest of M medius in
some respects but the two can be distinguished by the generally larger skull and more
massive teeth of the latter. Equally, the largest of medius approach or overlap the smallest of
M. schreibersii in some dimensions but medius has a narrower skull and braincase: in any
event, there appears to be no overlap when specimens from the same general area are
compared. Although the example of medius recorded here from New Guinea has a forearm
that is considerably longer than the forearms of medius from Java or from the more western
parts of its range, the specimen agrees very closely with Javan examples both in the structure
and the dimensions of its skull and teeth.

DISCUSSION. Peterson (198 \b} suggested that medius belonged to thefuscus species group, an
opinion confirmed by Maeda (1982). An examination of the holotype (BM(NH) 2.10.7.3) of
M.fuscus Bonhote, 1902 from Okinawa Island, Liu Kiu Islands shows that in all relevant
features it is clearly allied to M. medius and is also dimensionally very similar. Possibly with
or including yayeyamae Kuroda, 1924 from Ishigaki Island, also in the Liu Kiu Islands, it
may be a northern representative of medius: certainly on cranial features and size it has no
connection with M. schreibersii (Kuhl, 1819) as was suggested by Tate (1941?) or by
Ellerman & Morrison-Scott (1951) who, perhaps influenced by Tale's remark that it was
virtually equal to blepotis (Temminck, 1840) and to eschscholtzii (Waterhouse, 1845)
synonymised boihfuscus and yayeyamae with M. schreibersii blepotis.

Miniopterus schreibersii blepotis (Temminck, 1 840)

Vespertilio blepotis Temminck, 1840 : 212, pi. 53, figs 1, 2. Java, Timor and Amboina. Lectotype
tentatively designated and type locality restricted to Java by Tate (194 If): Maeda (1982) has
designated a further lectotype and a series of paralectotypes (including the specimen selected by Tate)
from Java.

SPECIMEN EXAMINED. C Sulawesi: d BM(NH) 82.141 Taronggo, 1°44' S, 121° 40' E (in alcohol, skull
extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. This appears to be the second record of M 5. blepotis from Sulawesi, Tate (1941?)
having recorded a specimen from the southern part of the island. In most dimensions (Table

INDO-AUSTRALIAN BATS 1 g 1

12) it agrees with specimens from western Java but has a longer forearm and longer skull
similar to those of specimens from Amboina Island. These latter were referred to a newly
described species M. oceanensis (see below) by Maeda (1982), who has made an extensive
study of Eurasian and Australasian Miniopterus and has provided a key. Use of this key,
which relies heavily on proportional ratios for discrimination, identifies the Sulawesian
specimen to the fuliginosus, blepotis or schreibersii groups and thence to M. fuliginosus or
M. eschscholtzii if the ratio forearm length/greatest length of skull is employed (and from
there to M. fuliginosus on forearm length and the ratio m3-mVcondylobasal length) but to
M. blepotis if the ratio mastoid width/greatest length of skull is used.

Measurements of BM(NH) 82.141: length of forearm 46-7; greatest length of skull 15-9;
condylobasal length 15-5; condylocanine length 14-5; least interorbital width 3-9; zygomatic
width 8-9; width of braincase 7-9; mastoid width 8-6; c'-c1 (alveoli) 4-5; m3-m3 6-8; c-m3 6-2;
length complete mandible from condyles 11-5; length right ramus from condyle 1 1-5; c-m3
6-8.

DISCUSSION. Until recently the conventional view of M schreibersii (Kuhl, 1819) of eastern
Asia and Indo-Australia included those forms with forearm lengths ranging from 44 or
slightly less to 50 or thereabouts, the condylobasal length of the skull varying from about
14-5 to a maximum of some 16, the exception being M. s. magnater Sanborn, 1931 from
New Guinea which exceeds these upper limits. Peterson (in Goodwin, 1979) considered
magnater to represent a distinct species while Kobayashi et al. (1980) have also raised
magnater to specific rank, recording it with schreibersii from Sabah in northern Borneo.
More recently Maeda (1982) has described the Sabah population initially referred to
magnater as a new species, M. macrodens, associating with it specimens from a wide variety
of localities from Hong Kong to South Australia. It is discussed in more detail below1. Tate
(194 la) suggested that the small forms fuscus Bonhote, 1902 from Okinawa Island, Liu Kiu
Islands and yayeyamae Kuroda, 1 924 from Ishigaki-Mura, Ishigaki Island, also in the Liu
Kiu Islands might be synonymous representatives of M schreibersii, equal to M. s. blepotis
with which both were synonymised by Ellerman & Morrison-Scott (1951). Specimens in the
British Museum (Natural History), including the holotype of fuscus, show clearly that these
do not represent M. schreibersii but instead are near to M. medius of Thailand, Malaya, Java
and New Guinea, as Maeda (1982) suggests, and of which they may be northern representa-
tives. Of these, fuscus is the earliest name.

The collections of the British Museum (Natural History) show that in fact the concept of
M. schreibersii hitherto widely accepted includes two sympatric forms, as the conclusions of
Peterson (1979) and Kobayashi et al. (1980) imply, and as Maeda (1982)1 has established,
in part from the same collections. One, regarded here as M. schreibersii, is smaller, with
a relatively narrower palate (Table 12). In eastern Asia and in the Indo- Australian region
it occurs from Afghanistan, India and Sri Lanka to Japan, the Philippine Islands and east
to the Solomon Islands and Australia. The other, consisting of larger examples, with longer
skull and toothrows and wider palate, but with the forearm on the whole only very slightly
longer (Table 13), is represented in the collections from Hong Kong, Burma, Thailand,
Malaya, Sumatra and Java1. Such specimens represent M. macrodens Maeda, 1982: they
agree closely in structure and size with M. magnater from New Guinea and are discussed
more fully under that species. All have been referred to M. schreibersii in the past when
this species was presumed to embrace a greater size range to include magnater as a subspecies
(e.g. BM(NH) 67.217 from Pahang by Hill (1972)) or before then (e.g. BM(NH) 9.1.5.460-
46 1 from Java by Thomas & Wroughton (1909) who in allocating these to blepotis remarked
upon their long forearms in comparison with other Javanese specimens) like those reported
and measured from Hainan by Allen (19386), which seem on size to be very similar to these
large examples and are referred without examination to macrodens by Maeda (1982). Speci-
mens from Java reported as blepotis by Sody (1930) are very large for this form and much
nearer in size to the large examples from Soekaboemi (BM(NH) 9. 1 .5.460-46 1 ) noted above
'See Addendum, p. 208

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J

Afghanistan; India (
Burma
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exMaeda, 1982)
Vietnam

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M. bismarckensis
Manus I, Bismarc

- D.

184 J. E. HILL

and are also referred to macrodens by Maeda, while M. ravus Sody, 1930, described concur-
rently, seems on measurements to be close to blepotis, to which Maeda refers its holotype
specimen.

Maeda (1982) has proposed wide ranging changes in the classification of Miniopterus from
Eurasia and especially Indo- Australia, many closely affecting that part of the genus formerly
held to include schreibersii, blepotis and a number of other related taxa. These changes may
be briefly summarized by comparison with the pre-existing classification, chiefly derived
from Tate (1941e) and from Ellerman & Morrison-Scott (1951), excluding purely Eurasian
taxa and ahofuscus and its allied forms:

Previous concept Maeda (1982)

M. schreibersii pallidus (Afghanistan) M. schreibersii

M. schreibersii fuliginosus M.fuliginosus

(Afghanistan to S China, Japan)

M. schreibersii chinensis (N China) M.fuliginosus

M. schreibersii eschscholtzii (Philippines) M. eschscholtzii

M. schreibersii blepotis (in part, Malaysia) M. blepotis

M. schreibersii blepotis (in part, Moluccas to Australia, M. oceanensis

Solomon Is)

M. schreibersii orianae (N Australia) M. eschscholtzii

To these he adds a further newly described species, M. haradai Maeda, 1982 from southern
Thailand.

As can be seen, the concept of one widely ranging polytypic species is replaced by Maeda
(1982) by five species. Moreover, the arbitrary statistical treatment used in this study
removes oceanensis, applied to specimens that formerly were referred to M. s. blepotis, to the
magnater subgroup of the fuliginosus group as Maeda defines them, while blepotis and
eschscholtzii form another group (the blepotis group), and schreibersii yet a third. There is no
doubt that the statistical analysis of metrical characters made by Maeda has identified and
codified many, if not all of the parameters used hitherto in the identification of the forms of
Miniopterus in Eurasia and Indo- Australia. The allocation of taxonomic rank to the group-
ings so characterised, however, seems very arbitrary and biased rather more heavily towards
the interpretation of metrical data than towards morphological and zoogeographical
considerations. Thus Maeda admits (p. 21) that there is no clear difference in general
morphological characteristics between all of the species of the fuliginosus group (i.e.
magnater, macrodens, oceanensis, fuliginosus} and those of the blepotis group (i.e. blepotis,
eschscholtzii, haradai), although he pointed out that there are some conspicuous differences
between the magnater subgroup (magnater, macrodens, oceanensis) and the blepotis group,
and also between the fuliginosus subgroup (fuliginosus) and the blepotis group. These state-
ments appear at least superficially to contain a contradiction. A comparison of Maeda's
distribution maps (figs 34, 35) for these taxa shows that (excluding magnater and macrodens,
discussed below) each replaces another in characteristic fashion across the region and
that none is sympatric although schreibersii and fuliginosus are nearly so in Afghanistan.
Maeda himself (p. 28) stated that according to his conclusions 'In many miniopterine bats,
the species belonging to the same group are usually allopatric in distribution'.

The interpretation by Maeda produces a number of curious distributional anomalies.
Thus oceanensis is said to occur in Australia, the Solomon Islands and tentatively in Burma
and Yunnan. However, the specimen from Burma (BM(NH) 67.2323) has a molar width
(m3-m3) of 6-8 and so, from his key, falls into the fuliginosus, blepotis or schreibersii groups,
while its toothrow (i'-m3) length of 7-3 and mastoid width of 8-7 take it to the fuliginosus
subgroup or the blepotis group. The length of the forearm is 48-0 and the greatest skull length
15-4, the condylobasal length 15-2: these values take it further in the key to fuliginosus.
Specimens from Yunnan (BM(NH) 12.7.25.4-5) have damaged skulls from which only the
palatal measurements can be drawn but these (Table 13) agree closely with those of
macrodens from Thailand and Koh Lak Island.

INDO-AUSTRALIAN BATS 185

A further anomaly is to be found in the distribution of eschscholtzii, as it is proposed by
Maeda. According to this author, it occurs in the Philippines whence it was originally
named, in northern and South Australia where in northern Australia according to Maeda it
displaces the name orianae Thomas, 1922, and on San Christobal Island in the Solomon
Islands. The South Australian record is drawn from Leche (1884) who originally identified
the material as M. schreibersii blepotis. The remaining Australian examples were originally
referred to M. s. orianae: those from San Christobal to M. s. blepotis by Hill (197 la).
However, the Australian examples referred initially to orianae do not always meet the
criteria laid down in Maeda's key for eschscholtzii, some (BM(NH) 22.10.8.2-3, 29.1.9.1)
keying more appropriately to blepotis and another (BM(NH) 29. 1 .9.3) producing some ratios
more appropriate to his parameters for fuliginosus. Similar considerations apply to speci-
mens (BM(NH) 54.901-903) from Rennell Island, from San Christobal Island (BM(NH)
67.1882-1883, 67.1885-1888) and one (BM(NH) 91.8.20.2) from Bugota Island, all in the
Solomon Islands.

A condition such as this might be expected within the distribution of a widespread poly-
typic species with populations scattered over numerous islands of different size: some
populations at least, even on widely distant islands, might be expected to resemble each
other in some features of size and proportion. A further point is that of the San Christobal
series (BM(NH) 67.1882-1888) identified formerly as blepotis, one example BM(NH)
67.1884) is determined by Maeda as oceanensis, while the remainder are referred to
eschscholtzii. This is apparently because this specimen has a slightly wider palate (m3-m3
7-0) than the others (m3-m3 6-6-6-9) and thus keys to the magnater subgroup of Maeda (and
hence to oceanensis) rather than to the schreibersii, fuliginosus or blepotis agglomerations, as
do the remainder of the series.

In Amboina Maeda (1982) records blepotis and oceanensis sympatrically. However,
members of the one series of specimens from Amboina Island in the British Museum
(Natural History) (BM(NH) 10.7.24.24-26) have very slightly wider palates and are very
slightly larger (Table 12) than specimens from the Kei Islands (BM(NH) 10.3.1.30,
10.3.1.70-71, 74, 94, 95) that otherwise they closely resemble. Consequently like BM(NH)
67.1884 from San Christobal they key to oceanensis while examples from the Kei Islands
are taken to blepotis, yet as can be seen (Table 12) actual differences are very small. The
Amboina specimens that Maeda refers to blepotis are in the Rijksmuseum van Natuurlijke
Historic: they include one skull (his population 47, p. 92). According to his table of measure-
ments (p. 127) this skull has a mastoid width of 7-0: it should therefore key out to his
magnater subgroup (including oceanensis) rather than to blepotis to which he refers it.

I am led to the conclusion that oceanensis from Amboina, New Guinea, the Solomon
Islands and eastern Australia should be considered a subspecies of M. schreibersii as here
understood, rather than be associated with M. magnater. Those from Timor reported as
magnater by Goodwin (1979) seem on size to be more properly referred to M. s. oceanensis:
all are linked to the smaller, more westerly populations by those from the Kei Islands. The
collections of the British Museum (Natural History) contain only a single example (BM(NH)
78.1.21.6) of M schreibersii from New Guinea: Maeda (1982) refers it to oceanensis and
records no other New Guinea specimen. However, I have examined two further examples (d1,
9 BBM 55803, 55893) from the Bernice P. Bishop Museum, Honolulu, that also come from
New Guinea. These are from Danowaria Cave, near Fak Fak, Fak Fak District, West Irian,
at 25 m.

In view of the various considerations advanced above, I have retained the view that all of
these medium-sized Miniopterus in Indo-Australia (excluding magnater and macrodens)
form a polytypic species extending from Afghanistan to Japan, the Philippine Islands, the
Solomon Islands and Australia. If allied to schreibersii of Eurasia then it extends westward
into western Europe and into Africa: there seems at present no reason to consider the Asiatic
and Indo-Australian populations distinct from schreibersii unless for example extensive
sympatry can be demonstrated in Afghanistan where M. s. pallidus is found at Kandahar and
M. s. fuliginosus at Jalalabad. In eastern Asia and in the Indo-Australian region this species

186 J. E. HILL

occurs from Japan (japoniae Thomas, 19066) and northeastern China (chinensis Thomas,
19080) to southern China and Vietnam (parvipes Allen, 1923), India and Sri Lanka
(fuliginosus (Hodgson, 1835)), the Philippine Islands (eschscholtzii (Waterhouse, 1845)),
Malaya and Sumatra east to Sulawesi and Java (blepotis (Temminck, 1 840)), the Molucca
Islands, New Guinea, the Solomon Islands and eastern Australia (oceanensis Maeda, 1982),
and in northern Australia (orianae Thomas, 1922c). Maeda (1982) associated oceanensis
with M. magnater but as already pointed out specimens referable to this form seem clearly
on comparison to represent the smaller and not the larger of the two similar sympatric species
although they are often rather larger than most of M schreibersii. No specimens from Thai-
land have been examined. Miniopterus haradai Maeda, 1982, from the southern part of the
country, however, is very close in size (Table 12) to specimens referred to M. schreibersii
from Vietnam and Malaya and may well be its local representative. For the present I would
consider it to be a subspecies of M. schreibersii.

Miniopterus magnater magnater Sanborn, 1931

Miniopterus schreibersii magnater Sanborn, 1931 : 26. Marienberg, 40 miles up Sepik R, Papua New
Guinea.

SPECIMENS EXAMINED. Papua New Guinea: d1 BM(NH) 78.893 No further locality (in alcohol; coll. P. A.
Morris); 2 rfd1, 8 99 BM(NH) 80.658-667 Mt Kainoi, Wau, Morobe, 7° 31' S, 146° 43' E; 2 d-d-,2 99
BM(NH) 82. 1 53-1 56 Cave at Avi, Mt Hagen (all in alcohol; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These specimens agree with M. m. magnater as it is represented by the limited
sample hitherto in the collections of the British Museum (Natural History).

DISCUSSION. The status of this species has been briefly discussed above. It is distinguished
from M. schreibersii by its generally longer forearm, larger skull, wider braincase and wider
palate. Maeda (1982) defined a magnater subgroup in some detail, but in including
oceanensis removed the basis of size characters in the definition of magnater, this newly
recognised taxon being little larger than other Indo- Australian specimens of M schreibersii
as here understood but having the palate very slightly widened. It is based on specimens
formerly allocated to M. schreibersii to which it is here allocated as a subspecies.

Specimens from localities from Hong Kong to Sumatra and Java have been briefly
mentioned under M. schreibersii blepotis to which they were once referred. With the
exception of some of the Thai and Sumatran examples they have been allocated to
macrodens by Maeda (1982) when describing that form. In detail these come from Hong
Kong (BM(NH) 65.1019, Old Mine, Silver Mine Bay), Burma (BM(NH) 32.11.1.3, Nam
Tisang), Thailand (BM(NH) 79.1425-1426, no further locality; (BM(NH) 17.2.6.3, Koh Lak
Island, off southeastern Thailand), Malaya (BM(NH) 67.217, Pine Tree Hill, Fraser's Hill,
Pahang), Sumatra (BM(NH) 0.8.2.11-14, Balighe) and Java (BM(NH) 9.1.5.460-461,
Soekaboemi, Preanger)1. The majority is male and all are fully adult or even old.

These specimens agree closely in structure and size (Table 13) with M. magnater from
New Guinea. According to Maeda (1982) M. macrodens differs from magnater in shorter
condylobasal length, shorter upper toothrow and narrower palate. A study of Maeda's
measurements shows that while macrodens is generally smaller in these respects than
magnater, the largest of the former overlaps the smallest of the latter, a circumstance
apparent in the specimens in London. Indeed, in some respects one Sumatran example of
macrodens exceeds magnater from New Guinea, yet in others falls at the lowest part of the
range of the New Guinea material. For this reason I consider macrodens Maeda, 1982 a
subspecies of M. magnater, distributed from Hong Kong and Burma at least to Sumatra and
Java. Maeda (1982) refers specimens without skulls from Amboina to macrodens but their
forearm length (populations 43, 45, pp. 110, 111) are compatible either with this form or
with magnater. specimens from Mount Flinders, South Australia, also referred to macrodens

'See Addendum, p. 208.

INDO-AUSTRALIAN BATS 187

by Maeda (population 81, pp. 1 14, 134) also have forearms of a similar length but the sole
skull measured agrees with macrodens.

Miniopterus tristis celebensis Peterson, 1981

Miniopterus tristis celebensis Peterson, 19816 : 841. Luwu, Wawondula, Sulawesi, 02° 38' S, 121° 21 '
E.

SPECIMENS EXAMINED. C Sulawesi: rfd BM(NH) 82.142-143 R Ranu, 1° 51' S, 121° 30' E (in alcohol,
skulls extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These specimens clearly support Peterson's contention that the Sulawesian
population of M. tristis is distinct from that of the Philippine Islands, from which
Sulawesian examples may be distinguished by their larger size and especially by the greater
length of the forearm and metacarpals. The examples recorded here bring the known total of
specimens of M. t. celebensis to four. Measurements: (BM(NH) 82.142, 82.143): length of
forearm 57-5, 57-1; IIIm 54-2, 53-3; IVm 51-9, 50-4; Vm 44-3, 44-0; length of tibia 24-1,
23-8; greatest length of skull 19-6, 19-7; condylobasal length 19-3, 19-2; condylocanine
length 18-4, 18-4; least interorbital width 4-1, 4-3; zygomatic width 1 1-0, — ; width of brain-
case 8-9, 9-0; mastoid width 1 0-0, 10- 1 ; d-c1 (alveoli) 6-0, 6- 1 ; m3-m3 8-8, 8-8; c-m3 8-3,8-1;
length of complete mandible from condyles 14-6, 14-7; length of right ramus from condyle
15-1, 15-l;c-m38-8, 8-7.

DISCUSSION. Hill (197 la) in recording M. tristis from the Solomon Islands and the New
Hebrides drew attention to the range of size variation exhibited by specimens referred to this
species over its then total range to the Philippine Islands. At that time, the collections of the
British Museum (Natural History) contained only the holotype of M. tristis sensu stricto,
BM(NH) 55.12.26.266, and from New Guinea there were specimens available from no more
than two widely separated localities. In view of the paucity of material this author left
unresolved the questions posed by the range of variation, except to associate robustior
Revilliod, 1914 from the Loyalty Islands with tristis as a distinct species rather than with M.
australis as a subspecies as it was described and had been considered hitherto in the
literature. Since then two widely divergent views of the classification of this group of bats
have emerged which although in agreement in accepting M. robustior as a distinct species
divide bats formerly regarded as M. tristis in different ways. Each of these treatments is based
to a considerable extent on the collections of the British Museum (Natural History).

Peterson (19816) recognised M. tristis in the Philippine Islands (M. t. tristis (Waterhouse,
1845)) and Sulawesi (M. t. celebensis Peterson, 198 16), with a second species, M.
propitristis, from western New Guinea (M. p. grandis Peterson, 19816), eastern New Guinea
(M. p. propitristis Peterson, 19816) and from the Bismarck Archipelago, the Solomon Islands
and the New Hebrides (M. p. insularis Peterson, 19816). This author had available a good
series of specimens from the Philippines and based his conclusions in part on a multivariate
analysis of a number of characters. Particular morphological features indicated as separating
propitristis from tristis include a greater degree of development of the sagittal crest over the
rear of the cranium, lower, flatter rostral profile, greater constriction, or less inflation, of the
posterior part of the braincase, a less conspicuous bulging of the supraoccipital (not the
supraorbital as stated by Peterson, p. 833) behind the lambdoidal crests, and a wider palate
in relation to the length of the toothrow.

Some of these features are not always especially effective in diagnosis. The sagittal crest,
for example is little different in the Sulawesian specimens (tristis) recorded here than in some
of those (propitristis) from western New Guinea, while the rostrum of some specimens from
western New Guinea is more steeply elevated as in Sulawesian specimens. The constriction
of the posterior part of the braincase varies slightly in New Guinea specimens and in a
number is little different from those from Sulawesi and although the posterior bulge of the

188 J. E. HILL

supraoccipital is more conspicuous in Philippine and Sulawesian specimens than in those
from western New Guinea, this condition is approached or equalled by others from the
eastern part of that island. As Peterson (fig. 7) shows, the width of the rear palate in relation
to the length of the toothrow provides a relatively useful diagnostic character, with little
overlap between specimens referred to tristis and those referred to propitristis. However, the
Sulawesian population seems equally referable on this feature to propitristis as to tristis:
indeed, with the addition of the specimens recorded here it could be regarded as corres-
ponding quite as closely to the propitristis grouping as it does to the Philippine examples.

Maeda (1982), on the other hand, regarded the Philippine population and those from both
western and eastern New Guinea and that on New Britain as referable to one species, M.
tristis, without subspecific separation. He removed the populations on the Solomon Islands
and the New Hebrides to a distinct species, M. melanesiensis, which he described as new on
account of its smaller size, the same grounds as those used by Peterson (198 1 b) in dividing M.
propitristis insularis from the New Guinea form M. p. propitristis, a feature also noted by
Hill (197 la). Maeda indicated in a footnote (p. 47) that melanesiensis is synonymous with
M. p. insularis, the paper by Peterson being unavailable when his study was in preparation.

Of these treatments that by Peterson (1981&) seems the more satisfactory. Maeda (1982)
examined only a single Philippine specimen and none from Sulawesi: it is apparent from the
tabulated measurements of Hill (197 la) and Peterson (1981&) that a good case can be made
for clinal variation among the populations from eastern New Guinea to the New Hebrides,
and both of these authors considered them conspecific. At the same time, their separation
from the Sulawesian and Philippine populations as a distinct species seems less strongly
based than Peterson (1981) implies. The morphological characters that he cites are not
wholly satisfactory and while they may be diagnostic between populations are not always so
when these populations are grouped in the way that Peterson suggested. Although in palatal
shape the two proposed species seem more definitely separable, there are indications that
the Sulawesian population may not fully conform to the criteria set by Peterson for tristis.
The representation remains small or limited in some instances: the Sulawesian population is
known from no more than four specimens, that from western New Guinea from one locality
only, and the population in eastern New Guinea only from three locations. Moreover,
Koopman (1982) considered only one species involved after examining the characters used
by Peterson in relation to specimens in the American Museum of Natural History. In these
circumstances I too prefer to regard all as subspecies of M tristis, i.e. M. t. tristis (Philippine
Islands), M. t. celebensis (Sulawesi), M. t. grandis (W New Guinea), M. t. propitristis (E New
Guinea) and M. t. insularis (with synonym M. melanesiensis Maeda, 1982 from San
Christobal Island, Solomon Islands) (Bismarck Archipelago to the New Hebrides).

Miniopterus tristis insularis Peterson, 1981

Miniopterus propitristis insularis Peterson, 1981& : 838. Kagilakulaku Cave, Hutuna, Rennell

Island, Solomon Islands 1 1° 45 ' S, 160° 15 ' E.
Miniopterus melanesiensis Maeda, 1982 : 19. St. ( = San) Christobal Island, Solomon Islands.

SPECIMENS EXAMINED. New Hebrides: 3 rfrf, 3 99 BM(NH) 73.1547-1552 Hog Harbour, Espiritu Santo
I, 1 50 ft (in alcohol, BM(NH) 73.1 548, 73. 1 552 without skulls; coll. Earl of Cranbrook, Royal Society
Expedition to the New Hebrides, 1971).

REMARKS. Specimens from Hog Harbour reported by Baker & Bird (1936) as M. schreibersii
were referred to M. tristis by Hill (197 la), this taxon being later divided (above) by Peterson
(198 \b). Others from St. Matthias Island in the Bismarck Archipelago were referred to

INDO-AUSTRALIAN BATS 189

insularis by Peterson (198 \b) but to M. oceanensis ( = M. schreibersii oceanensis as here
understood) by Maeda (1982). Further examination establishes them as belonging to
insularis.

DISCUSSION. Maeda (1982) has described Miniopterus bismarckensis from Manus I, Bis-
marck Archipelago as a new species in the tristis group, basing it on a series of five specimens
(BM(NH) 14.4.1.1 1, 1 1*-14) in the collections of the British Museum (Natural History). The
series appears as population 62 of Maeda (pp. 112, 130): although not noted as such by this
author, two (BM(NH) 14.4.1.12,14) are not fully adult. I am unable to agree with Maeda
that the length of the tibia in this series measures 16-5-17-5 (p. 112): in the two young
examples the tibia measures 17-4 and 16-9 respectively, in the three adults 19-0 (BM(NH)
14.4.1.11.11), 18-7(BM(NH) 14.4.1.1 1*) and 18-8 (BM(NH) 14.4.1.13). Forearm lengths in
these adults are 46-4, 45-0 and 44-4 respectively. Moreover, I fail totally to understand
whence Maeda obtained the total length for five skulls for this form (p. 1 30). The three adults
consist of the rostrum and mandible only, the whole of the cranium beyond the frontal region
missing. In BM(NH) 14.4.1.14 the basal part of the cranium and its posteriormost part except
the occiput is lost and the best value that can be obtained for total length is 15-1 while in
BM(NH) 14.4.1.12 the braincase has been damaged but the rear of the cranium survives.
It is thus the only more or less complete example but even so the premaxillae have been
damaged and the best value that can be obtained for total length is 16-3. It is difficult to
reconcile these facts with Maeda's published value for total length of skull of 15-8-17-3 with
a mean of 1 6-84 in the five examples. Other relevant values for BM(NH) 1 4.4. 1 . 1 2 are condy-
lobasal length 16-1, condylocanine length 15-6, least interorbital width 3-6, width of brain-
case 8-0, mastoid width 8-8 (these last two taken by eyepiece micrometer, the first of these
by extrapolation, one side of cranium damaged), m3-m3 c. 7-1 (left m3 missing), c-m3 7-0,
c-m3 7-3. Of the remaining specimens BM(NH) 14.4.1.1 1 has lost both rear upper molars
and cannot be used for palatal measurements, BM(NH) 14.4.1.11* (the holotype of bis-
marckensis) has the left upper molar displaced but the distance m3-m3 can be determined,
while BM(NH) 14.4.1.13 is palatally and dentally complete. Such palatal and dental
measurements as can be obtained appear in Table 13.

Although Maeda placed bismarckensis in the tristis group, the dimensions of the one
admittedly slightly young skull (BM(NH) 14.4.1.12) that provides enough measurements to
satisfy the requirements of his key take this specimen quite emphatically not to the tristis
group, but to thefuliginosus, blepotis or schreibersii groups, among which it then keys to the
magnater subgroup, fitting best with magnater or macrodens but perfectly with neither.
Peterson (198 \b) also apparently allocated four of these five specimens to M. propitristis
insularis ( = M. tristis insularis as understood here) but they are small for this form as he
defines it, a point confirmed by a specimen (BM(NH) 39.1407) from New Britain that
undoubtedly represents insularis and also by a long series of insularis from New Ireland that
I have been able to examine through the courtesy of Professor James D. Smith of California
State University at Fullerton, California. In these the greatest length of skull varied from
17-3-18-0 in 38 examples, the condylobasal length from 16-7-17-5 in 40 specimens, m3-m3
from 7-3-8-1 in 40 examples, and c-m3 from 7-1-7-6 in forty-one. Moreover, the values for
m3-m3 and c-m3 that can be established for the specimens from Manus fall considerably
outside the limits of M. t. insularis in the scatter diagram (p. 836, fig. 7) provided by
Peterson, although it appears that he has failed to plot the lowest values given for this form in
Table 5 (p. 839). In the relative width of the palate to the length of the toothrow the speci-
mens from Manus agree equally well with M. magnater magnater or M. magnater
macrodens. So far as can be determined the sagittal crest in the specimens from Manus is low
rather than well developed as in insularis, and the rostrum is less broadened than in that
form. While it is difficult to be definitive with such limited material, I am not convinced that
these specimens should be referred to the tristis group: it is possible that they represent a
form allied to M. magnater. The point can only be resolved by further, complete specimens
from Manus.

190 J. E. HILL

MURININAE

Murina suilla (Temminck, 1 840)

Vespertilio suillus Temminck, 1 840 : 224. Java.

SPECIMENS EXAMINED. Borneo: d" BM(NH) 82.548 Samusan Wildlife Sanctuary, between Cape Datu
and Gunong Pueh, W Sarawak, c. 1° 55 '-2° 00' N, 109° 34 '-109° 39' E (in alcohol; coll. N. A.
MacKenzie); d1 BM(NH) 82.555 Path to South Hitam, Gomantong, Sabah (in alcohol; coll. C. M.
Francis).

REMARKS. The few earlier records of M. suilla from Borneo are summarized by Medway
(1965, 1977). Length of forearm (BM(NH) 82.548, 82.555) 3 1 -4, 29-2.

Murina aurata Milne Edwards, 1 872

Murina aurata Milne Edwards, 1 872 : 250, pi. 37b, fig. 1 , pi. 37c, fig. 2. Moupin, Szechwan, China.
(l)Harpiocephalus feae Thomas, 1891 : 884; 1892 : 926-927. Biapo, Karen Hills, Burma.

SPECIMENS EXAMINED. Nepal: d BM(NH) 75.301 2 km E of Mukut, N of Dhaulagirui, 28° 50' N, 83°
25 ' E, 1 3,500 ft (skin, skull; coll. G. B. Corbet).

Thailand: d BM(NH) 82.162 Doi Ithanon, Chom Thong, Chiangmai, c. 2550 m, 18° 35 ' N, 98° 29'
E (in alcohol, skull extracted; coll. D. S. Melville).

REMARKS. These specimens agree closely with the detailed account of M. aurata by Maeda
(1980) who also examined and listed BM(NH) 75.301. The species has not before been
recorded from Thailand; it may be readily recognised by its small size (length of forearm (7)
28-5-32-0, condylobasal length (6) 12-3-13-1, c-m3 (7) 4-3^-6; values in part from Maeda,
1980) and distinctive coloration, the dorsal pelage basally brownish, the hairs with bright,
lustrous yellow brown tips, the ventral surface blackish, overlaid with white or greyish white
tipping. The upper surface of the forearm and thumb is sparsely covered with brown hairs
and the dorsal surface of the uropatagium, the tibiae and feet with a moderate cover of
longer, brownish or chestnut hairs. Externally it is very similar to M. aenea Hill, 1964 from
Malaya but this species has a longer forearm (length (2) 33-7-35-0) and a much larger skull
(condylobasal length (2) 1 5-2-1 5-9, c-m3 (2) 5-8-5-9) that is nearer in size to that of M. cyclo-
tis Dobson, 18726 and has bronze tipped pelage.

DISCUSSION. Ellerman & Morrison-Scott (1951) listed feae (Thomas, 1 89 1 ) as a subspecies of
M. aurata while Maeda (1980) tentatively synonymised the two but did not examine the
holotype of feae, which is probably in the Museo Civico di Storia Naturale Giacomo Doria in
Genoa. Certainly from the description feae is very like aurata, differing chiefly in the absence
of any distinct fringe of hairs along the edge of the uropatagium and in its duller, more
brownish coloration. Moreover, the holotype did not come from northern Burma as Maeda
indicated, but from the Carin or Karen Hills, northeast of Toungoo, in the more southerly
part of the country. Indeed, this location is not greatly distant from Doi Ithanon, whence
aurata is here recorded. In the restricted sense of Maeda (1980) who excluded specimens
from northeastern Asia and Japan as a distinct species only twelve examples (including the
holotype of feae) of M. aurata have been reported, their range extending through the
montane parts of northern India, Nepal, Burma and Thailand to Szechuan and Yunnan.

Murina tubinaris (Scully, 1881)
Harpiocephalus tubinaris Scully, 1881: 200, fig. 1 . Gilgit, Kashmir.

INDO-AUSTRALI AN BATS 1 9 1

SPECIMEN EXAMINED. Thailand: d BM(NH)82.163 Doi Inthanon, Chom Thong, Chiangmai,c. 1650 m,
1 8° 35 ' N, 98° 29 ' E (in alcohol, skull extracted; coll. D. S. Melville).

REMARKS. This is the first of M. tubinaris to be reported from Thailand. It agrees closely
with specimens of this species from northeastern India and northern Burma and is greyish
brown dorsally and greyish white ventrally, the upper surface of the forearms, tail, legs and
uropatagium sparsely or only moderately clothed with greyish brown hairs. The anterior
premolars (pmf) are considerably reduced to one half or less than one half the crown area of
the second premolars (pm|) and to about two thirds of their height. The species differs from
M. cyclotis and from M. huttonii (Peters, 1 872a) in its chiefly greyish and not brownish dorsal
colour, in more convergent upper toothrows, more reduced anterior premolars and in less
massive molar teeth. Although similar in length of forearm to M. aurata it has a slightly
larger skull (length of forearm (19) 29-0-32-0; condylobasal length (12) 13-3-14-2, c-m3 (15)
4-8-5-2).

DISCUSSION. Hill (1962) recorded M. tubinaris from northern Burma, considering it a species
distinct from M. huttonii with which it had been associated as a subspecies by Ellerman &
Morrison-Scott (1951), the same author later (1964) describing and discussing it in more
detail. Its discovery in the northern part of Thailand is not unexpected: it occurs otherwise
from Kashmir through northern India and Burma apparently to Vietnam.

Murina cyclotis cyclotis Dobson, 1 872

Murina cyclotis Dobson, 18726 : 210; 1873c : 206. Darjeeling, northeastern India.

SPECIMENS EXAMINED. Thailand: d BM(NH) 82.164 Doi Inthanon, Chom Thong, Chiangmai, c.
1 650 m, 1 8° 35 ' N, 98° 29 ' E (in alcohol); d BM(NH) 82.165 Doi Pha Horn Pok, c. 1 650 m (in alcohol,
skull extracted; both coll. D. S. Melville).

REMARKS. As with the earlier example of M. cyclotis reported from Chiangmai by Hill &
Thonglongya (1972) these specimens agree closely with the definition of the species by Hill
(1964) and with examples in the British Museum (Natural History) from Burma. Dorsally M.
cyclotis is generally some shade of rufous brown or ferrugineous, the ventral surface
predominantly greyish white: the forearms, tail, tibiae, feet and uropatagium have a dense
clothing of short, golden brown hairs on the upper surface. Young specimens are darker and
browner. Coloration apart, the nominate subspecies is larger cranially (length of forearm (9)
30-2-34-8, condylobasal length (8) 14-7-15-8, c-m3 (9) 5-0-5-7) than either M. aurata or
M. tubinaris: a yet larger subspecies M. c. peninsularis Hill, 1964 occurs in the Malay
Peninsula and in Borneo (vide infra) (length of forearm (9) 33-7-38-8, condylobasal length (9)
15-4-16-9, c-m3 (9) 5-5-6-2). Dentally, M. cyclotis may be distinguished by the lack of
reduction of the anterior upper premolar (pm2) which in crown area is equal to three quarters
or more of the crown area of the second upper premolar (pm4), and by the reduction of the
posterior triangles or talonids of its molar teeth, the paracones and protocones of the first
(m1) and second (m2) upper molars being slightly reduced, the hypoconids and entoconids of
the corresponding lower teeth very much more so, sometimes scarcely evident, and separated
from the protoconids and metaconids only by a very narrow trough.

DISCUSSION. Four species of Murina have now been reported from Thailand, namely M.
aurata, M. tubinaris, M. cyclotis and M. huttonii (Peters, 18720), the last being reported from
Chiangmai by Hill (1975) and Lekagul & McNeely (1977). This rather larger (length of fore-
arm (1 1) 33-2-37-0, condylobasal length (10) 15-3-16-7, c-m3 (1 1) 5-8-6-1) species is more
brownish than M. tubinaris and like M. cyclotis does not have markedly convergent upper
toothrows. The anterior upper premolar (pm2) is a little more reduced than in the latter
species, the tooth one half to two thirds the crown area of the second upper premolar (pm4),
while the hypoconids and entoconids of the first and second (m^) lower molars are well
developed and separated from the protoconids and metaconids by a comparatively wide
trough.

192 J. E. HILL

Murina cyclotis peninsularis Hill, 1964

Murina cyclotis peninsularis Hill, 1964: 55. Ulu Chemperoh, near Janda Baik, Bentong District,
Pahang," Malaya, c. 3° 18 ' N, 101° 50' E, 2000 ft.

SPECIMENS EXAMINED. Borneo: cf BM(NH) 78.1543 Melinau R, Gunung Mulu National Park, Sarawak
(in alcohol, skull extracted; coll. D. R. Wells, Royal Geographical Society Expedition to Gunung Mulu,
d BM(NH) 82.556 Sepilok Forest Reserve, Sabah (in alcohol, skull extracted; coll: C. M. Francis).

REMARKS. Only M. suilla among tube-nosed bats has been reported until now from Borneo,
M. cyclotis being known hitherto from the mainland of southeastern Asia and from single
specimens from the islands of Hainan off southeastern China and Mindanao in the Philippine
Islands. These Bornean specimens agree closely with M. c. peninsularis from Malaya, to
which they are referred.

Measurements (BM(NH) 78.1543, 82.556), with those of seven Malayan examples in
parentheses: length of forearm 37-0, 35-6 (33-7-38-8); greatest length of skull 18-8, 18-4
(17-4-18-9); condylobasal length 16-9, 16-8 (15-4-16-7); length orbit-gnathion 4-6, 4-5
(4-2-4-5); palatal length 9-3, 9-2 (8-2-9-3); rostral width at lachrymals 6-3, 6-3 (5-5-6-0); least
interorbital width 4-3, 4-5 (4-3^-6); zygomatic width 11-3, 10-9 (9-9-10-9); width of brain-
case 8-3, 8-3 (7-7-8-2); height ofbraincase 7-1,6-8 (6-6-6-9); mastoid width 9-4, 9-2 (8-3-9-1);
c'-c1 (cingula) 5-3, 5-2 (4-5-5-2), (alveoli) 5-1,5-1 (4-5-5-1); m3-m3 6-1, 6-0 (5-6-6-0); c-m3
6-1, 6-0 (5-5-6-2); length complete mandible from condyles 12-3, 12-0 (10-9-12-1); length
right ramus from condyle 12-8, 12-5 (1 1-2-12-7); c-m3 6-4, 6-2 (6-0-6-4).

DISCUSSION. There exist now apparently nine known examples of this large subspecies: in
addition to those recorded here from Borneo and those reported from Malaya by Hill (1964,
1972) the collections of the British Museum (Natural History) also include two further speci-
mens from Malaya, BM(NH) 73.630-63 1 , both male, from Telok Chempedak, Pahang.

Murina sp.

SPECIMEN EXAMINED. Molucca Islands: 9 BM(NH) 82.144 (juv.) Latuhalat, Amboina I (in alcohol; coll.
B. H. Gaskell, 'Operation Drake').

REMARKS. It is not possible to attempt any further identification of this juvenile example in
which the milk dentition is not yet completely lost and the adult dentition only partially
erupted.

DISCUSSION. Records of tube-nosed bats of the genus Murina from the region east of Borneo
are few (Van Deusen, 1961). The genus has yet to be reported from Sulawesi itself, although
it has been recorded from the nearby island of Peleng by Tate (\94\e): it is also known
from the Lesser Sunda Islands (Flores, Sumbawa), from the Moluccas (Ceram, Buru,
Goram), New Guinea and the Bismarck Archipelago (Ruk ( = Umboi) I). According to Van
Deusen (1961) the specimens forming the basis of these records total fourteen: more recently,
Richards (1981) and Richards et al. (1982) have reported a further example of the genus from
Queensland, Australia. The specimen recorded here is the first of Murina to be recorded
from Amboina, where clearly it might be expected to occur.

Three taxa have been named from the region: florium Thomas, 19086 from Flores, lanosa
Thomas, 19106 from Ceram, and toxopei( = toxopeusi) Thomas, 1923 from Buru. Laurie &
Hill (1954) considered lanosa and toxopeusi conspecific with florium, regarding specimens
from Sumbawa originally recorded by Mertens (1936) as M. suilla more probably
representative of M / florium. Tate (\94\e) found the specimens from Peleng Island
conformed closely to M. florium, while Hill (in Van Deusen, 1961) thought the example
from Ruk I, originally recorded by Thomas (\9\4a) without specific identification, to be

INDO-AUSTRALIAN BATS 193

close to toxopeusi. Van Deusen (1961) considered that an evaluation of the characters on
which these taxa were based seemed to indicate that only one good species existed east of
Wallace's Line, but through lack of comparative material did not attempt to apply a specific
name to the specimen that he recorded from New Guinea. Richards et al. (1982) referred the
specimen from Queensland to M.florium.

A further examination of the limited material in the collection of the British Museum
(Natural History) supports the view that only one species is involved, Jlorium, lanosa and
toxopeusi being almost certainly conspecific. All are larger, especially cranially, than M.
suilla from Java and its closely related congeners balstoni Thomas, 1908&, also from Java,
and canescens Thomas, \923a, from Nias Island, off the west coast of Sumatra, both very
similar to suilla and possibly no more than subspecifically different from it. Only the holo-
types offlorium (BM(NH)63.12.26.14) and toxopeusi (BM(NH)23.1 .2.27) are available, with
the holotype (BM(NH) 10.3.4.24) and one other example (BM(NH) 7.1.1 .482) of lanosa from
Ceram, together with a specimen (BM(NH) 10.3.4.115) from Goram referred to lanosa by
Laurie & Hill (1954) and the specimen (BM(NH) 14.4.1.33) recorded from Ruk Island by
Thomas (\9\4a). Removal of the skull from the specimen from Goram confirms that it rep-
resents lanosa rather than toxopeusi. The example from Ruk, however, has a shallow median

Table 14 Measurements ofMurina from east of Wallace's Line.

"T r-

<N <N

CO — '

O ff-t

•"• <N as

(N

oo

I
1

s u
'£ >>"

pl

o «5

5 ^

C3 CQ

i?i

s; — '

-2 — '

1^

O DO

•S o.

r. ?/lorium
BM(NH) 14.4.
uk I, Bismarck

ON C' 3
III Bl|

II'J •§•§!

^Q > £2 *
^•%Z ^x%

^ffitt,

^EU

^CQU

^ coO

^£ CQ

31 o(*

^>z ^cticx

Length of forearm

34-8

36-9

36-4

36-9

34-4

35-4

35 35-7

Greatest length

of skull

—

16-7

—

17-3

16-7

—

15-4 16-9

Condylobasal length

—

15-5

—

15-7

15-3

—

15-8

Condylocanine length

—

14-8

—

15-1

14-7

—

Palatal length

—

7-2

—

7-0

7-1

—

Width across

anteorbital foramina

4-5

4-3

4-5

4-5

4-2

4-3

Least interorbital width

4-2

4-5

4-3

4-5

4-2

4.4

4-3 4-6

Zygomatic width

—

9-2

—

9-5

9-3

—

8-9 10-2

Width of braincase

—

7-9

—

7-9

7-6

—

7-8 8-1

Mastoid width

—

8-2

—

8-4

8-0

—

c'-c1 (alveoli)

3-8

3-9

4-0

4-0

3-8

4-0

m3-m3

5-6

5-4

5-6

5-6

5-5

5-7

6-1

c-m3

5-3

5-5

5-6

5-6

5-5

5-6

5-3 5-8

Length complete

mandible from

condyles

10-8

11-3

11-1

11-2

11-0

11-3

Length right ramus

from condyle

11-1

11-4

11-6

11-5

11-4

11-6

c-m3

6-0

6-1

6-2

6-2

6-1

6-3

6-2

194 J. E. HILL

rostral sulcus and its anterior upper premolar (pm2) is longitudinally compressed and rather
short, in these respects agreeing more closely with the holotype of toxopeusi than with
lanosa. Although the Goram specimen has a slightly shallower sulcus than the examples
of lanosa from Ceram, its pm2 is less compressed than in the specimen from Ruk. These
specimens, with the exception of the skull of the specimen from Goram, were first examined
in 1961 at the instigation of the late Mr H. M. van Deusen of the American Museum of
Natural History.

KERIVOULINAE

Kerivoula hardwickei hardwickei (Horsfield, 1 824)

Vespertilio hardwickii Horsfield, 1 824 (unpaginated). Java.

SPECIMENS EXAMINED. C Sulawesi: rfcf BM(NH) 82.145-146 R Ranu, 1° 51' S, 121° 30' E (in alcohol,
skulls extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. The larger (BM(NH) 82.146) of these examples tends in some respects to bridge
the slight disparity in certain elements of size noted by Hill (1965) between specimens of K.
h. hardwickei from more easterly localities (Philippine Islands, Borneo and Sulawesi) and
those from more westerly locations (Java, Sumatra and Malaya). Measurements (BM(NH)
82.145, 82.146): length of forearm 31-7, 33-1; greatest length of skull 12-7, 13-6;
condylobasal length 12-3, 12-9; condylocanine length 1 1-9, 12-6; least interorbital width 3-1,
3-3; zygomatic width 7-7, 8-0; width of braincase 6-7, 6-7; depth of braincase 5-5, 5-7;
mastoid width 7-0, 7-3; c'-c1 (alveoli) 3-3, 3-3; m3-m3 4-9, 5-2; c-m3 5-1, 5-4; length complete
mandible from condyle 9-0, 9-6; length right ramus from condyle 9-2, 9-8; c-m3 5-4, 5-7.

Kerivoula papillosa (?) malayana Chasen, 1940

Kerivoula papillosa malayana Chasen, 1940 : 55. Ginting Bedai, Selangor-Pahang boundary, Malaya,
2300 ft.

SPECIMEN EXAMINED. C Sulawesi: rf BM(NH) 82.147 R Ranu, 1° 51' S, 121° 30' E (in alcohol, skull
extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. This is the first of K. papillosa to be recorded from Sulawesi, the species being
known hitherto from northeastern India to Sumatra, Borneo and Java. It agrees closely with
K. p. malayana, reported hitherto as far east as Borneo. The collections of the British
Museum (Natural History) do not include specimens of K. p. papillosa (Temminck, 1840)
from Java: according to Tate (\94\e), Bornean specimens agree in all essential respects with
the type from Java but Chasen (1940) remarked that the few skins that he had seen from Java
were rather greyer than those that he referred to malayana, with which he included Bornean
examples. Chasen noted further that the skulls of Javan specimens are smaller than those of
Malayan and Bornean examples, with a maximum length of about 17 but Tate gives
measurements of a 'cotype' in the Rijksmuseum van Natuurlijke Historic, Leiden that
suggest that this may not be so.

Measurements of the Sulawesian specimen: length of forearm 44-9; greatest length of skull
18-0; condylobasal length 16-5; condylocanine length 16-4; least interorbital width 3-6;
zygomatic width 11-5; width of braincase 8-4; depth of braincase 7-3; mastoid width 9-1;
c'-c1 (alveoli) 4-6; width inside m'-m1 3-1; m3-m3 6-6; c-m3 7-4; m1"3 3-9; length complete
mandible from condyle 12-6; length right ramus from condyle 13-0, c-m3 7-9.

INDO-AUSTRALIAN BATS 195

Kerivoula muscina Tate, 1 94 1

Kerivoula muscina Tate, \94\e : 586. Lake Daviumbu, 6 miles above mouth of Strickland R, middle
Fly R, Western Division, Papua, c. 20 m.

SPECIMEN EXAMINED. Papua New Guinea: d BM(NH) 80.668 Buso, Morobe, 7° 17' S, 147° 08' E (in
alcohol, skull extracted; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. In size, cranial architecture and dentition this specimen agrees closely with the
descriptions of K. muscina by Tate (\94\e) and Hill (1965). Measurements: length of
forearm 33-0; greatest length of skull 13-5; condylobasal length — ; condylocanine length — ;
least interorbital width 3-2; zygomatic width — ; width of braincase 6-8; mastoid width 7-2;
c'-c1 (alveoli) 3-2; m3-m3 5-0; width palato-pterygoid extension 1-7; width of cochleae 24;
width apart of cochleae 1 -3; c-m3 5-5; m1"3 2-7; length complete mandible from condyles 9-8;
length right ramus from condyle 10-0; c-m3 5-9.

MOLOSSIDAE

Tadarida (Chaerephon) plicata (Buchannan, 1800)

Vespertilio plicatus Buchanan, 1800 : 261, pi. 13. Bengal.

SPECIMENS EXAMINED. Sex indet. BM(NH) 78.2509 Lubang Rusa, Gunong Mulu National Park, 4th
Division, Sarawak (mummified, skull; coll. Earl of Cranbrook, Royal Geographical Society Expedition
to Mount Mulu); 2d<f, 9 (all yg.) BM(NH) 931-933 Mouth of Deer Cave, Gunung Mulu National
Park (in alcohol; coll. C. H. Fry, Royal Geographical Society Expedition to Mount Mulu); 4 d1 cf (3 yg.),
2 99 (1. yg.) BM(NH) 956-961 Deer Cave (in alcohol; coll. P. Chapman, Royal Society Expedition to
Mount Mulu); cf, 9 BM(NH) 79.1400-1401 Deer Cave (in alcohol; coll. S. Proctor, Royal Society
Expedition to Mount Mulu); sex indet. BM(NH) 79.953-954 Madai Cave, Sabah (skeletal parts); d
Sabah Museum No. NH 95 Madai Cave (in alcohol) (all from Sabah Museum).

REMARKS. There are relatively few published records of T. plicata from Borneo. The species
has been reported previously from Pengkalan Forest, 1 st Division in Sarawak by Medway
(1965) and in Sabah from Madai by Chasen (1931), Gomanton by Pryer (1884) and from
Sapagaya by Davis (1962). Length of forearm in 6 adults 40-4-43-5.

DISCUSSION. Specimens from Borneo are usually referred to T. p. plicata (Hill, 19616;
Medway, 1965). As Davis (1962) pointed out, however, the forearm in Bornean specimens
averages shorter than in those from Java (IT. p. dilatata Horsfield, 1822) and is also shorter
on the average than in the nominate subspecies. In forearm length they approach the
Philippine form T. p. luzonus Hollister, 1913 or the more doubtful T. p. tenuis (Horfield,
1 822) from eastern Java, recognised by Chasen ( 1 940), Tate ( 1 94 1 c) and Hill (1961 b).

Tadarida (Chaerephon) jobensis jobensis (Miller, 1902)

Nyctinomusjobensis Miller, 1902 : 246. Ansus, Jobi I, Geelvinck Bay, West Irian.

SPECIMENS EXAMINED. Papua New Guinea: 9 rfrf, 14 9$BM(NH) 80.669-679, 80.681-692 Buso,
Morobe, 7° 1 7 ' S, 1 47° 08 ' E (in alcohol; coll. B. H. Gaskell, 'Operation Drake').

REMARKS. These specimens with length of forearm (23) 43-3-47-9 confirm that in T. j.
jobensis the forearm is rather shorter than in T.j. colonicus (Thomas, 1906a) from northern
Australia which has a forearm length of (6) 47-0-52-2, but is generally longer than in T. j.
solomonis (Troughton, 193 1) from the Solomon Islands, for which the describer gave a fore-
arm length of (200) 40-5^5.

196 J. E. HILL

Tadarida (Chaerephon) jobensis bregullae Felten, 1964

Tadaridajobensis bregullae Felten, 1964 : 9. Malo I, New Hebrides.

SPECIMENS EXAMINED. New Hebrides: 9, d BM(NH) 73.1553-1534 Tan Lensingo Cave, Abounatori
Village, Malo I, 30 m; dd BM(NjH) 73.1555-1568 Abounatori Village, 30 m (BM(NH) 73.1553-1566
in alcohol, 73. 1 567-1 568 skins, skulls; obtained by the Earl of Cranbrook, Royal Society Expedition to
the New Hebrides, 1971).

REMARKS. These are the first of T. j. bregullae to be reported since the subspecies was
described. The length of the forearm in 1 5 fully adult examples ranges from 47-6-5 1 -4, rather
more widely than in the ten of the original specimens measured and reported by Felten. In
forearm length the subspecies is similar to T.j. colonicus but according to the describer has a
narrower skull, a contention supported by skulls from the present series.

Cranial measurements of BM(NH) 73.1567, 73.1568 with (in parentheses) those of 8 of
colonicus, from Felten, (1964) who also included measurements made by Hill (19616);
condylobasal length 18-5, 18-4 (18-3-19-7); least interorbital width 4-1, 4-0 (4-(M-4);
zygomatic width 11-8, 11-4 (11-9-13-1); width of braincase 9-5, 9-3 (9-5-10-4); mastoid
width 10-8, 10-7 (11-1-11-9); c'-c1 (alveoli) 5-4, 5-2 (5-2-6-1); m3-m3 (alveoli) 8-5, 8-3
(8-9-9-5); c-m3 (crowns) 7-5, 7-5 (— ), (alveoli) 7-2, 7-2 (7-3-7-7).

DISCUSSION. Freeman (1981) synonymised T.j. bregullae but whether into T.j. jobensis, T.j.
colonicus or T. j. solomonis is not clear. Its various dimensional differences from these,
however, suggest that at least for the present it should be retained.

Tadarida (Mormopterus) beccarii astrolabiensis (Meyer, 1899)

Nyctinomus astrolabiensis Meyer, 1899 : 19, pi. 10, figs. 19, 30, pi. 11, fig. 6. Bongu, Astrolabe Bay,
West Irian.

SPECIMENS EXAMINED. Papua New Guinea: 99 BM(NH) 80.680, 80.693, 80.695 Buso, Morobe, 7° 17'
S, 147° 08' E (in alcohol, skulls of BM(NH) 80.693, 80.695 extracted; coll. B. H. Gaskell, 'Operation
Drake').

Australia: 9 BM(NH) 78.2708 Mount Wando, 36 miles SSW of Emerald, C Queensland, 23° 56' S,
147° 46 ' E (coll. G. Gordon & B. C. Lawrie); d BM(NH) 78.2709 Barret's Lagoon, NE Queensland, 1 8°
03' S, 145° 58' E (coll. R. J. Grimes) (both in alcohol, skulls extracted; from F. R. Allison, through
National Parks and Wildlife Service).

REMARKS. Relatively few specimens of T. b. astrolabiensis have been recorded: these agree
closely with a small series (BM(NH) 11.11.11.18-20) in the British Museum (Natural
History) from the Mimika River, West Irian. The nominate subspecies T. b. beccarii (Peters,
1881) from Amboina Island is represented in London by no more than three examples
(BM(NH) 10.7.25.17-19) and has a slightly shorter and in some respects slightly narrower
skull with a shorter toothrow than T. b. astrolabiensis.

Measurements of specimens from New Guinea, including the Mimika River series
(BM(NH) 1 1.1 1.1 1.18-20) first measured by Hill (19616), with (in parentheses) correspond-
ing values for T. b. beccarii from Amboina (BM(NH) 10.7.25.17-19 in that order, skulls
BM(NH) 10.7.25.18-19): length of forearm (6) 33-3-35-9 (33-4, 34-3, 34-9); greatest length of
skull (5) 17-3-18-3 (16-6, 16-6); condylobasal length (5) 16-5-17-3 (16-0, 16-1); condylo-
canine length (5) 16-2-16-9 (15-7, 15-7); least interorbital width (5) 4-2^-7 (4-2, 4-2);
zygomatic width (2) 11-0, 11-7 (— , — ); width of braincase (5) 8-0-8-5 (8-0, 8-1); mastoid
width (5) 10-3-10-8 (9-7, — ); c'-c1 (alveoli) (5) 4-5-5-0 (4-6, 4-2); m3-m3 (alveoli) (5) 7-7-8-2
(7-5, 7-3); c-m3 (crowns) (5) 6-4-6-6 (5-9, 6-0), (alveoli) (5) 6-2-6-4 (5-8, 5-9); pm4-m3 (alveoli)
(5) 4-7-4-9 (4-5, 4-6); length complete mandible from condyles (4) 11-9-12-6 (11-2, 11-5);
length right ramus from condyle (4) 12-6-13-0 (11-7, 12-0); c-m, (crowns) (4) 6-8-7-0 (6-4,
6-4).

INDO-AUSTRALIAN BATS 197

The species does not appear to have been formally reported hitherto from Queensland or
indeed from Australia but F. R. Allison in Honacki et al. (1982) includes N Australia within
its distribution. The two specimens reported here are nearer in size to T. b. astrolabiensis
than to the nominate subspecies but have longer forearms and slightly more massive canine
teeth. It seems possible that adequate series of specimens might permit the recognition of an
Australian subspecies. Measurements BM(NH) 78.2708, 78.2709): length of forearm 37-9,
364; greatest length of skull 17-1, 18-1; condylobasal length 16-9, 17-9; condylocanine
length 16-1, 17-1; least interorbital width 4-0, 4-5; zygomatic width — , — ; width of braincase
8-2, 8-6; mastoid width — , 10-9; c'-c1 (alveoli) 6-2, 6-5; m3-m3 (alveoli) 7-3, 8-2; c-m3
(crowns) 6-4, 6-7, (alveoli) 6-2, 6-5; pm4-m3 (alveoli) 4-8, 5-1; length of complete mandible
fromcondyles 12-1, 12-9; length right ramus from condyle 12-8, 1 3-5; c-m3 (crowns) 6-8, 7-2.

The small anterior upper premolar (pm2) is absent from the left side of the jaw but present
in the other side in BM(NH) 80.693, the opposite condition occurring in BM(NH) 80.695,
while in BM(NH) 80.680 it is absent from both sides: it is present in both sides in the
Australian examples. The absence of this tooth from one of the Amboina specimens (T. b.
beccarii} and from two of those from the Mimika River (T. b. astrolabiensis) was noted by Hill
(19616).

DISCUSSION. Freeman (1981) raised Mormopterus to generic rank. Certainly it is one of the
most distinctive of the subgenera of Tadarida, its relatively delicate, angular ears of thin
integument, with small antitragal lobe, and its finely wrinkled upper lip setting it apart from
the other subgenera.

Tadarida sp.

SPECIMEN EXAMINED. Papua New Guinea: Sex indet. BM(NH) 80.696 (nurseling). Buso, Morobe, 7° 17
S, 147° 08 ' E (in alcohol; coll. B. H. Gaskell, 'Operation Drake').

Otomops papuensis Lawrence, 1948

Otomops papuensis Lawrence, 1948 : 413. Vailala R (flows into Gulf of Papua about 15 miles W of
Kerema).

SPECIMENS EXAMINED. Papua New Guinea: rf9 BM(NH) 73.136-137 Maul, Mount Suckling, 300m
(BM(NH) 73.136 skin, skull, 73.137 in alcohol; coll. J. I. Menzies).

REMARKS. These appear to be the first of O. papuensis to be reported since its description:
they conform closely to the original account. The dorsal surface is reddish brown, the hairs
slightly paler at the base, the overall colour a little darker on the crown, the lower back and
along the junction of the wings with the body. There is a band of paler, light brown hairs
across the shoulders. The ventral surface is a paler reddish brown than the back, with a
diffuse, slightly paler area across the base of the throat. The skull is as it is described by
Lawrence but the male skull examined has a low sagittal crest that extends posteriorly to join
slight lambdoidal ridges. As Lawrence points out, these project forward in an anteriorly
directed V-shape to leave the rear of the skull smoothly rounded.

External measurements (BM(NH) 73.136, 73.137): length of forearm 49-6, 50-2; cranial
measurements (BM(NH) 73.136): greatest length of skull 19-6; condylobasal length 17-9;
condylocanine length 1 7-2; basal length 16-1; length from posterior edge of narial aperture to
highest point of cranium 14-7; least interorbital width 4-3; zygomatic width 10-4; width of
braincase 9-4; mastoid width 10-6; bottom of bulla to highest point of braincase 9-7; height of
jugal projection of zygoma 2-1; c'-c1 (alveoli) 4-2; m3-m3 (alveoli) 7-9; i2-m3 8-2; c-m3 7-2;

198 J. E. HILL

length complete mandible from condyles 12-2; length right ramus from condyle 12-5; c-m3
7-5.

DISCUSSION. Two species of Otomops occur in New Guinea: the other, O. secundus Hay man,
1952, known so far from Tapu, Upper Ramu River Plateau, Papua New Guinea, differs from
O. papuensis chiefly in its longer forearm (length 57-58), dark chocolate brown general
coloration, the presence of a broad, well-defined buffy grey mantle and of a narrow but
conspicuous line of white hairs along the dorsal surface of the endopatagium at its junction
with the body. There is also a patch of white hairs at the anterior part of the inner or medial
base of each ear, flanking the interaural pocket, a point unremarked by Hayman in the
original account. A similar patch of hairs occurs in O. wroughtoni (Thomas, 1913) from
India but the base of the outer margin of the ear in O. secundus lacks the sprinkling of white
hairs found in this species. There is no white at the base of the ear in O. papuensis, the hairs
instead a rich reddish brown with slightly paler bases. Otomops secundus seems in fact
nearer in external size and coloration to O.formosus Chasen, 1939 from Java or to the larger
O. wroughtoni. The representation of all except O. wroughtoni is small: all have skulls
essentially similar in structure, while the skull of O. secundus is closely similar in size to that
ofO. papuensis.

