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Bulletin of the 

British Museum (Natural History 

Zoology series Vol 41 1981 

British Museum (Natural History) 
London 1982 

Dates of publication of the parts 

No 1 . . .27 August 1981 

No 2 . . 24 September 1981 

No 3 . 29 October 1981 

No 4 26 November 1981 

No 5 17 December 1981 

ISSN 0007-1498 

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

Zoology Volume 41 


No 1 Anatomy and phylogeny of the Chinese Major Carps Ctenopharyn- 
godon Steind., 1866 and Hypophthalmichthys Blkr., 1860 
By Gordon Howes ......... 1 

No 2 Morphology and systematics of some interior-walled cheilostome 
By P. L. Cook & P. J. Chimonides 53 

No 3 Notes on Atlantic and other Asteroidea. 1. Family Benthopectinidae 

By Ailsa M. Clark 91 

No 4 Miscellanea 

Observations on clonal cultures of Euglyphidae (Rhizopoda, Protozoa) 

By C. G. Ogden 137 

The Nuttall and British Museum (Natural History) tick collections: 
lectotype designations for ticks (Acarina: Ixodoidea) described by 
Nuttall, Warburton, Cooper and Robinson 

By J. E. Keirans & B. E. Brewster ... 153 

A revision of the spider genus Hispo (Araneae : Salticidae) 

By F. R. Wanless 179 

A revision of the spider genus Phaeacius (Araneae : Salticidae) 

By F. R. Wanless 199 

The protractor pectoralis muscle and the classification of teleost fishes 

By P. H. Greenwood & G. V. Lauder Jnr 213 

No 5 Miscellanea 

The ostracod genus Loxoconcha from Abu Dhabi lagoon, Persian Gulf 

By R. H. Bate & A. Gurney . 235 

A revision of the spider genus Cocalus (Araneae : Salticidae) 

By F. R. Wanless . 253 

The Phthiracarus species of C. L. Koch 

By B. W. Kamill ... ... .263 

The status ofLamingtona lophorhina McKean & Calaby, 1968 (Chirop- 
tera : Vespertilionidae) 

By J. E. Hill & K. F. Koopman 275 

The status of Hipposideros galeritus Carter, 1846 and Hipposideros 

cervinus (Gould, 1854) (Chiroptera: Hipposideridae) 

By P. D. Jenkins & J. E. Hill 279 

XV * 

Bulletin of the 

British Museum (Natural History) 

Anatomy and phylogeny of the Chinese 
Major Carps Ctenopharyngodon Steind., 
1866 and Hypophthalmichthys Blkr., I860 

Gordon Howes 

Zoology series Vol 41 No 1 27 August 1981 

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

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London SW7 5BD, 

World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 

Trustees of the British Museum (Natural History), 1981 

ISSN 0007-1498 Zoology series 

Vol 41 No 1 pp 1-52 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 27 August 1981 


Anatomy and phylogeny of the Chinese Major 
Carps Ctenopharyngodon Steind., 1866 and 
Hypophthalmichthys Blkr., 1860 

Gordon Howes 

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


Synopsis 1 

Introduction 1 

Abbreviations used in the figures 2 

Study material 3 

Comparison of selected characters in Ctenopharyngodon and Hypophthalmichthys 4 

Analysis of apomorph characters 4 

Phylogenetic relationships 38 

Relationships of Ctenopharyngodon 38 

Synopsis of taxa assigned to the Squaliobarbine group 40 

Relationships of Hypophthalmichthys 41 

Synopsis of taxa assigned to the Abramine group 45 

Discussion 47 

Classification of the Leuciscinae 47 

Leuciscine hybrids 48 

Classification of the 'true' carps 49 

Acknowledgements 49 

References 49 


The Chinese major carps, Ctenopharyngodon and Hypophthalmichthys, long held to belong 
to the subfamily Leuciscinae are each shown to represent separate lineages of a diphyletic 
Cyprinidae. This result is based on the analysis of 22 character complexes and the identifica- 
tion of supposed apomorphies. Ctenopharyngodon forms part of a monophyletic assembly 
termed the squaliobarbine group and Hypophthalmichthys is identified as the most derived 
member of the abramine group. The squaliobarbines are considered to be the sister group of 
all other barbelled Cyprinidae. The incongruity of character sets results in the presentation 
of two hypotheses for abramine intrarelationships. Hypophthalmichthys possesses a unique 
form of epibranchial organ; derived epibranchial structures are also identified in Xenocypris 
and Abramis. In the light of the data presented, the classification of the subfamily Leuciscinae is 
evaluated and many identifications of 'intergeneric' hybrids are falsified. Some comment is made con- 
cerning the phylogeny of barbelled cyprinids. 


The subjects of this study are the Chinese major carps, Ctenopharyngodon (the grass carp or 
white-amure) and Hypophthalmichthys (the silver and big-head carps). The object is to put 
forward a hypothesis of relationships of these taxa based on synapomorphic characters. 

Bull. Br. Mus. nat. Hist. (Zool.)41 (1) : 1-52 Issued27 August 1981 

2 G. J. HOWES 

The major Chinese carps are of considerable economic importance as sources of protein 
and, in the case of Ctenopharyngodon, a control agent in restricting pond and canal weed 
growth. That these carps are primary-consumers and breed well in captivity are additional 
assets in their culture. Although there is much information on their biology-breeding, 
physiology, biochemistry and culturing, there are very few papers describing their anatomy. 
Of these only two or three authors have made statements regarding possible phylogenetic 

Ctenopharyngodon is a monotypic genus included by Chu (1935) in the subfamily 
Leuciscinae. Berg (1949) considered the genus to be closely related to Scardinius and placed 
it in his all-embracing Cyprinini. Gosline (1978) placed Ctenopharyngodon, Mylo- 
pharyngodon and Squaliobarbus together, and although he was undecided whether they 
belonged to the subfamily Cyprininae or Leuciscinae, favoured retention in the latter. Even 
though the characters Gosline used for uniting these genera are considered by the present 
author to be a mosaic of plesio- and apomorphies, his hypothesis of relationships is not 
refuted in this study. 

Hypophthalmichthys includes three nominal species, H. molitrix (Val.), H. nobilis 
(Richardson) and H. harmandi Sauvage. Oshima (19 19) established the genus Aristichthys to 
contain nobilis, believing that differences in gill-raker form, abdominal keel length and 
pharyngeal dentition from those in molitrix were large enough to signify generic rank. 
Authors have been at variance in treating Aristichthys as a valid genus and many still regard 
nobilis as a species of Hypophthalmichthys. I would agree with the latter view. The taxa 
molitrix and nobilis possess unique synapomorphies and thus belong to the same higher 
category, ie. the genus Hypophthalmichthys. The species H. harmandi appears to have been 
overlooked by subsequent authors. To my knowledge it is known only from the type 
specimen from Hanoi, now in the Paris Museum. 

Giinther (1868) first introduced the category Hypophthalmichthina to contain Hypo- 
phthalmichthys which taxon Gill (1893) later recognized as a subfamily, the Hypophthal- 
michthinae. Regan (1911; 1922) thought Hypophthalmichthys to be related to Barilius a 
suggestion refuted by Howes (1980). Kryzanovskij (1947) referred Hypophthalmichthys to 
the Leuciscinae on evidence derived from ontogenetic development. Ryabov (1973) agreed 
with this subfamilial placement, using as his evidence the viability of larvae produced by 
cross-breeding Hypophthalmichthys and Abramis Abramis being considered a typical 
leuciscine. Gosline (1978), relying on a tri-lobed swimbladder as an indicator of this 
relationship, considered that there were 'clear indications of a cultrin derivation for 
Hypophthalmichthys and Aristichthys'. A tri-lobed swimbladder, however, has such a 
mosaic distribution throughout the Cyprinidae that it has no value as a character indicating 
shared common ancestry. Besides which, the third 'lobe' in Hypophthalmichthys is a mere 
constriction or 'tail' of the swimbladder and is variously developed in individual fish. 

There has been only one study of the comparative osteology of the major Chinese Carps, 
that by Watanabe (1951). Although this author pointed out significant differences between 
Ctenopharyngodon, Hypophthalmichthys and Mylopharyngodon, and noted the possible 
taxonomic value of certain characters, he made no attempt to utilize them in formulating 

Finally, it can be noted that artificial hybridization between Ctenopharyngodon and 
Hypophthalmichthys has been achieved and the morphology of the hybrids described by 
Berry & Low (1970). 

The consensus of ichthyological opinion it would seem, is that the two major carps, 
Ctenopharyngodon and Hypophthalmichthys belong to the same subfamily, the Leuciscinae. 
My thesis is that they do not, but that each represents a distinct monophyletic lineage, 
representative of a basic dichotomus division within the Cyprinidae. 

Abbreviations used in the figures 

AA Anguloarticular Aap Adductor arcus palantini muscle 

AEW Anterior wall of epibranchial organ Ah Adductor hyomandibularu ; muscle 


AMP Anterior medial maxillary process LEN 

BO Basioccipital LOF 

BOCB Basioccipital cartilaginous bloc ME 

BOK Basioccipital keel MET 

BON Basioccipital notch MF 

BP Basioccipital pharyngeal process MMP 

BUG Buccal cavity MP 

CB Ceratobranchial (numbered) N 

CF Ceratobranchial filaments NC 

CFE Carotid fenestra NS 

CP Coronoid process OC 

CPE Cartilaginous preethmoid OFT 

Dahm Dorsal section of adductor OP 

hyomandibularis OPE 

DPTC Dermopterotic canal PALH 

EB Epibranchial (numbered) PAR 

EBD Epibranchial duct (numbered) PE 

EBO Epibranchial organ PEW 

EC Ethmoid cartilage PF 

ECT Epibranchial connective tissue PFO 

EOD Epibranchial oesaphagal duct PH 

EOW Epioccipital wing PHF 

EP Epioccipital Php 

EPOP Epioccipital process PO 

ER Epibranchial rakers POC 

ES Extrascapular PPSF 

EXO Exoccipital PRO 

EXOF Exoccipital flange PS 

F Frontal PSAP 

FC Frontal canal PSCB 

FF Frontal foramen PTE 

FIC Internal carotid foramen PTES 

FJ Jugular foramen PTS 

FII Optic foramen PTT 

FV Foramen for trigeminal trunk PVP 

FVII Foramen for facial nerve trunk Q 

FVIII Foramen for acoustic nerve SE 

FIX Foramen for glossopharyngeal SEW 

GR Gill raker (globular) SM 

HMF Hyomandibular fossa SO 

HYO Hyomandibular SOCC 

HVII Hyomandibular branch of VII nerve SOP 

1C Intercalar SOR 

ICF Intercalar fossa SP 

IF Infrapharyngobranchials (numbered) V 

IO Infraorbitals (numbered) Vahm 

IOP Interoperculum 

LE Lateral ethmoid VPSR 

LEF Lateral ethmoid/frontal fenestra 

Supraethmoid notch for palatine head 

Lateral occipital foramen 



Maxillary foramen 

Midlateral maxillary process 

Masticatory plate 


Neural complex 

Neural spine 

Intramural opercular canal 

Olfactory foramen 


Ossified preethmoid 

Palatine head 



Posterior wall of epibranchial organ 

Palatal fold 

Prootic foramen 

Pharyngeal bone 

Fossa in pharyngeal bone 

Pharyngeal pad 


Preopercular canal 

Prootic/paraspenoid fenestra 



Parasphenoid ascending process 

Parasphenoid keel 


Pterotic spine 



Pterosphenoid ventral process 



Supraethmoid wing 

Ceratobranchial sifting membrane 


Supraoccipital crest 





Ventral section of adductor 

hyomandibularis muscle 

Ventral parasphenoid ridge 

Study material 

All specimens used in the preparation of this paper are in the osteological and alcohol 
collections of the British Museum (Natural History). Lists of comparative material will be 

4 G. J. HOWES 

found in previous publications (Howes, 1978; 1979). The catalogue numbers for specimens 
used in illustrating this work are given in the figure captions. 

Comparison of selected characters in Ctenopharyngodon and Hypophthalmichthys 

The following 22 characters were selected after a preliminary study had eliminated others 
which appeared to contain minimal information concerning their plesio- or apomorphic 
state (ie. intra- and interspecifically variable or mosaically distributed), and those which 
were considered to be symplesiomorphies. Guidance to the identification of plesiomorphic 
character state came both from previous studies on cyprinid phylogeny (Howes, 1978; 1979; 
1980) and wider outgroup comparisons outgroup in this case being other otophysans. 
Likewise, these earlier studies had led to the final selection of those characters which had 
previously been useful as phylogenetic indicators. Reasons for polarity assignment are 
present in the analysis below. 

Selected characters: 1. Ethmo-vomerine region; 2. Pterosphenoid; 3. Parasphenoid; 4. 
Prootic; 5. Subtemporal fossa and associated musculature; 6. Dilatator fossa; 7. Frontal; 8. 
Parietal; 9. Circumorbitals; 10. Supraoccipital; 11. Exoccipital; 12. Epioccipital; 13. 
Intercalar; 14. Basioccipital; 15. Posttemporal; 16. Opercular series; 17. Suspensorium; 18. 
Jaws; 19. Vertebral column; 20. Gill-arches; 21. Adductor hyomandibulae muscle; 22. 
Pectoral girdle. 

Analysis of apomorph characters 

1 . Ethmo- vomerine region 

Ctenopharyngodon (Fig. 1); supraethmoid broad and short with a shallow medial anterior 
notch and two lateral notches; preethmoids paired, largely cartilaginous; kinethmoid 
dorsally expanded; vomer widely V-shaped anteriorly; lateral ethmoid wall truncate. 

Hypophthalmichthys (Fig. 1); supraethmoid broad and short with a deep medial notch; 
preethmoid single, small, completely ossified; kinethmoid rod-shaped; vomer thickened 
anteriorly and forming the floor of the median ethmoid notch; lateral ethmoid contains 
entire olfactory foramen in a bony tube; the lateral lamella is prolonged distally and curves 
ventrally; fenestra between the dorsal surface of the lateral wall and the frontal. 

Both Ctenopharyngodon and Hypophthalmichthys possess a short, broad supraethmoid 
and depressed mesethmoid. This is hypothesized as the plesiomorph condition in cyprinids 
on the grounds of widespread occurrence. Within the Cyprinidae there are two basic forms of 
supraethmoid; (1) in which the anterior border of the bone is indented medially so as to 
accommodate the retracted kinethmoid (sometimes there may be indentations lateral to the 
medial one), and (2) where the supraethmoid is rostrally extended, its lateral border rounded 
and the tip of the extension bearing the median notch. These two types are illustrated in 
Fig. 2. 

The first morphotype is that most commonly encountered in non-barbelled Cyprinidae 
(ie. Leuciscinae), whilst the second is that found in almost all barbelled Cyprinidae (ie. 
Cyprininae; Barbus, Labeo etc). In Cyprinus, Carassius and several members of the 
Gobioninae, although there is a prominent rostral process there is no, or but a slight, medial 
notch the kinethmoid being attached to the base of the rostral process by a single ligament 
(oberes Russelband of Fiebiger, 1931). This is similar to the condition in the Catostomidae 
where the supraethmoid is elongate and without a medial indentation. 

A rostrally produced supraethmoid is the common condition amongst characoids and a 
comparison between the ethmoid region of Brycon and that of Catostomus shows a 
remarkable similarity. In siluroids the common condition, as in cyprinoids, appears to be 
one where the anterior ethmoid border bears a slight medial indentation. Within the 
cyprinids, although the extremes of the two ethmoid morphotypes can be readily 
appreciated, they are linked by a series of intergrades. Particular examples are in Garra 







Fig. 1 Ethmo-vomerine regions in dorsal view: (upper) Ctenopharvngodon idellus; (lower) 
Hypophthalmichthys molitrix. Specimens BMNH 1888.5.15:25; 1895.5.31:22. 
Scales = 10 mm. 

where the rostrally produced ethmoid shape is still discernible but the anterior border is 
deeply indented as in 'leuciscines'. Again, in some species of Barbus and Schizothorax the 
ethmoid is distinctly leuciscine-like. An extreme morphotype is encountered in Semiplotus 
and Scaphiodonichthys where the entire supraethmoid is almost vertically directed and it is 
the anterior border of the frontal which is medially indented. In general, it appears that 
cyprinids with a terminally placed mouth are those with a deeply notched ethmoid. On the 
other hand, the rostrally produced ethmoid is more characteristic of bottom feeding types 
where additional, almost ventral extension of the mouth is functionally desirable as in 
Semiplotus mentioned above. Fink & Fink (198 1) propose that a rostrally produced ethmoid 
is plesiomorphic for otophysans. It would appear that in cyprinids a supraethmoid with a 
slight, but broad rostral process is the plesiomorph state. This plan would offer the potential 
of producing either kind of morphotype. The variability of the supraethmoid in species of 
Barbus is discussed by Banister (in preparation). In that group, at least, the ethmoid appears 
to be a very labile structure and one responding to environmental influences. 

In Ctenopharyngodon there are lateral indentations on either side of the medial 
supraethmoid notch (LEN, Fig. 1). These lateral notches accommodate the enlarged anterior 
condyles of the palatines. A similar feature occurs in Squaliobarbus and Mylopharyngodon 
and is considered synapomorphic. 





Fig. 2. Ethmo-vomerine regions in dorsal view: (upper) Cyprinus carpio; (lower) Leuciscus 
cephalus. Composite drawing of several specimens. Scale = 5 mm. 

Concerning the posterior border of the supraethmoid, a transverse suture with the frontal 
appears to be the plesiomorph state. Only in some chelines is the posterior margin rounded 
or triangular and overlapped by the frontals (see Ramaswami, 19556, fig. 3, and Howes, 
1979). Amongst characoids the posterior ethmoid margin is triangular, that of siluroids is 
irregular, its medial part usually invaded by the anterior tip of the frontal fontanel. 

A rod-shaped kinethmoid is considered the plesiomorph type (see Howes, 1978). I am 
unaware of any detailed study concerning the possible derivation or homology of the bone. 
Goodrich (1909) thought it to be a specialized part of the ethmoid. The common occurrence 
of a bifurcated kinethmoid suggests that there has been a phylogenetic ossification into a 
single unit of two premaxillary-ethmoid ligaments. There is no cyprinoid known to me in 
which these ligaments remain unossified in the adult, or even in larval stages, and which 
would represent the most plesiomorph state. In its various derived states the dorsal part of 
the kinethmoid may be expanded in a transverse plane, or the medial part of the bone may be 

In Ctenopharyngodon the kinethmoid is dorsally expanded and thickened, a derived 
condition encountered in other cyprinid groups (eg. chelines; see Howes, 1979). In Hypo- 
phthalmichthys the bone remains as a slender rod. 

Ramaswami (1955a) noted the absence of a preethmoid in Hypophthalmichthys, but this is 
not the case. A preethmoid cartilage is present, although minute and only fully ossified in the 
adult. Likewise, in other genera which Ramaswami (1955a & b) cites as lacking a 
preethmoid, I have found it as a poorly ossified element. Ossification of the preethmoid is 
variable; specimens ofAlburnus show no sign of ossification until they are at least 56 mm SL; 



Fig. 3 Development of the ethmoid bloc in Barbus sp. Anterior view (top) from a specimen 
13mm SL; (centre) specimen 20mm SL; (bottom) specimen 35mm SL. Dashed areas = 
cartilage, stippled areas = ossification. Scales = - 5 mm. 

the preethmoid in Oreoleuciscus is entirely cartilaginous in a specimen of 1 14 mm SL. It is 
still recognizable as a preethmoid, being clearly differentiated from the ossified meseth- 
moidal process. 

According to Alexander (1966) the preethmoid is an ossification of the antero-lateral 
process of the ethmoid cartilage (mesethmoid). He notes that in Leuciscus (=Idus of 
Alexander), the maxillary ascending process articulates with the preethmoid cartilage, and 
with the palatine via a cartilaginous insert (submaxillary meniscus of Alexander). Corrobora- 
tion of Alexander's (1966) statement that the preethmoid is an ossification of the 
mesethmoid comes from an ontogenetic series of Barbus sp. (Fig. 3). In the smallest 
specimens, 12-14 mm SL, the preethmoid is not differentiated from the ethmoid cartilage 
and there is a sub-terminal ossification, in 20 mm specimens there is dorsal ossification of 
the lateral mesethmoid processes, greater area of terminal ossification and antero-lateral 
extension of the vomer. In specimens of between 25 mm and 27 mm SL there exists only a 
small area of cartilage between the mesethmoid, ossified preethmoid and vomer. At the 
30-35 mm stage, the antero-lateral mesethmoid processes have contacted the now large and 
completely ossified preethmoids and the vomer. A lateral mesethmoid-vomerine fossa is 
formed and the preethmoid contacts the fossa via a thin meniscus of cartilage. 

In all cyprinids I have examined, other than abramines (see below), the preethmoid is 
contained in a lateral fossa formed from the mesethmoid and the vomer (see Howes, 1 978). 

A lateral ethmo-vomerine fossa is lacking in Abramis, Rutilus, Chondrostoma and 
Xenocypris. In these taxa the vomer does not protrude laterally (see below) but extends 
forward; the preethmoid is extensive and covers the extended vomerine arm, contacting the 
mesethmoid only along its antero-ventral rim (see Figs 5A-D). The preethmoid is almost 
entirely cartilaginous with only the anterior tip being ossified in the case of Abramis, Rutilus 






Fig. 4 Hypophthalmichthys molitrix, lateral view of ethmo-vomerine region. Specimen 

1895.5.31 : 22. Scale =10 mm. 

and Chondrostoma, but a laterally ossified nub in Xenocypris. Although in Hypo- 
phthalmichthys there is a distinct reduction of the lateral fossa the preethmoid cartilage still 
contacts the lateral part of the mesethmoid (Fig. 4). Laterally, the preethmoid articulates 
directly with the palatine facet; only rarely is such a connection lacking (eg. Ochetobius); 
there is never direct contact with the maxilla. In some cyprinids, however, the preethmoid is 
extended anteriorly and may ossify into two parts. Such paired preethmoids were recorded in 
Cyprinus by Fiebiger (1931) as the lateral 'seitliches Knotchen' articulating with the 
palatine, and the anterior 'Sesamknoten' articulating with the maxilla. Paired preethmoids 
also occur in Ctenopharyngodon (Fig. 1), Mylopharyngodon, Squaliobarbus and some 
Barbus species. In these taxa, the anterior preethmoid is often unossified or only partially so. 
According to Ramaswami (1955a) it is the cartilage sandwiched between the maxilla and the 
preethmoid which ossifies to form a second preethmoid. However, in the taxa cited above, 
the additional, anterior ossification would appear to be within the original preethmoidal 

In gobioinids, cobitids and catostomids, the second preethmoid can be exceptionally well- 
developed and exists as an ossified strut (the septomaxilla of Regan, 1911 and Starks, 1926). 
Patterson (1975 : 501) reckons the preethmoids of cyprinoids to be neomorphs. As such 
they must be regarded as autapomorphic for Cyprinoidei. A single, lateral ossification of the 
ethmoid cartilage articulating with the palatine is hypothesized as representing the 
plesiomorph condition (see Howes, 1980). 

The mesethmoid is usually a triangular bloc, its apex forming the rostrum of the entire 
ethmoid bloc. In those taxa with a medially indented supraethmoid, the mesethmoid 
becomes thickened anteriorly and forms a somewhat hour-glass shape. In some leuciscines, 
the mesethmoid is laterally excavated to such an extent that it is virtually a medial septum. 
The posterior wall of the mesethmoid cartilage is, in one group of leuciscines and some 
cyprinines, excavated to form the anterior myodome (see below). 

The lateral ethmoid in cyprinids most often has a well-produced lateral wall with a 
broadly-triangular base; antero-medially it meets the mesethmoid and contains part of the 
olfactory nerve foramen in its leading edge; postero-medially it meets its fellow along the 
midline. In some cases the lateral ethmoids are indented into the mesethmoid cartilage so as 
to form an anterior myodome. Even though, as in aspinines, the anterior myodome extends 



Fig. 5 Ethmo-vomerine regions in dorsal and lateral views: A. Abramis brama; B. Xenocypris 
argenteus. The lateral views are semi-diagrammatic and show the anterior edge of the vomer as a 
dashed line, the cartilaginous preethmoid as cross-hatched and the ossified preethmoid as black. 
Composites, from skeletal and alizarin specimens; all drawn to same scale. 

well forward, the lateral ethmoids remain in medial contact. However, in Barbus barbus, 
Cyprinus and Carassius, the myodome invades the mesethmoid cartilage through a parting 
between the lateral ethmoids. Patterson (1975) was unable to conclude whether the absence 
of an anterior myodome was a primitive condition or due to secondary loss (apomorphic). Its 
presence does not seem necessarily to be correlated with any particular degree of ethmoidal 
elongation, nor its absence with any shortening. In noemacheilids, gobioinids and 
catostomids, an anterior myodome is lacking. Widespread, mosaic distribution in cyprinids 
and almost universal presence in characoids probably indicates that an anterior myodome is 
a plesiomorph feature. 

In Ctenopharyngodon and Mylopharyngodon the olfactory foramen is entirely counter- 
sunk within the lateral ethmoid, whereas in Hypophthalmichthys it projects forward as a 
bony tube (Fig. 4). In the latter genus there is a tubular opening between the medial dorsal 
part of the lateral ethmoid wall and the overlying frontal (LEF, Fig. 4). This tube contains a 
thick ligament which covers the top of the lateral ethmoid wall and the outer part of the 
supraorbital. A similar feature occurs in Xenocypris (Fig. 5B). In cyprinids the olfactory 
foramen is most often formed from the anterior margin of the mesethmoid. In some taxa, 
however, the foramen is confined entirely within the lateral ethmoid. This occurs in the taxa 
cited above, in alburnine genera (Alburnus, Chalcalburnus) and some Barbus. In Barbus 
tropidolepis the foramen is also housed in an anteriorly projecting tube (Banister, pers. 
comm.). On the basis of other sets of synapomorphic characters it is hypothesized that the 
occurrence of a tubular olfactory foramen in diverse taxa is of no phyletic significance. 

A short and broad vomer is hypothesized as plesiomorphic eg. as in Ctenopharyngodon 
and Hypophthalmichthys. However, in Hypophthalmichthys the vomer extends anteriorly to 
floor the median ethmoid notch, its anterior margin is almost straight and is curved upward 
so as to face the lower part of the ethmoid bloc. Laterally, the vomer is indented to contain 
the preethmoid. An anteriorly extended vomer, flooring the median ethmoid notch is a 
rarely encountered feature amongst cyprinids. In some cultrines (Culler) and in Ochetobius 
(currently included in the Leuciscinae), the vomer forms the floor of a shallow but wide 


Fll PVP 









Fig. 6 

Ctenopharyngodon idellus, ventral view of anterior otic region. Specimen BMNH 

ethmoid notch. In Xenocypris and Abramis, the vomer also floors the ethmoid notch but the 
mesethmoidal walls are deep, almost meeting one another ventrally and leaving only a 
narrow area of the vomerine floor visible from above (Figs 5A & B). As mentioned above 
(p. 7) Xenocypris, Abramis, Chondrostoma and Rutilus differ from other cyprinids in 
employing the vomer as the platform for the preethmoid and excluding the mesethmoid 
from taking part in its support. In these genera the vomer is deeply forked, each arm 
extending some distance anteriorly from the mesethmoidal border; in transverse plane the 
bone is markedly convex. 