Summary

In the Megachiroptera Rousettus amplexicaudatus stresemanni from New Guinea is considered a valid
subspecies; further material of the Sulawesian species Rousettus celebensis, Styloctenium wallacei and
Thoopterus nigrescens is reported and where appropriate discussed; a number of species of Dobsonia
are reviewed; Indo-Australian Cynopterus are examined in detail and C. titthaecheileus considered
specifically distinct; Javan Chironax melanocephalus is re-described and further examples are reported
from Sumatra and Sulawesi; a series of Harpyionycteris celebensis is reported and discussed; Nyctimene
celaeno is provisionally considered specifically distinct; Eonycteris in Indo-Australia is briefly reviewed
and the nomenclature of Macroglossus re-examined; Synconcyteris is reviewed and further specimens
ofNotopteris macdonaldi macdonaldi are reported and discussed.

Among the Microchiroptera Rhinolophus celebensis and R. borneensis are considered distinct
species and their subspecies allocated with R. importunus considered a subspecies of the latter. The
Sulawesian Hipposideros dinops pelingensis and H. inexpectatus are compared. Indo-Australian
Myotis have been examined in some detail with a review of M. muricola and M.mystacinus;M.ater'\s
considered a distinct species and the status of M. australis Dobson, 1878 discussed; the large-footed
species M. horsfieldii, M. hasseltii and M. adversus are reviewed and keyed and a new subspecies, M.
adversus orientis is described from the New Hebrides. Certain Indo-Australian Pipistrellus are briefly
reviewed, notably P. tennis, and the second known specimen of P. societatis is reported. A new species
of Hesperoptenus, H. gaskelli, is described from Sulawesi. Indo-Australian Miniopterus are reviewed in
detail and recent revisionary work on this part of the genus is discussed with a number of proposed
changes in its classification: M. solomonensis Maeda, 1982 is considered a subspecies of M. australis;
M. macrocneme Revilliod, 1914 a subspecies of M. pusillus; M. oceanensis Maeda, 1982 a subspecies
of M. schreibersii\ M. macrodens Maeda, 1982 a subspecies of M. magnater\ M. propitristis Peterson,
1981 conspecific with M. tristis; the status of M. bismarckensis Maeda, 1982 is discussed. Occurrences
of Murina east of Wallace's Line are examined and further specimens of Tadarida (Chaerephon)
jobensis bregullae and Otomops papuensis are reported.

The following new records and range extensions are reported in this paper: Dobsonia viridis (?) viridis
from Sulawesi; Cynopterus brachyotis brachyotis from Burma and C. titthaecheileus and Macroglossus
sobrinus sobrinus from Krakatoa I; Megaerops (?) ecaudatus from northeastern India; Aethalops alecto
from Java; Taphozous melanopogon from Sulawesi; Megaderma spasma niasense from Siberut I;
Rhinolophus pusillus pusillus, Hipposideros ridleyi and Coelops robinsoni from Borneo; Myotis muri-
cola (?) browni from Sulawesi; M. ater from Siberut I, M. hasseltii continentis from Burma; Pipistrellus
vordermanni from Borneo; Miniopterus australis tibialis from Sulawesi, M. pusillus and M. magnater
macrodens from Sumatra; Murina suilla and M. tubinaris from Thailand, M. cyclotis peninsularis from
Borneo; Kerivoula papillosa (?) malayana from Sulawesi and Tadarida (Mormopterus) beccarii from
Australia.

INDO-AUSTRALIAN BATS 199

Acknowledgements

In preparing this paper I have been very conscious of the debt owed by the British Museum
(Natural History) to the collectors and donors who obtain specimens for its collections, often
under considerable difficulties. Without their efforts, dedication and generosity studies such
as this would not be feasible and my thanks are due to the many individuals whose names
appear in the lists of specimens in the text.

I am indebted also to my colleagues Dr W. Bergmans of the Instituut voor Taxonomische
Zoologie, Amsterdam, who supplied details of specimens of Chironax from Sulawesi, and
Dr C. J. Smeenk of the Rijksmuseum van Natuurlijke Historic, Leiden, for the loan of
specimens. Dr K. F. Koopman of the American Museum of Natural History, New York and
Professor J. D. Smith of California State University at Fullerton have offered valuable
concepts and opinions in personal discussion.

Finally it is necessary to acknowledge the contribution made by the Executive Committee
of 'Operation Drake', whose initiative and organisation enabled Mr Ben Gaskell to collect
the specimens that led directly to this paper.

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1951. Harpyionycteris, a genus of rare fruit bats. Am. Mus. Novit. No. 1 522 : 1-9,4 figs, 2 tabs.

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INDO-AUSTRALIAN BATS 207

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1905a. On some Australasian mammals. Ann. Mag. nat. Hist (7)16 : 422^38.

19056. A new genus and two new species of bats. Ann. Mag. nat. Hist. (7) 16 : 572-576.

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208 J. E. HILL

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Manuscript accepted for publication 25 October 1 982

Addendum

While this paper was in press a further contribution by Maeda et al. (1982) became available
in London. On the basis of specimens from Sabah these authors considered magnater a dis-
tinct species and suggested that the specimens in the British Museum (Natural History) from
Hong Kong (BM(NH) 65.1017 (sic), Pahang (BM(NH) 67.217) and Upper Burma (BM(NH)
32.1 1.1.3) might represent it. They also suggested that the forearm lengths given by Lekagul
& McNeely (1977) for schreibersii in Thailand and by Medway (1978) in Malaya indicated
that specimens referable to magnater might have been included. This study clearly antedates
the more detailed work by Maeda (1982).

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Sabah, East Malaysia. Zool. Mag. Tokyo 91 : 125-134, 5 figs.

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Peter Humphry Greenwood

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Vol45 No 4 pp 209-231

British Museum (Natural History)

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London SW7 5BD Issued 25 August 1983

On Macropleurodus, Chilotilapia (Teleostei,
Cichlidae), and the interrelationships of African
cichlid species flocks

Peter Humphry Greenwood /

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5HD

Contents

Synopsis ........... 209

Introduction .......... 209

Materials and abbreviations . . . . . . . . 211

Dental characters and their ontogeny in Chilotilapia rhoadesi and Macropleur-
odus bicolor . . . . . . . . . . 211

A comparison and assessment of other anatomical features in Macropleurodus

bicolor and Chilotilapia rhoadesi ....... 222

Discussion and conclusions . . . . . . . . 227

Acknowledgements ......... 230

References . 230

Synopsis

The monotypic cichlid genera Macropleurodus bicolor from Lake Victoria, and Chilotilapia rhoadesi
from Lake Malawi share an outstandingly derived dentition of a kind not recorded in any other mem-
bers of the family.

Since both taxa share, uniquely, such a derived character complex, it might be thought that they
also share a recent common ancestry not shared with other members of the Lake Malawi and Lake
Tanganyika cichlid species flocks. A detailed consideration of other anatomical and morphological fea-
tures indicates that the dental similarity is a homoplastic feature. However, it seems probable that both
taxa are members of a monophyletic lineage within the haplochromine cichlids, and that representa-
tives of this lineage occur in Lakes Malawi, Victoria and Tanganyika.

The features previously used to establish the monophyly of the Lake Malawi haplochromine species
are found to be unsuitable for that purpose. Other characters suggest, on the contrary, that the species
flocks of Lakes Victoria, Malawi and Tanganyika are each composed of several distinct lineages, and
that members of at least some lineages occur in more than one lake.

Introduction

Little attention has been given to the problem of interrelating the endemic cichlid faunas
of the major African lakes, Tanganyika, Malawi and Victoria (but see Stiassny, 1981). In
part this lack of enquiry can be attributed to the size of the problem, over 500 species are
involved, in part to a long established idea that the African cichlids can be separated into
haplochromine and tilapiine lineages, hence some similarities were simply ascribed to con-
vergent evolution, and in part to another long standing assumption, that within a lake all
the taxa in either lineage are most closely related to one another (hence the greater concern
with intra- rather than interlake relationships; Regan, 1920, 19220 & b; Trewavas, 1935
& 1949; Fryer & lies, 1972; Greenwood, 1974).
Recent investigations have cast considerable doubt on the concept of a simple

Bull. Br. Mus. not. Hist. (Zool.) 45 (4): 209-231 Issued 25 August 1983

209

210 P. H. GREENWOOD

haplochromine-tilapiine dichotomy in the phylogeny of African cichlids (see discussions in
Fryer & lies, 1972, and Greenwood, 1978). This changed viewpoint has, in a sense, compli-
cated the issue. Lake Tanganyika species once thought to be tilapiine stock and therefore
convergent mimics of haplochromine taxa in Lake Malawi (see Regan, 19220: 675; Fryer
& lies, 1972: 513-8) must now have their phyletic relationships reassessed. Such reassess-
ment also involves species from Lake Victoria because convergence has been invoked to
explain similarities between certain species of that lake with others from Malawi and
Tanganyika (Regan, \922b: 159). Further complications arise from evidence which suggests
that the postulated monophyly of the haplochromine and the tilapiine species in each lake
is questionable (Greenwood, 1980), a suggestion which also questions the previously assumed
close relationships of the endemic species in a particular lake.

In short, the whole problem of phyletic interrelationships needs critical reexamination in
the light of information gathered since Regan's and Trewavas' pioneering researches on these
fishes. It also requires, because different levels of relationship are involved, that the search
for sister-taxa be extended beyond the confines of a single lake, a possibility that was not
critically examined in the past. Then, as more recently (Regan, 1920; \922a & b; Trewavas,
1935; 1949; Greenwood, 1974), the tendency has been to identify anatomically generalized
'ancestral' species for each lake fauna, rather than to search for congruent and uniquely
shared derived features, synapomorphies, which might indicate common descent irrespective
of a taxon's present geographical locality.

On reflection, it would seem that previous surveys have too readily invoked 'convergence'
and 'parallelism' to explain similarity between members of different lake faunas, and that
the assumptions were made without hard evidence based on critical analysis. Likewise, the
mono- or at most, oligophyletic origin of a particular lake's fauna, or elements of its fauna
when 'haplochromine' and 'tilapiine' lineages are involved, have also been assumed without
sufficient evidence.

Unfortunately, the large number of species involved, coupled with the very disparate levels
of information available on the anatomy and ontogeny of those species, requires that a piece-
meal approach to the problems be adopted. The limitations and pitfalls of such an approach
are serious. But, considering the size of the problem and the need to overhaul the bases
for previous and sometimes inhibiting assumptions, even a piecemeal approach can be
profitable.

This present paper centres around two monotypic genera, Chilotilapia rhoadesi Blgr.,
1908, from Lake Malawi, and Macropleurodus bicolor (Blgr) 1906, from Lake Victoria.

Both share a very characteristic dental pattern and tooth form neither of which is found
in any other cichlids, either African or American (see Figs 1-10). Ontogenetically, too, they
are alike in the way the dental pattern develops, and in having a dental succession involving
at least two, and probably three distinct types of teeth. Not surprisingly, larger individuals
in both taxa have similar feeding habits. Both crush molluscs between massive jaw teeth
before ingesting the soft parts of their prey (Fryer & lies, 1972; Greenwood, 1956; 1974).

Intergeneric similarities extend beyond these dental and trophic features, although some
are certainly correlates. The resemblances include marked similarity in the morphology of
the jaws, the syncranial architecture, and in certain myological details.

The two species would almost certainly be placed in single genus if they occurred in the
same lake basin. Thus, there would seem to be an a priori case for considering the genera
to be closely related.

The ranking of Chilotilapia and Macropleurodus as distinct and not particularly closely
related genera has been influenced by three major considerations. First, the belief that the
haplochromine species of any one lake are more closely related to one another than to species
from other lakes (Regan, 19226: 165). Second, the idea that, in Lake Malawi, this relation-
ship is manifest in certain marker characters carried by most of its endemic haplochromines
including Chilotilapia (Regan, 19226: 1 58; Trewavas, 1949), and thirdly, a misunderstanding
by some workers (Trewavas, 1935: 68; see also Fryer & lies, 1974: 75) about the fundamental
nature of the definitive tooth form in Chilotilapia.

ON MACROPLEURODUS & CHILOTILAPIA 21 1

Regan (1922/r. 159) did, however, notice a basic similarity in the dental morphology and
pattern of Macropleurodus and Chilotilapia. But, since he considered the cichlids of Lakes
Victoria and Malawi to be but distantly related, and the two taxa to be members of different
lineages, it was inevitable that he should attribute their similarities to convergent evolution.

Trewavas (1935: 69-70), on the other hand, recognized that Chilotilapia was in fact a
member of Regan's haplochromine group, as is Macropleurodus, but made no comments
on the close similarity of its dentition with that of Macropleurodus. Because of its peculiar
dentition in relation to that of other endemic Malawi haplochromines, Trewavas maintained
the generic status of Chilotilapia and suggested on other characters (op. civ. 110), that its
nearest living relative was a species in that lake, Haplochromis euchilus (see p. 222 et seq.}.

Before considering these different viewpoints and the whole question of relationships, if
any, between the cichlids of Lakes Victoria, Malawi and Tanganyika, detailed comments
must be made on the dentition and other anatomical features of both Chilotilapia and
Macropleurodus.

Materials and abbreviations

The entire British Museum (Nat. Hist.) collection of cichlid dry skeletal material and alizarin
transparencies used in previous studies on the interrelationships of African haplochromines,
was utilized in the preparation of this paper (see Greenwood, 1980; 1983).
Additional dry skeletons were made from:

Macropleurodus bicolor. 1955.2.10:3; 1955.2.10:63-64; 1955.2.10:74-79 two

preparations.

Chilotilapia rhoadesi: 1935.6.14: 2121-6; 1935.6.14:2103-2112 three preparations.

Dissections on the jaw and branchial muscles were made on:

Macropleurodus bicolor: 1955.2.10:74-79 three specimens.

Chilotilapia rhoadesi: 1935.6.14:2121-6, and 1935.6.14:2102-2112.
Additional but partial dissections were also made on specimens from this lot.

Dissections of the branchial musculature only were made from:
Astatotilapia burtoni: 1950.4.1:2176-2200
Aulonocranus dewindti: 1960.9.30:4629-4641
Grammatotria lemairei: 1930.4.1:3758-3785

Dissections of the jaw muscles were made on:
Ptyochromis annectens: 1956.10.9:84-87

The use of the letters M, T, or V in parentheses after a generic or specific name indicates
the taxon is from, respectively, Lake Malawi, Tanganyika or Victoria. For the purposes of
this paper, the haplochromine fauna of Lake Victoria is taken to include that of Lakes
Edward and George, Kioga, and Kivu since several of the genera discussed occur in, but
only in, all five lakes (see Greenwood, 1980).

All abbreviations used in text figures are amplified in the relevant figure captions.

Dental characters, and their ontogeny, in Chilotilapia rhoadesi
and Macropleurodus bicolor

The definitive, and most characteristic, dentition of Chilotilapia (Fig. 3) is apparent only
in fishes over 120 mm standard length (SL), but even in smaller fishes the teeth are already
distinctive (Fig. 2).

The outer row teeth in both jaws of fishes throughout the size range available, 48-200 mm
SL, are mostly stout bicuspids. The minor cusp is small but distinct, the major cusp slightly
protracted, the neck nearly cylindrical in cross- section, and the crown somewhat incurved

212

P. H. GREENWOOD

Fig. 1 Macropleurodus bicolor. Scanning electron micrograph of anterolateral teeth in the outer
row of the left premaxilla, seen in labial view (67-5 xN.S.). From a specimen 58 mm standard
length.

buccally (Figs 2 & 6). With increasing body size the teeth gradually become stouter, the
crown less compressed except at its extreme occlusal edge, and the minor cusp is reduced
further, partly it would seem through wear, but also through its absolute reduction in teeth
of successive replacement generations.

A second, and essentially tricuspid, tooth type (Fig. 3) first appears in fishes of between
105 and 1 1 5 mm SL. Such teeth sometimes predominate in the outer tooth rows of specimens
> 140 mm SL; but even when this is so a number of bicuspids persist and may be the predo-
minant form even in fishes as large as 220 mm SL.

In the tricuspid teeth the major cusp is still clearly the largest one, and has virtually the
proportions and shape of that cusp in bicuspids but it is slightly more protracted. The two
minor cusps are situated at the base of the major one, i.e. are labial in position, are of equal
or subequal size, and unless greatly worn, have their rounded tips at the same level as the
occlusal surface of the major cusp. A shallow groove, most marked in unerupted and newly
erupted teeth, lies between the minor cusps and extends for a short distance onto the occlusal
surface of the major cusp.

It is these teeth which Trewavas (1935:67) described as having '. . . crowns shaped like
a grain of wheat. . .' I cannot, however, agree with her statement (loc. cit) that in adult fishes
all the outer teeth are of this kind. Certainly in some large fishes the majority of teeth is
of that kind, but in other and even larger specimens tricuspids are poorly represented. Their
number generally increases with the size of the fish; none is present in specimens less than
105mmSL.

As noted before, the essentially tricuspid nature of 'wheat grain' teeth is most obvious in

ON MACROPLEURODUS& CHILOTILAPIA

213

Fig. 2 Chilotilapia rhoadesi. Scanning electron micrograph of anterior outer row premaxillary
teeth situated on either side of the premaxillary symphysis, seen in labial view (42 x N.S.). From
a specimen 65 mm standard length.

newly erupted teeth. With wear the distinctiveness of the minor cusps is lost and the groove
between them almost obliterated. The tooth then assumed the appearance of a conical
unicuspid in which the crown is aligned almost at right angles to the neck. Since, ontogeneti-
cally, predecessors of the tricuspid teeth are bicuspids, and since bicuspids occur together
with the 'wheat grain' tricuspids, I cannot agree with Fryer & lies' (1972:75) description of
the dentition in Chilotilapia as 'basically conical'.

In addition to size-related changes in tooth form, there are similarly correlated changes
in tooth alignment (see Fig. 6).

Most specimens less than 95 mm SL have the outer row teeth of the premaxilla arranged
so that the occlusal surface of the major cusp is aligned in parallel with the long axis of the
bone, the tips of the crowns following one another in a gently curving arc (Fig. 6). Exception-
ally, in some fishes from the upper part of this size range, i.e. 80-90 mm SL, the lateral and
posterolateral teeth, rarely only the anterior ones, are implanted so that the crown is angled
across the long axis of the bone. The tip of the major cusp points medially, that of the minor
one labially.

Specimens of standard lengths greater than 95 mm have the crowns of all teeth in the outer
premaxillary row directed medially, but with the anterior teeth less angled than the others
(Fig. 6).

Changes in the implantation angle of the outer row teeth in the dentary begin at a smaller
body size than do those of the premaxilla. At a standard length of ca 70 mm the posterior
and posterolateral teeth are distinctly angled, although even at a standard length of 1 10 mm
the crowns of the anterior and anterolateral teeth are but slightly displaced towards the mid-

214

P. H. GREENWOOD

Fig. 3 Chilotilapia rhoadesi. Scanning electron micrograph of anterior and anterolateral teeth
(outer and some inner rows) of the right premaxilla, seen in oblique and somewhat posterior
occlusal view(l l-6x N.S.). From a specimen 151 mm standard length.

line. In somewhat larger individuals, however, all the outer teeth on the dentary are distinctly
angled, the anterior teeth now directed medially at a greater angle than their counterparts
on the premaxilla.

It has not been possible to identify the underlying mechanical causes of these shifts in tooth
alignment. That is to say, whether succeeding teeth erupt and are then fused to the bone
at a different angle from that of their predecessors, or whether teeth already fixed to the jaw
undergo a relative change of angle as a result of differential growth changes in the underlying
bone. Certain features suggest, however, that differential bone growth may be responsible.
For example, changes in the orientation of individual teeth are coordinate over whole sec-
tions of the jaw, yet tooth replacement is irregular and should thus lead to individual teeth

ON MACROPLEURODUS&CHILOTILAPIA

215

Fig. 4 Macropleurodm bicolor. Occlusal surface of anterior and anterolateral outer and inner
row teeth from the right premaxilla, seen in oblique and somewhat lateral view (17-Ox N.S.).
From a specimen 120 mm standard length.

in any one section being at different angles depending on their time of eruption. Also, those
areas of the jaw in which the most marked angling occurs are coincident with regions
of the bone which undergo the most marked differential growth, that is, expansion and
curvature. These regions are the lateral and posterolateral sections of the premaxilla, and
the anterior and anterolateral regions of the dentary's alveolar surface.

Teeth forming the inner tooth rows in both jaws also show marked ontogenetic changes
in shape and size. The smallest individuals examined, 48-57 mm SL, have mostly small tri-
cuspid teeth in these series, although there are usually a few and slightly larger teeth situated
posterolaterally in the premaxilla, where there is but a single inner tooth row. All the inner
teeth in fishes of this size group are of the common tricuspid type. That is, with two small
lateral cusps flanking a larger central cusp, and the whole crown gently recurved.

Fishes between 60 and 105 mm SL show a gradual replacement of preexisting premaxillary
teeth by stouter and coarser ones. Posterolaterally the replacements are bicuspids closely
resembling, but smaller than, the teeth of the outer row (see p. 2 1 1 above). Elsewhere in
the upper jaw the replacements are either stouter versions of the earlier tricuspids, or, more
commonly, stout and weakly tricuspid teeth in which the median cusp is so dominant that
the tooth is all but unicuspid.

The inner rows of dentary teeth, which are virtually confined to the anterior and antero-
lateral parts of the bone, undergo similar changes in specimens of the same size ranges. Some
'wheat grain' teeth, smaller versions of those in the outer row, may be present in the outer-
most series of the inner dentary rows; one exceptional fish has almost the entire outermost
row composed of such teeth.

As body growth proceeds beyond about the 100 mm SL mark, so are increasing numbers
of inner row premaxillary teeth replaced by bicuspids which, although smaller than those

P. H. GREENWOOD

Fig. 5 Chilotilapia rhoadesi. Occlusal surface of anterior and anterolateral outer and inner row
teeth on the left premaxilla, seen in oblique and somewhat lateral view (12-8 x N.S.). From
a specimen 190 mm standard length.

of the outer row, closely resemble them in gross morphology. Laterally and posterolaterally
a few 'wheat grain' teeth make their first appearance in the inner tooth rows of the pre-
maxilla. Only the innermost one or two rows retain small and weakly tricuspid teeth, or
an admixture of these with small unicuspids.

In the largest fishes examined, i.e. 160-220 mm SL, the lateral and posterolateral regions
of the premaxilla have at least 2 or 3 rows of noticeably enlarged teeth. These may be of
the 'wheat grain' type, bicuspids or an admixture of both types.

Noticeably enlarged inner teeth on the dentary of large fishes are confined to the anterior
and anterolateral part of the outermost series. However, all teeth in the succeeding rows,
except the innermost one and those in the lateral part of the outermost row, are relatively
enlarged. Like all inner teeth on this bone, they are slightly smaller than their counterparts
on the premaxilla.

The number of tooth rows in both jaws increases, somewhat irregularly, with increasing
body length. The smallest specimen examined, 48 mm SL, has 2, in places 3, rows in both
jaws, the largest specimen, 220 mm SL, has 5 rows. Specimens in the middle size range may
have 3 or 4 rows in each jaw, that number not always correlated with the fish's length.

At first glance, larger fishes seem to have more teeth laterally and posterolaterally in the
premaxilla. This, however, is illusory, a result of the teeth in that region of the bone being
slightly larger and less closely arranged than elsewhere. In this region too, there is less space
between the tips of the teeth, the interspaces between them being taken up by the enlarged
crowns. Thus, this area of the premaxillary occlusal surface is, effectively, more compact
than it is elsewhere on the bone.

ON MACROPLEURODUS&CHILOTILAPIA

217

Fig. 6 Chilotilapia rhoadesi. Premaxilla showing dentigerous surface in occlusal view; from
specimens 65-0 (top) 98-0 and 190-0 mm standard length respectively. Scale = 3 mm.

218 P. H. GREENWOOD

Tooth number in the outer row of the premaxilla ranges from 20-30 in fishes < 100 mm
SL, with fishes in the 48-70 mm range having, in general, 20-24 teeth, and from 24-40 in
fishes 105-220 mm SL.

Ontogenetic changes in jaw shape were mentioned earlier (p. 215). These involve, princi-
pally, the premaxilla. In the smallest skeleton examined, from a fish 65 mm SL, there is,
relative to the generalized condition, an obvious broadening of the premaxilla and its alveolar
surface over the lateral and posterolateral extent of the horizontal arms (Fig. 6). This broad-
ening becomes more pronounced with growth, and the widened area extends from its former
anterior position to encompass the posterior parts of the arm as well (Fig. 6). Correlated
with these changes there is an overall thickening and inflation of the horizontal arms and
a change in the outline shape of the entire premaxillary occlusal surface. The latter changes
from one which is broadly U-shaped, to one in which the base of the U is flattened and
extended laterally (see Fig. 6).

Alterations in the shape and proportions of the dentary are somewhat less pronounced,
and involve, mainly, a thickening and outward bulging of the anterior and anterolateral
alveolar surfaces and the bone immediately underlying them. There is also a change in
occlusal outline correlated with the changing outline of the premaxilla (Fig. 7).

The distinctive dentition, dental pattern and jaw bone morphology characterizing Chilo-
tilapia are repeated, with only minor differences, in Macropleurodus bicolor, as are their
ontogenies (cf. Figs 1, 4, 8 & 9, with 2, 3, 5, 6 & 7).

Departures from the Chilotilapia condition occurring in Macropleurodus are:

1 . The major cusp in all bicuspid outer row teeth, and in the definitive tricuspid 'wheat-
grain' teeth, is more protracted than in Chilotilapia (cf. Figs 3 & 5 with 4). As in that genus,
however, the major cusp becomes relatively stouter and almost cylindrical in the largest
teeth.

2. The two minor cusps in a 'wheat-grain' tricuspid are lower than in Chilotilapia, and
the groove between them is shallower. The minor cusps are often worn away in larger fishes.

3. The widened posterior and posterolateral region of the premaxillary alveolar surface
in larger fishes is, relatively speaking, slightly narrower in Macropleurodus (cf. Figs 6 & 8).

4. There are fewer inner tooth rows laterally and posterolaterally in the premaxilla of
Macropleurodus, usually 1 or 2, compared with 2-5 in Chilotilapia; the number in that
taxon, but not Macropleurodus, being partially correlated with the fish's size.

5. One or occasionally both arms of the premaxilla are bowed in the vertical plane (see
Greenwood, 1956:306-7; 1980:80-81).

6. The different tooth forms, especially the 'wheat-grain' type, first appear in individuals
of a smaller size. However, it should be recalled that individuals of Chilotilapia reach a larger
maximum adult size, at least 220 mm SL, than do those of Macropleurodus, 1 50 mm SL,
modal range 11 0-1 30 mm. That Chilotilapia has more rows of premaxillary inner teeth
(see p. 216) may also be related to the larger adult sizes attained in that genus.

The buccal larvae of Macropleurodus bicolor, at a total length of about 9-0 mm, have slen-
der and conical outer teeth in both jaws (Greenwood, 1956: 308). Similar teeth occur in the
buccal larvae of Astatotilapia macrops at a comparable developmental stage; adults of that
species have the generalized bicuspid tooth type found in many haplochromine cichlids
(see Greenwood, 1979 & 1980). Unfortunately no larval Chilotilapia are available for
examination.

Compared with intergeneric dental differences in other cichlids, those distinguishing
Chilotilapia and Macropleurodus are very slight. Indeed, they are of a kind which, on current
taxonomic practice, would not be rated at more than 'specific-differences' if the two taxa
were sympatric.

From a phylogenetic viewpoint these uniquely shared and derived dental features of Chilo-
tilapia and Macropleurodus would seem to indicate a recent common ancestry for the two
taxa. To test that hypothesis, which contradicts current views on the relationships of the
two genera, see p. 2 1 1 above, a search was made for characters that might refute it.

ON MACROPLEURODUS&.CHILOTILAPIA

219

Fig. 7 Chilotilapia rhoadesi. Lower jaw in ventral view; from specimens 65-0 (top) 98-0 and
190-0 mm standard length respectively. ret = retroarticular. Scale = 3 mm.

220

P. H. GREENWOOD

Fig. 8 Macropleurodus bicolor. Premaxilla, showing dentigerous surface in occlusal view; from
specimens 58-0 (top), 1 15-0 and 144-0 mm standard length respectively. Scale = 3 mm.

ON MACROPLEURODUS&CHILOTILAP1A

221

Fig. 9 Macropleurodus bicolor. Lower jaw, in ventral view; from specimens 58-0 (top), 1 1 5-0 and
144-0 mm standard length respectively. ret = retroarticular. Scale = 3 mm.

222

P. H. GREENWOOD

A comparison and assessment of other anatomical features in
Macropleurodus bicolor and Chilotilapia rhoadesi

The superficial characters, body form and meristic characters of the two taxa (see Boulenger,
1915; Regan, 1922#), apart from the densely and almost completely scaled caudal fin in
C. rhoadesi, and differences in coloration, do not provide any features of value for testing
a hypothesis of close relationship.

The caudal fin character, which Regan (\922a & b) and Trewavas (1949) used as evidence
for close interrelationship of the Malawi haplochromines and their distinctness from those
of Lake Victoria, will be discussed later (p. 228).

In their live coloration (see Greenwood, 1956:308-310 for M. bicolor; Axelrod &
Burgess, 1977:176-177 for C. rhoadesi}, and in the colour-patterns of preserved specimens
(Greenwood, 1956:308-310; Trewavas, 1935:110) the taxa are quite distinct. Trewavas
(1935:1 10) suggested that the colour pattern of C. rhoadesi might indicate its relationship
with another Malawian endemic, Haplochromis euchilus Trewavas. This feature will be dis-
cussed later, together with Regan's ideas on colour- patterns as indicators of a monophyletic
origin for the Malawi haplochromines (Regan, 1922a:686).

Macropleurodus bicolor exhibits a form of sex-limited polychromatism in which about 30
per cent of females have a piebald coloration (Greenwood, 1956:309). Similar polychroma-
tism occurs in several endemic species of Lake Victoria haplochromines, not all of which
appear to be closely related to one another or to Macropleurodus (Greenwood, 1974:53-54;

Fig. 10 Chilolilapia rhoadesi. A. Dentary and B. Premaxilla; both in right lateral view, ret

retroarticular. Scale = 3 mm.

ON MACROPLEURODUS &. CHILOTILAPIA

223

also 1980, for revised views on the relationships of the various species). Sex-limited poly-
chromatism, involving both similar and different colour patterns, has also been recorded in
several Malawi endemics (Fryer & lies, 1972) but not in Chilotilapia.

Adult male Macropleurodus have, on the anal fin, a number of scarlet 'egg-dummies' of
the true ocellar type (see Greenwood, 1979:274-5). As far as I can tell from colour photo-
graphs and from written descriptions, true ocellar markings are lacking in male Chilotilapia.
Instead, the anal fin carries a few yellowish spots resembling those on the dorsal and caudal
fins, a not uncommon condition amongst Malawi haplochromines, but one not recorded in
Lake Victoria.

B

Fig. 11 Lower jaw, in ventral view, of: left, Pamlabidochromis crassilabris (105mm standard
length); right. 'Haplochromis' euchilus (9\ mm standard length). ret = retroarticular. Scale = 3 mm.

At a deeper anatomical level, intergeneric comparisons were made of the jaw and branchial
musculature, and of the cranial and axial skeletons. The latter will be considered first.

There is a close similarity in the morphology of the dentary and premaxilla in Chilotilapia
and Macropleurodus, as there is in the form of the maxilla, a stout and somewhat fore-
shortened bone in both genera.

The dentary (Fig. 10) too is short, deep and stout, with a noticeably inflated region
surrounding its division into ascending, coronoid, and horizontal arms. The principal
intergeneric difference lies in the greater posterior extension of the outer tooth row in
Chilotilapia, the row continuing backwards to a point almost two-thirds of the way up the
coronoid process. In Macropleurodus the teeth do not extend, or extend for only a short
distance, onto the process.

The premaxilla (Fig. 10) is basically similar in both genera, although in Macropleurodus
the dentigerous arm is somewhat less inflated. Also, in that genus one or both dentigerous
arms are distinctly arched in the vertical plane (see Greenwood, 1956:306-7; 1980:83); this
apparently is an autapomorphy peculiar to Macropleurodus.

Certain derived features in the dentary of Chilotilapia and that of Macropleurodus are
shared with some Malawi taxa, ('Haplochromis' euchilus Trewavas [see first para. p. 228]
and various Labidochromis species), with members of the Paralabidochromis-Ptyochromis
lineage in Lake Victoria (see below), and with at least one species, Lobochilotes labiatus Blgr,
from Lake Tanganyika.

These derived features, discussed by Greenwood (1980:93-94), are concerned chiefly with
the foreshortened lower jaw, the marked bullation of the dentary in the area surrounding

224

P. H. GREENWOOD

its bifurcation (see above), the pronounced medial curvature of its lateral walls, and the way
in which the anterior part of the outer tooth row has a strong ventral dip in its line of insertion
on the bone.

In Lake Victoria, these characters are shared by members of a lineage which includes
Macropleurodus and the genera Paralabidochromis and Ptyochromis. These taxa, in turn,
are part of a larger group, the so-called Psammochromis-Macropleurodus superlineage
(Greenwood, 1980:93-94).

Resemblances in lower jaw form between Chilotilapia and Macropleurodus on the one
hand, and the various Malawi, Victoria and Tanganyika taxa on the other, are most apparent
when specimens of Chilotilapia and Macropleurodus less than 70 mm SL are used as a basis
for comparison, irrespective of specimen size in the other species involved. Within the Lake
Victoria lineage, overall resemblance with Chilotilapia and Macropleurodus is closest when
comparisons are made with members of the genus Paralabidochromis (Fig. 1 1), less so when
the species of Ptyochromis are used (see Greenwood, 1980:60-72; 92-94). A level of simi-
larity equal to that existing with Paralabidochromis is found when small Chilotilapia and
Macropleurodus are compared with specimens of various Labidochromis species, 'Haplo-
chromis' euchilus and Lobochilotes (Fig. 1 1).

With growth, the dentary in both Chilotilapia and Macropleurodus changes its outline
shape (see above, p. 218), and thus, superficially, comes to resemble less closely the dentary
in these other species. The other derived features remain unaltered, however.

In the ontogeny of their jaws, at least over the size range of available specimens, Chilotil-
apia and Macropleurodus could be described as passing first through a Paralabidochromis-
Labidochromis stage and then an '//'. euchilus-Lobochilotes stage. From that point onwards
the jaws in Chilotilapia and Macropleurodus begin to broaden anteriorly, taking on the wide-
based U shape characteristic of the adults in both taxa. Simultaneously the autapomorphic
features of each genus begin to be manifest (see above p. 223).

Neurocranial shape and architecture is similar in Chilotilapia and Macropleurodus, but
with the former having a taller and more expansive supraoccipital crest. The crest also con-

Fig. 12 Chilotilapia rhoadesi Neurocranium in left lateral view. Scales= 10 mm.

ON MA CROPLEUROD US & CHILO TILAPIA 225

tinues further forward than it does in Macropleurodus, with the result that the slope of its
anterior margin is continuous with the slope of the ethmovomerine skull region (Fig. 12);
in Macropleurodus this line is interrupted (cf. Fig. 12 with fig. 54 in Greenwood, 1980).

The Macropleurodus-Chilotilapia skull can be considered derived relative to that in
Astatotilapia species and in several cranially generalized haplochromine lineages (see
Greenwood, 1979; 1980). It approximates closely to the skull form in some Ptyochromis
species of Lake Victoria, e.g. P. annectens (Regan), P. sauvagei (Pfeffer), and P. granti (Blgr).
Ptyochromis, it will be recalled, is a member of the Psammochromis-Macropleurodus super-
lineage to which the genus Paralabidochromis, mentioned above in connection with lower
jaw morphology, also belongs. Like Ptyochromis, both Chilotilapia and Macropleurodus are
mollusc-eaters which 'shell' their prey orally (see Fryer & lies, 1972:75; Greenwood, 1956;
1974:69-72; 1980:60-67). Thus, jaw form and skull shape in all these species could well
be correlated characters forming part of a functional unit. The similarities in jaw morphology
existing between non-mollusc eating members of the Psammochromis-Macropleurodus
superlineage (i.e., in this context, the Paralabidochromis species), 'Haplochromis ' euchilus,
the Labidochromis species and Lobochilotes labiatus on the one hand, and Macropleurodus
and Chilotilapia on the other, are not susceptible to this explanation.

There has, in the past, been some confusion about the nature of the pharyngeal apophysis
in Chilotilapia (see Regan, 19220:676; Trewavas, 1935:69-70; Greenwood, 1978:316). The
growth series of Chilotilapia neurocrania now available help both to clarify that situation
and to weaken further the idea of a rigid division between the 'Haplochromis' and 'Tilapia'
apophyseal types (see discussion in Greenwood, 1978:321).

In a Chilotilapia skull of 16 mm neurocranial length, i.e. skull length measured from the
anterior tip of the vomer to the posterior face of the basioccipital condylar surface, the apo-
physis is structurally intermediate between the 'Tilapia' and the 'Trop hens' types defined
by Greenwood (1978:299-305). Its articulatory facet is formed from the parasphenoid with
a small contribution from the basioccipital. The latter, however, has a rounded surface and
thus does not seem to form part of the functional articulatory surface. The basal part of the
apophysis, i.e. the prootic and basioccipital, is rather inflated; when viewed from behind the
entire structure is low and broad.

At a neurocranial length of 23 mm, the apophysis is intermediate between the 'Tropheus'
and 'Haplochromis ' types. The basioccipital makes a small but definite contribution to the
articulatory surface (about a third of its area), and the apophyseal base is narrow and less
inflated than in the 16 mm skull.

In the largest skull examined, 39 mm neurocranial length, the basioccipital contribution
to the articular surface has increased somewhat so that the surface is clearly of the 'Haplo-
chromis' type (see also Greenwood, 1978:316), as is the narrow, near-vertically walled
apophyseal base formed from the prootic and basioccipital.

The apophysis in the smallest available skull of Macropleurodus, 20 mm neurocranial
length, has an articulatory surface of the 'Haplochromis 'type, although the base of the apo-
physis is somewhat inflated. In larger skulls, to 25 mm neurocranial length, the base narrows,
its walls become more nearly vertical, and there is an increase in the extent to which the
basioccipital contributes to the articulatory surface.

The palatopterygoid arch and the suspensorium are similar in Chilotilapia and Macro-
pleurodus, although both are relatively deeper in Macropleurodus. The palatopterygoid arch,
when compared with that in Astatotilapia and in many other haplochromines, is derived.
It is foreshortened, with the lateral face of the metapterygoid deeply concave over most of
its area. Like other derived features of the syncranial architecture, this form of palatoptery-
goid arch occurs in several Lake Victoria species belonging to the Ptyochromis-Paralab-
idochromis assemblage, as well as in Hoplotilapia retrodens, another member of the
Psammochromis-Macropleurodus superlineage; Greenwood, 1980:82. It is also found in cer-
tain Lake Malawi taxa, e.g. Labidochromis and Cyathochromis, and in some species from
Lake Tanganyika as well.

226 P. H. GREENWOOD

Judging from that pattern of distribution, it seems that this type of palatopterygoid arch
is one correlated with a foreshortened skull and the development of a powerful adductor
mandibulae muscle complex.

The total number of vertebrae, excluding the fused PU, + U, elements, has the same range,
29 or 30, mode 30, in both genera. However, in the samples examined, Chilotilapia has
modal counts of 14 abdominal and 16 caudal centra, compared with 13 and 17 centra
respectively in Macropleurodus.

Myologically, the jaw and branchial musculature, and the associated ligaments in both
Chilotilapia and Macropleurodus are similar, and are close to the presumed generalized con-
dition found in Astatotilapia elegans (see Anker, 1978; Stiassny, 1981; Greenwood, 1983).

The A, and A2 divisions of the adductor mandibulae muscle complex in Chilotilapia and
Macropleurodus are relatively stouter than those in A. elegans, but are comparable with the
condition found in Ptyochromis species, which taxa are also oral-shelling mollusc eaters
(Greenwood, 1974; 1980).

The A3 division of the adductor mandibulae in Chilotilapia is more slender than in
Macropleurodus, and its outline is bobbin-shaped rather than triangular.

Chilotilapia differs from Macropleurodus in having three rather than two divisions to the
ethmo-palatine ligament. Macropleurodus, like the majority of Lake Victoria haplochro-
mines, has the long posterior division of the ligament running from the palatine bone to
the posterior, i.e. orbital, face of the lateral ethmoid, and the shorter division running to
the anterior face of that bone. In Chilotilapia, a third division lies between the other two,
whose attachments are as in Macropleurodus, and attaches to the ventral face of the lateral
ethmoid.

There is little comparative information on these ligaments in cichlid fishes, and thus the
significance of their intergeneric differences cannot be assessed. A tripartite ethmo-palatine
ligament does occur in several Malawi taxa I examined, and in some Tanganyika species
as well. The condition is rarely encountered in Lake Victoria haplochromines although, and
perhaps significantly, it is present in one oral-sheller, Ptyochromis annectens (Regan).

Gill-arch musculature in Chilotilapia and Macropleurodus is of the generalized type (see
Liem, 1981; Greenwood, 1983). There are slight differences in the degree to which certain
muscles are developed in the two genera. The levatores externi, especially the fourth pair,
are larger in Chilotilapia, and the transversus dorsalis epibranchialis 2 is more tendinous
in Macropleurodus. Both genera have the first levator externi muscles well-developed, with
musculose rather than tendinous insertions.

Because relatively little information is available on the comparative myology of the
Cichlidae, the information derived from Chilotilapia and Macropleurodus provides no data
of positive value in a phylogenetic context. Negatively, however, since their myology com-
pares closely with the generalized condition, it is possible to say that no uniquely apomorphic
features are shared by the two genera.

Indeed, the same can also be said for the apomorphic osteological features discussed
earlier; all are found in other taxa from Lakes Malawi, Tanganyika and, especially, in species
from Lake Victoria. This rather skewed geographical pattern of apomorphy distribution
could, however, be artefactual since more is known about the osteology in species from Lake
Victoria than in those from the other lakes.

Be that as it may, the only derived features uniquely shared by Chilotilapia and Macro-
pleurodus are those relating to tooth morphology and dental patterns in adult and near-adult
fishes (see pp. 211-218).

Since these dental apomorphies apparently are not congruent with any other derived char-
acters uniquely shared by the two taxa, they cannot be considered homologous, i.e., true
synapomorphies, and must therefore be treated as homoplasies (see discussion in Patterson,
1982).

It remains to be determined whether these homoplasies resulted from parallel or from con-
vergent evolution, and also to assess what phylogenetic significance can be given to those

ON MACROPLEURODUS&. CHILOTILAPIA 227

apomorphic features shared by Chilotilapia, Macropleurodus and the other taxa mentioned
above.

Discussion and conclusions

Before considering these various issues, it is necessary to review current ideas on the origin
and relationships of cichlids from Lakes Malawi, Victoria and Tanganyika.

Prior to Regan's (1920; \922a & b) revisionary studies on these fishes, and his fundamental
reorganization of their generic classification (Regan, 1920), no meaningful speculations or
hypotheses could be generated about the origin and relationships of the lake flocks.

Regan based his supraspecific grouping of African cichlids on the nature of the neurocra-
nial apophysis on which the upper pharyngeal bones articulate, a scheme he first worked
out on the taxa from Lake Tanganyika (Regan, 1920). On the basis of this character, the
majority of Malawian and Victorian species were placed in Regan's ' Haplochromis 'divison.
The Lake Tanganyika species, on the other hand, were more equally shared between this
division and the so-called 'Tilapia' one. Since Regan considered these two divisions repre-
sented a fundamental dichotomy in the phylogeny of the Cichlidae, his views on the relation-
ship of taxa, endemic and otherwise, from the three lakes were influenced accordingly. In
particular this meant that resemblances between any member of the 'Tilapia' group and
another of the 'Haplochromis' group could only be interpreted as the result of convergent
evolution.

Trewavas (1935) followed Regan's scheme of supraspecific grouping when she revised the
Malawi cichlids, although she transferred four taxa, one of which was Chilotilapia, from the
' Tilapia' io the 'Haplochromis' division.

Recent research (Wickler, 1963; Fryer & lies, 1972; Liem & Stewart, 1976; Greenwood,
1978, 1983) strongly indicates that a basic phylogenetic division of African cichlids on apo-
physeal structure cannot be substantiated (see also observations on the apophysis in Chilo-
tilapia, p. 225 above).

Abandoning the pharyngeal apophysis as a critical character for determining major cichlid
groupings has, in fact, little effect on the problem ofChilotilapia-Macropleurodus interrela-
tionships since definitive apophyseal structure in both genera is of the 'Haplochromis 'type.
It does, however, affect problems concerned with the relationship of other closely similar
taxon pairs, or trios, shared between Lakes Malawi, Victoria and Tanganyika. As examples
of taxa whose phyletic status requires reexamination on the basis of other features, one can
cite the supposedly convergent Petrotilapia (M) and Petrochromis (T), Tropheus (T) and
Labeotropheus (M), and Lobochilotes (T), 'Haplochromis' euchilus (M) and Paralabidochro-
mis chilotes (V).

Regan (19226:158), working within the framework of his strictly dichotomous divisions
of the cichlids, believed . . . 'The indications are that the endemic Nyassa ( = Malawi) cichlids
have originated in the lake from about half-a-dozen ancestral forms'.

Later, Trewavas (1949) supported Regan's concept of an oligophyletic origin for these
fishes. After quoting Regan's statement reproduced above, she carried the argument further
by concluding, with regard to the endemic Malawian genera, that the '. . . balance of evidence
(is) in favour of Nyassa relationship for all of them'. Indeed, Trewavas (op. cit.) specifically
rejects Myer's (1936) suggested relationship of the Malawian genus Pseudotropheus with the
Tanganyika Simotes, because the two taxa have different types of pharyngeal apophyses.

Regan (\922a & b) did not discuss the possible interrelationships of his suggested 'half-a-
dozen' ancestral species. He did, however, consider that the numerous endemic 'Haplochro-
mis' species of Lake Malawi '. . . appear to form a natural group and have evidently evolved
in the lake from one or a few ancestral forms' (Regan, 1 9226: 1 58). This idea was based largely
on one feature which, as far as Regan could ascertain, was shared by most Malawi 'Haplo-
chromis' species, namely a caudal fin densely covered in small scales. Regan contrasts this
condition with that in the 'Haplochromis' species of Lake Victoria where the '. . . caudal
fin is scaly only on its basal half.

228 P. H. GREENWOOD

The genus Haplochromis has now been restricted to five species, none of which occurs
in Lake Malawi (Greenwood, 1980). No attempt has yet been made to subdivide the Malaw-
ian species formerly classified in Haplochromis; these are referred to here under the epithet
'Haplochromis '.

In support of his ideas about the Malawi 'Haplochromis ' species being a natural group,
Regan (op. cit.) drew attention to the fact that the caudal fin in these species is more or less
distinctly emarginate (rounded or truncate in Victoria species) and that a few distinctive types
of coloration, none of which occurs in Lake Victoria, are prevalent amongst the Malawi
fishes. Here again, Trewavas (1935; 1949) accepted Regan's arguments, and expanded the
scope of the supposed natural group defined by these characters to include the endemic hap-
lochromine genera of Lake Malawi. Thus, at least by implication, all the haplochromine
taxa of Malawi were assumed to be members of a natural group. From this it would follow,
again by implication, that the ancestral forms were more closely related to one another than
to any other taxa. Also, but neither Regan nor Trewavas is explicit on this point, the argu-
ment would assume the caudal fin characters to have evolved within the Lake because no
extant fluviatile species show these features. Trewavas (1949) does, however, note, but makes
no further comment on, the absence of a fully scaled caudal fin in 'Haplochromis ' callipterus
(Giinther), a species which she thought to be a likely representative of the '. . . ancestor of
many ('Haplochromis') species of Nyasa'.

The validity of the caudal fin characters as evidence for a natural group in Lake Malawi
is weakened by a fact which neither author mentions, that is, the occurrence of a densely
and completely, or almost completely, scaled caudal fin in several endemic Tanganyika
genera, some of which also have the fin forked or emarginate, e.g. Petrochromis, Simochro-
mis, Tropheus, Lobochilotes, Eretmodus, Lamprologus, and Cyphotilapia; personal obser-
vations based on a by no means comprehensive sample of the Lake Tanganyika species.
Interestingly, four of these Tanganyika genera have been associated in a 'convergent'
relationship with taxa from Lake Malawi (see above, p. 227).

Finally, and perhaps most significantly, it must be noted that the four endemic species
of Oreochromis (=Tilapia of Trewavas, 1935 & 1949) in Lake Malawi also have densely
and completely scaled caudal fins. None of these species is in any way closely related to
the Malawi haplochromines.

Such a character distribution pattern, and especially its inclusion of the endemic Malawi
Oreochromis species, would seem to invalidate the character's use as a means of identifying
the Lake Malawi haplochromines as a 'natural group'.

The shape of the caudal fin margin is also of doubtful validity in this context. To start
with, it shows a quite considerable range of variation within the Malawi taxa, in some ol
which the nature of the fin margin scarcely differs from the condition found in certain Lake
Victoria species. In that Lake, pace Regan, a rounded caudal margin is rare, the modal con-
dition ranging from subtruncate to truncate; a few species even have a slight but distinctly
emarginate fin (compare figures in Greenwood, 1982a with those in Regan, \922a). Truncate
and emarginate fins, as well as subtruncate and rounded ones, occur amongst the Lake Tan-
ganyika endemics, sometimes with more than one type present in a single genus.

Again, one is apparently faced with a character of doubtful phylogenetic validity. Not only
is this so for the reasons given above but, like the scaly caudal fin, because it is not congruent
with other and well-substantiated synapomorphies snared by the various taxa involved.

The absence of a fully scaled caudal fin amongst the Lake Victoria cichlids, contrasted
with the near universal occurrence of that trait in Malawi cichlids, and its presence among
some but not all Tanganyika taxa, nevertheless requires explanation. The answer, however,
like that explaining the occurrence of predominant but different kinds of caudal fin margins
in each of the different lakes, is unlikely to be provided by the data currently available.

Regan's (1922a:686) view that the prevalence of a few distinctive types of coloration
among the Malawi haplochromines corroborated the other indications of their evolution
'. . . within the lake from a single ancestral form . . .', can also be challenged. Granted, there

ON MACROPLEURODUS & CHILOTILAPIA 229

are distinctive colour patterns, i.e. striping, barring etc., and some distinctive types of overall
coloration not found in the fishes of Victoria or Tanganyika. But, since these features are
not shared by every member of the Malawi flock, they cannot be taken to indicate mono-
phyly for all the species of the Lake. Their value as indicators of relationship lies at a much
lower level of universality.

The absence of true anal ocelli in male Chilotilapia, and in most Malawi haplochromines,
contrasted with their presence in males of, apparently, all Victoria haplochromines, poses
an insoluble question with regard to phylogenetic histories (see Greenwood, 1979:274-276).
It is a character that has not yet received sufficient study to permit its evaluation in that
context. Whether true ocelli be a derived or a plesiomorphic feature, the fact that both
ocellar and non-ocellar anal markings are present in Malawi haplochromines would argue
against a monophyletic origin for the flock as a whole.

In short, I would suggest that none of the characters so far proposed in support of a mono-
or even oligophyletic origin for the Malawi haplochromines is valid for that purpose. Since,
despite an extensive search, I can find no valid characters to replace them, the case for a
monophyletic origin of the Malawi haplochromines must, for the moment, remain unsub-
stantiated.

Research on the haplochromines of Lake Victoria has also failed to reveal a single synapo-
morphy indicative of a monophyletic origin for those taxa either (Greenwood, 1979; 1980).
What is suggested, however, is the presence of several distinct monophyletic lineages whose
distributions include not only Lake Victoria but Lakes George, Edward and Kivu as well.
Each of these lineages is distinguished by, at best, two or three synapomorphies. In other
words, there is a low level of interlineage differentiation, with the residual, i.e. plesiomorphic,
characters being common to most haplochromine taxa whatever their geographical distri-
bution may be.

Perhaps significantly, it is with one of the Victoria lineages, the Paralabidochromis-
Ptyochromis-Macropleurodus complex within the Psammochromis-Macropleurodus super-
lineage, that Chilotilapia shares the greatest number of apomorphic characters (see
Greenwood, 1980:91-94; & p. 223 above). Furthermore, there are at least two taxa in Lake
Malawi, Labidochromis and 'Haplochromis' euchilus, and one from Lake Tanganyika,
Lobochilotes labiatus Blgr, which also share a number of these features. In none of these
latter taxa can I identify any synapomorphies shared with Lake Victoria lineages other
than the Paralabidochromis-Macropleurodus one, nor any which might suggest a relation-
ship, above the sister-species level, with taxa from, respectively, Lake Malawi or Lake
Tanganyika.

Parenthetically it should be noted that some of the oral features shared by these taxa
also occur in two other Lake Victoria lineages, namely Neochromis and Lipochromis (see
discussion in Greenwood, 1980: 52 & 91-94; also p. 223 above). It has been argued
(Greenwood, op. cit.) on the basis of derived dental features shared by members of the
Psammochromis-Macropleurodus superlineage, and others uniquely characterizing Neo-
chromis and Lipochromis, that the similarities in jaw morphology shared by these two taxa
with members of the former group were independently evolved. Since the derived dental
features of the Psammochromis-Macropleurodus group taxa are manifest in Labidochromis,
'H'. euchilus, Lobochilotes and Chilotilapia, the same argument, that is convergence,
can be applied to account for their similarities in jaw morphology with Neochromis and
Lipochromis.

Work in progress supports the suggestion of several derived characters being shared by
the supposedly 'convergent' Malawi-Tanganyika taxon pairs mentioned on p. 227, and thus
that they too may be more closely related than was previously thought. Here, however, it
has so far proved impossible to identify any association with a particular lineage from Lake
Victoria.

In view of these character distribution patterns, and because the hypothesized monophyle-
tic origin for each lake flock cannot be substantiated, consideration should be given to an

230 P. H. GREENWOOD

alternative hypothesis. Namely, that the flocks are of polyphyletic origin and that many of
the lineages within a flock occur in more than one lake. The pattern of apomorph character
distribution in, for example, Macropleurodus (V), Chilotilapia (M), 'Haplochromis' euchilus
(M) and Lobochilotes (T) would be explicable on that basis.

Since Lakes Victoria, Malawi and Tanganyika are of different ages (Greenwood, 19826)
and are geographically distant from one another, it is probable that the progenitors of the
lineage representatives in each lake would have been different but closely related species.
This evolutionary pattern would mean that, despite the ultimate monophyly of a lineage,
at the lowest hierarchical levels sister taxa would occur sympatrically. For example, the Lake
Victoria members in the lineage cited above would be more closely related to one another
than to species in the other lakes, and similarly for the taxa from Malawi and Tanganyika.

On the basis of that argument, the homoplastic dental resemblances between Chilotilapia
and Macropleurodus (see p. 218) are parallelisms, and the nearest relatives of the two taxa
should be found in Lakes Malawi and Victoria respectively.

As has already been suggested, the relationships of Macropleurodus are apparently with
Paralabidochromis and Ptyochromis, although, for the moment no finer resolution is possible
(see Greenwood, 1980:92).

The closest relatives of Chilotilapia would, in my view, seem to be Labidochromis and,
as Trewavas (1935:1 10) suggested, 'Haplochromis' euchilus. Again, any finer resolution must
await further research, as must the resolution of relationships for Lobochilotes labiatus, at
present seemingly the sole representative of the lineage in Lake Tanganyika.

If this scheme of relationships is accepted, then the generic status accorded to Chilotilapia
and Macropleurodus by earlier workers is justified, despite the close similarity of the two
taxa (see p. 210). I would also agree with their suggestion that the resemblances are homo-
plastic ones, but not, as was also suggested, the results of convergent evolution.

What I would argue against is the reasoning used by those workers for separating the taxa
at more than the species level, namely the level and nature of phyletic differences between
the two cichlid faunas to which the taxa belong. There would seem to be no grounds for
assuming that each lake flock is of monophyletic origin, or that each flock is, phylogenetically
speaking, a natural group. On the contrary, there seem to be grounds for thinking that each
flock is composed of several lineages, and that most of these lineages have a geographical
distribution which cuts across the present day lake boundaries. This, of course is not to deny
that some lineages may have evolved in, and are endemic to, a particular lake, or that some
lineages may be common to only Lakes Malawi and Tanganyika (see p. 227).

Regan's (19226:158) postulated half-a-dozen ancestors for the Malawi haplochromine
flock could, in essence but perhaps not magnitude, be nearer the mark than his and other
workers' ideas on the flock's monophyly (Regan, 1922<2 & 6; Trewavas, 1935 & 1949;
Greenwood, 1974), and may well represent the situation in other lakes as well (Greenwood,
1980). Only a great deal more, and critical, work on the cichlids of all three major lakes
will serve to test those suggestions.

Acknowledgements

I am most grateful to all my colleagues in the freshwater fish section of this Museum, and
those in other institutions, who have discussed and argued various points with me. In particu-
lar I thank Gordon Howes for all his help in so many ways, including his skill in producing
the figures. The scanning electron micrographs are the work of Susan Barnes and her collea-
gues in the S.E.M. Unit of this Museum to whom I am also greatly indebted.

References

Anker, G. Ch. 1978. The morphology of the head muscles of a generalized Haplochromis species:
H. elegans Trewavas, 1933 (Pisces, Cichlidae). Neth. J. Zool. 28: 234-271.

ON MA CROPLEUROD US & CHILD TILAPIA 23 1

Axelrod, H. & Burgess, W. E. 1 977. African cichlids of lakes Malawi and Tanganyika. T.F.H. Publica-
tions, New Jersey.

Boulenger, G. A. 1915. Catalogue of the fresh-water fishes of Africa. 3, London.

Fryer, G. & lies, T. D. 1972. The cichlid fishes of the Great Lakes of Africa. Their biology and evol-
ution. Oliver & Boyd, Edinburgh.

Greenwood, P. H. 1956. The monotypic genera of cichlid fishes in Lake Victoria. Bull. Br. Mus. nat.
Hist. (Zool.) 3: 295-333.

1974. Cichlid fishes of Lake Victoria, east Africa: the biology and evolution of a species flock.

Bull. Br. Mus. nat. Hist. (Zool.) Suppl. 6: 1-134.

1978. A review of the pharyngeal apophysis and its significance in the classification of African

cichlid fishes. Bull. Br. Mus. nat. Hist. (Zool.) 33: 297-323.

1979. Towards a phyletic classification of the 'genus' Haplochromis (Pisces, Cichlidae) and related

taxa. Part I. Bull. Br. Mus. nat. Hist. (Zool.) 35: 265-322.
1980. Towards a phyletic classification of the 'genus' Haplochromis (Pisces, Cichlidae) and related

taxa. Part II. Bull. Br. Mus. nat. Hist. (Zool.) 39: 1-101.

19820 The haplochromine fishes of the east African lakes. Kraus International Publications,

Miinchen & British Museum (Nat. Hist.), London.

1982& Species flocks and explosive evolution. In: Greenwood, P. H. & Forey, P. L. (Eds) Chance,

change and challenge — The evolving biosphere, pp. 61-74. Cambridge University Press, Cambridge
& British Museum (Nat. Hist.), London.

1983. The Ophthalmotilapia assemblage of cichlid fishes reconsidered. Bull. Br. Mus. nat. Hist.

(Zool.) 44: 249-290.
I Jem, K. F. 1981. A phyletic study of the Lake Tanganyika cichlid genera Asprotilapia, Ectodus,

Lestradea, Cunningtonia, Ophthalmochromis, and Ophthalmotilapia. Bull. Mus. comp. Zool. Harv.

149: 191-214.
& Stewart, D. J. 1976. Evolution of the scale-eating cichlid fishes of Lake Tanganyika: a generic

revision with a description of a new species. Bull. Mus. comp. Zool. Harv. 147: 319-350.
Myers, G. S. 1936. Report on the fishes collected by H. C. Raven in Lake Tanganyika in 1920. Proc.

U.S. natn. Mus. 84 no. 2998 : 1-15.
Patterson, C. 1982. Morphological characters and homology. In: Joysey, K. A. & Friday, A. E. (Eds)

Problems of phylogenetic reconstruction: 21-74. Systematics Association Special Volume, No. 21.

Academic Press.
Regan, C. T. 1920. The classification of the fishes of the family Cichlidae — I. The Tanganyika genera.

Ann. Mag. nat. Hist. (9) 5: 33-53.

1922a. The cichlid fishes of Lake Nyassa. Proc. zool. Soc. Lond. 1921: 675-727.

\922b. The cichlid fishes of Lake Victoria. Proc. zool. Soc. Lond. 1922: 157-191.

Stiassny, M. L. J. 1981. Phylogenetic versus convergent relationship between piscivorous cichlid fishes

from Lakes Malawi and Tanganyika. Bull. Br. Mus. nat. Hist. (Zool.) 40: 67-101.
Trewavas, E. 1935. A synopsis of the cichlid fishes of Lake Nyasa. Ann. Mag. nat. Hist. (10) 16:

65-118.

1949. The origin and evolution of the cichlid fishes of the great African lakes, with special refer-
ence to Lake Nyasa. 13th Int. Cong. Zool. 1948: 365-368.
Wickler, W. 1963. Zur {Classification der Cichlidae, am Beispiel der Gattungen Tropheus, Petrochro-

mis. Haplochromis und Hemihaplochromis n. gen. (Pisces, Perciformes). Senckenberg biol. 44:

83-96.

Manuscript accepted for publication 14 December 1982

East African Cichlid Fishes

The haplochromine fishes of the east African lakes
P. H. Greenwood, British Museum (Natural History)

May 198 1 , 840 pp, 3 plates, 350 figures Hardback Price £25.00

This volume brings together, for the first time, Dr Greenwood's various papers (some now
out of print) on the taxonomy and biology of the species of haplochromine fishes from Lake
Victoria, east Africa. Also reprinted are his papers on the haplochromines from Lakes
Nabugabo, George and Turkana (Rudolph), two papers dealing with the classification of the
genus Haplochromis, and a recent essay on the explosive evolution of cichlid fishes in Africa.
An index to the 200 species dealt with in this book (and their current generic placement),
and a general introduction to the evolutionary and taxonomic problems posed by these bio-
logically complex and fascinating fishes, are included.

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Problems in catfish anatomy and phylogeny exemplified by the Neotropical
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Miscellanea

Bats (Mammalia: Chiroptera) from Indo- Australia. By J. E. Hill.

On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the
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Contents

Page

Deuterammina (Lepidodeuterammind) subgen. nov., and a redescription ofRotalina
ochracea Williamson (Protozoa: Foraminiferida). By P. Bronnimann & J. E.
Whittaker 233

The freeliving marine nematode genus Sabatieria (Nematoda: Comesomatidae). I. Two

new species from Stonington Island, Antarctica. By H. M. Platt .... 239

New species of marine nematodes from Qingdao, China. By Z. N. Zhang & H. M. Platt 253

Echinoderms of the Rockall Trough and adjacent areas. I. Crinoidea, Asteroidea and
Ophiuroidea. By J. D. Gage, Margaret Pearson, Ailsa M. Clark, G. L. J. Paterson
& P. A. Tyler 263

2 9 SEP 1983

cm

Deuterammina (Lepidodeuterammina) subgen.
nov., and a redescription of Rotalina ochracea
Williamson (Protozoa: Foraminiferida)

P. Bronnimann

Laboratoire de Paleontologie, Universite de Geneve, 13, rue des Maraichers, 1211 Geneve
4, Switzerland

J. E. Whittaker

Department of Palaeontology, British Museum (Natural History), Cromwell Road, London
SW7 5BD, U.K.

I. A lectotype for Rotalina ochracea Williamson

Rotalina ochracea Williamson (1858) was based on three specimens obtained from 'sand
from Shetland'. Spiral and umbilical views, probably of the same specimen, were illustrated
(figs 112, 113) and the species was described in the classic monograph On the Recent Forami-
nifera of Great Britain as follows (p. 55). . .

Shell trochoid, depressed; slightly convex and smooth superiorly, correspondingly
concave inferiorly; with about two and a half convolutions; arcuate; filled with brown
animal matter, which is pale and translucent in the ultimate segments, but very dark
and opaque in those near the umbilicus. Septa smooth in one specimen, a little elevated
in another; broad, and of light ochraceous yellow, contrasting richly in the dark tint
of the segments. The connecting spiral has the same ochraceous line. Inferiorly the
segments of the last convolution extend nearly to the umbilicus, concealing the rest.
Segments depressed or concave. Septal lines arcuate, flexuose and very prominent.

Balkwill & Millett (1884) recognised the agglutinated nature of the test and transferred
it to Trochammina. For a history of Trochammina, see Hedley et al, 1964 : 418. Although
Trochammina ochracea (Williamson) is one of the most frequently reported species of the
genus (see for instance, Heron-Allen & Earland, 1915; Cushman, 1920; Rhumbler, 1938;
Hoglund, 1947; Todd & Low, 1961; Murray, 1971; Haynes, 1973; Lutze, 1974) its detailed
morphology is for the most part poorly known. Williamson's figured specimen is, unfortu-
nately, not in the remaining part of his collection in the British Museum (Natural History)
and must be considered lost. The other two specimens (syntypes) are, however, extant and
were figured by Hedley et al. (1964; figs 2, 3). In spite of this work, the species continues
to be largely misidentified and it is now thought necessary to designate a lectotype and
describe and illustrate the species afresh, using the benefits of scanning electron microscopy
(SEM). Particular attention is paid to details of the apertures, in the light of recent advances
in trochamminid taxonomy (see p. 236, below). The lectotype chosen bears the BMNH reg.
no. 1963.2.19.16 and is illustrated in Figs 1-8, 10, herein.

DESCRIPTION. (Lectotype). Test a concavo-convex, strongly compressed ('watch-glass'
shaped) dextral trochospire comprising some 26 chambers arranged in almost 3 whorls, with
8 chambers in the final whorl. In outline the test is oval and slightly lobate; the periphery
is rounded-compressed. Proloculus ellipsoidal 15xlOum in diameter. The chambers
increase rapidly in size, those of the last whorl almost doubling in tangential diameter.
Sutures curved and weakly depressed in the last whorl. Sutures on umbilical side strongly

Bull. Br. Mus. nat Hist. (Zool.) 45(5): 233-238 Issued 29 September 1983

233

234

P. BRONNIMANN & J. E. WHITTAKER

Figs 1-6 Deuterammina (Lepidodeuterammina) ochracea (Williamson) subgen. nov. Lectotype
(1963.2.19.16). Figs 1-4, spiral, edge, oblique-umbilical and umbilical views; 5, details of umbilical
depression seen obliquely; 6, early enrolment (note ellipsoidal proloculus). Figs 1-4, x 175; fig. 5,
x 500; fig. 6, x!250.

Chosen from Williamson's remaining two syntypes, labelled 'Sand from Shetland', off N. Scotland,
in shallow water. Previously figured (drawing) by Hedley et al., 1964, fig. 3 : 2.

curved. Umbilicus narrow, open, star-shaped; chambers overlapping with inflated proximal
margins and pointed umbilical extensions. Two sets of apertures per chamber: a rounded
primary extraumbilical aperture situated peripherally and visible only in the last chamber;
and a secondary elongate aperture in an umbilical-sutural, posteriorly directed position,
below the upturned edge of the pointed umbilical lobe. Wall thin, agglutinated, single-
layered and imperforate. Spiral side composed of a rather smooth mosaic of minute,
well-cemented, rock flakes and mineral grains. Umbilical side more coarsely granular.

DIMENSIONS. (Lectotype). Maximum and minimum diameter 240x200 um; thickness (axial
height) 45 um.

REMARKS. Spirally, the sutures are not well defined and the thick spiral suture shown by
Williamson (1858: fig. 1 12) has not been observed. On the umbilical side the 8 crescentic
chambers each overlap strongly posteriorly, while anteriorly, and distal to the umbilical lobe,
the chamber outline undergoes a humpback-like enlargement quite different from other
'watch-glass' shaped forms (compare figs 9 and 10). This widening seems to be an essential
feature of the taxon. The umbilical lobes are upturned (fig. 5) and may be compared to the
blades of a turbine; underneath them the posteriorly directed secondary apertures open into
the umbilical (axial) cavity. The narrow gap between the lobes of the last two chambers is

DEUTERAMMINA (LEPIDODEVTERAMMINA) SUBGEN. NOV.

235

10

Figs 7, 8 Deuterammina (Lepidodeuterammina) ochracea (Williamson) subgen. nov. Lectotype
(1 963. 2. 1 9. 1 6). Fig. 7, spiral side showing the rapid increase in chamber size during growth, x 175.
Fig. 8, umbilical side showing the shape of the penultimate chamber (wide hatching), and the star-
shaped umbilical depression (stippled); adventitious matter (crossed hatched). Primary aperture
arrowed, x 175. Drawings based on SEM photographs, figs 1, 4.

Figs 9, 10 Diagrams showing form of chamber and umbilical lobe in species of Deuterammina
(Lepidodeuterammina). Fig. 9, D. (Lepidodeuterammina) sp., illustrated as Trochammina squamata
astrifica Rhumbler, in Levy et a/., 1974, pi. 1, fig. 9. Fig. 10, D. (Lepidodeuterammina) ochracea
(Williamson). Lectotype (1963.2.19.16). Shape of penultimate chamber marked by wide hatching
and primary aperture by arrows, in both cases. Based on SEM photographs. Fig. 9, x215; fig. 10,
X250.

about 45 jim long and 7 urn wide. The rim of the primary aperture is slightly damaged and
is difficult to see in figs 2-4, its position is indicated in the drawings, figs 8, 10. The chambers
of the umbilical side have a tendency to collapse which suggests that the agglutinant is less
well cemented than on the spiral side.

II. The systematic position of Rot a I'm a ochracea Williamson

Williamson's original coloured illustrations (1858, figs 112, 113) are somewhat stylised, in
particular the spiral side (fig. 1 12) which shows an unusually strong spiral suture. Whatever
their shortcomings they served as a standard reference for the identification of many morpho-
logically related, but discrete, small 'watch-glass' shaped trochamminids referred to Tro-
chammina ochracea, or to the group of forms typified by T. ochracea. Of the many references
to this species, only those of Murray (1970, pi. 1, figs 2, 3; 1971 :37,pl. 11, figs 1-5), Haynes

236 P. BRONNIMANN & J. E. WHITTAKER

(1973 : 40, pi. 5, figs 15-18) and Levy et al. (1974, pi. 1, figs 1,3) are conspecific with the
lectotype. Rhumbler (1938) and Hoglund (1947) attempted to differentiate taxonomically
between these small 'watch-glass' shaped trochamminids, but it is only the advent of SEM
that has made it possible to see and illustrate the often minute umbilical and apertural
features significant for the classification of this difficult group.

As a result of comparative SEM work on material from the Collections of the R.R.S.
Discovery, Bronnimann (1976) erected the new genus Deuterammina, type species: Tro-
chammina glabra Heron-Allen & Earland, to accommodate all previously described species
of Trochammina which were found to possess two apertures per chamber: a primary, inter-
marginal and a secondary umbilical opening, the latter situated at the umbilical tip of the
chamber in an umbilical-sutural, posteriorly directed, position. By this definition, the overall
shape of the test was not taken into consideration at all, and therefore not only forms with
rounded peripheries, subglobular chambers and little compression of the test, but also those
with almost angular peripheries, strongly compressed chambers and a flattened test, were
placed in Deuterammina. However, in the course of subsequent work on shallow water deu-
teramminids (Bronnimann & Whittaker, in preparation) we are now of the opinion that the
overall form of the test has also to be taken into account, but only as a classificatory factor
of second order to the apertural characteristics. The overall shape of the deuteramminid shell
seems to be in direct relationship to the environment in which it lives. In our South Atlantic
and Antarctic material flattened 'watch-glass' shaped tests occur in shallow water and tests
with subglobular chambers, in relatively deeper water and in the deep sea. The former live
attached to a substrate represented by sand grains, seaweed, fragments of bivalves, etc., a
moveable biotope in the sense of Rhumbler, 1938. The latter probably live free on or in
the uppermost layers of the sediment. These observations are substantiated by findings else-
where (Bronnimann, 1978; Bronnimann & Maisonneuve, 1980; Lutze, 1974) and by records
of Trochammina ochracea in the collections of the BMNH from British waters. We therefore
propose to formally erect subgenera of Deuterammina based on the degree of compression
of the text, this subdivision reflecting the environment in which the animals lived. Deuteram-
mina (Deuterammina) is proposed for all forms typified by D. glabra (Heron-Allen &
Earland) and Deuterammina (Lepidodeuterammina) for all the forms with flattened tests
typified by T. ochracea (Williamson).

Genus DEUTERAMMINA Bronnimann, 1976,
emended Bronnimann & Whittaker, herein.

EMENDED GENERIC DEFINITION. Test trochospiral, free or attached. Wall agglutinated, single
layered, imperforate. Chambers subglobular or axially strongly compressed; without
inner structures. Two sets of apertures: primary opening interiomarginal, umbilical-
extraumbilical; secondary or supplementary opening at tip of chamber, umbilical-sutural,
posteriorly directed.

TYPE SPECIES. Trochammina glabra Heron-Allen & Earland, 1932. Recent. Antarctic waters,
outer shelf species.

Subgenus DEUTERAMMINA (DEUTERAMMINA) Bronnimann, 1976

SUBGENERIC DEFINITION. Test trochospiral, free, Chambers subglobular. Primary aperture
interiomarginal, umbilical-extraumbilical or extraumbilical; secondary or supplementary
aperture at umbilical tip of chamber, umbilical-sutural, posteriorly directed.

TYPE SPECIES. As genus.

Subgenus DEUTERAMMINA (LEPIDODEUTERAMMINA) subgen. nov.

SUBGENERIC DEFINITION. Test trochospiral, 'watch-glass' shaped, attached. Chambers axially
strongly compressed. Primary aperture interiomarginal, extraumbilical; secondary or sup-
plementary aperture at umbilical tip of chamber, umbilical-sutural, posteriorly directed.

DEUTERAMMINA (LEPIDODEUTERAMMINA) SUBGEN. NOV. 237

TYPE SPECIES. Rotalina ochracea Williamson, 1858. Recent. North Atlantic seaboard.
Shallow-water species.

NAME. From the Greek, lepido + deuter+ammina, meaning scale-like (shell) with two (aper-
tures) and made of 'sand'. The prefix of the new subgenus in particular refers to the very
thin, flattened, concavo-convex nature of the test which we have termed 'watch-glass'
shaped.

REMARKS. The primary interiomarginal aperture of Deuterammina (Deuterammina) glabra
is an elongate narrow umbilical-extraumbilical slit, whereas that of D. (Lepidodeuteram-
mina) (e.g. the type species and D. (L.) labiosa (Hoglund) (1947, pi. 15, fig 6a-c) is either
subperipheral in position or (e.g. a new species of D. (L.) from Brazil, Bronnimann, in
press) of Trochammina-type, also a small rounded opening, but this time about halfway
between the umbilical cavity and the periphery of the test. Slit-like primary interiomarginal
umbilical-extraumbilical or extraumbilical openings have so far not been observed in the
'watch-glass' shaped, attached deuteramminids which seem to be characterised by small,
rounded extraumbilical interiomarginal primary openings. These differences in the form and
position of the primary openings are possibly taxonomically of more weight than here
accorded but only future work will show whether or not they are of generic, rather than sub-
generic significance.

In order to avoid confusion with other attached trochamminids, viz. Asterotrochammina,
Remaneica and Rotaliammina, Deuterammina (Lepidodeuterammina) differs in the follow-
ing respects: Asterotrochammina and Remaneica have invaginations (or secondary septa);
on the umbilical radial sutures in the former, in the latter, on the spiral side as well. Rota-
liammina and D. (Lepidodeuterammina) do not have invaginations but differ from each
other in apertural characters: the latter has a double deuteramminid aperture, a primary,
extraumbilical interiomarginal aperture, and a secondary opening at the umbilical tip of the
chamber. Rotaliammina possesses only a single, umbilically (axially) directed aperture. For
further discussion of these genera see Seiglie & Bermudez, 1977, and Bronnimann, 1978.

Acknowledgements

The text has benefited from being read by Drs C. G. Adams and L. R. M. Cocks (BM(NH)).
The SEM photographs, taken by one of us (J.E.W.) were printed by the staff of the Electron
Microscope Unit, BM(NH). The senior author's research was in part funded by the Fonds
National Suisse.

References

Balkwill, F. P. & Millett, F. W. 1884. The foraminifera of Galway. J. Micr. & nat. ScL, London,

3: 19-28,78-90, pis. 1-4.
Bronnimann, P. 1976. Two new genera of Recent Trochamminidae (Foraminiferida). Archs Sci.

Geneve, 29:215-218.
1978. Recent benthonic foraminifera from Brasil. Morphology and ecology. Part 3. Notes on

Asterotrochammina Bermudez and Seiglie. Notes Lab. Paleont. Univ. Geneve, 3: 1-8, pis 1-4.
& Maisonneuve, E. 1980. Revision of the trochamminid genus Remaneica Rhiimbler, 1938 (Fora-

miniferida). Notes Lab. Paleont. Univ. Geneve, 6: 1-14, pis 1-7.

& Whittaker, J. E. (in preparation). A revision of the Trochamminidae (Protozoa: Foraminiferida)

of the Discovery Reports, described by Heron- Allen & Earland, 1932-36. Bull. Br. Mus. nat. Hist.,

London (Zool.).
Cushman, J. A. 1920. The Foraminifera of the Atlantic Ocean. Part 2 — Lituolidae. Bull. U.S. natn.

Mus., Washington, 104(2): 1-111, pis 1-18.
Haynes, J. R. 1973. Cardigan Bay Recent Foraminifera (Cruises of the R. V. Amur, 1962-1964). Bull.

Br. Mus. nat. Hist., London, (Zool.) Suppl. 4: 1-245, pis 1-33.

238 P. BRONNIMANN & J. E. WHITTAKER

Hedley, R. H., Hurdle, C. M. & Burden, I. D. J. 1964. Trochammina squamata Jones and Parker

(Foraminifera) with observations on some closely related species. N.Z. Jl Sci., Wellington,

7:417-426, figs 1-3.
Heron- Allen, E. & Earland, A. 1915. The Foraminifera of the Kerimba Archipelago (Portuguese East

Africa). Part 2. Trans. Zool. Soc. Land., 20: 543-794, pis 40-53.
Hoglund, H. 1947. Foraminifera in the Gullmar Fjord and Skagerak. Zool. Bidr. Upps., 26: 1-328,

pis 1-32.
Levy, A., Mathieu, R., Poignant, A., Rosset- Moulinier, M. & Rouvillois, A. 1974. Les representants

des Ammodiscacea, Lituolacea, Nodosariacea, Buliminacea (Foraminiferes)dans les sables des plages

de Dunkerque, Remarques sur les especes signalees par O. Terquem. Revue Micropaleont., Paris.

17:127-133, pis 1,2.
Lutz, G. F. 1974. Benthische Foraminiferen in Oberflachen-sedimenten des Persischen Golfes. Teil

1: Arten. Meteor ForschErgebn., Berlin & Stuttgart, (C) 17: 1-66, pis 1-11.

Murray, J. W. 1970. Foraminifers of the western approaches to the English Channel. Micro-
paleontology, New York, 16: 471-485, pis 1, 2.

1 97 1 . An Atlas of British Recent Foraminiferids. xiii + 244 pp., 96 pis. Heinemann Educ., London.

Parker, W. K. & Jones, T. R. 1859. On the nomenclature of the Foraminifera: Part 2, on the species

enumerated by Walker and Montagu. Ann. Mag. nat. Hist., London, (ser. 3) 4: 333-351.
Rhumbler, L. 1938. Foraminiferen aus dem Meeressand von Helgoland, gesammelt von A. Remane

(Kiel). Kieler Meeresforsch., 2: 157-222, figs 1-64.
Seiglie, G. A. & Bermiidez, P. J. 1977. Notes on the foraminiferal genera Rotaliammina, Astero-

trochammina and Remaneica. J.foramin. Res., Washington, 7: 297-303, pis 1-3.
Todd, R. & Low, D. 1 96 1 . Near-shore foraminifera of Martha's Vineyard Island. Massachusetts. Contr.

Cushman Fdnforamin. Res., Ithaca, N.Y., 12: 5-21, pis 1, 2.
Williamson, C. W. 1 858. On the Recent Foraminifera of Great Britain. xx+ 107 pp., 7 pis. Ray Society,

London.

Manuscript accepted for publication 16 November 1982

The freeliving marine nematode genus Sabatieria
(Nematoda: Comesomatidae). I. Two new species
from Stonington Island, Antarctica

H. M. Platt

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Introduction

As a prelude to an eventual review of the genus Sabatieria, species from various geographical
locations and differing habitats are being examined in detail. This first paper describes two
new species from the constantly near-freezing Antarctic waters. The specimens were
obtained by SCUBA divers on 30 March 1974 from a small inlet, Back Bay, in the vicinity
of the former British Antarctic Survey base at Stonington Island; part of a series of collections
detailed elsewhere (Platt, 1979). The substrate was fine sand and stones underlayed at about
2 cm with gravel which prevented coring beneath this depth. The methodology employed
for studying the organisms and the abbreviations used in their description have been fully
described elsewhere (Platt, 1982).

Systematic descriptions

Sabatieria kelletti sp. nov.

(Figs 1-3)

MATERIAL STUDIED. Holotype: rfl BM(NH) 1982.5.26. Allotype: 9! BM(NH) 1982.5.32.
Paratypes: five males BM(NH) 1982.5.27-31; five juveniles BM(NH) 1982.5.33-37.

LOCALITY. 15m depth, Back Bay, Stonington Island, Antarctica. Lat. 68° 12-4' S, long.
66°59-5' W.

MEASUREMENTS (Table 1)

Holotype rf: — 350 M 2845 —p.,. ~, , 0 0 ,~ A c ,~

3095 um; a = 36; b = 8-8; c= 12-4; S = 63 urn

19 74 87 61

Allotype 9: - 330 1550 3035

— I / ^ s 3 _• _

DESCRIPTION. The body narrows suddenly in the region just posterior to the amphid: h.d.
26-33% of the posterior oesophagus c.d. Cuticle punctated: in the lateral field the dots are
larger and more irregularly arranged than medially, especially in the oseophagus and tail
regions (Fig. 3c, e). The medial dots are more or less arranged in transverse rows (Fig. 3d).
In the cylindrical portion of the tail the dots are present but are only very tiny. Somatic
setae in four longitudinal files, one on each side of the lateral field: 5-7 um long and spaced
50-130 um apart but closer together at the ends than in the middle of the body. In adults
there is a subventral row of three more closely spaced cervical setae (Figs Ic, d & 3a). In
stage-4 juveniles (Jl-4) there are ony two setae in this position (Fig. li) and no such setae
were observed in what is probably a stage-2 juvenile (J5). There are additional 7-10 um dor-
sal and ventral caudal setae in the males (Figs If, j & 2f). The tail tip has three terminal
setae, one dorsal and two subventral. Rl sensilla papilliform. R2 sensilla short but clearly
setiform. R3 sensilla in adults 42-47% h.d.: the same relative length in juveniles. Amphids

Bull. Br. Mus. nat. Hist. (Zool.) 45(5): 239-252 Issued 29 September 1983

240

H. M. PLATT

Fig. 1 Sabatieria kelletti: (a) rfl whole body; (b) d\ head; (c) d2 anterior region; (d) rf4 anterior region;
(e) cfl lateral cuticle pattern mid-body; (f) d4 tail region; (g) c?6 ventral caudal papilla; (h) 9! tail;
(i) Jl anterior oesophagus region; (j) cfl posterior region; (k) cfl copulatory apparatus. Bar scales:
a = 200 urn; j= 100 um; f=50 um; c,d, h = 30 um; i = 20(xm;b, e, g, k= lOum. Stars in c, dandi indi-
cate ventral side.

NEMATODE GENUS SABA TIER1A

241

Fig. 2 Sabatieria kelletti: (a) d 1 whole body; (b) amphid and R3 sensilla; (c) caudal cuticle punctations
and position of ventral papilla arrowed; (d) posterior region showing precloacal supplements, the
anteriormost (24th) is arrowed; (e) detail of supplements; (f) d1 tail, postcloacal papilla arrowed.

describe 2± to almost 3 turns; 60-70% c.d. in males; 40% c.d. in female; 45-60% in juveniles.
Anterior part of buccal cavity cup-shaped, posterior part not expanded. Oesophagus widens
in posterior 20% but is not set-off (Fig. 3b). Nerve ring at 52-54% of oesophagus length.
Excretory pore at 61-67% of oesophagus length with a conspicuous ampulla (Fig. 3b). Tail
conico-cylindrical; cylindrical part 50-60% of total with a distinct but not greatly swollen
tip; 3-6^4-3 a.b.d. in males, 5-0 a.b.d. in females.

242

H. M. PLATT

Fig. 3 Sabatieria kelletti: (a) anterior region showing row of three subventral cervical setae; (b) pos-
terior oesophagus region showing excretory pore (arrowed) and posterior expansion of the oesopha-
gus; (c) lateral cuticle pattern in post-amphid region; (d) lateral cuticle pattern mid-body; (e) lateral
cuticle pattern on tail.

NEMATODE GENUS SABATIERIA

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Spicules equal, curved; 1-0-1-2 a.b.d. as chord; 1-2-1-4 a.b.d. (68-72 urn) as arc.
Gubernaculum with strongly curved apophyses which proximally are caudally directed.
Ventral precloacal spine and 21-27 conspicuous precloacal supplements which extend
1 50-250 urn anterior to the cloaca measured along the ventral side of the body (measure-
ment a) or 225-320 urn (9-1 1% of total body length) as measured along the mid-line of the
body (measurement ft). There is a very slight but distinct swelling resembling a flattened
papilla between the conical and cylindrical parts of the male tail (Figs 1 f, g, j & 2c, 0
Two opposed testes: anterior to the left, posterior to the right of the gut.

Ovaries opposed and outstretched. Anterior ovary to the right, posterior to the left of the
gut.

DIFFERENTIAL DIAGNOSIS. Sabatieria kelletti sp. nov. belongs to the celtica-group in terms
of general body shape, cuticle punctation pattern, R3 sensilla length, amphid form and size,
number of precloacal supplements and general tail shape. Indeed, the new species seems to
be most similar to S. celtica Southern, 1914 itself, as redescribed by Lorenzen (1972), from
which it differs in the range of supplement number (21-27 vs 15-22), more conspicuously
narrowed and relatively smaller head (h.d. as percentage of posterior oesophagus c.d. 26-33%
vs 33-42%), relatively shorter cephalic setae (0-4-0-5 h.d. vs 0-6-1-2 h.d.) and the presence
in the male of a ventral caudal swelling or papilla.

ETYMOLOGY. The species is named after Mr Brian Kellett, my co-diver during the Antarctic
peninsula collecting trip.

REMARKS. Because of the differing amounts of curvature in the posterior region of the males,
measurement of the distance from cloaca to anteriormost supplement is subject to great vari-
ation (Table 1) even when an attempt is made to take into account the differing number
of supplements among the specimens. For example, the ratio of the a-measurement to the
number of supplements has a mean of 8-2 and a CV of 16-7%. However, this variation is
less when the /^-measurement is used, which should approximate the a of a straight worm:
the ratio of ft to supplement number has a mean of 1 1-8 and a CV of 7-8%. It would seem
therefore that ft would be the better measurement to report, although in view of the fact
that ft can be considerably larger than a, 1 -2 to 1 -7 times larger in S. kelletti, the exact method
of measurement should also be reported.

Sabatieria lawsi sp. nov.
(Figs 4-6)

MATERIAL STUDIED. Holotype: rfl BM(NH) 1982.6.38. Allotype: 9! BM(NH) 1982.6.44.
Paratypes: five males BM(NH) 1982.6.39-43; five females BM(NH) 1982.6.44^8; nineteen
juveniles BM(NH) 1982.6.49-68.

LOCALITY. See 5. kelletti.

MEASUREMENTS (Tables 2 and 3)

Holotype rf:— 233 M 2250 .... .. ,., , . c __

— — 2400 urn; a = 34; b=10-3; c=16-2; S = 73 urn

Allotype 9:— 233 1180 2395

15 52 83 50- 2595 um;a = 31;b=l 1-1; c= 12-9; V = 45%

Fig. 4 Sabatieria lawsi: (a) c?l whole body; (b) rfl head; (c) d 1 anterior region; (d) rfl posterior region;
(e) lateral cuticle pattern mid-body; (0 rfl copulatory apparatus; (g) cf copulatory apparatus, ventral
view; (h) en face view; (i) buccal cavity lumen shape level with R3 sensilla; (j) oesophagus lumen
shape just posterior to amphids. Bar scales: a = 200 urn; d = 50um; c = 30um; others = 10 urn. Star
in h indicates the ventral side.

NEMATODE GENUS SABATIERIA

245

246

H. M. PLATT

NEMATODE GENUS SABATIERIA 247

DESCRIPTION. The anterior region gradually narrows from the end of the oesophagus to the
head: h.d. 26-35% of the posterior oesophagus c.d. The anterior end is typically bent dorsally
in formalin-fixed specimens (Fig. 4c).. Cuticle punctated. Lateral differentiation of larger,
more irregularly arranged dots begins immediately posterior to the amphids and is especially
conspicuous in the oesophageal and caudal regions. In the middle region of the body the
lateral dots are not so markedly larger than the medial ones but they are still more irregularly
arranged (Fig. 6c). The decrease in the size of the lateral dots posterior to the oesophagus
region occurs gradually. In the anterior oesophagus, the dots appear somewhat stellate when
the microscope is focused at the surface (Fig. 6b). The dots are also conspicuously large
ventrally in the oesophagus region (Fig. 4b, c). Short but stout somatic setae are located at
the edges of the lateral field, relatively more numerous in the oesophagus and tail regions
where there are also some subdorsal and subventral somatic setae: no regular repeated or
distinctive patterns could be made out in the distribution of somatic sensilla. Rl sensilla
papilliform but conspicuous. R2 sensilla very short but setiform, about 1-5 urn (Fig. 6a). R3
sensilla 32^4% h.d.: similar relative length in juveniles. Amphids in adults describe almost
3 turns (Fig. 6a); similar in size in both sexes, 48-66% c.d. Buccal cavity cup-shaped
anteriorly, posterior part tubular and tri-radiate (Fig. 4i) but not expanded. Oesophagus
widens posteriorly: marginal tubes at the ends of the radii can be seen in optical cross-section
(Fig. 4j). Nerve ring at 50-54% of oesophagus length. Excretory pore at 53-66% of oesopha-
gus length. Tail conico-cylindrical with the cylindrical part about 33% of the total which
shows little variation (Fig. 5): 3-1-3-8 a.b.d. in males, 3-8-4-6 a.b.d. in females. There
appear to be separate subterminal openings to the caudal glands at the tail tip (Fig. 6f, g).

Spicules equal, curved, relatively long and slender with a proximal median partition
extending up to a third of the total spicule length (Fig. 4f): 1 -6-1 -7 a.b.d. (chord). Gubernacu-
lum apophyses long, straight and dorso-caudally directed. Ventral view of cloacal opening
shows it to be a double-bowed transverse slit (Fig. 4g). Ventral precloacal spine about 1 -5 um
(Fig. 6h). 17 inconspicuous tubular precloacal supplements extending 355-385 um from
cloaca (ft) or 15-17% of total body length. The posterior 4 or 5 supplements are situated
closer together than the remainder. Two opposed testes; anterior to left, posterior to right
of gut.

Ovaries opposed, outstretched. Vulva conspicuous; V = 45^9%. Eggs round. Receptaculum
seminis of mature females contain large hollow sperm (Fig. 6d, e).

DIFFERENTIAL DIAGNOSIS. Sabatieria lawsi sp. nov. belongs to the celtica- group of species
in terms of general body shape, cuticle punctation pattern, R3 sensilla length, amphid form
and size, number of precloacal supplements and general tail shape. In several respects, the
new species appears to be similar to S. heterura (Cobb, 1898), a species originally found
in Australia and subsequently redescribed by Wieser (1954) on specimens from Chile. How-
ever, the species may be distinguished most easily by the detailed structure of the spicules
and gubernaculum.

ETYMOLOGY. The species is named after Dr R. M. Laws, Director of the British Antarctic
Survey.

REMARKS. Sabatieria lawsi would appear to be the more common of the sympatric Sabatieria
species: they could be readily distinguished even in the juvenile stages by the markedly differ-
ent shape of the oesophagus region.

The hollow sperm in the females were similar to those previously reported from Sabatieria
by Riemann (1983).

Fig. 5 Sabatieria lawsi: (a-e) tails of c?5, rf6, rf4, cf2 and cf3, respectively; (f-j) tails of 95, 9 U ?4,
98 and 92 respectively; (k-1) tails of fourth-stage Jl and J4; (m) tail of third-stage J7; (n) tail of
second-stage J15. Bar scale = 50 urn for all figures.

248

H. M. PLATT

Fig. 6 Sabatieria lawsi: (a) anterior region showing amphid, R3 sensilla and lateral R2 sensilla;
(b) lateral cuticle punctations in post-amphid region; (c) lateral cuticle pattern mid-body; (d) mid-
body region of 9 showing vulva and two eggs, box labelled e shows position of following figure; (e)
large hollow sperm in receptaculum seminis, see Fig. 6d; (f-g) tail tips showing exit pores of caudal
glands, arrowed; (h) precloacal region showing spine (solid arrow) and first two supplements (hollow
arrows).

NEMATODE GENUS SABATIERIA

249

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Fig. 7 Relationships between body length and oesophagus length, and body length and maximum
body diameter for 5. lawsi: curves fitted by eye.

Discussion

Although several species of Sabatieria have previously been reported from Antarctic
waters (as denned in Platt, 1979) only one can be considered valid: Sabatieria mawsoni
Wieser, 1954 which was originally described from Commonwealth Bay, Adelie Land, as
Parasabatieria antarctica by Cobb (1930). Cobb provided a small and rather inadequate
figure but described it as having six supplements, which clearly separates it from the two
species described here. Sabatieria antarctica Cobb, 1914, not Parasabatieria antarctica, was
described from a juvenile and was considered doubtful by Wieser (1954). Allgen's (1929)

N EM ATODE GENUS SA BATIERIA 251

S. australis was also described from juveniles and is here considered dubious as are several
other species either described, S. curvispiculum, S. dorylaimopsis and S. heterospiculum: all
from South Georgia and unfortunately too poorly described ever to be recognised again with
any degree of certainty, or reported from Antarctic waters by Allgen (1953, 1959).

Measurements of S. lawsi juveniles and adults suggest that during post-embryonic growth,
or more strictly during post-juvenile-stage-4 growth, most of the anatomical features other
than those of the reproductive system remain in similar proportion. There is some indication
from the 'a' ratio that the worms become slightly thinner relatively (Tables 2 & 3) although
a straight line can be fitted to the points of the relationship between body length and width
(Fig. 7); that this line does not pass through the origin suggests a curved line might be more
appropriate. Geraert (19780, \919a) also concluded that various freeliving soil forms
became relatively thinner during post-embryological growth but became thicker again during
maturation.

A less equivocal change can be seen in the relative length of the oesophagus, with the 'b'
ratio increasing (i.e. the oesophagus becoming relatively shorter) with no overlap in the
ranges for the various developmental stages, assuming these have been correctly identified
(Tables 2 and 3). Geraert (19780, 19786, 19790) found that for freeliving soil forms there
were many different kinds of oesophageal growth patterns: the shape of the curve in
Fig. 7 suggests that for S. lawsi the oesophagus continues to grow in all stages but progress-
ively slows down, as Geraert found occurred in rhabditids.

In his study of relative tail length, Geraert (19796) concluded that the 'c' ratio was of minor
importance and that tail length varied independantly and was taxonomically unimportant.
However, the relatively low CV for these measurements in marine nematodes together with
low infraspecific variability in shape (Fig. 5) indicates that at least for these comesomatids,
characters of the tail are of greater importance than in the soil forms studied by Geraert,
possibly because in marine nematodes the tail plays a greater functional role than in soil
forms.

A final point of interest to emerge from this study is the general range of infraspecific vari-
ation found in these organisms in view of what would appear to be very stable and cold
environmental temperatures. The CV for most characters was under 10% (Tables 1-3), simi-
lar to but not any less than the range previously reported for Sabatieria species from warmer
waters (Jensen & Gerlach, 1977; Jensen, 19790, 19796). The maximum body diameter CV
can exceed 10%, especially in females (Table 2) but this, as pointed out by Jensen (19790)
is influenced by the state of gonad development.

Acknowledgements

The specimens described were collected whilst employed by the British Antarctic Survey:
permission to continue the analysis of this material is acknowledged. Thanks are due to the
Master and crew of the RRS Bransfield for logistic support for the diving operations and
especially to my co-diver Mr Brian Kellett for invaluable companionship both above and
below water.

References

Allgen, C. 1953. Uber eine bemerkenswerte neue Siidsee-Art der Nematodengattung Sabatieria De
Rouville, S. heterospiculum von Siid-Georgien. K. nor. Vidensk. Selsk, Forh. 26: 4-6.

1959. Freeliving marine nematodes. Further zool. Results Swed. Antarct. Exped. 5 (2): 1-293.

Cobb, N. A. 1898. Australian free-living marine nematodes. Proc. Linn. Soc. N. S. W. 23: 383-407.
Geraert, E. 1978<a. On growth and form in Nematoda: oesophagus and body-width in Tylenchida.

Nematologica 24: 137-158.

19786. On growth and form in nematodes: II. Oesophagus and body width in Dorylaimida.

Nematologica 24: 347-360.

252 H. M. PLATT

1979a. Growth and form in nematodes: III. Comparison of oesophagus and body shape.

Nematologica 25: 1-21.

19796. Growth and form in nematodes: IV. Tail length and vulva position. Nematologica 25:

439^44.

Jensen, P. 1979a. Nematodes from the brackish waters of the southern archipelago of Finland. Benthic

species. Ann. Zool. Fennici 16: 151-168.

19796. Revision of Comesomatidae (Nematoda). Zoologica Scr. 8: 81-105.

& Gerlach, S. A. 1977. Three new Nematoda-Comesomatidae from Bermuda. Ophelia 16: 59-76.

Lorenzen, S. 1972. Die Nematoden fauna im Verklappungsgebiet fur Industrieabwasser nordwestlich

von Helgoland. II. Desmodorida und Chromadorida. Zool. Anz. 187: 283-302.
Platt, H. M. 1979. Freeliving marine nematodes of Antarctica. A current appraisal. Annales Soc. r.

Zool. Belg. 108:93-101.
1982. Revision of the Ethmolaimidae (Nematoda: Chromadorida). Bull. Br. Mus. nat. Hist. (Zool.)

43(4): 185-252.
Riemann, F. 1983. On spermatozoa in aquatic nematodes. In A. R. Stone, H. M. Platt & L. Khalil

(Eds), Concepts in nemalode systematics. pp. 85-93. Academic Press, London.
Southern, R. 1914. Nemathelmia, Kinorhyncha and Chaetognatha (Clare Island Survey, part 54). Proc.

R. Ir. Acad. 31: 1-80.
Wieser, W. 1954. Free-living marine nematodes II. Chromadoroidea. Acta Univ. lund(N.F. 2) 50 (16):

1-148.

Manuscript accepted for publication 15 November 1982

New species of marine nematodes from Qingdao,
China

Z. N. Zhang

Shandong College of Oceanography, Qingdao, Shandong Province, China

H. M. Platt

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Introduction

This paper is the first taxonomic report on freeliving marine nematodes from China. The
three new oncholaimid species described here were extracted from intertidal sand samples
taken from beaches adjacent to the city of Qingdao in summer, 1981.

Material and methods

The specimens came from two intertidal localities: mid-tide level on Qingdao bathing beach
and mean low water level east of Qingdao pier. Cumulative curves of the sediment grain
size analysis data are shown in Fig. 1. The pier beach is considered polluted by sewage
whereas the bathing beach is relatively clean. Methods of microscopic study, abbreviations
and style of description were as set out in Platt & Zhang (1982).

Systematic descriptions

Oncholaimus qingdaoensis sp. nov.

Fig. 2

MATERIAL STUDIED. Holotype: cf, BM(NH) 1982.1.2. Allotype: 9 BM(NH) 1982.1.5. Para-
types: two males BM(NH) 1982.1.3^1.

LOCALITY. Qingdao bathing beach, Shandong, China.

DIMENSIONS

cf.: — 372 M 2590 ~,Ar. 00 , -

2640um; a = 98; b = 7; c = 53

20 25 27 20

~ 384 M 237°
22 21 25 21

2420 urn; a = 97; b = 6; c = 48

cf • — 340 M 2331 ~~or. n~ , _

3 - 2380um; a = 92; b = 7; c = 49

20 21 26 20

9: — 365 M 2300

22 31 32 24 2350 urn; a = 73; b = 6; c = 47

DESCRIPTION. Cuticle smooth. R, sensilla papilliform. R2 + R3 sensilla at the same level and
equal in size: 8-9 urn, 40-45% h.d. Head constricted immediately posterior to cephalic setae.
Amphid 9 jam wide, 40% c.d.: in some individuals the amphids are difficult to distinguish.
Buccal cavity 25-27 urn deep. Large left subventral, small right subventral and small dorsal

Bull. Br. Mus. nat. Hist. (Zool.) 45(5) : 253-261 Issued 29 September 1983

253

254

Z. N. ZHANG & H. M. PLATT

100 H

o

Fig. 1

-20246

Particle diameter (Phi)

Cumulative curves from the sediment analysis data for: (A — A) Qingdao bathing beach; (

Qingdao pier beach sediments.

teeth. Oesophagus without bulb. Excretory pore situated 48-64 urn from the anterior, about
2-2-5 times the length of the buccal cavity (Fig. 2a, b). Nerve ring situated at 43^18% of
oesophagus length. Tail conical, slightly curved ventrally; 2-0 a.b.d. in female, 2-5 a.b.d. in
male.

Spicules equal, 34—36 urn (1-8 a.b.d.), proximally cephalate and pointed distally. Guberna-
culum absent. Pairs of 3-7 um special subventral stout spine-like setae: 4 pairs precloacal,
4 pairs adcloacal and 3 pairs on the tail. Two pairs of precloacal spines, not conspicuously
situated on papillae. Conspicuous ventral papilla about 12 urn from the tail tip (Fig. 2c, d).
Two opposed testes.

Vulva at 69% of total length. Single anterior ovary. A demanian system could not be dis-
tinguished in the only female available for study.

DIFFERENTIAL DIAGNOSIS. Oncholaimus qingdaoensis sp. nov. belongs to a group of species
which have a conspicuous papilla near the male tail tip. Of these species, in terms of tail
shape, the new species most closely resembles O. cobbi (Kreis, 1932), O. domesticus
(Chitwood & Chitwood, 1938), O. hyrcanus Tchesunov, 1979 and O. longus (Wieser,
1953). Amongst several minor differences, O. qingdaoensis may be differentiated from O.
longus by its less slender body (male a = 92-98 vs 1 30-1 50) and more anterior excretory pore
(2-2-5 vs 4 buccal cavity lengths from anterior); from O. cobbi by the fewer number of
adcloacal pairs of spines (4 vs 10-15), shape of the postcloacal papilla and the presence of
precloacal spines; from O. domesticus by the lack of a well-developed demanian system with
adanal or postanal openings; from O. hyrcanus by the lack of unequal spicules, lack of
conspicuous precloacal papilla and different shape of the female tail. In addition, O.
qingdaoensis appears to differ from all of these four species in having its larger left subventral
tooth more conspicuously anterior to the dorsal and right subventral teeth.

ETYMOLOGY. The species name refers to the city near the type locality.

DISCUSSION. The genus Oncholaimus Dujardin, 1845 and the related genera are
currently in some considerable state of confusion and badly need revising. Several sets of
Oncholaimus-\ikQ species have been placed in separate genera, the bases of which appear
to be breaking down in the light of more recent findings. Genera separated from Oncholaimus
include Metaparoncholaimus De Coninck & Stekhoven, 1933 on the presence of two large

MARINE NEMATODES FROM CHINA

255

Fig. 2 Oncholaimus qingdaoensis: (a) cf, head; (b) c?2 head; (c) d1, tail region; (d) <53 tail region:

(e) 9 tail region. Bar scales: 10 urn,

256 Z. N. ZHANG & H. M. PLATT

equal subventral teeth; Oncholaimium Cobb, 1930 on (originally) the presence of a pre-
cloacal papilla and a demanian system without exit pores; Pseudoncholaimus Kreis, 1932
on the absence of a demanian system. Rachor (1969) considered the last two to be synony-
mous with Oncholaimus. So, for the purposes of this analysis, species previously assigned
to Oncholaimium and Pseudoncholaimus have been considered along with Oncholaimus
sensu stricto. Altogether, this group includes 91 known species, of which 26 are considered
to be dubious on the grounds of inadequate description.

Several Oncholaimus species have been reported as having teeth of equal or almost of
equal length, e.g. O. nigrocephalatus Cobb, 1930, O. opisthonchus Filipjev, 1927 and
O. problematicus Coles, 1977. Indeed, drawings of these species differ little from De Man's
original (1876) drawing of Metaparoncholaimus campylocercus, the genotype. Therefore,
any future revision of this group should carefully consider whether Metparoncholaimus
should be maintained as a distinct genus or whether the relative tooth sizes should be con-
sidered a variable infrageneric character within Oncholaimus, albeit of great use in practical
identification.

Oncholaimus sinensis sp. nov.
Fig. 3

MATERIAL STUDIED. Holotype: d1, BM(NH) 1982.1.6. Paratypes: cT2 BM(NH) 1982.1.6, c?3
in the collection of Z. N. Zhang and one juvenile BM(NH) 1982.1.7.

LOCALITY. East of Qingdao pier, Shandong, China.

DIMENSIONS

cf,: — 310 M 1990

2080um:a = 52;b = 7;c = 23

19 3g 4Q 22
'°

2010nm;a=50;b=6;c=25

W 4 23

rf3: — 340 M 2074

— — 4Q 42 23 2160um;a = 51;b = 6;c = 25

Juvenile: — 405 M 2480

21 35 35 23 2570 urn; a = 73; b = 6; c = 29

DESCRIPTION. Cuticle smooth. R, sensilla papilliform. R2 + R3 sensilla at the same level and
almost the same length: 5-6-5 um, 25-33% h.d. Amphid 7 urn wide, 29% c.d., situated just
anterior to dorsal tooth. Buccal cavity 24-26 um deep, armed with large left subventral tooth
and smaller right subventral and dorsal teeth. Excretory pore situated 80-93 um from
anterior, about 3 times the length of the buccal cavity (Fig. 3a). Nerve ring situated at 45-51%
of oesophagus length. Tail conico-cylindrical, 3-5^4 a.b.d. with slightly swollen tip.

Spicules equal, 26-27 um (1-2 a.b.d.), proximally slightly cephalate, distally pointed but
with some striations on the ventral side. Gubernaculum absent. Relatively conspicuous
'fleshy' precloacal papilla present. There is also a ventral postcloacal papilla situated at about
60% of the tail length from the cloaca with two pairs of 3 um setae. About 1 1 pairs of
4-5-7 um adcloacal setae in addition to subdorsal caudal setae.

DIFFERENTIAL DIAGNOSIS. Oncholaimus sinensis sp. nov. most closely resembles O. appendi-
culatus (Cobb, 1930) and O. campylocercoides De Coninck & Stekhoven, 1933 in the pres-
ence of a prominent precloacal papilla and a conico-cylindrical tail. However, O.
campylocercoides as originally described and as redescribed by Gerlach (1952) apparently
lacks the postcloacal papilla, although Wieser (1959) mentions and figures the presence of
two setose papillae in the middle of the tail. O. appendiculatus has a single ventral postcloacal
papilla but the spicules are much longer than those of the new species. Three other species

MARINE NEMATODES FROM CHINA

257

Fig. 3 Oncholaimus sinensis: (a) <$l anterior region; (b) cf, head; (c) d1, tail region; (d) juvenile tail

region. Bar scales: a = 20 um; b-d= 10 urn.

have been described with single postcloacal papillae and similarly shaped tails, O, manilius
Gerlach, 1957, O. martini Wieser, 1959 and O. olium Belogurov, Belogurova & Pavluk,
1975: all lack the conspicuous precloacal supplement and have their amphids situated more
posteriorly than in O. sinensis.

ETYMOLOGY. The species name refers to the country of the type locality.

258 Z. N. ZHANG & H. M. PLATT

DISCUSSION. It is hoped that an analysis of the female demanian system will be given in the
near future when new material is available. In the meantime, the assessment of these Chinese
specimens as belonging to a distinct species rests on the male characters alone. Judging from
descriptions in the literature, and in view of the need for a revision of the genus as discussed
above, it is considered that there are sufficient minor points of difference between O. sinensis
males and those of species which most closely resemble them to support the Chinese species
as being new. In any event, to place them in an already extant species would be to risk sub-
merging what might be useful information. The striated nature of the tip of the spicule, for
example, does not appear to have been reported hitherto, but may possibly have been over-
looked in some other species.

Metoncholaimus moles sp. nov.
Fig. 4

MATERIAL STUDIED. Holotype: rf, BM(NH) 1982.1.8. Allotype: 9, BM(NH) 1982.1.9. Para-
types: two males BM(NH) 1982.1.8. and two females BM(NH) 1982.1.9.

LOCALITY. East of Qingdao pier, Shandong, China.
DIMENSIONS

<f,: — 420 M 3540

25 54 5S 39 3700nm;a = 67;b=9;c = 23

<j • _ 446 M 3292

' 24 54 54 38 3440^m; a = 64; b = 8; c = 23

24 59 61 36
93:^ - 440 M 3204
~Y1 57 58 36

3820um;a = 63;b = 9;c = 28
3340um;a = 58;b = 8;c = 25

DESCRIPTION. Cuticle smooth. Six papilliform R, sensilla. R2 + R3 sensilla at the same level:
R2 sensilla 7-8 urn, 28-32% h.d.; R3 sensilla 5-7-5 urn, 20-30% h.d. Somatic setae 5-7 urn,
most numerous in the anterior oesophageal region. Head somewhat attenuated. Amphid
usually distinct, 10 urn wide (33% c.d.) and situated level with the dorsal tooth. Buccal cavity
26-30 urn deep. Large left subventral tooth and smaller right subventral and dorsal teeth.
Oesophagus widens slightly towards the posterior, but no bulb. Excretory pore just posterior
to the base of the buccal cavity, 29-36 urn from the anterior. Nerve ring at 44-49% of the
oesophagus length. Tail conico-cylindrical, 3-6-4-2 a.b.d. with a characteristic ventral inflec-
tion of the tip in the male (Fig. 4d).

Spicules equal, 43-44 um (chord), proximally cephalate and distally pointed (Fig. 4e).
Gubernaculum present with a well-developed dorsally directed apophysis, 2 1-22 urn long.
There are 9-1 1 pairs of stout adcloacal setae. Precloacally, there are an additional 4-6 thicker
but shorter setae (Fig. 4e): a further 3 pairs of similar setae are found subventrally near the
tail tip (Fig. 40- Two testes.

Vulva situated posterior to the mid-body; V = 68-78%. Single anterior ovary. Demanian
system appears to be relatively simple, typical of other Metoncholaimus species (Rachor,
1969). The terminal duct opens as a transverse slit about 65 um anterior to the anus. The
body is constricted immediately posterior to the pore (Fig. 4g), and in this region there are
characteristic subdorsal and subventral pairs of setae.

MARINE NEMATODES FROM CHINA

259

Fig. 4 Metoncholaimus moles: (a) d1, anterior region; (b) d1, anterior region; (c) 9, head region;
(d) cf, tail region; (e) d1, copulatory apparatus; (0 <?, tail tip; (g) 9, posterior region. Bar scales: a,
d, g = 20 |im; b, c, d, e= 10 |im.

260 Z. N. ZHANG &H. M. PLATT

DIFFERENTIAL DIAGNOSIS. Metoncholaimus moles sp. nov. is unique among the thirteen pre-
viously described species in having a well-developed dorsally directed gubernaculum.

ETYMOLOGY. The species name refers to the type locality being near a pier (moles, Latin =
pier).

DISCUSSION. The traditional distinction between the genera Oncholaimus Dujardin, 1845
and Metoncholaimus Filipjev, 1 9 1 8 is another that is beginning to break down. Characteristi-
cally, Oncholaimus species have short spicules and no gubernaculum whilst Metoncholaimus
species have extremely long spicules and a small gubernaculum lying parallel with the distal
end of the spicules, e.g. M. demani Zur Strassen. Already a number of Oncholaimus species
have been described with simple gubernacula, e.g. O. gladius Gerlach, 1956, O. longispiculo-
sus Gerlach, 1955 and O. dujardini sensu Wieser, 1953. Metoncholaimus species with
long spicules but without a gubernaculum have also been reported, e.g. M. intermedius
Wieser & Hopper, 1967 and M. scissus Wieser & Hopper, 1967. However, none of the
Oncholaimus or Metoncholaimus species, nor any species belonging to other genera of
the Oncholaiminae, have, to the best of our knowledge, been described as having a dorsally
directed gubernaculum.

References

Belogurov, O. I., Belogurova, L. S. & Pavluk, O. N. 1975. Morphology and systematic position of

two new species of marine nematodes (Nematoda: Oncholaimidae) from far-eastern seas: Oncholai-

mium olium sp. n. and Pseudoncholaimus vesicarius (Wieser, 1959) comb. nov. Biol. Morya,

Vladivostok 2: 25-30 (in Russian).
Chitwood, B. G. & Chitwood, M. B. 1938. Notes on the "culture" of aquatic nematodes. /. Wash.

Acad. Sci. 28:455^*60.
Cobb, N. A. 1930. The demanian vessels in nemas of the genus Oncholaimus; with notes on four new

oncholaims. J. Wash. Acad. Sci. 20:225-241.
Coles, J. W. 1977. Freeliving marine nematodes from southern Africa. Bull. Br. Mus. nat. Hist. (Zool.)

31: 1^9.
De Coninck, L. A. & Schuurmans Stekhoven, J. H. 1933. The freeliving marine nemas of the Belgian

coast. II. Mem. Mus. r. Hist. nat. Belg. 58: 1-163.
Dujardin, F. 1845. Histoire naturelle des Helminthes ou vers inestinaux. Libraire encyclopedique de

Roret, (Suites a Buffon). Paris 1845. 652 pp.
Filipjev, I. N. 1918. Free-living marine nematodes of the Sevastopol area. Trudy osob. zool. Lab.

Sevastop. biol. Sta. 4: 1-350 (in Russian).
1927. Les Nematodes libres des mers septentrionales appartenant a la famille des Enoplidae. Arch.

Naturgesch. 91 A (6): 1-216.
Gerlach, S. A. 1952. Nematoden aus dem Kiistengrundwasser. Abh. math.-naturw. Kl. Akad. Wiss.

Mainz 6: 31 5-372.
1955. Zur Kenntnis der freilebenden marinen Nematoden von San Salvador. Z. wiss. Zool. 158 :

249-303.

1956. Brasilianische Meeres- Nematoden I. Bolm Inst. Oceanogr. S. Paulo 5: 3-69.

1957. Die Nematodenfauna des Sandstrandes an der Kiiste von Mittelbrasilien (Brasilianische

Meres- Nematoden IV). Mitt. zool. Mus. berl. 33:41 1-459.
Kreis, H. A. 1932. Freilebende marine Nematoden von den Sunda-Inseln. II. Oncholaiminae. Vidensk.

Meddr dansk naturh. Foren. 93: 23-69.
De Man, J. G. 1876. Contribution a la connaissance des Nematodes marins du Golf de Naples.

Tijdschr. ned. dierk. Vereen. 3: 88-1 18.
Platt, H. M. & Zhang, Z. N. 1982. New species of marine nematodes from Loch Ewe, Scotland. Bull.

Br. Mus. nat. Hist. (Zool.) 42(4): 227-246.
Rachor, E. 1969. Das de Mansche Organ der Oncholaimidae, eine genito-intestinale Verbindung bie

Nematoden. Z. Morph. Tiere 66: 87-166.
Tchesunov, A. V. 1979. Oncholaimus hyrcanus nom. n., a new name for Oncholaimus orientalis

Tchesunov, 1976. Zool. Zh. 58(2): 260 (in Russian).

MARINE NEMATODES FROM CHINA 261

Wieser, W. 1953. Free-living marine nematodes I. Enoploidea. Ada Univ. /wm/(N.F.2) 49(6): 1-155.
1959. Free-living nematodes and other small invertebrates of Puget Sound beaches. 179 pp.

University of Washington Press, Seattle.
& Hopper, B. 1967. Marine nematodes of the east coast of North America. Bull. Mus. comp.

Zool. Harv. 135: 239-244.

Manuscript accepted for publication 15 November 1982

Echinoderms of the Rockall Trough and adjacent
areas

I. Crinoidea, Aster oidea and Ophiuroidea

J. D. Gage & Margaret Pearson

Dunstaffnage Marine Research Laboratory, Scottish Marine Biological Association, P.O.
Box 3, Oban, Argyll PA34 4AD

Ailsa M. Clark & G. L. J. Paterson

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

P. A. Tyler

Department of Oceanography, University College of Swansea, Swansea SA2 8PP

Synopsis

Four species of crinoid, forty species of sea star and thirty-six species of brittle star are identified from
recent benthic sampling in the deep-sea area to the west of the British Isles, mainly in the Rockall
Trough. Of these, one crinoid, nine asteroids and eighteen ophiuroids have not previously been
recorded from the seas around the British Isles. The zoogeographical distribution and bathymetric
range of each species is summarised as far as was previously known giving the range extension provided
by the present records. Notes are also provided on observations on the biology, including the mode
of reproduction, of the more abundant species. The records demonstrate a broader bathymetric distri-
bution of juvenile and post larval stages than of the adult populations of some of the more abundant
species. A greater diversity of species also is evident from the small number of samples from the
western, probably current- swept, side of the Rockall Trough compared with the much larger number
of samples collected at similar depths in the east.

Introduction

The cruises of H.M.S. Lightning and H.M.S. Porcupine in the years 1868-70 in the Rockall
Trough and other deep sea areas lying west of the British Isles and of the Iberian peninsula
asserted beyond doubt the presence of life at great depths.

Echinoderm data figure prominently in the literature (Lyman, 1882; Sladen, 1889;
Thomson, 1872, 1873, 1874) resulting from these cruises that covered deep-sea areas not
subsequently sampled by the Challenger expedition of 1872-76. Many useful echinoderm
records also were made on the cruises of H.M.S. Knight-Errant in 1880 and H.M.S. Triton
in 1882 in the course of exploration of the submarine sill bounding the northern end of the
Rockall Trough that was subsequently named the Wyville Thomson Ridge (e.g. Hoyle, 1884;
Sladen, 1882). To the authors' knowledge most subsequent records of deep-sea echinoderm
fauna from this area have emanated from opportunistic extension into deeper water of essen-
tially coastal sampling programmes. Notable among these are the cruises of the Irish Fish-
eries Department cruiser Helga in the early years of this century. Exploration of some of
the outer banks and shelf areas around the northern perimeter of the Rockall Trough has
also resulted in many useful records of fauna including echinoderms (Danielssen & Koren,
1884; Siissbach & Breckner, 1911; Pawsey & Davis, 1924). Other records have come from
stations worked in the Rockall area in the course of wider ranging cruises, such as those
of the Thor, and the Michael Sars in her Atlantic expedition of 1910, and more recently

Bull. Br. Mus. not. Hist. (Zool.) 45(5): 263-308 Issued 29 September 1983

263

264

62°N

J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

^VQ, \M

Bathymetry

200m

1000 m

2000 m

3000m

4000m

Other isobaths at
500m intervals

/ V Porcupine

18°W 16 U 12 10 8 6 4 2 0°W

Fig. 1 Bathymetric chart showing Permanent Station and Station 'M' and all other sample stations

(crosses).

ROCKALL TROUGH ECHINODERMS I 265

by the French oceanographic ship Jean Charcot in 1969 (Cherbonnier & Sibuet, 1973) and
in 1976 (INCAL expedition). Although not extending into the Rockall Trough the sampling
in neighbouring northern areas by the major expeditions to the N. Atlantic such as the cruises
of the Voeringen Norwegian North Atlantic Expedition, 1876-1878, and the Danish Ingolf
expedition, 1895-6, should also be mentioned in this context.

The Scottish Marine Biological Association (SMBA) in 1973 initiated exploratory biologi-
cal sampling in the deep waters off the continental shelf as an extension of programmes pre-
viously restricted to coastal environments. Subsequent sampling has been undertaken on
many cruises mainly on R.R.S. Challenger, in the Rockall Trough and the adjacent continen-
tal slope off the west of Scotland (Fig. 1 ). This work has been focussed both as quantitative
and seasonal sampling programmes in studies of the structure, reproduction and growth of
midwater, demersal and benthic populations offish and invertebrate species at fixed stations
located in the middle and southern part of the Rockall Trough (Gordon, 1979; Gage et al,
1980; Gage & Tyler, 1982; Kawaguchi & Mauchline a, b in press). In addition there has
been some sampling by the SMBA in the N Feni Ridge and Wyville Thomson Ridge areas,
in the Porcupine Seabight and its vicinity, and on the S. Feni Ridge (Fig. 1 ).

The present account concentrates on echinoderm distributions, because populations of
echinoderm species are conspicuous elements of the benthic megafauna, animals easily
recognisable in sea-bed photographs). Echinoderms hence may frequently occur in the fish
catches obtained by coarse-meshed demersal trawls, such as the large commercial pattern
Granton trawl and smaller single-warp otter trawls. Echinoderms also dominate catches
made with a 3 -metre wide Agassiz trawl. This trawl has been deployed in poor weather that
has precluded use of other gear employed in the SMBA programme, such as the epibenthic
sled (Hessler & Sanders, 1967) and USNEL 0-25 m2 Box Corer (Hessler & Jumars, 1974)
both of which have the advantage of retaining benthic larval and juvenile stages, many of
them identifiable. Results indicate that juvenile stages may extend over a wider area and
greater bathymetric range than that of adult specimens caught in trawls. Therefore in describ-
ing the distributional range of species it is necessary to distinguish records where larvae or
juveniles only have been found from those where adults have been collected, and where
presumably a breeding population exists.

In the present account, the geographical distribution and bathymetric data relating to both
past and present records are interpreted in relation to the properties and dynamics of water
masses, which are relatively well known in the Rockall area (Ellett & Martin, 1973; Roberts,
1975).

Station List

Details of bottom sampling stations worked are listed below. Except for benthic stations
118-125 that were worked from R.R.S. Shackleton, all stations listed were worked from
R.R.S. Challenger. Benthic stations and fishing stations, worked by Dr J. D. M. Gordon of
SMBA, are listed separately in chronological order with a prefix to the station number that
identifies gear used: ES, epibenthic sled; ABD, anchor-box dredge; SBC, \ m2 spade box corer;
AT, 3 m Agassiz trawl; GT, Granton trawl; SWT, single warp otter trawl, OTSB, semi-
balloon otter trawl 14. For most fish trawls (GT, SWT and OTSB) depths at mid-point of
haul is unknown and total depth range of haul is given.

Position

(at mid-point of
track on bottom
St. No. Date if applicable) Depth (m)

Benthic stations

ES2 4Junel973 55°04'N, 12°33'W 2857

ABD 3 5 June 1973 56°46 'N, 10°02 'W 1997

266

St. No.