2. Pterosphenoid 

Ctenopharyngodon (Fig. 6); contact between the pterosphenoid and ascending process of the 
parasphenoid is via a thin strut; medial contact with the prootic is by a long suture. On the 
lateral face of the pterosphenoid is a small, ventrally directed process (autapomorphic 
feature; PVP, Fig. 6). 

Hypophthalmichthys (Fig. 7); contact with the parasphenoid is via a thick, anteriorly 
directed pillar-like process. The part to the anterior trigemino-facialis foramen, and a medial 
portion of the bone provides the inner wall of the anterior part of the chamber; this is a 
unique condition in cyprinids. 

Howes (1979) presented a classification of the various types of contact between the 
parasphenoid and pterosphenoid, assuming that this classification reflected a phylogenetic 
sequence for the Cyprinidae as a whole. Further analysis has shown that similar sequential 
series can be detected in all groups so far identified as monophyletic, ie. the plesiomorphic 
condition is one with no contact between para- and pterosphenoid (Type 1 and derived Type 
1 in Howes, 1980) and the most apomorphic condition involves extensive lateral contact 
between the pterosphenoid and parasphenoid (Type 5 in Howes, 1980). 

A pterosphenoid pedicle is present in various teleosts and may or may not contact the 







Fig. 7 Hypophthalmichthys molitrix, lateral aspect of anterior otic region, viewed slightly antero- 
ventrally. Drawn from specimen BMNH 1895.5.31 : 22 with additions from alizarin prepar- 
ations and dissections from BMNH 1980.5.21 : 5-14. Scale = 10mm. 

parasphenoid ascending process. Patterson (1975) regards this feature as a plesiomorphic 

3. Parasphenoid 

In both Ctenopharyngodon and Hypophthalmichthys the parasphenoid is anteriorly broad, 
narrowing posteriorly in Hypophthalmichthys but remaining broad in Ctenopharyngodon in 
which genus it bears a marked ventral medial range. In Hypophthalmichthys the 
hypophyseal portion of the parasphenoid is ventrally extended into a long deep process, or 
keel the posterior edge of which is developed into a transverse ridge. The ventral keel serves 
both as the anterior point of attachment for a medial cartilaginous insert dividing the forward 
part of epibranchial diverticula (p. 23) and as lateral sites of attachment for the adductor 
arcus palatini muscle (p. 37). 

In the majority of cyprinids the central surface of the parasphenoid is grooved, rarely is 
there a midline ridge or process, and these features are regarded as derived. The anterior 
broadening of the parasphenoid, although extreme in the Chinese major carps, is a common 
feature in cyprinids (see Ramaswami, 19556). Weisel (1960) notes that in Catostomus the 
anterior part of the parasphenoid is considerably wider than that of cyprinids. However, this 
is not so when compared with the two taxa discussed here. 

Ctenopharyngodon shares with Mylopharyngodon the developed ventral midline ridge 
and deep anterior concavity of the parasphenoid, both features regarded as apomorphic. The 
hypophyseal keel of Hypophthalmichthys is a feature shared with Xenocypris and Abramis, 
although in these taxa it is developed to a lesser extent (see below). 

The internal carotid fenestra in Ctenopharyngodon is small and indents the posterior 
border of the parasphenoid ascending process and the antero- ventral margin of the prootic. 
In Hypopohthalmichthys the internal carotid artery passes through a small foramen situated 
in the basal part of the parasphenoid keel (Fig. 7). 

Patterson (1975) considered that the plesiomorph condition of the carotid opening in 


teleosts is for it to occur as a foramen within the parasphenoid. In nearly all cyprinids I have 
examined, and in all characoids, the internal carotid passes through a fenestra between the 
parasphenoid and the prootic, a condition Patterson (1975) thought derived and due to the 
eroding of the parasphenoid ascending process. In catostomids (see Weisel, 1960) and 
siluroids (?all) the foramen is situated entirely within the parasphenoid. Although similarly 
there is a parasphenoid carotid foramen in Hypophthalmichthys I consider this to be a 
derived condition in this taxon and for the following reasons. 

In Alburnus, Rutilus, Chondrostoma and Abramis, the internal carotid fenestra is 
extensive (Fig. 8); it is covered laterally by a fascia of tissue and plugged by a wedge of fat 
leaving a small ventral opening to allow passage of the carotid vessel. In Abramis there is a 
tendency for the fenestra to elongate and in Abramis sapa the 'fenestra' is divided; the lower 
opening being confined to the parasphenoid and carrying the internal carotid artery, the 
upper indenting the ventral margin of the prootic and not providing passage for any vessel 
(Fig. 8). It is hypothesized that the condition of the carotid fenestra in Abramis sapa is a 
derived one and represents a transitional stage toward that in Hypophthalmichthys. That this 
is the directional path and not the reverse, viz the incorporation of the foramen into a 
fenestra, as indicated by the morphology of the parasphenoid. In Abramis there is a ventral 
extension of the hypophyseal part of the parasphenoid similar to that in Hypophthal- 
michthys (see above), and this is most highly developed in A. sapa (Fig. 8). Furthermore, the 
carotid foramen in Hypophthalmichthys is situated in the ventral keel, rather than 
(plesiomorphically) in the ascending process (Patterson, 1975). Similarly, an internal carotid 
foramen occurs in the same position in Xenocypris but is a larger hole than in Hypo- 

On the basis of this evidence I suggest that the foramen for the internal carotid artery in 
Hypophthalmichthys and Xenocypris is secondarily derived in its position in the 
parasphenoid. This positioning has come about both as a consequence of the ventral 
prolongation of the parasphenoid and the re-orientation of the first branchial arch (p. 31) 
whereby the plane of the efferent artery would be sagittal rather than transverse. Thus, this 
shared feature in Hypophthalmichthys and Xenocypris is considered a synapomorphic 

Modifications to the ventral border of the parasphenoid are present in a unit of leuciscine 
taxa represented by Abramis, Rutilus and Chondrostoma. In these genera, there is a deep 
ventral channel in the posterior part of the parasphenoid. The channel is confluent with a 
similar feature in the basioccipital and the whole is filled with a cartilaginous block (Fig. 27). 
Discussion of this character is reserved for inclusion with that of the basioccipital (p. 23). 

Mention should be made here of medial parasphenoid processes. In Labeo and related 
taxa (see Howes, 1980 : 152, fig. 26 and p. 16 below) a process arises from either side of the 
medial wall of the parasphenoid just anterior to the optic foramen. Each process contacts the 
ventral border of each respective pterosphenoid. These medial processes vary from thin 
struts to thick pillars. Only in Pelecus is there a single, medial parasphenoid pillar. These 
structures are not to be interpreted as homologues of the basisphenoid. Harrington (1955) 
reported that Holmgren & Stensio (1936) recognized a basisphenoid in some cyprinids, eg. 
Abramis. However, the structure reported by Holmgren & Stensio is the lateral 
pterosphenoid strut that makes contact with the parasphenoid wing. The medial 
parasphenoid extensions of Pelecus and Labeo are considered as neomorphs. 

4. Prootic 

Ctenopharyngodon (Fig. 6); contact with the parasphenoid ascending process is via a long 
suture. The posterior opening of the trigemino-facialis chamber is large and lies ventrally. 
The upper posterior part of the bone is deeply concave and forms the entire inner face and 
part of the roof of the subtemporal fossa. 

Hypophthalmichthys (Fig. 7); no contact with the parasphenoid ascending process. The 
posterior trigemino-facialis foramen is small and situated laterally. The upper posterior half 






Fig. 8 Lateral views of anterior otic regions: (top) Rutilus rutilus; (centre) Abramis brama; 
(bottom) Abramis sapa. Semi-diagrammatic, compiled from several specimens. 

of the bone is slightly concave and forms only the lower inner face of the subtemporal fossa. 
The lower part of the anterior border forms the posterior rim of the trigeminal exit fenestra. 

The plesiomorphic condition of the prootic in cyprinids is hypothesized as an almost 
square bone, its lateral face hardly depressed, its anterior border interrupted by the 
trigeminal exit foramen and with a short lateral commissure (see for example, Opsariichthys 
in Howes, 1980). Derived conditions are those where the anterior trigeminal foramen pierces 
the lateral face; the lateral commissure is long, and the prootic contributes a ventral portion 
to the apophyseal platform. 

The general depression of the otic part of the cranium in Ctenopharyngodon has resulted 
in the forepart of the prootic being laterally extended (Fig. 6), which, as a consequence, has 
shifted the orientation of the posterior trigeminal foramen. This opening faces ventrally 




Fig. 9 Subtemporal fossae: (left) Hvpophthalmichthvs; (right) Ctenopharyngodon. Specimens 
BMNH 1888.5.31 : 22 and 1888.5.15 : 25. Scales = 10 mm. 

instead of laterally, so the nerve trunk and jugular vein enter without passing across the 
prootic face. This derived condition occurs also in Mylopharyngodon and Squaliobarbus. 

The most outstanding feature of the prootic in Hypophthalmichthys is the exclusion from 
the bone of the anterior trigeminal foramen. Only the lower part of the prootic forms the 
border of the opening, the upper part of the foramen being confined to the pterosphenoid. Jn 
one specimen, the upper part of the foramen, through which mdV and mxV exit is, on one 
side of the cranium, entirely enclosed in the pterosphenoid. Reduction of the anterior part of 
the prootic in Hypophthalmichthys appears to have occurred with the posterior movement of 
the pterosphenoid, itself associated with hypertrophy of the adductor hyomandibularis 
muscle. The form of the prootic in Hypophthalmichthys is an autapomorphic feature. 

5. Subtemporal fossa 

Ctenopharyngodon (Fig. 9); extensive, hemiovoid with a deep conical anterior portion, or 
chamber, that extends into the autosphenotic; the levator posterior muscle arising from that 
bone. The inner wall of the fossa is formed from the prootic, the roof from the pterotic and 
epioccipital, and anteriorly from the sphenotic. None of these bones meet dorsally and the 
ventral surface of the parietal is exposed thus also contributing to the Subtemporal fossa roof. 

Hypophthalmichthys (Fig. 9); almost pyramidal, lacking an anterior sphenotic extension. 
The posterior, exoccipital border of the fossa is indented by a notch. The bones forming the 
inner wall of the fossa, prootic, exoccipital and epioccipital, do not meet along their respec- 
tive dorsal and anterior borders, the space between being a thin cartilage wall. 

The most frequently occurring Subtemporal fossa morphology amongst the Cyprinidae is 
one which is deep and circular or ovate. The fossa is the site of origin for levator posterior 
muscles running to the pharyngeal bone and its size is correlated with that of the muscles' 
size and their orientation. It is thus difficult to assess the polarity of some features such as the 
posterior notch, which occurs not only in Hypophthalmichthys but in a variety of Meuciscine' 
and 'barbine' taxa. There is generally a space between the bones forming the inner wall of the 
Subtemporal fossa (viz between the posterior-dorsal edge of the prootic, posterior edge of the 
sphenotic, and antero-dorsal edge of the epioccipital) across which is a thin cartilage and 
which forms the apex of the fossa. In this feature Ctenopharyngodon and Hypophthal- 
michthys are each autapomorphic. In the former taxon the space is covered by the parietal 


and in the latter taxon the exoccipital contributes substantially to the inner face and its 
antero-dorsal edge contacts the cartilaginous sheet. 

The vast subtemporal fossa of Ctenopharyngodon is a feature shared only with 
Mylopharyngodon and Squaliobarbus. Likewise, in these two genera the fossa extends 
anteriorly deep into the sphenotic from which the levator posterior musculature originates. 
The size, shape and sphenotic contribution of the subtemporal fossa in these three genera is 
considered synapomorphic. The anterior contribution of the sphenotic to the subtemporal 
fossa occurs also to a greater or lesser degree in Cyprinus, Carassius and in some Barbus, and 
in the two former taxa part of the levator posterior musculature originates in that anterior 
extension. This may thus represent a synapomorphy linking all these genera (see p. 49). 

The shape and depth of the Hypophthalmichthys subtemporal fossa are shared with 
Xenocypris. However, these parameters are treated with caution as synapomorphies; 
variations in subtemporal fossa depth cover a wide range throughout the cyprinids. Abramis 
has what is possibly the deepest subtemporal fossa and again, its pyramidal shape greatly 
resembles that of Hypophthalmichthys. Neither in Xenocypris nor Abramis does the 
exoccipital contribute substantially to the inner wall of the subtemporal fossa. Only in 
Rutilus does the anterior edge of the exoccipital penetrate forward so as to separate the 
prootic and epioccipital. 

6. Dilatator fossa 

Ctenopharyngodon (Fig. 10); formed mostly from the sphenotic which extends laterally with 

a small anterior contribution from the frontal; the frontal border forms a medial roof to the 


Hypophthalmichthys (Fig. 10); formed in almost equal parts from the frontal and 
sphenotic and sloped antero-laterally as a broad shelf. 

The plesiomorph condition of the dilatator fossa in cyprinids is considered to be one in 
which the fossa is small and formed from nearly equal portions of the frontal, sphenotic and 
pterotic, and to be roofed or partially roofed by the frontal. This type of fossa morphology is 
widespread amongst cyprinids and is represented in Ctenopharyngodon. An almost identical 
type of dilatator fossa morphology in characoids is regarded by Vari (1979) as plesiomorphic. 
In siluroids there is no cranial fossa, the dilatator operculi muscle originating from the 
ventral cranial surface (see Alexander, 1964). Similar ventral muscle origin is to be found in 
some cyprinids and characoids (see below; Alexander, 1964; Howes, 1978). In most 
catostomids the dilatator fossa is well developed and involves the sphenotic and the medial 
part of the frontal. Cobitids (sensu lato) possess a small fossa of the type which involves 
mostly the sphenotic. 

It would seem that from the plesiomorphic type of dilatator fossa two avenues of develop- 
ment were open. One was toward extended coverage of the cranial roof, involving mostly the 
frontal. The second was for the frontal to become invaginated ventro-medially and be 
pierced by the dilatator operculi. The most derived state of the first morphotype in cyprinids 
is that in Hypophthalmichthys where the major contributing elements to the fossa are the 
frontal and sphenotic, the pterotic being virtually excluded. The sphenotic extends laterally 
as a broad, ventrally sloping shelf confluent with the frontal. The lateral part of the frontal 
also slopes and medially bears a ridge which forms the border of the fossa. A similar form of 
dilatator fossa occurs in Xenocypris and Abramis, but in these genera the frontal and 
sphenotic have a more acute ventral slope. 

The most derived state of the second morphotype is that where a divided dilatator operculi 
muscle passes through a frontal foramen and a fenestra between the frontal and sphenotic 
(see below). 

These two morphotypes embrace the four enumerated by Howes (1978). Type 1 is 
described above as the plesiomorph condition and Types 2 and 3 are morphoclinal stages in 
the encroachment of the cranial surface. 

Within the second lineage (Type 4 of Howes, 1978), in which the dilatator operculi 



Fig. 10 Dilatator 

fossae: (upper) Ctenopharyngodon; (lower) 
Scales = 5 mm. Specimens as previous figure. 


originates from the ventral surface of the frontal, a possible transitional sequence can be 
distinguished. An early stage is hypothesized as being represented by Squaliobarbus. Here 
the frontal is deeply indented anteriorly to the sphenotic and contains a small foramen which 
enters the orbital cavity. In Barbus and Capoeta the muscle has 'broken through' the frontal 
to attach to its ventral surface inside the orbital cavity. A previous statement (Howes, 1978) 
to the effect that the frontal and sphenotic in Barbus contacted each other only along their 
lateral margins is incorrect. The anterior margin of the sphenotic is bordered by the frontal, 
albeit in some instances an extremely thin segment (Fig. 1 1). Carassius and Cyprinus appear 
to mark a further stage in development of the foramen for there is in some specimens an 
almost complete separation between the frontal and sphenotic, the ventro-lateral part of the 
frontal extending only part way along the sphenotic margin. Cyprinus is the more derived 
taxon in this respect since it has an almost complete frontal-sphenotic foramen (Fig. 1 1). 
Labeo and Garra appear to represent the most derived state, with the dilatator operculi being 
divided, the upper section passing through a frontal foramen, the lower through a foramen 
between the frontal and a strut-like sphenotic process (Fig. 1 1 ). The doubly foraminate forms 





Fig. 11 Dilatator fossae: (upper) Cvprinus carpio (uncatalogued); (centre) Capoeta capoeta 
BMNH 1 879. 11.14:19; (lower) Labeo coubie BMNH 1 907. 1 2.2 : 3744. 

of dilatator channel appear to be rewarding characters for the classification of those taxa in 
which they occur and work in preparation discusses the feature in more detail (Banister & 
Howes, in preparation). 

7. Frontal 

Ctenopharyngodon (Fig. 12); broad and short, forming 50% of the cranial length (anterior tip 
of the ethmoid to the posterior border of the parietal), 5-6 pores in the embedded sensory 
canal, lateral part sloping ventrally at 27 to the transverse plane, a triangular depression 
occurs above the supraorbital. 

Hypophthalmichthys (Fig. 12); broad anteriorly, narrow posteriorly, forming 75% of the 
cranial length; anterior lateral part sloped ventrally at 45; sensory canal is an osseus tube on 
the surface, with 9-10 pores. 

In cyprinids, the plesiomorph condition of the frontal is considered to be short and broad 
(Howes, 1978 : 31). Only in derived members of monophyletic lineages are the frontals 
elongate, with an increased number of sensory pores. The raised tubular sensory canal, as in 
Hypophthalmichthys, is a feature Tretiakov (1946) used in marking Tinea as a primitive 
cyprinid related to catostomids which also possess this character. However, in catostomids 










Fig. 12 Dorsocrania: (left) Ctenopharyngodon; (right) Hypophthalmichthys. Scales = 20 mm. 

Specimens as previous figures. 

the situation is rather different for here the canal is mostly detached from the frontal and runs 
at the surface of thick subcutaneous tissue covering the cranium (see comments by Sagemahl, 
1891 : 508 and Weisel, 1960 : 122). Although raised frontal canals are an uncommon feature 
in Cyprinidae they occur in abramine genera (see p. 45 for included genera), some cultrines 
and some chelines (eg. Oxygaster). This pattern of ossification has possibly been derived 
independently in these several groups. 

The frontal morphology in Ctenopharyngodon is generally of the plesiomorphic type but 
sloped and laterally depressed, extended border of the bone is a derived state and shared with 
Mylopharyngodon. The markedly sloped frontal in Hypophthalmichthys is shared only with 
Xenocypris and Abramis. 

8. Parietal 

Ctenopharyngodon (Fig. 12); as broad as the frontal and half its length. 

Hypophthalmichthys (Fig. 12); half the greatest width of the frontal and 65% of its length. 

Commonly in cyprinids, the parietal is short, being 30%-50% of the frontal length. A 
parietal fontanelle is usually absent in Cyprinidae but may be found in specimens of 
Cyprinus and various gobioines (see Ramaswami, 1955a). This feature, widespread in other 
otophysans is possibly a plesiomorph character and closure of the fronto-parietal fontanelle 
is to be regarded as apomorphic for the Cyprinidae (see also Fink & Fink, 1981). 

9. Circumorbital bones 

Ctenopharyngodon (Fig. 13); 1st infraorbital thick and curved outward, 2nd and 3rd 




Fig. 13 Circumorbital bones: (upper) Ctenopharyngodon; (lower) Hypophthalmichthys. a and b 
are anterior views of the respective 1st infraorbitals. Scales = 10 mm. 

infraorbitals elongate, 4th almost perpendicular, 5th long, crossing the sphenotic; supra- 
orbital extensive. 

Hypophthalmichthys (Fig. 13); 1st infraorbital thin, vertically aligned; 2nd, 3rd and 4th 
elongate, 5th, an almost right-angled bone, the vertical portion crossing the frontal, the 
horizontal part across the sphenotic; supraorbital extensive. 

In both genera the supraorbital is extensive and has the effect of extending ventrally the 
lateral border of the cranial roof so that the ventral edge of the supraorbital comes to lie in 
the same horizontal plane as that of the parasphenoid. In Ctenopharyngodon the supra- 
orbital slopes laterally at the same angle as the frontal, its medial portion is thickened and 
anteriorly bears a wide groove. The lower anterior border of the bone contacts the postero- 
dorsal edge of the 1st infraorbital. In Hypophthalmichthys the supraorbital is directed almost 
perpendicularly and its anterior part contacts half the dorsal margin of the 1st infraorbital. In 
small specimens (up to 100 mm SL), only the canal tube of infraorbitals 2-5 is ossified, the 
upper part of the 4th being fragmented into as many as four portions (Fig. 14). 

Comments made previously concerning infraorbitals (Howes, 1978; 1980) suggested that 
both extreme reduction and expansion of the elements could be seen as derived states. Of 
these the most often encountered condition in cyprinids is for there to be a reduction of 
infraorbital ossification, particularly that of the 5th bone, and also this bone's dissociation 
from the supraorbital. 

Gosline (1974) made much of the type of infraorbital-supraorbital sensory canal 
connection, dividing cyprinids into two groups on the basis of whether or not there was a 
continuous connection between the two canal systems. Howes (1978) refuted this suggestion 
by noting that there was always some kind of connection between the canals. What is 





Fig. 14 


Circumorbital bones and opercular-pterotic canal system of Hypophthalmichthys 
molitrix. Drawn from an alizarin specimen 65 mm SL. Scale = 3 mm. 

possibly of more significance is the form of the 5th infraorbital and its association with the 
dermosphenotic. Contrary to a previous statement (Howes, 1978) the dermosphenotic is 
present in many cyprinids (the element labelled as a dermosphenotic in Oreoleuciscus 
[Howes, 1978, fig. 22] appears to be the fragmented 5th infraorbital, and that in 
'Chelaethiops' [Fig. 23] part of the dermopterotic). The distribution of the dermosphenotic 
amongst cyprinids and the morphology of associated canal systems is discussed in a separate 
paper (Banister & Howes, in preparation). 

The thickened 1st infraorbital of Ctenopharyngodon is an autapomorphic feature, 
although there is a tendency towards thickening of that bone in Mylopharyngodon. Both 
genera, however, share the derived, massive ridged supraorbital. Similarly, the enlarged 
supraorbital of Hypophthalmichthys is shared with Xenocypris. But, again the infraorbital 
pattern of Hypophthalmichthys is autapomorphic. 

10. Supraoccipital 

Ctenopharyngodon (Fig. 12); short with posterior lamellate process. 

Hypophthalmichthys (Fig. 12); long with dorsally thickened crest. 

The supraoccipital of Ctenopharyngodon is the type most commonly present in 
Cyprinidae viz an almost square bone with a low crest. The broad, thick supraoccipital crest 
of Hypophthalmichthys is not encountered elsewhere in the cyprinids. Another autapo- 
morphic feature occurring in this genus is that the anterior portion of the supraoccipital is on 
a raised platform, the sides of which are contributed by the parietals. 

1 1 . Exoccipital 

Ctenopharyngodon (Fig. 1 5); contains a fossa posterior to the subtemporal fossa; the postero- 
ventral border of the bone is horizontal. 

Hypophthalmichthys (Fig. 15); without supplementary fossa; postero-ventral border 
sloping laterally and caudally. 

An earlier statement (Howes, 1978 : 38) claimed that in cyprinids the exoccipital showed 
little variability. This is not the case in the two genera now under discussion. 

In Ctenopharyngodon the posterior border of the exoccipital lateral wall is horizontal a 
condition considered plesiomorphic on account of its widespread distribution. The dorsal 
part of the bone, immediately posterior to the subtemporal fossa, bears a shallow depression 
(also contributed to by the intercalar see below). From this supplementary fossa there arises 
a posterior section of the levator posterior muscle. 










Fig. 15 Posterior cranial region: (left) Ctenopharyngodon; (right) Hypophthalmichthys. 
Scales = 10 mm. Specimens as previous figures. 

In Hypophthalmichthys the posterior exoccipital wall slopes ventrally at an angle of 45 to 
the horizontal, its ventral border is also directed laterally at a similar angle with the result 
that the entire wall forms a triangular wing. The postero-lateral flaring of the exoccipital wall 
is so great that its distal edge contacts the distal tip of the pterotic spine a feature otherwise 
present only in Xenocypris. In all other cyprinids examined the pterotic spine is separated 
from the exoccipital margin by a substantial part of the pterotic floor or by the intercalar. 
The dorsal surface posterior to the subtemporal fossa is flat, the levator posterior taking its 
origin entirely from the subtemporal fossa. 

A levator posterior with a section originating from a supplementary subtemporal fossa as 
in Ctenopharyngodon is known elsewhere only in Mylopharyngodon and Squaliobarbus and 
is considered synapomorphic. 

The lateral occipital fenestra in Ctenopharyngodon is narrow and when viewed posteriorly 
is obscured by the exoccipital medial border; a feature which otherwise occurs only in 
Mylopharyngodon. In contrast, there is an extensive fenestra in Hypophthalmichthys. In the 
Cyprinidae there is quite some variation in the size and shape of the fenestra although 
usually it is small and ovate. Rarely does it have the proportions of that in Hypophthal- 
michthys, which are approached only by Xenocypris and Abramis. To my knowledge, the 
fenestra is never reduced to the extent of Ctenopharyngodon and Mylopharyngodon. Both 
these conditions may be recognized as derived states. 

12. Epioccipital 

Ctenopharyngodon (Fig. 1 5); bullate with a slight posterior ridge; posttemporal fossa exists as 

a mere depression between the epioccipital and the pterotic. 

Hypophthalmichthys (Fig. 1 5); triangular with a prominent posterior lip below which is a 
depression of the epioccipital wall; posttemporal fossa well-developed, extending someway 
anteriorly between the epioccipital and pterotic. 

The epioccipital in Ctenopharyngodon is greatly developed and contributes the major part 
to the formation of the subtemporal fossa. As in all cyprinids the epioccipital roof provides 
the major site of origin for the levator posterior muscles (see Eastman, 1971). The bone in 
Hypophthalmichthys is virtually pyramidal and posteriorly has a broadly pointed lip. This 
lip or shelf serves for the attachment of hypaxial musculature and although variously 
developed throughout cyprinids in none is it more so than in this taxon. 




Fig. 16 Intercalar: (upper) Squaliobarbus curriculum BMNH 1888.5.15:29; (lower) Mylo- 
pharyngodon piceus BMNH 1895.5.31 : 40. Scales in mm divisions. 

13. Intercalar 

Ctenopharyngodon (Fig. 16); extensive and forming the posterior corner of the cranium 
between the basioccipital and pterotic. Its ventral surface contributes to the supplementary 
subtemporal fossa (see above) and provides the site of origin for the posterior section of the 
levator posterior muscle. 

Hypophthalmichthys (Fig. 16); smaller than the former taxon, when viewed posteriorly is 
seen as a triangular wedge intruding into the lateral border of the exoccipital (Fig. 1 5). Its 
ventral surface is broad and lies between the exoccipital and medial edge of the pterotic spine 
(Fig. 9). The intercalar morphology of 'Hypophthalmichthys is autapomorphic. 

In cyprinids the intercalar is transitory, as for example in the aspinine group (see Howes, 
1978) where it is well-developed in most members, but absent in the more derived taxa. 
Rarely is the intercalar developed to the extent that it is in Ctenopharyngodon and occurs 
similarly only in Mylopharyngodon and Squaliobarbus (Fig. 16) where it also provides the 
site of origin for part of the levator posterior muscle. 

In Capoeta the intercalar also contributes to the area of origin of the levator externus (Fig. 
17). However, in this genus the intercalar forms the lateral border to the exoccipital. The 
posterior wall of the exoccipital is strongly concave and provides the site of origin for the 
remainder of the posterior segment of the levator muscle. The exoccipital wall is strongly 
concave and its lateral margin is separated by a wide gap from the pterotic spine. In Cyprinus 
and Carassius the posterior wall of the exoccipital is virtually identical to that of Capoeta. 




Fig. 17 Postero-ventral region of the cranium: (upper) Cyprinus carpio (uncatalogued); (lower) 
Capoeta capoeta BMNH 1879.11.14: 19. Scales = 5 mm. 

However, in these two genera, the intercalar is minute (or may even be absent) and is 
confined to the lateral edge of the pterotic; it plays no part in providing an area of attachment 
for the levator posterior muscle. 

14. Basioccipital 

Ctenopharyngodon (Fig. 19a); ventral keel shallow, concave anteriorly; masticatory plate 
large and ovate with a shallow ventral depression; pharyngeal pad thick, lozenge shaped with 
a longitudinally ridged surface; pharyngeal process broad and thick, its distal end rounded 
and upwardly curved; basioccipital facet circular. 

Hypophthalmichthys (Figs 1 8a & 1 9e); ventral keel deep with markedly concave anterior 
border; masticatory plate cardiform, cancellous with a thick anteriorly directed point; 
pharyngeal pad thin, keratinized, its surface bearing a median ridge; pharyngeal process 
elongate, cylindrical and ventrally directed; basioccipital facet ovate. 

Although there is quite some variation in the morphology of the basioccipital keel and its 
associated masticatory plate, that of Hypophthalmichthys is outstanding in its length and 
shape. Only in Xenocypris and Distoechodon is there a similar degree of development of the 
basioccipital keel (see illustrations in Tomodo, 1979 : 96). The ventral prolongation of the 
basioccipital in Hypophthalmichthys is associated with the epibranchial organs and serves 
for the attachment of the epibranchial duct and a medial ligament that stems from the 
cartilaginous bloc running along the midline. Genera which most closely approach 
Hypophthalmichthys and Xenocypris (including here Distoechodon) are Rutilus and 
Chondrostoma (Figs 18d & c). In these genera and Abramis, the ventral border of the 
basioccipital just anterior to the aortic foramen is formed into a deep channel. The most 
derived condition is in Abramis where a rounded notch indents the ventral border of the 
bone. The basioccipital channel is confluent with a similar feature of the parasphenoid (see 
p. 12) and is filled with cartilage. 



Fig. 18 Basicrania in lateral view: a. Hypophthalmichthys molitrix; b. Xenocypris argenteus; c. 
Abramis brama; d. Rutilus rutilus; e. Chondrostoma nasus. Abramis (c) is also shown in ventral 
view. The solid arrows indicate considered apomorphic features of the parasphenoid- 
basioccipital. Composites, all drawn to same scale. 

The shape and size of the basioccipital masticatory plate is variable in the Cyprinidae. 
Being part of the pharyngeal feeding mechanism it would be expected that a particular 
pharyngeal dentition would be associated with a specific plate morphology. By and large this 
appears to be the case; the most common type of plate is a small triangular one with a 
compressed triangularly shaped posterior process. This morphology is associated with 
omnivorous or piscivorous dental types (simple, often single-rowed, recurved teeth). The 
pharyngeal plate of Ctenopharyngodon greatly resembles that of Squaliobarbus and Rutilus 
(Figs 19a, c & d), both regarded as being plesiomorphic. However, the posterior pharyngeal 
process is of a unique type whereas that in Rutilus and Squaliobarbus is of the plesiomorphic 
form. Mylopharyngodon exhibits a derived, bowl-shaped plate associated with molariform 




Fig. 19 Basioccipital masticatory plates: a. Ctenopharyngodon idellus BMNH 1888.5.15 : 25; b. 
Mylopharyngodon piceus BMNH 1895.5.31:40; c. Squaliobarbus curriculus BMNH 
1888.5.15:29; d. Rutilus rutilus uncatalogued skeleton; e. Hypophthalmichthys molitrix 
BMNH 1895.5.31 : 22; f. Abramis brama uncatalogued skeleton. Scales = 10mm. 

dentition but one obviously little modified from the plesiomorphic pattern and seemingly 
achieved by lateral and posterior expansion (Fig. 19b). A similarly developed plate type is 
found in Labeo species but here, the pharyngeal process is differently modified, forming a 
roof over the aortic channel (Reid, 1978, unpublished thesis). Another derived state occurs 
in Abramis where the pharyngeal plate has virtually disappeared (Fig. 190- Extreme 
reduction is also found in Semiplotus, but a characteristic surface and aortic channel 
morphology is apomorphic for this taxon. 

It is thus concluded that particular details in the morphology of the basioccipital plate and 
posterior process can be useful indicators of relationship. 

15. Posttemporal 

Ctenopharyngodon (Fig. 12); broad, straight with long dorsal arm, basal part of the bone 
applied to the pterotic and intercalar, its upper part bridges the shallow posttemporal fossa to 
lie against the epioccipital. The anterior margin of the dorsal extension is bordered by a 
broad extrascapula. 

Hypophthalmichthys (Fig. 12); elongate with a broad paddle-shaped basal portion, its 
upper half narrow and pointed distally. The anterior border of the bone curves round the 
posterior margin of the extrascapula. The extrascapula is elongate with an irregular anterior 

The Ctenopharyngodon type of posttemporal and extrascapula is widespread throughout 
the cyprinids and is regarded as plesiomorphic. The curved, spine-like posttemporal and 
narrow extrascapula of Hypophthalmichthys is otherwise present only in Xenocypris. 




Fig. 20 Opercular bones: (upper) Ctenopharyngodon idellus BMNH 1861.1.17:6; (lower) 
Hypophthalmichthys molitrix BMNH 1 895.5.3 1 : 22. Scale = 10 mm. 

16. Opercular bones 

Ctenopharyngodon (Fig. 20); preoperculum with a shallow horizontal limb containing 6 

pores of the sensory canal system; operculum large, almost square. 

Hypophthalmichthys (Fig. 20); preoperculum with deep horizontal limb containing 15 
sensory pores; operculum large, deeper than long, its outer surface strongly ridged. 

In both genera the basal part of the antero-dorsal opercular process (to which is attached 
the tendon of the dilatator operculi muscle) contains a tubular canal connecting the 
preopercular canal with the pterotic canal. The connection between the operculum and the 
pterotic in Hypophthalmichthys is effected through an ossified canal lying vertically above 
the opercular process. This small canal-carrying bone appears to be part of the 

There appears to be some variability in the morphology of the opercular sensory canal. 
Gosline (1974) noted that the preopercular canal ends 'blindly' or passes across the opercular 
face. However, where there is a connection between the preopercular and pterotic canals, it 
can be via a dermal opercular canal or 'suprapreopercula', an opercular tube or an epidermal 
connection. These varying conditions and their taxonomic significance are discussed in a 
separate paper (Banister & Howes, in preparation). 

The opercular series, and particularly the size of the operculum are somewhat variable 
throughout the Cyprinidae but certain types of opercula can be associated with particular 
monophyletic groups. Examples are the bariliines where the opercular bones are almost 
always deep and narrow; the neobolines (Chelaethiops, Neobolus) where the opercula have 
an attenuated posterior margin; the chelines and aspinines where the opercula possess a long 
dorsal border. Caution must be exercised, however, in using opercula bone morphology as a 
phyletic character. Large opercular and expanded preopercular bones are to be found in such 
widely different trophic specialists as Luciobrama, Barbus, Ctenopharyngodon and 


Hypophthalmichthys. In these taxa the similarity of opercular bone shapes appear to be the 
result of different functional demands which are at present not understood. 

17. Suspensorium 

Overall, there is a great similarity in the suspensorial elements of Ctenopharyngodon and 

The palatine is broad anteriorly in both genera. However, in Ctenopharyngodon articula- 
tion of the palatine head is with the lateral supraethmoid notch (see p. 5; Fig. 1). The more 
usual condition is for the palatine head to be at a lower level, almost in the plane of the 
preethmoid and to be attached by a ligament to either the anterior rim, the ventral surface of 
the supraethmoid, or to the upper part of the mesethmoid. Only in Mylopharyngodon and 
Squaliobarbus is the palatine head elevated as in Ctenopharyngodon and closely articulated 
with a lateral supraethmoid notch. The posterior part of the palatine in Ctenopharyngodon 
is rounded and articulates in a deep entopterygoid facet. The posterior region of the palatine 
in Hypophthalmichthys is laterally compressed and articulates with a shallow, narrow 
entopterygoid facet. Both types of posterior palatine articulation are encountered widely in 
cyprinids and it is not possible at present to say which is the more derived. 

In both genera the hyomandibula is produced laterally and its articulatory dorsal border is 
steeply angled antero-ventrally. However, in Ctenopharyngodon there is a lateral shelf on 
the upper face of the hyomandibula and the ventral limb of the bone is short and broad, 
whereas in Hypophthalmichthys it is elongate. In both genera the entopterygoid and 
metapterygoid are depressed medially with the metapterygoid having a strongly concave 
dorsal border and a well-defined lateral ridge; the quadrate bears a wide ventro-lateral shelf, 
but in Ctenopharyngodon it is a larger bone than in Hypophthalmichthys. 

There are differences in the form of the symplectic which is elongate in Hypophthal- 
michthys but short and deep in Ctenopharyngodon. This latter type appears to represent the 
more derived condition (see Howes, 1980), the elongate symplectic being common in 

In general, there is uniformity of the suspensorial elements in the Cyprinidae; the varia- 
bility is one of degree, usually involving elongations, deepening or medial depression of the 

18. Jaws 

Ctenopharyngodon (Figs 2 Id, e, 0; maxilla deep with a triangular midlateral ascending 
process; the anterior medial process is directed almost horizontally to contact its partner via 
a ligament; maxillary distal portion is broad, medially curved and produced into a ventral 
flange which contacts the distal part of the premaxilla. Approximately mid-way along the 
maxilla is a foramen sited below a lateral ridge (which serves as the attachment for the 
palatine ligament). The premaxilla is deep with only a short anterior ascending process its 
ventral edge is chisel-like. The dentary has a short, dorsally rounded coronoid process; the 
anguloarticular, a convex dorsal border. 

Hypophthalmichthys (Figs 2 la, b, c); maxilla shallow with a high, square midlateral 
ascending process; the anterior medial process is compressed and directed ventrad toward 
the midline; maxillary posterior portion is laterally compressed and ventrally extended to 
articulate with the premaxilla. The premaxilla is exceedingly shallow and thin with a short, 
broad, anterior ascending process and although its ventral edge is sharp, it is not bevelled as 
in Ctenopharyngodon. The dentary is shallow at the symphysis but deepens posteriorly and 
has a high, backwardly sloped coronoid process. The anguloarticular is short and deep with a 
long, steep dorsal border (60 slope to the horizontal). 

The plesiomorphic cyprinid maxilla appears to be of the Ctenopharyngodon type, viz 
deep, with a large mid-lateral, or posteriorly placed ascending process with convex anterior, 
and concave posterior borders. Apomorphic derivatives involve the formation of a concave 
anterior border to the mid-lateral process as in Hypophthalmichthys, or complete reduction 






Fig. 21 Jaw bones in lateral view: Hypophthalmichthys, a. left dentary, b. right maxilla and c. 
right premaxilla; Ctenopharyngodon, d. left dentary, e. right maxilla and f. right premaxilla. 
Scales = 10 mm. 

of the process as in some bariliines (see Howes, 1980). Another apomorphic feature of the 
maxilla appears to be the presence of an anterior foramen. This foramen provides the 
channel for a branch of the facial nerve which usually innervates an associated barbel. 
Although such a foramen is present in Ctenopharyngodon, there is no associated barbel and 
it will be argued later that this is due to secondary loss of the barbel (see p. 39). For the most 
part, however, it is true to say that those cyprinids with a maxillary foramen also have a 
rostral barbel. One cyprinid group which lack the maxillary foramen but have barbels are 
some bariliines (see Howes, 1980). However, in these taxa the barbel is not associated with 
the maxillary bone. The phylogenetic significance of these various conditions is discussed 
elsewhere (Banister & Howes, in preparation), but here it can be stated that the presence of a 
maxillary foramen is an apomorphic feature in one group of Cyprinidae (see also Banister & 
Bunni, 1980: 157). 

An L-shaped premaxilla without any dorsal or lateral protruberances is thought to repre- 
sent the plesiomorph condition. 

Although superficially alike, the lower jaws of Ctenopharyngodon and Hypophthal- 
michthys differ in the form of the coronoid process and anguloarticular. In the Cyprinidae 
the most common type of lower jaw morphology is one where the coronoid process occupies 
a central position and is formed entirely from the dentary. In these cases the anguloarticular 
has an almost horizontal dorsal border (eg. Barbus, Leuciscus, Aspius, Culler; see fig. 36 in 
Howes, 1978). Contribution of the anguloarticular to the coronoid may be considered 
derived and associated with orientation of the jaw and a shift in mandibular muscle 
insertions. A lower jaw element which may be worthy of more critical attention in cyprinids 
is the coronomeckelian bone. It commonly occurs as a deep, short bone with a well- 
developed shelf along its medial face to which attaches the tendon of part of the adductor 




Fig. 22 Anterior dorsal vertebral elements shown in lateral, dorsal and anterior views of: a. 
Ctenopharyngodon; b. Hypophthalmichthys', c. Abramis brama. Semi-diagrammatic, compiled 
from several specimens; all drawn to same scale. 

mandibulae complex. There is some variation in the shape of this bone and certainly in one 
group of 'leuciscines' its spear-shaped appearance seems to offer an apomorphic character of 
phyletic value (Howes, in preparation). The morphology of the lower jaw is possibly the 
most variable anatomical unit in Cyprinidae, relative growth of its various parts probably 
being a major contributing factor to this variability. 

1 9 . Vertebral column 

Ctenopharyngodon (Fig. 22a); 1st vertebra compressed and with laterally directed processes, 
2nd with an elongate lateral process, 4th with anteriorly sloping pleural rib and backwardly 
sloping neural spine. Weberian complex: tripus broad, sickle-shaped; ossa suspensoria 
making extensive medial contact and forming a posteriorly curved transverse septum. Total 
number of vertebrae 43 (4 + 22 + 1 6 + 1 ). 

Hypophthalmichthys (Fig. 22b); 1st vertebra with caudally directed lateral processes, 2nd 
with short, triangular, caudally directed processes, 4th with anteriorly sloping pleural rib and 
upright neural spine. Weberian complex: tripus elongate with mesially curved tip, ossa 

30 G. J. HOWES 

suspensoria extensive forming an antero-ventrally sloping transverse septum, neural 
complex bifurcated and articulating with supraoccipital. Total number of vertebrae 39-40 

The most striking differences in the morphology of the vertebral column in Cteno- 
pharyngodon and Hypophthalmichthys occur in the anterior vertebrae forming the Weberian 
complex. The neural complex of Ctenopharyngodon is antero-dorsally expanded and curved 
forward so that its anterior tip is above the 1 st vertebra (Fig. 22a); there is no contact with the 
supraoccipital. In Hypophthalmichthys the neural plate is deep and lamellate with a concave 
anterior border. The antero-dorsal prong is bifurcated and the supraoccipital process con- 
tained within the fork (Fig. 22b). The dorsal margin of the neural plate is deeply grooved for 
the midline ligament, as it is in the aspinines (see Howes, 1978 : 19). 

In the Cyprinidae contact between the supraoccipital crest and the neural complex is 
uncommon. It occurs in Labeo, Catla, Osteobrama and Semiplotus, but in these taxa there is 
no deeply grooved dorsal margin to the neural complex, and the anterior bifurcation is either 
shallow or lacking. Only in Xenocypris does the neural complex have nearly the same 
morphology as in Hypophthalmichthys. However, a well-developed lamellate neural plate is 
commonly encountered in the Cyprinidae and may be a plesiomorphic character. In 
Abramis, Rutilus and some species of Chondrostoma the neural complex has no antero- 
dorsal extension but is perpendicular or anteriorly inclined and deeply forked; a condition 
unique in cyprinids (Fig. 22c). In some catostomids (Carpoides) there is contact between the 
supraoccipital and neural complex. Here the condition is similar to that in Labeo where the 
neural plate is lamellate and without a dorsal groove or anterior bifurcation. 

It seems likely that in the latter cases expansion of the neural complex (? fused 3rd and 4th 
supraneurals) and its contact with the supraoccipital is associated with a deepened body and 
the forceful forward probing movements of the head when feeding. Reinforcement of the 
anterior part of the vertebral column and rigidity of the posterior cranium is provided by this 
arrangement. In Hypophthalmichthys which is essentially a mid-water or surface feeder, this 
feature must serve a different purpose and may be associated with some elevation of the 
cranium during feeding. 

The common (plesiomorph) condition of the first four (Weberian) vertebrae in Cyprinidae 
is as in Ctenopharyngodon (the typical condition is seen in Opsaridium microcephalus, 
Howes, 1980, fig. 39D). Similar development of the transverse processes, tripus and ossa 
suspensoria similar to those of Hypophthalmichthys is encountered elsewhere in members of 
the 'Leuciscinae' (ie Leuciscus, Abramis) and the aspinine group. However, elongation of 
these elements is also found in some barbines, chelines and neobolines (eg. Chelaethiops) and 
its usefulness as an indicator of phyletic relationship is doubtful. 

20. Gill- arches 

Ctenopharyngodon has no particular modifications of the gill arches, which are similar to 
those of other cyprinids. The gill-rakers are short and spiny, biserially arranged and almost 
horizontally placed on the arch so that they intermesh with those on the neighbouring arch 
to form a filter. 

Hypophthalmichthys displays marked modification of the gill arches and possesses a 
complex epibranchial organ (Figs 24-26). 

The gill structures of Hypophthalmichthys have been described in varying detail by a 
number of authors. Boulenger ( 1 90 1 ) first described the epibranchial organ; Fang ( 1 928) gave 
a detailed description of the gill arches and epibranchial organ, and speculated on their 
function. Zambriborshch (1955; 1957) described and figured the gross morphology of the 
branchial apparatus and histology of the gill-rakers, including comments on function. 
Verigin (1957) described and figured the epibranchial organ, gill-rakers, upper pharyngeal 
pad and roof of the pharynx. He also studied the ontogeny of the branchial apparatus. 
Wilamovski (1972) made a description of the epibranchial organ and gill-rakers, comparing 
them with the branchial system of Cyprinus. Taken in toto these descriptions give an 




Fig. 23 Posterior gill-arch elements: (left) Hypophthalmichthys molitrix, from dissected specimen 
BMNH 1895.5.31:22; (right) Xenocypris argenteus from alizarin specimen BMNH 
1 889.6.8 : 19. Left scale = 5 mm, right = 2 mm. 

adequate account of the organ's morphology, and it is necessary here only to emphasize 
certain of its apomorph characters. 

The ceratobranchials are elongate, bearing numerous biserially arranged slender gill- 
rakers. The two rows of rakers are set at a diverging angle to one another thus producing an 
'interbranchial space' (Fang, 1928). There are approximately 13-14 rakers per 1 mm of 
ceratobranchial. Fang (1928) recognized two types of gill-rakers, broad and narrow, the 
broad type occurring between each 4th and 5th narrow type. The gill-rakers are connected 
by a reticulate mucuous membrane (sifting membrane of Fang, 1928) which forms a network 
or filter, whose pores are about 20 // in diameter. The gill-rakers continue onto the 

The 1st and 4th epibranchials are expanded dorsally into axe-shaped elements. The 1st 
forms the anterodorsal wall, and the 4th the posteromedial wall of the epibranchial organ 
(the branchial arches are so orientated that the 4th epibranchial comes to lie in a medial 
(longitudinal) plane rather than a posterior (transverse) one). Although not entirely ossified, 
a 5th epibranchial is present as a well-developed, broad, lamellate element, and has a 
complex articulation with the 4th ceratobranchial (Fig. 23). 

There are two large well-ossified infrapharyngobranchials recognized as IF2 and IF3; IF4 
is represented by an extensive cartilaginous bloc bordering the posterior rim of IF3 and 
contacting epibranchials 3 and 4 (Fig. 24). In front of IF2 there is another large cartilaginous 
cap onto which articulates epibranchial 1. This no doubt is IF1. In adult specimens the 5th 
epibranchial is joined to the capsule of the epibranchial organ via an interposed cartilage 
which may represent the 5th infrapharyngobranchial. 

The epibranchial organ is formed by four curled cartilaginous channels or ducts, which 
are prolongations of the four epibranchials (Fig. 26). Each duct is lined with two rows of 
biserially arranged gill-rakers, the posterior row of one arch intermeshing with the anterior 
row of the adjacent arch and in this way running into each duct (Fig. 24). The outer row of 
the 1st arch enters a small chamber of the first duct and becomes reduced in size and number. 
The first duct houses the gill-raker rows of the 1st and 2nd epibranchials, the 2nd those of the 





Fig. 24 Hypophthalmichthys epibranchial organ: a. anterior view of left organ, ossified areas of 
the infrapharyngobranchials are stippled; b. dorsal view of left organ to show orientation, dashed 
line represents the ventral midline of the head, and solid arrow indicates the direction of view 
depicted in a. Scales = 10 mm. 

2nd and 3rd epibranchials, the 3rd those of the 3rd and 4th, and the 4th duct those of the 4th 
and 5th epibranchials. Zambriborshch (1957) referred to the organ ducts as supra- and 
infrapharyngobranchials. However, the infrapharyngobranchials are here identified as 
elements distinct from the cartilaginous ducts. 

The entire epibranchial structure is invested in palatal connective tissue. The tissue below 
the basal part of the organ is produced into four folds (palatal folds of Fang, 1928) which 
hang down into the interbranchial channels (Figs 25 & 26). The lateral face of the 
epibranchial organ, also formed from palatal tissue, is connected to the medial face of the 
preoperculum via a segment of the adductor hyomandibulae muscle (= pharyngopraeo- 
percularis muscle of Zambriborshch, 1955); see Fig. 25. 

The morphology of the epibranchial arches in Hypophthalmichthys is unique. Modified 
gill-arches and gill-rakers are, however, found elsewhere in cyprinids. In Xenocypris 
argenteus the epibranchials are rounded over a thickened palatal tissue pad, the gill-rakers 
are numerous, approximately 25 per 1 mm of gill-arch, and extend to the proximal part of 
the epibranchial. The infrapharyngobranchials extend into the dorsal surface of the palatal 
tissue pad. The pads or cushions have a hollow interior, the internal surface of the tissue is 
highly papillose and convoluted. Similar development of the epibranchials, gill-rakers and 
palatal cushions are to be found in Plagiognathops, although the gill-rakers on the 1st arch 
are 'normal' and the palatal cushion is more papillose. 

An hypothesized plesiomorph condition of the epibranchial organ is encountered in 
Abramis. Here the epibranchials are markedly rounded and the palatal tissue greatly 
thickened. This is particularly evident in Abramis sapa where the palatal cushion occupies a 
large part of the pharyngeal cavity (Fig. 27b). Anteriorly it is rounded and forms a cup- 
shaped depression in the palate. The palatal cushion resembles that of Xenocypris in having 
a hollow interior, although not as cavernous. The outer surface of the cushion shows slight 
ridging between the gill-arches, reminiscent of the palatal folds in Hypophthalmichthys. The 
cartilaginous proximal tips of the epibranchials are highly developed and lie embedded in a 
cartilaginous casing which extends along the basioccipital (Figs 27a, b). Again, as in 









Fig. 25 Hypophthalmichthys, postero-lateral view of epibranchial organ and gill-arch elements. 
The ceratobranchials have been partly cut away to show the organ's folded tissue surface. 
Schematic drawing based on two dissected specimens. 

Xenocypris the cartilaginous tips of the epibranchials extend ventrally into the floor of the 
palatal cushion roof. 

Elsewhere in the Cyprinoidei an epibranchial organ is present in Catostomidae. In these 
taxa as in Hypophthalmichthys and the xenocyprinids, the epibranchials are rounded over a 
complex pharyngeal pad and are medially prolonged (Fig. 28a). In catostomids the 
pharyngeal pad more closely resembles that in Xenocypris being a medially undivided 
structure surrounding the basioccipital process, and not in the form of paired diverticula as 
in Hypophthalmichthys. The catostomid basioccipital is expanded laterally and is fretted, 
playing an integral part in the structure of the pharyngeal cushion. This is unlike the 
cyprinid condition where the basioccipital acts as a support for connective tissue dividing the 
organ (see Weisel, 1960 for an account of the catostomid pharyngeal organ, and Eastman, 
1977 for a description of catostomid pharyngeal anatomy). 

When compared to the epibranchial organs of other teleosts which possess them, that of 
Hypophthalmichthys most closely resembles the continuous tube organs of some clupeids 
(see Miller, 1969). 

It is possible that a form of epibranchial organ occurs in the south-east Asian cyprinid 
genus Thynnichthys. This fish shows a remarkable superficial resemblance to Hypophthal- 
michthys, particularly in its low positioned eye. As in the latter, the gill-arches are elongate 
but they lack gill-rakers. There does, however, appear to be a swollen area of pharyngeal 







Fig. 26 Hypophthalmichthys, ventral dissection of the epibranchial organ, ceratobranchials cut 
away. Scale=10mm. Drawn from specimens BMNH 1895.5.31 : 36-37 and uncatalogued 
Swinhoe specimen. 

tissue with a single posterior median duct leading into the oesophagus. Only a single 
specimen is available for superficial dissection but those osteological features which are 
observable are unlike those of Hypophthalmichthys and more closely resemble Ambly- 

Although Berry & Low (1970) described certain morphological features in hybrids 
between Hypophthalmichthys and Ctenopharyngodon, no mention was made of the develop- 
ment of an epibranchial organ. 

A specialization of the upper branchial arch shared by Hypophthalmichthys and 
Xenocypris is hypertrophy of the 5th epibranchial. In Xenocypris this element is broadly 
lamellate and has a form of articulation with the 4th ceratobranchial similar to that of 
Hypophthalmichthys (Fig. 28a). In cyprinids the 5th epibranchial is greatly reduced or 
appears as a small cartilaginous element above the 4th epibranchial (Banister, pers. comm.). 
The hypertrophy of this element and its articulation with the 4th epibranchial are seen as 
synapomorphies uniting Hypophthalmichthys and Xenocypris. 