J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

Date

Position

(at mid-point of
track on bottom
if applicable)

Depth (m)

ES4

5 June 1973

56°52'N, 10°01'W

1993

ESS

5 June 1973

56°43 'N, 09°05 'W

763

ES6

2 July 1973

55°03'N, 12°29'W

c. 2900

ES8

3 July 1973

54°45'N, 12°10'W

c. 2900

ES 10

4 July 1973

56°37'N, 10°04'W

c. 2540

ES 12

20 Sept. 1973

56°49'N, 10°15'W

2076

ES 14

22 Sept. 1973

56°45'N, 09°46'W

1770

ES 15

22 Sept. 1973

56°44'N, 09°28'W

1632

ES 18

22 Sept. 1973

56°44'N, 09°20'W

1392

ES20

23 Sept. 1973

56°46'N, 09°17'W

1271

ES22

23 Sept. 1973

56°41'N, 09°11'W

1028

ES23

23 Sept. 1973

56°37'N,09°10'W

704

ABD24

23 Sept. 1973

56°36'N, 09°13'W

810

ABD25

23 Sept. 1973

56°36'N, 09°10'W

588

ES27

3 Nov. 1973

54°40'N, 12°16'W

c. 2880

ES28

3 Nov. 1973

54°33'N, 12°21'W

c. 2880

ES31

20 Mar. 1975

55°03'N, 12°01'W

2875

ES32

22 Mar. 1975

55°16'N, 12°58'W

2871

ES33

23 Mar. 1975

56°35'N, 07°16'W

148

ES34

10 May 1975

56°36'N, 11°30'W

2515

SBC 46

6 Sept. 1975

55°04'N, 12°06'W

2875

SBC 47

6 Sept. 1975

55°03'N, 12°03'W

2875

SBC 48

7 Sept. 1975

55°04'N, 12°04'W

2875

SBC 49

7 Sept. 1975

55°03'N, 12°05'W

2875

SBC 50

7 Sept. 1975

55°04'N, 12°02'W

2875

SBC 51

7 Sept. 1975

55°03'N, 12°03'W

2875

ES52

15 Nov. 1975

c. 54°40'N, 12°16'W

2886

ES53

15 Nov. 1975

c. 54°40'N, 12°16'W

2886

ES54

17 Nov. 1975

c. 54°40'N, 12°16'W

2878

ES55

17 Nov. 1975

c. 54°40'N, 12°16'W

2878

ES56

1 Mar 1976

c. 54°40'N, 12°16'W

2886

ES57

21 June 1976

c. 54°41'N, 12°23'W

c. 2900

SBC 58

21 June 1976

54°41'N, 12°17'W

c. 2900

ES59

21 June 1976

54°40'N, 12°20'W

c. 2900

SBC 60

23 June 1976

56°35'N, 11°03'W

c. 2500

SBC 61

23 June 1976

57°08'N, 12°09'W

c. 2000

SBC 62

23 June 1976

57°28'N, ITOO'W

c. 610

SBC 63

25 June 1976

56°37'N, 09°49'W

c. 1800

SBC 64

26 June 1976

56°38'N,09°29'W

c. 1400

SBC 65

26 June 1976

56°39'N, 09°40'W

c. 1600

SBC 66

26 June 1976

56°39 'N, 09°23 'W

c. 1200

SBC67

26 June 1976

56°39'N, 09°13'W

c. 1000

SBC 68

1 July 1976

58°42'N,09°43'W

c. 1800

AT68A

2 July 1976

59°13'N, 08°01'W

1330

ES69

2 July 1976

59°39'N, 07°12'W

1050

ES87

4 July 1976

61°13'N, 03°59'W

1050

ES90

7 July 1976

60°05'N, 05°55'W

1040

AT90A

7 July 1976

60°05'N, 05°57'W

1040

ES99

9 July 1976

60°00'N, 10°35'W

1160

ES 105

10 July 1976

58°27'N, 12°35'W

1600

AT 107 A

11 July 1976

57°07'N, 12°06'W

c. 2000

ES 111

22 Oct. 1976

54°40'N, 12°16'W

2886

ES 112

25 Oct. 1976

55°12'N, 15°50'W

1900

ROCKALL TROUGH ECHINODERMS I 267

Position

(at mid-point of
track on bottom
St. No. Date if applicable) Depth (m)

ES 113

27 Oct. 1976

57°18'N, 14°07'W

168

AT 114

28 Oct. 1976

56°48'N, 10°54'W

2400

ES 115

28 Oct. 1976

56°29'N, 10°22'W

1000

ES 118

28 Jan. 1977

54°39'N, 12°14'W

2910

AT 119

28 Jan. 1977

54°40'N, 12°14'W

2908

ES 120

28 Jan. 1977

54°39'N, 12°16'W

2911

AT 121

29 Jan. 1977

54°37'N, 12°09'W

2910

ABD 125

30 Jan. 1977

52°56'N, 12°58'W

394

ES 129

7 April 1977

54°39'N, 12°17'W

c. 2900

AT 131

7 April 1977

55°03'N, 12°20'W

c. 2900

AT 132

8 April 1977

53°14'N, 14°16'W

225

AT 133

8 April 1977

52°30'N, 14°03'W

315

AT 134

9 April 1977

54°05'N, 12°06'W

800

ES 135

7 Aug. 1977

54°39'N, 12°16'W

2900

ES 137

22 Feb. 1978

54°34'N, 12°19'W

2900

AT 138

23 Feb. 1978

55°31'N, 10°24'W

2450

AT 139

24 Feb. 1978

55°35'N, 10°25'W

2450

ES 140

13 April 1978

54°40'N, 12°16'W

2912

AT 141

14 April 1978

54°44'N, 12°14'W

2909

ES 143

14 April 1978

54°41'N, 12°14'W

2892

AT 144

19 April 1978

57°13'N, 10°20'W

2240

ES 147

2 June 1978

54°36'N, 12°19'W

2921

SBC 150

3 June 1978

54°37'N, 12°13'W

2916

AT 151

6 June 1978

57°21'N, 10°22'W

2175

ES 152

13 Jan. 1979

54°42'N, 12°02'W

2900

AT 153

15 Jan. 1979

57°20'N, 10°27'W

2200

AT 154

21 May 1979

57°08'N, 10°22'W

2264

SBC 155

4 Aug. 1979

48°27'N, 10°20'W

1330

AT 157

6 Aug. 1979

49°31'N, 13°11'W

1752

AT 161

8 Aug. 1979

50°52'N, 12°22'W

2055

AT 162

9 Aug. 1979

51°50'N, 13°00'W

992

SBC 163

10 Aug. 1979

54°41'N, 12°24'W

2910

ES 164

11 Aug. 1979

54°37'N, 12°22'W

2925

SBC 166

12 Aug. 1979

57°07'N, 10°20'W

2274

AT 167

13 Aug. 1979

57°05'N, 10°25'W

2300

AT 169

28 Feb. 1980

54°40'N, 12°17'W

2910

AT 171

3 Mar 1980

57°16'N, 10°17'W

2225

ES 172

27 May 1980

54°39'N, 12°17'W

2910

AT 175

28 May 1980

57°19'N, 10°16'W

2210

ES 176

28 May 1980

57°15'N, 10°26'W

2245

AT 177

29 May 1980

57°18'N, 10°16'W

2220

ES 178

29 May 1980

56°33'N, 09°17'W

997

ES 180

15 Sept. 1980

54°42'N, 12°H'W

2886

AT 181

16 Sept. 1980

57°19'N, 10°28'W

2220

ES 182

16 Sept. 1980

57°19'N, 10°28'W

2170

ES 184

17 Sept. 1980

57°14'N, 10°24'W

2260

ES 185

10 April 1981

54°44'N, 12°15'W

2907

AT 186

12 April 1981

57°22'N, 10°19'W

2170

ES 190

16 Aug. 1981

54°41'N, 12°18'W

2898

AT 191

17 Aug. 1981

56°00'N, 13°58'W

2190

AT 192

18 Aug. 1981

57°21'N, 12°02'W

1862

AT 194

18 Aug. 1981

57°27'N, 11°10'W

630

AT 195

18 Aug. 1981

57°23'N, 10°27'W

2190

268

St. No.

J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

Date

Position

(at mid-point of
track on bottom
if applicable)

Depth (m)

ES 197

19 Aug. 1981

57°21'N, 10°29'W

2220

AT 198

15 Oct. 1981

57°15'N, 10°20'W

2215

ES200

6 Feb. 1982

57°20'N, 10°32'W

2220

Fishing stations

mean

range

GT 1

19 Mar. 1975

56°37'N, 09°09'W

713

650-805

GT2

19 Mar. 1975

56°37'N, 09°15'W

1027

972-1084

GT7

3 Sept. 1975

56°28'N, 09°11'W

689

640-780

GTS

3 Sept. 1975

56°25'N, 09°20'W

1054

1024-1072

GT 11

7 April 1976

56°32'N, 09°17'W

1014

934-1054

GT 13

8 April 1976

56°33'N, 09°07'W

523

508-543

GT 14

9 April 1976

56°33'N, 09°12'W

761

713-788

GT 15

24 June 1976

56°28'N, 09°11'W

720

672-740

GT 16

25 June 1976

56°25'N, 09°20'W

958

916-992

GT 17

25 June 1976

56°28'N, 09°27'W

1237

1190-1296

SWT 10

5 Aug. 1977

56°51'N,09°58'W

2018

2000-2030

SWT 11

6 Aug. 1977

56°03'N, 11°05'W

2530

2500-2560

SWT 12

9 Aug. 1977

53°25'N, 15°24'W

2996

2980-3000

SWT 13

9 Aug. 1977

52°04'N, 16°09'W

3463

3425-3500

SWT 15

11 Aug. 1977

49°30'N, 16°12'W

4810

constant

AT 1

19 Oct. 1977

56°24'N,09°12'W

750

—

AT 3

20 Oct. 1977

56°23'N, 09°18'W

1000

—

SWT 16

21 Oct. 1977

56°12'N, 10°57'W

2441

2400-2490

SWT 17

22 Oct. 1977

56°49'N, 09°55'W

1977

1940-2020

SWT 18

22 Oct. 1977

56°46'N,09°42'W

1809

1785-1845

SWT 27

4 May 1978

54°26'N, 12°52'W

2965

constant

SWT 32

31 Oct. 1978

56°48'N,09°56'W

2006

1995-2020

OTSB51001

30 April 1981

54°45'N, 12°24'W

2890

2885-2895

(starting position)

List of Species

Class Crinoidea
Order Millericrinida
Family Bathycrinidae

Rhizocrinus lofotensis M. Sars
Order Comatulida

Family Pentametrocrinidae

Thaumatocrinus jungerseni A. H. Clark
Family Antedonidae
Heliometra glacialis (Owen)
Hathrometra sarsi (Diiben & Keren)

Class Asteroidea
Order Paxillosida
Family Luidiidae
Luidia sarsi (Diiben & Keren)
Luidia ciliaris (Philippi)
Family Astropectinidae
Astropecten irregularis (Pennant)

Bathybiaster vexillifer (Wyville

Thomson)

Plutonaster bifrons (Wyville Thomson)
Dytaster insignis (Perrier)
Psilaster andromeda (Miiller & Troschel)
Psilaster patagiatus Sladen
Family Porcellanasteridae
Porcellanaster ceruleus Wyville

Thomson

Hyphalaster inermis Sladen
Order Notomyotida

Family Benthopectinidae
Benthopecten simplex (Perrier)
Pect master filholi Perrier
Pontaster tenuispinus (Diiben & Keren)
Order Valvatida

Family Odontasteridae

ROCKALL TROUGH ECHINODERMS I

269

Hoplaster spinosus Perrier
Odontaster sp.

Family Radiasteridae
IRadiaster tizardi (Sladen)

Family Goniasteridae
Pseudar chaster parelii (Diiben & Koren)
Pseudar chaster gracilis (Sladen)
Paragonaster subtilis (Perrier)
Plinthaster dentatus (Perrier)
Evoplosoma scorpio Downey

Family Poraniidae

Porania pulvillus O. F. Miiller
Order Spinulosida

Family Pterasteridae
Pteraster militaris (O. F. Miiller)
Pteraster pulvillus M. Sars
Pteraster reductus Koehler
Pteraster sp. aff. P. acicula (Downey)
Hymenaster membranaceus Wyville

Thomson
Hymenaster pellucidus Wyville

Thomson

Hymenaster gennaeus H. L. Clark
Hymenaster rex Perrier

Family Solasteridae
Crossaster squamatus (Doderlein)

Family Echinasteridae
Henricia sanguinolenta (O. F. Miiller)
Henricia abyssicola (Norman)
Order Forcipulatida

Family Brisingidae
Brisinga endecacnemos Asbjornsen
Brisingella coronata G. O. Sars
Freyella spinosa Perrier
Freyella sexradiata (Perrier)

Family Asteriidae
Hydrasterias sexradiata (Perrier)
Stichastrella rosea (O. F. Miiller)

Family Zoroasteridae
Zoroaster fulgens Wyville Thomson

Class Ophiuroidea
Suborder Euryalida
Family Asteronychidae
Asteronyx loveni Miiller & Troschel
Astrodia tenuispina Verrill

Family Asteroschematidae
Asteroschema inornatum Koehler

Family Gorgonocephalidae
Gorgonocephalus caputmedusae

(Linnaeus)
Order Ophiurida

Family Ophiacanthidae
Ophiotrema alberti Koehler
Ophiomyces grandis Lyman
Ophiacantha abyssicola G. O. Sars
Ophiacantha bidentata (Retzius)
Ophiacantha crassidens Verrill
Ophiacantha cuspidata Lyman
Ophiacantha simulans Koehler
Ophiacantha aculeata Verrill
Ophiolimna bairdi (Lyman)

Family Ophiactidae
Ophiactis abyssicola (M. Sars)
Ophiactis balli (Thompson)

Family Amphiuridae
Amphiura otteri Ljungman
Amphiura fragilis Verrill
Amphiura griegi Mortensen
Amphipholis squamata (Delle Chiaje)

Subfamily Amphilepidinae
Amphilepis ingolfiana (Mortensen)

Family Ophiochitonidae
Ophiochiton ternispinus Lyman

Family Ophiuridae

Subfamily Ophiurinae
Ophiopleura inermis (Lyman)
Ophiopleura borealis Danielssen &

Koren

Homalophiura tesselata (Verrill)
Amphiophiura convexa (Lyman)
Amphiophiura saurura (Verrill)
Stegophiura macrarthra H. L. Clark
Ophiocten gracilis G. O. Sars
Ophiocten hastatum Lyman
Ophiura affinis Liitken
Ophiura carnea Liitken
Ophiura irrorata (Lyman)
Ophiura imprudens (Koehler)
Ophiura ljungmani (Lyman)
Ophiura ophiura (Linnaeus)

Subfamily Ophiolepidinae
Ophiomusium lymani Wyville Thomson

Systematic Account

The following account aims to summarise known distributional records for the individual
species together with observations related to the biology of the more abundant ones, includ-
ing the colour of fresh material if this information is not given by Mortensen (1927).

Throughout, taxonomic references are kept to a minimum by citing, where available, one
or more works containing a good description from which the original description and syn-
onymy may be traced. After these citations the sampling stations from which each species
has been collected are listed together with the total number of specimens, adults and juven-

270 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

iles, in parentheses. Records from benthic stations are followed by those for fishing stations,
which include two Agassiz trawl hauls. Where juveniles only were collected, this is indicated
but the term juvenile is used only as a guide and is based arbitrarily on size. The number
of specimens has no quantitative significance in comparisons between samples because for
most epibenthic sledge hauls a partial sorting only of the catch has been achieved. Total
depth range of the present records is given in square brackets after this listing. Details of
hauls, including depth range and positional coordinates are listed on pp. 265-268.

Systematic structure of the class Crinoidea follows A. M. Clark (19706); that of the
Asteroidea, Downey (1973), Clark (1981) and Blake (1981); while for the Ophiuroidea,
including family and subfamily names, we follow Fell (1960). References to the size of
animals refer to disk diameter of brittle stars and arm radius, R (measured from arm tip
to the centre of the disk) and the interradius, r, of sea stars.

Although all co-authors of the present study have been involved in the preparation and
revision of the text, taxonomic responsibility for the identifications of the crinoids and most
of the asteroids is taken by A.M.C. and for the ophiuroids by G.L.J.P.

Examples of all species referred to in the present account have been or will be deposited
with the British Museum (Natural History).

Class CRINOIDEA
Order MILLERICRINIDA
Family BATHYCRINIDAE

Rhizocrinus lofotensis M. Sars, 1868
See: A. M. Clark, 19706 : 19-22, fig. 4, map 2.
SAMPLES. ES 18 (3), ES 20 (8). [1271-1392 m]

DISTRIBUTION. Recorded from the W coast of Norway, the Rockall Trough area, Iceland,
Greenland and the eastern side of North America. Depth range off Norway includes the
minimum, 140-700 m, while those from both sides of Greenland include the maximum
of 3135m (Ingolf st. 37) but other records east of Iceland rarely much exceed 1200m.
[Carpenter's record from NE Brazil mentioned by A. M. Clark (19706) was referred to
Democrinus by A.M.C. (1977)].

Previous records from the Rockall Trough area include Lightning and Knight Errant
material from 6 1°02'N, 12°04'W, 1990 m and c.60°N, 07°W, c.960 m (Carpenter, 1884) and
Thor st. 99, 61°15 'N, 09°35 'W, 872 m, recorded by A. H. Clark (1923) who earlier (1913)
recorded one specimen from Helga st. CXX, 53°58 'N, 12°24'W, 687 m. The latter record
probably validates Kemp's (1905) provisional record from '64 miles NW by N of Cleggan
Head' (c.54f N, 1 If W), 364 m.

The present sample from the Hebridean slope (56°46'N), consists only of calyces and
separate stalk fragments.

REMARKS. We retain the familiar and historic generic name Rhizocrinus M. Sars, 1868,
although according to Roux (1976) it should probably be treated as a junior synonym of
Conocrinus d'Orbigny, 1850, because of the fused calyx plates in both nominal genera.
However, in 1978 Roux listed Rhizocrinus with R. lofotensis among the Bathycrinidae.
Conocrinus was known only from lower Caenozoic fossil species until Roux ascribed to it
two recent species from the Bay of Biscay which he named C. cherbonnieri and C. cabiochi.
The former is based on a single incomplete specimen from the French research ship Thalassa
st. Z397, 47°33-8 'N, 07°12-6 ' W, 5 1 1 m; superficially very similar to R. lofotensis but said
to differ in the joint structure of the columnals.

ROCK ALL TROUGH ECHINODERMS I 271

Order COMATULIDA
Family PENTAMETROCRINIDAE

Thaumatocrinus jungerseni A. H. Clark, 1923
See: A. H. Clark, 1923 : 13-17, figs 2-4; A. M. CLark, 1980 : 204-205.
SAMPLES. SWT 12 (8), SWT 13 (2). [2980-3000 m; 3425-3500 m]

DISTRIBUTION. Previously known from the Denmark Strait, NW and SW of Iceland and far
to the west of Ireland (c.53°N, 20°W) in 823-2734 m.

The present new records from the Porcupine Seabight from between 2980 m and 3500 m
depth extend the geographical range eastwards and the lower limit of the bathymetric range.
However, an unpublished record exists of four specimens from R.R.S. Discovery station
9640/1, (50°05 ' N, 13°51 ' W, 3749-3757 m depth, soft mud) that were identified by A.M.C.

Family ANTEDONIDAE

Heliometra glacialis (Owen, 1833)

See: A. M. Clark, 1970/7 : 24-27, fig. 5, map 3.
SAMPLE. ES 112(1). [1900m]

DISTRIBUTION. Previously known primarily from the Arctic where it is circumpolar, extend-
ing south in the NW Atlantic to George's Bank off Cape Cod, but in the east only to the
Faroe Channel and northernmost Norway; 14-1358 m; usually on gravel with sand and mud
or loose stones.

The present record from the South Feni Ridge extends both the known range southwards
to latitude 55°N in the NE Atlantic and the lower limit of the bathymetric range.

REMARKS. An arcturid amphipod, identified as such by Dr R. J. Lincoln of the British
Museum (Natural History), was found associated with this specimen.

^Hathrometra sarsi (Diiben & Koren, 1 846)

See: A. M. Clark, 19706: 42-45, figs 15, 16 for Poliometra prolixa, 39-42, fig. 14 for Hathrometra
sarsi, 36-39, fig. 12 for Leptometra celtica.

SAMPLE. ES 20(1). [1271m]

REMARKS. The single badly broken specimen has lost all the cirri completely and the arms
from the first or second syzygy (brachials 3 or 9). An arm fragment from the same sample
has pinnules with very long pinnulars. This fact and the blunt conical form of the centrodor-
sal suggest identity with Poliometra prolixa (Sladen) or Hathrometra sarsi (Diiben & Koren),
possibly Leptometra celtica (Barrett & McAndrew) but the depth and latitude are both too
high to make this last identity likely. A. H. Clark (1913) recorded P. prolixa from Silver
Belle st. 9, 60°18'N, 04°43 'W, 905 m and referred to 'Hathrometra sp.' single specimens
from Helga sts SR 353 and 506 at c.SOfN, 1 1°W, down to 990 and 1230 m. In 1967 (in
Clark & Clark) he cited both of these as H. sarsi without question, prompting A.M.C. (1970&)
to repeat this southern record, though clearly it needs confirmatory material. Otherwise, the
southernmost record of H. sarsi to the west of the British Isles is from Triton st. 2,
59°37 '30 " N, 06°19 ' W, 969 m very close to Porcupine st. 54, 633 m where P. prolixa was
taken (Clark & Clark, 1967). Of the two species, H. sarsi is much more likely to extend
further south since it extends in Scandinavia to Danish waters whereas P. prolixa is limited
to northernmost Norway. Morphologically, the two are distinguished by the more regular
arrangement of the cirri in Poliometra, tending to form vertical lines on the centro-
dorsal, especially in larger specimens; on this count, the present specimen inclines towards
Hathrometra.

272 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

Class ASTEROIDEA

Order PAXILLOSIDA

Family LUIDIIDAE

Luidia sarsi Diiben & Koren, 1 845

See: Sussbach & Breckner, 191 1 : 210-214; A. M. Clark, 1982 : 175-80, figs Ic, 2c, g, 3m-o, v, 6.
SAMPLES. ES 33 (4), ES 1 13 (juvenile, 1). [148 m, 168 m]

DISTRIBUTION. From the Faroe Bank Channel eastwards to Trondheim, Norway and south-
wards to the Mediterranean, Cap Blanc and the Azores; 9-1300 m; usually on muddy rather
than clean sand, or on clay. Luidia sarsi was taken by the Helga at many stations worked
off the west coast of Ireland.

The present records from the Hebridean Sea and the Rockall Bank are well within the
known bathymetric range and its present occurrence further north and west is to be expected
(Mortensen, 1927).

Luidia ciliaris (Philippi, 1837)

See: Sussbach & Breckner, 1911: 209-210; Mortensen, 1927 : 70, fig. 39a.
SAMPLE. GT 1 (1). [650-805 m]

DISTRIBUTION. From the Faroe Channel east to southern Norway and south to the Mediter-
ranean, Canary Is. and Azores; previously recorded from 1 to 400 m (Mortensen, 1927).

The present record surprisingly extends the deeper limit to at least 650 and possibly 805 m
depth; substratum usually fine gravel, often with little or no mud.

Family ASTROPECTINIDAE

Astropecten irregularis (Pennant, 1777)
See: Sussbach & Breckner, 1911: 203-207; Mortensen, 1927 : 57-58, fig. 32.

SAMPLES. AT 133 (7 var. pentacanthus). GT 1 (1), GT 7 (3 var. pentacanthus). [315m to
650-805 m]

DISTRIBUTION. Coastal distribution from Norway south to Morocco; 10-1000 m. It is said
to prefer a sandy bottom (Mortensen, 1927).

The presence of this well-known species on the upper Hebridean slope down to between
650 and 805 m depth is not unexpected. The specimens from GT 7 and AT 133 lack the
large spine on the superomarginals and therefore agree with the variety pentacanthus (Delle
Chiaje).

Bathybiaster vexillifer (Wyville Thomson, 1873)
See: Grieg, 1921 [1932] : 16-18; Mortensen, 1927 : 61-63, figs 34, 35 (as B. robustus and B. vexillifer}.

SAMPLES. ES 10 (1), ES 34 (3), AT 90A (1), AT 107A (2), AT 1 14 (6), AT 138 (juvenile, 1), AT 139
(6), AT 144 (23), AT 151 (15), AT 153 (22), AT 154 (34), AT 161 (6), AT 162 (8), AT 167 (9), AT
171 (28), AT 175 (10), ES 176 (4), AT 177 (11), ES 178 (2), AT 181 (25), ES 182 (1), ES 184 (7),
AT 186 (10), AT 191 (juvenile, 1), AT 192 (6), AT 195 (17), ES 197 (20), AT 198 (9), ES 200
(3). SWT 10 (1), SWT 1 1 (1), SWT 16 (28), SWT 17 (3). [992 m-2540 (72560) m]

DISTRIBUTION. This purely circumpolar Arctic/N boreal species is known to extend south
to North Carolina in the west and the Bay of Biscay in the east Atlantic; 630-3 100m
(shallower depths only within the Arctic); on soft muddy sediments.

Our shallowest record at 992 m on the Porcupine Bank does not accord with a distribution
limited by a temperature of 1-4°C suggested by Cherbonnier & Sibuet (1973) since

ROCKALL TROUGH ECHINODERMS I

273

temperatures in the Rockall Trough between 900 and 1 000 m, although showing little
seasonal variation, may exceed 8°C (Ellett & Martin, 1973).

REPRODUCTION. Histological examination of the gonads of individuals collected in a time
series from St. 'M' show no evidence of reproductive synchrony within or between samples
taken at different times of the year; the sexes are separate and the large size of ripe oocytes
(c. 1000 urn diameter) suggests direct, external lecithotrophic development (Tyler, Pain &
Gage, 1982#). There is no evidence of brooding of young.

Plutonaster bifrons (Wyville Thomson, 1873)
See: Grieg, 1921 [1932]: 13-14; Mortensen, 1927 : 63-64.
SAMPLES. ES 4 (4), ES 6 (juvenile 1), ES 18 (1), ES 20 (3), AT 68A (1), ES 69 (1), ES 105 (1), AT

58°N

58° N

57-

U°W

9°W

.Plutonaster bifrons

D Porcellanaster ceruleus • Paragonaster subtilis

m Psilaster andromeda *Be"t»opecten simplex OHymenastermembranaceus
A Bothy blaster vexillifer T pect master f,lholi + Zoroaster fulgens

O Pseudarchaster parelii

Fig. 2 Chart of central part of Rockall Trough showing station locations for records of the ten most
abundant sea star species. Larger symbols denote records from more than one sample taken on or
near the position.

274 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

107A (6), AT 144 (10), AT 151 (7), AT 153 (20), AT 154 (5), AT 161 (10), AT 167 (15), AT 171
(17), AT 175 (5), ES 176 (2), AT 177 (12), AT 181 (15), ES 182 (1), ES 184 (2), AT 186 (24), AT
191 (4), AT 192 (69), AT 195 (17), ES 197 (2), AT 198 (3), ES 200 (1). GT 17 (4), SWT 10 (40),
SWT 1 1 (12), SWT 17 (3), SWT 18 (38), SWT 27 (4), SWT 32 (12), AT 3 (5). [1000-2965 m]

DISTRIBUTION. Previously known from the Faroe Channel to South Africa; 800-2500 m; on
soft muddy sediments.

Our records for this species extend the lower limit of the known range to 2965 m. The
bathy metric range of these records covers a temperature range of around 3-5-7°C in the
Rockall Trough (Ellett & Martin, 1973). Farran (1913) indicates from trawlings in the Porcu-
pine Seabight that P. bifrons does not usually occur off the Irish coast at depths less than
500 fathoms (9 14m).

REPRODUCTION. The sexes are separate, and preliminary examination of the serially arranged
gonads lying on the underside of the dorsal arm surface of females shows numerous small
oocytes with a maximum size of about 120jim diameter. There is also evidence of repro-
ductive seasonality, with spawning in spring (Tyler, Pain & Gage, 1982ft; Tyler & Pain,
1982a). Sibuet & Cherbonnier (1972) describe, with SEM figures, the post-metamorphosal
stages.

REMARKS. Specimens of P. bifrons from St. 'M' contain a small flatworm in the body cavity;
this appears not to affect gonadal development nor to harm the host in any way.

Dy taster ins ignis (Perrier, 1884)
See: Perrier, 1884 : 253, pi. 9, fig. 5; Downey, 1973 : 35-36, pi. 9, figs C, D.

SAMPLES. ES 53 (1), ES 55 (2), ES 56 (1), ES 120 (2), AT 121 (15), ES 129 (1), AT 131 (2), AT 153
(1), AT 181 (1), ES 182(1). SWT 13(3), SWT 27(1). [2170 m to 342 5-3 500m]

DISTRIBUTION. Downey (1973) has synonymised several previously described nominal N
Atlantic species of this exclusively deep-water genus, including D. biserialis Sladen, the
oldest name being D. insignis. This has a wide distribution throughout the N Atlantic deep
sea and possibly extends over the entire Atlantic; on soft deep-sea ooze. Dytaster biserialis
was recorded from the NE Atlantic from depths of 3645 m and 4360 m (Mortensen, 1927).
The present records confirm that this species has an essentially abyssal distribution.

REPRODUCTION. Histological study of the gonads of D. insignis, which are serially arranged
along the underside of the dorsal surface of the arm, suggests a high fecundity, probably in
excess of 106 eggs/individual. Maximum egg size is 120 urn, suggesting indirect, possibly
pelagic, development (Tyler, Pain & Gage, 1982ft; Tyler & Pain, 19820). The latter authors
also indicate that this species shows some form of gametogenic synchrony in the population
sampled.

REMARKS. Species of the genus Dytaster are thought to be omnivorous scavengers; fragments
of algae, foraminiferous worm tubes and molluscs having been found in the stomach
(Downey, 1973) while Bruun & Wolff (1961) give a photograph with a specimen ingesting
a large fish bone.

Psilaster andromeda (Miiller & Troschel, 1842)

See: Sussbach & Breckner, 1911 : 208-209; Grieg, 1921 [1932]: 18-19 (?St. 70); Mortensen, 1927:
59-60, fig. 33; D'yakonov, 1950 : 31.

SAMPLES. ES 23 (3), AT 68A (1), AT 107A (5), AT 186 (3), AT 192 (29; ? Duveniles] 5). GT 2 (1),
GT7(1),GT 11 (5),GT 14(2),GT 15(1),GT 16(4), SWT 18 (7), SWT 27 (4), AT 1 (3). [640-780 m
to 2965 m]

DISTRIBUTION. Previously known from the southern Barents Sea south to Senegal and
from the Azores; 20-1853 m. If P. jlorae Verrill, 1878 proves to be a synonym, then also
from the western N Atlantic. On ooze and other soft muddy sediments.

ROCKALL TROUGH ECHINODERMS I 275

The present records extend the lower limit of the known range of P. andromeda to 2965 m
depth. Our shallowest record, between 640 and 780 m, roughly accords with that of speci-
mens collected by the Helga off the west coast of Ireland (Farran, 1913). In more northerly
latitudes it may extend into considerably shallower depths.

REPRODUCTION. The sexes are separate. Mortensen (1927) suggested that the large yolky eggs
of this species indicate a direct development without a larval stage. Tyler, Pain & Gage
(19826) and Tyler & Pain (\9S2a) have noted a maximum egg size of 950 urn in histological
preparations of the gonad of females. Hence, direct development with a benthic stage seems
likely.

REMARKS. Our specimens were as described by Mortensen (1927), being slimy-skinned and
pale pink or dirty white in colour, although D'yakonov (1950) describes Russian specimens
as a bright red in life.

Psilaster andromeda is known as the host of an unidentified species of the parasitic
barnacle genus Dendrogaster which causes severe regression of the gonads (P.A.T. personal
observation).

Psilaster patagiatus Sladen, 1889

See: Grieg, 1921 [1932]: 19 (as Psilasteropsis patagiatus); Mortensen, 1927: 65-66, fig. 37 (as
Persephonaster patagiatus)', Downey, 1973 : 32, pi. 8, figs A, B.

SAMPLES. ES 15(2), ES 18(3), AT 192(1). SWT 18 (1). [1392-1862 m]

DISTRIBUTION. From the Rockall Trough, Porcupine Seabight (Farran, 1913) to the Azores
and Cape Verde Islands (Koehler, 1909); and in the Gulf of Mexico and Caribbean Sea off
Venezuela where the depths recorded are only 730 m and 915 m; on soft muddy sediments.
Mortensen (1927) gives the lower bathymetric limit as 2165 m.

There is, however, a complication about the limits of this species. Although Downey
(1973) has referred West Indian specimens to P. patagiatus, H. L. Clark (1941) used another
name spinulosus, which he referred to Persephonaster together with patagiatus, for Cuban
specimens from c. 1000 m. He also thought the Azores specimens belonged to a third species.
A.M.C. (MS) has provisionally named as P. patagiatus a Discovery specimen from the
Canaries. Downey (1973) queried the Cape Verde area as type locality for P. patagiatus but
this identification, if correct, would support it.

Family PORCELLANASTERIDAE

Porcellanaster ceruleus Wyville Thomson, 1877

See: Lieberkind, 1935 : 5-19, pi. 2 figs 1-8, pi. 3 fig. 12, pi. 5 figs 16-17; Madsen, 1961 : 1 123-142,
figs 22-24.

SAMPLES. ES 2 (18), ES 4 (5), ES 6 (835), ES 8 (22), ES 10 (101), ES 27 (41), ES 34 (4), SBC 49 (1),
ES 52 (7), ES 55 (28), ES 56 (50), SBC 58 (1), ES 59 (20), ES 1 1 1 (26), ES 1 18 (1 1), AT 121 (6), ES
129 (39), ES 135 (29), ES 137 (53), ES 140 (150), AT 141 (2), ES 143 (12), ES 147 (46), ES 152 (61),
ES 164 (91), ES 169 (68), ES 172 (71), ES 176 (2), ES 180 (110), ES 184 (3), ES 185 (195), ES 190
(63), AT 191 (3), ES 197(2). SWT 13 (1). [1993 m to 3425-3500 m]

DISTRIBUTION. A cosmopolitan deep-sea species, except in Antarctic waters; known from
1 160-6040 m depth; on soft deep-sea ooze.

Individuals of this small species (max. R c. 36 mm, Madsen, 1961) were most numerous
in epibenthic sled hauls from the Permanent Station, decreasing in number in sled hauls
from shallower stations.

REMARKS. The specific name of ceruleus, unnecessarily amended to 'caeruleus ' by Sladen
(1889), was derived from the sky blue colouring of the sediment filling the stomach of the
type specimen from Challenger st. 45, off Delaware, NW Atlantic. The stomachs of this

276 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

deposit- feeding species in the recent samples were likewise filled with mud, but of the khaki
brown colour of the Globigerina ooze characteristic of the deep-sea sediment in the Rockall
Trough.

Hyphalaster inermis Sladen, 1 883
See: Madsen, 1961: 58-72, figs 6, 7, pis 1-3, 13, figs 3, 4.
SAMPLES. ES 147 (? [juvenile] 1). SWT 13(1). [2921 m to 3425-3500 m]

DISTRIBUTION. Previously known from the West European Basin north to c. 47°N, 08°W,
south to the Guinea Basin, also from the Labrador Basin and mid N Atlantic (c. 54°N,
27-28°W) and from the Indo-Pacific; 2280-5430 m; on mud, Globigerina ooze or calcareous
clay.
The present records extend the known range slightly in the NE Atlantic.

REPRODUCTION. Egg size reaches 0-5-0-6 mm with presumably direct development (Madsen,
1961).

REMARKS. The specimen from SWT 13 has R 39 mm, r 19 mm. The specific identity of the
single juvenile from ES 147 remains uncertain.

Order NOTOMYOTIDA
Family BENTHOPECTINIDAE

Benthopecten simplex (Perrier, 1881)
See: Mortensen, 1927 : 74-75, fig. 41 (as B. armatus (Sladen)); A. M. Clark, 1981 : 130-134.

SAMPLES. ES 4 (13), ES 10 (juvenile 1), ES 34 (2), AT 107 A (3), AT 1 14 (5), AT 138 (5), AT 139
(10), AT 144 (117), AT 151 (25), AT 153 (71), AT 154 (49), AT 161 (47), AT 167 (33), AT 171
(110), AT 175 (62), ES 176(56), AT 177(171), AT 181 (98), ES 182 (7), ES 184(16), AT 186(139),
AT 191 (juveniles, 45), AT 192 (93), AT 195 (113), ES 197 (27), AT 198 (20; ? [juveniles], 10), ES
200 (11). SWT 13 (2), SWT 16 (1), SWT 17 (1), SWT 18 (19), SWT 32 (10). [1785-1845 m to
3325-3500 m]

DISTRIBUTION. East of Cape Cod south to the Gulf of Mexico, Colombia and Guyana Basins,
and south of Iceland (c. 62°N, 19°30 'W) and the Rockall Trough south to the Gulf of Guinea;
1 175 m to 2735-3670 m, this lower limit being derived from Pillsbury st. 681, off Surinam.

Our deepest record is from a trawling taken at the deep water confluence of the Rockall
Trough with the Porcupine Abyssal Plain. The most northerly record was from the Feni
Ridge at c. latitude 57°20'N in c. 1860 m depth close to Michael Sars st. 101 where Grieg
(1921) also reported collecting 12 specimens of Benthopecten. Both the present and previous
records strongly suggest a distribution centred on depths of c. 2000 m at temperatures below
about 4°C in the N Atlantic, on soft muddy sediments.

Benthopecten simplex is fairly abundant in Agassiz hauls from st. 'M' and these samples
usually include substantial, though varying, numbers of juveniles with a range of sizes (Pain,
Tyler & Gage, 19826).

Maximum size (R) recorded is c. 130mm (Tyler, Pain & Gage, 19826) whereas that of
the few Sarsia specimens from the Bay of Biscay (E. C. Southward, personal communication)
is only 30-35 mm. Sladen's Challenger expedition specimens, of armatus, are also smaller
than equatorial Atlantic ones where R max. = c. 150 mm (A. M. Clark, 1981).

REPRODUCTION. The sexes are separate and no seasonal periodicity is evident from the histo-
logical comparison of the gonads of specimens taken at different times of the year; maximum
oocyte size is 950 urn, suggesting non-pelagic, possibly lecithotrophic, direct larval develop-
ment (Tyler, Pain & Gage, 19826 as B. armatus; Pain, Tyler & Gage, 19826).

ROCKALL TROUGH ECHINODERMS I 277

REMARKS. Farran (1913) and Mortensen (1927) used the name B. armatus (Sladen, 1889)
while Grieg ( 1 92 1 ) used B. spinosus Verrill ( 1 884). A. M. Clark (1981) believes all NE Atlan-
tic and some NW Atlantic specimens of Benthopecten to be conspecific with the tropical
Atlantic B. simplex (Perrier, 1881) of which armatus is a synonym but B. spinosus, only
known from the NW Atlantic, is a distinct species.

Pectinasterfilholi Perrier, 1885

See: Sladen, 1889: 43-47, pi, 8, figs 3, 4, pi. 12, figs 3, 4 (as Pontaster forcipatus); A. M. Clark,
1981 : 119-121.

SAMPLES. ES 10 (6), ES 34 (7), AT 1 14 (13), AT 138 (3), AT 139 (14), AT 141 (juvenile 1), AT 144
(14), AT 153 (5), AT 154 (11), AT 157 (1), AT 161 (7), AT 167 (3), AT 171 (3), AT 175 (3), ES
176 (1; ? [juvenile] 13), AT 177 (4), AT 181 (5), ES 182 (1), ES 184 (4; ? [juveniles] 14), ES 185 Guven-
iles, 2), AT 186 (1), AT 198 (8). SWT 16 (38), SWT 17 (7). [1752-2909 m]

DISTRIBUTION. Previously recorded (A. M. Clark, 1981) from all four quadrants of the
Atlantic, north to c. 60°N, 20°W (S from Ireland); 1260 m^850 m, the maximum being the
isolated record for the holotype of Pontaster pristinus Sladen, off the River Plate, the deepest
other record being 3430 m. Cherbonnier & Sibuet (1973) cite a minimum depth of 1 152 m
but details of this record have not been traced; on mud or Globigerina ooze.

This species was present in Agassiz trawls from St. 'M'. Elsewhere, adult specimens were
taken from a number of locations in the Rockall Trough and Porcupine Seabight area
between 1752 m and 2540 m. Juveniles were also taken between 2200 m and 2900 m.

Cherbonnier & Sibuet (1973) recorded P.filholi sensu stricto from NW Africa, the Azores
and the southern Bay of Biscay, their record on map 4 of the results of the Noratlante
Expedition being misplaced to west of Brittany, if the station list data are correct. A. M.
Clark (1981) returned Pontaster forcipatus Sladen, 1889, as well as P. pristinus, to the
synonymy of Pectinaster filholi, bringing in records from eastern North America and South
Africa, as well as two Discovery records from the central North Atlantic to the far west of
Ireland and south from Iceland. Although no Helga specimens were recorded by Farran
(1913), a small one was found in the British Museum collections from the sample of
Benthopecten armatus [simplex] from st. SR 944, SW of Ireland.

REPRODUCTION. The sexes are separate; preliminary studies on the gametogenic cycle (Pain,
Tyler & Gage, 1982/7) indicate a maximum oocyte size of 850 um, suggesting a non-pelagic,
possibly lecithotrophic or direct development. Brooding has not been recorded. Individuals
become mature at c. r 3 mm. Size frequencies in the sample time series from St. 'M' indicate
a low, probably year-round recruitment by young of around r 1 mm. Adult size structure
is variable in the samples which suggests a possibly patchy demographic composition on
the bottom (Pain, Tyler & Gage, 19826).

Pontaster tenuispinus (Diiben & Koren, 1 846)

See: Sladen, 1889 : 28-29, 29-30, pi. 6, fig. 7, pi. 7, figs 3, 4 (var. platynota), 35-38, pi. 6, figs 3, 4,
pi. 7, figs 1, 2 (as P. limbatus); Sussbach & Breckner, 1911 : 201-202; Mortensen, 1927: 72-73,
fig. 40.

SAMPLES. ES 23 (4), ABD 25 (1), ES 87 (5 var. platynota), AT 90A (? [juveniles Var. platynota] 3),
AT 1 07 A ( 1 ). GT 1 (4), GT 7 (4), GT 1 3 ( 1 ), GT 1 4 (2), GT 1 5 (4), AT 1 (74), AT 3 (2). [508-543 m
to 2000 m]

DISTRIBUTION. Circum-arctic extending south in the Atlantic to the latitude of Cape Cod
in the west and the eastern slope of the Bay of Biscay (c. 44°N) in the east, 1 6-c. 2000 m,
(2330m according to Perrier, 1894), rarely <200m, except in arctic waters. The records
of Farran (1913) are from c. 400-800 m in the Porcupine Seabight. The Porcupine record
(Sladen, 1889) from 100-150 fathoms, 1 83-274 m, off Valentia, SW Ireland is surprisingly
shallow. The round numbers, in fathoms, quoted suggest that this record might be regarded

278 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

with suspicion. Found on soft mud, muddy sand or clay, sometimes fine sand or sand with
shell.

Specimens of P. tenuispinus forma tenuispinus were collected from hauls on the Hebridean
Slope from 523 m to 1000 m depth and one only from the North Feni Ridge area in 2000 m,
whereas others with basally broadened rays conforming to Sladen's variety platynota were
identified in the 'cold' area north of the Wyville Thomson Ridge, the Faroe-Shetland
Channel in 1040 m and 1050 m depth. The five specimens from ES 87 (61°13 'N, 03°59'W)
ranged from R 35 mm to R 5-8 mm. In the smallest specimen however, the bivalved adambu-
lacral pedicellariae are not developed.

REPRODUCTION. The sexes are separate; maximum oocyte diameter is 800 um, suggesting
a non-pelagic possibly lecithotrophic or direct development (Pain, Tyler & Gage, 19826).

Order VALVATIDA
Family ODONTASTERIDAE

Hoplaster spinosus Perrier, 1882

See: Perrier, 1894 : 324; Sladen, 1889 : 275 (as Pentagonaster lepidus).
SAMPLES. AT 121 (3), AT 167(1). [2910m, 2300 m]

DISTRIBUTION. Previously known from the vicinity of the Azores and in the Porcupine
Seabight, recorded in an appendix by Mortensen 1927 : 439; 1800-2995 (73307) m; soft mud
or ooze.

The present records extend the known range north to 57°. The largest specimen, from the
2910 m haul, measured R 14 mm, whereas in the very few specimens previously described
R did not exceed R 7-5 mm (Mortensen, 1927 : 77).

REMARKS. The monotypic genus Hoplaster does not have the reflected hyaline-tipped oral
spines otherwise characteristic of the family Odontasteridae but does show the distinctive
odd interradial marginal plates.

Odontaster sp.

See: von Marenzeller, 1893 : 6, figs 4A-C (Gnathaster medi terra neus); Mortensen, 1927 : 77-78, figs
42, 43 (? O. mediterraneus); Verrill, 1899 : 206-207, pi. 29, fig. 3 (O. hispidus).

SAMPLE. AT 162(1). [992m]

REMARKS. The single (dried) specimen, from the N slope of the Porcupine Seabight, has R/r
29/11-5 mm = 2-5/l and a relatively narrow frame of marginal plates. The Atlantic speci-
mens recorded as O. mediterraneus by Koehler (1909) from the Bay of Biscay (Princesse
Alice st. 1450, c. 45°N, 3°W, 1804 m, R 15 mm) and Mortensen (1927) from the Porcupine
Seabight (Helga, 51°26 'N, 1 1°45 'W, 927 m, 'small'), as well as a record from the Seabight
(unpublished, IOS st. 50601, 51°20'N, 1 1°42'W, 927 m, R/r 20/13 = 1-5/1), all have more
nearly pentagonal form and disproportionally broad marginals, though this is probably at
least partly attributable to the relatively small size. Odontaster hispidus Verrill, 1880 has
only been recorded from the NW Atlantic in 73-1 160 m. The body shape is very variable
in Odontaster and the mean R/r ratio for both species is c. 2-5/1 in larger specimens,
R > 20 mm. The main differences in mediterranean specimens appear to lie in broadened
paxillar columns on the distal abactinal plates, frequent rather than occasional glassy 'crystal
bodies' embedded in the distal superomarginals, both visible when the plates are denuded,
short capitate superomarginal spinelets and three or four rather than two, occasionally three,
furrow spines. The present specimen has the distal abactinals no broader than the proximal
ones, no crystal bodies and tapering slender superomarginal spinelets. However, it does have
four furrow spines on a few plates, otherwise three; also the actinal armament is less crowded

ROCKALL TROUGH ECHINODERMS I 279

than is usual in hispidus, though the dry condition may contribute to this. In the IOS speci-
men the abactinal areas are not at all prolonged on to the rays and the more distal plates
still have round columns, there are no crystal bodies on the marginals and the furrow spines
usually number three but the superomarginal armament is very short and slightly capitate.
Clearly further material and comparison is needed for the specific identity of these specimens
to be established.

Family RADIASTERIDAE

? Radiaster tizardi (Sladen, 1 882)
See: Mortensen, 1927 : 97, fig. 55.
SAMPLE. ES 10 (juvenile, 1). [2540 m]

DISTRIBUTION. Although according to Mortensen (1927) this species is previously known
only from the Faroe- Shetland Channel (Triton and Knight Errant) and the Porcupine
Seabight (Helga) in 930 (? 730)m-1320 (? 1630) m, Sladen (1882) gives the single Knight
Errant record as from 59°33 'N, 07° 14 'W in 1015m depth on the 'warm' side of the Wyville
Thomson Ridge in the northern Rockall Trough; mud or ooze.

REMARKS. The single specimen measures R = 3-5 mm, r = 2-0 mm. The abactinal arma-
ment appears clustered and the margins thick. The present record must be uncertain in view
of the small size of the specimen, specimens of tizardi are not yet known with R < c. 60 mm,
and the considerable depth (2540 m) from which it was collected.

Family GONIASTERIDAE

Pseudarchaster parelii (Diiben & Koren, 1846)

See: Sussbach & Breckner, 1911 : 202-203 (as Tethyaster parelii); Mortensen, 1927: 87-88, fig. 49;
Halpern, 1972: 366-370.

SAMPLES. ES 55 (? Duveniles] 9), ES 56 (? [juveniles] 2), ES 87 (? [juvenile] 1), AT 1 14 (1), AT 132
(1), AT 138 (1), AT 139 (2), AT 144 (1), AT 151 (1), AT 154 (1), AT 167 (1), AT 175 (1), ES 176
(1), AT 177 (2), AT 181 (1), ES 184 (1), AT 191 (10), AT 192 (1), AT 195 (2), AT 198 (juvenile,
1), ES 200 (1). GT 1 (2), GT 7 (2), GT 13 (1), GT 14 (2), SWT 10 (2), SWT 1 1 (1), SWT 16 (4),
SWT 17(1), SWT 1 8 (8), SWT 27(1). [225-2965 m]

DISTRIBUTION. Previously known from Greenland south to the West Indies in the western
Atlantic, and from northern Norway to the Bay of Biscay and the Azores in the east;
15-3000 m, southern records being from deep water; preferring waters with a temperature
slightly above zero (D'yakonov, 1950) or from 4-8°C (Cherbonnier & Sibuet, 1973)
and l-5-6°C (Halpern, 1972); on bottoms of soft mud to sand, sometimes with pebbles
(D'yakanov. 1950).

Our present records from the Rockall Trough range from 225 m on the Porcupine Bank
to 2965 m in the vicinity of the Permanent Station, where juveniles only were collected, and
hence apparently show an ability to tolerate temperatures from 2 to 5°C up to c. 9-5-10°C
from hydrographic data in Ellett & Martin (1973).

REPRODUCTION. Preliminary examination of the gonads of some of these specimens indicates
that the sexes are separate, while the presence of large oocytes (c. 1000 urn) suggests direct
or demersal development (Tyler, Pain & Gage, 19826; Tyler & Pain, 19826).

REMARKS. The colour of fresh specimens was a rose pink; Mortensen (1927) and D'yakonov
(1950) describe freshly collected specimens as bright red to red-brown on the dorsal side.

Pseudarchaster gracilis (Sladen, 1889)
See: Halpen, 1972 : 360-366, figs 1,2; Downey, 1973 : 59-60, pi. 23, figs C, D.

280 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

SAMPLE. SWT 18 (1). [1785-1845 m]

DISTRIBUTION. Previously known from just south of Cape Cod and from the Gulf of Mexico
to Surinam, in the W Atlantic; from the Azores and from Mauritania to the Gulf of Guinea
in the east; 320-2736 m. (Only in the West Indies area from < 1000 m).

This record on the Hebridean slope extends the known distribution of P. gracilis to British
waters. Although it is known to occur on both sides of the North Atlantic, the most northerly
of previous records in the east were from the Azores.

Paragonaster subtilis (Perrier, 1881)
See: Halpern, 1972 : 374-378, figs 5 (pt), 6 (pt); Downey, 1973 : 57, pi. 22, figs A, B.

SAMPLES. ES 6 (juvenile, 1), ES 27 (1), ES 28 (2), ES 34 (1), ES 53 (2), ES 56 (1), ES 111 (1), AT
121 (17), ES 129 (2), AT 131 (2), AT 138 (1), AT 139 (1), ES 140 (2; ? [juvenile] 1), AT 141 (6),
ES 147 (1), ES 164 (1), AT 171 (juveniles, 2), AT 177 (juvenile, 1), ES 180 (3), AT 186 (5). SWT
16 (17), SWT 17 (5), SWT 18 (4), OTSB 51001 (1). [1785-1845 m to 2925 m]

DISTRIBUTION. Previously known from south of Cape Cod and the Gulf of Mexico, in the
W Atlantic, the Azores and mid-equatorial Atlantic, and from the Bay of Biscay, C. Verde
Is. area and Gulf of Guinea in the east; 2455^4700 m; soft ooze. This N Atlantic species
has an abyssal distribution not previously known to extend further north than Biscay in the
NE Atlantic.

The present records, although mostly from the vicinity of the Permanent Station (c.
2900 m depth) confirm the essentially abyssal distribution of P. subtilis, although the
shallowest record of 1785-1 845m from 56°46'N on the Hebridean slope is less than the
minimum depth of 2455 m quoted by Halpern (1972). The temperature range quoted by
Halpern is 1-5^°C.

REPRODUCTION. The sexes are separate with a maximum egg size of about lOOOum in
females, suggesting direct development (Tyler, Pain & Gage, 19826; Tyler & Pain, 19826).

Plinthaster dentatus (Perrier, 1884)
See: Halpern, 1970 : 244-252, figs 17, 18, 19; Downey, 1973 : 52-53, pi. 19, figs A, B.

SAMPLES. ES 59 (juvenile, 1), AT 68A (2), ES 169 Guvenile, 1), ES 185 Guvenile, 1) SWT 18
(1). [1331-2190 m]

DISTRIBUTION. Previously known from both sides of the north and central Atlantic, from
North Carolina to northern Brazil in the west and from the west of Ireland to Liberia in
the east and from the Azores; 229-2 1 1 7 m; on muddy sediments.

Although previously collected in the Porcupine Seabight (Farran, 1913; Grieg, 1921), the
present records extend the known northward distribution of this N Atlantic species to lati-
tude 59° and extend the lower limit of its depth range from 1804 m (Halpern, 1970) in the
eastern Atlantic to 2910 m.

Evoplosoma scorpio Downey, 1981
See: Downey, 1981 : 561-563, fig. 1.
SAMPLE. ES 112(3). [1900m]

DISTRIBUTION. Previously only known from the holotype taken by the Sarsia off the Western
Approaches to the English Channel (c. 48°N, 10°W); c. 1600 m (Downey, 1981).

The new record provides a northward extension of range to the Feni Ridge in Rockall
Trough. Two other species of Evoplosoma are known, one from the Pacific near Hawaii,
the other south of Sri Lanka in the Indian Ocean.

REMARKS. The present specimens are large and robust, R 90-130 mm, R/r 3-3-3-6/1.
Evoplosoma is closely related to Hippasteria but differs from H. phrygiana in the long

ROCKALL TROUGH ECHINODERMS I 281

narrow arms (R/r c. 3-5/1), continuous irregular granulation interspersed with conical spines
and pedicellariae with the two valves elongate, projecting and spatuliform rather than
extremely short but very broad and almost flush with the body surface, as in Hippasteria.

Family PORANIIDAE

Porania pulvillus (O. F. Miiller, 1766)

See: Doderlein, 1900: 217-218, pi. 8, figs 10, lOa; Sussbach & Breckner, 191 1 : 218-219; Mortensen,
1927 : 90-92, fig. 51.

SAMPLES. ES 33 (1), ES 113 (1, juveniles, 2). [148 m, 168 m]

DISTRIBUTION. Previously known from Scandinavia, the Lousy Bank and west of Ireland
south to the Bay of Biscay; 10-1000 m but rarely deeper than 300 m, on the usually mixed,
sandy sediments of commercial trawling grounds.

The present records span the Rockall Trough and fall within the known bathymetric range
although P. pulvillus apparently has not been previously recorded from Rockall Bank (ES
113).

REMARKS. The two small specimens (R c. 6-5 mm) from ES 1 13 resemble the holotype of
Marginaster fimbriatus Sladen, 1889 from Porcupine st. 31, c. 56°N, 1 1°30'W, 2487 m, in
the prominent inferomarginal fringe but have relatively sparse abactinal and actinal arma-
ment, only a single actinal spinelet in each interradius. Marginaster fimbriatus was referred
to the synonymy of the Mediterranean M. capreensis (Gasco) by Ludwig, 1897 followed
by Mortensen (1927) (see fig. 54) but A.M.C. believes it more likely to be the juvenile form
of another poraniid from this vicinity. The large specimen from the same station, R 55 mm,
is unusual in having the inferomarginal spines mostly aborted, there being only one or some-
times two on five to seven proximal plates of each interradial arc.

Order SPINULOSIDA
Family PTERASTERIDAE

Pteraster militarised. F. Miiller, 1776)

See: Sussbach & Breckner, 1911: 226; Mortensen, 1927 : 104-105, fig. 60.
SAMPLES. ES 87 (juveniles, 2). GT 2 (2), GT 1 1 (1). [934-1054 m to 1050 m]

DISTRIBUTION. This is an arctic-boreal species with a circumpolar distribution in the Arctic
and a NW and NE Atlantic distribution along the American coast to Cape Cod, off the north
west of Scotland and in the Faroe Channel, and off the entire west coast of Norway
(D'yakonov, 1950); 10-1 100m.

The present records are all from around 1000 m and extend the known distribution slightly
(and predictably) to the Rockall Trough.

Pteraster pulvillus M. Sars, 1861

See: Sussbach & Breckner, 1911 : 227; Mortensen, 1927 : 103; D'yakonov, 1950 : 81.
SAMPLES. ES 1 13 Guvenile, 1), ES 1 15 (? [juvenile] 1). [168 m, 1000 m]

DISTRIBUTION. This circumpolar N boreal species is known to range southward along the
Atlantic coast of N America to 42°N (Massachusetts Bay and Bay of Fundy) and on the
Norwegian coast down to 60°N in the vicinity of Bergen (D'yakonov, 1950). The record from
Rockall Bank together with that recently recorded on the West Irish shelf in 122 m depth
(O'Connor, 1981), confirm Mortensen's (1927) prediction that this species would be found
in areas round the British Isles; 36-3700 m (202 1 fins [= 3696 m] according to Sladen, 1 889).

282 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

REMARKS. The specimen from ES 115 is in poor condition and has the stomach everted,
damaging the oral furrow armament so that the characteristic continuous webbing of the
oral furrow spines across the apex of each jaw has been ruptured, if it was present. The good
specimen from ES 1 1 3 has R/r 8/6 mm.

Pteraster reductus Koehler, 1 907

See: Koehler, 1909 : 96-97, pi. 3, figs 8, 9, pi. 20, figs 10; Grieg, 1921 : 28-29, pis 5, 6, 7; Mortensen,
1927: 103.

SAMPLE. ES 1 1 2 (juvenile, 1 ). [1 900 m]

DISTRIBUTION. This species is known only from the Azores and a single specimen from the
South Feni Ridge in 2215 m (Cherbonnier & Sibuet, 1973); 1920-2900 m.

It is interesting that the present specimen was also collected from the Feni Ridge at a depth
nearly within the previously recorded bathymetric range.

Pteraster sp. aff. P. acicula (Downey, 1973)
See: Downey, 1973 : 79, pi. 34, figs C, D.
SAMPLE. GT 16(1). [958m]

DISTRIBUTION. P. acicula has only been recorded from a few small poorly- preserved speci-
mens (R up to 8 mm) from the northern Gulf of Mexico in 1 060-1 3 1 6 m, so the identification
of the Rockall Trough specimen remains to be confirmed.

REMARKS. The complete absence of webbing from the oral furrow spines, characteristic of
the subgenus Apterodon, and the very large number of paxillar spinelets (c. 25 on one
midradial paxilla) distinguish this specimen from all the NE Atlantic Pterasters hitherto
recorded.

HYMENASTER Wyville Thomson

At present there is some confusion over the taxonomy of Hymenaster in the N Atlantic.
We here retain the specific names membranaceus, pellucidus, gennaeus and rex according
to the original identifications of the specimens made from Sibuet's tabular key to the Atlantic
species (1976), pending a forthcoming comprehensive review of the family by M. E. Downey,
in Clark & Downey (in prep.). A.M.C. dissociates herself from the identification of N Atlantic
specimens as H. gennaeus, believing that Sibuet (1976) had a mistaken concept of H.
giganteus as a membranous rather than a fleshy species as Farran (1913) describes the type
and Helga specimens.

Hymenaster membranaceus Wyville Thomson

See: Sladen, 1889: 521-2, pi. 92, figs 6, 7, pi. 93, figs 10-12; Mortensen, 1927: 106; Sibuet, 1976:
315-318, fig. 6 A, B.

SAMPLES. ES 12 (2), AT 141 (juvenile, 1), AT 144 (287), AT 151 (30), AT 153 (203), AT 154 (111),
AT 161 (3), AT 167 (137), AT 171 (556), AT 175 (180), ES 176 (60), AT 177 (319), AT 181 (354),
ES 182 (16), ES 184 (79), AT 186 (423), AT 191 (? [juveniles] 47), AT 195 (410), ES 197 (37), AT
198(133), ES 200 (15;? [juveniles] 4). SWT 32 (1). [1995-2020 m to 2909 m]

DISTRIBUTION. Known from the E, N and S Atlantic from a depth range of 1 000-3000 m;
on soft ooze. Results of recent intensive sampling in the Bay of Biscay indicate a somewhat
restricted bathymetric range (21 19-2878 m, Sibuet, 1976).

H. membranaceus was collected in large numbers from St. 'M' (c. 2200 m depth), apart
from a single juvenile specimen from the Permanent Station (2909 m), a sample of three
from the Porcupine Seabight (c. 2055 m) and two records (2; 1 specimen) from the Barra
Fan in the northern Rockall Trough (2076 m; 1995-2020 m).

ROCKALL TROUGH ECHINODERMS I 283

REPRODUCTION. The eggs of this species grow to 1 1 00 um diameter, suggesting direct devel-
opment (Pain, Tyler & Gage, 1982a). These authors also indicate that there is no synchrony
in gametogenic development in the population, and that breeding probably occurs year-
round at a low level. Size frequency data from St. 'M' suggest a low possibly continuous
recruitment to the population which consists mainly of mature adults. The sexes are separate
and, although fertilization may take place in the supradorsal chamber, there is no evidence
of brooding in the St. 'M' population.

REMARKS. H. membranaceus is thought to live partially buried in bottom sediment, with
only the valves of the central opening protruding above the sediment in order to allow venti-
lation of the supradorsal cavity (Mortensen, 1927). However, we note here that in photo-
graphs of the bottom in the Porcupine Seabight in Rice et al. (1982), specimens of
'Hymenaster sp.', that are probably the same as the present species, are shown lying on the
sediment surface. Possibly these animals were in the process of changing station where they
would normally be buried out of sight.

Hymenaster pellucidus Wyville Thomson, 1873

See: Danielssen & Koren, 1884: 72-80, pi. 13, figs 1-17, pi. 15, figs 7-8; Grieg, 1921 [1932]: 29;
Mortensen, 1927: 107- 108, figs 62, 63 1,2.

SAMPLE. ES 87(1). [1050m]

DISTRIBUTION. Previously known from the Siberian and Norwegian Seas, extending to the
Faroe Channel in c. 15-2800 m.

This very delicate, perfect specimen, R 22 r 15 mm, R/r 1-5/1, is from the Faroe-Shetland
Channel.

Hymenaster gennaeus H. L. Clark, 1923
See: H. L. Clark, 1923 : 302-303, pi. 10; Sibuet, 1976 : 31 1-314, figs 4, 5B.

SAMPLES. AT 144 (1), AT 154 (1), AT 171 (1), AT 181 (1), AT 195 (? 1). SWT 11 (1). [2190m
to 2500-2560 m]

DISTRIBUTION. Known previously only from the holotype (Clark, 1923) taken by Pieter
Faure in the SE Atlantic near Cape Point in 1400 m and one specimen from c. 47°35 'N,
8°39 'W in 2245 m (Sibuet, 1976) but see note on p. 282.

Our records are from St. 'M' c. 2200 m, except for one specimen from 2500-2560 m at
c. 56°N, 1 PW that was taken in a fish trawl.

REMARKS. The queried specimen from AT 195 is probably not conspecific with the rest,
having a subpentagonal rather than pentagonal body form, R/r nearer 2/1 than 1-5/1, con-
sistently single paxillar spines rather than usually three, but three rather than usually one,
sometimes two adambulacral spines. It agrees better in these characters with H. regalis
Verrill, 1 895, known from Carolina to Cuba, but that species is supposed to lack the smooth
interbrachial arcs found in this specimen.

REPRODUCTION. Preliminary observations of Pain, Tyler & Gage (1982#) indicate an egg size
and mode of development similar to that of H. membranaceus.

Hymenaster rex Perrier, 1 894

See: Perrier, 1894: 186-189, pi. 13, fig 2; Grieg, 1921 [1932]: 29-30.
SAMPLE. AT 121 (1). [2910m]

DISTRIBUTION. Previously known from the Bay of Biscay (c. 46°N, 07°W) south to off NW
Africa, 1 140-2285 m, so the present record provides extensions northwards and downwards.

REMARKS. This specimen has R/r 1 1/8-5 mm, the form being almost pentagonal. There are
3 adambulacral and oral furrow spines and 2 suboral spines on each plate.

284 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

Family SOLASTERIDAE

Crossaster squamatus (Doderlein, 1900)

See: Doderlein, 1900: 208-209, pi. 6, figs 5-5c (as Solaster papposus var. squamata); Greig, 1921
[1932]: 26-27 (as Solaster squamatus); Mortensen, 1927: 1 14 (as S. squamatus).

SAMPLE. ES 87 (9). [1050m]

DISTRIBUTION. Known from west of Greenland to the Barents Sea, including most of the
cold area of the Norwegian Sea south to the Faroe Channel (c. 61°N, 4°30'W, M. Sars)\
100-1600 m. Previous records from the Faroe Channel area are all c. 1000 m.

Family ECHINASTERIDAE
HENR1C1A Gray

The treatment of the genus Henricia provisionally follows Mortensen (1927) pending further
study by A.M.C. in Clark & Downey (in prep.).

Henricia sanguinolenta (O. F. Miiller, 1776)

See: Sussbach & Breckner, 1911 : 224-226 (as Cribrella sanguinolenta); Mortensen, 1927: 118-120,
fig. 70.

SAMPLE. ES 23(1). [704m]

DISTRIBUTION. Owing to confusion about the specific limits, it is impossible to give a precise
range. Henricia sanguinolenta sensu stricto appears to range from the north east of the
U.S.A. to Iceland, the Faroes, northern British Isles, Scandinavia and the White Sea in depths
possibly from c. 10-1000 m.
The present record (R/r 57/13 mm) of this coastal species is from the Hebridean Slope.