A superficially similar gill-arch morphology to that in Hypophthalmichthys occurs in 
Gibelion and Catlacarpio (Fig. 28d). In these taxa also the ceratobranchials are elongate and 
bear numerous gill-rakers. The epibranchials are sickle-shaped, their medial 'handle' 
portion being dorso-ventrally expanded and antero-posteriorly compressed. The two infra- 
pharyngobranchials are thickened, IF2 being virtually reorientated from the usual horizontal 
plane to a vertical one. The entire upper branchial basket is surrounded by a thick, papillose 
pharyngeal cushion and this, together with the fretted basioccipital pharyngeal process 
greatly resembles the upper pharyngeal anatomy of catostomids. It may be that the catline 
taxa employ similar feeding methods to those of catostomids (see Weisel, 1960). Again, 
similarly developed epibranchials are present in "Labeo" 1 stoliczkae (Fig. 28c), but they are 







Fig. 27 Upper pharyngeal anatomy ofAbramis. Dorsal branchial elements of: a. Abramis brama, 
left side BMNH 1 974.9. 5 : l-3l; b. Abramis sapa, right side BMNH 1967.7.17 : 3^1, drawn in 
situ, scale = 1 mm; c. lateral view of cartilaginous basicranial bloc and pharyngeal pad in A. 
brama, scale = 10 mm; d. pharyngeal cavity of A. sapa showing 4th ceratobranchial from which 
most of the gill filaments have been removed, scale = 5 mm. 

not so acutely curved medially, and neither are the infrapharyngobranchials orientated in 
the same way. Furthermore, the basioccipital process is thick and broad. Other neurocranial 
characters suggest that 'Labeo ' stoliczkae has as its closest relative, Cirrhina, whilst Gibelion 
and Catlacarpio seem closely related to Thynnichthys, Amblypharyngodon and Osteobrama 

Pharyngeal bones and teeth of both genera have been described and figured by Chu (1935). 
The pharyngeal bone of Ctenopharyngodon has a greatly attenuated lower limb and there is 
a prominent process (for the attachment of the posterior portion of the levator posterior) at 
the curvature of the upper limb. The teeth are in two rows (2, 4-5), greatly compressed and 
with their grinding surfaces serrate, the serrations being the lateral margins of grooves on the 
tooth's outer face. 



Fig. 28 Posterior views of the right 4th epibranchial of: a. Xenocypris argenteus BMNH 
1889.6.8 : 19-22 (alizarin); b. Abramis ballerus (uncatalogued skeleton); c. Labeo stoliczkae 
BMNH 1 89 1 . 1 1 .30 : 286 (skeleton); and d. Gibelion catla BMNH 1 908. 1 2.28 : 1 1 22 (skeleton). 
Scales for a, b, and c = 2 mm, d = 5 mm. 

According to Chu (1935), the teeth and pharyngeal bone of the Hypophthalmichthinae 
'. . . are singularly unique, without parallel in other cyprinids'. The anterior limb of the 
pharyngeal bone is dorsoventrally depressed and the upright posterior limb is antero- 
posteriorly compressed. There is a large fenestra within the expanded dentigerous part of the 
bone. In situ this fenestra is covered laterally by a translucent tissue, medially it serves as a 
cavity into which the expanded edge of the basioccipital masticatory plate inserts (Fig. 26). 
The 4 teeth of Hypophthalmichthys are in a single row, are compressed, have a concave 
occlusul surface with a transverse median groove and lateral striation. 

Chu (1935) is correct in describing the pharyngeal dentition of Hypophthalmichthys as 
unique. There are, however, similarities between the pharyngeals of this species and those of 
Xenocypris and Plagiognathops, where there is also an extensive foramen in the pharyngeal 
bone, and the teeth have similar grinding surfaces (see Chu, 1 935). 

2 1 . Adductor hyomandibulae muscle 

In Ctenopharyngodon the adductor hyomandibulae is developed as in the majority of 
cyprinids, namely, a thick dorsal element arising from the face of the prootic and inserting, 
musculously, on the upper medial face of the hyomandibula. 

In Hypophthalmichthys the muscle is expanded and is continuous (see below) with the 
adductor arcus palatini, forming a muscular 'wall' which effectively separates the branchial 
and buccal cavities. The upper part of the muscle (Dahm, Fig. 29) arises from the prootic and 
inserts on the upper medial face of the hyomandibula as in other cyprinids. However, a 
posterior segment attaches to the antero-dorsal wall of the epibranchial organ and another 
part of the (? same segment) of muscle runs from the lateral wall of the epibranchial organ to 
the preoperculum (pharyngopraeopercularis of Zambriborshch, 1955). Anterior to the 
dorsal part of the muscle another element can be distinguished originating from the lower 
part of the prootic and the pterosphenoid spur to insert on the medial section of the 
hyomandibula. Anteriorly this second portion is continuous with the adductor arcus palatini 










Fig. 29 Hypophthalmichthvs, ventro-posterior view of adductor hyomandibularis musculature, 
based on specimens BMNH 1895.5.31 : 36-37. Scale = 10 mm. 

muscle which originates from the parasphenoid ventral process and inserts along the lower 
limb of the hyomandibula and, anteriorly, along the dorsal rim of the metapterygoid. 
Winterbottom (1973) points out that the adductor hyomandibulae and, at least in certain 
taxa, the adductor arcus palatini are parts of a single muscle. Certainly, in this case it is only 
the orientation of the fibres that distinguishes the upper medial section of the muscle as part 
of the adductor hyomandibulae rather than as part of the adductor arcus palatini. 

Such a highly developed adductor hyomandibularis musculature occurs elsewhere only in 
Abramis, Rutilus and Chondrostoma. In Abramis the upper medial portion of the muscle is 
strongly attached to the pharyngeal tissue, and thence to the posterior border of the 
hyomandibula. Although in Xenocypris the adductor hyomandibulae is not as expansive as 
in the other genera, the upper medial section is attached, along its entire ventral surface, to 
the tissue covering the branchial arches, and some fibres appear to extend on to the 
preoperculum. Morphologically, this can be considered a forerunner of the derived 
condition realized in Hypophthalmichthys where part of the adductor hyomandibulae 
becomes attached to the epibranchial organ and preoperculum and possibly acts as the 
power unit forcing water through the epibranchial ducts. The adductor hyomandibulae 
morphology in Hypophthalmichthys, Xenocypris, Abramis, Rutilus and Chondrostoma is 
considered synapomorphic. 

38 G. J. HOWES 

22. Pectoral girdle 

The pectoral girdle of Ctenopharyngodon is remarkably similar to that of Barbus and 

Cyprinus, except that the posterior cleithral lamina is extended posteriorly. The horizontal 

cleithral limb is broad, with a concave anterior border. The cleithra contact each other only 

at their anterior tips. The coracoid is shallow and rounded posteriorly; the mesocoracoid is 


The pectoral girdle of Hypophthalmichthys displays many modifications relative to the 
presumed plesiomorph pattern. The horizontal cleithral lamina is greatly expanded and rises 
in the vertical plane to join its partner for the anterior third of its length. The posterior 
border of the coracoid is square. The mesocoracoid limb is broad. In both genera the 
postcleithra are elongate and curved mesially, ie. plesiomorphic state; see Howes (1980). 

In no other cyprinid is the horizontal cleithral lamina expanded to the same extent as in 
Hypophthalmichthys. Only in Abramis and Xenocypris is there an approachable condition. 
In these taxa, however, the medial cleithral border is deeply notched at a point just below the 
dorsal articulation of the mesocoracoid. The function of this notch is to contain the 
expanded border of the pharyngeal bone. The mesocoracoid limb is broad in all these taxa. 

Phylogenetic relationships 

From the foregoing descriptions and analyses of Ctenopharyngodon and Hypophthal- 
michthys, a series of apomorph characters can be identified, none of which is shared by both 
taxa. Instead, these characters serve to relate the respective genera to other taxa and thus 
Ctenopharyngodon and Hypophthalmichthys must be considered members of two distinct 
monophyletic assemblages. 

Relationships of Ctenopharyngodon 

The following characters are identified as apomorphies. 

1 . Subtemporal fossa extending anteriorly to invade the autosphenotic on which part of 
the pharyngeal musculature originates (p. 1 5) 

2. Lateral notch in the supraethmoid with which the palatine head articulates (p. 5) 

3. Extensive prootic with ventrally directed trigemino-facialis foramen (p. 1 3) 

4. Extensive intercalar forming posterior corner of cranium and contributing to an 
accessory subtemporal fossa (p. 22) 

5. Divided levator posterior muscle, the posterior segment originating from the intercalar 
(P. 22) 

6. Extensive and bullate epioccipital (p. 2 1 ) 

7. Reduced lateral occipital fenestra (p. 2 1 ) 

8. Olfactory foramen countersunk entirely within the lateral ethmoid (p. 9) 

9. Frontal broad and sloping ventrally (p. 18) 

1 0. Massive and shelved supraorbital (p. 20) 

1 1 . Orbital portion of parasphenoid broadened and with ventral ridge (p. 11) 

12. Ventrally directed pterosphenoid process (p. 10) 

1 3 . Thickened 1 st infraorbital (p. 20) 

14. Distally broadened and upwardly curved basioccipital pharyngeal process (p. 23) 

Characters 1-6 are synapomorphies with Mylopharyngodon and Squaliobarbus; 7-1 1 with 
Mylopharyngodon; 12-14 are autapomorphic. 

From trie distribution of these apomorph characters, Mylopharyngodon is considered to be 
the sister taxon of Ctenopharyngodon with both forming the derived sister group of 
Squaliobarbus. These three taxa constitute a monophyletic lineage termed the Squalio- 
barbine group. 

Mylopharyngodon (Fig. 30b) is a monotypic genus (type species M. piceus (Richardson, 
1845)), characterized by the autapomorphic features of an extensive bowl-like masticatory 



Fig. 30 Outline drawings of members of the squaliobarbine group: a. Ctenopharyngodon idellus; 
b. Mylopharyngodon piceus; and c. Squaliobarbus curriculus. 

plate (Fig. 19b); large molariform pharyngeal teeth; small basioccipital facet dorsally over- 
lapped by an exoccipital ridge (Fig. 31); and narrow, mesially curved posterior cleithral 

Squaliobarbus (Fig. 30c) is the plesiomorph member of the lineage differing from its sister 
taxa in possessing two pairs of small barbels. The anterior pair of barbels are minute and 
appear superficially as mere appendages of the ventral infraorbital skin fold. However, the 
barbel is innervated by a sub-branch of the VII facial nerve. The main nerve branch passes 
through a foramen in the maxilla just below a shelf which forms the attachment point of the 
palatine ligament. The barbel originates some distance posterior to the maxillary foramen 
and the main nerve branch supplies the lip fold. 

It was stated on p. 28 that a maxillary foramen carrying a nerve supply to an anterior barbel 
is an apomorph character uniting those taxa which possess it. It was also noted (p. 28) that 
Ctenopharyngodon has a maxillary foramen but both nerve and barbel are lacking. In 
Mylopharyngodon there is no trace of a maxillary foramen or barbels. 

Two hypotheses are proposed to account for the distribution of barbels and associated 
foramina amongst squaliobarbines: 

1 . That Ctenopharyngodon and Mylopharyngodon represent a plesiomorph condition in 
which the foramen and/or barbel has not formed. 

2. That the absence of barbels and associated nerve in these genera represents secondary 




Fig. 31 

Mylopharyngodon piceus; posterior view of basioccipital from specimen BMNH 
1895.5.31 : 40. Scale = 5 mm. 

It is assumed that the second hypothesis is more likely to be the correct one as the plesio- 
morph lineage (Squaliobarbus) possesses barbels, and that in this taxon their reduction and 
that of the associated nerve signifies transition to loss state. Also, reduction of barbels is 
recorded in other cyprinid taxa (see Banister & Bunni, 1980). 

The distribution of cyprinid barbels; anterior and posterior, whether they are associated 
with a foramen, the type of nerve supply and form of proximal attachment is a subject dealt 
with elsewhere (Banister & Howes, in preparation). For the purposes of this discussion it is 
considered that the squaliobarbine assemblage form part of that series of taxa which possess a 
maxillary foramen carrying a nerve supply to an anterior barbel. 

The suite of apomorph characters defining the squaliobarbines involves, for the most part, 
the occipital region of the cranium the expansion of the subtemporal fossa and the inter- 
calar. These osteological features are concomitant with an increased evolutionary develop- 
ment of the levator posterior muscle which extended forward into the autosphenotic and was 
divided posteriorly, the posterior section originating from the intercalar. These specializ- 
ations could be associated with the essentially phtyophagous habits of the squaliobarbine 

The proposed theory of relationships is to regard the squaliobarbine group as a divergently 
specialized sister lineage to all other barbelled Cyprinidae (i.e. Cyprininae); see Fig. 32. 

Synopsis of taxa assigned to the Squaliobarbine group 

Ctenopharyngodon Steindachner, 1866 

Pristiodon Dybowskii, 1877 

Type and only species: C. idella (Valenciennes, 1844). 
Distribution: Pacific drainage of the USSR and China, lat. 50-23N. 

Introduced into many tropical and temperate areas including Europe, North America, India, 
New Zealand, Malaysia. 

Principal bibliographic sources concerning biology: Lin, 1935; Nikolsky, 1954; Inaba & 
Nomura, 1956; Hickling, 1967;Okada, 1960; Cross, 1969; Edwards, 1974. 




Fig. 32 Cladogram representing the phyletic relationships of the squaliobarbine group. 
Ci = Ctenopharyngodon, My = Mylopharyngodon, Sq = Squaliobarbus. Node 1. Possession 
of maxillary foramen. Node 2. Synapomorphies 1-7 (page 38). Node 3. Synapomorphies 
8-1 1 (page 38), also loss of maxillary foramen in Mylopharyngodon. 

Mylopharyngodon Peters, 1873 

Myloleucus Gunther, 1873 
Myloleuciscus Garman, 1912 
Leucisculus Oshima, 1920 

Type and only species: M. piceus (Richardson, 1 845). 

Distribution: Amur, China, Formosa. 

Principal bibliographic sources: Nikolsky, 1954 & 1956. 

Squaliobarbus Gunther, 1868 

Type species: S. curriculus (Richardson, 1845). 

Sauvage (1884) described another species, S. caudalis, from Tonkin. Pellegrin (1934), when 

commenting on the Hanoi collection of Sauvage, made no mention of this species but 

included in his list, S. curriculus. 

Distribution: Amur, China, Western Korea, Hanoi. 

Principal bibliographic source: Nikolsky, 1956. 

Relationships of Hypophthalmichthys 

The following characters are identified as apomorphies: 

1 . Medial ethmoid notch floored by the vomer (p. 9) 

2. Frontal with an acute lateral slope, providing an extensive cranial dilatator fossa (p. 1 5) 

3 . Extensive lateral occipital fenestra (p. 2 1 ) 

4. Modification of the adductor hyomandibulae muscle involving partial attachment to 
the preoperculum and upper pharyngeal tissue (p. 36) 

5. Expanded horizontal cleithral lamina (p. 38) 

6. Markedly rounded epibranchials (p. 33) 

7. Deep basioccipital keel with marked concave anterior border (p. 23) 

8. Carotid foramen contained in ventrally produced parasphenoid process (p. 12) 

9. Lateral ethmoid-frontal fenestra (p. 9) 

1 0. Expanded 1 st and 4th epibranchials, the 5th present and hypertrophied (p. 34) 


1 1 . Extensive fossa in pharyngeal bone (p. 36) 

12. Exoccipital wing extended distally to contact pterotic spine (p. 21) 

13. Curved posttemporal (p. 25) 

14. Pterosphenoid contributing to anterior trigemino-facialis foramen (p. 10) 

1 5. Cardiform masticatory plate with tubular posterior extension (p. 23) 

16. Intercalar invading border of exoccipital (p. 22) 

1 7. Complexly developed epibranchial organ (p. 3 1 ) 

18. Neural complex bifurcated and articulating with supraoccipital (p. 30) 

Characters 1-6 are synapomorphic with Abramis and Xenocypris (1 Plagiognathops and 
Distoechodon); 7-13 with Xenocypris; 14-18 are autapomorphic. 

From the distribution of these apomorph characters Xenocypris is regarded as the twin 
taxon to Hypophthalmichthys. Together they form the sister group to an assemblage 
including Abramis. 

Xenocypris (Fig. 33b). The taxonomy of this genus is confused, two other monotypic taxa, 
Plagiognathops and Distoechodon being regarded by some authors as subgenera (see 
Banarescu, 1970). All these taxa are presently placed in the subfamily Xenocyprinidinae. 

Two previous comments regarding Xenocypris are in need of modification. The first 
(Howes, 1979), was to the effect that the Xenocyprinidinae was polyphyletic, viz that the 
included genera, Xenocypris, Plagiognathops and Distoechodon shared no derived 
characters. In fact they do (see below). The second statement (Howes, 1980) was that the 
closest relatives of Xenocypris are amongst the hemicultrines (Hemiculter, Hemiculterella 
and Ochetobius). This assumption was based on what are now deemed to be symplesio- 
morphies, and indeed, the hemicultrines as earlier conceived represent a paraphyletic 

Plagiognathops and Distoechodon share with Xenocypris (as represented by the type 
species X. argenteus) apomorph characters involving the gill-arches and basioccipital, 
although they are not so highly derived as in Xenocypris. Lack of osteological material 
prevents a complex diagnosis of the character complexes and the three genera are considered 
together as an unresolved trichotomy. 

Abramis (Fig. 33c), here including the 'genera' Vimba and Blicca\ see p. 46, shares with 
Xenocypris and Hypophthalmichthys synapomorphic characters 1-6, involving the ethmoid 
region, cranial roof, pharyngeal roof covering and associated musculature. However, a 
simple sister-group relationship between Abramis and Xenocypris + Hypophthalmichthys is 
doubtful in the light of other apomorphies possessed by Abramis and exclusively shared with 
Chondrostoma and Rutilus (Figs 33d & e). These are: 

(i) confluent ventral channelling of the parasphenoid and basioccipital; 
(ii) columnar, dorsally bifurcated neural complex; and 
(iii) gill-rakers with globular, papillate bases. 

The underlying synapomorphy (Saether, 1979) uniting Rutilus, Chondrostoma, Abramis, 
the Xenocypris complex and Hypophthalmichthys is a modification of the basicranium, viz a 
parasphenoid notch or confluent ventral channelling of the parasphenoid and basioccipital, 
and deepening of the basioccipital process. From the distribution of synapomorphies within 
this assemblage termed the Abramine group two hypotheses of generic relationships are 
available and represented as cladograms A & B (Fig. 34). 

In cladogram A, Abramis is reckoned to be the sister group of Rutilus and Chondrostoma 
(the similarity between the parasphenoid-basioccipital morphology links these two genera as 
sister taxa) on the basis of characters (i) and (ii) cited above. In this case, the characters 
Abramis shares with Hypophthalmichthys and the Xenocypris complex (listed as synapo- 
morphies 1-6 on p. 41) must be considered as convergent. 

In cladogram B, Abramis is considered as the sister group to Hypophthalmichthys + the 

Fig. 33 Outline drawings of members of the abramine group: a. Hypophthalmichthys molitrix; b. 
Xenocypris argenteus; c. Abramis brama\ d. Chondrostoma nasus; e. Rutilus rutilus. 




Ab Hy 




Fig. 34 Cladograms expressing two schemes of phyletic relationships of the abramine group. 
Ru = Rutilus, Ch = Chondrostoma, Ab = Abramis, Hy = Hypophthalmichth ys, X-P- 
D = Xenocypris-Plagiognathops-Distoechodon trichotomy. 

Cladogram A. Node 1. Modification of basioccipital-parasphenoid; development of pharyngeal 
pad; transversely convex vomer; hypertrophy of adductor hyomandibulae muscles. Node 2. 
Basioccipital-parasphenoid channel; columnar neural complex; globular based gill-rakers. (Con- 
vergence of 6 characters in Abramis with Hypophthalmichthys and Xenocypris.) Node 3. 
Synapomorphic basioccipital morphology (see Fig. 18). Node 4. Synapomorphic gill-arch struc- 
ture; development of epibranchial organ and associated basicranial deepening. (Loss of convex 
vomer and deep ethmoid in Hypophthalmichthys.) 

Cladogram B. Node 1. Combination of nodes 1 and 2 in cladogram A. Node 2.= node 3 in 
cladogram A. Node 3. Synapomorphies 1-6 (p. 41). Node 4. Synapomorphies 7-13 (p. 41); also 
loss of 5 characters through nodes 1-3 (see text, p. 43). 

Xenocypris complex. In this case, five characters must be recognized as having been 
secondarily lost in the two latter taxa, viz: parasphenoid-basioccipital channel; columnar 
neural complex; globular based gill-rakers (lost in both Hypophthalmichthys and 
Xenocypris); convex and deeply notched ethmoid (lost in Hypophthalmichthys). 

It may be argued that in their vertebral elements Hypophthalmichthys and Xenocypris are 
conservative and have retained the plesiomorphic lamellate neural complex. Likewise, 
Hypophthalmichthys may have retained the plesiomorphic ethmo-vomerine bloc. (No 
evidence is available to show that apomorphic modifications of the ethmoid region are 
linked with those of the basicranium.) If this is so, then the more likely hypothesis is that 


Rutilus, Chondrostoma and Abramis form the sister group to Hypophthalmichthys and the 
Xenocypris complex, as one has only to postulate the loss of the apomorphic ethmo- 
vomerine assembly in Hypophthalmichthys (cladogram A). This is also the more likely 
hypothesis of relationships if the autapomorphic parasphenoid-basioccipital and neural 
complex of Abramis are seen as terminal points of a possible evolutionary sequence 
represented by the more plesiomorphic types in Rutilus and Chondrostoma. 

Although it may appear that Abramis displays an incipient form of the epibranchial struc- 
ture in Xenocypris, this is misleading. In fact, the gill-arch morphology and pharyngeal 
dentition suggests that the pharyngeal pad of Abramis differs functionally from that of 
Xenocypris and its relatives. There is indeed, no 'morphoclinal' or 'transitional' series 
represented in extant abramine genera which might reflect the evolutionary development of 
an epibranchial organ. However, the basic architectural modification of the basicranium 
(deepening of the parasphenoid and basioccipital) can be seen as a preadaptation to the 
formation of an epibranchial structure which in the abramines appears to have followed two 
evolutionary pathways hypothesized to be most closely represented by the relationships 
expressed in cladogram A. Falsification of either hypothesis will be forthcoming when more 
complete anatomical data are available for Plagiognathops and Distoechodon and a 
functional analysis of pharyngeal/epibranchial mechanisms available for all abramine taxa. 

Synopsis of taxa assigned to the Abramine group 

Hypophthalmichthys Bleeker, 1860 

Cephalus Basilewsky, 1855 
Abramocephalus Steindachner, 1869 
Onychodon Dybowski, 1872 
Aristichthys Oshima, 1915 

Type species: H. molitrix (Valenciennes, 1 844). 

Two other species, H. nobilis (Richardson, 1844) and H. harmandi Sauvage, 1884 are 

included. Reasons have already been given for synonymizing Aristichthys (p. 2) 

Distribution: Amur, China. Introduced into Taiwan, Thailand, Japan, Sri-Lanka, Europe 

and North America. 

Principal bibliographic sources concerning biology: Tsui, 1936; Nikolsky, 1956. 

Xenocypris Giinther, 1868 

Type species: X. argentea Giinther, 1868. 

Another 9 or 10 species are recognized by various authors; see Banarescu (1970) for most 

recent revision, and Nikolsky ( 1956) for biological data. 

Plagiognathops Berg, 1907 

Type and only species: P. microlepis (Bleeker, 1871). 

Distribution: Yangtze, middle Amur, Ussuri, Lake Khanka (see Berg, 1949). 

Distoechodon Peters, 1880 

Type and only species: D. tumirostris Peters, 1880. 
Distribution: Southern China. 

Rutilus Rafmesque, 1 820 

Scardinius Bonaparte, 1837 
Hegerius Bonaparte, 1845 
Acanthobrama Heckel, 1843 
Leucos Heckel, 1843 
Pigus Bonaparte, 1845 
Cenisophius Bonaparte, 1845 
Gar danus Bonaparte, 1845 

46 G. J. HOWES 

Pararutilus ; Bonaparte, 1845 

Pachychilon Steindachner, 1 866 

Rubellus Fitzinger, 1873 

Mirogrex Goren, Fishelson & Trewavas, 1973 

Type species: R. rutilus Rafmesque, 1820. 

Numerous nominal species with a wide distribution throughout Europe, USSR, Israel to the 

Tigris and Euphrates. 

The characters used to define Leuciscine genera are criticized below (p. 48). Typical are the 
use of modal numbers of fin rays, lateral line scales, gill-rakers and plesiomorphically low 
vertebral counts. No comparisons of the osteology of the various 'genera' given in the above 
synonymy have been made previously. I have compared the osteology of Rutilus rutilus with 
a selection of species currently assigned to this genus, to Scardinius, to Acanthobrama and to 
Mirogrex and am unable to find any noticeable differences. Indeed, all these taxa possess 
those apomorph characters which identify Rutilus, viz form of the median ethmoid notch; 
extension of the vomerine arms; preethmoid not, or barely contacting the mesethmoid; 
columnar neural complex; globular, papillate based gill-rakers. 

Scardinus is usually separated from Rutilus on the basis of its having the pharyngeal teeth 
in two rows instead of a single row, and the presence of a ventral body keel. I would consider 
that the differences in pharyngeal teeth are of a specific nature, as in Abramis (see Shutov, 
1967 and Eastman & Underbill, 1973); likewise, the ventral keel is variously developed in 
Rutilus rutilus. Wheeler (1976) concluded that of the features used to characterize Rutilus 
and Scardinius the morphology of the pharyngeal teeth was probably the most significant. If 
Chu's (1935) hypothesis is accepted, namely that three rows of pharyngeal teeth represent 
the primitive (plesiomorph) condition, then the two rows of Rutilus and the single row of 
Scardinius are seen as successively derived states. In the majority of Rutilus x 'Scardinius' 
hybrids a second tooth row is apparent which may indicate a dominant genetical pattern for 
the plesiomorph condition. 

Concerning the synonymy of Acanthobrama and Mirogrex; Goren, Fishelson & Trewavas 
(1973) separate these two genera on modal differences in the number of lateral line scales, 
gill-raker counts and the number of vertebrae. These authors compare the two 'genera' with 
Capoetobrama (see below), but not with Rutilus, and they state that the generic position of 
the presumed ancestral taxon to Acanthobrama, 'Rutilus' tricolor '. . . has not been 
determined'. I would agree that the species tricolor does not belong to Rutilus and should 
properly be placed amongst a complex including Leucaspius. However, Acanthobrama and 
Mirogrex share all the apomorph characters of Rutilus sensu stricto, they are thus to be 
regarded as belonging to that genus. The genus Capoetobrama (see above) does not belong to 
this assemblage at all and may be more closely allied to the alburnine group (unpublished 

Abramis Cuvier, 1817 

Ballerus Heckel, 1843 
5//cca Heckel, 1843 
Zopa Fitzinger, 1873 
Vimba Fitzinger, 1879 
Leucabramis Smitt, 1895 
Sapa Kazanskii, 1928 

Type species: A. brama (Linnaeus, 1 758). 

Five or six species with wide distribution in Europe, the Baltic, Black, Aral and Caspian seas; 

Asia Minor, Caucasus. 

Principal bibliographic source: Backiel & Zawisza, 1968. 

Shutov (1969) considered Blicca synonymous with Abramis. His analysis of populations of 
Blicca bjoerkna revealed a morphometric range encompassing that of Abramis brama and a 
decrease in the numbers of teeth in, and ultimate loss of the second tooth row across the 
range of the samples. Similarly, I find no differences in the osteology of Abramis and Vimba. 


Both genera possess an identical form of parasphenoid-basioccipital border with ventral 
channel and notch and highly developed pharyngeal tissue pads. They also share the 
apomorphic feature of a scaleless pre-dorsal fin ridge. Characters formerly used to separate 
these 'genera', such as the position of the anal fin origin and the presence of a dorsal ridge on 
the caudal peduncle are continuously variable and meaningless as taxon defining criteria. All 
Abramis species display considerable morphometric variability (see Berg, 1949 and 
Banarescu, 1963). 

Chondrostoma Agassiz, 1835 

Type species C. nasus Agassiz, 1835. 

Berg (1949) records 18 species, having a wide distribution throughout Europe south to the 

Tigris and Euphrates. 

Chondrostoma is characterized by those autapomorphic characters associated with 
forshortening and deepening of the upper jaw bones and a forwardly inclined coronoid 
process on the dentary. The posterior extension of the pharyngeal plate is thin and broad 
with a grooved dorsal surface. Assertions made concerning Chondrostoma in the anatomical 
descriptions presented here refer only to the type species, C. nasus. 


Earlier comments (Howes, 1976) that few, if any, presently recognized cyprinid subfamilies 
are monophyletic (in the sense of Hennig, 1966) have been amply reinforced by this study. 
Two genera, Ctenopharyngodon and Hypophthalmichthys previously included in the 
Leuciscinae are here identified as representative of two monophyletic lineages only distantly 

As noted above (p. 28), the presence of a maxillary foramen (or its suggested past presence) 
associated with a rostral barbel is considered apomorphic for one branch of a diphyletic 
division of the Cyprinidae. It is to this assemblage that the monophyletic lineage 
(squaliobarbine group) containing Ctenopharyngodon belongs. Hypophthalmichthys is 
contained within the abramine group and belongs to that assemblage of taxa lacking the 
maxillary foramen. It is the majority of these taxa which are provisionally recognized as the 
subfamily Leuciscinae. 

Nikolsky (1954) made a similar dichotomous classification of the Cyprinidae. In one 
branch he included leuciscines, Aspius '. . . and other fish', the other he represented by 
Barbus, Cyprinus and Gobio. Nikolsky did not give formal recognition to these divisions and 
retained the formerly recognized subfamilies without assigning them to one or other lineage. 

The following discussion is concerned mainly with the classification and intrarelation- 
ships of the non-barbelled cyprinids (Leuciscinae). 

Classification of the Leuciscinae 

The 'subfamily' Leuciscinae (originally a division introduced by Bonaparte, 1846) has long 
been used as a category for containing 'generalized' cyprinids, including approximately 40% 
of Old-World genera. Not all authors have accepted the Leuciscinae; Berg (1949) assigned 
the majority of Old-World genera to the Cyprininae. On the other hand, Hubbs & Lagler 
(1958) extended the geographical range of the Leuciscinae to embrace New- World genera. 
Gosline's (1978) concept of the Leuciscinae embraced also the Cultrinae. These decisions 
demonstrate a lack of methodology in determining cyprinid supraspecific relationships. 

As it stands, the Leuciscinae includes at least four monophyletic assemblages; aspinine 
(Howes, 1978), abramine (reported herein), alburnine and phoxinine (unpublished) all of 
which are interrelated to other non-barbelled cyprinids presently recognized as four mono- 
phyletic lineages; bariliine (Howes, 1980), cheline (Howes, 1979), hemicultrine (Howes, 
1978, part) and cultrine (Howes, 1978, part). It seems, however, that none of these groups is a 
likely candidate for the status of sister-group to the abramines. 

48 G. J. HOWES 

Identification of the abramine sister-group is hampered by the non-resolution of the 
monophyly of the nucleus of the Leuciscinae, the genus Leuciscus. It appears that part of 
Leuciscus is related to the aspinine group and another possibly forms the sister-group to the 
abramines. Characters supporting an abramine-Leuciscus relationship are the presence in 
some species of Leuciscus of an incipient ventral basioccipital channel and a similarity in the 
ethmovomerine region. 

Abramis has been thought by some authors to be related to the American cyprinid genus 
Notemigonus (Miller, 1958:203; Banarescu, 1973). Contrary to this idea I find no 
synapomorphies linking Notemigonus to any member of the abramine group. At this point it 
must be stated that New- World Cyprinidae have not been surveyed in depth in an attempt to 
formulate their relationships with any element of the Old-World Leuciscinae. Only 
Ptychocheilus has been examined in detail and would seem (together possibly with 
Algansea) to share apomorphies with the Old-World aspinine group (work in preparation). 
Whilst it may well be that New- World genera such as Notropis and Chrosomus are closely 
related to the Eurasian Phoxinus (see Mahy, 1975), it is quite wrong to consider them as 
synonymous (Banarescu, 1973). It must be borne in mind that Phoxinus as it stands is also a 
paraphyletic group and such is likely to be the status of Chrosomus and Notropis. 

Christopher & Linder (1979) note the close similarity between Chondrostoma and the 
American Acrocheilus. Certainly the jaw morphology of both taxa are alike but no other 
cranial features demonstrate a close relationship. 

At some level, of course, the New- World cyprinids must form sister groups of the Old- 
World. Whether they represent a single monophyletic sister group to one or other of the 
monophyletic lineages contained within the 'Leuciscinae' or comprise several lineages 
forming twin taxa with other non-barbelled Old-World groups (cultrines, chelines, etc) are 
theories which remain to be tested. 

Leuciscine hybrids 

Futuyama (1979) notes that '. . . typological thinking focuses first on the "pure" forms and 
then interprets intermediate phenotypes as secondary products. If we focused on the 
continuum of phenotypes as a unit, we might not worry as much about which are the species 
and which the hybrids'. 

Futuyama's statement is supported by a revised concept of leuciscine genera. Former 
hypotheses for the relationships between leuciscine genera have depended upon degree of 
genetic compatibility as demonstrated by hybridization. The measure of relationship being 
determined by terminal development stage of offspring, survival rate and reproductive 
viability. A contrary opinion regarding the validity of hybridization in a phylogenetic 
context has been expressed by Rosen (1979). Rosen regards reproductive compatibility as a 
plesiomorphy marked by mosaic distribution within a group. This hypothesis would 
certainly account for those conflicting cases amongst cyprinids where supposedly phylo- 
genetically closely related taxa fail and distantly related ones succeed in being reproductively 
compatible. Such inconsistencies are usually explained away as misidentification of parental 
stocks. Suzuki's (1968) explanation for one such conflicting result was that '. . . Some Soviet 
species of Leuciscinae are probably more closely related to Cyprininae than Japanese species 
of Leuciscinae'. Such a statement disregards any kind of rigorous methodology in elucidating 
phylogenetic relationships. 

A greater number of naturally occurring intergeneric hybrids have been reported for 
Leuciscine genera than for possibly any other group of teleosts (see Schwartz, 1972). A more 
rigorous (cladistic) definition of these genera reduces many cases of intergeneric to 
interspecific hybridization (ie. crosses between 'Vimba' 'Blicca'x Abramis (Berg, 1949), 
'Aristichthys'x Hypophthalmichthys (Voropayev, 1969), 'Scardinius'x Rutilus (many 
reports, see Schwartz, 1972). Naturally occurring intergeneric hybrids are judged to be so on 
the basis of intermediacy of characters, notably the pharyngeal dentition. A typical example 
of the use of 'intermediate' characters is given by Berg (1949 : 88-89) for a 'hybrid' popula- 


tion of Leuciscus cephalus x Chalcaburnus chalcoides. As enumerated, these characters 
could well describe a populational variant of one of the presumed parental species. 

A similar argument can be applied to many so-called interspecific hybrids. The likelihood 
is that they are not crosses but clinal or other populational fragments. The numerous sub- 
species and varieties described in Leuciscine genera by Berg (1949) indicate a considerable 
degree of populational variability. This of course is not to deny that interspecific hybridiza- 
tion does occur. 

Wheeler (1976) notes that high levels of hybridization amongst cyprinid taxa occur in 
areas where recent introductions have been made. Certainly, nearly all the records I have 
checked for 'intergeneric' European cyprinid hybrids have been cases involving one or both 
of the taxa being recently introduced into a 'new' ecological situation. 

I conclude that 'hybridization' amongst cyprinid taxa must be treated with caution and is a 
useless character as a measure of phylogenetic affinity. 

Classification of the 'true' carps 

The squaliobarbine group belongs to the great assemblage of taxa characterized by the 
presence of a maxillary foramen and (usually) an associated rostral barbel (see p. 28). As yet 
no accounts of monophyletic groups, other than the squaliobarbines (dealt with herein), have 
been published, although Reid (1978, unpublished thesis) has identified a labeoine group. 
The hypothesis proposed above (p. 40) that the squaliobarbines form the sister group to all 
other barbelled carps is a temporary expedient. But, with available data I am unable to offer 
a more worthwhile solution. Such genera as Capoeta, Cyprinus and Carassius appear to be 
more closely related to the squaliobarbines than do other investigated taxa. Work in prepara- 
tion (Banister & Howes) seeks to establish monophyletic grouping amongst the barbelled 
cyprinids. In view of the suspected non-monophyly of so many currently recognized 
cyprinid assemblages, the reconstruction of ichthyogeographical relationships (eg. 
Banarescu, 1973) is fruitless. 


I am most grateful to my colleagues in the Fish Section of this museum for their help and 
advice. Particular thanks are due to Drs P. Humphry Greenwood and K. E. Banister for their 
critical comments of the manuscript and discussions of cyprinid phylogeny; to Alwynne 
Wheeler for his helpful comments concerning British cyprinids; to Margaret Clarke for her 
help with synonymies and literature searches; to Robert Travers for preparing alizarin 
specimens; and to Alice Parshall for providing me with unpublished information on cobitid 

Sara and Dr William Fink generously allowed me to see their unpublished manuscript on 
ostariophysan anatomy. 

To Dr Chu Xin-Luo of the Kunming Institute, China go my warmest thanks for supplying 
many small specimens of Hypophthalmichthys. 

Finally, my thanks to Gina Sandford for typing the manuscript. 


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Manuscript accepted for publication 28 November 1 980 

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Morphology and systematics of some interior- 
walled cheilostome Bryozoa 

P. L. Cook and P. J. Chimonides 

Department of Zoology, British Museum (Natural History), Cromwell Road, London 
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Synopsis 53 

Introduction 53 

Material and methods 54 

Measurements and abbreviations used 57 

Key to species described 58 

Systematic section 58 

Family Euthyrisellidae Bassler 58 

Family Didymosellidae Brown 68 

Discussion 81 

Summaries in French and German 83 

Acknowledgements 85 

References 85 


The colony structure and zoodial morphology found in two cryptocystidean cheilostome bryozoan 
Families, the Euthyrisellidae and the Didymosellidae are described. All calcified walls are interior, and 
colony-wide (extrazooidal) basal coeloms are present in nearly all species. In the Family 
Euthyrisellidae, interzooidal visceral-to-hypostegal communication complements or even replaces the 
more usual, intrazooidal kind, which is that found in the Family Didymosellidae. In the Euthyrisellidae 
an extension of the hypostegal coelom on the basal side of the calcified frontal shield is present in most 
species. One new species, Tropidozoum burrowsi, is introduced. The Family Didymosellidae has a 
fossil record extending from the Middle Eocene to the Recent, and a wide Tertiary-to-Recent distri- 
budon. One new Recent species, Didymosella pluma, is introduced, and Tubiporella boninensis Borg is 
redefined. Tubiporella levinseni Borg is redescribed and assigned to the umbonuloid ascophoran genus 


The calcified and uncalcified body walls defining the member zooids of bryozoan colonies 
may conveniently be regarded as belonging to two kinds. The first is the exterior, limiting or 
boundary wall, which is primarily cuticular and which, if calcified, almost invariably has an 
outermost layer of cuticle throughout ontogeny. This kind of wall is actually or potentially 
capable of dividing the internal coelomic tissues of zooids from the environment. Zooids 
delimited by this kind of wall develop as uncalcified buds by expansion and extension of 
coelomic tissue through septulae in the calcified wall of an existing zooid. The boundary 
wall of cuticle and epidermis expands intussusceptively as growth proceeds. The earliest 
known, Late Jurassic cheilostomes had zooidal walls almost entirely of this kind (see 
Boardman & Cheetham, 1973), and large numbers of species now living have the same, or 
similar zooidal structure and budding pattern. Many lateral vertical walls of contiguous 

Bull. Br. Mus. nat. Hist. (Zool.) 41 (2) 53-89. Issued 24 September 1 98 1 



zooids in expanses of more integrated species are also exterior walls, even though they may 
appear to be "inside' the colony. The second kind of wall is the interior wall, which may or 
may not possess any cuticular element, and which partitions existing coeloms and is not 
directly in contact with the environment. Zooids with interior walls are differentiated from a 
colony-wide expansion of coelom bounded by intussusceptively expanding cuticle and 
epidermis, by growth of a calcified lamina which, in earlier stages of ontogeny at least, is 
surrounded by coelom (see Boardman & Cheetham, 1973). Both kinds of wall have arisen 
several times in the long evolutionary history of the Bryozoa, and both kinds are usually to 
be found in any cheilostome colony. In some colonies, the distinction between the two kinds 
of wall is not clear cut. However, the astogenetic and ontogenetic sequences reflected by 
interior and exterior walls, and the intrazooidal and interzooidal communications associated 
with them, are important in the study of colony structure, of integration, and of systematic 
relationships through time. 

Colonies with a high proportion of their calcified walls of the interior kind are well 
known among the 'free-living' anascan cheilostomes (Hakansson, 1973, Chimonides & 
Cook, 1981). Like these, several ascophoran Families, such as the Conescharellinidae, and 
genera such as Sphaeropora, as well as the Euthyrisellidae and Didymosellidae, have 
virtually no exterior calcified walls at all (Cook & Lagaaij, 1976, and Cook & Chimonides, 
1981). The ascophoran forms show a much larger range of colony morphotypes than the 
anascan forms. 

The species of Euthyrisellidae and Didymosellidae have cryptocystidean frontal shields 
(see Banta, 1970). These frontally placed parts of zooids in which the orifice is situated, are 
interior calcified walls which partition the visceral from the hypostegal coelom. The exterior 
frontal wall above the hypostegal coelom is completely cuticular, with underlying epidermal 
cells. In some species (e.g. Pleurotoichus clathratus) the frontal, cuticular wall is a 
continuous, colony-wide expanse, connected only intermittently with the vertical walls and 
the frontal shields. The lateral and distal vertical calcified walls are also interior walls, which 
partition a colony-wide coelom at the growing edge or tip of an expanse or branch. Basal 
calcified walls are also interior walls in the majority of species described here. These too, 
have communication with an extrazooidal basal coelom, which is protected from the 
environment by a cuticular and epidermal wall. 

Intrazooidal, interzooidal and extrazooidal communication is an important feature in 
these colonies (Figs 1A, B) and its skeletal expression, in the form of septulae, and often in 
subsequent thickening of calcification, enables inferences to be made for fossil and Recent 
specimens in which the cuticular walls and coelomic tissues are no longer present. 

Material and methods 

The possibility of restoring recognizable structures, including muscles and viscera, in 
previously dried material, and the advantages of study of epoxy-resin mounted thin-sections 
of 'hard' and 'soft' tissues in situ have been demonstrated (Cook, 1975). Dried specimens 
were previously soaked in 10% trisodium phosphate solution for 48 hours, gently cleaned in 
detergent solution, and rinsed in water before staining. The accession of fixed, well-preserved 
material of Pleurotoichus, Tropidozoum, Neoeuthyris and Didymosella has enabled us to 
make sections which may be compared with those made from restored material. This com- 
parison has tested the inferences drawn previously, and will allow further inferences to be 
made with greater confidence in the future. 

Specimens for scanning electron microscopy were treated with sodium hypochlorite, 
rinsed and coated with gold-palladium. Specimens to be thin-sectioned were heavily stained 
in borax carmine or safranin, impregnated with epoxy-resin, cut and polished attached to 
3" x 1 " microscope slides. 

Early attempts at embedding followed Nye el al (1972) and Sandberg (1971). Three resin 
types were available: Reichold Epotuf 37 128 with hardener 37 614, Shell Epon 812 and 




Fig. 1 Generalized diagrams to show residual communication after development of interior 
calcified walls in: (A) Euthyrisellidae; (B) Didymosellidae. Intercoelomic communications 
arrowed: h hypostegal coelom, sf subfrontal hypostegal coelom, v visceral coelom, b basal 


Ciba Araldite CY212 with their complementary additives MNA, DDSA, BDMA, DMP 30 

The Reichold resin, well tested by Nye et al (1972) was found to be difficult to use. The 
high viscosity of the components hindered obtaining the intended ratio of 4 parts resin to 1 
part hardener and their subsequent mixing. The mix was found to be sensitive to heat and 
U.V. light (e.g. sunlight) and there was a tendency for thermal runaway to occur. This 
reaction was more readily provoked if the mix contained slightly excess hardener, even 
though the plug size was only 30 cm 3 and curing was initiated at room temperature, 20C. It 
was found that surface temperatures in excess of 80C were reached, coupled with emission 
of phenol-smelling fumes, bubbling and volumetric expansion. When curing could be con- 
trolled by refrigeration a volumetric contraction, estimated as 8% total, was noted. No 
significant change in behaviour was noted with prolonged storage of the components at 
5-10C. The recommended life of hardener 37 614 is given as 6-12 months in a tightly 
closed container at room temperature or less. Denaturing is characterized by progressive 
discolouration to a dark brown colour. Epon 812 was used as described by Sandberg (1971) 
using only MNA as hardener. DMP 30 and BDMA (Glauert, 1965, 1974) were used as 
accelerators in parallel series. Araldite CY212 was used in the same way. The preferred 
ratios of the components were: resin 54%, hardener 44% and accelerator 2%. Both resins 
were found to have tolerant curing temperatures of 50-80C. Epon 812, with its low 
viscosity and production of relatively clear and hard plugs promised to be the ideal medium. 
It was found, however, that as lapping of thin-sections progressed, small, granular particles of 
resin (and sometimes of specimen) would break away leaving score marks on the polished 
surface. This was more noticeable as the thickness of the preparation reached 40 u or less. It 
was almost impossible to produce an intact section below 20 ji. Freeman & Spurlock (1962), 
in electron microscopical studies, reported Epon 812 as being inherently granular. This 
suggests that thin-section disintegration may be a larger scale manifestation of the same 
characteristic. The disintegration problem was more pronounced at the lower curing 
temperatures. In addition, it was found that the resin in sections became 'milky' in aqueous 
solutions (even without the aquon component), or in the lapping lubricant used, Engis 
Hyprez type W. The resin would also accept stains which were difficult to remove. Further, 
as the section was thinned, it would distort and lift from the slide. These effects were noted 
regardless of resin and additive batch and of precautions to ensure uniform mixing of com- 
ponents and cleanliness of the glass slides. A noticeable volumetric contraction (of about 
6-7%) was again found, regardless of recipe and of curing procedure. 

Araldite CY212, in spite of its high viscosity, was found to be the most suitable resin. Its 
total curing contraction was estimated as 1-2%, regardless of recipe. Stability during 
lapping was high, and it was possible to obtain sections in the order of 3 u thickness. One 
disadvantage was that xylene affected the cured resin and coverslides mounted with Canada 
balsam were therefore avoided. Where required, coverslides were mounted using more 

In all cases propylene oxide was used as the antemedium. Final embedding was vacuum 
assisted after first soaking the specimens in an equal mixture of propylene oxide and 
accelerated resin mix. One well known advantage of the use of propylene oxide is that any 
residual quantities trapped in the specimen should polymerize with the resin. 

In all resins used, behaviour appeared equally divided between pre- and post-gel 
contraction. Reichold Chemicals Inc. (pers. comm. 16.2.1977) suggested that a more typical 
contraction behaviour of their resin would be approximately 5% total, distributed as 3% 
pre-gel and 2% post-gel phase. Potter (1970 : 171) identified post-gel shrinkage as the phase 
at which stresses may be introduced which can lead to damage, although his account relates 
to electrical component encapsulation and is without reference to scale. It was indicated that 
post-gel contraction for epoxy resins is normally in the region of 2-3% and that contraction 
overall is affected primarily by the accelerator used. 

In making thin-sections the effects of resin contraction were most apparent when attaching 
preparations to microscope slides, when bowing of the glass, and in many cases, even 


breaking was noted. The amount of resin was therefore kept to a minimum, and Araldite 
resin obviously showed to advantage. 

Within the specimen itself, shrinkage should have little effect providing that complete 
penetration by the resin has been achieved. Although penetration may appear to be com- 
plete, it has been found, during cutting and polishing stages, that calcified walls may drop out 
of the section completely, as opposed to partial fracturing and partial loss. This indicates a 
lack of specimen support. 

If a final surface stain was applied, there was a tendency for the stain to delineate the 
boundary of the calcification, although it should be borne in mind that the micro-relief 
produced by lapping may have been a contributory factor. In sections of 10 u thickness or 
more, the stain sometimes soaked into the boundary irregularly. Support failure and 
staining characteristics suggest an unpredictable penetration of the resin into the calcifi- 
cation, together with a degree of incomplete adhesion which may be exacerbated by resin 
shrinkage. The evidence is at present considered to be inconclusive, especially as the effects 
were observed most often when using the structurally unsatisfactory Epon 812 resin. For 
these reasons staining before embedding was preferred. 

Measurements and abbreviations used 

The value of comparative measurements among, for example fossil and Recent colonies of 
the same or different species, varies considerably among bryozoans, both with the structures 
measured and with the species groups being investigated. For example, some species retain 
recognizable traces in frontal view, of structures such as the tops of vertical walls, through- 
out ontogeny and in spite of wear. In such species comparisons of zooidal length and width 
among colonies are easily made and are significant. Primary calcified orifices with distinct 
boundaries are also capable of yielding reasonably accurate measurements. In the species 
considered here, one family, the Euthyrisellidae, which has no fossil record, retains 
throughout ontogeny recognizable features which allow morphometric analysis. The 
Didymosellidae, on the other hand, includes species in which continuous ontogenetic 
changes throughout colony life obscure zooidal boundaries and primary orifices, which in 
themselves are not easily definable. Fossil (and some Recent) specimens, which have been 
damaged or are worn, are therefore not strictly comparable with well preserved colonies. 
Other micro-environmental effects, such as those influencing the length of peristomes, may 
sometimes be recognized, but are difficult to quantify. 


Measurements in millimetres 

Length of autozooid Lz Width of autozooid lz 

Length of brooding zooid Lbz Width of brooding zooid Ibz 

Length of primary orifice Lo Width of primary orifice lo 

Length of brooding z. orifice Lbo Width of brooding z. orifice Ibo 

Length of rostrum Lr Width secondary orifice 12o 

Length of mandible Lm 

BMNH British Museum (Natural History) MM Manchester Museum 

NMV National Museum of Victoria AM Australian Museum, Sydney. 

ZMA Zoological Museum, Amsterdam SAM South African Museum, Cape Town. 


Key to species described 

1 Raised peristomes absent, opercula well-developed, often with two layers and with 
complex sclerites. Frontal pores and septulae absent or rare; large frontal foramina present 
or absent. Avicularia and ovicells usually absent. Secondary calcification slight. Brooding 

zooids strongly dimorphic EUTHYRISELLIDAE 2 

Raised peristomes present, opercula simple, not usually visible. Frontal pores and septulae 
numerous. Avicularia and ovicells present. Secondary calcification extensive. Dimorphic 
zooids not obvious DIDYMOSELLIDAE 6 

2 Colonies encrusting, avicularia and ovicells present, basal coelom absent 

Neoeuthyris woosteri p. 67 

- Colonies erect. Avicularia and ovicells absent, basal coeloms present .... 3 

3 Colonies flustriform, unilaminar, with lateral and basal coeloms 4 

Colonies cellariiform, with kenozooidal nodes. Internodes with zooids in curved, laterally 
contiguous series. Basal coelom forming a central column . . . Tropidozoum 5 

4 Frontal shield complete, smooth. Lateral coeloms with series of calcified plates 

Euthyrisella obtecta p. 60 

Frontal shield with calcified bars and large, slit-like foramina. Lateral coeloms without 
series of calcified plates Pleurotoichus dathratus p. 63 

5 Autozooidal orifices not sinuate, frontal shield with large foramina 

Tropidozoum cellariiforme p. 64 

- Autozooidal orifices with a deep, triangular sinus, frontal shield with small foramina 
T. burrowsisp.nov. p. 65 

6 Peristomes with large, paired proximal foramina, spiramen absent . Didymosella 

- Peristomes with one, or no foramen, spiramen present . . . . Tubiporella 13 

7 Colonies erect, ligulate, unilaminar. Zooids in 2-5 transverse rows. Basal pores 
present 8 

- Colonies encrusting, semi-encrusting or repent, or bilaminar or cylindrical ... 9 

8 Branches flat, 4-5 zooids wide, basal pores numerous Didymosella porosa p. 72 
Branches laterally curved towards the basal side, 2-3 zooids wide, basal pores rare 

D. clypeata p. 72 

9 Colonies bilaminar or cylindrical D. /arva/w-complex (fossil populations) p. 73 
Colonies unilaminar or semi-encrusting, with large basal foramina, or basal pores and/or 
calcified processes 10 

10 Colonies encrusting or semi-encrusting. No coelom basally, but zooid walls uncalcified or 
with a large, central foramen. Peristomes deficient distally, raised proximally 

D. /arva/w-complex (Recent populations) p. 73 
Colonies repent, with basal pores and/or long calcified processes. Peristomes tubular . 1 1 

1 1 Zooids large Lz>0'70 mm), avicularian rostra with an open channel terminally 12 
Zooids smaller (Lz < 0'70 mm), avicularian rostra very small and acute parviporosa p. 75 

1 2 Zooids with more than 25 frontal pores. Basal surface with pores (fossil) D. crassa p. 7 1 
Zooids with less than 25 frontal pores. Basal surface with long, calcified processes. 
Avicularian mandible expanded laterally (Recent) . D. pluma sp. nov. p. 76 

13 Colonies forming broad, repent expansions. Peristomes with a single foramen early in 
ontogeny. Avicularia with acute, hooked mandibles Tubiporella magnirostris p. 77 

- Colonies ligulate, 4-7 zooids wide, repent. Peristomes entire, very long and tubular. 
Avicularia elongated, with laterally expanded mandibles . T. boninensis p. 79 

Systematic Section 

Euthyrisellidae Bassler, 1953 : G226. 
TYPE GENUS. Euthyrisella Bassler, 1936. 

DESCRIPTION. Interior walled cryptocystidean ascophorans, usually with extrazooidal, 
cuticle-bounded basal coeloms. Colonies encrusting algae, or erect, flexible, rooted. Zooids 


Table 1 Average measurement (mm) of zooids of Euthyrisellidae. 

Lz Iz Lbz Ibz Lo lo Lbo Ibo 

E. obtecta 









P. clathratus 









T. cellariiforme 









T. burrowsi 









N. woosteri 








0:3 1 

with relatively little secondary calcification frontally, frontal septulae restricted or absent. 
Hypostegal coeloms extensive, often extending basally to the calcified frontal shield, and 
communicating interzooidally with visceral coeloms of zooids of the previous astogenetic 
generation. Externally obvious brooding structures and avicularia usually, but not ex- 
clusively absent. Orifices of brooding zooids enlarged, dimorphic. 

REMARKS. The family was introduced by Bassler (1953) for the genera Euthyrisella, 
Pleurotoichus, Neoeuthyris and Urceolipora. The relationships among these genera have 
been discussed by Levinsen (1909), Harmer (1957) and Hastings (1964). Urceolipora shows 
many characteristics in common with Euthyrisella and Neoeuthyris on the one hand, and 
with the genera Calwellia and Icthyaria on the other. These generic groups, together with 
others which have some similarities in their colony organization, such as Onchoporella and 
Euthyroides all require detailed analysis, but are beyond the scope of this paper. Urceolipora 
is therefore excluded here, but the genus Tropidozoum is included in the family. 