Henricia abyssicola (Norman, 1869)
See: Mortensen, 1927 : 120-121, fig. 71.
SAMPLES. ES 99 (2), ES 1 12 Guvenile, 1), ES 1 13 (juvenile, 1). [168-1900 m]

DISTRIBUTION. Previously recorded off SW Ireland (Helgd) and from the 'warm' area of the
Faroe Channel (Triton) in c. 930-1400 m.

The present records lie at widely varying localities and depths; two are from within the
Rockall Trough, while one is from Rockall Bank (ES 113). They are considerably shallower
than previous records of this species, which is apparently not recorded from localities other
than off the British Isles.

Order FORCIPULATIDA
Family BRISINGIDAE

Brisinga endecacnemos Asbjornsen, 1856
See: Farran, 1913 : 27-28; Mortensen, 1927 : 125-127, fig. 73.

SAMPLES. AT 107A (1), AT 151 (1), AT 181 (1), AT 186 (8), AT 192 (14), ES 200 (2). SWT 17
(12), SWT 18(1), SWT 32 (13). [1785-1845 m to 2220m]

DISTRIBUTION. Previously recorded from the Faroe Channel and southern Norway, south
to the Cape Verde area and from the Porcupine Seabight, the last by Farran (1913);
286-2200 m.

FEEDING. Bottom photographs and submersible observations have led to speculation on the

ROCKALL TROUGH ECHINODERMS I 285

mode of life of members of this very singular deep-sea family. A bottom photograph in Rowe
& Staresinic (1979) from the Tongue-of-the-Ocean (Bahama Is.) at 2000 m shows an asteroid
(with arms raised up in a filter feeding posture) which probably belongs to this family.
Pawson (1978) gives photographs of brisingids in a similar posture and comments that they
are presumably capturing particulate matter from the water and transporting it to the mouth
by ciliary currents.

REPRODUCTION. The oocytes grow up to 1250 urn in diameter (P.A.T., personal obser-
vations); this suggests a direct mode of reproductive development.

REMARKS. This species is thought to prefer a rocky bottom (Mortensen, 1927). Some of
the present records of this very large species are of disks only. This is unfortunate, as the
serial gonads in the arms ofBrisinga endecacnemos provide the most obvious distinguishing
feature from Brisingella coronata.

Brisingella coronata (G. O. Sars, 1871)
See: Ludwig, 1897 : 418; Mortensen, 1927 : 127.
SAMPLES. AT 138 (1), AT 162 (5), AT 167 (4), ES 176 (1), AT 177 (1). [992-2450 m]

DISTRIBUTION. Recorded from the Rockall Trough by Grieg (1921) and off northern Norway
south to the Cape Verde Is. area and the Azores; 100-2600 m.

The present records at 992 m in the Porcupine Seabight, where it has previously been
collected by Farran (1913), and at St. 'M' hence are not unexpected.

REPRODUCTION. The eggs grow up to 1200 um diameter (P.A.T., personal observations); this
suggests direct development.

Freyella spinosa Perrier, 1894
See: Perrier, 1894 : 85-89, pi. 8; Mortensen, 1927 : 129.
SAMPLE. SWT 15 (6). [4810m]

DISTRIBUTION. Previously known from the Bay of Biscay south to the Azores and Angola;
1 884-4060 m.

The present record extends the range northwards into the Porcupine Abyssal Plain and
downwards to 48 10 m depth.

REMARKS. Pawson (1978) mentions that he has observed from the submersible Alvin a species
of Freyella feeding with '. . . arms upraised, and it tends to raise itself as far as it can into
the water column by climbing to the tops of nearby rocks.'

The stronger articulation at the base of each arm ensures that most specimens of Freyella
are preserved with at least the proximal parts of some arms still attached to the disk, unlike
Brisinga and Brisingella. F. spinosa has 8-13 arms.

Freyella sexradiata (Perrier, 1885)

See: Perrier, 1894 : 89-90, pi. 3, fig. 2; Grieg, 1921 [1932]; 30-31, fig. 10; Mortensen, 1927 : 129.
SAMPLE. SWT 1 5 ( 1 ). [4810m]

DISTRIBUTION. Previously recorded from the Bay of Biscay south to the latitude of Gibraltar
(c. 35°N) and west to the Azores; 4020-51 10 m.

The single specimen collected in the Porcupine Abyssal Plain along with F. spinosa like-
wise extends the known range northwards from the Bay of Biscay.

Family ASTERIIDAE

Hydrasterias sexradiata (Perrier, 1882)
See: Perrier, 1894 : 100-102, pi. 9, fig. 2 (as Pedicellaster sexradiatus).

286 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

SAMPLES. AT 1 38 (arms), AT 1 39 (arms), AT 1 54 (7), AT 1 67 ( 1 ), ES 1 76 (juvenile, 1 ). [2245-2450 m]

DISTRIBUTION. Previously known from the Bay of Biscay south to the Cape Verde Is. area
and the Azores; 600-4260 m.

This record extends the known distribution of this species northward in the NE Atlantic
to the Rockall Trough.

ONTOGENY. The juvenile specimen is at the armless 'Stellosphaera mirabilis' stage of Perrier
(1894). The identification has been confirmed by Miss M. E. Downey of the Smithsonian
Institution, Washington, U.S.A., partly on the basis of the structure of the crossed pedicellar-
iae, which are much more compact than those of Freyella sexradiata, the only other six-
rayed forcipulate known from this vicinity.

Stichastrella rosea (O. F. Muller, 1 776)
See: Siissbach & Breckner, 1911: 228-229 (as Stichaster roseus); Mortensen, 1927 : 136, fig. 77.

SAMPLES. ES 15 (1 var. ambigua), ES 33 (2), ES 1 13 (4; juvenile, 1). GT 1 (2 var. ambigua), GT
7 (1 var. ambigua), GT 14 (3). [168-1632 m]

DISTRIBUTION. Previously known from the Lofoten Is., Norway and the north and west of
the British Isles south to the Bay of Biscay; 4-1 150 (? 1330) m.

The present records are from the Hebridean Slope, adjacent shelf and the Rockall Bank
and show a considerable depth range. Mortensen (1927) cites records of this species known
only from the NE Atlantic from 4 m to 430 m depth, while in 586 (?-393) m-1 146 (? 1 33 1 ) m
in the Porcupine Seabight Farran (1913) records a variety ambigua with shorter and stouter
arms which differs also in the even more regular arrangement of the dorsal skeletal plates
than is present in S. rosea sensu stricto, itself a relatively regular asteriid. On this account
together with the bathymetric difference, A.M.C. considers that a specific rank for ambigua
may prove to be more appropriate. The problem will be considered in Clark & Downey
(in prep.).

REMARKS. The juvenile specimen from St. ES 113 has R only 3 -9 mm, R/r 1-6/1. The
relatively large semicircular terminal plates are 1 -3 mm broad and each bears a conspicuous
horizontal fan of fourteen flat petaloid spines similar to the two spines on each of three
ambital inferomarginals in each series, the first inferomarginal being spineless and the fifth
and distalmost with a single smaller spine.

Family ZOROASTERIDAE

Zoroaster fulgens Wyville Thomson, 1873

See: Farran, 1913: 19-22, pi. 1, fig. 3 (as Z. fulgens var. Ackleyi); Mortensen, 1927: 132-133,
fig. 75.

SAMPLES. ES 10 (3), ES 18 (3), ES 20 (1), ES 34 (26), AT 107A (2), AT 114 (17), AT 138 (9), AT
139 (18), AT 171 (7), AT 175 (3), AT 177 (9), AT 181 (3), ES 182 (1), AT 186 (7), AT 191 (juveniles,
11), AT 192 (7), AT 195 (1). GT 17 (2), SWT 10 (1), SWT 11 (2), SWT 12 (1), SWT 13 (1), SWT
15(1), SWT 16(366), SWT 17(128), SWT 18 (21), SWT 27 (2), AT 3 (2). [c. 1000-4810 m]

DISTRIBUTION. From its original discovery by the Porcupine in the N Rockall Trough in
1869 (W. Thomson, 1873), this species has been found from hauls on soft ooze all over the
deep N Atlantic.

The deepest of the present records (SWT 15) from the Porcupine Abyssal Plain extends
the previously known bathymetric range of 367-3660 m (Downey, 1973) to 4810 m. How-
ever, if Z. longicauda Perrier, 1885, from NW of Africa and the Azores, is conspecific with
Z. fulgens, as Downey (1970) notes is probable, then the previously known depth range must
be extended to 4255 m. Specimens were numerous in the vicinity of St. 'M' at depths around
2200 m. The shallowest station was on the adjacent Hebridean Slope.

ROCKALL TROUGH ECHINODERMS I 287

A photograph showing Z.fulgens in situ on the bottom was taken during a RRS Challenger
haul ate. 49°22'N, 12°49'Win 1398-1404 m depth (Rice et al, 1982; fig. 2(e)).

REPRODUCTION. Z. fulgens has a maximum egg size of 950 um (P.A.T., personal obser-
vation).

REMARKS. Specimens examined from St. 'M' occasionally contained one or more specimens
of an unidentified species of the parasitic ascothoracid barnacle, Dendrogaster (P.A.T., per-
sonal observations).

Class OPHIUROIDEA

Suborder EURYALIDA

Family ASTERONYCHIDAE

Asteronyx loveni Miiller & Troschel, 1842

See: Sussbach & Breckner, 1911 : 262; Mortensen, 1927 : 158-160, fig. 90; D'yakonov, 1954 : 21-22,
fig. 4.

SAMPLES. SWT 18 (1), SWT 32 (1). [1785-1845 m to 1 995-2020 m]

DISTRIBUTION. This is a widespread species found in all the major oceans except the Arctic;
previously recorded bathymetric range, c. 100-2963 m. Mortensen (1927) lists several
records around the coast of Scotland and Grieg (1921) took a specimen from the Porcupine
Seabightat 1797m.

The present specimens are from the Hebridean Slope and thus extend the known distri-
bution into the Rockall Trough.

REMARKS. The long prehensile arms and climbing habit on gorgonians and pennatulids is
typical of euryalids. One specimen collected was entwined in characteristic posture around
the pennatulid Distichoptilum gracile Verrill. Grassle, Sanders, Hessler, Rowe & McLellan
(1975) observed Asteronyx loveni from the submersible Alvin at c. 1800m depth off New
England, the ophiuroid being present coiled in the branches of every gorgonian seen.
Hartman (1963) photographed A. loveni on the ocean floor using its long arms to feed from
the water column above. Mortensen (1927), in notes on the biology of this species, states
that adults are plankton eaters; two or three arms are wound around the pennatulid, while
the others wave freely in the water catching the small pelagic animals, mainly copepods,
on which it feeds. Mortensen also thought that this species may feed on the polyps of the
pennatulid to which it clings.

Astrodia tenuispina Verrill, 1884

See: Verrill, 1884: 219; Koehler, 1907 : 304, pi. 21, figs 48-50.
SAMPLE. SWT 13 (147). [3425-3500 m]

DISTRIBUTION. Previously recorded from the N Atlantic deep-sea region; 2365-3300 m.

This unusually large sample was recovered from the deep water connecting the southern
Rockall Trough with the Porcupine Abyssal Plain, so being the first record for this species
in the deep sea around the British Isles and from a greater depth than previously recorded.

REMARKS. Most specimens when examined were entwined around the pennatulid
Scleroptilum grandiflorum Kolliker.

The stomach was devoid of recognisable contents except in one of ten individuals
examined. In this specimen, remains of the uropods of a mysid crustacean were found. It
seems likely that this species feeds on plankton in a manner similar to that which Mortensen
(1927) supposed for Asteronyx loveni.

288 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

REPRODUCTION. Measurements of disk diameter show a unimodal frequency ranging from
5-25-1 1-25 mm, peaking in the 7-50-7-75 size interval. Of the ten individuals dissected, all
appeared to have female interradial gonads, each gonad containing mainly large yolky-
looking eggs. Fecundity, expressed as the number of visible gonads, may vary widely between
individuals. In one specimen, the gonads extended dorsally over the large stomach.

Family ASTEROSCHEMATIDAE

Asteroschema inornatum Koehler, 1 906
See: Koehler, 1909 : 206, pi. 3, figs 46, 47.
SAMPLE. ES 112(1). [1900m]

DISTRIBUTION. Known only from the NE Atlantic deep sea; 1480 m (Mortensen, 1933 : 114).
The present record from the Feni Ridge extends the bathymetric and northern geographic
range of this species.

REMARKS. The single specimen was entwined in characteristic fashion on a gorgonian pro-
visionally identified as Paramuricea biscaya Grasshoff, 1977.

Family GORGONOCEPHALIDAE

Gorgonocephalus caputmedusae (Linnaeus, 1758)
See: Mortensen, 1927 : 162-163, figs 91, 92.
SAMPLES. GT (1), AT 3 (1). [972-1084 m]

DISTRIBUTION. West of the British Isles this species has been recorded off Corn wall, off SW
Ireland and Lousy Bank; 1 1 16-1214 m. Elsewhere it is known from scattered localities all
over the N Atlantic; full bathymetric range c. 150-1200 m.
The present samples were recorded from the Hebridean Slope west of Barra.

REMARKS. The recovery of a single specimen entangled around a sounding line cast to 800
fathoms (1463 m) in Baffin Bay during Sir John Ross' attempt to find the North West Passage
represents probably the first ever record of a deep-sea animal (Menzies, George & Rowe,
1973). Gorgonocephalus caputmedusae uses its branched arms to ensnare plankton.
Mortensen (1927) records that it is often found aggregated on rocks in current-swept areas.

Suborder OPHIURIDA
Family OPHIACANTHIDAE

Ophiotrema alberti Koehler, 1896
See: Koehler, 1909 : 196-198, pi. 28, figs 1, 2.
SAMPLE. AT 192 (2). [1862 m]

DISTRIBUTION. Previously recorded only from deep water off the Azores and in the S Atlantic
(Mortensen, 1927); 3785-4060 m.

The present record from the Feni Ridge is considerably shallower than any previous record
for this species.

REMARKS. The dorsal side of the disk was missing from both specimens and large orange-
yellow gonads were visible.

Ophiomyces grandis Lyman, 1879
See: Lyman, 1879:46, pi. 14 figs 383-385; 1882 : 241-242, pi. 19 figs 13-15.

ROCKALL TROUGH ECHINODERMS I 289

SAMPLE. SBC 62 (3). [610m]

DISTRIBUTION. Previously recorded from off Tristan da Cunha (1800m) and in the Bay of
Biscay; 230-800 m.

These specimens were collected from the Hebridean Slope west of Barra. This is the first
record of O. grandis off Britain and is a northern extension of its known previous range.

REMARKS. The scarcity of occurrence may reflect a lack of sampling in the present pro-
gramme at depths between 230-800 m where it may be expected (Cherbonnier & Sibuet,
1973).

Ophiacantha abyssicola G. O. Sars, 1871
See: Mortensen, 1927 : 194, fig. 105; 1933 : 23-24, fig. 9.
SAMPLES. ES 23 (4), ES 1 1 5 (4; ? O'uveniles] 6). [704-1000 m]

DISTRIBUTION. This species is distributed throughout the N Atlantic. Although a wide
bathymetric range, c. 35-3500 m, is quoted (Mortensen, 1933; see also D'yakonov, 1954 and
Cherbonnier & Sibuet, 1973), it appears to be essentially an upper bathyal species extending
into the shallow coastal areas of the Norwegian Sea. It has also been recorded on the Irish
shelf off Eagle Island at a depth of 301 m (Farran, 1913). Samples providing unpublished
records of O. abyssicola collected by the Porcupine from Rockall and SW Ireland are to be
found in the British Museum (Natural History).

Ophiacantha abyssicola was recovered only from the Hebridean Slope off Barra and on
the Hebridean Terrace Seamount but, like Ophiomyces grandis, the apparent rarity of this
species in the samples may result from lack of sampling at favourable depths.

Ophiacantha bidentata (Retzius, 1 805)
See: Mortensen, 1927 : 196, fig. 105; 1933 : 20-22, figs 6-8.

SAMPLES. ES 10 (39), ES 12 (? 1), ES 34 (1 1), ES 55 (juvenile, 1), ES 56 (juveniles, 2), ES 69 (? [juveniles]
2), AT 107A(7), ES 112 (7), AT 121 (14), AT 139(8), AT 141 (12), AT 144(211), ES 147 (3), AT
151 (72), ES 152 (juvenile, 1), AT 153 (96), AT 154 (69), SBC 155 (juvenile, 1), AT 157 (2), ES 164
(?Duvenile]l),AT 167 (264), AT 171 (168), ES 172 (juvenile, 1), AT 175 (85), ES 176 (10; ? Duveniles]
17), AT 177 (259), AT 181 (128), ES 184 (55), AT 186 (217), AT 191 (49), AT 192 (168), AT 195
(186), ES 197 (104), AT 198 (34), ES 200 (45). SWT 10 (1), SWT 1 1 (2), SWT 16 (1), SWT 18 (1),
SWT32(14). [1 330-2925 m]

DISTRIBUTION. A widespread Arctic species the range of which extends into the N Atlantic.
A large bathymetric range of 10^4500 m is quoted (Mortensen, 1933), but in the 'warm
water' areas of the N Atlantic the bathymetric distribution is more restricted and it rarely
occurs above 500 m.

Ophiacantha bidentata occurs abundantly in the Rockall Trough. Agassiz trawls from the
Feni Ridge and from the repeat station St. 'M' at depths of 1900-2200 m have collected large
numbers of this species. It was also recorded from the Porcupine Seabight at 1765 m depth
by Farran (1913) under the name O. hibernica.

REMARKS. Ophiacantha bidentata was collected with its arms typically entwined in the
branches of the gorgonian Acanella arbuscula (Johnson), which occurs in abundance on the
west and east sides of the Anton Dohrn Seamount. It was also found occasionally attached
to the calcareous bases of both living and dead specimens ofFlabellum alabastrum Moseley.
Grassle, Saunders, Hessler, Rowe & McLellan (1975), observed an Ophiacantha sp. (either
O. bidentata, O. aculeata or O. simulans} from the submersible Alvin, on rocks and small
cobbles but not on the mud bottom, at c. 1800m off New England. Acanella seems to
occur on relatively current-swept bottoms on the continental margin where it may show a
consistent orientation to the current direction (Laubier & Sibuet, 1979). At St. 'M', near-
bottom currents may reach almost 50 cm/sec (Edelsten, 1980). Hence it seems likely that
Ophiacantha bidentata, in common with other ophiacanthids, feeds on suspended material

290

J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

using arms held out into the current in a similar way to that described by Warner & Woodley
(1975) for Ophiothrix fragilis (family Ophiotrichidae).

The colour of the arms of fresh specimens was not drab brownish as Mortensen (1927)
describes, but a fresh salmon pink, while the disk was a similar colour and greyish green
where the underlying large green-coloured stomach showed through.

REPRODUCTION. Studies on the breeding of this species show that the St. 'M' specimens are
protandrous hermaphrodites with probably direct or abbreviated development, whereas
re-examination of the shallow Arctic population, originally studied by Thorson (1936),
confirms his finding that these individuals have separate sexes (Tyler & Gage, 1982a). A
much needed revision of the genus Ophiacantha may well separate the N Atlantic deep-sea
populations as a distinct species from the shallow Arctic populations.

58°N

58° N

U°W 13 12 11 10 9°W

Oph/opleura inermis
v Homalophiura tesselata
*0phiomusium lymani

OOphiura irrorato

Fig. 3 Chart of central part of Rockall Trough showing station locations for records of the ten most
abundant brittle star species. Larger symbols denote records from more than one sample taken on
or near the station position.

Ophiacantha bidentata
Ophiactis abyssicola
Amphilepis mgolfiana

Ophiura ljungmani
Ophiocten gcacilis
Ophiocten hastatum

ROCKALL TROUGH ECHINODERMS I 291

Ophiacantha crassidens Verrill, 1885
See: Mortensen, 1927 : 192-193, fig. 106.
SAMPLE. AT 192(6). [1862m]

DISTRIBUTION. Previously known only from the N Atlantic deep-sea region (Mortensen,
1933); c. 970-3 120 m. This species was taken by the Helga in the upper Porcupine Seabight
from 986 m to 1861 m depth (Farran, 1913) but was not previously known from the Feni
Ridge where these specimens were collected in a single haul.

REMARKS. The colour of fresh specimens is a pale yellowish pink with slightly greyish darker
areas on the disk. The underlying stomach is orange coloured.

Ophiacantha cuspidata Lyman, 1878
See: Mortensen, 1933 : 31-33, figs 17, 18.
SAMPLE. ES 112(2). [1900m]

DISTRIBUTION. Previously known from scattered localities in the eastern and central North
Atlantic from Iceland and off the Azores; 785-2460 m.

Two specimens were collected from the southern Feni Ridge making this the first record
of O. cuspidata in the deep water off the British Isles.

Ophiacantha simulans Koehler, 1 896
See: Mortensen, 1933 : 26-29, figs 1 1, 13.
SAMPLE. ES 112(26). [1900m]

DISTRIBUTION. Previously known in the eastern and central North Atlantic from off the
Azores, the Bay of Biscay and S and SW of Tee land from 1480 m to 3018 m (Mortensen,
1933); in the western north Atlantic from off North Carolina.

The present record of twenty-six individuals from a single sled haul on the southern Feni
Ridge represents an extension of the known range of this species to the British seas.

REMARKS. It is interesting to note that an apparently discrete zonation of this species centred
on 1900 m depth is described by Menzies, George & Rowe (1973) off the Carolinas (NW
Atlantic). These authors also provide, from the basis of bottom photographs, a 'habitat
sketch' of this animal lying on the sediment surface in a characteristic posture with the
greater length of the arms raised vertically in the water, presumably feeding.

Ophiacantha aculeata Verrill, 1885
See: Mortensen, 1933 : 28-29, figs 14c, 15.
SAMPLE. SWT 13 (1). [3425-3500 m]

DISTRIBUTION. Previously recorded from the NW Atlantic with a bathymetric distribution
of 2425-25 10m.

This solitary record from the Rockall Trough is the first from the E Atlantic and extends
the bathymetric range.

Ophiolimna bairdi (Lyman, 1883)

See: Lyman, 1883: 256, pi. 5 (as Ophiacantha); Liitken & Mortensen, 1899; 177, pi. 18 (as
Ophiacantha); D'yakanov, 1954 : 24, fig. 6.

SAMPLE. ES 118(1). [2910m]

DISTRIBUTION. Ophiolimna bairdi has a circumpolar arctic, N Atlantic and Pacific range;
previously known bathymetric range 620-2600 m. The vertical limits of this species appear
to be restricted in the warm water area of the N Atlantic to bathyal or abyssal depths and

292 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

this species, like O. bidentata, seems to demonstrate Ekman's (1953) 'submergence' phenom-
enon.

This is the first record of this species in the Rockall Trough although Mortensen (1933)
stated that this species can 'be expected to occur all over the deeper parts of the North
Atlantic'.

Family OPHIACTIDAE

Ophiactis abyssicola (M. Sars, 1861)
See: Mortensen, 1933 : 47-50, figs 28-30.

SAMPLES. ES 18 (1), ES 34 (1), ES 1 12 (700), ES 113 (juvenile 1), ES 115 (7), AT 144 (2), AT 151
(1), AT 157 (43), AT 167 (3), AT 171 (2), AT 177 (9), ES 184 (2), AT 191 (16), AT 192 (26), AT
195 (9), ES 197 (1). GT 1 (3), GT 8 (1), SWT 12 (10), SWT 32 (2). [168 m to 2980-3000 m]

DISTRIBUTION. This species has a wide distribution throughout the Atlantic including the
Norwegian Sea; 12 5^4000 m. Ophiactis abyssicola was previously taken in the Rockall
Trough by the Michael Sars at 1856 m depth (Grieg, 1921).

Although occurring in small numbers in hauls from St. 'M', this species appears most
abundant on the Feni Ridge from where in a single haul at 1900m depth (ES 112) O.
abyssicola dominated the ophiuroid catch. It also occurred in small numbers on the Hebri-
dean slope as shallow as 650 m depth, and on the Hebridean Seamount at around 1000 m.
Both locations are thought to be somewhat current-swept, as is also the area of the Feni
Ridge sampled by ES 1 12 and the bottom at St. 'M'.

REMARKS. This species previously has been collected amongst branching corals and alcy-
onarians, and especially on the Bird's Nest Sponge, Pheronema, off western Ireland (Farran,
1913). The present records are mainly from St. 'M' where it was found entwined, along with
Ophiacantha bidentata, amongst the bushy growths of the gorgonian Acanella arbuscula.
It was occasionally also found in the base of both living and dead specimens of the solitary
coral Flabellum alabastrum.

Freshly collected specimens are a fresh salmon-pink or orange colour, with grey colouring
on the dorsal disk where the underlying stomach shows through the body wall.

Ophiactis balli (Thompson, 1 840)

See: Sussbach & Breckner, 1911: 252; Mortensen, 1927 : 200-202, fig. 1 12.
SAMPLE. ES 113(1). [168m]

DISTRIBUTION. Appears to be limited to the NE Atlantic where it has been recorded from
both the Rockall and Lousy Banks, from off the south and west coasts of the British Isles
and in the North Sea (Mortensen, 1927); mainly 60-400 m (extending to 1765 m). Farran
(1913) lists records on the Porcupine Bank indicating that it is 'common ... in water from
about 30 fathoms downwards, wherever it can find crevices in stones or coral in which to
insert itself.

The present record on the Rockall Bank is not unexpected for this essentially upper
bathyal-shelf species.

Family AMPHIURIDAE

Amphiura otteri Ljungman, 1871

See: Mortensen, 1933 : 59-61, fig. 35 (as A. palmeri Lyman).
SAMPLES. SBC 67 (1), ES 143 (1). [1000-2892 m]

DISTRIBUTION. This species is widespread, being found on both sides of the Atlantic; pre-
viously known bathymetric range, 210-1425 m.

ROCKALL TROUGH ECHINODERMS I 293

These records from the Hebridean Slope and the Permanent Station appear to be the first
published for this species from the deep sea around the British Isles, and the lower known
depth range is extended.

REMARKS. The identification of this species was mistakenly given by Mortensen (1933) and
Koehler (1907) as A. palmeri Lyman. Furthermore, we can find no difference between speci-
mens described by Koehler (1896) as A. grandis and A. otteri Ljungman, 1871; A. otteri
takes precedence.

Amphiura fragilis Verrill ,1885

See: Mortensen, 1927 : 214, fig. 121 (as A. denticulata Koehler).
SAMPLES. ABD 24 (2). [810m]

DISTRIBUTION. A. fragilis has been recorded from both sides of the N Atlantic. In the west
from off Martha's Vineyard north to the Davis Strait and W. Greenland. From the east it
has been recorded from the Faroe Channel; 430-2640 m.

This sample is from the Hebridean Slope and is the first record of this species from the
'warm' water off the west of the British Isles.

Amphiura griegi Mortensen, 1920
See: Mortensen, 1927 : 210, fig. 1 19.
SAMPLE. ABD 24(1). [810m]

DISTRIBUTION. Previously known from Norwegian fjords at 60 m to 300 m depth (Mortensen,
1920) and from the Bay of Biscay at 328-562 m (Cherbonnier, 1969; 1970). Amphiura griegi
appears to be sporadically distributed from Norway south to the Bay of Biscay.

The present record of this little known species from the Hebridean Slope represents an
interesting addition to its known geographic and bathymetric range.

Amphipholis squamata (Delle Chiaje, 1828)

See: Siissbach & Breckner, 1911: 253 (as Amphiura elegans); Mortensen, 1927: 221-222, fig. 125; A.
M.Clark, 1970a: 30.

SAMPLES. ES 5 (5), ES 23 (juvenile, 2), SBC 66 (juveniles, 5), ES 99 (juveniles, 28), ES 1 15 (juvenile,
1), SBC 155 (juvenile, 1). [704-1 330m]

DISTRIBUTION. This species is known to have a cosmopolitan distribution throughout tem-
perate and tropical coastal waters, which extends into the intertidal in the British Isles
(Mortensen, 1927; Marine Biological Association, 1957); previously known from 0-809 m
(Mortensen, 1933; Hartman & Barnard, 1958).

The present records on the Hebridean Slope, N Feni Ridge and Whittard Canyon consider-
ably extend the lower limit of the previously known bathymetric range.

REMARKS. The microphagous suspension feeding mechanism of this species has been de-
scribed by Pentreath (1970) from New Zealand material.

REPRODUCTION. A. squamata is known to brood its young (Mortensen, 1927; Fell, 1946)
and hence it is difficult to explain the occurrence of individuals in deep water as sterile out-
liers resulting from fall-out of pelagic larvae emanating from shallower depths. We therefore
infer a possibly continuous distribution from the intertidal zone (Loch Creran, personal
observations of J.D.G.) to bathyal depths off the west of Scotland and probably elsewhere.

Subfamily AMPHILEPIDINAE

Amphilepis ingolfiana (Mortensen, 1933)
See: Mortensen, 1933 : 54-56, figs 31-33.

294 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

SAMPLES. ES 4 (134), ES 6 (10), ES 8 (1), ES 10 (29, ? 10), ES 12 (juvenile, 5), ES 27 (? [juveniles]
2), ES32 (1), ES34(17), ES 55 (? 1), ES 56 (2), ES 1 1 1 (1), ES 118 (1), AT 121 (2), ES 135(1), ES
140 (4), ES 143 (2), AT 151 (1), AT 153 (2), SBC 163 (1), ES 164 (2), ES 169 (juveniles, 3), ES 172
(juveniles, 5), ES 176 (18; ? [juveniles] 5), AT 177 (4), ES 180 (1), ES 184 (13), ES 185 (12), AT 186
(1), ES 190 (5), AT 192 (1), ES 197 (7, ? 1), ES 220 (1). [1862-2925 m]

DISTRIBUTION. Although Mortensen (1933) pointed out that the published records and
descriptions of A. norvegica from the Atlantic (Lyman, 1882; Verrill, 1884; Koehler, 1924)
refer to A. ingolfiana, it is likely that N Atlantic deep-sea records of A. norvegica made
subsequent to publication of Mortensen's description of A. ingolftana also should be referred
to this species, notably those of Cherbonnier & Sibuet (1973) from the Bay of Biscay down
to 4760 m depth.

Amphilepis norvegica then remains as an essentially shallow Arctic species with records
from the northern Norwegian Sea down to the southern North Sea (Siissbach & Breckner,
1911) and the Faroe-Shetland Channel.

Amphilepis ingolfiana is probably widely distributed throughout the N Atlantic (see also
Schoener, 1967) but because of the confusion existing between the two species, it is difficult
to give an accurate geographic or bathymetric range. Mortensen (1933) quoted the depth
range of A. ingolfiana as 957 to 2580 m.

REMARKS. Some specimens collected in our samples had a ferruginous staining that is often
found on tubes or hard parts of burrowing fauna that generate a ventilatory current to their
burrow. It is interesting to note that such currents have been observed in burrows of other
amphiurid species (Woodley, 1975; Ockelmann & Muus, 1978).

Mortensen (1927) mentions that Amphilepis norvegica lives '. . . probably burrowed in the
mud in the same way as the Amphiuras.' However, it seems unlikely that A. ingolfiana is
a deep burrower in deep-sea sediments or else it would not be collected in Agassiz trawls
or epibenthic sled hauls which disturb only the superficial sediment (Gage, 1975; Gage et
al, 1980). Barham, Ayer & Boyce (1967) observed and photographed a small ophiuroid,
which they referred to as Amphilepis sp., from the bathyscaphe Trieste in the San Diego
Trough at 1234 m depth: they describe it as normally buried '. . . with only its thin, sediment-
colored rays splayed out on the surface.'

By analogy with Woodley's (1975) observations on five species of coastal amphiurids, it
seems highly possible that Amphilepis is a shallow-burrowing deposit feeder, using the
arms and tube feet as undulatory pumps that create a ventilating current, thus carrying
mucus-trapped food particles from the tips to the mouth.

Schoener (1967) describes the morphology of the post-larvae and subsequent juvenile
stages of this species, which would appear to have a widespread distribution in the N Atlantic
deep sea (Mortensen, 1933).

Family OPHIOCHITONIDAE

Ophiochiton ternispinus Lyman, 1883

See: Lyman, 1883 : 255, pi. 5, figs 67-69; Mortensen, 1933 : 67-69, fig. 40, pi. 3, figs 25-26.
SAMPLES. AT 157(2), AT 177 (1), AT 191 (1), AT 192 (1). SWT 18(1). [1 752-2200 m]

DISTRIBUTION. Previously recorded from the Faroes, W Ireland and N America; 425-1850
(? [2220) m. Mortensen (1933) considers that this large species probably is widely distributed
in the N Atlantic deep sea. However, it seems likely that nowhere is it abundant, previous
records being of single specimens in trawl hauls. The original description of Lyman (1883)
was based on a specimen dredged by the Porcupine in the Porcupine Seabight. Grieg (1921)
records a single specimen collected by the Michael Sars from a position in the Rockall
Trough close to St. 'M'.

The present records extend the known deeper bathymetric range, although records for O.
grandis Verrill, 1884, which Mortensen (1933) synonymises with O. ternispinus, and for O.

ROCKALL TROUGH ECHINODERMS 1 295

solutus Koehler 1907, which Mortensen (1927) considers may well represent the present
species, extend the previous known range to 2220 m depth.

REMARKS. The colour of fresh material was a drab olive on the disk with fawn coloured arms.

Family OPHIURIDAE
Subfamily OPHIURINAE

Ophiopleura inermis (Lyman, 1878)
See: Mortensen, 1927 : 251-252, fig. 137 (as O. aurantiaca (Verrill)).

SAMPLES. ES 20 (1), ES 99 (1). GT 1 (3), GT 2 (1), GT 8 (5), GT 1 1 (1), GT 16 (6). [650-805 m
to 1271 m]

DISTRIBUTION. On comparing the syntypes of Ophiura inermis (Lyman) with specimens of
Ophiopleura aurantiaca held in the collections of the British Museum (Natural History), we
can find no difference between them. We therefore agree with Mortensen (1933) that the
two names are synonymous, the species being distributed in both the N and S Atlantic with
a bathymetric range of 280-1875 m. In the NE Atlantic, this species has been collected from
depths of 567 m and greater off Iceland and around 1000 m depth in the N Rockall Trough
(Hoyle, 1884) to c. 1200 m in the Porcupine Seabight and 1490-1740 m in the Bay of Biscay
and the Azores (Mortensen, 1927).

The present records may reflect an intermediate level of progressive submergence in the
range of this species from high to lower latitudes in the N Atlantic.

Ophiopleura borealis Danielssen & Koren, 1877
See: Mortensen, 1927 : 249-250, fig. 136; 1933 : 94-96.
SAMPLE. ES 87 (4). [1050m]

DISTRIBUTION. Known previously from Arctic seas and off Britain from the cold area of the
Faroe Channel; 10-1885 m.

The occurrence of this impressively large species amongst the present records is confined
to a single haul from the cold water to the north of the Wyville Thomson Ridge.

REMARKS. The four specimens collected measured from 33 to 35 mm in disk diameter (cf.
the size frequency distribution in Mortensen (1933)).

Homalophiura tesselata (Verrill, 1894)
See: Koehler, 1898 : 37, pi. 7, figs 34, 36; 1909 : 156, pi. 25, figs 12-13; Mortensen, 1927 : 231-232.

SAMPLES. AT 144(2), AT 151 (1), AT 154 (2), AT 171 (3), ES 176(1), AT 181 (1), ES 184(1). SWT
18(4). [1785-1 845m to 2264m]

DISTRIBUTION. H. tesselata has previously been found throughout the N Atlantic;
458-3720 m.

The present records, which extend the distribution of this species to the Rockall Trough,
are all from St. 'M' (c. 2200 m depth) or the adjacent Hebridean Slope and lie within the
bathymetric range quoted (Mortensen, 1927). H. tesselata remains unknown beyond the N.
Atlantic (Cherbonnier & Sibuet, 1973).

Amphiophiura convexa (Lyman, 1878)

See: Lyman, 1878 : 84, pi. 3, figs 83, 84; 83, pi. 3, figs 85, 86 (as A. bullata); Mortensen, 1927 : 231.
SAMPLES. ABD 3 (juvenile, 1), ES 10 (10). [1997-2540 m]

DISTRIBUTION. This species may have a cosmopolitan distribution in the abyssal of the world
ocean; 2160-5280 m in Atlantic (Cherbonnier & Sibuet, 1973).

296 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

REMARKS. Specimens were collected from two stations in the Rockall Trough. Cherbonnier
& Sibuet (1973) found specimens possessing certain characteristics of both A. convexa and
A. bullata (Wyville Thomson, 1873) suggesting that these are conspecific; they note that both
have a nearly identical distribution in the Atlantic. Amphiophiura convexa was first recorded
in the Pacific.

Amphiophiura saurura (Verrill, 1894)

See: Koehler, 1898 : 40, pi. 6, figs 19-21; Mortensen, 1927 : 231; 1933 : fig. 49, pi. 3, figs 9, 10.
SAMPLE. ES 112(11). [1900m]

DISTRIBUTION. Previously collected in the Bay of Biscay but unknown in British Seas.
Although apparently uncommon, Mortensen (1933) suggests that this species has a wide dis-
tribution in the N Atlantic deep sea; 848-2167 m. The depth of the present record on the
Feni Ridge falls within this range.

? Stegophiura macrartha H. L. Clark, 1915
See: H. L. Clark, 1915 : 315, pi. 19, fig. 7, 8.
SAMPLE. ES 112(1). [1900m]

DISTRIBUTION. Previously recorded only once before off Georgia (NW Atlantic) at a depth
of 424m.

This find on the Feni Ridge represents the first record in the eastern Atlantic and extends
the bathymetric range to 1900 m.

Ophiocten gracilis (G. O. Sars, 1871)
See: Paterson, Tyler & Gage, 1982: 115-117.

SAMPLES. ES 4 (juveniles, 3315), ES 5 Guveniles, 265), ES 10 (juveniles, 43400), ES 12 (juveniles, 1240),
ES 18 (juveniles, 4441), ES 20 (7465), ES 22 (6761), ES 23 (5665), ES 27 (juveniles, 3), SBC 46 Guven-
iles 3), ES 54 (juvenile, 1), SBC 58 (juveniles, 33), ES 59 (juveniles, 2281), SBC 60 (? Duvenile) 1),
SBC 64 (juvenile, 1), SBC 65 (juveniles, 21), SBC 66 (57), SBC 67 (26), SBC 68 (juveniles, 11), ES
69 (9946), ES 90 (10), AT 90A (1), ES 1 1 1 (juvenile, 1), ES 99 (? 6; juveniles, 1627), ES 1 1 1 (juvenile,
1), ES 1 15 (312), ES 129 (juveniles, 5), ES 135 (juveniles, 16562), ES 137 (juveniles, 35), AT 141 (2),
ES 147 (juveniles, 10263), ES 164 (juveniles, 417), ES 169 (juvenile, 1), ES 172 (juveniles, 1694),
ES 176 (juveniles, 10380), ES 178 (147), ES 180 (juveniles, 8), ES 184 (juveniles, 650), AT 1
(77). [704-2925 m]

DISTRIBUTION. The synonomy and status of species of the genus Ophiocten have recently
been revised by Paterson, Tyler & Gage, (1982). Identifications of populations of O. gracilis
previously have been confused both with the shallow arctic species O. sericeum (Forbes)
and with O. abyssicolum (Forbes) which is re-defined as a lusitanian species, the Atlantic
distribution of which becomes increasingly submerged northwards, and does not appear to
extend into the Rockall Trough (Paterson et al, 1982).

We have found adults (2-50-4-75 mm disk) of O. gracilis only from upper bathyal depths
on the Hebridean slope (stations ES 20, ES 22, ES 23, SBC 66, SBC 67 in the north Rockall
Trough at 1040 m depth (ES 90) and on the Hebridean Seamount at 1000 m depth (ES 1 1 5).
The remaining records listed above refer only to post larvae of this species found in samples
from greater depths where a massive but non-viable settlement occurs from pelagic larvae
probably originating from the upper slope populations (Gage & Tyler, 198 la), Semenova,
Mileikovsky & Nesis (1964) and Tyler & Gage (19826), show that a form of Ophiopluteus
ramosus' which Geiger (1963) and the above authors indicate to be abundant in the surface
zooplankton in spring in parts of the N Atlantic, is most probably the pelagic larval stage
of this species. We suggest that O. gracilis may have a ribbon- like distribution along the
upper continental slope, and on submerged banks around the 1000 m depth level, around
the N Atlantic.

ROCKALL TROUGH ECHINODERMS I 297

REMARKS. The evidently high population densities (Semenova et ai, 1964; Gage & Tyler,
198 la) suggest that O. gracilis might be extensively preyed on by bottom feeding fish:
remains of this species are identifiable and commonly found in stomachs of demersal fish
trawled on the Hebridean slope (Mauchline & Gordon, personal communication).

Close-up bottom photographs taken at around 1000 m on the Hebridean slope only rarely
show brittle stars (A. J. Southward, Marine Biological Association, Plymouth, personal com-
munication). Possibly the individuals of this abundant population are shallow burrowers,
or lie with the mottled coloured disk closely pressed against the sediment or with a dusting
of particles covering it. The markedly long oral and arm tube-feet of this species may be
associated with a microphagous feeding mechanism.

Ophiocten hastatum Lyman, 1878
See: Paterson, Tyler & Gage, 1982: 117-1 19.

SAMPLES. ES 6 (9; ? Ouveniles] 4), ES 10 (1; ? Ouveniles] 2), ES 27 (1; ? [juvenile] 1), ES 31 (1), ES
52 (2), ES 55 (1), ES 56 (2; ? [juvenile] 1), AT 107A (10), ES 1 1 1 (1), AT 119 (1), AT 121 (187),
ES 129 (1), ES 137 (4; ? [juvenile] 1), ES 140 (7), ES 143 (? 4), ES 147 (7), ES 152 (2), ES 164 (? 1),
ES 185(5). OTSB51001 (1). [2000 m to 2921 (? 2925) m]

DISTRIBUTION. This species may now be regarded as cosmopolitan in distribution, with a
bathymetric range in the lower abyssal from 1 1 30 to 5000 m, but concentrated on the lower
abyssal zone. Paterson, Tyler & Gage (1982) in re-examining species of the genus Ophiocten
have found no significant differences between O. hastatum and O. latens, records of the latter
being confined to the Atlantic.

Ophiura affinis Liitken, 1858

See: Sussbach & Breckner, 1911: 244-247; Mortensen, 1927 : 244-245, fig. 132.
SAMPLE. ES 113(10). [168m]

DISTRIBUTION. Although not apparently recorded previously on Rockall Bank this record
is not unexpected in view of the known distribution in the NE Atlantic and around the British
Isles (e.g., Farran, 1913; Mortensen, 1927; D'yakonov, 1954; Ursin, 1960); 8-550 m.

Ophiura carnea Liitken, 1858
See: Mortensen, 1927 : 243, fig. 131.
SAMPLES. ES 2 (1), AT 194 (1). [630-2857 m]

DISTRIBUTION. Ophiura carnea is an eastern Atlantic species found from Norway south to
the Bay of Biscay, previously collected also from the upper Porcupine Seabight by the Helga
from 215 m to c. 400 m (Farran, 1913); 50-1260 m.

The present records are the first for this species in the Rockall Trough, that from the Anton
Dohrn Seamount, AT 194 in 630m depth, is well within the previously recorded range;
the one from ES 2 in 2857 m depth represents a considerable extension to the known
bathymetric range.

Ophiura irrorata (Lyman, 1 878)
See: Lyman, 1882 : 47, pi. 5, figs 7-9; Mortensen, 1933 : 86-87, fig. 48.

SAMPLES. AT 151 (2), AT 175(1), AT 177(1), AT 181 (1), AT 191 (59), AT 192 (220), AT 195(1),
AT 198(1). [1 862-2220 m]

DISTRIBUTION. This is a cosmopolitan abyssal species; Mortensen (1933) gives the known
bathymetric range as 600-43 1 5 m.

A few specimens were found in Agassiz hauls at St. 'M'. However, hauls on two positions
on the Feni Ridge yielded by far the largest samples indicating a more numerous develop-

298 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

ment of the population there. All specimens agreed with Mortensen's (1933) description
of O. irrorata sensu stricto. When compared with localities worked on the eastern side of
Rockall Trough, the small numbers or apparent absence of this species seems striking.
Possibly, O. irrorata is associated with conditions, such as strong bottom currents, that are
best developed on Feni Ridge.

REMARKS. The colour of fresh specimens is a pale pink with a dark blue-purple colouration
where the stomach shows through the dorsal and ventral disk. The tube feet on the arms
are a pale orange colour which immediately distinguishes them from the bright red tube feet
of 0. ljungmani when found in the same haul.

Ophiura imprudent (Koehler, 1906)
See: Koehler, 1907 : 256, pi. 18, figs 9-10.
SAMPLE. ES 113(6). [168m]

DISTRIBUTION. This species has only been recorded once before, off the Azores; 560 m.

The present record from Rockall Bank of this evidently rare species is somewhat shallower
at 168 m.

Ophiura ljungmani (Lyman, 1878)
See: Mortensen, 1927 : 240-242, fig. 130.

SAMPLES. ES 2 (55), ABD 3 (? 7), ES 4 (1464), ES 6 (1439), ES 8 (? 36), ES 10 (juveniles, 6120), ES
12 (548), ES 14 (32), ES 15 (243), ES 20 (juvenile, 1), ES 27 (170), ES 28 (25), ES 31 (25), ES 32
(1) ES 34 (1 164), SBC 46 (3), ES 52 (53), ES 53 (15), ES 54 (1), ES 55 (851), ES 56 (688), ES 57 (921),
SBC 58 (5), ES 59 (425), SBC 60 (juveniles, 4), SBC 68 (juvenile, 1), ES 69 (juveniles, 2), ES 99 (juven-
iles, 8), AT 107 A (2), ES 1 1 1 (205), ES 1 12 (44), ES 1 18 (201), AT 1 19 (2), AT 121 (640), ES 129
(512), ES 135 (973), ES 137 (787), AT 138 (2), AT 139 (8), ES 140 (1056), AT 141 (50), ES 143(107),
AT 144 (3), ES 147 (965), SBC 150 (4), AT 151 (8), ES 152 (360), AT 153 (47), AT 154 (12), ES
164 (530), AT 167 (49), ES 169 (323), AT 171 (17), ES 172 (270), AT 175 (15), ES 176 (289), AT
177 (35), ES 180 (291), AT 181 (35), ES 184 (352), ES 185 (74), AT 186 (25), ES 190 (73), AT 191
(65), AT 192 (38), AT 195 (21), ES 197 (5), AT 198 (1), ES 200 (1). SWT 1 1 (2), SWT 13 (2). [1050 m
to 3425-3500 m]

DISTRIBUTION. Commonly occurring throughout the N Atlantic; 309^4070 m (Cherbonnier
& Sibuet, 1973).

Ophiura ljungmani was abundant in samples taken at the Permanent Station (2900 m),
becoming less numerous at shallower stations where Ophiomusium lymani became abun-
dant, although adult specimens were found as shallow as 1632 m on the Hebridean slope.
At the latter position, the bottom haul also included adults of Ophiocten gracilis occurring
near the lower end of the bathyal range of the breeding population. The deepest record of
Ophiura ljungmani was from 3425-3500 m in the Porcupine Seabight.

REMARKS. The colour of fresh specimens is pale rose to white, with conspicuously bright
red tube feet. The stomach shows through the disk as a dark blue-purple colouration.

REPRODUCTION. Ophiura ljungmani has separate sexes in roughly equal proportions. Study
of the gametogenic cycle shows a marked seasonal periodicity with rapid vitellogenic
growth in autumn and maximum development and probable spawnout by late January/early
February at the Permanent Station (Tyler & Gage, 1979, 1980). These authors suggest that
egg size (max. 90 um) and fecundity (up to 5700 eggs per individual) indicate planktotrophic
larval development.

Analysis of disk-size frequencies from the Permanent Station indicates low survival of the
annually recruited postlarvae and a high though constant rate of mortality amongst older
year classes. The time series studied at the Permanent Station also suggests both a seasonal
growth pattern, with maximal growth in spring, and a variation in year-class strength (Gage
& Tyler, 19816). Reproductive maturity is thought to be reached in the third year at a disk

ROCKALL TROUGH ECHINODERMS I 299

diameter of 3-5^4-0 mm with individuals thereafter reproducing annually (Tyler & Gage,
1980).

Ophiura ophiura (Linnaeus, 1758)

See: Sussbach & Breckner, 1911: 238-241 (as O. ciliaris); Mortensen, 1927 : 236-238, fig. 128 (as O.
texturata).

SAMPLE. ES 23(1). [704m]

DISTRIBUTION. Previously recorded all round the coasts in British seas; 0-200 m.

The presence of this well-known coastal species on the upper Hebridean Slope represents
a surprising extension of its known bathymetric range.

Subfamily OPHIOLEPIDINAE

Ophiomusium lymani Wyville Thomson, 1873
See: Mortensen, 1927 : 253-254, fig. 138.

SAMPLES. ES 4 (153), ES 10 (juveniles, 13), ES 12 (32), ES 14 (10), ES 15 (juveniles, 33), ES 18 (1),
ES 20 (juveniles, 4), ABD 24 (1), ES 34 (15), SBC 68 (1), AT 68A (1), AT 107A (192), AT 1 14 (42),
AT 138 (8), AT 139 (14), AT 141 (juveniles, 14), AT 144 (1686), ES 147 (juveniles, 2), AT 151 (233),
AT 153 (610), AT 154 (369), AT 157 (3), AT 161 (119), SBC 166 (juvenile, 1), AT 167 (944), AT
169 (juvenile, 1), AT 171 (1565), AT 175 (735), ES 176 (402), AT 177 (1245), AT 181 (1711), ES
182 (140), ES 184 (1 174), AT 186 (1027), AT 191 (64), AT 192 (714), AT 195 (1434), ES 197 (308),
AT 198 (724), ES 200 (192). SWT 10 (505), SWT 1 1 (1), SWT 16 (3), SWT 17 (6), SWT 18 (129),
SWT32(35). [8 10-292 1m]

DISTRIBUTION. Ophiomusium lymani is now known to have a cosmopolitan distribution;
130-3435 m. It was first described by Wyville Thomson (1873) from dredgings by the
Porcupine in the Rockall Trough. Elsewhere this species has been recorded from between
700 m and 4700 m and at temperatures varying from 2° to 9°C (Cherbonnier & Sibuet, 1973).
Adult specimens in our samples were taken over a wide area and in depths from 810m
on the Hebridean Slope to 2540 m in the central Rockall Trough. Postlarvae of this species
are distinctive and easily recognised (see Schoener, 1967); they were common in epibenthic
sled hauls taken between these depths and were also found in hauls down to 292 1 m depth
in the vicinity of the Permanent Station. This suggests that postlarvae dispersed to depths
significantly greater than about 2500 m do not survive to adulthood.

REMARKS. The numbers of specimens collected in Agassiz hauls at different depths indicate
that the population is best developed around the 2000 m isobath in Rockall. At St. 'M', the
population density visible in close-up bottom photographs is a little less than 1 m~2 (Dr A.
J. Southward, personal communication). Size frequencies from Agassiz hauls at this station
show a marked dominance by adults, with a postlarval peak apparent from the fine-meshed
epibenthic sled hauls. Although finding no seasonal pattern in oogenesis from comparison
of oocyte size frequencies in samples taken at different times of the year, Gage & Tyler (1982)
interpret differences in the relative size of the postlarval peak in spring and summer samples
as evidence for a seasonal recruitment to the population. A model age structure consisting
of two fast growing juvenile year classes with a slowing of growth on attaining reproductive
maturity at c. 20 mm disk diameter and a consequent 'stacking up' of adult year classes is
deduced from study of a time series of disk-size frequencies in large samples of Ophiomusium
lymani from St. 'M'.

Heavy mortality is thought only to apply to post-larvae; the relatively heavily calcified
body and large size of adults protecting them from heavy predation. However, a small per-
centage (c. 2-5%) of specimens show healed, or partially healed, lesions usually in the dorsal
interradial area. They may be caused by attacks from biting predators, probably fish. Sub-
sequent regeneration of the wound may give rise to 'monstrous' specimens sometimes found.

300 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

Bottom photographs and submersible observations (Barham, Ayer & Boyce, 1967; Wigly
& Emery, 1967; Grassle, Sanders, Hessler, Rowe & McLellan, 1975) indicate that this species
moves over, or nestles into, the sediment surface without burrowing. Some photographs
show the disk raised well clear of the sediment, supported by the proximal part of the arms.
Barham et al. (1967) suggested that O. lymani is a filter feeder; however, the absence of
obvious ciliary tracts, long spines and arm podia equipped with mucus producing glands
as found in other brittle stars known or likely to filter feed, such as members of the families
Amphiuridae, Ophiactidae, Ophiotrichidae and Ophiocomidae (see Fell, 1966 and Reese,
1966 for reviews of pre-1965 literature; Fontaine, 1965; Pentreath, 1970; Warner & Wood-
ley, 1975) seem to render a filter feeding mode of nutrition unlikely for Ophiomusium
lymani. Preliminary study of the stomachs of specimens from St. 'M' has yielded few recog-
nisable contents, the stomachs of most of 100 specimens examined appearing empty. How-
ever, fragments of mysid uropods and small gastropod shells were found along with
polychaete setae, indicating that O. lymani may rather be an opportunistic scavenger or
carnivore.

Discussion

For many relatively well known species that are also abundant in our samples a bathymetric
range of adults is recorded that is somewhat narrower than that known over its full range.
This is not unexpected considering the comparatively small area of the present study
compared to the full geographical range of the deep-sea distribution of a species, when, as
seems likely, these distributions are determined chiefly by the temperature of water masses
(Ekman, 1953). Examples include the asteroid Porcellanaster ceruleus and the ophiuroids
Ophiacantha bidentata, Ophiactis abyssicola, Ophiura ljungmani and Ophiomusium
lymani. In addition there are many uncommon species known from great depths to the south
that are recorded in the Rockall area at somewhat shallower depth. Examples include the
ophiuroids Ophiotrema alberti, Ophiacantha crassidens and the asteroids Pteraster reductus,
Hoplaster spinosus. This may be explained in terms of the general trend towards progressive
submergence of the cold, deep ocean water, expressed as a trend to tilting of isotherms in
longitudinal sections of the Atlantic Ocean (Wiist, 1936), that is related to the continual
formation of deep water from the polar oceans. The species Ophiacantha cuspidata provides
an illustration of this trend towards an increasingly submerged distribution towards lower
latitudes, which is analogous to the bipolar-equatorial submergence seen in distributions of
many coastal species in the Atlantic that are described by Ekman, 1953. Mortensen (1927,
1933) cites records made by the IngolfoiO. cuspidata from SW of Iceland in 1461 m depth,
and by the Challenger at 785 m depth in the S Atlantic off Ascension Island; while the pre-
sent records from the Rockall Trough at 55°12 'N in 1900 m fall between these depths and
the Princesse Alice record at 42°53 'N in 2460 m off the Azores (Koehler, 1909).

Topographic boundaries such as the Wyville Thomson Ridge give rise to locally sharp
step-like increases in the physical gradient where cold arctic water enters the Atlantic deep
sea by a process of intermittent overflow (Ellett & Roberts, 1973). The southerly route of
this overflow is thought to lie along the sedimentary feature, Feni Ridge, which describes
a sinuous path along the eastern margin of the Rockall Bank. The consequent latitudinal
difference in hydrographic regime on the west and east sides of Rockall Trough may well
be connected with differences seen in the echinoderm fauna around the 2000 m level (Table
1).

Although, as shown in Table 1 , more samples have been taken on the east side, a consider-
ably richer echinoderm fauna appears in the few samples taken in the west. Some species
recorded there were not taken at all in the east around St. 'M' and the adjacent lower Hebri-
dean slope despite the relatively intense sampling effort made in this area. Many of the
species recorded only in the west are uncommon and have, as already discussed, only been
previously taken in deeper water to the south. Other more common species, such as Ophiura
irrorata, appear to be far more numerous on Feni Ridge.

ROCKALL TROUGH ECHINODERMS I

301

Table 1 Distribution of species on west and east sides of the Rockall Trough. *species present only
as juveniles; tprobable suspension feeders (see text).

Species taken only from
west (on Feni Ridge at
1600-2 190m depth, 5 hauls)

Species common to
both areas

Species taken only from
east side (on St. 'M' or
adjacent Hebridean Slope at
1600-2200 m depth, 23 hauls)

Heliometra glacialis

*Psilaster andromeda
Evoplosoma scorpio
*Pteraster reductus
~\Asteroschema inornatum
^Ophiotrema alberti
"fOphiacantha crassidens
"fO. cuspidata
fO. simulans
Amphiophiura saurura
? Stegophiura macrarthra
Ophiocten has to. turn

Porcellanaster ceruleus

Bathybiaster vexillifer
Plutonaster bifrons
Benthopecten simplex
Pseudarchaster parelii
Hymenaster membranaceus
Ophiacantha bidentata
^Ophiactis abyssicola
^Amphilepis ingolfiana
Ophiochiton ternispinus
Ophiura ljungmani
Ophiura irrorata
Ophiomusium lymani

Rhizocrinus lofotensis
? Hathrometra sarsi
Dy taster ins ignis
Pectinaster filholi
Hoplaster spinosus
*Paragonaster subtilis
^Asteronyx loveni
Homalophiura tesselata

It is tempting to account for the presence of these species as essentially abyssal elements
that find the cooler water overflowing from the Arctic to their liking. However, hydrographic
data given by Ellett & Martin (1973) although showing some east-west tilting of deep-water
temperature and salinity isopleths indicates that this explanation may be insufficient:
samples from the Permanent Station at 2900 m where the bottom water generally is as cool
as in shallower water to the west on Feni Ridge have not yielded the missing species.

It seems more likely that hydrographic conditions other than temperature may also be
important in the west. Since overflow is thought to be associated with fast bottom currents
and high turbidity (Jones, Ewing, Ewing & Eittreim, 1970), it seems possible that the richer
fauna may reflect the better feeding conditions, particularly for microphagous suspension
feeding, in the nepheloid layer at the benthic boundary. Hence, it may be significant that
six out of the eleven species found as adults only from the Feni Ridge are likely, on
morphological criteria, to be suspension feeders (Table 1 ). However, one difficulty with this
explanation is that the apparently higher species richness of echinoderm groups covered by
the present account on Feni Ridge is also associated with physical conditions that, because
of the episodic nature of overflow, are more variable than elsewhere. Although admittedly
we are dealing with only a fraction of the total benthic fauna, the present data do not accord
with the stability-time hypothesis invoked by Sanders (1968, 1969) to explain why species
richness in the supposedly stable environment of the deep sea appears to be higher than in
comparable coastal habitats where conditions for life are more variable. It is interesting to
note that results from two other studies of deep-sea faunas (Rex, 1973; 1976; Thistle, in press)
do not fit in with the prediction of a simple positive relationship between increasing physical
stability and species richness. Furthermore, the results of intensive sampling in the Bay of
Biscay indicate that species diversity of asteroids on the lower continental slope decreases
with increasing depth after a maximum at 2200m (Sibuet, 1977), despite a presumed
increasing physical stability with increasing depth. Rex (1981) has re-analysed data available
from the NW Atlantic to show a parabolic response of species diversity of both macrofauna
and megafauna to depth, with maxima at levels from 1900 m to 2800 m.

The present study also demonstrates a greater geographic and bathymetric range of juven-
iles and post- larvae of some relatively abundant species compared to the adult range. Juven-

302

J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

lies of the following species of sea star have been identified over a wider bathymetric range
than adults: Plutonaster bifrons, Pectinaster filholi, Pseudarchaster parelii and Hymenaster
membranaceus. Juveniles and post-larvae of brittle stars usually are smaller than those of
sea stars. However, Gage & Tyler (198 la) have shown that a massive non- viable settlement
of the upper bathyal brittle star Ophiocten gracilis occurs seasonally over a wide area of the
Rockall Trough. The easily recognisable post-larvae and juveniles of Ophiomusium lymani
also are found in samples from a wider range of depths than adults. Post-larvae are typically
found in small numbers in epibenthic sled hauls from the Permanent Station (c. 2900 m)
at a depth several hundred metres deeper than the lower limit of adults. We suggest that
the probably demersal lecithotrophic larvae are dispersed by currents to areas sometimes
inimical to adult survival. Post-larvae of other ophiuroid species have also been ident-
ified in the Permanent Station samples; one of these being an ophiacanthid, probably
Ophiacantha bidentata, that probably is derived from an adult population in shallower
water, since no adult ophiacanthid species has as yet been collected from the Permanent
Station.

Summary

Four species of sea lily, forty species of sea star and thirty-six species of brittle star are ident-
ified from recent extensive sampling conducted by the Scottish Marine Biololgical Associ-
ation from R.R.S. Challenger in the deep sea area lying to the west of the British Isles. The
following species have not previously been recorded from the British seas.

Crinoidea Thaumatocrinus jungerseni

Asteroidea Hyphalaster inermis

Pseudarchaster gracilis
Paragonaster subtilis
Pteraster sp. aff. P. acicula
Hymenaster gennaeus (provisionally)
H. rex

Freyella spinosa
F. sexradiata
Hydrasterias sexradiata

Ophiuroidea Astrodia tenuispina

Asteroschema inornatum

Ophiotrema alberti

Ophiomyces grandis

Ophiacantha cuspidata

O. simulans

O. aculeata

Ophiolimna bairdi

Amphiura otteri

A. griegi

Amphilepis ingolftana

Homalophiura tesselata

Amphiophiura convexa

A. saurura

? Stegophiura macrarthra

Ophiocten hastatum [Hoyle (1884) records this species but under the name O.

sericeum]

Ophiura irrorata

O. imprudens

Ophiura imprudens is known from only one specimen from the Azores and Ophiacantha
aculeata only from N America and SW Iceland.

Extensions of bathymetric range are recorded (in metres) for the following:

ROCKALL TROUGH ECHINODERMS I

303

Crinoidea

Thaumatocrinus jungerseni
Heliometra glacialis

Asteroidea
Luidia ciliaris
Plutonaster bifrons
Psilaster andromeda
? Radiaster tizardi
Paragonaster sub tills
Plinthaster dentatus
Evoplosoma scorpio
Pteraster reductus
Hymenaster gennaeus
H. rex

Henricia abyss ico la
Brisinga endecacnemos
Freyella spinosa
Stichastrella rosea
Zoroaster fulgens

Ophiuroidea
Astrodia tenuispina
Asteroschema inornatum
Ophiotrema alberti
Ophiacantha aculeata
Ophiolimna bairdi
Amphiura otter i
A. griegi

Amphipholis squamata
Amphiophiura convexa
? Stegophiura macrarthra
Ophiura carnea
O. imprudens
O. ophiura (syn. O. texturata)

Upper limit

2455-(1785) 1845
1920-1900
c. 930-168

3785-1862

2160-1997

560-168

Lower limit

2734-3425 (3500)
1358-1900

400-650 (805)
2500-2965
1853-2965
1320(? 1630)-2540

2117-2190
c. 1600-1900

2245-2500 (2500)
2285-2910

2200-2220
4060-4810
1150(? 1330H632
3660-4810

3300-3425 (3500)
1480-1900

2510-3425(3500)
2600-2910
1425-2892
300-810
740-1330

424-1900
1260-2857

200-704

In this list the depths given are the minimum possible extensions of the previously known
bathymetric limits. Figures in brackets give the maximum possible from the depth range of
hauls (fish trawlings only) although of course specimens may not have been taken throughout
the depth range of the tow.