ONTOGENY OF ZOOID WALLS. In all species, the primary ontogenetic changes in calcification 
of the zooid walls and formation of the orifice, operculum and ascus, appear to be very rapid, 
as few intermediate stages between undifferentiated buds and complete zooids are present. In 
addition, growth appears to proceed in distinct episodes, alternating with periods of 
astogenetic quiescence, as many colonies show little evidence of any zones of developing 
zooids at the growing edges or tips, and some colonies show distinct 'growth lines' (see 

Where present, the growing zones consist of an undifferentiated area of coelomic tissue 
bounded by thin cuticle. The calcification of interior walls proceeds distally, and the lateral 
walls become intermittently attached to the frontal cuticle. At the same time the opercular 
cuticle becomes thickened and differentiated. Where there is a basal coelom, the calcified 
lateral walls converge, forming a basal zooid wall which may be linear, or merely a curved 
continuation of the lateral walls. Contact with the cuticular, basal colony-wide wall occurs 
later in ontogeny in some species. The development of the lophophore and viscera is com- 
plete 2-3 zooid generations from the growing zone, just in advance of the final stages of ascus 
development (see also Harmer, 1902). 

In Tropidozoum, particularly T. burrowsi (see p. 65, there is some subsequent 
thickening of the calcification of the frontal shield and basal walls. In Euthyrisella, Pleuro- 
toichus and Neoeuthyris, thickening is confined to the development of processes attaching 
the cuticle to the frontal shields, or to the development of calcification within the lateral 
extrazooidal coeloms, when present. 

Coelomic systems and communication 

The intercommunication among the hypostegal, visceral and extrazooidal coeloms is con- 
siderable, and reflects the varying extent and patterning of these coeloms throughout the 
colony (Fig. 1A). The virtual isolation of the hypostegal coelom, particularly in 
Tropidozoum, is a direct result of the rarity of the frontal septulae (areolae) usually found in 
cryptocystidean ascophorans, and is accompanied in most species by an extension of this 
coelom beneath the calcified frontal shield (see also Cook, 1975). Transfer of nutrients in 


coelomic fluids is inferred by the presence of communication pores in the distal-terminal or 
distal-lateral calcified walls, which connect the visceral coelom of one zooid with the 
hypostegal coelom of the next successive, distal zooid. This system, too, is unusual in 
cheilostomes, although it probably occurs in the genera Urceolipora, Calwellia and 
Onchoporella as well (see above). The basal and lateral extrazooidal coeloms are extensive in 
all the erect genera, but absent in Neoeuthyris, which is encrusting. Where present, they are 
inferred to function connectively and, as their cuticular walls thicken with ontogeny, to 
become involved in the anchoring and support of the colony. 


The opercula are large and double layered, with strengthening sclerites in most species. The 
morphology of the complex opercula of Euthyrisella and Pleurotoichus has been fully 
described by Harmer (1902). Those of Tropidozoum and Neoeuthyris are simpler in con- 
struction, but also have strong sclerite systems. 

Brooding zooids 

All species show dimorphism of brooding zooids, the larger dimorphs having enlarged 
orifices and, in the erect species, swollen basal walls. Embryos have now been observed in all 
species and without exception, are associated with the larger dimorphs. They are not readily 
seen except by transmitted light (cf. Harmer, 1902), but when present, usually fill the central 
and proximal parts of the cystid and are presumably contained in ovisacs. 

The ovicells of Neoeuthyris differ considerably in appearance from the brooding zooids of 
the other genera. However, the brooding cavity may be regarded as a displacement of the 
swollen basal cavity in a distal and frontal direction. This is correlated with the encrusting 
mode of life, which does not allow basally directed expansion. 


Euthyrisella Bassler, 1936 : 161 (for Euthyris Hincks preocc.) 
TYPE SPECIES. Euthyris obtecta Hincks, 1 882. 

DESCRIPTION. Colony flustriform, branching, multiserial and unilaminar, with basal and 
marginal extrazooidal coeloms bounded by cuticular walls, which become thickened to form 
anchoring, rooting structures, with internal calcified plates late in ontogeny. Zooids with 
depressed, curved, thinly calcified frontal shields, with few pores. Lateral walls converge 
basally and are attached to the basal cuticular wall by intermittent calcified processes. 
Dimorphic brooding zooids inflated basally, with enlarged orifices and opercula. 

Euthyrisella obtecta (Hincks) 

Euthyris obtecta Hincks, 1882 : 164, pi. 7, fig. 3, 1893 : 177. Harmer, 1902 : 277, pi. 16, figs. 32-37. 
Levinsen, 1909 : 272, pi. 15, figs 2a-f. Canu & Bassler, 1929 : 464, Figs 199 A-M. 

Figs 2-7 Scanning electron micrographs of Euthyrisellidae: (2) Euthyrisella obtecta (Hincks) 
BMNH 1890.3.24.22, Torres Straits, lateral view of zooids showing lateral and oral septulae 
and basal processes x65; (3) E. obtecta, zooids viewed from distal end, showing distal septulae 
(arrowed) and frontal processes x!20; (4) Tropidozoum cellariiforme Harmer BMNH 
1964.3.10.1, Celebes, frontal view of primary zooids of internode, showing calcified tube at 
distal end of nodal kenozooid x72; (5) T. burrowsi sp. nov. BMNH 1975.1.4.1, Madagascar, 
basal view of proximal part of internode, showing autozooids and basal coelomic column, with 
basal septulae and bars of calcification x30; (6) T. burrowsi, frontal view of autozooids and 
brooding zooid x53; (7) Neoeuthyris woosteri (MacGillivray) BMNH 1979.2.1.1, Western 
Australia, frontal view of autozooids and avicularia x30. 




SPECIMENS EXAMINED. BMNH. North Australia, 1897.5.1.445 (possibly type material); 
Broughton Id., New South Wales, 1883.11.29.3; Port Denison, East Queensland, 
1883.11.29.43; Murray Id., Torres Straits, 1890.3.24.22, 16-36 metres; 1890.7.23.38, 
17-36 metres; 1916.8.23.115, 9-36 metres; Baleine Bank, Northwest Australia, 
1892.1.28.34A, 17-22 metres; Holothuria Bank, Northwest Australia, 1892. 1.28.61 A, 
22 metres; Long Reef, 5 miles North of Sydney, New South Wales, 1975.1.2.3, 101 metres. 

DESCRIPTION (Figs 2, 3, 33, 35). Euthyrisella with zooids intercommunicating by 6-8 lateral 
septulae (8-10 in brooding zooids) and 5-6 distal septulae. Zooids with 12-20 basal septulae 
(45-50 in brooding zooids) and 2-8 septulae distal to the orifice, which communicate with 
the hypostegal coelom of the next distal zooid. Frontal shield thinly calcified, with 2-5 pores 
proximal to the orifice (10-15 in brooding zooids), some of which are septulae communi- 
cating with the part of the hypostegal coelom basal to the frontal shield. Orifices with small 
lateral condyles. Dimorphic (brooding) zooids about twice the size of the autozooids. 

REMARKS. The hypostegal coelom beneath the frontal shield is not nearly as extensive as in 
Pleurotoichus and Tropidozoum (see below), but forms a small expansion proximally to the 
orifice (Fig. 35). The dimorphic brooding zooids occur in a proportion of approximately 
1 : 20 autozooids. Large embryos are present in one colony from Torres Straits 

The colonies are profusely branched and reach a height of 1 50 mm. The branches are 
2-3 mm wide and are usually formed by simple bifurcation, but some, astogenetically late, 
lateral branches occur. Before a bifurcation the width increases to 4 mm, and the number of 
zooids in a transverse series from 3-4 to 5-6. There is a tendency for one branch of a bifur- 
cation to include more zooids than its pair. At irregular intervals there is an increase in the 
width which does not result in a bifurcation, but a sudden decrease in zooid numbers 
between successive generations. This seems to be associated with a period of cessation of 
growth, as the basal cuticle is much thicker on the proximal side of the interruption. 

The early astogeny has been inferred from one colony (1883.11.29.3). A single, erect 
kenozooid is followed by a distal zooid which itself buds one distal and paired distal-lateral 
zooids. Thereafter, increase in zooid rows originates from bifurcations of series distal to 
marginal zooids. At the base of fronds zooids have the series: Lz 0'50, 0'50, 0'67, 0'50, 0'53, 
0'58, 0'60 mm. The only other astogenetic changes are confined to brooding zooids which do 
not appear until the branch is 100 mm in length (300 zooid generations). Brooding zooids 
are central and often paired in the same transverse series. 

The extrazooidal marginal coelomic systems are extensive and resemble those formed by 
lateral kenozooids in several anascan, flustriform genera and the ascophoran species 
Euthyroides episcopalis and Onchoporella buskii. The changes during colony growth 
may be regarded as ontogenetic, in that they occur within the bounding walls of an existing 
structure. The primary changes are found 40 zooid generations back from the growing tip. 
The lateral cuticle expands intussusceptively and thickens considerably. About 1 50 zooid 
generations back, calcareous spicules derived from the lateral walls of the marginal zooids 
expand and form a series of plates within the lateral coeloms. Subsequent series of plates, 
alternating with cuticle, are formed at the base of the colony, and the investing cuticular wall 
continues to expand and thicken (see Harmer 1902). At the earliest astogenetic level there 
are 9-12 series of thickened plates on each side of the branch. The growth of anchoring 
rootlets is also an ontogenetic rather than an astogenetic change. The marginal elements 
extend basally and divide into rhizoids which penetrate the substratum, incorporating 
particles of sediment within a meshwork. Some calcification occurs so that the rhizoids are 
not only flexible but adherent. It is possible that further extensions are astogenetic and 
consist of kenozooids budded from the extrazooidal coeloms. It is interesting that an almost 
identical series of calcareous plates occurs in the kenozooidal, lateral rooting systems of 
O. buskii. The rooting system of the colony (1883.11.29.3) has incorporated the 
substratum of shell fragments, sand grains, foraminiferans and dead bryozoans. Although 
these last are worn, and may have been transported, it is interesting that they comprise 



Figs 8 and 9 (8) Pleurotoichus clathratus (Harmer) BMNH 1976.1.2.1, Heron Island, 
sketch of frontal surface of autozooids and brooding zooids, showing opercula and irregular bars 
of calcification of frontal shield beneath the cuticular frontal wall; (9) Didvmosella larvalis 
(MacGillivray) BMNH 1890.4.17. l-10pt., Spencer Gulf, sketch of ancestrula. Scale bars = 
0-50 mm. 

rooted 'sand fauna' species (see Cook, 1965, 1966, \919b). They include colonies of 
Conescharellina sp., the internodes of jointed species of cyclostomes and of the cheilostome 
anascan, Exostesia didomatia (see Brown, 1948 : 111). Other epizoic forms include a ramify- 
ing hydroid and numerous egg sacs of some unknown animal. 


Pleurotoichus Levinsen, 1909 : 270. 

TYPE SPECIES. Euthyris clathrata Harmer, 1902. 

DESCRIPTION. Colony flustriform, branching, multiserial, unilaminar, with marginal and 
basal extrazooidal coeloms. Zooids with flat, depressed frontal calcified shields consisting of 
irregular bars and foramina, each bar surrounded by cuticle. Lateral walls curved and 
converging, basal walls linear, reaching the extrazooidal basal wall in discontinuous series. 
Dimorphic zooids slightly larger than autozooids, with larger opercula, inflated basally. 
Lateral coeloms slightly calcified late in ontogeny. 

Pleurotoichus clathratus (Harmer) 

Euthvris clathrata Harmer, 1902 : 266, pi. 16, figs 18-31. 

Pleurotoichus clathratus : Levinsen, 1909 : 270. Canu & Bassler, 1929 : 464, Figs 200 A-N. 

SPECIMENS EXAMINED. BMNH. Australia, 1899.7.1.2606; Port Jackson, New South Wales, 
1903.5.1.2, sublittoral; Swan River, South Australia, 1935.10.18.1; Middle Harbour, 
Port Jackson, 1975.8.1.7; Watsons Bay, 1975., under rock ledges, low tide; Heron 
Id., Queensland, 1976.1.2.1. 

DESCRIPTION (Figs 8, 34). Pleurotoichus with zooids communicating by 7-9 lateral and 
7-10 distal septulae. Zooids with 12-15 basal septulae, and 3-5 distal to the orifice, 


communicating with the hypostegal coelom of the next distal zooid. The frontal shield is 
formed by irregular cuticle-covered bars, which anastomose, leaving large slit-like foramina. 
The hypostegal coelom is extensive both above and below the shield. The dimorphic 
brooding zooids are only slightly larger than the autozooids; they are inflated basally, and 
occur in larger numbers than in E. obtecta, in a proportion of 1 : 5 autozooids. 

REMARKS. The presence of cuticle surrounding the bars of calcification of the frontal shield, 
and the layers of cuticle on either side of the lateral and distal calcified walls, together with 
the general delicacy of the interior structures, makes it almost impossible to prepare calcified 
skeletons of P. clathratus for scanning electron microscopy. Investigation of specimens has 
therefore been almost entirely confined to examination of thin-sections. The hypostegal 
coelom, both frontal and basal to the calcified shield is very extensive. The convergent 
lateral walls reach the basal cuticular extrazooidal wall in one short, linear insertion. The 
calcified processes are cuticular terminally at the point of contact. Zooids are thus almost 
suspended from the circum-orificial region in a bag of tough, flexible cuticle. This flexibility 
is probably correlated with the turbulence of the very shallow waters in which the species 
lives. The growth form is analogous to that of Chartella papyracea (see Ryland & Hay ward, 
1977 : 80), which also inhabits rocky ledges in the sublittoral region. 

Colonies may reach a height of 185 mm (Harmer, 1902 : 268). The specimens examined 
have a maximum height of 50 mm, and have very few bifurcations. A few lateral sub- 
branches occur late in astogeny. The earliest stages are missing, but may be inferred from a 
colony from Port Jackson (1975.8.1.7), which has 4 zooids in a transverse row at its base. 
Extrapolating from this, the early astogeny is probably similar to that of E. obtecta, and the 
zooids show a similar series of increase in size. Interruptions of growth, involving a sudden 
decrease in the number of zooids in transverse series from 6-7 to 3-4, occur frequently, 
between 6 and 1 zooid generations apart. 

Dimorphic zooids are frequent (Fig. 2), and are first found 25 generations from the base of 
the colony. They are placed in the central zooid series, and large embryos are present in the 
colony from Heron Island (1976. 1.2.1.). 

The marginal coeloms become inflated with ontogeny, and the cuticular walls become 
thickened. Small calcified processes develop from the lateral walls of adjacent zooid series, 
but calcified plates do not occur. 

The colony from Heron Island has a large number of epizoic forms, particularly colonies 
of hydroids. On the basal cuticle there are small colonies of jointed cyclostome bryozoans 
and ofCellaria sp. The lower part of the colony is completely invested by a colonial ascidian. 

Tropidozoum Harmer, 1957 : 1 106. Cook, 1975. 
TYPE SPECIES. T.cellariiforme Harmer, 1957. 

DESCRIPTION. Colony cellariiform, internodes connected by complex nodes of partially 
calcified kenozooids and investing extrazooidal basal coelom. Basal coelomic channel 
confluent among internodes. Central zooids of internodes with extended basal calcification 
forming a keel. Zooids with depressed frontal shields, with large foramina. Hypostegal 
coelom extensive, frontal septulae absent. Brooding zooids large, inflated basally, with 
dimorphic orifices. 

Tropidozoum cellariiforme Harmer 

Tropidozoum cellariiforme Harmer, 1957:1106, pi. 67, figs 15-17, 20. Cook, 1975:162, pi. 2, 
pi. 3, Figs 2A, 3 (explanation of plates 1 and 3 transposed). Cook, 1979a : 200, pi. l,fig. 1. 

SPECIMENS EXAMINED. BMNH. Siboga Stn 144, South of Halmaheira, Djilolo, East Celebes, 
0-45 metres, coral and lithothamnion bottom, 1964.3.10.1 and 1979.1.4.1. 
ZMA, as above, TYPE and other material. 


DESCRIPTION (Figs 4, 1 1, 30). Tropidozoum with internodes 8 mm long and l - 6 mm wide, 
comprising 70 zooids. Longest, usually the central series, of 12 zooids; widest, usually the 
penultimate transverse series, of 6 zooids. Zooids occupying an arc of 240 in section. The 
proximal zooid of each internode is wider (Lz 0'37 mm), and the ultimate zooids shorter and 
narrower (Lz 0'50, Iz 0'30 mm), than other zooids. Orifice rounded distally with large 
complex condyles delimiting a proximal part which is straight or curved distally. Frontal 
shield with up to 30 large, irregular foramina. Hypostegal coelom extending beneath the 
frontal shield. Communication between the visceral coelom of one zooid and the hypostegal 
coelom of the next distal zooid inferred to be through a pair of minute pores passing through 
the distal-lateral calcification of the orifice (Fig. 30). Communication with the basal coelom 
through numerous pores in the confluent lateral and basal walls. Brooding zooids dimorphic, 
large, inflated basally, placed in the central zooid series, occurring in the proportion of 1 : 10 
autozoids. Brooding zooid orifices larger, but similar to those of autozoids. 

REMARKS. The earliest astogenetic stages are missing from the colonies examined, but it is 
inferred that they are similar to those of T. burrowsi (see below). The cuticle of the basal 
coelom becomes increasingly thickened and inflated with ontogeny, and eventually forms a 
supportive and rooting structure. 

The septulae connecting the visceral and hypostegal coeloms of sequential zooids are very 
small, and apparently pass through the thick calcification distal and lateral to the orifice. 
The tubular communication traces a curved path, at an angle to the longitudinal axes of the 
zooids, and is impossible to expose in its entirety in either longitudinal-lateral or in 
tangential thin-section. The pattern of other zooidal and extrazooidal communications has 
been described by Cook (1975). 

A few, small embryos have been seen in the brooding zooids. The distal part of the orifice 
of these zooids appears to be formed by the proximal calcification of the next distal zooid 
(see Fig. 30). 

Epizoic forms include other erect, jointed bryozoans such as Cellaria sp., small colonies of 
a Bowerbankia-like ctenostome, encrusting colonies of Robertsonidra argentea (Hincks), 
and foraminiferans belonging to the informal 'HomotremcC group. 

Tropidozoum burrowsi sp. nov. 

Tropidozoum sp. Cook, 1975 : 165, pi. 1, figs A-C, Fig. 2B (explanation of Plates 1 and 3 transposed). 
Hayward & Cook (in press). 

SPECIMENS EXAMINED. BMNH. HOLOTYPE, Tulear, TU 24, Madagascar, exterior Grande 
Vasque, 23.9.69, 15 metres, 1975.1.4.2. Paratypes, Tulear, Pichon D25, 1975.1.4.1; 
other material: South of Durban, Umtwalume River mouth North by West 7 miles, 12271, 
shell fragment bottom, 90 metres, 1949.11.10.184, and 12272, 1949.11.10.250, trans- 
ported, dead internodes. 

SAM. 'Meiring Naude\ Southeast Africa, Stn 131, 780 metres, Stn 151, 900 metres, 
transported, dead internodes. 

ETYMOLOGY. Named after Mr H. W. Burrows, whose manuscript notes and drawings on the 
Gilchrist Collection from South Africa are stored at the BMNH. 

DESCRIPTION (Figs 5, 6, 10). Tropidozoum with internodes 7 mm long and M mm wide, 
comprising 70 zooids. Longest longitudinal series of 16 zooids; widest, penultimate trans- 
verse series of 6 zooids. Zooids occupying an arc of 300 in section. Proximal and ultimate 
zooids wider than other zooids. Central basal keel thickened by base of calcification during 
ontogeny. Orifices rounded distally, with large condyles delimiting a deep, triangular sinus. 
Frontal shield with 30 small, irregular foramina, occluded during ontogeny. Dimorphic 
brooding zooids inflated basally, larger than autozooids, with wide, non-sinuate orifices. 

REMARKS. The early stages in astogeny of erect branches arise from a confused mass of 



rooting-kenozooids, and it is not known if these represent one, or a large number of colonies. 
Some kenozooids are rootlets, others are shorter, and from these small erect internodes 
originate. The maximum number of internodes in an erect branch is 1 5, reaching a height of 
130 mm. The primary internodes are nearly all small, including only 3 zooids, and are 
rounded in shape. Later internodes become progressively more elongated and include more 
zooids; a typical sequence has 3, 6, 25, and 36 zooids. Occasionally, a small internode 
occurs later in astogeny, between two longer internodes. Zooid length at the base of an 
internode forms a series Lz 0*55, 0'47, 0*60, 055, 062, 0'58, 0'60 mm (cf. E. obtecta). 

Brooding zooids appear by the fifth internode and are placed in the centre of a transverse 
zooidal series; a maximum of 10 occur in one internode. T. burrowsi is known from direct 
observation (Dr J.-G. Harmelin, pers. comm. 1974), to be a shallow-shelf species, and all the 
deep water records given above are of worn, dead internodes found in sediments which had 
been transported. The colonies from Madagascar were observed alive in situ; they have 
several epizoic and epiphytic forms, including sponges, small gorgonians, hydroids, 
serpulids, the bryozoan Cellaria sp. and small algae. The coelomic pigment of living colonies 
was bright red, and they were found growing together with gorgonians etc., and forming 
patches 20^40 cm across on calcareous accretions covered by sand. 


Neoeuthyris Bretnall, 1921 : 157. 

TYPE SPECIES. Euthyris woosteri MacGillivray, 1 89 1 . 

DESCRIPTION. Colonies encrusting, basal walls of zooids uncalcified except at the margins. 
Frontal shields curved, hyaline. Lateral walls prominent. Orifices with a wide sinus and 
paired condyles. Avicularia unilateral and oral, mandible hinged on a complete bar. 
Brooding zooids with large, dimorphic orifices and a brood chamber formed by a large 
endozooidal ovicell, closed by the operculum. 

REMARKS. The characters of N. woosteri, the only species, differ from those of other genera of 
Euthyrisellidae in the encrusting habit, lack of basal coelom, and presence of ovicells and 
avicularia. The essential structure of the colony, the nature of the frontal shield, the 
interzooidal communications and the dimorphism of brooding zooids, are, however, all 
similar to those found in the other genera. 

Neoeuthyris woosteri (MacGillivray) 

Euthyris woosteri MacGillivray, 1891 : 77, pi. 9, fig. 2. 

Neoeuthyris woosteri: Bretnall, 1921 : 158, Fig. 1. Hastings, 1960; 1964 : 245, figs 1,2. 

SPECIMENS EXAMINED. BMNH. Fremantle, Western Australia, 1948.3.12.1; Western 
Australia, 1938.8.10.1; Cosy Corner, Torbay Head, Western Australia, 1 979.2. 1.1. 
AM and NMV: U875 HOLOTYPE (see Hastings, 1964 : 246). 

DESCRIPTION (Figs 7, 12, 28, 29, 31, 32). Neoeuthyris encrusting algae (usually 
Metamastophora flabellatd), zooid basal walls uncalcified. Zooids communicating by 3-5 
lateral and 2-3 distal septulae placed at the base of the vertical walls. Visceral-to- 
hypostegal communication by paired groups of 4-5 septulae in the upper part of the distal 

Figs 10-13 Scanning electron micrographs of brooding zooids of Euthyrisellidae and 
Didymosellidae: (10) Tropidozoum burrowsi sp. nov. BMNH 1975.1.4.1, Madagascar x!03; 
(11) T. cellariiforme Harmer BMNH 1964.3.10.1, Celebes x!27; (12) Neoeuthvris woosteri 
(MacGillivray) BMNH 1979.2.1.1, Western Australia x50; (13) Didymosella larvalis 
(MacGillivray) BMNH 1979.7.5.2, near Melbourne, showing partially formed ovicells at 
growing edge x3 5. 


wall behind the orifice (Fig. 32). Frontal shield smooth, very thinly calcified, with small 
protuberances, but no suboral septulae, and with no extension of hypostegal coelom basally. 
Two pairs of frontal septulae beside the orifice, one pair giving rise to an avicularium (Fig. 
28). Avicularia with acute mandibles, directed proximally. Brooding zooids large, with 
dimorphic orifices, ovicell apparently formed by an expansion of the lateral and distal walls, 
protruding into the cavity of the next zooid, which is very short (Lz 0*45 mm). 

REMARKS. Avicularia on the right hand side of a lobe tend to occur on the right hand side of 
the zooids (viewed frontally), and vice versa. The algal substratum is influenced during 
growth by the presence of the bryozoan (see Hastings, 1964). 

The extreme fragility of the colonies makes investigation very difficult. Scanning electron 
microscopy of small fragments, and thin sections, have enabled us to reconstruct the 
interzooidal and intrazooidal communication systems, which are hardly visible in whole 
preparations using a light microscope. 

N. woosteri has rarely been reported, and the ancestrula and early astogeny is unknown. 
Most specimens are known to have encrusted M.flabellata which had been washed up on the 
shore, and both species probably occur together in deeper water. Nearly all records are 
from Western Australia, but MacGillivray (1891) reported specimens from Cooktown, 

Table 2 Measurements (mm) of autozooids and avicularia of Didymosellidae 




Lr Lm 

No. of 

Didymosella sp. 







D. irregularis 






D. acutirostris 






D. crassa 






D. porosa 






D. clypeata 






D. 'larvalis'- fossil 






(New Zealand) 

D. 'larvalis '-fossil 







D. larvalis- Recent 




0-20-0-30 0-15-0-25 


D. parviporosa 






D. pluma 




0-28-0-32 0-70-0-80 


T. magnirostris 




0-25-0-28 0-23-0-30 


T. magna 






T. boninensis 




0-15-0-20 0-50-0-75 


Didymosellidae Brown, 1952 : 194. 
TYPE GENUS. Didymosella Canu & Bassler, 1917. 

DESCRIPTION. Interior-walled cryptocystidean ascophorans, usually with extrazooidal basal 
coeloms, and considerable basal calcification. Zooid frontal shields with pores and marginal 
frontal septulae, becoming thickly calcified during ontogeny, obscuring the lateral walls. 

Peristomes elongated, with or without foramina and/or spiramina frontally. Primary 
calcified orifice not well defined, operculum a flap-like extension of the distal part of the 
ascus wall. Septulae in vertical and basal zooid walls numerous. Avicularia adventitious, 


arising from oral frontal septulae, usually unilateral, with large subrostral chambers and 
acute rostra. Mandibles orientated laterally, hinged on a complete bar. Brood chambers 
formed by an expansion of the distal part of the peristome wall, opening above the 

Ontogeny of zooid walls 

As in the Euthyrisellidae, ontogenetic changes in the Didymosellidae are apparently rapid. 
Few colonies show complete series of frontal shield development, and partially uncalcified 
buds are contiguous proximally with fully formed zooids in which secondary calcification is 
well advanced (see Fig. 21). For a brief period, the prominent lateral walls and depressed 
crypyocystal shield resemble those of Pleurotoichus or Tropidozoum. The presence of 
numerous frontal marginal septulae, however, means that the frontal calcification soon 
thickens, obscuring all zooidal boundaries. Funnels in the calcification common to tubular 
extensions above septulae of adjacent zooids are developed, as in the umbonuloid genus 
Tremogasterina (see Cook, 1977). 