For some of the more abundantly occurring species from fine-meshed epibenthic sled
hauls, the records demonstrate a wider bathymetric distribution of juveniles and post-larvae
than of adults, suggesting that externally developing larvae often are dispersed to areas where
they do not survive to adulthood.

In general, records of adults of the most abundantly occurring species cluster well within
the vertical range recorded over their, often circumoceanic, known distribution. This may
reflect a trend towards progressive submergence of isotherms in the deep ocean from high
to low latitudes.

Despite a concentration of sampling effort to the east, comparison between the east and
west sides of Rockall Trough has indicated that a richer echinoderm fauna, particularly of
filter feeding species, is present in the west on Feni Ridge. This may be related to a greater
availability of suspended food in the west associated with the passage of fast-flowing turbid
water at the benthic boundary as a result of overflow from the Faroe Channel over the
Wyville Thomson Ridge.

304 J. D. GAGE, M. PEARSON, A. M. CLARK, G. L. J. PATERSON & P. A. TYLER

Acknowledgements

We record our gratitude to Miss B. Rae of Dervaig, Mull, for valuable help in sorting
samples, and Dr J. D. M. Gordon of S.M.B.A. and to Mr C. Palmer and Dr F. Woodward
of the Glasgow City Museum for their cooperation in making available material originally
collected mainly by Dr Dietrich Burkel on Dr Gordon's deep-water fishing cruises on R.R.S.
Challenger. We are also grateful to Miss Maureen Downey, Dr Myriam Sibuet and Dr
Geoffrey Smalden for their expertise in identifying, giving advice on particular species or
for advancing information prior to publication. We thank also Dr Gert Konnicker of Univer-
sity College, Galway and Dr Paul Cornelius of the British Museum (Natural History) for
kindly identifying pennatulid and gorgonian specimens found carrying brittle stars, Dr John
Mauchline, S.M.B.A. for identifying crustacean fragments in Ophiomusium stomachs, and
Mr James Wright of University College of Swansea for loan of colour slides of freshly
collected material.

We wish to express our special gratitude to the captain, officers and crew of R.R.S.
Challenger and R.R.S. Shackleton and to colleagues participating in cruises, too numerous
to name individually, without whose cooperation and many kindnesses our work would not
have been possible. P.A.T. gratefully acknowledges support from N.E.R.C. grant no.
GR3/4131.

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Manuscript accepted for publication 1 1 November 1982

British Museum (Natural History)
The Red Notebook of Charles Darwin

Edited by Professor Sandra Herbert

Charles Darwin used the pocket-sized Red Notebook to record various
observations and ideas over the course of the year from mid- 1 836 to
mid- 1 837. It was an important year, spanning the last months of the voyage
of H. M.S. Beagle and the first months back in England. The notebook
contains observations on points visited, reading notes, and speculations on
theoretical questions. The theoretical questions Darwin considered in the
notebook pertain primarily to geology, where he was interested in finding
an explanation for the rise and fall of the earth's crust, and to the subject
of the mutability of species. Most significantly, the Red Notebook contains
the earliest known evidence of Darwin's adoption of an evolutionary
hypothesis. The notebook also reveals Darwin's dependence on professional
zoologists working in London for technical judgements decisive for his
adoption of an evolutionary position.

Sandra Herbert is Associate Professor in History at the University of
Maryland Baltimore County and is the author of several studies of various
aspects of the early career of Charles Darwin.

Published in Bulletin of the British Museum (Natural History) Historical series, Volume 7
(paper covers), 1980.

Co-published by the British Museum (Natural History) and Cornell University Press in hard
bound edition. 1980.

Titles to be published in Volume 45

Problems in catfish anatomy and phylogeny exemplified by the Neotropic
Hypophthalmidae (Teleostei: Siluroidei). By G. J. Howes

Miscellanea

Bats (Mammalia: Chiroptera) from Indo- Australia. By J. E. Hill

On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the
interrelationships of African cichlid species flocks. By P. H. Greenwood

Miscellanea

The cranial muscles of loricarioid catfishes, their homologies and value as
taxonomic characters (Teleostei: Siluroidei). By G. J. Howes

Miscellanea

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

Bulletin of the

British Museum (Natural History

The cranial muscles of loricarioid catfishes
their homologies and value as taxonomic
characters (Teleostei: Siluroidei)

Gordon J. Howes

Zoology series Vol 45 No 6 27 October 1983

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British Museum (Natural History)
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The cranial muscles of loricarioid catfishes, their
homologies and value as taxonomic characters
(Teleostei: Siluroidei)

Gordon J. Howes

Department of Zoology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Contents

Introduction 309

Muscle nomenclature 310

Specimens examined 311

Description of loricariid cranial muscles 313

Summary of muscle systems 322

Out-group comparisons and homologies of loricarioid jaw muscles . . 323

Interrelationships of loricariids based on myological characters . . . 329

Congruent osteological features 330

Myological + osteological characters 337

Classification of the Loricariidae 338

Discussion 341

The interrelationships of the Loricariidae 341

Functional considerations 343

Acknowledgements 344

References . 344

Introduction

The Loricariidae are a family of Neotropical siluroids (catfishes) with two outstanding exter-
nal features: an 'armoured' body and a sucker-like ventrally situated mouth. The bodies of
the catfishes are virtually encased in a series of bony plates or scutes reflecting the muscle
segmentation (see Alexander, 1965: 136) and the heads are covered with a mosaic of bony
plates. The bony plates generally run in an overlapping series of 5 longitudinal rows, but
in those forms with an attenuated and compressed caudal peduncle, the series may be
reduced posteriorly to two. All the plates, most of the head covering, the fin spines and rays
bear spines. In some taxa, particularly in males, the spines may form dense clumps on the
tops and lateral parts of the head. The operculum and interoperculum also bear spines which
on the latter, may be attenuated and hooked.

The spines, or denticles all appear to have a similar structure, namely dentine surrounding
a pulp cavity and with an enamel cap (see Bhatti, 1938). Thus, to all intents and purposes
they are to be regarded as teeth. 0rvig (1977) used the term 'teeth' for those dental elements
confined to the jaws and 'odontodes' for similar elements occurring on the extra-oral dermal
skeleton. In this paper, 'odontodes' will be used to refer to such spines, prickles and bristles
on the scutes, head and fin rays.

The sucker-like mouth is in the form of reflected upper and lower, usually papillate, lips.
They surround highly mobile jaws (see p. 343) which often bear long, usually bifid, hooked
teeth. The jaws may be long and shallow with the teeth numerous, arranged in regular rows
e.g. Chaetostoma, short and deep with the teeth few and clumped together e.g. Pseudacanthi-
cus, or lamellate and edentulous, some Loricaria.

Bull. Br. Mus. nat. Hist. (Zool.) 45(6): 309-345 Issued 27 October 1983

310 G. J. HOWES

Alexander (1965) has described the mouth and its functional morphology. He showed how
the modified mouth, body scutes and fins are all features associated with the torrential and
benthic environments of loricariids.

In terms of numbers of taxa, the Loricariidae is the largest siluroid family, containing ca
70 genera and 600 nominal species (Isbriicker, 1980). Whether or not this number of taxa
are a reflection of the taxonomic concepts of various authors is a matter for debate (see.
p. 339) but even if many proved to be synonyms, it is likely that the family would retain
its numerical supremacy.

Of the currently recognized genera (Isbriicker, 1980) approximately 50% occur in the
Amazon basin, 30% in Guianas, 20% in both the Orinoco and trans- Andean regions, 25%
in Parana, 15-16% in the Magdalena and Andes, and 10% in south-eastern Brazil.

The biology of loricariids is poorly known. Most appear to feed on algae or perhaps organ-
isms that live in algal mats. Angelescu & Gneri (1943) found that different taxa fed on
molluscs, chironomid larvae, mud and diatoms. Many species live amongst stones and roots
and some Hypostomus live in tunnels in muddy river banks (Vaz-Ferreira & Senorans,
1971).

The development of long odontodes on the head and anterior borders of the pectoral fin
spines are sexually dimorphic characters in some taxa, Ancistrus, Loricaria and Lithoxus;
their function is discussed later, p. 343.

There exists no comprehensive work on loricariid anatomy. Regan (1904; 1911) listed a
series of osteological characters by which he defined the family and its subunits. Leege (1922),
Peyer (1922), Starks (1926), Gregory (1933), Angelescu & Gneri (1943), Chardon (1968),
Alexander (1964; 1965) and Lundberg & Raskin (1969) are the principal sources of infor-
mation on various osteological features. Of these papers, only those of Alexander, Chardon
and Lundberg & Baskin have provided anything approaching a comparative analysis of the
family with other siluroids. Even so, the number of taxa whose anatomy has been described
has been exceedingly small.

Only one paper, that of Alexander (1965), has described the musculature of loricariid fishes
in any detail. The earlier paper by Leege (1922) has negligible myological information.
Alexander limited his description of jaw musculature to Hypostomus, = Plecostomus in
Alexander, and the opercular musculature (part) of Ancistrus. Alexander's work was aimed
primarily at a functional description, and detailed myological description was lacking.

The principal aim of this paper is to go some way towards filling the lacunae of knowledge
about loricariid myology. By making out-group comparisons with other siluroids and
otophysans the muscular system of loricariids are viewed in terms of character states and
used to construct a cladogram. This scheme is compared with others utilizing selected osteo-
logical characters and the resultant information is used to produce a cladogram of nested
synapomorphies for those taxa studied. No detailed treatment of loricariid interrelationships
is presented. As yet no cladistic framework is available for any group of siluroid fishes and
so any comments on interrelationships must be regarded as highly speculative.

Some attention is given to the possible function of the muscular systems described.

Muscle nomenclature

The adductor mandibulae muscle complex of teleostean fishes is usually recognised as being
divisible into several elements each identified by a lettering sequence A,-AW, following Vetter
( 1 878). As various subdivisions of these elements have been recognised, they have been identi-
fied by addition of a subscript letter or numeral. This system of nomenclature has, in some
instances, resulted in complex notations (see for example Matthes', 1963, system for cypri-
nids). More important, however, is whether those elements similarly labelled in different taxa
represent homologues — a question discussed later (p. 323).

In most siluroids, the morphology of the adductor mandibulae is complex, and this is par-
ticularly so in loricariids. In order to clarify the following descriptions the muscles are shown
schematically in Fig. 1 .

CRANIAL MUSCLES OF LORICARIOID CATFISH

311

man

Fig. 1 Schematic diagram of loricariid jaw-palatine muscles and their insertions.

The outer part of the adductor muscle complex, which in more advanced teleosts runs
to the maxillary bone of the upper jaw, designated as A,, inserts, in siluroids, on to the lower
jaw. The muscle having its insertion on to the maxilla is recognised as the retractor tentaculi.
The phylogenetic origin of this muscle is possibly from a medial part of the adductor
mandibulae; see Howes (1983), and below, p. 327 for discussion of homology. The number
of medial divisions of the adductor complex in loricariids exceeds that of other siluroids,
and possibly that of other otophysans. This multiplication of elements leads to difficulties
in using the generally applied system of notation since their respective homologies are uncer-
tain (see p. 327). Thus, for descriptive purposes all jaw muscles are designated a-f.

Muscle a is that inserting on the maxilla; muscles b-d are various divisions of the adductor
mandibulae; muscles e-f attach to the palatine bone and are derivatives of the adductor arcus
palatini.

Other suspensorial, opercular and hyoid muscles are named according to the nomenclature
of Winterbottom (1974) and their homologies with those muscles so named in other teleosts
are accepted.

Specimens examined

The specimens listed are those dissected for this study. In addition, dissections, skeletons,
alizarins and radiographs of specimens representing nearly all siluroid families have been
examined.

Loricariidae: Ancistrus occidentalis 1981.1.5:722-748; 749-797
Ancistrus stigmaticus 1901.5.8:10 (skeleton)
Ancistrus spp Uncatalogued (skeleton)
Chaetostoma anomalum 1903.6.30:102 (skeleton)
Chaetostoma breve 1898.1 1.4:38 (skeleton)
Chaetostoma dermorhynchus 1880.12.8:64-66
Chaetostoma microps 1981.1.5:821-867

312 G. J. HOWES

Cochliodon cochliodon 1981.1.5:777-787

Cochliodon sp. Uncatalogued (alizarins)

Farlowella knerii 1880.12.5:206 (skeleton)

Farlowella spp Uncatalogued (inc. alizarins)

Hemiancistrus megacephalus 1972.10.17:872-875

Hemiodon ichthys acipenserinus 1879.12.1:75-79; 1977.3.10:186

Hemipsilichthys calmoni 1929.2.1:66-71

Hypoptopoma carinata 1926.10.27.367-73

Hypostomus plecostomus 1981.1.5:760-776

Hypostomus spinossisimus 1 898. 1 1 .4:30-3 1

Hypostomus verraes 1895.3.29:38 (skeleton)

Ixinandria montebelloi 1 98 1 . 1 1 .30: 1 3-1 5

Lipopterichthys carrioni 1935.5.20:18-27

Lithoxus lithoides 1972.7.17:66-1 15

Loricaria cataphracta 1926.3.2: 765-82; 1860.11.10:15 (skeleton)

Loricaria brunneus 1981.6.9:1074-1081

Loricaria jubata 19 14.5. 1 8:68-72

Loricaria labialis 1934.8.20:368-388

Loricaria lanceolata 1981.1 .5:873-904

Loricaria microlepidogaster 1929.2.1:29-50

Loricaria platystoma 1974.5.22:544-545

Loricaria simillima 1934.8.20:326-46

Loricaria strigilata 1929.2.1:31^0

Loricaria teffeana 1895.5.17:103 (skeleton)

Loricaria variegata 1914.5.18:73

Loricaria venezeuelae 1947.7.1:228-232

Loricaria spp Uncatalogued (alizarins)

Metaloricaria nysseni 1982.4.7:7

Rhadniloricaria macromystax 1 869.5 .2 1 :8

Neoplecostomus granulosus 1896.7.4:3-5; 6 (skeleton)

Otocinclus nigricauda 1934.8.20:220-240 (alizarin)

Panaque nigrolineatus 1974.8.14:13-15

Peckoltia pulcher 1980.7.1:87; Uncatalogued

Pogonopoma wertheimeri 1904.1.28:6

Pogonopomoides parahybae 1904. 1 .28:7

Pseudacanthicus serratus 1870.3.10:4 (skeleton)

Pseudancistrus barbatus 1972.7.17:49-50

Pseudohemiodon sp. Uncatalogued

Pseudohemiodon laticeps 1895.5.17:113-4

Pterygoplichthys anisitsi 1934.8.20:255-64

Pterygoplichthys multiradiatus 1897.12.1:280 (skeleton)

Stoniella leopardus 1973.7.17:188-198

Sturisoma panamense Uncatalogued

Sturisoma lyra 1897.12.1:72 (skeleton)

Astroblepidae: Astroblepus festae 1894.11.4:18 (skeleton)

Astroblepus heterodon 1910.7.11:138 (skeleton)
Astroblepus grixalvii 1898.11 .4: 1 2-1 7
Astroblepus cf. praeliorum 1969.1.27:35-46

Callichthyidae: Callichthys callichthys 1981.6.9:995-1005; Uncatalogued
Hoplosternum littorale 1934.9.12:413^420

Diplomystidae: Diplomystes papillosus 1889.11.14:33; 1896.10.2:22 (skeleton)
Doradidae: Amblydoras hancocki 1971.4.14:15-30
Mochokidae: Euchilichthys royauxi 1909.4.29:98-102
Nematogenyidae: Nematogenys inermis 1883.1 1.27:45-48; 49 (skeleton)

CRANIAL MUSCLES OF LORICARIOID CATFISH 313

Scoloplacidae: Scoloplax dicra UMMZ 198967, Paratypes (inc. alizarins)

Sisoridae: Glyptosternum cous 1968.12.13:471-76
Glyptothorax major 1978.3.20:286-289

Trichomycteridae: Branchioica magdalenae 1947.7.1:203-8 (alizarins)
Eremophilus mutisii 1978.2.22:16
Trichomycterus cordovense 191 1.1 1.25:1-10
Trichomycterus rivulatus 1944.6.16:142-5

Description of loricariid cranial muscles

Jaw muscles

Whilst the jaw muscles of loricariid fishes differ in their arrangement from those in other
siluroids (see p. 323), within the group there is a high degree of uniformity. Differences are
marked by the relative size of each muscle and by varying points of insertion. Both variables
are diagnostic for sub-groupings within the family.

In order to present a succinct comparative analysis, the jaw musculature of one taxon will
be described in detail and attention drawn to taxa which differ from that plan. The represen-
tative taxon is Pterygoplichthys anisitsi. Choice of this species is determined by its repre-
senting the largest group of loricariids — the subfamily Hypostominae (see p. 338), and by
what may be termed its relatively generalised morphology. This latter impression does not
imply that the taxon is primitive or plesiomorphic.

When the thick, plate-like epidermis is removed from the side of the head in
Pterygoplichthys three large muscles are exposed. The medial, band-like muscle (a, Fig. 2A)
originates from the lateral margin and ventral surface of the lateral ethmoid passes laterally
to the palatine and inserts on the maxilla. The maxillary ramus of the trigeminal nerve runs
along its lateral border and sends a branch into the muscle's ventral surface. The buccal
ramus passes across the dorsal surfaces of the muscle and the palatine to enter the maxilla.

The outer muscle (b), originates below the eye and inserts on the lower jaw. Its lateral
fibres originate from the fascia of the cheek armour, the posterior fibres from the pre-
operculum, and the medial fibres from the hyomandibula and pterygoid. There is a conti-
nuity of origin across all the bony medial elements and it is not possible to differentiate
individual muscle segments. The lower jaw insertion is somewhat complex (Fig. 2B). Prior
to insertion the outer bloc of fibres bifurcates, one bundle inserting on the medial margin
of the dentary, the other on the coronoid process. Both insertions are musculose. An inner
slip of fibres, arising from the pterygoid is also differentiable from the anterior part of the
main muscle mass. This muscle slip turns medially around the anterior border of the
pterygoid to insert tendinously on the inner aspect of the dentary.

Lying medial to muscle b is a long, almost cylindrical muscle, c, which originates from
the hyomandibula; its anterior part passes medially below muscle a and inserts tendinously
onto the premaxilla.

Medial to muscle c, and extending along the pterygoid, is a thin, narrowly triangular
muscle (d, Figs 2B & 3 A). Its origin is from the hyomandibula and its insertion is into a
complexly divided connective tissue sheet. This sheet has attachments to the ventral surface
of the palatine, to the upper surface of the premaxilla, and to the dentary (Fig. 3 A).

In Pterygoplichthys there is a tendinous insertion on the premaxilla, although the dorsal
part of the muscle is firmly attached to the palatine. The mandibular trunk of the trigeminal
nerve passes across the face of muscle c, to make a slight loop below it and attach to its
ventral border by connective tissue. From this point the nerve curves downward into muscle
b and passes between the bifurcation of that muscle into the dentary-anguloarticular foramen
to innervate the outer tissues of the lower jaw.

314

G. J. HOWES

rmxV

v" -VJ< ^f>'€^**^C->^^

f>y.'r

mx

pmx

Fig. 2 Jaw muscles of Pterygoplichthys anisitsi (BMNH 1234.8.20: 255-64). A, superficial muscula-
ture, dorsal view; B, deeper muscles with muscle a removed, dorso-lateral view. Scale=10mm.
eml = ethmo- maxillary ligament: ep = epidermis; pal = palatine.

Muscles e and f (Fig. 3B) operate the palatine; muscle e originates from the medial face
of the pterygoid and from the ventral surface of the lateral ethmoid, its insertion is onto the
postero- ventral margin of the palatine. The fibres deriving from the pterygoid are horizon-
tally aligned whereas those stemming from the lateral ethmoid are aligned at angles varying
between 20°-45° to the horizontal. Muscle f originates from the ventro-medial surface of
the lateral ethmoid and from the ventral surface of the frontal. It inserts on the posteromedial
aspect of the palatine. A strong ligament connects the posterior lateral face of the palatine
with the anterior border of the pterygoid.

In Pterygoplichthys muscle e lies in a pterygoid channel. The lateral wall of the channel
rises to meet the lateral ethmoid margin and so totally encloses the muscle which exits
through an anterior vertical slit. Alexander (1965) described a similar feature in Hypostomus.

Variants to the jaw muscle pattern described above occur in a group of taxa represented
by Hemipsilichthys and Chaetostoma where muscle a is broadened and fan- like so that it
completely covers muscle c (Fig. 4 A). In species of this group the suspensorium is orientated
into an almost horizontal plane, so that muscle b bulges laterally.

CRANIAL MUSCLES OF LORICARIOID CATFISH

315

pmx

Fig. 3 Pterygoplichthys anisitsi, A, insertion of inner jaw muscle d, lateral view; B, configuration of
extensor tentaculi muscle in dorsal view. The cut surfaces of the pterygoid are shown hatched.
Scale = 3 mm. lpt = pterygoid-palatine ligament.

More marked differences occur in a group containing Loricaria, Sturisoma and Farlowella
(Figs 5 A; 6). Used here, Loricaria corresponds to the broad concept of Regan (1904) rather
than the restricted one of Isbriicker (1981); see further discussion on p. 338. In these genera
muscle c inserts on the premaxilla via a thick tendon whose length is correlated with that
of the snout. The tendon passes through a loop of connective tissue at the rostro- ventral
margin of the palatine, to turn through a right angle before its premaxillary insertion (Fig.
6, tc). Muscle d is directed medially, its tendon of insertion running into the palatine-
premaxillary connective tissue and almost meeting its antimere in the midline before joining
the premaxilla. In Sturisoma, the muscle is attached to the ventro-posterior part of the
palatine as in Pterygoplichthys.

In some Loricaria species there are distinct dorsal and ventral portions of muscle a. The
lower portion (av, Fig. 5A) inserts on the wide, convex dorsal surface of the palatine, its
postero-medial fibres are inseparable from those of muscle e; the upper part (ad) runs forward
to insert on the maxilla as in other loricariid taxa.

316

pmx

B

pmx

Fig. 4 Hemipsilichthys calmoni (BMNH 1929.2.1: 66-71). A, superficial musculature; B, deeper
muscles with muscle a removed; both in dorsal view. Scale = 10 mm.

Farlowella differs from Loricaria and Sturisoma in that muscle a is enclosed within a ptery-
goid channel similar to that in Pterygoplichthys. Pseudohemiodon and Hemiodonichthys
depart in several respects from the above arrangements (Fig. 5B). Muscle a is a narrow, strap-
like element; muscle c does not insert directly on the maxilla but runs into an extensive tissue
sheet connecting the pterygoid, palatine, premaxilla and maxilla; muscle d inserts, via a long
tendon, on the premaxilla and maxilla; muscle e inserts tendinously onto the ventral centre
of the lamellate palatine.

Two other genera which depart from the generalized jaw muscle plan are Hypoptopoma
and Otocinclus (Fig. 7). In these genera, muscle a occurs as the most lateral of the superficial
jaw muscles. It runs from the laterally shifted lateral ethmoid (see p. 332) to insert
tendinously into the centre of the maxilla. Muscles b and c have their 'normal' insertions
onto the lower jaw and premaxilla respectively. Muscle d joins a thick connective tissue band
linking the palatine and premaxilla.

CRANIAL MUSCLES OF LORICARIOID CATFISH

av

317

mx

Fig. 5 Jaw and palatine muscles of A, Loricaria strigilata (BMNH 1929.2.1: 31^40) in dorso-lateral
view; B, Pseudohemiodon sp. (uncatalogued) in dorsal view. Scale = 5 mm.

Hyoid muscles

The muscles serving the lower jaw, and having their origins from the hyoid bar, have a
uniform morphology in the Loricariidae.

The intermandibularis is relatively feeble compared with that in other siluroids. It is most
highly developed in Pterygoplichthys (Fig. 8A) where its lateral insertion is bifurcate, the
inner part joining the dentary, the outer attaching to the loose connective tissue at the rictus
of the mouth. In other loricariid taxa the intermandibularis is a thin band of muscle fibres
which in broad-mouthed genera, e.g. Chaetostoma, Pseudancistrus, Lipopterichthys, attaches
laterally to the reflected lip tissue (Fig. 8B).

318

G. J. HOWES

-tc

mx

Fig. 6 Metaloricaria nysseni (BMNH 1982.4.7:7) showing insertions of deeper muscles; lateral view.
lf= lower labial tissue; pmf=premaxillary medial labial fold; tc = insertion tendon of muscle c.
Scale = 5 mm.

eth

pmx

mx

Fig. 7 Hypoptopoma carinatum (BMNH 1973.3.27: 3) jaw muscles in dorsal view. Scale= 1 mm.

CRANIAL MUSCLES OF LORICARIOID CATFISH

im A

319

a.

hyi

im

Fig. 8 Hyoid muscles of A, Pterygoplichthys anisitsi (BMNH 1934.8.20: 255-64); B, Chaetostoma
acquinoctialis (BMNH 1978.7.14: 20), both in ventral views. Scales=5 mm. ph, a, b = divisions of
protractor hyoidei; hya, hyi = hyohyoideus adductores and inferioris; \m = intermandibularis.

The protractor hyoideus is well-developed and extends between the posterohyal and
anterohyal to the lower jaw (Fig. 8 A). The fibres from the left and right hyoid bars do not
meet in the midline as in other siluroids and indeed, in most teleosts. Each part of the muscle
is divided for almost its entire length into two segments (Pha, Phb, Fig. 8A). One lies dorso-
medially and attaches to the dentary, the other is ventro- lateral and attaches to the dorsal
face of the reflected lip tissue.

The largest hyoid muscle in loricariids is the hyohyoides inferioris which extends laterally
from the midline of the urohyal to the anterohyal and posterohyal. In Cochliodon, Panaque
and Hypostomus, the lateral border of the muscle has a marked concave indentation to allow
for the bypass of the protractor hyoideus. The hyohyoidei adductores extend from the upper
medial face of the operculum, between the branchiostegal rays, to a midline raphe above
the sternohyoideus.

320 G. J. HOWES

Opercular and suspensorial muscles

The muscles having the greatest degree of variability in size and position are those controlling
the opercular apparatus.

In Pterygoplichthys the dilatator operculi muscle is a thin, narrow element running along
a hyomandibular groove. It originates from a pterotic fossa and inserts on the antero-medial
border of the operculum (do Fig. 9A). The levator operculi muscle originates from the
pterotic-posttemporal = temporal plate of Regan, 1904, and inserts on the dorso-medial
margin of the operculum (lo, Fig. 9A). The adductor operculi originates from a cavity formed

do

?

Fig. 9 Opercular and suspensorial muscles of A, Pterygoplichthys anisitsi; B, Loricaria simillima; C,
Loricaria cataphracta; D, Ancistrus occidentalism E, Pseudohemiodon sp. Scale = 5 mm. A-D in
dorso- lateral, E in dorsal views. ah = adductor hyomandibular is; po = preoperculum; bsr =
branchiostegal ray.

CRANIAL MUSCLES OF LORICARIOID CATFISH 321

between the hyomandibula and pterotic, to run almost vertically behind the hyomandibula
and insert on the central dorso-medial aspect of the operculum (ao, Fig. 9 A).

In Pterygoplichthys the levator arcus palatini is a thin, vertically aligned muscle extending
between the sphenotic and hyomandibula (lap, Fig. 9A). The adductor arcus palatini muscle
in all loricariids examined runs laterally and almost horizontally from the posterior part of
the parasphenoid to the medial face of the hyomandibula. It is probably more correct to refer
to this muscle as the adductor hyomandibulae (see Winterbottom, 1974: 239) as it is a
separate element from the anterior part of the adductor arcus palatini which is modified,
as in most siluroids, to form the extensor tentacularis muscles (see p. 328).

Apart from minor variations in the orientation of the muscles, the above descriptions of
opercular and suspensorial muscles in Pterygoplichthys are applicable to Hypostomus and
Hemiancistrus.

In Neoplecostomus, the dilatator operculi inserts on the dorsal rather than the anterior
aspect of the opercular margin. The interoperculum in Neoplecostomus is also larger than
that in other loricariid genera.

Pseudacanthicus, Pseudancistrus, Stoniella, Cochliodon and Panaque show a departure
from the hypostomine opercular muscle pattern. The dilatator operculi is shifted more
anteriorly so as to run across the posteromedial aspect of the orbit. The muscle passes,
laterally, beneath a bridge formed by the preoperculum and pterotic-posttemporal plate
before inserting on the reduced operculum. In Panaque the dilatator operculi is situated so
far anteriorly that it crosses the medial centre of the orbit (Fig. 1 0). In these five taxa the
levator operculi originates from a ventral cavity in the pterotic-posttemporal plate and inserts
via a thin tendon along the thickened dorso-medial edge of the operculum. The adductor
operculi appears to be represented by a small slip of muscle running horizontally from the
pterotic-posttemporal to insert on the posterior border of the preoperculum, thus running
along the top of the bridge beneath which the dilatator operculi emerges. There is no connec-
tion of the adductor operculi with the operculum. In Pseudacanthicus, Stoniella and Panaque
there are evertible 'interopercular' spines (see p. 334).

do

cl

Fig. 10 Opercular muscles of Panaque nigrolineatus (BMNH 1974.8.4: 13-15), in lateral view.
Scale =10 mm. cl = cleithrum; io= interoperculum; hyo= hyomandibula; po= preoperculum.

322 G. J. HOWES

In Hemipsilichthys and Chaetostoma the dilatator operculi is thin and tendinous. It origi-
nates from the pterotic and runs outward at an angle of 30° to the horizontal in a lateral
hyomandibular groove. The adductor and levator operculi muscles are of similar morphology
to those in Pterygoplichthys. The levator arcus palatini is, however, smaller, occupying a
postorbital space between the sphenotic and hyomandibula.

In Ancistrus the opercular muscles are hypertrophied. The dilatator operculi muscle is
extensive, originating from the roof of the cranium and the supraoccipital (Fig. 9D). The
central tendon runs antero-laterally at an angle of about 45°c to the vertical and attaches
to the postero-dorsal aspect of the operculum. The operculum is modified to form a
mechanism for rotating the 'interoperculum' laterally; described below (p. 334).

The levator and adductor operculi muscles are well developed, the adductor originating
from a ventral process of the fused vertebral complex and directed ventro-laterally at 35°
to the vertical. The levator arcus palatini, instead of inserting on the suspensorium, as in
other loricariids, joins the insertion tendon of the dilatator operculi and thus appears to
reinforce the adduction of the operculum.

In Loricaria and Sturisoma, the dilatator operculi originates principally from the hyoman-
dibula, a few antero- ventral fibres take their origin from the posterior rim of the preopercu-
lum. In some species of Loricaria, e.g. L. simillima, the muscle's origin is shared in part
with that of the levator arcus palatini which is also well developed (Fig. 9B). In other species,
e.g. L. cataphracta, the levator arcus palatini is absent (Fig. 9C).

In all Loricaria species examined the levator operuli is absent; the adductor operculi
originates from a pterotic-posttemporal cavity. In Sturisoma a few tendinous fibres passing
along the dorso- medial surface of the operculum may represent the levator operculi, and the
adductor operculi may be represented by a few fibres emanating from the anterior fascia of
the body musculature, which at this point is directed laterally to insert on the pterotic-
posttemporal plate.

Farlowella possesses a similar morphology of the dilatator operculi and levator arcus
palatini muscles to that of Loricaria. In this genus both adductor and levator operculi muscles
are present but weakly developed.

In Pseudohemiodon and Hemiodonichthys the dilatator operculi is reduced virtually to a
tendon which runs in a lateral hyomandibular groove. Insertion is on to the antero-dorsal
corner of the operculum. Levator and adductor operculi muscles are absent. Instead, the
medial wall of the operculum provides the site of attachment for lateral fibres of the
hyohyoideus adductores muscle (9E). There is no levator arcus palatini muscle in these taxa.

Hypoptopoma and Otocinclus both have a feeble dilatator operculi originating from the
hyomandibula and inserting on the antero-dorsal point of the operculum. A levator operculi
is present but weakly developed and runs forward ventro-laterally from a medial fascia of
the epaxial musculature.

The levator arcus palatini originates from a sphenotic fossa, runs outward at an angle
of 45° to the vertical and inserts on the hyomandibula just anterior to the ventral part of
the dilatator operculi.

Summary of muscle systems

From the data presented above, the following groups of loricariid catfishes are discernible
on the basis of correlated myological features:

1 . Jaw muscles: muscle a narrow, band-like, lying straight and lateral to the mesethmoid,
inserting on the maxilla; muscle b inserting entirely on the lower jaw; muscle c inserting
on the premaxilla; muscle d inserting on the premaxilla via a connective tissue sheet which
attaches to the dentary and the palatine; muscle e running through a well-developed ptery-
goid channel to insert on the palatine. Opercular muscles: usually well-developed; levator
arcus palatini (LAP) small. Hyoid muscles: intermandibularis well-developed, sometimes
bifurcated laterally, hyohyoidei inferioris with concave lateral border, protractor hyoideus

CRANIAL MUSCLES OF LORICARIOID CATFISH 323

with bifurcate insertion to the lower jaw, hyohyoidei adductores attaching to the medial face

of the operculum.

Included genera: Neoplecostomus, Hypostomus, Pterygoplichthys, Hemiancistrus,

2. Jaw and hyoid muscles as (1). Opercular muscles: dilatator operculi shifted anteriorly
medial to orbit; adductor operculi bridges pterotic-posttemporal plate and preoperculum.
Included genera: Panaque, Cochliodon.

3. Jaw and hyoid muscles as (1), but no pterygoid channel accommodating muscle e.
Opercular muscles as (2).

Included genera: Stoniella, Pseudacanthicus, Pseudancistrus, Acanthicus.

4. Jaw and hyoid muscles as (1 ), but muscle a expanded concomitantly with increased maxil-
lary length; muscle e running in pterygoid channel. Opercular muscles: dilatator operculi
mostly tendinous and obliquely angled laterally. LAP considerably reduced.

Included genera: Hemipsilichthys, Lipopterichthys, Chaetostoma, Lasiancistrus.

5. Jaw and hyoid muscles as (1). Opercular muscles hypertrophied, dilatator operculi
originating from supraoccipital, levator operculi from the complex vertebrae.

Included genera: Ancistrus, Lithoxus.

6. Jaw and hyoid muscles as (1), but muscle d directed medially, its tendon of insertion
almost meeting its antimere; muscle a in some Loricaria partly inserting on palatine; muscle
e in Farlowella and Loricaria platystoma enclosed in a pterygoid tunnel. Opercular muscles:
dilatator operculi variably developed; levator and adductor operculi muscles always reduced,
sometimes absent. LAP absent in some Loricaria species.

Included genera: Loricaria, Farlowella, Sturisoma.

1. Jaw and hyoid muscles as in (1), but muscle d with long insertion tendon; muscle e

inserting into the centre of the palatine ventrally. Opercular muscles reduced, levator and

adductor operculi absent. LAP absent.

Included genera: Pseudohemiodon, Hemiodonichthys, Crossoloricaria.

8. Jaw muscles differ from all the above in that muscle a lies laterally to the other adductor

musculature, otherwise as in (1). Opercular muscles reduced.

Included genera: Hypoptopoma, Otocinclus.

Out- group comparisons and homologies of loricarioid jaw muscles

A broad comparison of siluroid jaw muscles is presented elsewhere (Howes, 1983) and analy-
sis is here restricted to the following taxa:

Diplomystes, the single genus comprising the family Diplomystidae, considered by most
authors to be the most 'primitive' extant siluroid.

Nematogenys, the single genus comprising the family Nematogenyidae. Recent authors,
Gosline (1945), Chardon (1968), Arratia et al. (1978) include this genus within the Tricho-
mycteridae. For reasons given here (p. 329), I follow Eigenmann (1928) in recognising the
genus as representing a distinct family.

Astroblepus, the single genus comprising the family Astroblepidae. Regan (191 1) and Gosline
(1947) both considered the astroblepids a subfamily of the Loricariidae. I follow Eigenmann
(1910) in regarding them as a distinct family (see p. 341). Differences in dentition and cranial
osteology suggest that there are possibly two genera contained within Astroblepus.

Representatives of the Trichomycteridae and Callichthyidae, both families believed by
Chardon (1968) to be related to loricarioids. The terms 'loricarioid' refers to the members
of the Loricariidae plus the Astroblepidae, and 'loricariid' to members of Loricariidae only.

In the majority of siluroids the adductor mandibulae inserts on the lower jaw. Plesiomor-
phically, as in Diplomystes and Nematogenys, divisions of the muscle block are absent or
poorly differentiated. The outer, most massive part of the muscle originates from the pre-
operculum, hyomandibula and usually the quadrate, and inserts onto the outer face of the
anguloarticular-dentary. This arrangement has been found in all siluroids examined and it

324

G. J. HOWES

was the contention of Fink & Fink (1981) that a lower jaw insertion of the outer part of
the muscle (A,) was an apomorph character of siluroids (but see below, p. 327).

Although in loricariids a major portion of the adductor mandibulae, termed muscle b in
this text, does insert on the lower jaw, medial portions, muscles c and d, pass via thin tendons
running through the palatal connective tissue to the premaxilla (see Fig. 3 A).

As stated by Alexander (1965: 135), in the Callichthyidae the superficial part of the
adductor mandibulae, which he believed to be a combined A, + A2, inserts partly on the
lower jaw and partly on the anguloarticular-maxillary ligament, whilst a medial part inserts
on the outer face of the lower jaw. The situation is rather more complex than that outlined
by Alexander since the dorsal part of the adductor can be distinguished as a separate element.
Thus, the adductor morphology in callichthyids may be described as follows: An element
originating from the hyomandibula, just postero- ventral to the orbit, inserts tendinously on
the antero- medial process of the maxilla (rt, Fig. 11). The larger part of the adductor origi-
nates posteriorly from the preoperculum, medially from the hyomandibula and laterally
from the 3rd infraorbital. Its antero-ventral fibres insert on the medial and dorsal aspects
of the anguloarticular, the dorsal fibres share the points of insertion on the maxilla with those
of the more dorsal muscle, ?A, (Fig. 1 1 B). As noted above, the deeper part of the muscle
inserts entirely on the lateral face and dorsal margin of the anguloarticular.

pal

rt

pmx

mx

B

Fig. 11

Jaw muscles of Hoplosternum littorale(BMNH 19434.9.12: 413-420). A, superficial muscula-
ture; B, deep muscles; both in lateral view, Scale = 5mm. ri = retractor tentaculi.

CRANIAL MUSCLES OF LORICARIOID CATFISH

325

As in loricariids, the superficial part of the principal muscle is separated from its medial
portion by the ramus mandibularis nerve trunk. In this respect, the outer portion, which
has its insertion on the upper and lower jaws, should be considered as A,, whilst the medial
part having its insertion entirely on the lower jaw should be considered as A2. The dorsal
muscle, which has its insertion on the maxilla, is equivalent to the retractor tentaculi in
loricariids and other siluroids, except that its origin is posterior rather than anterior to the
orbit. That this muscle appears to be a dorso-medial part of the adductor, lends credence
to the idea that phylogenetically, the retractor tentaculi originated from a medial part of the
adductor muscle complex (see Alexander, 1965; Howes, 1983).

In Diplomystes and Nematogenys the adductor mandibulae is relatively undifferentiated,
the outer part of the muscle running on to the lateral aspect of the anguloarticular, its dorsal
fibres inserting via a thick tendon on to the dorsal rim of the dentary. The tendon is continu-
ous with a thick 'ligamentous' tissue attaching to the maxilla (see Howes, 1983).

In the Trichomycteridae, the adductor mandibulae is also relatively simple (Fig. 12 A).

am

tmx

B

op

am

pom

Fig. 12 Trichomycterus rivulatus (BMNH 1944.6.16: 142-45). A, superficial jaw muscles in dorsal
view. B, posterior jaw and opercular muscles, la lateral view. Scale = 5 mm. am -adductor
mandibulae; pom = preopercular muscle; man = mandible; rb = buccal ramus of facial nerve;
rmd = mandibularis nerve; sp = sphenotic; tmx = tendon of adductor muscle to maxilla.

326

G. J. HOWES

The outer portion inserts on the lower jaw but a tendon runs forward into connective tissue
attaching to the maxilla. A medial section of the muscle, ?A2, inserts into the inner aspects
of the dentary.

In the Astroblepidae, the adductor mandibulae, muscle b, is as in the Loricariidae, with
the outer portion being divided from the inner by the mandibularis nerve (Figs 13A & B).
The outer portion inserts on the rim of the anguloarticular, whilst the inner inserts on
the medial face of the dentary. Muscle c is also present but is deeper and thicker in the
Loricariidae. Two portions of this muscle are discernible, a broad dorsal segment and a deep
medial one. The ventral fibres of the medial segment originate from the inner part of the
muscle b, the remainder from the ptergyoid. The dorsal fibres of the upper portion originate
from the lateral ethmoid and the ptyergoid. Medially the two portions are inseparable but
insert on short, separate tendons which join prior to their insertion on the premaxilla (Fig.
13B).

mx

B

et c(part'd')

pmx

Fig. 13 Astroblepus cf. praeliorum (BMNH 1969.1 .27: 35^46). A, superficial jaw musculature; B, deep
muscles, both in dorsal view. Scale = 5 mm. man-mandible; no = nasal organ.

CRANIAL MUSCLES OF LORICARIOID CATFISH 327

The morphology of the adductor mandibulae in other siluroids is discussed elsewhere
(Howes, 1983), but it is pertinent to note here that in none is there a direct connection
between the upper jaw and the outer muscle element. It is also argued (Howes, 1983) that
this condition is plesiomorphic for otophysans, and that encroachment of lateral fibres to
the upper jaw is a derived feature.

The premaxillary insertion of part of the adductor mandibulae in the Loricariidae and
Astroblepidae, and its maxillary insertion in the Callichthyidae, are viewed as apomorph
features. In all three groups, the articulation between the premaxilla and the mesethmoid
is a mobile one, a condition not encountered elsewhere in siluroids. In other siluroids, includ-
ing Diplomystes, the premaxilla is firmly united to the lateral extensions of the mesethmoid.
Alexander (1965: 134) showed that in Corydoras the mouth is protrusile, and this protusi-
bility is partly the result of having a movable premaxilla. It seems likely that because of
its restricted distribution amongst siluroids, a movable ethmoid-premaxillary joint is more
likely to be a derived character, particularly if Diplomystes is viewed as the most plesiomorph
extant siluroid (see Fink & Fink, 1981). Suffice it to say that in other otophysans, mobile
premaxillary-ethmoid articulations are regarded as derived characters (see Vari, 1979: 272
concerning characoids).

If the plesiomorph conditions of adductor mandibulae A, in teleosts is regarded as that
in which the muscle is inserted on the lower jaw and lies lateral to the mandibular nerve
trunk, then muscle b in loricariids and astroblepids must be the homologue of A,. This
pattern is, however, further complicated in these two groups by the mandibular nerve trunk
entering the muscle mass and passing between two bundles of fibres before entering the lower
jaw (see p. 313). Thus, on the strict definition given above, those fibres lying medial to the
nerve trunk must be regarded as A2. This suggestion is supported by the insertion of the
medial fibres of muscle b on to the coronoid process and medial face of the dentary — which
is the usual place of insertion for A2. Whether the inner slip which in loricariids inserts on
the medial face of the dentary should be regarded as A3 is doubtful, as by definition, A3 inserts
on the anguloarticular. However, it must be borne in mind that the loricariid lower jaw has
rotated outward so that the inner surface has come to lie in a horizontal plane with the
dentigerous surface forming the anterior border. It may be that coordinate with this rotation,
the 'A3' slip has moved forward from the anguloarticular to the dentary. Thus, it may be
concluded that muscle b in loricarioids comprises the two elements identified in other
otophysans as A, and A2 with the possibility that an inner segment present only in Lori-
cariidae represents A3.

Muscle c lies dorso-medially to the A,, A2, A3 complex and at its hyomandibular origin
cannot be separated from the outer combined element. Insertion is via a tendon running
through connective tissue to the premaxilla. This muscle appears to be the unlabelled dorsal
element shown in Alexander's (1965; fig. 17) figure of Hypostomus. This element has prob-
ably derived from a dorso-medial separation of the adductor mandibulae. In Astroblepus,
the lower portion of muscle c is not entirely separated from the outer adductor complex (see
above, p. 326 & Fig. 13). This condition suggests that muscle c is a derivative of the medial
part of the adductor series which has become attached to the premaxilla via the connective
tissue sheet that connects the lower and upper jaws, and the palatine.

Alexander (1965) applied the name retractor premaxillae to the element designated here
as muscle d. However, it would seem that Alexander overlooked the fact that part of the
adductor mandibulae, here named muscle c, also joins the upper jaw. Concerning functional
roles, it would appear that muscle c performs the action which Alexander ascribed to muscle
d, namely being the antagonist of the extensor tentaculi muscles, e and f, and thus retracting
the upper jaw. For this reason I am applying the name retractor premaxillae to the dorso-
medial element designated in this text as muscle c.

Muscle d which Alexander (1965) recognised as the 'retractor premaxillae', appears to
perform a different function in that, whilst it ultimately inserts into the premaxilla, it is also
connected with the lower jaw and with the palatine. In this respect its functional role is not

328 ' G. J. HOWES

at all clear, but manipulation of preserved specimens indicates that a significant downward
movement of the palatine occurs when the muscle is pulled posteriorly. Thus, I will refer
to it as the retractor palatini. Again, this muscle seems to be a de novo development from
the inner part of the adductor mandibulae complex.

In the Diplomystidae, Nematogenyidae and Trichomycteridae, there is no separate medial
muscle attaching to the upper jaw. Only in the Astroblepidae are there elements recognisable
as homologues of muscles c and d in the Loricariidae (p. 326). In most loricariids (e.g. Ptery-
goplichthys, Fig. 3 A) muscle d (retractor palatini) has a plesiomorphic attachment to the
underside of the palatine prior to its insertion on the upper jaw. It is hypothesised that phylo-
genetically the inner part of the adductor mandibulae has, in loricariids, shifted from the
lower to the upper jaw via a connection with the palatine.

Muscle a is regarded as the homologue of the retractor tentaculi in other siluroids. Along
the lateral border of the muscle runs the ramus maxillaris of the trigeminal nerve which bifur-
cates anteriorly, one branch innervating the dorsal, and the other the ventral fibres of the
muscle. The dorsal face of the muscle is crossed by the buccal ramus which terminates in
the maxillary tissue. Similar innervations and nerve course are to be found associated with
the retractor tentaculi of other siluroids (see for example Mithel, 1964). The origin of the
retractor tentaculi has been the subject of debate (see Winterbottom, 1974), some authors
believing the muscle to be derived from A, and others that it is a de novo development from
the medial aspect of the adductor. A more detailed discussion of the matter (Howes, 1983)
concludes that the latter is a more likely hypothesis to account for its derivation. The course
of the trigeminal nerve rami in Nematogenys and the trichomycterids, taxa lacking a retractor
tentaculi, reinforces this view. Both the buccal and maxillary rami lie medial to the principal
muscle mass (Fig. 12 A). The only way it is possible for the nerves in loricariids to have come
to occupy their present position would have been for a muscle segment to have developed
ventro-medially to them and for the nerves to have shifted laterally in relation to the muscle.

The most extreme shift of the retractor tentaculi is that in Hypoptopoma and Otocinclus
where the lateral ethmoid has extended so far laterally that the muscle makes an angle of
about 45° to the midline.

Muscles e and fare homologous with the extensor tentaculi of other siluroids. Both sections
are derivatives of the adductor arcus palatini and form the antagonists of the retractor
tentaculi (see Winterbottom, 1974: 240). In Astroblepus, the extensor tentaculi is of a
primitive, single element whose fibres run laterally from the lateral ethmoid and insert on
the posterior portion of the palatine (Fig. 13B). The muscle is floored by a thick tendinous
sheet connecting the parasphenoid with the lateral ethmoid processes.

Opercular muscle morphology in loricariids is correlated with the size and position of the
opercular bones. In the majority of taxa the operculum is small, with its dorso-posterior
border sealed to the pectoral girdle by a membrane, leaving only a ventral opening. Such
immobility of the operculum has, in some taxa, led to atrophy or disappearance of the levator
and adductor operculi muscles (see above, p. 322). Although the dilatator operculi is some-
times, as in Pseudohemiodon, markedly reduced, it is always present and in Ancistrus it
extends from .the dorsal midline of the cranium and cranial roofing bones. In some taxa the
dilatator operculi originates from the hyomandibula rather than the cranium. The hyoman-
dibula is firmly and immovably articulated with the cranium, dorsally via an interdigitation
with the prootic, and posteriorly with the pterotic-posttemporal plate; the loss of hyoman-
dibular cranial fossae is synapomorphic for loricariid taxa. The sites of origin of the levator
and adductor operculi muscles are similar to those in other siluroids, although somewhat
modified owing to the ventral extension of the pterotic-posttemporal plate. What appears
to be the adductor operculi in one group of loricardiids (e.g. Panaque, Pseudacanthicus; p.
321) has shifted its point of insertion from the operculum to the preoperculum.

The opercular muscles in Astroblepus (Astroblepidae) have a relatively plesiomorphic ar-
rangement. The dilatator operculi is strongly developed and runs postero-ventrally from the
pterotic to its insertion on the antero-dorsal part of the operculum. Levator and adductor

CRANIAL MUSCLES OF LORICARIOID CATFISH 329

operculi muscles also originate from the pterotic and insert on the dorso-medial surface of
the operculum.

The levator arcus palatini is a stout muscle originating from the pterotic and sphenotic,
and inserting on the hyomandibula.

In Nematogenys (Nematogenyidae) the opercular muscles are complex, the dilatator
operculi is large and plesiomorphic in its angle of insertion, i.e. postero-ventral, cf. the
antero- ventral, apomorphic angle, in Loricariidae. The levator operculi inserts on the lateral
face of the operculum; the adductor operculi is in its more usual position of the medial face.
The levator arcus palatini is a large element with its lateral fibres interdigitating with the
antero- ventral fibres of the dilator operculi.

In the Trichomycteridae the dilatator operculi is hypertrophied and is separable into
lateral and medial parts. It originates from the underside of the frontal to pass beneath a
sphenotic bridge where its lateral profile becomes convex before inserting on the lateral side
of the long dorsally directed opercular process. The strap-like medial part of the muscle joins
the medio-distal area of this process. The ventral portion of the dilatator operculi joins an
aponeurosis with the dorso-posterior part of a muscle extending along the lateral face of the
preoperculum. This muscle may be either a disassociated part of the adductor complex, or
yet another segment of the dilatator operculi which has shifted ventrad.

The levator and adductor operculi also are hypertrophied and insert on the dorso- lateral
and medial margins of the operculum respectively. The hypertrophy and lateral insertion
of the dilatator operculi in the Nematogenyidae and the Trichomycteridae is viewed as
synapomorphic - such a condition is unknown elsewhere in siluroids.

Interrelationships of loricariids based on myological characters

Although of restricted taxonomic range in terms of numbers of 'genera' examined (but, see
below, p. 338), the anatomical data obtained from this investigation suffice to hypothesise
phylogenetic relationships for the groups of loricariid genera. The scheme is based on what
are considered to be derived states of jaw and opercular muscle characters. Two myological
synapomorphies unite the Loricariidae and Astroblepidae as a monophyletic lineage; these
are:

1 . Presence of a medial muscle stemming from the suspensorium and inserting on the upper
jaw (premaxilla). The muscle is recognized as a retractor pre maxillae (p. 327).

2. Complete medial separation of the protractor hyoideus (p. 319). Within this lineage, the
astroblepids retain a more generalized cranial muscle pattern and are considered to represent
the plesiomorphic sister group to the loricariids. The loricariidae are defined myologically
by the presence of a further medial adductor element inserting on the palatine (retractor
palatini; p. 327) and by the orientation of the dilatator operculi which is angled antero-
ventrally rather than postero-ventrally as in other siluroids and, indeed, other teleosts.

Within the Loricariidae, the following sequences of derived states in jaw and opercular
muscles are discernible and are represented as a cladogram in Fig. 14.

1. Dilatator operculi angled antero- ventrally; retractor palatini present:
LORICARIIDAE.

2. Reduction of opercular muscles and levator arcus palatini.

3. Lateral shift of retractor tentaculi: Hypoptopoma, Otocinclus.

4. Expansion of retractor tentaculi: Chaetostoma, Lipopterichthys, Hemipsilichthys.

5. Retractor palatini muscles meet in midline: Loricaria (part), Farlowella, Sturisoma.

6. Loss of levator arcus palatini: Loricaria (part).

7. Ventral division of retractor tentaculi inserts on palatine: Loricaria (part).

8. Loss of levator and adductor operculi: Pseudohemiodon, Hemiodonichthys.

9. Dilatator operculi originating from cranial roof, levator operculi from complex vertebrae:
Ancistrus, Lithoxus.

330

G. J. HOWES

10. Anterior shift of dilatator operculi; adductor operculi spans posttemporal-pterotic plate
and preoperculum: Panaque, Cochliodon, Stoniella, Pseudancistrus, Pseudacanthicus,
Acanthicus.

Because it is based on a single character complex, this synapomorphy scheme has limited
information content and, for the most part, is a series of unresolved polychotomies. In order
to test, and more fully resolve the scheme, various osteological characters have been utilised.

Fig. 14 Cladogram of loricariid taxa based on jaw and opercular muscle synapomorphies (see text,
p. 329). a = Hypoptopoma, Otocinclus; b = Chaetostoma, Lipopterichthys, Hemipsilichthys; c =
Loricaria (part); d = Pseudohemiodon, Hemiodonichthys, Crossoloricaria; e = Loricaria (part);
f—Farlowella, Sturisoma; % = Ancistrus, Lithoxus; h = Pseudacanthicus group; \ = Hypoptostomus,
Neoplecostom us. Pterygoplichthys.

Congruent osteological features in Loricariidae

Four osteological characters are analysed. Three that are functionally integrated with the
facial and opercular muscles, and one that has no direct relationship with the cranial muscu-
lature. They are: the lateral ethmoid; pterygoid channel; opercular bones and the lateroptery-
gium of the pelvic girdle.

Lateral ethmoid and nasal capsule

Alexander (1965) noted that in most siluroids the nasal cavities are bounded medially by
the ethmoid and posteriorly by the lateral ethmoid. In the majority of siluroids the palatine
lies adjacent to the mesethmoid and contributes no part to the nasal cavity. In Nematogenys,
(Nematogenyidae) the palatine is large and widely separated from the mesethmoid, forming
an angle between the lateral ethmoid and the maxilla. The nasal cavity is thus bounded later-
ally by the palatine (Fig. 15B).

CRANIAL MUSCLES OF LORICARIOID CATFISH

331

B

Fig. 15 Neurocrania, dorsal view of anterior region and upper jaw bones of A, Astroblepus festae
(BMNH 1864.11.4:18); B, Nematogenys inermis (BMNH 1883.11.27:49). Scales = 5mm. eth =
ethmoid; fr= frontal; le = lateral ethmoid; mx = maxilla; pmx = premaxilla; pal = palatine.

Synapomorphic for Astroblepus (Astroblepidae) species is an expanded palatine which
occupies the area between the lateral ethmoid and the mesethmoid. Its dorsal surface is exca-
vated and accommodates the entire nasal capsule (Fig. 15 A). Elsewhere this feature occurs
only in Trichomycterus rivulatus (Trichomycteridae). If it is regarded as synapomorphic with
that of Astroblepus, then the monophyly of Trichomycterus is in doubt.

In the Loricariidae the nasal capsule is contained within the lateral ethmoid. Entire encap-
sulation within the lateral ethmoid occurs also in the Callichthyidae (see Alexander, 1965).
In some species of Doradidae and Auchenipteridae the posterior part of the nasal cavity is
housed within the bone. Two conditions of encapsulation occur in loricariids; one in which
the nasal cavity is incompletely walled and the other where the cavity is entire.

The former is exemplified by Hypoptopoma where the nasal cavity, whilst being entirely
within the lateral ethmoid opens anteriorly as a shelf adjacent to the mesethmoid (Fig. 16 A).
The anterior border of the lateral ethmoid shelf meets the posterior border of the palatine.
A similar situation occurs in Sturisoma, Pterygoplichthys, Chaetostoma (Fig. 18B), Hemipsi-
lichthys and Pseudacanthicus. However, in all these taxa the anterior opening of the nasal
cavity is partially occluded by a medially directed process extending from the lateral wall
of the cavity (Fig. 16B). In Ancistrus the cavity is entire although a medially directed process
protruding from its lateral wall is discernible (Fig. 17). The nasal cavity is entire in Loricaria,
Farlowella (Fig. ISA) and Neoplecostomus.

The width of the anterior rim of the cavity is variable, being narrow in Farlowella and
long and shelf-like in Ancistrus (Figs 17 and ISA). The length of the pre-nasal cavity part
of the lateral ethmoid seems not to be coordinate with any elongation of the mesethmoid.
Indeed, topographical relationship between the nasal cavity and its surrounding elements
is maintained. In those taxa where there is mesethmoidal elongation, Farlowella and
Sturisoma, there is no corresponding proportional lengthening of the lateral ethmoid. The
lateral ethmoid serves as the site of origin for the retractor tentaculi muscle and it is likely
that extension of the bone's surface area corresponds with hypertrophy of the muscle, as
in Chaetostoma.

332

G. J. HOWES

Fig. 16 Neurocrania, dorsal view of anterior and orbital regions of A, Hypoptopoma carinatum
(uncat.); B, Pterygoplichthys multiradiatus (BMNH 1897.12.1:280). io = infraorbital; so = possible
supraorbital. Scale = 5 mm.

In Hypoptopoma and Otocinclus the lateral ethmoid margin is extended laterally with the
consequence that the retractor tentaculi muscle is also shifted laterally with respect to the
other adductor muscles. Unlike other loricariids, the eyes in these genera are entirely lateral.
It may be mentioned here that Gosline (1947) appears to have confused epidermal plates
with neurocranial elements. His figures of the skull of Hypoptopoma show a "dermospheno-
tic' lying lateral to the nasal cavity, i.e. forming the anterior border of the true neurocranium.
In fact Gosline's 'dermosphenotic' is the lateral ethmoid, and the unlabelled bone shown
lateral to it is perhaps the 1st infraorbital (Fig. 16 A). It is also pertinent to draw attention
to a dermal bone in some loricariids lying in the position of a supraorbital. One defining
character of siluroid fishes is that they lack a supraorbital; see Fink & Fink, 1981.

A naive hypothesis to account for the position of the nasal cavity in the lateral ethmoid
would be to assume, during phylogeny, a posterior migration of the nasal organ from its
plesiomorphic position, adjacent to the anterior mesethmoid, via an anterior lateral ethmoid
bridge. There is no extant 'transitional series' to indicate that this was the phylogenetic
history of the nasal cavity shift. On the contrary, development of an anterior extension to
the lateral ethmoid appears to be an apomorphic feature connected with the retractor
tentaculi muscle, rather than indicating a plesiomorphic retention of the pathway along
which the nasal organ shifted. Indeed, in assumed plesiomorphic members of the Loricarii-
dae, e.g. Neoplecostomus, the nasal cavity is entirely enclosed within the lateral ethmoid.

CRANIAL MUSCLES OF LORICARIOID CATFISH

333

ior4

Fig. 17 Neurocranium dorsal view of anterior and orbital regions of Ancistrus stigmaticus (BMNH
1901.5.8:10). Scale = 5 mm. Abbreviations as in previous captions.

In Astroblepus the lateral ethmoid has a plesiomorph otophysan morphology i.e. short,
and triangular; see Fink & Fink, 1981; Howes, 1981. The morphology and position of the
palatine is, however, derived and might be viewed as the precursory condition of nasal organ
reorientation in loricariids. This hypothesis is refuted, however, by the essentially plesiomor-
phic rod-shaped morphology of the palatine in the Loricariidae.

In the Callichthyidae the nasal organ is also enclosed within a cavity in the lateral ethmoid.
Unlike the loricariids, the nasal cavity is well- separated from the mesethmoid, which is short
and triangular, regarded as plesiomorphic in otophysans; see Fink & Fink, 1981; Howes,
1981. A superficial investigation of callichthyid osteology indicates that the lateral ethmoid
encapsulation of the nasal organ has been derived independently from that in loricariids.
More detailed studies of callichthyid and loricariid osteology are necessary before the feature
can be accepted as a synapomorphy.

Pterygoid channel

In the Loricariidae the extensor tentaculi muscles originate from the lateral face of the ptery-
goid. The term 'metapterygoid' is avoided here as the elements of the siluroid suspensorium
appear to have been misinterpreted by several authors (see Howes, 1983). The pterygoid in
loricariids contacts the lateral ethmoid along its anterior length. In some taxa there is a ptery-
goid ridge or channel and it is the medial face of this ridge that provides the site of origin
for the outer extensor tentaculi muscle, muscle e in the above text. In its more derived state,
the pterygoid ridge extends dorsally to contact the lateral ethmoid and so completely encloses
the extensor tentaculi in a tunnel.

A pterygoid ridge is weakly developed in Pseudacanthicus, Stoniella and Acanthicus; well-
developed in Pseudancistrus, Hemipsilichthys, Lipopterichthys and Chaetostoma, and
tunnel-like in Ancistrus, Pterygoplichthys, Panaque and Farlowella. In Hypostomus and
Cochliodon, the anterior and posterior parts of the ridge are produced dorsally, with con-

334

G. J. HOWES

B

Fig. 18 Neurocrania, dorsal view of anterior and orbital regions of A, Farlowella knerii (BMNH
1880.12.5: 206); B, Chaetostoma anomalus (BMNH 1903.6.30:102). Scales = 5 mm. Abbreviations
as previous captions.

nective tissue covering the intervening concavity. No development of the ridge is evident
in Hypoptopoma, Otocinclus, Loricaria, Sturisoma, Pseudohemiodon and Neoplecostomus.

In the Astroblepidae and Nematogenyidae, the pterygoid is extensive, and in astroblepids
its posterior margin, the 'metapterygoid' is sutured with the hyomandibula, as it is in
loricariids. There is no lateral pterygoid ridge present in either family.

The pterygoid channel character states are incongruent with the cladogram of muscle
characters. If the presence of a pterygoid ridge-tunnel is accepted as a synapomorphy, then
that lineage containing Chaetostoma and Hemipsilichthys must be closely united with that
represented by Hypostomus. So too, must Farlowella be united with Ancistrus. These seem
unlikely combinations in the light of other synapomorphies (p. 338) and the most parsimo-
nious explanation is to accept the pterygoid channel as having been derived independently
in several lineages.

The opercular series

Some loricariid genera possess a mechanism whereby the spine-bearing 'interoperculum' can
be rotated outward and the spines produced horizontally. Gosline (1947) was unable to find
an interoperculum in Ancistrus and was of the opinion that what had been referred to as
that bone was a patch of dermal plates. Gosline was only partially correct. He is certainly
right that in Ancistrus the spine-bearing element does not appear to be the interoperculum

CRANIAL MUSCLES OF LORICARIOID CATFISH

335

but a partially ossified ligament. None the less, this ligament actually connects the operculum
with the interoperculum which is immovably sutured to the preoperculum. A similar situ-
ation is present in Pseudacanthicus (Fig. 19B).