The calcification of the primary orifice merges into the development of the peristome and 
no clear-out boundary is obvious. Avicularia are formed as lateral-oral frontal buds, their 
subrostral chambers deriving from a flattened plate of the frontal shield, bordered by 3-8 
marginal septulae (see Fig. 20). In D. pluma particularly, funnels derived from these 
septulae, and frontal pores, remain prominent features of the large avicularian chambers 
quite late in ontogeny (see Fig. 2 1 ). 

Coelomic systems and communication 

The presence of frontal septulae is correlated both with the absence of interzooidal 
visceral-to hypostegal communication (Fig. 1 B), and with the massive calcification of the 
frontal shields. In nearly all species, basal septulae, which communicate with a thin 
extrazooidal coelom, have been observed. Basal calcification, too, is considerable, and forms 
processes or massive pillars, which contact the substratum, allowing a semi-repent mode of 
life, or strengthening erect branches. The nature of the porous basal calcification has long 
been noted by authors (Stoliczka, 1865, MacGillivray, 1895, Waters, 1885, Levinsen, 1909, 
and Brown, 1952), who 'have either compared it to that of Selenaria (which also has an 
extrazooidal, basal coelom, see Chimonides & Cook, 1981), or have suggested a radicular 
function for the pores. 

Opercula, mandibles and peristomes 

In correlation with the elongated peristomes (see Harmer, 1957 : 652), and in contrast to the 
Euthyrisellidae, opercula are hardly differentiated, and are flap-like expansions of the ascus 
wall. Mandibles are acute, and hooked in D. larvalis and T. magnirostris. Those of D. pluma 
and T. boninensis are elongated and resemble the mandibles of the anascan genus Smittipora 
(see Harmer, 1926), in having a central rachis, and expanded wings of cuticle. 

Brooding zooids 

In some species, brooding zooids and their orifices are larger than those of autozooids, but 
there seems little evidence of dimorphism in other characters. In some species, the brooding 
zooids may have smaller avicularia, or none, and there is evidence of patterning of the 
avicularia of surrounding zooids. 

Unlike the Euthyrisellidae, the family Didymosellidae has an extensive fossil record (see 




DWYMOSELLA Canu and Bassler 

Didymosella Canu & Bassler, 19 1 7 : 43. 

TYPE SPECIES. Lepralia larvalis MacGillivray, 1869. 

DESCRIPTION. Zooidal peristomes with large, paired foramina, spiramina absent. 

REMARKS. Didymosella was included in the family Escharellidae by Canu & Bassler 
(1929 : 33), and in the Exochellidae by Bassler (1953 : G205). The distinctive characters of 
the genus were discussed by Brown (1952 : 194), who noted that the Italian fossil species 
Porina bioculata Waters (1891 : 26, pi. 3, fig. 15), the type species of Bimicroporella Canu 
(1904 : 12), was not closely related to Didymosella. B. bioculata (Waters), B. ventricosa Canu 
(1904) and B. watersi Brown (1958) all have some superficial similarity with Didymosella, 
but have no suggestion of a basal coelom, and have hyperstomial ovicells. 

Didymosella has an extensive fossil record. The earliest known species is an undescribed 
form listed by Labracherie (1972 : 44), from the Middle Eocene of North Aquitain, France. 
Specimens of this species have been examined (Bazas-rural, 200 metres, Labracherie Coll.). 
The material is fragmentary and comprises only 30 zooids. Most of the fragments appear to 
have been encrusting, but some have basal grooves and a few have minute pores, which 
together with the thick, basal calcification are interpreted to indicate the former presence of a 
basal coelom. The frontal shields have 3-6 pores and 7-9 funnels indicating frontal 
septulae. Zooids communicate through 1-2 distal, and 2-3 lateral septulae. The paired 
peristomial foramina are distinct, as are the avicularian rostra, some of which have a 
complete bar. The rostra are open terminally which, by analogy with D. pluma (see p. 76) 
probably indicates that the mandibles were elongated. 

Another fossil species, D. irregularis, was described from the Upper Eocene (Lower 
Jacksonian) of north-western Florida by Cheetham 1963:66, pi. 2, fig. 13). This had 
unilaminar colonies and larger zooids with up to 20 frontal pores and very small, acute 
avicularia. The orifices of the brooding zooids; which had broken peristomes exposing the 
ovicells, were slightly dimorphic. 

In view of Labracherie's (1972) records, it appears that D. acutirostris Faura y Sans & 
Canu (1917 : 90, pi. 7, figs 3-5), from the Upper Eocene of Monresa, Barcelona, is also 
attributable to Didymosella. The illustration is not conclusive, but the species was described 
with paired, peristomial foramina and peristomial ovicells. The zooids were larger than 
those of Labracherie's species. 

The wide Eocene distribution of Didymosella continued during the Oligocene and 
Miocene (see below), and is maintained in Recent seas. 

Didymosella crassa Canu and Bassler 

Didymosella crassa Canu & Bassler, 1920 : 1 16, pi. 88, figs 1-7. 

SPECIMENS EXAMINED. BMNH. Oligocene (Vicksburgian), west bank of Conecuh River, 
Escambia Co., Alabama. Canu & Bassler Coll. 

Figs 14-20 Scanning electron micrographs of fossil and Recent Didymosellidae: (14) 
Didymosella clypeata Canu & Bassler BMNH Lower Miocene, Victoria, basal side of branch, 
showing septulae x31; (15) D. clypeata, frontal side of branch x34; (16) D. porosa (Stoliczka) 
BMNH D 36784-92, Middle Oligocene, New Zealand, part of frontal margin of branch x21; (17) 
D. 'larvalis' BMNH D36783-7, Miocene, New Zealand, x40; (18) Tubiporella boninensis Borg 
BMNH 1889.8.21.105, Recent, China Sea, lateral view of young colony, ancestrula at right, 
showing basal extrazooidal calcification x25; (19) T. boninensis zooids at growing edge, 
showing spiramina, frontal pores and avicularian rostra x57; (20) T. magnirostris 
(MacGillivray) BMNH 1899.7.1.2686, Recent, Victoria, zooids viewed from distal end, 
showing spiramen (arrowed) and base of avicularian subrostral chamber (bottom right) x57. 


DESCRIPTION (Fig. 23). Didymosella with large zooids and tubular peristomes. Zooids with 
25-35 frontal pores and 6-8 marginal septulae. Avicularia large, with channelled rostra 
open terminally. Basal surface thickened, with grooves and pores. 

REMARKS. In all respects D. crassa and D. pluma (see p. 76) are so alike that it is possible to 
postulate a direct genetic link between the Oligocene and Recent populations from the 
central West Atlantic. The skeletal characters differ from those of D. pluma only in the larger 
number of frontal pores and in the absence of elongated calcified basal processes, which last 
may be due to wear during preservation. The open channel at the terminal end of the 
avicularian rostrum allows the inference that the mandible was elongated, and may have 
even been expanded laterally like that of/), pluma (see p. 78). 

Didymosella porosa (Stoliczka) 

Semiescharipora porosa Stoliczka, 1865 : 128, pi. 19, figs 10-13. 
Didymosella porosa : Canu & Bassler, 1929 : 328. 
Didymosella aft. porosa : Brown, 1952 : 198, Figs 139, 140. 

SPECIMENS EXAMINED. BMNH. Middle Oligocene, Pareora, Tarahoke Quarry, N.W. tip of 
South Island, New Zealand, D36788-92. 

DESCRIPTION (Figs 16, 27). Didymosella with erect, ligulate branches, with 4-5 transverse 
zooid series. Branches flat and bifurcated. Lateral marginal zooids curved outward, their 
peristomes forming serrations. Secondary orifices with paired spine bases. Marginal 
avicularia large. Basal calcification with numerous pores. 

REMARKS. Branches are composed of 20 sequentially distal zooid series. The marginal 
zooids are curved so that their orifices are at an angle of over 45 to those of the central series. 
At the same time, the orifices are curved towards the frontal side of the branch. The paired 
peristomial foramina are separated from the proximal edge of the peristome by a distance 
greater than 0*20 mm. 

Stoliczka (1865) mentioned the thickening of the basal calcification and the porous 
surface, which was also noted by Brown (1952), who suggested that the pores might have a 
radicular function. In spite of Brown's doubtful assignment of his material to D. porosa, it is 
so close in general characters, age and locality to Stoliczka's specimens (which were from the 
Lower Miocene of Orakei (Hauraki) Bay, North Island), that it may be confidently included 
in the species. 

Didymosella clypeata Canu & Bassler 

Didymosella clypeata Canu & Bassler, 1935 : 32, pi. 9, figs 7, 8. 

SPECIMENS EXAMINED. BMNH. Lower Miocene, Mount Martha, Victoria. Anticline Creek 
(Janjukian), Dartmoor, Victoria D34200-01. 

DESCRIPTION (Figs 14, 15). Didymosella with erect ligulate branches, with 2-3 transverse 
series of zooids. Branches curved towards the basal side so that orifices of the marginal series 
are at an angle of 70 to those of the central series. The marginal zooids also curve outwards 
slightly at about 1 5 to the central axis. Secondary orifices with paired spine bases. Frontal 
pores are stellate (in basal view) and the paired peristomial foramina are close to the 
proximal edge of the peristome, at a distance of 0- 10-0- 1 5 mm. Basal calcification thickened, 
with a few irregularly spaced pores. 

REMARKS. The marginal avicularia are large, and occur about halfway up the zooids. 

Although very similar to D. porosa, D. clypeata differs in the number of zooids in a 
transverse series, the curvature of the branches and the placing of the peristomial foramina. 


The combination Porina clypeata Waters (1881 : 332, pi. 17, fig. 67, 1882a : 268) was used 
for an entirely different species from the Miocene of Mount Gambier, Victoria. 

The Didymosella /ai*va/is-complex 

The Recent species D. larvalis is here confined to algal-associated, encrusting colonies, with 
no evidence of basal coeloms. Two fossil populations, one from the Australian Miocene, the 
other from the Oligocene to Miocene of New Zealand, have been ascribed by several authors 
to D. larvalis. Neither of these populations is assignable to D. clypeata or D. porosa, and 
they differ considerably from Recent D. larvalis in their colony forms. 

A. Didymosella aff. larvalis (fossil specimens) 

Porina larvalis: MacGillivray, 1895:104, pi. 14, fig. 26, Waters, 1882a:269, pi. 8, fig. 19; 

1882b: 509; 1887: 189, pi. 6, fig. 8. 
Didymosella larvalis: Brown, 1952, part: 195, Figs 135-138; 1958 : 56. 

SPECIMENS EXAMINED. BMNH. Oligocene, Tarakohe Quarry, Waitapu, New Zealand, 
D36787; Miocene, Weka Pass, New Zealand, D36783-6, D36938, D36961-2; Miocene 
Muddy Creek, Victoria, Australia, D34255; Mount Gambier, Victoria, D32981-3 (Waters 
MM. Bairnsdale, Victoria, T42 (figured Waters, 1882a, pi. 8, fig. 19). 

DESCRIPTION (Fig. 17). Didymosella with erect colonies with bilaminar expansions or 
cylindrical branches. Peristome slightly raised, thickened proximally forming a transverse 
ridge which overhangs the paired, peristomial foramina. Frontal pores stellate. 

REMARKS. The bilaminar colonies (from Weka Pass, New Zealand), have basal walls forming 
a double layer. No pores connecting the layers can be seen. The erect fragments also have 
complete basal walls, and have a central cavity. One fragment (D36786) has a very long 
avicularium with a channelled rostrum which is open terminally. In general, the specimens 
are poorly preserved and worn. The calcification is, however, considerably thicker than that 
of Recent specimens. The stellate frontal pores can be seen from the inner surface of broken 

B. Didymosella larvalis (MacGillivray) 

Lepralia larvalis MacGillivray, 1869 : 134. 1879 : 30, pi. 37, fig. 5. 
Escharoides larvalis: Levinsen, 1909 : 318. Livingstone, 1926 : 170, pi. 11. 
Didymosella larvalis Brown, 1952, part: 195. 

SPECIMENS EXAMINED. BMNH. Spencer Gulf, 1890. 4. 17.1 Opt. Australia, 1897.5.1.883, 
1963.4.18.16; Port Phillip Heads, 1897.5.1.881.882; Adelaide, 1899.7.1.5036A; Cowes 
Phillipps Id., 20 miles S.E. Melbourne, beach sand, 1979.7.5.2; No locality, 1963.2.12.63. 

DESCRIPTION (Figs 9, 13, 36). Didymosella with colonies encrusting algae, sometimes rising 
into small, unilaminar expansions. Zooids communicating by 2 distal and 2-3 lateral 
multiporous septulae, surrounded by chambers of calcification. Basal wall uncalcified in 
encrusting zooids, peripherally calcified, or with a large, uncalcified, cuticle covered window 
in erect zooids. Proximal side of peristome elongated, calcification thin and hyaline. Distal 
side of peristome deficient. Lateral peristome with spinous processes. Frontal pores stellate, 
frontal septulae numerous. Avicularia large, with acute mandibles, hooked terminally. 
Ancestrula with a complex frontal shield of spinous processes. 

REMARKS. The thin frontal calcification and lack of basal calcification are similar to that of 




the algal-associated Neoeuthyris woosteri (see p. 67), and are also usually present in 
numerous other cheilostome species which are specific to this type of substratum. In erect 
zooids, the uncalcified basal area is progressively decreased with astogeny, eventually 
becoming confined to a window in the basal wall. No evidence of a basal coelom or of 
radicular expansions has been seen. 

Specimens are small, but a few parts of colonies (1897.5.1.882) comprising more than 
100 zooids, on narrow fronds of algae, show a distinct central zone of brooding zooids which 
have minute, paired avicularia. They are flanked by lateral series of autozooids in which 
most of the avicularia are placed with the closed mandible directed medially (cf. T. 
magnirostris, p. 77). In other colonies, there is no obvious patterning of brooding zooids or 
of avicularia. 

Ancestrulae (1890.4. 17. 1. -10 pt., 1963.2.12.63) are large (La 0'52 mm, la 0'40 mm), 
and distinctly 'cribrimorph' in character. There is a well developed gymnocyst and large, 
oval opesia, overarched by 4 robust lateral spines. The oral shield is formed by a pair of large, 
spinous processes, which are fused centrally, leaving an open foramen. The shield terminates 
distally in 5 spines, and a further, laterally placed pair of bifurcated spines is closely apposed 
to the lateral walls of the ancestrula and the distal-lateral primary zooids of the post- 
ancestular triad (see Fig. 9). The systematic significance, if any, of 'cribrimorph' ancestrulae 
in ascophoran species is unknown. The ancestrulae of other species of Didymosella have not 
been found, and that of Tubiporella boninensis (see below) is not 'cribrimorph', but 
resembles subsequent zooids. Most cribrimorph species have anasciform, 'tata' ancestrulae, 
but 'cribrimorph' ancestrulae are known in the umbonuloid ascophorans Triporula 
bidenticulata (Canu & Bassler), see Cook (1967:341) and Romancheina asymmetrica 
Moyano (1975 : 63, Figs 1-8). Temachia opulenta Jullien (1883 : 509, pi. 14, figs 26-29) also 
has a 'cribrimorph' ancestrula and is so similar to R. asymmetrica that it too may be 
postulated to have umbonuloid ontogeny. Apart from D. larvalis, the only cryptocystidean 
species with 'cribrimorph' ancestrulae known at present is Schizoporella kiiensis (Okada & 
Mawatari), see Mawatari (1952 : 279, fig. 13). 

Didymosella parviporosa Canu & Bassler 

Didymosella parviporosa Canu & Bassler, 1929 : 327, pi. 39, fig. 1. 
Didymosella costulata Canu & Bassler, 1929 : 327, pi. 39, fig. 2. 

SPECIMENS EXAMINED. BMNH. Philippines, Albatross Stn D5 145, 42 metres, 1931.12.30.119 
(D. parviporosa}; Stn D55141, 53 metres, 1931.12.30.120. (D. costulata}. 

DESCRIPTION. Didymosella with small, narrow zooids, each with an average of 20 frontal 
pores. Secondary thickening forming interzooidal funnels, with bars of calcification. Basal 
calcified wall with furrows and a few scattered pores. Avicularia, small, with acute rostra. 

REMARKS. The specimens include one fragment of D. costulata and three of D. parviporosa, 
together comprising only 40 zooids. Both species were described from the same region and D. 
costulata appears to be a slightly worn, ontogenetically thickened stage of development of D. 
parviporosa. This conclusion is strengthened by the locality data of the BMNH specimens, 
which were labelled by Bassler. One, labelled D. parviporosa, is from Stn 5145, the original 
locality for D. costulata; the other labelled D. costulata, is from Stn 5141, one of the original 
localities for D. parviporosa. 

Figs 21-24 Scanning electron micrographs of fossil and Recent Didymosellidae: (21) 
Didymosella pluma sp. nov. BMNH 1899.7.1.5324, Recent, off Brazil, zooids at growing edge, 
showing raised lateral walls, frontal pores and channelled avicularian rostra x71 ; (22) D. pluma, 
lateral view of zooids showing lateral septulae and basal calcified processes x62; (23) D. crassa 
Canu & Bassler BMNH Oligocene, Alabama, showing channelled avicularian rostra x59; (24) 
Tubiporella magnirostris (MacGillivray) BMNH 1899.7.1.2686, Recent, Victoria, growing 
edge viewed from distal end of zooids, showing distal and basal septulae and basal calcified 
processes x7 5. 


D. parviporosa differs from D. larvalis in the presence of a basal coelom, and its small, 
narrow zooids. It differs from D. pluma in its much smaller zooid size, and in its avicularian 
mandibles, which may be inferred to have been acute. 

Didymosella pluma sp. nov. 
Didymosella sp. Brown, 1952 : 198. Cook, 1981 (in press). 

SPECIMENS EXAMINED. BMNH. HOLOTYPE, John Adams Bank (=Victoria Bank, off Brazil, 
Approx. 21S, 37W, depth less than 183 metres), 1899.7.1.5322. Paratypes, 1899.7.1.5323, 
5324,4476,4477, 176; Havana, 1911.10.1.1702, 146 metres. 

ETYMOLOGY. Pluma (L) a feather, referring to the expanded avicularian mandible. The 
name was given as a manuscript label by Busk. 

DESCRIPTION (Figs 21, 22, 25). Didymosella with semi-repent colonies forming irregular 
expansions. Frontal shield with 15-23 pores and 6-8 marginal septulae, the distal pair 
enlarged. Zooids communicating by 2-4 distal and lateral septulae. Peristomes raised, 
tubular, with paired, marginal spinous processes. Basal calcified wall becoming extrazooidal 
with septulae and long, calcified extensions. Avicularian subrostral chambers very large, 
rostra with a channel open terminally. Mandible very long, expanded and fimbriated 

REMARKS. The thickening of the frontal calcification above and around the distal pair of 
frontal septulae forms large, slit like foramina, one either side of the orifice. Two large lobes 
of about 300 zooids each, show areas in which groups of zooids with 'left-handed' avicularia 
are opposed to groups with 'right-handed' avicularia. Unlike the colonies of T. magnirostris 
(see p. 79), these groups are not obviously associated with the presence of brooding zooids. 
The avicularian rostra have pores which are visible late in ontogeny. The remarkable 
mandibles resemble those of Smittipora (see Cook, 1964), but are fimbriated marginally. 
When open, each mandible covers the frontal foramina of the zooid; when closed, it usually 
covers the foramina of the adjacent, lateral zooid (see Fig. 25). Presumably the mandibles 
ensure that the foramina do not become blocked by detritus, which would restrict water flow 
into the ascus, and the laterally directed water currents produced by mandibular movement 
may assist in clearing deposits from the frontal surface of the zooids (see Cook, 1981). 
Expanded mandibles are also present in Tubiporella boninensis (see below). 

D. pluma is so similar to D. crassa that it is highly probable that Miocene-to-Pliocene 
specimens, linking the two forms may eventually be found from sediments in the Gulf of 
Mexico region. 

Tubiporella Levinsen, 1909 : 304. 
TYPE SPECIES. Lepralia magnirostris MacGillivray, 1883. 

DESCRIPTION. Colonies semi-encrusting, with free expansions. Zooids with long, tubular 
peristomes and a frontal spiramen. Frontal and basal septulae numerous, and extrazooidal 
calcification considerable. Avicularia lateral, arising from several frontal septulae, subrostral 
chambers large. Rostra acute, directed laterally, mandible slung on a complete bar. Brooding 
zooids with a slightly enlarged orifice. 

REMARKS. Tubiporella was introduced with no formal description. Levinsen (1909 : 304) 
listed it as 'n.g.' in a key to the family Tubucellariidae; T. magnirostris was the only species 
included. Species of Tubucellaria (or Margaretta, see Harmer, 1957 : 824), have an 
ascopore, not a peristomial spiramen, and have erect, jointed colonies. Tubiporella was 
assigned to the Tubucellariidae by Bassler (1953 : G210), and to the Porinidae by Harmer 


(1957 : 844). Finally, Brown (1952 : 194) recognized its close affinities with Didymosella and 
included it in the family Didymosellidae. 

Tubiporella, like Didymosella, has an Australian Tertiary record, although this extends 
only to Miocene deposits. The characters of fossil specimens are very similar to those of 
Recent T. magnirostris, but there are some doubts in accepting the identity of these forms 
without examination of type specimens. Brown (1952) noted that Recent colonies had the 
larger zooidal dimensions (see Table 2), and in view of the degree of difference between 
Recent and fossil populations of the D. larvalis-complex, it seems advisable at present to 
regard the Tertiary forms as distinct taxa. Several fossil forms have been described. Waters 
(1882# : 268, pi. 9, figs 33, 34) introduced Micropella introversa from the Miocene of Mount 
Gambier, Australia with two drawings which he stated were magnified at x!2 and x25 
respectively. The (secondary) orifice measurements he gave on p. 268 were: Lo - 16 mm, lo 
0'20 mm. Using the magnifications given, the average dimension of zooids for Fig. 33 would 
be: Lz l - 75 mm, Iz I'OO, Lo 0'25 mm, lo 0'33 mm. These are obviously too large; very few 
cheilostome zooids are more that 1'5 mm in length, and Waters did not mention that the 
zooids of M. introversa were particularly large. A magnification of x20 would give Lz 
O90 mm, Iz O60 mm, Lo 0'15 mm, lo 0*20 mm, which is not only in accordance with the 
given orifice measurements, but is more feasible. The spiramen, which Waters regarded as 
'microporellid' ascopore, was described and figured as being very close to the proximal 
border of the secondary orifice. When describing further fossil specimens from the River 
Murray Cliffs, Waters (1885 : 296) called this a 'central pore'. In a still later paper, Waters 
(1887 : 55) included M. introversa in the synonymy of T. magnirostris although the species 
had originally been described by him a year before MacGillivray's species. Waters 
(1885:295, pi. 7, fig. 7) also described, as Microporella magna, fossil specimens from 
Aldinga and Mount Gambier, and identified them with Lunulites magna Tenison Woods 
(1880), which he stated was 'very closely allied' to T. magnirostris. Tenison Woods (1880 : 7, 
pi. 1, figs 6 a-d), had described an irregularly domed, hollow, almost circular colony. His 
figure (6c) shows zooids with raised peristomes, numerous frontal pores and acute, laterally 
directed avicularia with a complete bar. This species is obviously ascophoran, and is in no 
way referable to the free-living, anascan genus Lunulites. Two specimens in the British 
Museum Collections confirm the existence of this species in the Australian Tertiary. The 
first (labelled 'Lunulites magna' . D33050, Janjukian, Aldinga) has a domed colony 25 mm 
in diameter and 10 mm high, somewhat larger than Tension Woods' colonies. The zooids 
have raised peristomes, extrazooidal frontal thickening with funnels and pores, and a distinct 
proximal pore which may be a spiramen. The avicularia are directed almost distally, and are 
much larger than those figured by Tenison Woods (1880, pi. 1, fig. 6c). They have a 
complete bar, and the rostra are acute, with a terminal channel which may indicate that the 
mandibles were elongated. The basal surface of the colony is obscured by matrix. The second 
colony (D31940, River Murray Cliffs) forms an almost truncated cone 20 mm in diameter 
and 1 3 mm high, but the zooids are not arranged in regular radial series. They are very worn, 
but the peristomial orifices and avicularia can be seen. The hollow basal side of the colony 
has numerous pores and irregular masses of calcification and may be inferred to have 
possessed an extrazooidal coelom in life. 

Pachystomaria parvipuncta MacGillivray (1895:97, pi. 8, fig. 24), another Miocene 
Australian form, appears to be a further species belonging to this complex. MacGillivray 
noted that the porous basal surface was similar to that ofSelenaria, an observation also made 
by Waters (1885) when describing M. magnirostris. At present it is not known whether any 
or all of these nominal species are synonymous, or if they are conspecific with Recent T. 

Tubiporella magnirostris (MacGillivray) 

Lepralia magnirostris MacGillivray, 1883 : 134, pi. 1, fig. 6. 





Tessaradoma magnirostris: MacGillivray, 1889 : 21 1, pi. 175, fig. 1. 

Porina magnirostris: Hincks, 1884 : 279 (sep. p. 129), pi. 9, fig. 6. 

Tubiporella magnirostris: Levinsen, 1909 : 307, pi. 16, figs 5a-d. Canu & Bassler, 1920 : 549, Figs 161 


Tubiporella magnirostris part, 'f. \ictoriensis": Borg, 1940 : 416, Figs 1-3. 

SPECIMENS EXAMINED. BMNH. Port Phillip Heads, Victoria, Australia, 1887.6.27.1, 
1887.12:10.120, 1893.8.11.17, 46 metres; 1927.8.4.24, 1979.1.12.3. Port Phillip, 
1899.5.1.1214, 1899.7.1.5409, 5410, 5411, 2333, 2338, 2686, 2687, Victoria, 1979.7.2.1. 
Amirante Islands, 64 metres, 1 936. 1 2.30. 1 99. 
NMV. Port Phillip, 63876. 

DESCRIPTION (Figs 20, 24). Tubiporella with colonies forming broad, foliaceous expansions 
attached above the substratum by thick, extrazooidal, columns of porous calcification. 
Zooidal peristomes tubular, but not very prominent, with a small proximal, cuticle-covered 
foramen early in ontogeny. Spiramen originating on the proximal edge of the base of the 
peristome, apparently migrating proximally with thickening calcification. Avicularian 
subrostral chambers without pores, mandible hooked but not greatly elongated. Lateral 
frontal septulae surrounded by large uncalcified areas, which increase in size with calcifi- 
cation. Brooding zooids with larger secondary orifices than autozooids, with no avicularia, 
or paired, small avicularia. Avicularian mandibles of autozooids surrounding brooding 
zooids are directed away from brooding zooid when closed. 

REMARKS. The primary orifice can be seen early in ontogeny, and the developing peristome 
has a small foramen which rapidly becomes obscured. The 'migration' of the spiramen with 
thickening of the frontal shield is similar to that described in the umbonuloid family 
Adeonidae (see Cook, 1973). The patterning of brooding zooids and avicularia is similar to 
that found in D. pluma. The peristomial orifices of the brooding zooids are slightly larger 
than those of the autozooids, and the distal expansion of the ovicell can be seen. The lateral 
frontal septulae of the brooding zooids remain visible at the base of the funnels in the 
calcification, resulting in wide lacunae beneath the investing cuticle (cf. D. pluma}. Levinsen 
(1909 : 308) mentioned that the 'very thick basal wall of the colony ... is ... perforated by 
pore canals' which he noted were derived from multiporous basal septulae. Harmer 
(1957:1012) described a colony of Emballotheca subimmersa (MacGillivray) which 
encrusted the basal wall of a large specimen of T. magnirostris. 