In Chaetostoma the situation is quite different. Here, the interoperculum is completely
movable and articulates with the condyle of the opercular ventral extension (Fig. 19 A). It
is ligamentously attached to the preoperculum. In all taxa with a spine-erecting mechanism
the operculum is elongate with an antero-ventral extension. There are, however, marked

op

Fig. 19 Pterygoid and opercular bones in medial views of A, Chaetostoma anomalus, B, Pseuda-
canthicus serratus. The hatched portion of the operculum in A indicates the insertion area of the
dilatator operculi muscle. hyo = hyomandibula; ih = interhyal; io = interoperculum; liop =
interopercular-opercular ligament; op = operculum; phc = pterygoid cavity for interhyal; po =
preoperculum; pte = pterygoid; ptt = posttemporal-pterotic plate; q = quadrate.

336 G. J. HOWES

differences in morphology distinguishing two groups of taxa. In Pseudacanthicus, Ancistrus
and Lithoxus there is a single articulation of the operculum with the hyomandibula at the
point of the dilatator operculi muscle insertion. In Chaetostoma and Lasiancistrus there is
a double articulation, one with the hyomandibular condyle and the other, via a dorsal
process, with the medial surface of the preoperculum (Fig. 19 A). These two groups of taxa
also differ in the topographical arrangement of the suspensorial bones (cf. Figs. 19A & B).
From these observations it is suggested that the 'interopercular' spine-erecting mechanism
has been derived independently in two lineages of loricariids.

In some Trichomycteridae there is also an an tero- ventral process of the operculum which
articulates with a socket on the medial face of the spine-bearing interoperculum, e.g.
Branchioica, Pareiodon. If the Trichomycteridae is a monophyletic assemblage (see remarks
on p. 331 concerning the genus Trichomycterus) then it is clear that a similar mechanism
to that of loricariids is the result of parallelism (sensu Holmes, 1980: 49). The function of
interopercular spines is discussed on p. 343.

The lateropterygium

In the Loricariidae and Astroblepidae the pelvic girdle bears a lateral, anteriorly directed
process named by Regan (1904) a 'rudimentary ventral ray' and by Shelden (1937) a 'laterop-
terygium', the name adopted here. The lateropterygium articulates in condylar fashion with
the lateral process of the basipterygium and is anchored laterally via a ligament to an adjacent
body scute.

Shelden (1937) made a thorough study of the morphology of the lateropterygium as well
as the pelvic muscles in Astroblepus and various loricariids. He found that in Farlowella
a lateropterygium was lacking, a condition Shelden thought might be an artefact of pre-
paration. I can confirm that this is not an artefact and that the lateropterygium is absent also
in Loricaria, Sturisoma, Hemiodonichthys and Pseudohemiodon. As these taxa are admitted
within the Loricariidae on the basis of other synapomorphies, e.g. specific form of scute
development; structure of swimbladder encapsulation; caudal fin skeletal morphology and
jaw musculature, the absence of a lateropterygium is viewed as a secondary loss in this
lineage, = Loricariinae.

In the Astroblepidae, the lateropterygium is a disc-like plate and the basipterygium has
posteriorly directed, elongate processes. The lateropterygium in Loricariidae varies from
short and broad, as in Neoplecostomus, to long and rod-shaped as in Pterygoplichthys. There
are no posterior processes of the basipterygium.

Regan (1904) considered the lateropterygium to be a rudimentary anterior ventral fin ray
which had become '. . . internal and directed forwards'. Shelden (1937) on the other hand,
thought the element to be homologous with the lateral cartilage in other siluroids.

Arratia et al. (1978) discussed the lateropterygium and the 'pelvic splint' in Nematogenys
and the Trichomycteridae. They concluded that the elements are not homologous. However,
they then considered that as neither lateropterygium nor pelvic splint has been recorded from
other siluroids this was ' . . . evidence of some relationship between these groups of South
American fishes', thus implying homology. These authors further confuse the issue by
referring to the primitive nature of the 'pelvic splint' on the grounds of its occurrence in
'lower' teleosts (vide Patterson, 1964).

A pelvic splint occurs in many characoids and cyprinoids and from its shape and position
appears to be homologous with that similar element in other teleosts (Patterson, 1964: 444).
Among siluroids a supplementary pelvic element is recorded only in the Trichomycteridae,
Astroblepidae, Nematogenyidae, Loricariidae and Diplomystidae. In the Trichomycteridae
the morphology of the bone is similar to that in other otophysans except that it is associated
with the pelvic musculature and is detached from the first pelvic ray (see Arratia et al., 1978).

CRANIAL MUSCLES OF LORICARIOID CATFISH

337

Within the Loricariidae, the lateropterygium appears in its simplest form in the Hypopto-
pominae where in Otocinclus it resembles the pelvic splint of trichomycterids and diplomys-
tids but, unlike the conditions in those taxa, it is orientated antero-dorsally instead of
latero-posteriorly. This condition suggests that the lateropterygium is a pelvic splint which
has become reorientated and articulated with the pelvic bone (as suggested by Regan, 1904;
see above). Thus, the lateropterygium would be the homologue of the pelvic splint (rudimen-
tary spine) in other teleosts, but its peculiar nature and articulation with the pelvic bone
is a derived condition limited to the Loricariidae and Astroblepidae only.

Gosline (1974) was of the opinion that the lateropterygium was not a 'good classifying
character, and that its presence, size and shape depend more or less on the extent of the
plating of the sides and abdomen'. There must, as Gosline suggests, be a correlation between
body shape, scute morphology and the development of the lateropterygium. Nevertheless,
the presence and similarities of the bone in various taxa must indicate a shared ancestral
condition. Thus, it is hypothesised that especially elongate rod-like lateropterygia are
derived, whereas the short, broader morphology represents the plesiomorph condition.

Shelden (1937) concluded from his study of loricariid pelvic girdle morphology that
Ancistrus and Farlowella represented two separately advanced states, a conclusion supported
by this study. Shelden accounted for the function of lateropterygium as increasing the
manoevrability of the pelvic girdle.

Myological + osteological characters

Utilizing derived character states from both myological and osteological complexes detailed
above, together with those already noted in the literature (particularly Gosline, 1947), the
following scheme of relationships is hypothesized and shown in Fig. 20.

1 . Lateropterygium present

2. Retractor premaxillae muscle present

3. Protractor hyoideus muscle medially divided

4. Nasal capsule confined to palatine

5. Gill-rakers absent

6. Lateropterygium disc-like

7. Lack of hyomandibulae fossae

8. Nasal capsule confined to lateral ethmoid

9. Dilatator operculi muscle antero-ventrally
orientated; retractor palatini muscle present

10. Body encased in scutes

1 1 . Reduction of opercular muscles, dilatator

operculi confined to hyomandibula (Neoplecostomus)

12. Expansion of retractor tentaculi muscle

13. Reduction of posterior cranial elements (e.g. supraoccipital, posttemporal-
pterotic plate)

14. Jaws lengthened and widened

15. Lateral cheek spines (Hemipsilichthys)

16. Interopercular spine mechanism (Chaetostoma, Lasiancistrus,
Lipopterichthys

17. The single character uniting Hypoptopoma and Otocinclus with the
loricariine genera is the midline contact of the ventro-lateral scutes between
the anus and anal fin origin (Gosline, 1947: 96) versus a single, azygous prea-
nal plate in other loricariids. Synapomorphic for this lineage is the lateral
position of the retractor tentaculi. On the basis of the characters used, it is
not possible at this point to resolve a trichotomy of the neoplecostomine-
chaetostomine-hypoptopomine (loricariine) lineages.

Astroblepidae

+

Loricariidae

Astroblepidae

Loricariidae

Neoplecos-

tominae

Chaetostominae

Hypopto-

pominae

338 G. J. HOWES

18. Retractor palatini muscles meet in the midline

19. Caudal peduncle depressed, caudal vertebrae elongated, lateral scutes
reduced.

20. Lateropterygium lost.

21. 'Adipose' fin lost; reduction of caudal fin ray number (10-14). These
characters are of dubious value; the 'adipose' fin is also absent in some
neoplecostomines, but it is noted here because of its complete absence in all
loricariine taxa; the caudal fin ray number may be reduced in other lineages
but appears consistently so in loricariines.

22. Mesethmoid elongated (Sturisoma + Farlowella)

23. Extensor tentaculi muscle enclosed in pterygoid tunnel (Farlowella +
Loricaria platystoma); tendon of retractor ischii muscle runs through body-
armour canals (Farlowella; see Shelden, 1937: 63).

24. Palatine lamellate, usually convex; 3rd, 4th and often 5th ceratobranchials
expanded (?all taxa)

25. Ventral part of retractor tentaculi inserts on palatine (Loricaria, part)

26. Postorbital notch

27. Jaw teeth reduced or absent (Loricaria, part)

28. Palatine laterally compressed, extensor tentaculi inserts on its mid-
ventral border; papillae developed over pterygoid. (Pseudohemiodon,
Hemiodonichthys)

29. Lower pharyngeal bones edentulous (?all taxa; see Gosline, 1947: 86)

30. Lateropterygium well-developed

31. Pterygoid channel or tunnel (Hypostomus, Pterygoplichthys, Cochliodon)

32. Pseudointeropercular mechanism entirely or partially developed (Peckoltia,
Hemiancistrus)

33. Anterior shift dilatator operculi medial to orbit, adductor operculi bridges
posttemporal-pterotic plate and preoperculum; upper jaw reduced, pre-
maxillae coalesced (Panaque, Stoniella, Pseudacanthicus, Acanthicus,
Pseudancistrus)

34. Dilatator operculi originates from cranial roof, levator operculi from complex
vertebrae; levator arcus palatini joins tendon of dilatator operculi. (Ancistrus,
Lithoxus)

Loricariinae

Hypostominae

The cladogram constructed from nested sets of myological and osteological characters
is, for the most part, congruent with that constructed from myological characters alone. It
will be necessary to utilize other character sets, particularly those involving the morphology
of the gill-arches, swimbladder encapsulation and the pectoral girdle in order to more highly
corroborate this proposed phylogeny.

Classification of Loricariidae

The most recent synopsis of the Loricariidae is that of Isbriicker (1980) wherein are recog-
nized six subfamilies, the Neoplecostominae, Lithogeneinae, Hypostominae, Ancistrinae,
Hypoptopominae and Loricariinae. The genera included in these subfamilies follows, for the
most part, those of Gosline (1947), Isbriicker, however, retains Regan's (1904) concept of
the Neoplecostominae as monotypic by referring to it only Neoplecostomus, whereas Gosline
has included 12 genera in the subfamily. Few representatives of those genera recognised by
Gosline have been available for my study and their placement in one or other of the sub-
families recognised here is open to question. I have recognised Neoplecostomus as a sole
representative of the Neoplecostominae purely on its retention of plesiomorph characters.

In the cladistic scheme presented in Fig. 20 the major lineages are equivalent to subfamilies
and the included taxa are broadly those currently contained in the Hypostominae, Hypopto-
pominae and Loricariinae. Genera formerly incorporated in the Ancistrinae are here divided
between the Hypostominae and Chaetostominae = Chaetostomidi, Fowler, 1958. No speci-

Neo

CRANIAL MUSCLES OF LORICARIOID CATFISH 339

Chaeto Loricariinae Hypt Hypostominae

J

m

34-

-33

32-

29

-31

-10

1-3

Fig. 20 Cladogram of the Astroblepidae and Loricariidae based on muscle and skeletal synapo-
morphies (see text, p. 337). a = Neoplecostomus (Neo = Neoplecostominae); b — Hemipsilichthys,
c=Chaetostoma, Lasiancistrus, Lipopterichthys, (Chaeto = Chaetostominae); d = Slurisoma\
t = Farlowella\ {—Loricaria (part, strigilata, simillima), Metaloricaria; g = Pseudohemiodon,
Hemiodonichthys, Crossoloricaria; h = Loricaria (part, e.g. cataphractd); \ = Hypoptopoma,
Octocinclus, Parotocinclus, IMicrolepidogaster, (Hypt = Hypoptopominae); ] — Pekoltia, Hemiancis-
trus\ k = Ancistrus, Lithoxus; \-Panaque, Stoniella, Pseudacanthicus, Acanthicus, Pseudancistrus;
m = Hypostomus, Pterygoplichthys, Cochliodon.

mens of Lithogenes have been examined and thus no comment can be made concerning the
status of the monotypic subfamily Lithogeneinae.

Isbriicker (1980) lists 70 loricariid genera; it is likely that many are synonyms, particularly
those recognised on the basis of single specimens. It is pertinent to note that of the 70 genera,
2 1 are monotypic and so might merely reflect the generic concepts of their respective authors.
The characters used to define many loricariid genera appear to depend on meristic and dental
differences, varying numbers of body scutes and lower lip morphology. Most of these
characters appear to be gradal. A character that appears to be significant in the classification
of Loricaria species is the presence in some of a posterior orbital notch or fossa. The fossa
is variously developed, varying from a slight indentation to a deep, medially directed
channel. There is no organ or vessel contained within the fossa and it is floored with connec-
tive tissue. Its function is unknown but may serve to enlarge the posterior visual field.
Presence of a post-orbital fossa is regarded as synapomorphic for those Loricaria species in

340 G. J. HOWES

which it occurs. In a sample of 16 species belonging to 7 'genera' the following distribution
of the fossa is recorded:

Absent: Crossoloricaria, Spatuloricaria, Loricaria, Loricariichthys
Slightly developed: Paraloricaria, Loricaria, Ricola
Well developed: Rineloricaria, Loricaria, Loricariichthys

One genus, Loricaria, appears in all three morpho-groups and Loricariichthys appears in
two. This result indicates that the present generic classification of the Loricariinae is possibly
a paraphyletic one. In support of this statement I find that in Loricaria sensu stricto as repre-
sented by the type species, L. cataphracta, the retractor tentaculi muscle inserts partly on
the palatine (see p. 315). This feature is restricted to those taxa which lack the post-orbital
fossa, or at best have it slightly developed, e.g. 'Limatulichthys'punctatus; 'Pseudoloricaria'
laeviuscula.

Other characters which override current generic allocations of loricariine species are the
ligamentous suspension of the upper jaw, the length of the lateral ethmoid and morphology
of the palatine.

There are two basic types of ligamentous jaw suspension in Loricaria, (1 ) where the maxilla
is attached to a ventral process of the vomer via a single, folded ligament; there is also a
small rostral cartilage, and (2) where there are several ligaments connecting the maxilla with
the vomer and there is a large rostral cartilage (Figs 2 1 A & B). The former type of jaw suspen-
sion is present in Loricaria labialis, assigned by Isbriicker to Loricariichthys, L. cataphracta,
Sturisoma, Farlowella and Hemiodonichthys. The latter occurs in Loricaria jubata and L.
brunneus which Isbriicker refers to Rineloricaria and Loricariichthys respectively.

pmx

mx

vo

vo

rca

pmx

B

Fig. 21 Ligamentous connections of the upper jaws in A, Loricaria labialis (BMNH 1934.8.20:
368-388) and B, L. jubata (BMNH 1914.5.18: 68-72). Ventral views, both drawn to same scale.
mx = maxilla; pmx = premaxilla; rca = rostral cartilage; vo = vomer.

CRANIAL MUSCLES OF LORICARIOID CATFISH 341

The length of the lateral ethmoid and morphology of the palatine in Loricaria species are
perhaps correlated. For example, in L. cataphracta, the lateral ethmoid is long and shelf-like,
the palatine narrow and lamellate with a long posterior process whereas in 'Rineloricaria'
teffeana the lateral ethmoid is short and broad and the palatine is also short and broadly
convex, lacking a posterior process.

Isbriicker (1981) in discussing the Loricariini notes that there are ' . . .several different
types of secondary sexual dimorphism . . . ' within the group. If this is so, then here is a
further indication of the group's paraphyly.

Discussion

The interrelationships of the Loricariidae

In the latest classification of the order Siluroidei, Chardon (1968) includes the Loricariidae,
Astroblepidae and Callichthyidae in the superfamily Loricarioidae within the suborder
Loricarioidei. Chardon recognises two other superfamilies as constituting the suborder, the
Aspredinoidae and the Trichomycteroidae.

It is doubtful that the aspredinids are closely related to the loricariids and trichomycterids.
Chardon used an assortment of apo- and plesiomorph characters involving the swimbladder
and Weberian apparatus to unite his groups. There are marked differences in the structure
of the swimbladder- Weberian system between the Aspredinidae and the Loricariidae as
follows:

Aspredinidae Loricariidae

Swimbladder Not encapsuled encapsuled

Anterior centra Elongate short

4th parapophysis Short contributes to swimbladder

capsule

On the other hand, there are several synapomorphies of the swimbladder- Weberian system
and caudal fin skeleton that link the Loricariidae with the Astroblepidae and Callichthyidae
(see Alexander, 1964, 1965; and Lundberg & Baskin, 1969; also below).

As noted above p. 323 the astroblepines have been variously treated as a subfamily of
the Loricariidae (Regan, 1911; Gosline, 1947) and as a distinct family (Eigenmann, 1910).
There is no doubt that on the basis of synapomorphies presented here the astroblepines must
be treated as the sister group to the Loricariidae. Following Chardon's (1968) scheme of
classification both families would form the super- family Loricarioidae — this is not, how-
ever, in accordance with the cladistic scheme presented in Fig. 22.

Bailey & Baskin (1976) described a new taxon which they regarded, provisionally as a sub-
family of the Loricariidae, the Scoloplacinae. The authors pointed out that its inclusion in
the family would necessitate a 'significant expansion' of the Loricariidae. Baskin's (1973,
unpublished) cladogram places Scoloplax as the sister group to the Loricariidae + the Astro-
blepidae, i.e. at family rank. With this I would concur and also follow Isbriicker (1980) in
recognising the Scoloplacidae.

The pattern of phylogenetic relationships between the Loricariidae + Astroblepidae and
the Nematogenyidae, Trichomycteridae, Callichthyidae and Scoloplacidae has still to be
resolved. Baskin (1973, unpublished thesis) has produced a phylogenetic scheme which is
essentially like that presented here (Fig. 22). There are, however, several conflicting pre-
sumed synapomorphies not taken into account by Baskin. For example the Nematogenyidae
shares only with the Trichomycteridae a derived configuration of the opercular musculature
(p. 329), but shares only with the Diplomystidae a similarly papillate branchiostegal
membrane and buccal roof tissue. Again, the Callichthyidae shares what appear to be
derived features of swimbladder encapsulation with the Astroblepidae, Loricariidae and
Scoloplacidae, but a premaxillary-ethmoid hinge joint only with the two former families.
Lateral ethmoid encapsulation of the nasal organ occurs only in the Loricariidae and
Callichthyidae.

342

G. J. HOWES

f

<V

11 • »

10- •

9i»

12"

Fig. 22 Hypothesised relationships of the Loricariidae and Astroblepidae. Node I =swimbladder
encapsuled, divided into separate vesicles; some part of the cranium contributing to encapsulation;
odontodes on fin rays and/or body. Node 2=Claustrum and intercalarium lacking from the
Weberian vertebral series. Node 3 = Posttemporal contributing to distal portion of the swimbladder
capsule; derived hypural fusion pattern; low number of principal caudal fin rays. Node 4 =
connecting bone between the 1st rib and the 2nd pterygiophore. Node 5 = muscle characters listed
in this paper, associated with reverted lip; lateropterygium (see p. 336; 6 fused anterior centra. Line
6 = papillate branchiostegal membrane and buccal roof tissue. Line 7 = lateral insertion of the
dilatator operculi muscle. Line 8 = odontodes on opercular bones. Line 9 = Ethmoid-premaxillary
hinge joint. Line 10 = nasal organ encapsuled in lateral ethmoid. Line ll=rows of lateral plates
partly or entirely encasing body. Line 12 = rostal epidermal ossifications bearing odontodes.
Characters 6-12 are incongruent with 1-5 and so indicate either independent derivation (9, 10 in
Callichthyidae) or secondary loss (8, 11, 12 in Astroblepidae).

The characters listed above are compiled from Gosline (1947), Lundberg & Baskin (1969), Bailey
& Baskin (1976), Baskin (unpublished thesis, 1973) and personal observation.

CRANIAL MUSCLES OF LORICARIOID CATFISH 343

The absence in the Astroblepidae of certain characters linking the Loricariidae, Scolo-
placidae, Callichthyidae and Trichomycteridae (i.e. opercular odontodes, body plates) is
most parsimoniously explained by secondary loss (see opinions of Bhatti, 1938 and Gosline,
1947 concerning loss of armour in Astroblepus).

As yet, comparative anatomical studies on the various genera presently included in the
Callichthyidae are lacking. Such data as are available are summarized in the cladistic scheme
presented here (Fig. 22).

Functional considerations

Several pointers have emerged in the course of this study as to the functional significance
of certain myological features. In particular, the medially divided protractor hyoideus signi-
fies independent mobility of the separate halves of the lower jaw. Aquarium observations
on a variety of loricariid taxa have confirmed this (pers. obs.). The intermandibularis is only
weakly attached to the two halves of the mandible and the bulk of the muscle runs through
the lip tissue. It would appear that in loricariids the principal function of the muscle is to
activate the lip as a sucker. Alexander (1965) has given an account of lip movement associ-
ated with respiration.

The diversity of jaw morphology ranges from long, broad upper and lower jaws (Chaeto-
stoma) to the short, deep inwardly curved lower jaw and coalesced, almost circular, upper
jaw of Pseudacanthicus. In both morphotypes the jaws appear highly mobile, but the angles
of their respective muscle insertions indicate quite different manipulative abilities.

In astroblepids and loricariids, the upper jaw is free from the restraint imposed by the
ankylosed mesethmoid, common to the majority of siluroids. Only in callichthyids has the
upper jaw a similar, ligamentous, attachment to the mesethmoid, although here its vertical
rotation appears limited through firm attachment to the palatines.

The palatine appears to play a somewhat different functional role in loricariids as com-
pared with other siluroids. Its lack of firm attachment to the ethmoid seems to impart a
greater degree of mobility to the bone, and the postero- ventral attachment of part of the
extensor tentaculi muscle provides a degree of downward movement lacking in other silur-
oids. This is particularly marked in the Pseudohemiodon group where the extensor tentaculi
has shifted its position to the centre of the centre of the palatine and there is a tendinous
palatine-maxillary-premaxillary linkage (see Fig. 5B).

An inferior, sucker-like mouth comparable with that of loricariids occurs elsewhere
amongst siluroids in the African genus Euchilichthys and the Asian Glyptothorax and Glyp-
tosternon. In these taxa the muscular insertions to the upper jaw differ from those in lori-
cariids in that the retractor tentaculi inserts for the most part on the premaxilla with a
tendinous outer portion running to the maxilla. The palatine is short and is actuated by an
extensor tentaculi passing into its medial face. The premaxilla is overlapped by an anterior
extension of the mesethmoid and appears to have less mobility than in loricariids.

As noted above, p. 342, loricariids and their relatives have a head and body covering of
odontodes and in some taxa these have become elongate and collected in certain areas to
form clumps of spines. The spines are of two kinds, those that are embedded in the cheek
tissue to which the pediments are attached by ligament (see Bhatti, 1938) and those that
are attached to the interoperculum, embedded in small plates. Both types of spine can be
shed; according to Moodie & Power (1982), in Loricaria uracantha, cheek spines develop
over a 64 day period and are lost within 148 days. Scars on the interoperculum of male
Chaetostoma specimens indicate that the spines have been shed (pers. obs.). Allison, (1980;
pers. comm.) reports aquarium observations on the use of interopercular spines in Ancistrus
sp. When two males sight one another their initial reaction is to open the interopercular
flap and evert the spines rapidly on both sides simultaneously. If one or other fish remains
undeterred a 'combat' situation develops with the individuals aligning themselves side by
side, everting their opposing spines (those on the blind side remaining withdrawn) and engag-
ing them by sideways movements of the head. These movements are made by strong lateral

344 G. J. HOWES

curvatures of the caudal peduncle and tail. These observations suggest that at least in Ancis-
trus the interopercular spine function is one of display. It is noted that the spines are not
as well-developed in females ofAncistrus who nevertheless do display their spines by lateral
movements when approached by conspecifics whilst feeding (Allison, pers. comm.).

The fixed cheek spines of Hemipsilichthys and various Loricaria species is also a sexually
dimorphic character. In males the cheeks are fringed with long, closely packed spines, whilst
those of the females bear small, widely spaced spines. Moodie & Power (1982) report on
the use of pectoral fin and cheek spines in Loricaria uracantha where the male has the ability
to rotate forward its pectoral spines so that they lie adjacent to the cheek spines and so
probably act as clasping organs during spawning.

According to Eigenmann (1918) trichomycterids use their opercular spines as holdfasts
to the substrate. The same function would seem to operate in parasitic trichomycterids, the
only difference being the nature of the substrate. Such may be the plesiomorph function of
the interopercular spines in loricariids. Aquarium observations suggest that some loricariids
use their spines to secure a position in the substrate (Allison, pers. comm.; pers. obs.).

The dilatator operculi muscle is well-developed only in those taxa with a pseudointeroper-
cular spine erecting mechanism, Ancistrus, Pseudacanthicus etc., and its role in activating
the operculum in respiratory movements seems restricted. Respiratory activity probably falls
largely on the adductor hyohyoidei which extend well up the medial face of the operculum.
Often the last branchiostegal ray is attached to the opercular face by the musculature or at
least an intervening tissue fascia. The tight seal of the operculum to the cleithrum also
suggests its rather passive role in respiration.

Acknowledgements

My sincere thanks are due to Drs Jon Baskin, Isaac Isbriicker and Mary Power for most
informative discussions on loricariid fishes, and to Dr Reeve Bailey (University of Michigan,
Museum of Zoology) for the loan of Scoloplax specimens.

Dr Humphry Greenwood's critical reading and discussion of the manuscript has made
for a great deal of improvement, and to him I am most grateful.

Special thanks go to David Allison of the Catfish Association of Great Britain for providing
me with so much information on the behaviour of loricariids.

References

Alexander, R. McN. 1964. The structure of the Weberian apparatus in the Siluri. Proc. Zool. Soc.

Lond., 142 (3): 4 19-440.

1965. Structure and function of the catfish. J. Zool. Lond., 148: 88-152.

Allison, D. 1980. Family Loricariidae, continued. Catfish Assn. Great Britain Magazine 25: 16-17.
Arratia, F., Chang, G., Menu- Marque, S. & Rojas, G. M. 1978. About Bullockia gen. nov., Tricho-

mycterus mendozensis n.sp. and revision of the family Trichomycteridae (Pisces, Siluriformes). Stud.

Neotrop. Fauna & Environ., 13: 157-194.
Angelscu, V. & Gneri, F. S. 1949 Adaptaciones del aparato digestive al regimen alimenticio en algunos

peces del rio Uruguay y del Rio de la Plata. I. Revta. Inst. nac. Invest. Cienc. nat. Mus. argent. Cienc.

nat. Bernardino Rivadavia. Ciencias Zool. 1 (6): 161-272.
Bailey, R. M. & Baskin, J. N. 1976. Scoloplax dicra, a new armoured catfish from the Bolivian

Amazon. Occ. Pap. Mus. Zool. Univ. Mich. 674: 1-14.

Baskin, J. N. 1973. Structure and relationships of the Trichomycteridae. Unpublished thesis.
Bhatti, H. K. 1938. The integument and dermal skeleton of Siluroidea. Trans. Zool. Soc. Lond., 24,

(1): 1-102.
Chardon, M. 1968. Anatomic comparee de 1'appareil de Weber et des structures connexes chez les

Siluriformes. Annls Mus. Afr. cent. Ser. 8° Sci. Zool., 169: 1-277.

CRANIAL MUSCLES OF LORICARIOID CATFISH 345

Eigenmann, C. H. 1910. Catalogue of the fresh- water fishes of tropical and south temperate America.

Rep. Princeton Univ. Exped. Patagonia 3(4) (Zool.): 375-51 1.

1918. The Pygidiidae, a family of South American catfishes. Mem. Carnegie Mus, 1 (5): 259-398.

1928. The fresh-water fishes of Chile. Mem. Nat. Acad. Sci. 22: 1-63.

Fink, S. V. & Fink, W. 1981. Interrelationships of the ostariophysan fishes (Teleostei). Zool. J. Linn.

Soc. 72 (4): 297-353.

Fowler, H. W. 1958. Some new taxonomic names of fishlike vertebrates. Notul. nat., 310: 1-16.
Gosline, W. A. 1947. Contributions to the classification of the loricariid catfishes. Arq. Mus. Nac. Rio

de Janeiro, 41: 79-134.
Gregory, W. K. 1933. Fish skulls. A study of the evolution of natural mechanisms. Trans. Amer.

philos.Soc.,23(2):15-48\.
Holmes, E. B. 1980. Reconsideration of some systematic concepts and terms. Evolutionary Theory,

5: 35-87.
Howes, G. J. 1981. Anatomy and phylogeny of the Chinese major carps Ctenopharyngodon Steind.,

1866 and Hypophthalmichthys Blkr, 1860. Bull. Br. Mus. nat. Hist. (Zool.) 41 (1): 1-52.
1983. Problems in catfish anatomy and phylogeny as exemplified by the Neotropical Hypophthal-

midae (Teleostei: Siluroidei). Bull. Br. Mus. nat. Hist. (Zool.), 45(1): 1-39.
Isbrucker, I. J. H. 1980. Classification and catalogue of the mailed Loricariidae (Pisces, Siluriformes).

Versl. Techn. Geg., Inst. Taxon. Zool. (Zool. Mus.) Univ. Amsterdam, 22: 1-181.
1981. Revision of Loricaria Linnaeus, 1758 (Pisces, Siluriformes, Loricariidae). Beaufortia, 30

(3): 5 1-96.
Leege, C. O. 1922. Der Rumpfpanzer der Panzerwelse und seine Skelettbeziehungen (Plecostomus

angipinnatus n. sp., Callichthys callichthys L. und Corydoras paleatus Jen.). Jena Z. Naturw., 58

(NS)51 (2): 145-270.
Lundberg, J. G. & Baskin, J. N. 1969. The caudal skeleton of the catfishes, order Siluriformes. Am.

Mus. Novitates, 2398: I^t9.
Matthes, H. 1963. A comparative study of the feeding mechanism of some African Cyprinidae (Pisces,

Cypriniformes). Bijdr. Dierk., 33: 3-35.

Mithel, M. 1964. The cranial nerves of the sisorid catfish Bagarius bagarius. Copeia 1964 (4): 673-678.
Moodie, G. E. E. & Power, M. 1982. The reproductive biology of an armoured catfish, Loricaria

uracantha, from Central America. Env. Biol. Fish., 1 (2): 143-148.
0rvig, T. 1977. A survey of odontodes ('dermal teeth') from developmental, structural, functional, and

phyletic points of view. In: Problems of vertebrate evolution ed. S. Mahala Andrews, R. S. Miles

& A. D. Walker. Linn. Soc. Symp. Ser., 4: 53-75.
Peyer, B. 1922. Uber die Flossenstacheln der Welse und Panzerwelse, sowie des Karpfens. Morph.

Jahjr. 51: 493-554.
Regan, C. T. 1904. A monograph of the fishes of the family Loricariidae. Trans, zool. Soc. Lond.,

17(3): 191-350.
1911. The classification of the teleostean fishes of the order of Ostariophysi. 2. Siluroidea. Ann.

Mag. nat. Hist., 8 (8): 553-577.
Starks, E. C. 1926. Bones of the ethmoid region of the fish skull. Stanford Univ. Publ. Biol Sci., 46:

137-338.
Vari, R. P. 1979. Anatomy, relationships and classification of the families Citharinidae and Disticho-

dontidae (Pisces, Characoidea). Bull. Br. Mus. nat. Hist. (Zool.), 36 (5): 261-344.
Vaz-Ferreira, R. & Senorans, J. S. 1971. Oviposicion e incubacion de Plecostomus alatus Castelnau,

en cuevas. Bol. Soc. Zool. Uruguay, 1: 12-17.
Vetter, B. 1878,. Untersuchungen zur vergleichende Anatomic der Kiemen- und Kiefermuskulatur

der Fische II. Jena Z. Naturw., 12 (3): 431-550.
Winterbottom, R. 1974. A descriptive synonymy of the striated muscles of the teleostei. Proc. Acad.

nat. Sci. Philad., 125 (12): 225-317.

Manuscript accepted for publication 13 December 1982

)

:

-.

L / UL \ : - |

I

' •

British Museum (Natural History)

The Tilapiine fishes of the genera

Sarotherodon, Oreochromis and Danakilia

Dr Ethelwynn Trewavas

The tilapias are cichlid fishes of Africa and the Levant that have become the subjects of
fish-farming throughout the warm countries of the world. This book describes 41 recognized
species in which one or both parents carry the eggs and embryos in the mouth for safety.
Substrate-spawning species, of the now restricted genus Tilapia, are not treated here.

Three genera of the mouth-brooding species are included though in one of them, Danakilia,
the single species is too small to warrant farming. The other two, Sarotherodon, with nine
species, and Oreochromis, with thirty-one, are distinguished primarily by their breeding
habits and their biogeography, supported by structural features. Each species is described,
with its diagnostic features emphasised and illustrated, and to this is added a summary of
known ecology and behaviour. Conclusions on relationships involve assessment of parallel
and convergent evolution. Dr Trewavas writes with the interests of the fish culturists, as well
as those of the taxonomists, very much in mind.

About 480pp, 1 88 illustrations include halftones, diagrams, maps and graphs.
Extensive bibliography. Publication due 1983. 0 565 00878 1

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Problems in catfish anatomy and phytogeny exemplified by the Neotropical
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Bats (Mammalia: Chiroptera) from Indo- Australia. By J. E. Hill

On Macropleurodus, Chilotilapia (Teleostei: Cichlidae) and the
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Miscellanea

The cranial muscles of loricarioid catfish es, their homologies and value as
taxonomic characters (Teleostei: Siluroidei). By G. J. Howes

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Miscellanea

24NOVI983

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Contents

Page

A lectotype for Deuterammina (Deuterammina) rotaliformis (Heron-Allen &
Earland) and new trochamminids from E. Ireland (Protozoa: Foraminiferida). By
P. Bronnimann & J. E. Whittaker 347

Notes on Atlantic and other Asteroidea. 3. The families Ganeriidae and Asterinidae,

with description of a new asterinid genus. By Ailsa M. Clark ...... 359

Peniculus haemuloni, a new species of copepod (Siphonostomatoida: Pennellidae)

parasitic on Haemulon steindachneri from Ubatuba, Brazil. By P. D. Alexander 381

A lectotype for Deuterammina (Deuterammina)
rotaliformis (Heron- Allen & Earland) and new
trochamminids from E. Ireland (Protozoa:
Foraminiferida)

P. Bronnimann

Laboratoire de Paleontologie, Universite de Geneve, 13, rue des Maraichers, 1211 Geneve
4, Switzerland

J. E. Whittaker

Department of Palaeontology, British Museum (Natural History), Cromwell Road, London
SW7 5BD

Introduction

In their 'Report on some Recent Foraminifera found off the coast of Dublin and in the Irish
Sea', Balkwill & Wright (1885) illustrated four views, two umbilical and two spiral (pi. 13,
figs 11 a, b and 12a, b) of agglutinating foraminifera which they refer to as 'Trochammina
inflata Montagu sp. var.'; no formal description was given. Twenty-six years later Heron-
Allen & Earland (1911 : 309) encountering in shore sands of Selsey Bill, Sussex, a single
trochamminid which they regarded as identical with that reported by Balkwill & Wright,
sent it to Joseph Wright in Ireland for a second opinion. Wright indeed agreed with their
identification. In their paper, Heron- Allen and Earland cite it as 'Trochammina rotaliformis
J. Wright MS' and write '. . . as Mr. Wright will, we understand, describe and figure the
variety shortly under the name rotaliformis, we refrain from further particulars of the variety
which has not hitherto been accorded a distinctive name, although presenting well-marked
characteristics'. Wright did not publish this description, but Heron-Allen & Earland by
referring to Balkwill & Wright's figures, while citing Wright's manuscript name, themselves
made the name available. Article 12 of the International Code of Zoological Nomenclature
states 'that names published before 1931 ... must have been accompanied by a description,
definition or indication [Art. 16]'; an indication being constituted under Article 16a (i) by
'a bibliographic reference to a previously published description, definition or figure'. An
'indication' was therefore clearly given, furthermore, Heron-Allen and Earland are the
authors of the name, not Wright, with 191 1 being the date of availability (R. V. Melville,
pers. comm.). This is contrary to subsequent references to the species as 'Trochammina
rotaliformis J. Wright MS' by Heron-Allen & Earland (1913), Cushman (1920), and in the
Catalogue of Foraminifera (Ellis & Messina, 1940 et seqq.}.

Cushman (1920 : 77, pi. 16, figs 1, 2) re-illustrated Balkwill & Wright's four figures, and
at the same time made the first formal description of the species. . .

Test small, trochoid, spire somewhat elevated, composed of about three volutions,
gradually increasing in diameter; chambers distinct, four in each volution, sutures
oblique and curved, slightly depressed, ventral side irregular, the last formed chamber
occupying nearly one-half the area of the test, smoothly finished; aperture elongate, at
the base of the chamber in the umbilical region, with a sort of lip-like projection above;
color reddish or yellowish brown. Diameter, up to 0-45 mm.

He however omitted to select a type-specimen, which is unfortunate, for as will be seen three
of the four individuals illustrated by Balkwill & Wright (and Cushman), though of similar
size are morphologically distinct, belonging even to separate genera. As long as the species

Bull. Br. Mm. nat. Hist. (Zool.) 45(7): 347-358 Issued 24 November 1983

347

Figs 1-8 Deuterammina (Deuterammina) rotaliformis (Heron-Allen & Earland). Figs 1-3, Lectotype
(N.M.I, no. 1.1980). Umbilical, edge and oblique-umbilical views. Specimen figured by Balkwill &
Wright, 1885, pi. 13, fig. 12b. Figs 4, 5 Paralectotype (N.M.I, no. 2.1980). Edge and spiral views.
Figs 6-8, Paralectotype (N.M.I, no. 3.1980). Oblique-umbilical, oblique and spiral views.
All from Killiney Bay, E. Ireland, 8-14 fathoms (15-25 m). Ex Wright collection (13-1921), labelled
Trochammina inflata van, x220.

FORAMINIFERA 349

lacked a type-specimen, no clear morphological concept existed and hence subsequent
workers interpreted T. rotaliformis differently, for instance Hoglund (1947), Murray (1971),
and Todd& Low (1981).

In order to stabilize the concept of T. rotaliformis Heron- Allen & Earland, its morpho-
logical features need to be determined on the basis of a lectotype. Then, and only then can
this species be properly identified and used meaningfully in taxonomic and ecological work.

At first glance Cushman's description (1920: 77) of T. rotaliformis apparently refers to
the slightly smaller specimen shown in Balkwill & Wright's drawings (1885, pi. 13, fig. 1 Ib)
by an umbilical view, because he mentions four chambers per whorl and that the final
chamber occupies nearly half of the umbilical side. Other features of this description referring
to the spiral side, the aperture and the smoothly finished surface, on the other hand, seem
to be derived from the other three specimens shown in the original illustrations (pi. 13, figs
1 la; 12a, b) which clearly are morphologically different from the first mentioned individual.
Cushman apparently regarded, as did Balkwill & Wright, the four specimens as conspecific.
The supposition is confirmed by the grouping of Balkwill & Wright's four figures in
Cushman's pi. 16 — as fig. 1 (putting erroneously together as belonging to the same form,
their original figs 12a and b) and, as fig. 2 (doing the same for their figs 1 la and b). Cushman's
description is therefore a composite one.

Recently Balkwill & Wright's figured specimens have been found in the National Museum
of Ireland (N.M.I.) preserved in the Joseph Wright Collection (13-1921) and in the light of
the above confusion, it is opportune to select a lectotype and describe the species afresh.
Of their figured material, those specimens representing pi. 13, figs 1 Ib, 12a and 12b (1885)
have been photographed using a Scanning Electron Microscope and are re-illustrated by our
respective Figs 1-3, 25 ( = fig. 12b); Figs 13-16, 26 ( = fig. 12a) and Figs 17-20, 27 (-fig.
1 Ib). The specimen shown in their fig. 1 la has not been scanned; it has, we can confirm,
the same umbilical features as that shown in fig. 12b.

Our micrographs clearly demonstrate that the original illustrated specimens were not
conspecific, but in fact belong to three different species which are placed according to their
apertural characteristics into the genera Deuterammina (D enter ammina) (Balkwill &
Wright, 1885, pi. 13, figs 1 la; 12a, b) and Paratrochammina (Paratrochammina) (figs 1 Ib).

We designate Balkwill & Wright's specimen N.M.I. 1.1980, shown originally in umbilical
view (fig. 12b) as lectotype of D. (D.) rotaliformis (Heron- Allen and Earland). Their fig. 1 la
is also conspecific and as it is by far the commonest form in the material labelled 'T. inflata
var.' or 'T. rotaliformis MS' in the Wright Collection from E. Ireland, it would therefore
seem the logical choice. The remaining two specimens from the original figured material
are described as D. (D.) balkwilli ( = fig. 12b) and Paratrochammina (P.) wrighti ( = fig. 1 Ib),
while a further species has been isolated in Wright's faunal slide collection labelled as T.
rotaliformis. Previously not illustrated, it has distinctive apertural characters and is described
as Deuterammina (Centrodeuterammina) dublinensis subgen. et sp. nov.

Systematics

The specimens illustrated in this paper are deposited in the National Museum of Ireland,
Dublin, and registered as 1.1980-3.1980 and 107.1981-1 1 1.1981, inclusively.

Genus DEUTERAMMINA Bronnimann, 1976, emended Bronnimann & Whittaker, 1983

Subgenus DEUTERAMMINA Brdnnimann, 1976
TYPE-SPECIES. Trochammina glabra Heron-Allen & Earland, 1932.

Deuterammina (Deuterammina) rotaliformis (Heron- Allen & Earland)

(Figs 1-8, 25)

1885 Trochammina inflata (Montagu) var. Balkwill & Wright (pars): 33 1, pi. 13, figs 1 la, 12b only.
1889 Trochammina inflata var. Balkwill & Wright (sic); Halkyard: 63, pi. 1, fig. 10.

350 P. BRONNIMANN & J. E. WHITTAKER

1892 Trochammina injlata (Montagu) var. Balkwill & Wright; Chaster: 58.

191 1 Trochammina rotaliformis J. Wright MS: Heron-Allen & Earland: 309 (availability of name).
1920 Trochammina rotaliformis J. Wright; Cushman (pars): 77, pi. 16, fig. 1 (right). 2 (left) only. [After
Balkwill & Wright, 1885].

1973 Trochammina astrifica Rhumbler; Haynes (pars): 34, pi. 4, fig. 18 only (non Trochammina
squamata Jones & Parker astrifica Rhumbler, 1938).

1974 Trochammina rotaliformis Wright; Levy el al: pi. 1, figs 4, 5.

MATERIAL. 9 syntypes, 3 from Killiney Bay, 6 from off Drogheda, E. Ireland. Wright Collec-
tion (13-1921), National Museum of Ireland (N.M.I.), Dublin.

LECTOTYPE. N.M.I, no. 1.1980, from slide labelled Trochammina injlata var., Killiney Bay,
E Ireland, long. 6°8'W, lat. 53°15'N, depth 8-14 fathoms (15-25 m). Specimen figured by
Balkwill & Wright, 1885, pi. 13, fig. 12b. Refigured in Figs 1-3, 25.

DESCRIPTION (LECTOTYPE). Test free, a low dextral trochospire, subcircular in umbilical and
spiral views and slightly lobate. In edge view, broadly domed on the spiral side though some-
what flattened axially; shallow-concave umbilically; periphery sharply rounded, but not
carinate. Umbilical depression star-shaped, open and deep with five branches. On spiral side,
small subglobular proloculus followed by about 20 chambers arranged in at least three whorls
with five chambers in the final whorl. Radial and spiral sutures not well defined in the early
volutions and thus exact numbers of chambers difficult to determine. In final whorl, slightly
curved intercameral sutures are but little depressed spirally; rather more so on umbilical side.
Septum not infolded. Umbilically, shape of chamber broadly triangular; spirally, low and
elongate. Two sets of apertures consisting of an interiomarginal extraumbilical primary
opening of Trochammina-type and a secondary opening in a umbilical-sutural, posteriorly
directed position. Secondary openings lead directly into the umbilicus and probably remain
functional throughout ontogeny. Primary openings, on the other hand, not in direct connec-
tion with umbilical depression, only the last-formed connecting with the exterior. Final
primary opening devoid of any lip-like structure but secondary openings accompanied by
distinct extensions which normally are well separated from septal wall. Wall single-layered
and imperforate. Agglutinant of quartz flakes arranged in mosaic-like pattern. Colour reddish
brown.

DIMENSIONS (LECTOTYPE). Maximum spiral and umbilical diameter 260 urn, minimum diam-
eter 230 um; thickness (axial height) lOOum. Maximum diameter of star-shaped umbilical
depression 60 um, length of each branch of star c.20 um. Length and height of primary
aperture, 27 and 9 um respectively. Extension associated with secondary apertures (final
chamber) 30x15 um.

VARIATION (PARALECTOTYPES). 4 paralectotypes have been chosen from the remaining
syntypic series, two each from Killiney Bay, and from off Drogheda, (long. 6°15'W, lat.
53°45'N), N.M.I, no. 2.1980, 3.1980, and 107.1981, 108.1981, respectively. Their overall
morphology is similar to that of the lectotype, two are dextral, two, sinistrally coiled. All,
together with the remaining unfigured specimens in the Wright Collection, are characterized
by 5 chambers in the final whorl and by the same, 5-branched star shaped, open and deep
umbilical depression, into which the secondary openings of the chambers of the final whorl
lead under the hook-shaped lip-like extensions. Paralectotype2.1980 (Figs 4, 5) is made up
of about 20 chambers and a subglobular proloculus of about 10 um diameter. The number
of chambers of the spirally shown paralectotype (3.1980: Figs 6-8) cannot be determined,
the early volutions being masked by foreign matter.

DIMENSIONS (PARALECTOTYPES)

Diameter Thickness

N.M.I, no. (Fig.) Maximum Minimum (axial height)

2. 1980 (Figs 4, 5) 240 um 220 um lOOum

3. 1980 (Figs 6-8) 220 um 200 um

107.1981 (Figs 9-11) 250um 220um lOOum

108.1981 (Fig. 12) 230um 210um

Figs 9-12 Deuterammina (Deuterammina) rotaliformis (Heron-Allen & Earland). Figs 9-11,
Paralectotype (N.M.I, no. 107.1981). Umbilical, edge and oblique-umbilical views. Fig. 12,
Paralectotype (N.M.I, no. 108.1981). Umbilical view.

Both from off Drogheda, E. Ireland, 16 fathoms (30 m). Ex Wright collection (13-1921), faunal slide
no. 34. x220.
Figs 13-16 Deuterammina (Deuterammina) balkwilli sp. nov. Holotype (N.M.I, no. 109.1981).

Oblique-umbilical, spiral, edge and umbilical views. Specimen figured by Balkwill & Wright, 1885,

pi. 13, fig. 12a.

From Killiney Bay, E. Ireland, 8-14 fathoms (15-25 m). Ex Wright collection (13-1921), labelled

Trochammina inflata var., x220.

352 P. BRONNIMANN & J. E. WHITTAKER

REMARKS. D. (D.) rotaliformis (Heron- Allen & Earland) has been compared with the deep-
water deuteramminids found in the Discovery material from the S. Atlantic and Antarctic
waters — D. (D.) discorbis (Earland), D. (D.) glabra (Heron-Allen & Earland), D. (D.) grisea
(Earland) and other, so far undescribed species (Bronnimann & Whittaker, in preparation).
It is easily distinguishable by its overall morphology, the position and shape of the primary
aperture, plus the hook-shaped, lip-like extensions (see Fig. 25) under which the secondary
openings occur.

D. (D.) rotaliformis under the name 'Trochammina rotaliformis Wright' or 'Wright MS'
is one of the most frequently cited taxons in the Trochamminidae, notwithstanding the fact
that, lacking up till now a type specimen, no-one could have known its true morphology
(Heron-Allen & Earland, 1916 : 228; 1922 : 1 14; 1932 : 344; Earland, 1933 : 85; 1934 : 99;
Parr, 1950: 276; Collins, 1958 : 354; and others). Furthermore in almost all cases where
it has been illustrated (Heron-Allen & Earland, 1913 : 52, pi. 3, figs 11-13; Cushman &
McCulloch, 1939 : 107, pi. 87, figs 2a-c; Cushman & Todd, 1947 : 7, pi. 1, fig. 14; Cushman,
1948:42, pi. 4, fig. 16; Todd & Low, 1967: 418, pi. l,figs 18a,b; 1981 : 18, fig. 49; Murray,
1971 : 39, pi. 12, figs 1-5; Van Voorthuysen, 1973 : pi. 13 (pi. 14 on plate explanation),
figs 3a-c) it invariably does not correspond with the lectotype as selected in the present paper.
Only two records subsequent to the original citation appear definitely conspecific; one is by
Haynes (1973 : 34, pi. 4, fig. 18 (but not figs 19 and 20) who illustrated a specimen from
Cardigan Bay, W. Wales (as Trochammina astrifica Rhumbler), which is typical of D. (D.)
rotaliformis. The other is by Levy et al. who illustrated (1974 : 128, pi. 1, figs 4, 5) from
Dunkirk beach sand, N. France, an umbilical and spiral view, of the true rotaliformis. The
umbilical view of their specimen, in particular, shows beautifully the double aperture.

Deutemmmina (Deuterammina) balkwilli sp. nov.
(Figs 13-16,26)

1885 Trochammina inflata (Montagu) var. Balkwill & Wright (pars): 331, fig. 12a only.

1920 Trochammina rotaliformis J. Wright; Cushman (pars): 77, pi. 16, fig. 1 (left) only. [After Balkwill

& Wright, 1885].
71971 Trochammina rotaliformis Heron-Allen & Earland; Murray (pars); 39, pi. 12, fig. 5 only.

DIAGNOSIS. A 6-chambered, compressed Deuterammina. Star-shaped umbilical depression
small with 6 very short radial branches. Primary and secondary apertures poorly separated.

NAME. For F. P. Balkwill, co-author with J. Wright of the paper 'Recent Foraminifera off
the coast of Dublin and Irish Sea', 1885.

MATERIAL. 1 specimen, ex J. Wright Collection (13-1921), National Museum of Ireland
(N.M.I.), Dublin.

HOLOTYPE. N.M.I, no. 109.1981, from slide labelled Trochammina inflata var., Killiney,
E. Ireland, long. 6°8'W, lat. 53°15'N, depth 8-14 fathoms (15-25 m). Specimen figured by
Balkwill & Wright, 1885, pi. 13, fig. 12a. Refigured in Figs 13-16, 26.

DESCRIPTION (HOLOTYPE). Test free; a low sinistral trochospire consisting of about 16 axially
compressed chambers arranged in 3 whorls with 6 chambers in the final whorl. Suboval in
umbilical and spiral views and slightly lobate. In edge view, gently convex spirally and
shallow-concave umbilically; periphery rounded. Umbilically, shape of chambers roughly
triangular separated by almost straight sutures; spirally, chambers are elongate and relatively
low, separated by curved sutures. Radial and spiral sutures well defined. Umbilical
depression open and deep, characterized by star-shaped outline with 6 short radial branches,
the result of incomplete overlap of the apertural rim of the primary aperture on that of the
preceding chamber. Two sets of apertures consisting of primary interiomarginal opening in
umbilical-extraumbilical position and a minute posteriorly directed secondary opening in
sutural position, hence of Deuterammina-lype. Primary opening an elongate slit accompa-
nied by an upturned rim, the prolongation of the septal wall. It rests with its border on the

FORAMINIFERA 353

first and penultimate chamber of final whorl (Figs 13, 26). Secondary openings not well-
separated from large slit- like interiomarginal primary openings: however, recognizable as
minute sutural slits between ends of apertural rims of primary openings and those of pre-
ceding chambers; they occur below proximal sides of radial branches and are posteriorly
directed. All primary and secondary openings are in direct communication with the umbilical
cavity and hence the exterior. Wall agglutinated, smooth, and apparently single- layered and
imperforate. Surface appears finely granular on both sides, but granulation slightly coarser
umbilically.

DIMENSIONS (HOLOTYPE). Maximum spiral and umbilical diameter 260 um, minimum diam-
eter 220 um, thickness (axial height) 90 um. Maximum diameter of star-shaped umbilical
depression 40 um, length of each branch of star c.lOum. Length and height of primary
aperture, 60 and 22 um, respectively; width of apertural rim c.5 um.

REMARKS. D. (D.) balkwilli differs from D. (D.) rotaliformis (Heron-Allen & Earland) essen-
tially in the apertural features. In the latter, the primary and secondary openings are separ-
ated, the secondary sutural, posteriorly directed openings are much longer and better
developed than in D. (D.) balkwilli. As the primary and secondary openings in our new
species cannot be clearly separated, except by assuming that the extent of the apertural rim
marks the extent of the primary aperture, there is some evidence for a new subgeneric
deuteramminid taxon distinct from Deuterammina (Deuterammina) Bronnimann, 1976 and
D. (Lepidodeuterammina) Bronnimann & Whittaker, 1983. However, this would be prema-
ture with the material available. D. (D.) balkwilli has 6 chambers in the final whorl as against
5 in D. (D.) rotaliformis and a slightly different aspect in edge view. Murray's photograph
(1974, pi. 12, fig. 5) of a six-chambered specimen of what he calls Trochammina rotaliformis
Heron-Allen & Earland could be conspecific with ours. It is however only shown in edge
view, the spiral and umbilical views are of different specimens and a different species (see
p. 357 below).

Genus DEUTERAMMINA Bronnimann, 1976
Subgenus CENTRODEUTERAMMINA subgen nov.

TYPE SPECIES. Deuterammina (Centrodeuterammina) dublinensis Bronnimann & Whittaker,
1983.

DEFINITION. Test free, trochospiral, low, but not 'watchglass' shaped. Wall agglutinated,
single-layered, imperforate. Chambers axially compressed; without inner structures.
Aperture double: primary opening interiomarginal, extraumbilical; secondary or supplemen-
tary opening at umbilical tip of chamber; directed centrally into umbilical cavity.

NAME. Subgeneric name refers to centrally directed secondary opening.

REMARKS. D. (Centrodeuterammina) differs from D. (Deuterammina) and D. (Lepidodeuter-
ammina) (see Bronnimann & Whittaker, 1983), both of which possess secondary openings
posteriorly directed in an umbilical-sutural position. The secondary openings of D. (Centro-
deuterammina) are centrally directed and occur at the umbilical tip of the chambers.
D. (Lepidodeuterammina) differs further from both D. (Deuterammina) and D. (Centrodeu-
terammina) by the attached, 'watchglass' shaped test.

Deuterammina (Centrodeuterammina) dublinensis sp. nov.
(Figs 2 1-24, 28-30)

DIAGNOSIS. A 7-chambered deteramminid, compressed in edge view; spiral side piano-,
convex. Strongly lobate in umbilical/spiral views, chambers slightly inflated, spiral and radial
sutures well defined. Umbilical depression narrow and deep.

NAME. After the type locality, Killiney Bay, near Dublin, Eire.

Figs 17-20 Paratrochammina ( Paratrochammina) wrighti sp. nov. Holotype (N.M.I, no. 1 10.1981).

Spiral, edge, umbilical and oblique-umbilical views. Specimen figured by Balkwill & Wright, 1885,

pi. 13, fig. lib.

From off Drogheda, E. Ireland, 16 fathoms (30m). Ex Wright collection (13-1921), labelled

Trochammina inflata var., x 220.
Figs 21-24 Deuterammina (Centrodeuterammina) dublinensis subgen. et sp. nov. Holotype (N.M.I.

no. 111.1981). Oblique-umbilical, spiral, edge and umbilical views.

From Killiney Bay, E. Ireland, 8-14 fathoms (15-25 m). Ex Wright collection (13-1921), faunal slide

no. 27, x220.

FORAMINIFERA 355

MATERIAL. 1 specimen, ex Wright Collection (13-1921), National Museum of Ireland
(N.M.I.), Dublin.

HOLOTYPE. N.M.I, no. 111.1981, from faunal slide no. 27, Killiney Bay, E. Ireland, long.
6°08'W, lat. 53°15'N, depth 8-14 fathoms (15-25 m). Specimen illustrated in Figs 21-24,
28-30.

DESCRIPTION (HOLOTYPE). Test free, trochospiral; coil very low, sinistral, consisting of 20
chambers, including the proloculus, arranged in three whorls with 7 chambers in the last
whorl. Chambers of at least the last two whorls, axially compressed. Subcircular in umbilical
and spiral views and strongly lobate. In edge view, spiral side almost plano-convex, umbilical
side shallow-concave, periphery rounded. Radial and spiral sutures slightly curved, in early
whorls straight-oblique; umbilically, slightly sinuous. Umbilical depression narrow, deep,
showing at the sutures short sutural branches formed by the overlap of the rims of the
secondary umbilically-directed openings on those of the preceding chamber. Each chamber
with two apertures: a primary interiomarginal opening in an extraumbilical position of
Trochammina-type (that is with its borders resting completely on the wall of the first
chamber of the final whorl) and a secondary opening at umbilical tip of chamber, directed
into axial cavity. Primary apertures covered by succeeding chambers leaving only ultimate
one to communicate directly with exterior, whereas all secondary apertures open into
umbilical depression and remain functional. Primary interiomarginal opening an elongate
slit devoid of any border lip but edged with granules (Fig. 28). Secondary openings arranged
in a low trochospire within umbilical cavity (Fig. 30): in the early part of the coil, short
and semicircular; final ones, elongate with more or less flat walls. Borders of early openings
a series of discrete granules and flakes arranged more or less perpendicularly to the borders,
later ones have thin walls devoid of granules. Agglutinated wall apparently single-layered
and imperforate. Spiral side with granules, umbilical side with flakes and granules, the latter
surface of distinctly coarser texture.

DIMENSIONS (HOLOTYPE). Maximum diameter in umbilical and spiral view 240 urn, mini-
mum diameter 220 um, thickness (axial height) 80 um. Maximum diameter of umbilical
cavity 40 um. Length and height of primary aperture, 55 and lOum, respectively; ditto,
secondary apertures (early coil), 10x5 um.

REMARKS. This species was not figured by Balkwill & Wright, 1885, but was contained in
Wright's faunal slide collection and from his MS identification was clearly considered by
him to fall within the concept of T. rotaliformis.

Genus PARATROCHAMMINA Bronnimann, 1979 emended Bronnimann & Whittaker

EMENDED GENERIC DEFINITION. Test trochospiral, free. Wall agglutinated, single layered, im-
perforate. Chambers devoid of inner structures (septal infoldings). Aperture single, interio-
marginal, umbilical-extraumbilical; border of opening resting on umbilical wall of first
chamber of last whorl and on that of penultimate chamber; asymmetric in respect to axis
of coiling, with or without an umbilical flap.

TYPE SPECIES. Paratrochammina madierae Bronnimann, 1979. Recent. South Atlantic
(Brazilian Shelf).

Subgenera of Paratrochammina

In the present classification of the Trochamminidae only the position of the single interio-
marginal aperture is accepted as a generic criterion. Because the presence of an umbilical
flap can be variably developed this character is here reduced to second order of importance
and, unlike formerly (Bronnimann & Whittaker, 1981), now accepted only as of subgeneric
significance. The genus is divided into P. (Paratrochammina) and P. (Portatrochammina).

356 P. BRONNIMANN & J. E. WHITTAKER

Subgenus PARATROCHAMMINA (PARATROCHAMMINA) Bronnimann, 1979

SUBGENERIC DEFINITION. Aperture single, interiomarginal, umbilical-extraumbilical, border
of opening rests on umbilical wall of first chamber of last whorl and on that of penultimate
chamber; asymmetric in respect to axis of coiling; without umbilical flap.

TYPE-SPECIES. As genus.

Subgenus PARATROCHAMMINA (PORTATROCHAMMINA) Echols, 1971

SUBGENERIC DEFINITION. Aperture single, interiomarginal, umbilical-extraumbilical, border
of opening rests on umbilical wall of first chamber of last whorl and on that of penultimate
chamber; asymmetric in respect to axis of coiling, with umbilical flap more or less covering
umbilical depression.

Type-SPECiES. Portatrochammina eltaninae Echols, 1971 ( = Paratrochammina (Portatro-
chammina) antarctica antarctica (Parr, 1950). Recent. Soctia Sea, Antarctica.

Paratrochammina (Paratrochammina) wrighti sp. nov.
(Figs 17-20,27)

1885 Trochammina inflata Montagu var. Balkwill & Wright (pars): 331, pi. 13, fig. 1 Ib only.
71913 Trochammina rotaliformis J. Wright MS; Heron-Allen & Earland: 52, pi. 3, figs 1 1-13.
1920 Trochammina rotaliformis J. Wright; Cushman (pars): 77, pi. 16, fig. 2 (right) only. [After

Balkwill & Wright, 1885].
71971 Trochammina rotaliformis Heron-Allen & Earland; Murray (pars): 39, pi. 12, figs 1-4 only.

DIAGNOSIS. A 4-chambered paratrochamminid. In edge view, convex spirally, shallow-
concave umbilically. Chambers 'broad bean' shaped, spiral and radial sutures well defined.
Central umbilical depression shallow, aperture with small lip-like plate.

NAME. After Joseph Wright, co-author with F. P. Balkwill of the first and only comprehen-
sive and illustrated account of the foraminifera of the western Irish Sea.

MATERIAL. 1 specimen, ex Wright Collection (13-1921), National Museum of Ireland
(N.M.I.), Dublin.

HOLOTYPE. N.M.I, no. 110.1981, from slide labelled Trochammina inflata var., off
Drogheda, E. Ireland, long. 6°15'W, lat. 53°45'N, depth 16 fathoms (30m). Specimen
figured by Balkwill & Wright, 1885, pi. 13, fig. 1 Ib. Refigured in figs 17-20, 27.

DESCRIPTION (HOLOTYPE). Test free, trochospiral; coil moderately low, dextral, consisting of
13 chambers arranged in three whorls, with four chambers in final whorl and at least in last
two whorls axially compressed. Broadly oval in umbilical and spiral views. In edge view,
convex spirally, obtusely pointed axially; shallow-concave umbilically; periphery rounded.
Chambers 'broad bean' shape umbilically, elongate crescentic spirally. Radial and spiral
sutures well-defined. Intercameral sutures strongly curved spirally and somewhat sinuous
umbilically. Umbilical depression, shallow, open. Single interiomarginal aperture of crescen-
tic shape with central lip-like plate. Border of aperture rests on wall of first and third chamber
of final whorl (of P. (Paratrochammina) type). Agglutinated wall appears single-layered and
imperforate. Surface, fairly rough, composed of flakes and grains of rock fragments; texture
of spiral surface coarser than umbilical.

DIMENSIONS (HOLOTYPE). Maximum spiral and umbilical diameter 250 urn, minimum
diameter 220 um, thickness (axial height) 120um. Maximum diameter of umbilical
depression c.50um. Length of aperture 90 urn, length and width of apertural plate 50 and
9 um, respectively.

REMARKS. The single Paratrochammina-lype aperture readily distinguishes this species from
the deuteramminids described above, all with double apertures. The central lip-like apertural

FORAMINIFERA

357

Figs 25-30 Apertural detail. Fig. 25, Deuterammina (Deuterammina) rotaliformis (Heron-Allen &
Earland). Lectotype (N.M.I, no. 1.1980). Oblique view showing primary and secondary apertures,
X530. Fig. 26, Deuterammina (Deuterammina) balkwilli sp. nov. Holotype (N.M.I, no. 109.1981).
Oblique view showing primary and secondary apertures, x 360. Fig. 27, Paratrochammina (Paratro-
chammina) wrighti sp. nov. Holotype (N.M.I, no. 1 10.1981). Oblique view of umbilicus and aper-
ture, x350. Figs 28-30, Deuterammina (Centrodeuterammina) dublinensis sp. nov. Holotype
(N.M.I, no. 1 1 1.1981). Fig. 28, edge view of primary aperture, x950. Fig. 29, oblique view of pri-
mary and secondary apertures, x375. Fig. 30, oblique close-up view of secondary apertures within
open umbilicus, x 1000.

plate does not cover the umbilical depression as it does in Paratrochammina (Portatrocham-
mina). Murray's 1971 illustrations (pi. 17, figs 1-4 only) of T. rotaliformis could belong to
this form.

Acknowledgements

We are extremely grateful to Dr Colin Harris (Trinity College, Dublin) who first brought
the whereabouts of the Joseph Wright Collection to our attention. The Director of the
National Museum of Ireland kindly allowed us to borrow the material and gave permission
for Scanning Electron Microscopy. Mr Mark Holmes, Natural History Division, N.M.I.,
facilitated the loan. Mr R. V. Melville, International Commission of Zoological Nomen-
clature, once again advised us on taxonomic procedure. Drs C. G. Adams and L. R. M.
Cocks, British Museum (Natural History) commented helpfully on the text. The work of
P. Bronnimann is in part funded by the Fonds National Suisse.

References

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Manuscript accepted for publication 7 March 1983

Notes on Atlantic and other Asteroidea. 3. The
families Ganeriidae and Asterinidae, with
description of a new asterinid genus

Ailsa M. Clark

Department of Zoology, British Museum (Nat. Hist.), Cromwell Road, London SW7 5BD

Introduction

Examination of new and old material of these two closely related starfish families has
prompted some additional observations on their limits and relationships, with consequent
modifications to the diagnoses. A notable addition in the Ganeriidae is the abyssal genus
Vemaster Bernasconi, 1965, formerly assigned to the Goniasteridae, but now shown by dis-
section of new specimens to be more closely related to Ganeria, a move supported by
Downey and Blake (pers. comms). Several of the nominal genera proposed by Verrill (1913)
for the Asterinidae are reevaluated, resulting in the synonymy of Patiria with Asterina and,
conversely, in the revival ofCallopatiria. Some recently collected asterinids from the Atlantic
coast of Panama proved to represent a new genus remarkable for having many of the
abactinal plates paxilliform.

A difficulty arose in formally citing the type species of Asterina itself, for which a possible
solution to the nomenclatural problem is given in an appendix.

Systematic account
Families GANERIIDAE Sladen and ASTERINIDAE Gray

Blake (1981) has reviewed the history of these two families which he transferred to the order
Valvatida from the Spinulosida. Briefly, Gray (1847) originally included Ganeria itself in
his Asterinidae, being followed in this by Perrier (1875) and Viguier (1878). However, Sladen
(1889) distinguished the subfamily Ganeriinae within the Asterinidae, on the basis of
enlarged marginal plates, for Ganeria and Cycethra Bell, 1881 and this was raised to family
rank by Perrier ( 1 894). Later authors, notably Fisher ( 1 9 1 1 a) and Blake (1981), have empha-
sized the close affinity between the Ganeriidae and the Asterinidae. Fisher followed Sladen
in distinguishing them on the basis of large size of the marginal plates in ganeriids but this
does not take account of those individuals of Cycethra and all the species of the later- added
Perknaster in which the marginals are reduced and the series may be barely, if at all, dis-
tinguishable over part of their extent. Bernasconi (1964) and Blake (1981) have separated
them on the form of the abactinal plates, with well denned columns of various height in
the Ganeriidae but with ridge-like elevations or gently rounded convexities in the Asterinidae
(though Blake termed both plate forms 'metapaxilliform' since the basal part of each plate
is variously enlarged and the elevation often excentric) but this again does not take into
account the amazing variations of Cycethra, some individuals of which have simply convex
or crescentic elevations indistinguishable from those of certain Asterinidae instead of the
usual well-defined but low columns ranging from circular through oval to crescentic in sec-
tion. Blake also utilized the arrangement of these plates, said to be in ill-defined rows in
Ganeriidae but well-defined ones in Asterinidae; however, some asterinids, especially those
of large size comparable to that of many specimens of Ganeria and Cycethra, have an essen-
tially alternating abactinal skeleton, though alternate plates may have a secondarily
longitudinal alignment, for instance in Patiria granifera (Fig. 6, below) and Nepanthia
crassa.

In an attempt to assess the relationships of the various plate systems of Atlantic asteroids,

Bull. Br. Mm. nat. Hist. (Zool.) 45(7): 359-380 Issued 24 November 1983

359

360

AILSA M. CLARK

Fig. 1 Partial cross sections near base of ray viewed from proximal side of : (a) Leilaster radians
(Perrier), Gerda st. 1270. Yucatan Channel, W of Cuba, 318m, R 11-5-14-5 mm (with isolated
inferomarginal plate viewed from above); (b) Lophaster furcifer (Duben & Koren), BM reg. no.
91.5.20.1, Trondheim fjord, R 30mm; (c) Cycethra verrucosa (Philippi), 1971.9.1.4, N Argentina,
R 40 mm; (d) C. verrucosa, formerly named Patina obesa, 1948.3.16.356, mid-Argentina, R 33 mm;
(e) Asterina gibbosa (Pennant), 98.5.3. 1 7 1 , Falmouth, Cornwall, R 1 5-5 mm; (0 Anseropoda placenta
(Pennant), 88.10.8.1, W of Scotland, R 31^0 mm (oblique section following the lines of abactinal
and actinal plates). Plates not in the plane of the section shown by discontinuous lines except in
(0- The scale measures 2 mm for (a), 2-5 mm for (e) and (0, 3 mm for (bHd).

GANERIIDAE & ASTERINIDAE 361

I have made cross- sections of the rays of some taxa. Where the Ganeriidae and Asterinidae
are concerned, these do show considerable resemblances between some species of both fami-
lies but I find there is always a more or less well-defined ventro- lateral angle at the level
of the inferomarginal plates in the Asterinidae, whereas in ganeriids the contour of the ray
in this area is curved, the actinal surface being slightly convex and merging into the side
of the ray. Inevitably, in asterinids such as Nepanthia, where the well-defined arms are nearly
cylindrical, the angle is least, but it is still distinct. Coupled with the superficial shape, the
marginals of the Ganeriidae, when enlarged, are block-like, these of the two series similar
in form and not wedged in by internal extensions of the lateralmost abactinal and actinal
plates. In the Asterinidae the marginals of the two series are consistently small and often
only recognizable by their regularity along the angle (the superomarginals sometimes barely
distinct), the flattened inferomarginals appearing pearshaped or pegshaped in the sectional
view and wedged in by the inner ends of the adjacent plates or some supernumerary internal
plates which form a series of buttresses or a more continuous structure reinforcing the ventro-
lateral angle (see Fig. le, 5b, c). In the extremely flattened asterinid Anseropoda the ventro-
lateral angle is very narrow and the reinforcement for the upper side spans almost the entire
body cavity of the ray, except for a narrow midradial area, taking the form of angled struts
in obliquely transverse series, each made up by a slender flattened process from an abactinal
plate overlapping the upper end of a similar process from the corresponding actinal plate
(Fig. If).

One consequence of this study is realization that Leilaster A. H. Clark, 1938 is not a
ganeriid. When established by Clark for type species Korethraster radians Perrier, 1881, from
the Caribbean, Fisher evidently supported a proposal to include it in the Ganeriidae. He
reiterated this in 1940 (pp. 152-153) when discussing the antarctic Mimstrella, which he
referred to the Asterinidae though commenting that the 'produced inferomarginals' also sug-
gested affinity with the Ganeriidae. Presumably he could then only have had Leilaster in
mind, not the true ganeriids. A cross-section of a ray of the small Leilaster radians
(Fig. la), where R is up to only c. 1 5 mm, shows a flat underside formed mainly by the rela-
tively large cigarshaped inferomarginal plates which abut directly on to the outer ends of
the ambulacral plates (the single actinal plate in a proximal section being superimposed).
As viewed from above, the inferomarginals are slippershaped, tapering outwardly, and have
a notch on the inner (adradial) half forming a socket into which fits the lower end of the
vertically aligned corresponding superomarginal plate. The coarse abactinal plates imbricate
adradially (the more lateral ones) or proximally (the midradial ones) and each is raised into
a ridge or rounded column and crowned with coarse capitate spinelets. The formation of
a lateral flange to the body made by the inferomarginal plates and the imbricating abactinals
are characters shared with most asterinids but in Leilaster the inferomarginals are relatively
much larger and nothing like their articulation with the superomarginals is shown in the
asterinids studied, nor do they directly contact the ambulacral plates in asterinids without
multiple series of intervening actinal plates — even at this relatively small size. Further, in
Leilaster the proximal articulating face of the adambulacrals (which provides characters
given considerable weight by Blake) is rectangular and has a complex near- horizontal groove
unlike any of the squarish, slightly hollowed adambulacrals of the asterinids and other
Valvatida studied.

These internal structures have yet to be studied in Mirastrella biradialis Fisher when more
material is available. Consultation with Miss M. E. Downey of the Smithsonian Institution,
co-author of the review of Atlantic Asteroidea for which the present note is a preliminary,
yielded her opinion that Leilaster and Mirastrella are closely related. They share two unusual
characters not shown by any ganeriids or asterinids. For one thing, there is a distinct groove
(or fasciole) in each interradius running vertically from the margin up to the flat top of the.
disc between the two first superomarginals and the interradial abactinal plates above.
Secondly, there are two longitudinal (but bowed) series of enlarged abactinal plates (the 'first
adradial' series of Fisher) on each ray framing the three (Leilaster) or two (Mirastrella) longi-
tudinal series of midradial plates like the two prongs of a wishbone.

362 AILSA M. CLARK

The complex articulating face of the adambulacrals in Leilaster shows some resemblance
to that of the spinulosan Lophaster furcifer (Diiben & Koren) of the family Solasteridae
(Fig. Ib). Lophaster also has reduced actinal areas, though the section made shows one
actinal interposed between the inferomarginal and the ambulacral/adambulacral articula-
tion, at least in the proximal part of a ray at R c.30 mm. The superomarginal in Lophaster
has a tall paxillar column and a triradiate base, two lobes of which articulate with the upper
side of the expanded adradial part of the inferomarginal straddling a median hollow which
does not form a socket. The marginals as well as the abactinal plates in Lophaster and other
solasterids are highly paxilliform with slender rounded columns, rather than being meta-
paxilliform as in Leilaster. Even so, I think Verrill (1915) had some justification for
treating a specimen with R 35 mm, three times that of the type of L. radians, as conspecific;
his specimen was renamed Lophaster verrilli by A. H. Clark in 1938. Whether or not the
small specimens which have been referred to Leilaster radians and Mirastrella biradialis
should be referred to the Spinulosida remains to be determined from a review of that order.

Another genus of uncertain affinities is the diminutive abyssal Vemaster Bernasconi, 1965,
from the Argentine Basin (maximum R only 1 5 mm). Because of some resemblance in the
tabulate abactinal plates to Cladaster and Nectria Bernasconi referred Vemaster to the
Goniasteridae. However, examination of some additional material of Vemaster shows that
the superomarginal plates are undeveloped and the inferomarginals displaced to a ventro-
lateral position and not at all blocklike (both marginal series being essentially well-developed
and blocklike in Goniasteridae), there is no regularity in the arrangement of the abactinal
plates and the actinal plates are very reduced and irregular, being possibly secondary calcifi-
cations. Also, internally Vemaster has an archlike calcified interbrachial column arising from
near the oral plates in each interradius, curving laterally at its upper end to meet the lateral
wall of the disc, this compares with a solid calcified interbrachial buttress fused to the side
of the disc in Nectria, which genus also has superambulacral plates developed (according
to Fisher, 191 la), not observed in Vemaster, besides having regularly arranged abactinals.
However, just such an inclined columnar interbrachial support is found in Ganeriidae such
as Cycethra and Perknaster (indeed in the specimen of P. sladeni studied, this structure is
arched over as in Vemaster), which genera also show a marked tendency for reduction of
the marginals and loss of their blocklike form, especially Perknaster, where the abactinal
plates are irregular and the actinal areas somewhat reduced.

Apart from the much finer abactinal skeleton of Perknaster, the main difference between
it, together with the other southern genera of Ganeriidae, and Vemaster is the absence of
a ventrolateral angle — emphasized above as a characteristic feature of the family in com-
parison with the Asterinidae — whereas the inferomarginals of Vemaster form a distinct
projection in this area, the arms being D-shaped in cross-section with the ventral side quite
flat. The asterinid Nepanthia does have an almost D-shaped arm section but here actinal
plates are developed in regular series on the arms and superomarginals combine with the
inferomarginals to make the ventrolateral angle.

Both Downey and Blake (pers. comms) support reference of Vemaster to the Ganeriidae.
However, in view of the marked difference in arm section, I think that its isolation in a
separate subfamily from the closely related Ganeria, Cycethra and Perknaster may prove
to be desirable. This problem must await comparison of all these with other genera from
parts of the world outside the Atlantic which have been referred to the Ganeriidae but which
seem to me less closely interrelated than are the three Southern Ocean genera. These other
genera are: Aleutiaster A. H. Clark, 1939, from the Aleutian Is, North Pacific, Hyalinothrix
Fisher, 191 Ib, from the Hawaiian Is, Knightaster H. E. S. Clark, 1972, from New Zealand
and Tarachaster Fisher, 1913, from the Philippines with a second species from New Zealand
of McKnight, 1973, the rest being monotypic.

Another taxon which the study of arm sections shows to have been misplaced is Patiria
(originally Parasterina} obesa (H. L. Clark, 1910), recorded from both sides of southern
South America. In 1962 I renamed as P. obesa four Discovery specimens from southern
Argentina (which Fisher (1940) had identified as Cycethra verrucosa forma nitida) since they

GANERIIDAE & ASTERINIDAE

363

••••••••••••••I

••••••••••••••••••••••••••CM'

Fig. 2 Cycethra verrucosa (Philippi), BM reg. no. 1948.3.16.356, Argentina, R 33mm (formerly
named Patina obesa), x2 (above) and 1948.3.16.344, W Falkland I, R 39 mm, with more abrupt
elevations on the abactinal plates but reduced marginals, x 1£ (below).

364 AILSA M. CLARK

closely resemble the photographs of H. L. Clark and of Madsen (1956) of P. obesa from
Chile and have notably asterinid-like abactinal plates with only low crescentic convexities,
though some of them have the distal superomarginals enlarged in an 'un-asterinid-like' way
(fig. 2 top). Madsen also recorded an Argentinian specimen as obesa and noted that the
development of the marginals varies from barely distinct (as in the holotype) to fairly
conspicuous, as in the Discovery specimen sectioned (Fig. Id) and even more the one photo-
graphed. These marginals are block-like and those of the two series similar in shape, forming
a curved surface with no sign of an angle at the inferomarginal. Bernasconi too (1973) found
the marginal area of obesa to be thick and rounded. Such a form matches that of some speci-
mens of Cycethra verrucosa, in which species the marginals show a similar wide range of
development, whereas in asterinids the marginals variously developed have been found to
have the abactinal plates variously developed with elevations ranging from ill-defined
columns to crescentic rounded convexities. I must therefore concede the correctness of
Fisher's identification of the Discovery specimens, from which it follows that Patina obesa
is yet another synonym of Cycethra verrucosa (Philippi, 1857), a taxon so variable that I
think it futile to try and treat Cycethra as other than a monotypic genus.

Madsen and Bernasconi both compared P. obesa with P. bellula Sladen of South Africa,
the type species of Callopatiria Verrill, 1913, synonymized with Patina by Fisher (1940);
Madsen suggested 'perhaps this genus [Callopatiria] ought to be retained and also include
the species obesa\ an idea supported by Tortonese (pers. comm.). I too agree that
Callopatiria deserves recognition, as discussed below, and think it comes closest of all the
asterinids to obesa, though I am convinced that the true place of obesa is among the
Ganeriidae.

In his drastic summary revision of the Asterininae (1913), Verrill recognized no less than
12 nominal genera in addition to Stegnaster and Tremaster, since isolated in a separate
subfamily, four of them unconnected with the Atlantic fauna, leaving the following for
consideration:

Asterina Nardo, 1834. Type species (see appendix, p. 378): 'Asterias minuta = A. gibbosa

(Pennant, 1777)' by subsequent designation by Fisher, 1906.
Asterinides Verrill, 1913. Type species: Asteriscus folium Lutken, 1860.
Patiria Gray, 1840. Type species: P. coccinea Gray, 1840, a synonym of Asterias miniata

Brandt, 1835.

Patiriella Verrill, 1913. Type species: Asterina regularis Verrill, 1867.
Asterinopsis Verrill, 1913. Type species: Asterias penicillatus Lamarck, 1816.
Enoplopatiria Verrill, 1913. Type species: Asteriscus marginatus Hupe, 1857, a nom. nud.

conspecific with A. stellifer Mobius, 1859.
Callopatiria Verrill, 1913. Type species: Patiria bellula Sladen, 1889, a synonym of P. grani-

/mzGray, 1847.
Allopatiria Verrill, 1913. Type species: P. ocellifera Gray, 1847.

Of these, Asterinides has been synonymized with Asterina by Fisher (1919), in his
Philippine book, on the grounds that the absence of pedicellariae (the only distinction given
by Verrill) is not a character of generic weight. However, Brito (1968) apparently overlooked
this and Tommasi (1970) disregarded it, both of them retaining Asterinides for folium with-
out further justification. Although occurrence of pedicellariae is unreliable even as a specific
character, the name Asterinides might be utilized, at least at the subgeneric level, for a group
of species with the body relatively low and near pentagonal (though this alone is not of
supraspecific weight) correlated with very fine, easily detached abactinal spinelets. Besides
A. folium it could also include the Indo- Pacific sarasini group of species discussed in Clark
&Rowe(1971 : 67-68).

Verrill (1913; 1914a & b) differentiated his nominal genera only in two versions of a
dichotomous key, the 1913 one confused by the omission of one option: DD. 'Interactinal
plates without a fan of spines', which I think should have come on p. 480 after Enoplopatiria
and been immediately followed by b and bb (for Patiriella and Asterinopsis) though Verrill

GANERIIDAE & ASTERINIDAE

365

(\9\4a) proposed its insertion on p. 479 after EE and before x for Patina which does have
fans of spines on the actinal plates. It appears that Enoplopatiria, resulting from xx, must
have been an afterthought, since the sequence in the key is otherwise alphabetical from A
to G. The 1 9 1 4b version shows Verrill's true intention and this was clarified by Fisher (1919),
who adopted Patiriella and produced a simplified key to the Pacific genera of Asterininae,
differentiating Patiriella and Patiria from Asterina by the abactinal structure, simpler in
Asterina with the plates of the papular areas of only one kind and rounded, elliptical, cordate
or shieldshaped (in Verrill's words), as opposed to plates of two or more diverse kinds, the
larger arched or crescentic and notched for the papulae and the smaller ones ovate or
pear-shaped and inserted endwise (vertical to the surface) between the larger.

Fig. 3 Denuded abactinal plates from proximal part of midradial area of: (a) Asterina gibbosa
(Pennant), BM reg. no. 1960.8.4.4, Syra, Aegean Sea, R 27mm; (b) Asterina (formerly Patiria}
miniata (Brandt), 1954.9.8.43, San Pedro, S California, R 16mm; (c) Patiriella regularis (Verrill),
1970.10.8.111, Nelson, New Zealand, R 21 mm; (d) Asterina folium (Liitken), USNM no. 38811,
Sand Key Reef, S Florida, R 8 mm (a few surviving spinelets shown at the proximal ends of two
plates); (e) Callopatiria (formerly Patiria) granifera (Gray), 1903.8.1.47, Mossel Bay, S Africa, R
40 mm; (f) Asterina (formerly Patiria) stellifera (Mobius), 1936.10.14.3, Accra, Ghana, R 41 mm;
(g) Allopatiria ocellifera (Gray), 1969.12.16.13, Cape Lever, Mauritania, R 38mm. The scale
measures 4 mm for (g) and 2 mm for the rest; (a) and(f) are from wet specimens, the pores appearing
relatively small.

However, examination of a relatively large specimen of Asterina gibbosa (R 27 mm) (Figs
3a, 4 top) shows closer agreement of the abactinal skeleton with the second option than the
first, there being two or even three small secondary plates in some of the papular areas and
the primaries in the papular areas are mostly crescentic. The type species of both Patiria
and Patiriella reach a considerably larger size, R>60mm, and develop many more sec-
ondary plates though at R<30 mm these are relatively few in number (Fig. 3b, c). The form
and arrangement of the other plates, including the internal ones buttressing the lateral angles
of the arms (see Fig. le), and the armament of A. gibbosa and the other two type species,
do not seem to me significantly different, while the difference in maximum size alone seems
inadequate for the distinction of genera. Verrill's keys tried to distinguish Patiriella from
Asterina also by the reduced actinal armament of usually only a single spine, rarely more

366

AILSA M. CLARK

Fig. 4 Dorsal views, x 1£, and enlargements of denuded rays, x3, of: Asterina gibbosa (Pennant),
BM reg. no. 1960.8.4.4, Syra, Aegean Sea, R 27 mm (above); Asterina (formerly Patirid) miniata
(Brandt), 1954.9.8.43, San Pedro, California, R c.32 mm (middle); Patiriella regularis (Verrill),
1970. 10.8. 11 2, Golden Bay, New Zealand, R 30 mm (below).

GANERIIDAE & ASTERINIDAE 367

than two on a plate. Dartnall (1971) in diagnosing Patiriella also noted that the abactinal
armament is short, granular and bluntly capitate, the carinal (midradial) plates are doubly
notched to accommodate two longitudinal rows of papulae and pedicellariae are lacking.
The other characters he mentioned are common to Asterina gibbosa and indeed some of
the midradial plates in that species also subtend two pores though the three proximal lobes
are less marked than in Patiriella regularis. Small specimens of A. gibbosa and many A. bur-
toni Gray, for instance, also have no more than two actinal spines. However, the particularly
coarse and abbreviated abactinal armament in the australasian species P. regularis and P.
gunni (Gray), as well as the pan-oceanic P. exigua (Lamarck), and perhaps the frequent
development in these of a regular pair of secondary plates proximal to some of the trilobed
midradial primaries (not observed in A. gibbosa}, may serve to justify retention of Patiriella
with generic (or perhaps subgeneric) rank. Dartnall (pers. comm.) shares my doubts about
the generic distinction of Patiriella. Since Australia is rich in species of both Patiriella and
Asterina, the resolving of this problem seems best left to one of the several interested
Australian specialists, the name Patiriella being retained here for P. exigua which extends
to St Helena.

Patiria miniata from the NE Pacific was only shown to be the type species of Patina in
1933 when Mortensen synonymized with it P. coccinea Gray, mistakenly recorded from
South Africa. Fisher (19 11 a), in his N Pacific asteroid monograph, had included miniata
in Asterina and this combination has been perpetuated in the Pacific literature for both
miniata and the closely related Japanese pectinifera (Miiller & Troschel). These two differ
from Asterina gibbosa (apart from the greater maximum size) in lacking pedicellariae but
this is not true of P. stellifera (Mobius), the type species of Enoplopatiria, from E South
America and W Africa, which is otherwise very similar to P. miniata in shape, plate structure
and armament. Enoplopatiria was synonymized with Patiria by Fisher (1919), discounting
the taxonomic weight of the occurrence of pedicellariae, as with Asterinides, but Enoplopa-
tiria was similarly retained by some South American authors, in this case Bernasconi (1955),
as well as Brito (1968) and Tommasi (1970). However, Madsen (1950) had used Asterina
for stellifera and this was followed by me in 1955. Subsequently I expressed doubts (quoted
by Tommasi) about this position for stellifera because of the many secondary abactinal plates
and, in Clark & Courtman-Stock (1976), like Tortonese (1956) and Bernasconi (1966), used
Patiria stellifera. I am now forced to the reverse, rejecting greater maximum size and its
commensurate elaboration of the abactinal skeleton as justifying a generic distinction; both
Enoplopatiria and Patiria itself are therefore to be synonymized with Asterina.

Of the remaining species currently included in Patiria, P. obesa (H. L. Clark, 1910) is
referred above to the synonymy of Cycethra verrucosa, P. granulosa (Perrier, 1875),
supposedly from the Hawaiian Is, was based on specimens which appear to be conspecific
with P. miniata (indeed, one sample is so referred in the catalogue of the Paris Museum
collections) and P. chilensis (Liitken, 1859) is also close to miniata and so falls within
Asterina. This leaves only P. granifera Gray, 1847 and P. formosa (Mortensen, 1933), both
from South Africa, P. obtusa Gray, 1847, supposedly from Pacific Panama and P. pulla and
P. rosea both of Koehler & Vaney, 1906, from Mauritania, these last two dealt with below
under Allopatiria.

Patiria granifera is type species (by synonymy with C. belluld) of Callopatiria, which was
formally synonymized with Patiria by Fisher in 1941, since Mortensen (1933) cited
C. bellula in the references to [Parasterina] bellula and Fisher himself (1940) returned bellula
to Patiria. In 1956 I synonymized bellula with granifera. Comparison of arm sections of this
species with Patiria (or Asterina} miniata now shows a significant difference in the alignment
of the superomarginal plates, those of P. granifera and the closely related P. formosa having
both superomarginal and inferomarginal plates projecting to a similar degree together
forming the ventrolateral angle (Fig. 5b) — whereas in Asterina sensu lato, Patiriella and
Allopatiria, the angle is sharper and formed only by the inferomarginal series projecting
beyond the inset superomarginals which may be hardly distinguished from the adjacent
abactinal plates and can alternate in position with the inferomarginals along the arm. Of

368

AILSA M. CLARK

Fig. 5 Partial cross sections near base of ray viewed from proximal side, with lateral views of marginal
area immediately distal to section, of: (a) 'Patiria ' obtusa Gray, holotype, BM reg. no. 1938.6.23.24,
Panama, R up to 54 mm (the arrow shows the plane of the section where the internal part of the
inferomarginal is exposed); (b) Callopatiria (formerly Patiria) granifera (Gray), 81.6.22.14, Cape
Town, R 29 mm; (c) Asterina (formerly Patina) stellifera (Mobius), USNM, Angola, R c.35 mm; (d)
Nepanthia crassa (Gray), 61.7.8.32, Fremantle, Western Australia, R 40mm; (e) N. brevis
(Perrier), 81.10.26.172, Albany I, N Australia, R 28mm. The parallel rows of dots on the
ambulacral plates show the hyaline areas over which the tube foot slides; hatching shows cut surfaces
of plates and discontinuous lines indicate plates not in the plane of the section: in (d) the outlines
of the plates buried in the body wall are hypothetical. The scale measures 3 mm.

GANERIIDAE & ASTERINIDAE

369

Fig. 6 Nepanthia crassa (Gray), BM reg. no. 61.7.8.32, Fremantle, Western Australia, R c. 38 mm
(above left); Callopatiria (formerly Patina) formosa (Mortensen), 1975.10.29.62, False Bay, South
Africa, R 36 mm (above right); Callopatiria (formerly Patina) granifera (Gray), 1948.3.16.798,
Saldanha Beach, South Africa, R 45 mm (below). All x 1^.

370 AILSA M. CLARK

the Asterininae studied, only in the Indo-Pacific Nepanthia does the marginal structure
approximate to that in P. granifera and formosa, the sides of the ray being approximately
vertical and the angle rounded, especially in species such as N. belcheri (Perrier) with thicker
body walls and more numerous secondary abactinal plates.

Several other differences mark off P. granifera and formosa, together with Nepanthia, from
the rest of the Asterininae under discussion. Firstly, the primary abactinal plates along the
midradial line in the former are all crescentic or rounded and have an alternating or irregular
arrangement whereas in larger specimens of Asterina gibbosa and other Asterininae some
of these plates are proximally trilobed and tend to form at least a partial single longitudinal
row, though this may be double for some of its length (Figs 3a-c, 0- Secondly, some of the
secondary abactinal plates are elongated in the superficial plane instead of vertical to it in
the South African species and the Nepanthia species where such plates are numerous. Lastly,
and perhaps most importantly, the transverse sections show that there are internal ventral
plates bracing the ambulacrals by linking them with the actinals and other additional plates
integrated with the ventrolateral angle, the adradialmost of which are equivalent to super-
ambulacral plates. In Nepanthia brevis (at least in the holotype) the superambulacrals are
fairly regular in size and position and, in the proximal part of the arm at least, link through
two other internal plates overlying the actinals to the lateral bodywall (Fig. 5e). However,
in the specimen of Patiria granifera sectioned, these plates are less well developed and not
regularly aligned, being also partly immersed in the fleshy internal part of the ventral body-
wall (Fig. 5b). In Nepanthia crassa (formerly Parasterina but provisionally referred to
Nepanthia by Fisher (1941) and confirmed there by Rowe & Marsh, 1982) there is massive
ventral thickening of the body wall reinforced with irregular plates, some acting as super-
ambulacrals, more or less deeply embedded in the soft tissue (Fig. 5d). N. crassa also carries
to an extreme for Nepanthia the number of secondary (and even tertiary) abactinal plates.
At the same time, its primary abactinals develop a more rounded than crescentic form com-
pared with other Nepanthia species such as N. belcheri, such a modification having a parallel
in P. formosa as opposed to the P. granifera (and indeed in Allopatiria ocellifera as opposed
to the geographically adjacent Asterina stelliferd). Marsh (pers. comm.) confirms the presence
of obscured superambulacrals in N. crassa and has also found them in N. maculata and
belcheri. This structure contrasts with the other Asterininae where the reinforcing plates for
the ventrolateral angle do not extend adradially to help support the ambulacral arch (see
Figs le, 5c). Blake (pers. comm.) suggests that the development of superambulacrals in this
instance is correlated with the relatively narrow and more nearly cylindrical arms, compared
with most asterinids. The two South African species approximate to Nepanthia in body form,
though the arms broaden slightly at the base to make blunt-angled or rounded interradial
arcs.

Clearly, if these characters are as important as I believe (particularly the last), then the
affinities of P. granifera and formosa are with Nepanthia (and especially with species such
as TV. crassa and N. belcheri with thicker body- walls and more numerous secondary abactinal
plates) rather than with Patiria sensu Gray, 1 840 — now synonymized with Asterina. How-
ever, this contradicts Fisher's conclusion of 1941 that granifera is close to Patiria miniata
but crassa is allied with Nepanthia. If Fisher's evaluation of the affinities is correct and at
least a generic distinction should be recognized between granifera and crassa, then
Callopatiria needs to be revived for P. granifera and formosa. The zoogeographical
separation, with the westernmost record for Nepanthia being from Burma, supports such
a distinction but the morphological characters are individually relatively unimportant. They
include the following: In the two South African species the arms are less well defined basally
so that the interradial arcs are rounded or only bluntly angled; the superomarginal plates
appear to match the inferomarginals in size during growth, the successive ones remaining
contiguous whereas in most Nepanthias seen the superomarginals are smaller, some of them
being separated by interstitial abactinal plates and in N. briareus, which has particularly
cylindrical arms and ill-defined ventrolateral angles, the superomarginals outnumber and fail
to match the inferomarginals; the actinal armament is much more coarse while the spinelets

GANERIIDAE & ASTERINIDAE

371

Fig. 7 Allopatiria ocellijera (Gray), BM reg. no. 1969.12.16.13, Cape Lever, Mauritania, R 38 mm
(the uppermost ray denuded), x 1^ (above); 'Patiria ' obtusa Gray, holotype, 1938.6.23.24, Panama,
R up to 54 mm (the lower right ray partly denuded, and a few plates lost) x \\ (middle), side view
of detached arm, partly denuded, of the same, x3 (below).

372 AILSA M. CLARK

of at least the proximal plates usually number less than ten and form single or double fans,
especially informosa, rather than being very fine and forming dense clusters as in the species
of Nepanthia; lastly, the development of the superambulacral plates is less advanced, as
outlined above. Cumulatively, these differences probably do justify the revival of Callopatiria
with generic rank but the relationship with Nepanthia merits further attention. That genus
has already been broadened in concept considerably by the inclusion of Parasterina crassa
with its very thick body wall and complex abactinal skeletal structure.

The only remaining species of Patiria is P. obtusa Gray, 1847. The poorly preserved
holotype ((Fig. 7 below) with its relatively long arms (R/r 53/17 mm = 3-1/1) and irregular
abactinal plating (possibly at least partly due to the condition) appears very unlike an
Asterina but superficially resembles Callopatiria granifera, with which Verrill (1913) con-
sidered it congeneric. However, the similarity does not extend to the internal characters since
the broken arm shows a complete absence of any superambulacrals or other internal rein-
forcing plates. The inferomarginals are relatively large, appearing pearshaped in the section
view (Fig. 5a) but flattened and tapering externally to appear long oval on the surface; their
relatively ventral position is probably due to the unnatural flattening of the specimen and
this may also account for the rounded ventro- lateral profile. The species is not a ganeriid
since the marginals are not at all blocklike, the adjacent abactinal and actinal plates are
strongly imbricating so as to reinforce the ventrolateral area, quite unlike those individuals
of Cycethra with marginals reduced to this same small superficial size. Without more
material in better condition, a proper assessment of the position of 'Patiria' obtusa among
the asterinids cannot be made. The species is known only from the holotype, Verrill's near-
pentagonal specimen from Panama recorded under this name (1 869) with its relatively sparse
actinal armament (rather than dense clusters of spinelets) being some other asterinid (perhaps
Enoplopatiria siderea Verrill, 1913, also from Panama). Neither H. L. Clark (1940) in the
echinoderms of the Zaca expedition to west Central America, nor A. H. Clark (1946) in
the echinoderms of the Pearl Islands, Bay of Panama, recorded obtusa, so there must be an
element of doubt about its source.

The name Asterinopsis has been declared invalid by H. L. Clark (1938), since Mortensen
(1933) noted that Lamarck's material of the type species is lost and its precise locality was
uncertain. H. L. Clark noted that the two Atlantic species included in Asterinopsis by Verrill:
Asterina lymani and A. pilosa of Perrier, 1881, are not congeneric with the Indo- Pacific
species subsequently included, leaving them in limbo when he transferred some of the
non- Atlantic species to Paranepanthia Fisher, 1917.

Finally, Allopatiria was for long of uncertain value, owing to ignorance of the type locality
of the only species, Patiria ocellifera Gray, Verrill's supposition of Australia being unsub-
stantiated. In 1963 I redescribed the holotype, prompted by Professor Tortonese who had
just discovered a similar specimen from the Mediterranean and who subsequently (1963)
synonymized with A. ocellifera the Mauritanian Parasterina africana Engel & Croes, 1960.
I believe that both Patiria rosea and P. pulla Koehler & Vaney, 1906, again from Mauritania,
are also synonyms of A. ocellifera. Apart from colour differences in the type specimens
(pink/red as opposed to grey with yellow spots) these two were supposed to differ from one
another by minor differences in the body shape, the denuded condition of some of the
enlarged abactinal plates in the holotype of rosea and different degrees of prominence of
the furrow spines — the last two almost certainly due to artefacts of preservation. Madsen
(1950) has already suggested that these differences are only individual variations when he
provisionally synonymized pulla with rosea. The figures of rosea (1906, pi. 5, figs 3, 4) show
a considerable resemblance to the holotype of A. ocellifera illustrated in A.M.C. (1963, pis
1, 4), despite the much larger size of the latter (R c.80 mm as opposed to 43 mm in the
holotype of rosed).

Allopatiria ocellifera has a very distinctive abactinal skeleton with extremely numerous
secondary plates (at least at R>30 mm), many of them superimposed on the primary plates
(or at least their peripheral parts), so that only rounded areas of most of the radial 'field'
primaries of the rays are not overlain (Fig. 3g). The armament of the primaries (when intact)

GANERIIDAE & ASTERINIDAE 373

is a dense covering of short blunt spinelets, though the secondaries often bear a high propor-
tion of two- or three- valved spiniform pedicellariae. Asterina stellifera also has similar pedi-
cellariae, body form and large maximum size (R max. > 50 mm) and occurs in West Africa
to the south of Cape Verde, as well as on the east coast of South America. Despite these
factors, I think that Verrill's generic distinction is well justified. A. stellifera and the other
species of Asterina have short but broad midradial primary abactinals, many of them with
a more or less well marked double concavity in the proximal side formed by three lobes
of varying prominence and the other primaries of the pore areas are clearly crescentiform.
A relatively small specimen ofAllopatiria ocellifera (a paratype of Parasterina africand) with
R 1 7 mm has three longitudinal series of apparently round primary plates on the radial
'fields', the more proximal ones revealed as rhombic when the spinelets are removed, the
widest part being at the middle of the length. These plates just imbricate at the angles and
the secondaries are limited to the interstitial rounded popular areas between them, number-
ing up to five in each area. The distal primaries, especially of the two outer series, become
relatively broad and short and their armament resolves itself into two transverse linear series
of spinelets. As growth proceeds, the secondaries become much more numerous, encircling
the primaries and then encroaching progressively on their upper surface so that at R 80 mm
(possibly before) only some of the distalmost primaries (which have become relatively large
and rounded) are exposed except for their spinelets (A.M.C., 1963). This highly modified
abactinal skeleton I believe fully justifies retention of a generic distinction for Allopatiria
as suggested by Tortonese (1963).

Before revised diagnoses can be given of the families Ganeriidae and Asterinidae in the
light of the present comments, it is necessary to describe a new aberrant genus of Asterinidae.

Genus PAXILLASTER1NA nov.

DIAGNOSIS. A genus of Asterinidae with very flat, almost pentagonal, body form; primary
abactinal plates not arranged in distinct radial and lateral 'fields' but regularly arranged, thin
and flat except for an abrupt, short, flat-topped central column on many plates, including
the midradial row of proximally 3-lobed plates, the columns replaced on others — especially
the first adradial row each side of the midradial row — by a smaller rounded convexity, all
these elevations crowned by a tall pompom or rounded tuft of numerous slender spinelets
which are embedded in a soft matrix so as to radiate from a central conical core and not
articulate directly with the elevation on the plate; only a few small secondary abactinal plates
present proximally; margin very thin, formed by the inferomarginal plates only, the supero-
marginals small and inset dorsally; actinal and adambulacral plates armed with relatively
few (<6) slender spinelets webbed together into fans; no pedicellariae observed.

TYPE SPECIES. Paxillasterina pompom sp. nov.

REMARKS. Superficially the small asterinid on which this new genus is based (R only c. 1 5 mm
in the four specimens) resembles the flatter, more nearly pentagonal species of Asterina such
as A. folium (Liitken) but is marked off on closer scrutiny by the many paxilliform abactinal
plates and their peculiar armament. It should be noted that in poorly-preserved and
especially in dried specimens the pompoms contract and may become appressed on to the
surface and the low paxillar columns can only be appreciated when the skeleton is denuded.
Possibly similar specimens have been collected and attributed to the common Caribbean
A. folium, this modification having been overlooked. It necessitates an alteration to the
diagnosis of the Asterinidae in which truly paxilliform plates were not previously known.
Conversely the diagnosis of the Ganeriidae also needs to be qualified to allow for the non-
-paxilliform plates of individuals of Cycethra verrucosa such as were previously referred to
Patiria obesa.

374

AILSA M. CLARK

Paxillasterina pompom sp. nov.
Figs 8, 9

HOLOTYPE. Korbiski Reef, San Bias Is, off Atlantic Panama east of the Canal Zone, c.6 m.,
under coral rubble. Collected by Dr G. Hendler.

R/r 14/1 1 mm — 1-3/1. The form is pentagonal, the body thin with central height only
5 mm (including the armament). The primary abactinal plates are very regularly arranged
in rows parallel to the radii (or in chevrons if regarded interradially), and there is no distinc-
tion of radial and lateral 'fields' of plates, though the more adradial plates have the superficial
part crescentic visible when denuded, subtending a single papular pore, except for the plates
of the mid-radial row which have a median proximal lobe and a double concavity subtending
two pores, the more lateral plates rhombic or fan-shaped; all these plates rather thin with
only an inconspicuous texturing of crystal bodies and flat except for a central elevation which
takes the form of a well-defined but low flat-topped column on many plates, including all
of the midradial row and most of the second adradial each side, as well as scattered other
plates; these columns crowned by a tall ovate pompom of numerous fine needle-like spinelets
numbering up to c.50 and embedded in a soft matrix so that their bases arise from a central
uncalcified cone and not directly from the column; other primary plates, including the first
adradial row, have a small rounded central convexity bearing a smaller pompom in propor-
tion, some plates adjoining the superomarginals having as few as 6 spinelets in the cluster;
one or two small secondary abactinal plates occur around some of the proximal papular
pores; there are 10 longitudinal rows of pores on each ray, the outermost of only 4 pores.
The superomarginal plates are mostly smaller than the adjoining abactinal plates and are
rounded or D-shaped; each bears a small pompom, the more interradial ones with up to
c. 1 5 spinelets. The inferomarginal plates project well beyond the superomarginals to form
the edge of the body, their adradial parts are flattened dorso-ventrally but the free part is
prolonged and constricted so as to be rounded in cross section and separated from the adjac-
ent inferomarginals; each bears a laterally compressed cluster of numerous fine spinelets,
the apical ones slightly coarser than the abactinal spinelets. The actinal plates are in regular

Fig. 8 Paxillasterina pompom gen. & sp. nov., holotype, USNM, Korbiski Reef, Atlantic side of
Panama, R 14 mm (a) and (b) parts of denuded and intact proximal abactinal midradial areas; (c)
three paxillar spinelets; (d) vertical section of one pompom (detached from column); (e) and (0 two
pairs of marginal plates in dorsal view, denuded and intact (proximal to the left). The scale measures
1 mm for (c) and 2 mm for the rest.

GANERIIDAE & ASTERINIDAE

375

Hiiiiiiiiii

Fig. 9 Paxillasterina pompom, holotype, from above and below, x4.

376 AILSA M. CLARK

chevrons and each has two or three adradial lobes (distinct when denuded); each bears a
tangentially-aligned fan of up to 5 spinelets, mostly 3 but the admarginal ones with 5,
joined by skin. The adambulacral plates each have a shallow abradial step bearing a fan of
subambulacral spinelets aligned slightly obliquely to the furrow and close behind the fan
of furrow spines, which are slightly longer but arise at a higher level so that the tips project
to about the same extent; each fan is usually of 4 spines or spinelets and webbed. Each jaw
is armed with a larger fan of up to 16 spines, 7 or 8 on each oral plate, the apical ones about
twice as large as the outermost; there is also a pair (or trio) of small suboral spinelets on
each plate. The gonopores cannot be distinguished.

PARATYPES: Three other specimens are also available from the Atlantic side of Panama. One
is also from Korbiski Reef but in 3 metres depth; it has R/r 15/1 1-5 mm and is similar to
the holotype but for having usually 5 furrow spines. Another, from Mackerel Reef, also San
Bias Is, was only collected two years ago and still shows traces of yellow colouration in spirit.
The third is from 'Landing Craft Reef, south of the Galeta Marine Laboratory, Canal Zone,
'under rock in the coralline zone or higher'; it is dried and distorted with the pompoms
mostly appressed horizontally; R is c.lOmm and the subambulacral spinelets usually
number only 3.

Family GANERIIDAE Sladen

Asterinidae: Ganeriinae Sladen, 1889 : 375-376.

Ganeriidae: Perrier, 1894: 171; Fisher, 191 la: 251 (in key); Verrill, 19146: 365-366; Fisher, 1940:
127; A. M. Clark, 1962 : 23-24; Bernasconi, 1964 : 59-60; Spencer & Wright, 1966 : U69; Blake,
1981 : 380-381 (pt). [Non Ganeriidae: A. H. Clark, 1938, Downey, 1973 (Leilas ter)]

Asterinidae (pt): Madsen, 1956 : 22-23; Bernasconi, 1973 : 335-336 (Patina obesa).

Goniasteridae (pt): Bernasconi, 1965 : 333-335 (Vemaster).

A family of Valvatida with body stellate, rarely almost pentagonal, interradial arcs rounded,
arms normally five, rounded in cross-section except in the aberrant abyssal Vemaster, abacti-
nal plates imbricating, usually paxilliform but the columns sometimes broadened and not
well-defined, or coarse and tabulate, or with the basal lobes reduced and the plates obscured
by thickened pustular body wall (in wet specimens of Perknaster), crowned with clusters
of spinelets, which may be short and superficially appearing granuliform in Ganeria, some
specimens of Cycethra and in Vemaster, papulae in small groups in the meshes between
the primary plates, more numerous adradially; marginal plates ranging from blocklike and
conspicuous to indistinguishable from the adjacent plates, aligned laterally or towards the
ventral side when smaller (as usually in Perknaster), reduced to one slightly projecting series
(presumed inferomarginals) in Vemaster, armed with oval groups of spinelets or spines, or
one or two vertical combs when coarser, as in Ganeria; actinal plates imbricating adradially,
tending to form oblique transverse as well as longitudinal rows, especially the latter, each
armed with a group of spinelets or one or two larger spines, except in Vemaster where arma-
ment is rare and the plates are irregular and possibly secondary and limited to the interradii;
adambulacral plates small, matching the adradial row of actinal plates, armed with two (or
sometimes one) furrow spines and several subambulacral spines, often forming pairs parallel
(or oblique) to the furrow or all the spines forming one (or two) series transverse to the
furrow; pedicellariae absent (rarely a few spinelets forming a simple fascicle); internally a
proximal interradial calcified pillar supporting the abactinal body wall dorsolaterally, no
superambulacral plates.

Family ASTERINIDAE Gray

Asterinidae Gray, 1840: 228; Perrier, 1875: 27-28[292]; Sladen, 1889: 374-376; Fisher, 191 la:
253-254; Verrill, 1913: 477-481; 19146: 262-263; 364-365; Fisher, 1919: 409; Madsen, 1956:
22-23; Spencer & Wright, 1966 : U68; Bernasconi, 1973 : 335-336; A. M. Clark & Courtman-Stock,
1976 : 75; Blake, 1981 : 380-391 (pt).

GANERIIDAE & ASTERINIDAE 377

A family of Valvatida with body form usually pentagonal to stellate but sometimes the rays
narrow and more or less finger-like, usually numbering five but occasionally more, the
surface completely flat below (unless the whole body is arched upwards when it is concave)
but variously convex above, ranging from very low, the body leaf-like, to high with almost
cylindrical rays but still with a distinct ventro-lateral angle supported internally by over-
lapping extensions of the lateralmost abactinal and actinal plates, the angle culminating in
the relatively small inferomarginals, or sometimes blunted by similar prominence of both
marginal series (in Nepanthia and Callopatiria); abactinal plates superficially flattened or
convex, sometimes with a low ridge bearing the armament and in one case (Paxillasterina
nov.) with a low circular paxillar column, the primary radial and interradial plates of the
disc often distinguishable by their regular shape, the larger lobed more adradial plates of
the papular areas appearing more or less crescentic, imbricating adradially (the more lateral
plates) or proximally (the more midradial plates), often two magnitudes of plates, at least
adradially, with much smaller secondary plates subdividing or restricting the papular spaces
proximal to the primary plates, often with considerable regularity of arrangement of all the
plates, though this tends to become obscured in large specimens; abactinal armament of
multiple small spinelets, sometimes granules, on the middle or raised part of the plate, or
armament lacking when the skin is more or less thickened; papulae usually single, sometimes
in small clusters, absent near the margin; superomarginal plates small, often distinguishable
from the adjacent abactinal plates only by their position above (sometimes alternating with)
the inferomarginal plates which are seen in transverse sections of the ray to be wedge- or
pear-shaped or peg-like, appearing as a reversed keystone wedged in among the overlapping
inner ends of the adjacent lateralmost abactinal and actinal plates, but in external view
appear oval or rectangular; the very flattened genera Anseropoda (and probably also
Stegnaster and Tremaster) with less massive reinforcement for the very narrow angle, the
entire body except for a narrow midradial band strengthened by overlapping struts projecting
from the abactinal and corresponding actinal plates to form a latticework; actinal plates
imbricating adradially, forming more or less regular transverse or oblique series from furrow
to margin, also tending to form longitudinal rows parallel to the furrow but the abradial plates
sometimes irregular, armed with one or several spinelets, when multiple arranged in fans
or clusters: adambulacral plates relatively small, matching the innermost actinal rows and
with armament usually of similar magnitude, furrow spines in a fan or comb; pedicellariae,
if present, usually simple alveolar with two, occasionally three, slightly curved spiniform
valves, sometimes (Nepanthia) fasciculate with more numerous valves; internally a proximal
interradial pillar supporting the abactinal body wall; superambulacral-like plates developed
more or less regularly in the narrow-rayed genera Nepanthia and Callopatiria but absent
elsewhere.

Acknowledgements

I am indebted to Dr W. Decraemer of the Institut Royal des Sciences Naturelles, Brussels,
for the loan of specimens, also to Professor E. Tortonese, Professor D. B. Blake, Mrs L. M.
Marsh, Miss M. E. Downey, Dr A. Dartnall and Dr M. Jangoux for constructive exchange
of ideas and, or information.

Synopsis

Leilaster A. H. Clark, 1938 (family Ganeriidae) and Mirastrella Fisher, 1940 (family

Asterinidae) are abstracted from these families and provisionally referred to the order

Spinulosida, family yet to be determined.
Vemaster Bernasconi, 1965, is referred from the Goniasteridae to the Ganeriidae but may

warrant isolation in a suprageneric taxon when the whole family is reviewed.
Patina obesa (H. L. Clark, 1910) is found not to be an asterinid but is synonymized with

Cycethra verrucosa (Philippi, 1857), family Ganeriidae.

378 AILSA M. CLARK

Patiria Gray, 1840, type species: P. miniata (Brandt, 1835) is synonymized with Asterina

Nardo, 1834.
Patina miniata (Brandt, 1835), P. pectinifera (Muller & Troschel, 1842), P. chilensis

(Liitken, 1859) and P. stellifera (Mobius, 1859), are referred to Asterina.
Patiria granulosa (Perrier, 1875) is a synonym of Asterina miniata (Brandt, 1835).
Patiria granifera Gray, 1847 and P. formosa (Mortensen, 1933) are referred to Callopatiria

Verrill, 1913, revived.
Patiria pulla and P. rosea Koehler & Vaney, 1906, are synonymized with Allopatiria

ocellifera (Gray, 1 847).
Paxillasterina gen. nov. for type sp. P. pompom sp. nov. from Atlantic Panama is described.

The following taxa are thought to be in need of reassessment:
Ganeriidae: Aleutiaster A. H. Clark, 1939

Hyalinothrix Fisher, 19116

Knightaster H. E. S. Clark, 1972

Tarachaster Fisher, 1913.
Asterinidae: Patiriella Verrill, 1867

'Patiria ' obtusa Gray, 1 847

'Enoplopatiria ' siderea Verrill, 1913

Appendix — The generic name Asterina and its type- species

The first use of the name Asterina was by Nardo (1834 : 716) for the two species Asterias
minuta Linnaeus and A. exigua Lamarck, neither one designated as type species. In 1836
L. Agassiz adopted Asterina Nardo and cited only Asterias minuta Linnaeus. The first formal
type designation now traced is the apparently provisional one of Fisher (1906 : 1087) stating
'Type (?) Asterina gibbosa (Pennant). Asterina minuta (Olivi) Nardo = Asterias gibbosa',
repeated without the query and abbreviated to 'Asterias minuta Olivi = Asterias gibbosa
Pennant' by Fisher in 191 la: 254, while Verrill (1913: 479) modified it to 'A. minuta
Nardo = A. gibbosa (Pennant)'. Asterias minuta Linnaeus dates from the Fauna Svecica
(1761 : 512) and was described from Norwegian waters in terms mainly comparative with
Asterias rubens. In the 12th and 13th (Gmelin) editions of the 'Systema naturae' (1767 and
1791) the concept of A. minuta was extended to include first the Asterina-like seastar from
American waters shown in pi. 5, figs 14 and 15 of Seba (1758) and then A. minuta: O. F.
Muller, 1776, again from Scandinavia, and A. minuta: Fabricius, 1780, from Greenland. In
none of these localities does the common European starlet widely known as Asterina gibbosa
occur and Verrill (1914#) notes of [A] minuta Linnaeus, 1761 There is nothing in the few
words of description to show that it is not the young of A. rubens or some similar species.'
At first sight, this appears to render Asterina Nardo a synonym of Asterias Linnaeus.

However, it is evident that Fisher was aware of this anomaly, hence his introduction of
'Asterias minuta Olivi' (1792), a record from the Adriatic where gibbosa is found (also the
recently recognized Asterina phylactica Emson & Crump but that diminutive species may
be ignored in this context). Articles 69 and 70 of the Code of Nomenclature, on subsequent
designation of type-species and deliberate use of misidentification, are intended to allow for
this sort of problem so that the species which Fisher had in mind — namely Asterias
gibbosa Pennant, 1777 — is usable as type-species of Asterina Nardo. Evidence that this
concept of Asterina is in accord with Nardo 's lies in Forbes' use of Asterina gibbosa
already in 1839.

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Manuscript accepted for publication 14 December 1982

Peniculus haemuloni, a new species of copepod
(Siphonostomatoida: Pennellidae) parasitic on
Haemulon steindachneri from Ubatuba, Brazil

P. D. Alexander

*c/o The Department of Zoology, British Museum (Natural History), Cromwell Road,
London SW7 5BD.

Introduction

The genus Peniculus was established by Nordmann, 1 832, to accommodate Peniculus fistula,
a parasite of the teleost fish — Diplodus annularis Linnaeus (as Sargus annularis). To date
17 nominal species exist although according to Kabata (1979) at least two of them should
be regarded as species inquirendae: P. calamus Nordmann, 1864 and P. fissipes Wilson,
1917, the latter probably being a synonym of the type species. The new species is the first
record of Peniculus from Brazilian waters and the first record of Haemulon as a host for
Peniculus. The structure of the appendages within the Pennellidae is so uniform that they
are of little taxonomic value for species recognition. The key to the species of Peniculus
is based on gross body morphology.

Description of new species
Peniculus haemuloni n.sp.

Adult female: The cephalothorax is about one tenth as long as the trunk (Fig. 1A), ovoidal,
and flattened dorsally but convex ventrally (Fig. IB) with a pair of rounded swellings anter-
iorly bearing the second antennae (Fig. ID). The prominent mouth tube is directed postero-
ventrally (Fig. 1C). The four thoracic segments form a relatively short, uniformly slender
neck comprising about one sixth of the trunk length. The neck is slightly flattened dorso-
ventrally and has a conspicuous internal system of longitudinal and transverse chitinous
thickenings (Fig. 1C). Posteriorly the neck bears three pairs of swimming legs. The elongate
subcylindrical trunk is slightly dorsoventrally flattened and comprises four-fifths of the total
body length (Fig. 1 A). A pair of terminal ventrolateral swellings bears the oviduct openings.
The egg sacs are straight and uniseriate (Fig. 2B). The small rounded, dorsally placed abdo-
men bears two groups of four short setae (Fig. 2C) which are presumed to represent the caudal
rami. The first antennae are absent. The second antennae are stout with a strongly curved
terminal claw. The mouth tube is well developed extending almost to the posterior margin
of the cephalothorax. The apex of the mouth tube has a circular opening surrounded by
a thin membrane. The swimming legs comprise an oval flattened protopod and a 2-
segmented exopod. The first segment is devoid of armature elements, the second has five
setae on both the first and the third legs, the second leg armature was missing. Each limb
pair is connected by a sternal bar (Fig. 2 A). Dimensions of the holotype female: total length
(exc. egg sacs) 5-6 mm; trunk length 4-4 mm; trunk width, 0-6 mm; neck length, 0-7 mm; neck
width, 0- 1 mm; cephalothorax length, 0-5 mm; cephalothorax width, 0-35 mm; egg sac width,
0-16 mm.

MATERIAL EXAMINED. Two adult females from the tail fin of the teleost Haemulon steindach-
neri collected at Ubatuba, Brazil (23°27 'S: 45°06 'W) by Dr K. Rohde in March 1979. The

*Work completed during a six month secondment from the Department of Applied Biology, U.W.I. S.T. King
Edward VII Avenue, Cardiff.

Bull. Br. Mus. nat. Hist. (Zool.) 45(7): 381-385 Issued 24 November 1983

381

382

P. D. ALEXANDER

0.1mm

Fig. 1 Peniculus haemuloni n.sp. A, Female, ventral; B, cephalothorax, lateral; C, cephalothorax and
neck showing mouth tube and internal chitinous framework, ventral; D, Anterior of cephalothorax
showing second antennae.

NEW COPEPOD FROM BRAZIL

383

Fig. 2 Peniculus haemuloni n.sp. A, Structure and position of the three pairs of limbs, ventral; B,
posterior region of trunk and egg sacs, lateral; C, posterior region of trunk showing caudal setae,
ventrolateral.

ovigerous female is the holotype, registration number 1979.683; paratype female, registration
number 1979.684. Both are deposited in the collections of the British Museum (Natural
History), London.

REMARKS. The presence of only three pairs of swimming legs distinguishes Peniculus haemu-
loni n.sp. from most known species of the genus. Two other species, Peniculus sp.
Kazatchenko & Avdeev, (1977) and P. sciaenae Gnanamuthu (195 la), have been described
with three pairs of swimming legs. The former has the first two close together with a marked
gap before the last pair at the base of the neck, whereas in P. haemuloni all three legs are
close together at the base of the neck. It is probable that leg 1 of P. sciaenae was overlooked
by Gnanamuthu (195 la) because this species appears to be closely related to P. trichiuri,
P. stromatei, P. theraponi and P. scomberi although the last species apparently lacks legs
in the adult female. All of these species were described by Gnanamuthu (195 la, b) each com-
ing from a host belonging to a different family of teleost fish found in Indian waters. All

384 P. D. ALEXANDER

possess a posterior swelling on the ventral surface of the cephalothorax and exhibit the same
distribution of swimming legs, where present, along the neck and trunk. The differences
between these species are relatively minor and it is possible that they represent forms of a
single variable species.

Another useful character distinguishing P. haemuloni is the large prominent mouth tube
which is similarly well developed in only two other species, P. clavatus Kreyer, 1863 and
P. fistula Nordmann, 1832. The length to width ratio of the trunk (6-7 : 1) is most closely
matched by that of P. scomberi (6-5 : 1) and P. fistula (6-3 : 1). These characters apart the
new species shares many features with other Peniculus species, such as cephalothorax shape,
the small rounded abdomen and the presence of a chitinous framework in the neck.

Key to the species of Peniculus (adult females only)

1 Cephalothorax with 4 large holdfast processes P. asinus Kabata

- Cephalothorax without such processes 2

2 Cephalothorax with rounded swelling on ventral surface posterior to mouth tube ... 3
Cephalothorax without posterior swelling on ventral surface 6

3 Swimming legs apparently absent P. scomberi Gnanamuthu

- Swimming legs present 4

4 3 pairs of swimming legs present P. sciaenae Gnanamuthu

4 pairs of swimming legs present 5

5 Trunk about 1 1 times longer than wide P. trichiuri Gnanamuthu

Trunk about 8 times longer than wide . P. stromatei GnanamuthuAP. theraponi Gnanamuthu*

6 3 pairs of swimming legs present -. 7

4 pairs of swimming legs present 8

7 All 3 pairs of legs positioned close together at the base of the neck ... P. haemuloni n.sp.
Last 2 pairs of legs separated by a gap Peniculus sp. Kazatchenko & Avdeev

8 Legs 3 and 4 closer together than legs 1 and 2 P. communis Leigh-Sharpe

Legs 3 and 4 further apart than legs 1 and 2 9

9 Trunk more than 6 times longer than wide P. fistula Nordmann

Trunk between 3 and 4-5 times longer than wide 10

10 Mouth tube forming a massive posteriorly-directed proboscis P. clavatus Kr0yer

Mouth tube not forming posteriorly-directed proboscis 11

11 Cephalothorax ovoid 12

Cephalothorax widest near posterior margin and tapering anteriorly . P. elegans Leigh-Sharpe

12 Abdomen well developed; longer than wide P. minuticaudae Shiino

- Abdomen minute; posterior margin of trunk more-or-less straight 13

13 Trunk 4-3 times longer than wide; neck less than half as long as cephalothorax

P. ostraciontis Yamaguti

Trunk 3-3 times longer than wide; neck more than half as long as cephalothorax

P. truncatus Shiino

- N.B. There appear to be no reliable characters separating these two species.

Acknowledgements

I would like to thank Dr K. Rohde of the Department of Zoology, University of New
England, Armidale (Australia) for providing the specimens upon which this study is based.
I am also grateful to Dr G. A. Boxshall and Dr R. J. Lincoln of the Department of Zoology
(Crustacea), British Museum (Natural History), London for their help and advice with this
paper.

References

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NEW COPEPOD FROM BRAZIL 385

Gnanamuthu, C. P. 195 la. Three new species of Lernaeid copepods parasitic on South Indian fishes.

Annals and Magazine of Natural History, (12) 4 : 77-86.
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of Zoology 55: 1988-1991.

Kabata, Z. 1979. Parasitic Copepoda of British Fishes. The Ray Society, London, 468 pp. 199 pi.
Kazatchenko, V. N. & Avdeev, G. V. 1977. Parasitic copepods (Crustacea) collected during the 57th

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Instituta Okeanologii 107 : 35-37 (In Russian).

Leigh-Sharpe, H. 1934. Commensal and parasitic copepods. Part II. Siboga Expeditie XXIX: p. 35.
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2. G. Reimer, Berlin, pp. 1-150.
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Manuscript accepted for publication 14 December 1982

British Museum (Natural History)

An Atlas of Freshwater Testate Amoebae

C. G. Ogden & R. H. Hedley

1980, Hardcovers, 222pp, £1 7.50 (£1 8.00 by post). Co-published by British Museum
(Natural History) and Oxford University Press.

This book illustrates, using scanning electron micrographs, most of the common
species of testate amoebae that are found in freshwater habitats. Information on
the biology, ecology, geographical distribution and a classification are followed by
descriptions of ninety-five species. Each of these is illustrated by several views of
the shell.

The text is designed not only to enable biologists to identify species of testate
amoebae, but to serve as an introduction to students interested in the taxonomy
and biology of these freshwater protozoa. It will be of special interest to
protozoologists, ecologists, limnologists, water treatment specialists and
micropalaeontologists interested in recent sediments.

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