Tubiporella boninesis Borg 

Tubiporella magnirostris part, 'f. boninensis': Borg, 1940 : 420, Fig. 4, (operculum). 

SPECIMENS EXAMINED. BMNH. Tizard Reef, 64 metres, 1889.21.27; 68 metres, 1892.8.8.26; 
Macclesfield Bank, China Sea, 66 metres, 1889.8.21.95,105; 64-79 metres, 1934.1 1.6.23; 46 
metres, 1 893. 8. 1 1 . 1 7; China Sea, 66 metres, 1979. 1 .6.2; S. of Bua Bua passage, Falafuti, 264 
metres, 1903.1.29.60. Fuafatu, 109 metres, 1903.1.29.61. 

DESCRIPTION (Figs. 18, 19, 26). Tubiporella with narrow, ligulate, semi-erect colonies, with 
zooids in 4-7 transverse series. Zooids with long, tubular peristomes, imperforate early 
in astogeny. Spiramen at the base of the peristome, migrating only slightly with ontogeny. 
Basal calcification produced into a massive, central keel in erect specimens, which produce 
branched, supporting pillars of extrazooidal, porous calcification 5-7 mm deep. Avicularian 
subrostral chambers large, imperforate. Rostra acute, with an open channel terminally. 

Figs 25 and 26 (25) Didymosella pluma sp. nov. BMNH 1899.7.1.5322, off Brazil, sketch of 
four zooids and two avicularia with open (upper) and closed (lower) mandibles. Scale bar = 
0-50 mm; (26) Tubiporella boninensis Borg BMNH 1893.8.11.17, China Sea, sketch of three 
zooids and two avicularia with open (upper) and closed (lower) mandibles Scale bar = 0- 50 mm. 


Mandibles elongated, directed laterally towards the margins of the colony; expanded 
laterally, with notched, fimbriated edges. 

REMARKS. Although Borg (1940) did not consider that his specimens were specifically 
distinct from T. magnirostris and did not formally introduce the name 'boninensis' for them, 
the name is available and is therefore used here. Borg noted that the operculum was hardly 
differentiated from the ascus wall, and that the avicularian mandible was 'distinctly broader 
than that of the Australian colonies and nearly straight, but with its tip rather sharply bent'. 
He also described the cuticular expansions and noted that 'this border is often broader on one 
side'. The mandibles are similar to those of D. pluma; when open they cover the area above 
the frontal spiramen. (see Fig. 26). 

The ancestrula resembles later-budded zooids and is unusual in having a lateral 

Generally, T. boninensis differs from T. magnirostris in its colony form, shorter, more 
erect zooids, longer peristomes, and avicularian mandibles. 

Note on Tubiporella levinseni Borg, 1 940 

In his description of the genus Tubiporella, Borg (1940) described a new species, T. 
levinseni, which had large, encrusting zooids with long, tubular peristomes, multiserial 
marginal pores and a proximal spiramen. The centrally placed avicularium had an 
elongated, almost setiform mandible. Borg mentioned that he had shown his specimens to Sir 
Sidney Harmer, who had recognized them as belonging to the same species as specimens 
from the Paternoster Islands which were present in the Siboga Expedition Collections. 
Harmer was not aware of Borg's description, which was subsequently published in Sweden 
during the 1939-1946 war (see Explanatory Note by Dr A. B. Hastings in Harmer, 
1957 : xiv). The specimens from the Paternoster Islands were described as Reptadeonella 
flagellifera by Harmer (1957), who, however, had provisionally labelled them as 

The structure and relationships of T. levinseni differ from those of Tubiporella as 
represented by the type species, T. magnirostris, and by T. boninensis. The specimens 
examined by Harmer from the Siboga Collections, together with one large, additional colony 
without locality, but comprising more than 2000 zooids (which was found fortuitously in the 
unnamed collections of the BMNH, and was not seen by Harmer), have been examined. A 
few zooids show a cuticular frontal wall with operculum, which is overarched by calcifi- 
cation, indicating that the ontogeny of the frontal shield is umbonuloid. There are no frontal 
pores, and the frontal septulae are multiserial and extend round the distal side of the orifice 
(see also Borg, 1940). The long tubular peristome is thus subterminal, and the primary 
orifice is well defined, wih minute condyles and distinct operculum (see also Borg, 
1940 : 422, Fig. 8). The avicularium is central in position and is derived from one lateral- 
oral septula. The subrostral chamber is small, and the rostrum is acute and directed distally 
and laterally. The mandible is elongated, setiform and slung between condyles. Zooids 
communicate through septulae at the base of the vertical walls, but there is no basal coelom 
and the basal wall is completely calcified. The zooids are nearly all encrusting, but where 
they stretch across irregularities in the substratum, have produced small, peg-like 
expansions. There are however, no extrazooidal pillars as in Tubiporella. 

All the character correlations are similar to those found in the umbonuloid family 
Adeonidae (see Cook, 1973), and they confirm that Harmer (1957) was correct in attributing 
his specimens to the genus Reptadeonella, which has encrusting colonies. 

Both Borg (1940) and Harmer (1957) mentioned that some zooids with short peristomes 
may have been brooding zooids. A few zooids have wide, rather slit-like orifices, but their 
peristomes do not differ from those of other zooids. Generally, brooding zooids in the 
Adeonidae are strongly dimorphic, and it is probable that as yet, no brooding zooids have 


been found in this species. The synonymy, distribution and measurements of R, levinseni 
(Borg) are as follows: 

Reptadeonella levinseni (Borg) 

Tubiporella levinseni Borg, 1940 : 420, pi. 1 and Figs 5-8. 
Reptadeonella flagellifera Harmer, 1957 : 817, pi. 54, figs 4-7. 

SPECIMENS EXAMINED. BMNH. Siboga Stn 315, N. of Sumbawa, Paternoster Islands, 0-36 
metres, 1979. 1.2.1; no locality, 1979.1.2.2. 
ZMA. Stn 3 1 5 and Stn 60, 303, Hangsisi, Samau Islands, W. Timor, 0-36 metres. 

DISTRIBUTION. Jaluit, Marshall Islands, 2 metres (Borg), Paternoster Islands and West 

MEASUREMENTS (in mm). Lz 0-80-1-30 Iz 0-50-0-90 Lo 0-13 lo 0-16-0-20 Lr 0-16-0-25 
Lm 0-30-0-50. 


Study of Bryozoa emphasizing the concept of colonies as entities composed of cooperative 
members has increased during recent years as a result of observation of living colonies and 
the application of new techniques. Species show many kinds and degrees of structural and 
functional integration. Colonies may be composed of almost autonomous member zooids, or 
have various correlations of morphology and behaviour which in some cases amount to 
virtual colony control (see Cook, \919a, and Ryland 1979). The relationships of zooid body 
walls, intercommunications, zooidal and extrazooidal coeloms, astogenetic and ontogenetic 
changes, polymorphism and the patterning of polymorphs, all express the degree of such 
colony control and may be analysed and arranged in series of increasing integration 
(see Boardman & Cheetham, 1973 and Cook, 19790). 

Expressed in terms of such series, the degree of integration of the families Euthyrisellidae 
and Didymosellidae is considerable. It is reflected in the zonation and polymorphism of 
brooding and avicularian zooids, and in the supporting and attachment structures of 
colonies. Most significantly, it is expressed in the preponderance of interior calcified 
walls, and in the various combinations of intrazooidal, interzooidal and extrazooidal 
coelomic communications. 

Relationships both between the Euthyrisellidae and the Didymosellidae, and between 
them and other ascophoran groups are hard to evaluate. Although they share crypto- 
cystidean frontal shield ontogeny, the differences between the two kinds of shield suggest that 
these structures have arisen more than once in cheilostome evolutionary history. The 
appearance through time of more than one kind of frontal shield in the 'catenicellid' genera 
(see Banta and Wass, 1979), indicates that many more cryptocystidean families will require 
detailed investigation before the degrees and kinds of relationships among apparently 
similar groups are established. 

The presence of extrazooidal basal coelom and of interior basal zooid walls is another 
shared character. However, these structures are also known in such diverse systematic groups 
as the lunulitiform, free-living anascan Selenariidae (see Chimonides & Cook, 1981) and the 
cryptocystidean Petraliidae. Petralia undata has erect, reteporiform, rooted colonies with a 
high degree of organization and an extensive basal coelom. 

The lateral and basal rooting coleomic systems of the Euthyrisellidae have very close 
parallels among the many erect, flustriform genera, especially Onchoporella (see p. 59). 

Peristomal spiramina, too, are found in a diversity of Recent and fossil ascophoran genera; 
for example, Adeonella (see Cook, 1973), Gigantopora (see Harmer, 1957), and compare 
Pachydera rarepunctata Voigt (1967 : 67, pi. 24, figs 1, 2, from the Upper Cretaceous of 
West Kasachstan, USSR). 

Within the Euthyrisellidae, there is great diversity in frontal shield calcification and in 




orifice shape which is not strongly correlated with colony form. This diversity suggests that 
the family has a long evolutionary history, but unfortunately, no fossil species are known. 
Internodes of Tropidozoum are fairly robust, and have been found in transported deposits; 
they might be expected to be recognizable in Tertiary sediments. Even the thinly calcified 
zooids of Euthyrisella could be preserved, by analogy with the equally fragile fragments of 
Selenariopsis, which have been found, albeit rarely, in the fine fractions of Australian 
Miocene sediments (see Cook & Chimonides, 1981). Lagaaij (1968, 1973) has docu- 
mented records of similar fragile species, several of which were not previously known as 
fossils. It is perhaps possible that the ancestral forms of Euthyrisellidae had more heavily 
calcified colonies, which have not yet been recognized as being systematically related to the 
Recent species. 

In contrast, the long fossil record of the Didymosellidae might be expected to provide some 
evidence of evolutionary trends and relationships. Diversity among the known species, both 
in zooidaP morphology and colony form is, however, far lower than that found in the 
Euthyrisellidae. Generally, the species have very similar patterning of avicularia and 
autozooids, and the repent or semi-encrusting colony form is know in both Eocene and 
Recent species. The erect, branching colonies of D. porosa and D. clypeata are not found 
among Recent species, and the algal association of D. larvalis (like that of Neoeuthyris 
woosteri), may have been evolved fairly recently. 

To a certain extent, the distribution of species in time and space shows strong collection 
bias, and reflects the large number of faunal descriptions of the abundant bryozoan 
sediments of the Tertiary-to-Recent of the Gulf of Mexico and Australasian regions. The 
Euthryisellidae seems to have a purely Indo-West Pacific and Australian range. Most species 
are from shallow shelf or sublittoral waters, and several are rooted in sediments. The 
Didymosellidae are also shallow shelf species, although D. pluma and T. boninensis are 
found from the deeper end of the range. All species are from warm waters. Labracherie 
(1972) noted that the occurrence of Didymosella (together with other species) allowed the 
inference of warm marine conditions in the Middle Eocene of south Western France. The 
general pattern of distribution of the family in time and space follows that illustrated by 
Lagaaij & Cook (1973) for other warm water, shallow shelf species, and even for some deeper 
water forms (see Cook & Lagaaij, 1976). This comprises a very wide (sometimes 
circumtropical) distribution during the Eocene and Oligocene followed by a progressive 
restriction of range in a south-easterly direction through time. The occurrence of Recent D. 
pluma off Brazil and in the Gulf of Mexico is of importance, as it indicates that, like the 
genus Adeonellopsis (see Cook, 1973), groups which are supposed to be 'extinct' in the 
western Atlantic may be found to be still living in this area. 

Summaries in French and German 

Nous decrivons la structure de la colonie et la morphologic zooidale trouvees dans les deux 
families cheilostome-cryptocystidiennes des Bryozoaires, la Euthyrisellidae et la 
Didymosellidae. Tous les murs calcifies sont internes, at les coelomes basals (extra- 
zooidales) sont presents dans presque toutes les especes. Dans la famille Euthyrisellidae, la 
communication interzooidale viscerale- a hypostigienne complemente ou meme remplace 

Figs 27 -30 Scanning electron micrographs of communication septulae: (27) Didymosella porosa 
(Stoliczka) BMNH D36788-92, Middle Oligocene, New Zealand, basal septulae (left) xlOO 
(right) x600; (28) Neoeuthyris woosteri (MacGillivray) BMNH 1979.2.1.1, Recent, Western 
Australia, lateral frontal septulae at base of developing avicularian subrostral chamber x315; 
(29) N. woosteri, visceral-to-hypostegal septulae (arrowed) viewed from distal side x!97; (30) 
Tropidozoum cellariiforme Harmer BMNH 1964.3.10.1, Recent, Celebes, visceral-to- 
hypostegal septula in distal-lateral wall of brooding zooid, viewed from proximal side. Note 
suture in wall showing contribution of distal zooid calcification to brooding zooid orifice x368. 




a s 

33 --, 


la sorte intrazooidale la plus commune, cells qu'on trouve dans la Didymosellidae. Dans la 
Euthyrisellidae, une extension du coelome hypostigienne au-dessous du cote basal de la 
paroi frontale cacifiee, est presente dans la majorite des especes. Une espece nouvelle est 
introduite, Tropidozoum burrowsi. La famille Didymosellidae possede un record fossil qui 
s'etend d'Eocene Moyen et a une distribution tres grande du Tertiare au Recent. Nous 
introduisons une nouvelle espece Recente, Didymosella pluma. Tubiporella boninensis Borg 
a ete redefmie. Tubiporella levinseni Borg a ete redecrite et attribue au genre umbonuloide 

Die Koloniestruktur und Morphologic der Zooide der Euthyrisellidae und Didy- 
mosellidae, zweier cryptocystider Familien der Bryozoa Cheilostomata, werden beschrieben. 
Alle verkalkten Wande sind intern, und kolonieweite (extrazooidale) Basalcoelome fmden 
sich bei fast alien Arten. Bei den Euthyrisellidae erganzt oder sogar ersetzt die interzooidale 
(viscerale bis hypostegale) Kommunikation die gewohnlichere intrazooidale, die sich bei 
den Didymosellidae findet. Bei den meisten Arten der Euthyrisellidae ist eine Verlangerung 
des hypostegalen Coeloms an der Basalseite des verkalkten Frontalschildes zu beobachten. 
Eine neue Art, Tropidozoum burrowsi, wird eingefuhrt. Die Familie Didymosellidae ist 
fossil seit dem mittleren Eozan nachgewiesen und ist vom Tertiar bis zur Gegenwart weit 
verbreitet. Eine neue recente Art, Didymosella pluma, wird eingefuhrt und Tubiporella 
boninensis Borg neu defmiert. Tubiporella levinseni Borg wird wiederbeschrieben und in die 
umbonuloide Gattung Reptadeonella venviesen. 


These investigations would not have been possible without the collection and donation to 
the British Museum (Natural History) of well-preserved specimens of 'rare' species. We are 
therefore particularly grateful to Dr P. Bock (Royal Melbourne Institute of Technology, 
Victoria), Dr J.-G. Harmelin (Station Marine d'Endoume, Marseille), Dr J. S. Ryland 
(University College of Swansea), Dr R. E. Wass (University of Sydney, New South Wales) 
and Dr W. J. Woelkerling (La Trobe University, Victoria) for their help. We should also like 
to thank Dr M. Labracherie (Universite de Bordeaux) and Dr P. Taylor (British Museum, 
Natural History) for the loan of fossil specimens. 


Banta, W. C. 1970. The body wall of cheilostome Bryozoa. Ill, The frontal wall of Watersipora arcuata 

Banta with a revision of the cryptocystidea. J. morph. 131 : 37-56. 
Banta, W. C. & Wass, R. E. 1979. Catenicellid cheilostome Bryozoa: 1. Frontal walls. Aust. J. Zool. 

Suppl. Ser. 68 : 1-70. 
Bassler, R. S. 1936. Nomenclatorial notes on fossil and Recent Bryozoa. /. Wash. Acad. Sci. 

24(4): 156-162. 
Boardman, R. S. & Cheetham. A. H. 1973. Degrees of colony dominance in stenolaemate and 

gymnolaemate Bryozoa. In R. S. Boardman, A. H. Cheetham & W. A. Oliver, Animal Colonies. 

Borg, F. 1940. On the genus Tubiporella and on a new boring Bryozoan. Zool. Bidrag. Upps. 

Bretnall, R. W. 192 1 . Neoeuthyris: A new genus to accommodate Euthyris woosteri MacGillivray. Rec. 

Aust. Mm. 13(4): 157-162. 

Figs 31-33 Micrographs of longitudinal thin-sections of epoxy resin embedded specimens of 
Euthyrisellidae: (31) Neoeuthyris woosteri (MacGillivray) BMNH 1979.2.1.1, Western 
Australia, zooids with cuticular basal walls encrusting algae, autozooid (left) and brooding zooid 
with large ovicell (centre), arrow indicates area enlarged in Fig. 32 x30; (32) N. woosteri visceral- 
to-hypostegal communications (arrowed) x!65; (33) Euthyrisella obtecta (Hincks) BMNH 
1890.7.23.38, Torres Straits, brooding zooids (without ova), cuticular basal wall detached at 
right x70. a ascus, o ovicell, op operculum, /frontal process, / tentacles, g gut, h hypostegal 
coelom, v visceral coelom, b basal coelom, as algal substratum. 







Brown, D. A. 1948. Six new Recent and Tertiary Genera of cheilostomatous Polyzoa from New 

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1965. Notes on some Polyzoa with conical zoaria. Cah. Biol. mar. 6 : 435^54. 

1966. Some 'sand fauna' Polyzoa (Bryozoa) from Eastern Africa and the Northern Indian Ocean. 

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1967. Polyzoa (Bryozoa) from West Africa. The Pseudostega, the Cribrimorpha and some 

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Figs 34-36 Micrographs of longitudinal thin-sections of epoxy resin embedded specimens of 
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Queensland, showing large subfrontal hypostegal coeloms x60; (35) Euthyrisell obtecta (Hiricks) 
BMNH 1890.7.23.38, Torres Straits, visceral-to-hypostegal communication (arrowed) and 
minute subfrontal hypostegal coelom x!70; (36) Didymosella larvalis (MacGillivray) BMNH 
1 890.4. 1 7. 1-1 Opt., Spencer Gulf, zooids with cuticular basal walls encrusting algae x40. a ascus, 
o ovicell, op operculum, / frontal process, p primary orifice, 5 secondary orifice, h hypostegal 
coelom, s/subfrontal hypostegal coelom, v visceral coelom, b basal coelom, as algal substratum. 


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1957. The Polyzoa of the Siboga Expedition, Pt. 4. Cheilostomata Ascophora 11. Siboga Exped. 

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Notes on Atlantic and other Asteroidea. 
1. Family Benthopectinidae 

Ailsa M. Clark 

Zoology series Vol 41 No 3 29 October 1981 

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Notes on Atlantic and other Asteroidea. 1. 

Ailsa M. 

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


Synopsis 91 

Introduction 91 

Systematic discussion 91 

Family Benthopectinidae 91 

Genus Cheiraster 107 

Genus Pectinaster 118 

Genus Benthopecten 122 

Acknowledgements 134 

References 134 


Various taxa of the family Benthopectinidae, mainly from the Atlantic, are reviewed. The genera 
Pectinaster and Cheiraster are redefined, Pectinaster being restored to Perrier's concept of 1894 by 
transfer of several nominal species without specialized papularia to Cheiraster, which is enlarged by 
inclusion in it of Luidiaster Studer as a subgenus. Two further subgenera of Cheiraster are described, 
Barbadosaster for Cheiraster echinulatus (Perrier) and Christopher aster for C. mirabilis (Perrier), also 
including C. blakei sp. nov. and two other species. On the basis of older collections and new material, 
mainly from the Pillsbury and Gerda collections of the University of Miami, the morphological and 
geographical ranges of certain species of these two genera and of Benthopecten are extended and several 
names are put in synonymy, tabular keys to the Atlantic species are also included. 


In the course of a major review of the Atlantic species of Asteroidea, assessment of the 
neglected deep-sea family Benthopectinidae proved so complicated as to necessitate a 
preliminary paper to dispose of some of the more complicated taxonomic problems. 

The ordinal position of the Benthopectinidae will be discussed in the main study on 
Atlantic asteroids (Clark & Downey). The family was removed from the Paxillosida to the 
revived order Notomyotida Ludwig, 1 9 1 by McKnight (1975). 

Systematic discussion 
Family BENTHOPECTINIDAE Verrill, 1894 

Archasteridae Benthopectininae Verrill, 1894 : 245-246, 268. 

Archasteridae Pontasterinae Verrill, 1894 : 246-247, 268. 

Benthopectinidae Verrill, 1899:200, 217; Ludwig, 1910:442, 458-461; Fisher, 1911:120-122; 

Verrill, 1914 : 310-31 1; Spencer & Wright, 1966 : U48. 
Plutonasteridae Pontasterinae Verrill, 1899 : 200. 
Cheirasteridae Ludwig, 1910 : 442, 444-447. 

Bull. Br. Mus. nat. Hist. (Zool.) 41 (3) : 9 1-1 35 Issued 29 October 1 98 1 









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This family of primarily deep-water Asteroidea is in need of review, the last comprehensive 
survey being by Ludwig (191 0), intended only as a preliminary treatment, modified by Fisher 
(191 1), while Macan (1938) simply listed the species of Pectinaster and Benthopecten (the 
genera represented in the Murray Expedition collections), summarizing earlier conclusions 
about these two genera. 

Since the present study concentrates on the Atlantic fauna, the genera Acontiaster, 
Myonotus and Nearchaster known only from the Indo-Pacific have not merited attention 
here but are listed below with the other taxa currently recognized together with nomen- 
clatural modifications now proposed. Saraster A. H. Clark, 1916, type and only species, S. 
insignis A. H. Clark, is now synonymized with Nearchaster. It was described without 
comparative remarks or figures and the holotype (from Lower California) has disintegrated 
but the description leaves no doubt in my mind that it was conspecific with N. aciculosus 
(Fisher, 1910), taken at 14 Albatross stations off California (and further north), any 
differences being accountable to the smaller size of the type. 

Additionally, the following names formerly reduced to synonyms are now otherwise 


Benthopecten armatus (Sladen, 1889), NE America Portugal, 2290-2470 m., synonymized 
with B. spinosus Verrill by Macan, 1938, following Verrill, 1899 and Grieg, 1921, treated 
as valid by Farran, 1913 and Mortensen, 1927 but now synonymized with B. simplex 
(Perrier), following Ludwig, 1910. 

B. fischeri Perrier, 1894. NW Africa, 1060-1430 m, synonymized with B. spinosus by 
Macan, 1938, following Grieg, 1921, but now provisionally treated as valid or a possible 
synonym of B. simplex. 

Cheiraster planus Verrill, 1915, West Indies (no details), synonymized with C. mirabilis 
(Perrier) by Downey, 1 973, but now considered as valid. 

Pontaster perplexus Perrier, 1894, NW Africa, 2320-2330 m., synonymized with P. 
tenuispinus (Diiben & Koren) by Ludwig, 1910, but now synonymized with Pontaster 
sepitus Verrill, which is itself now revived from the synonymy of (C.) echinulatus (Perrier), 
to which it was referred by Ludwig, 1910, and newly referred to Cheiraster. 

Pontaster venustus Sladen, 1889, Azores Portugal, 1250-4000 m., with var. robusta 
Sladen, 1889, Cape Verde Is area, depth ?, synonymized with Pectinaster filholi Perrier by 
Macan, 1938, following Ludwig, 1910, but now synonymized with Cheiraster sepitus. 

B. semisquamatus (Sladen, 1889) was treated by Goto, 1914, without explanation as a 
synonym of B. spinosus Verrill, apparently extending Verrill's synonymy of B. semi- 
squamatus var. occidentals to the Pacific semisquamatus itself, despite Ludwig's 
retention in 1910 of this as valid, followed by Fisher (1919 : 210) and Doderlein (1921). 
Okutani (1969) has followed Goto. 

The genera of Benthopectinidae can be distinguished by the tabular key (Table 1). 

Some of the generic limits within the Benthopectinidae are still ill-defined. In Ludwig's 
review (1910), on the strength of the unusual dorsal arm muscles, the present family was 
interpreted as a suborder Notomyota including two families: Cheirasteridae (for Cheiraster 
Studer, 1883,LwzW/asterStuder, 1 884, Pontaster Sladen, 1885 restricted, Pectinaster Perrier, 
1885, Marcelaster Koehler, 1907 and Gaussaster nov., of which Marcelaster was 
synonymized with Luidiaster by Fisher, 1940) and Benthopectinidae (for Benthopecten 
Verrill, 1884 and Pararchaster Sladen, 1885). The latter group was distinguished by the 
presence of odd interradial marginal plates, the superomarginal ones with unusually large 

Fisher (1911) described two new genera, Nearchaster and Myonotus, both with the 


Table 1 Tabular key to the genera of Benthopectinidae. Entries in brackets indicate a slight 
development or tendency. 

12 3 4 5 6 7 89 






M/R + 









M' + 









M + 









R(M) + 









M (+) 




Cheiraster (Luidiaster) 





M -(+) 




Cheiraster (Barbadosasler) 









Cheiraster (Christopher aster) 





M + 




Cheiraster (Cheiraster) 














M/R - 













1 . Interradial suture between the marginal plates: 

A-absent, an odd plate present in each series, the superomarginal one inset on to the disc 

I-often irregular, plates asymmetric or odd plates in some or all interradii but the superomarginal 

one on the periphery of the disc 

R-regular, no odd interr'adial plates (though the sutures are occasionally asymmetric in one or two 

2. Papular areas: 

b-bilobed distally (except in small specimens, R < 30 mm), limited to the arm bases, initiated from a 

single median proximal primary pore 
d-initially double, two lateral areas on each arm base merging proximally in large specimens, R 

> 80 mm, to form a bilobed area, pores relatively small and interstitial 
o-oval and swollen with deepened plates forming well-defined median papularia 
w-widespread over most of disc and at least the proximal parts of the arms 

3. Superomarginal plate alignment: 

D-partly or mainly dorsal, framing the paxillar area in dorsal view 
L-mainly lateral, inconspicuous dorsally 

4. Abactinal plates of disc and arm bases: 

f-flat, usually scale like, with only a small median elevation, if anything 
p-(para-)paxilliform with a more or less well defined median column or convexity 

5. Armament of most abactinal plates: 

M-multiple, with a cluster of spinules, often surrounding a larger median spinelet or spine, 

sometimes totally lacking 
R-reduced, often to a single spinule or spinelet throughout 

6. Central armament of primary and some other proximal plates 
+-large spines 2+ mm long present 

only 1 (rarely 2,3) central spinelets of larger plates enlarged up to c. 1'5 mm 

7. Subambulacral spine number 

8. Dorsal arm muscles: 

p-paired, attached proximally to one or two ambulacral plates and the adjacent marginals, or to the 

abactinal body wall, or both 
u-united in a single mass, attached proximally only to the body wall 

9. Pedicellariae (if present): 

B-bivalved with only two elongated valves, on adambulacral plates only 

F-fasciculate with multiple narrow tapering valves, usually in two, occasionally three, combs on 

adjacent plates (pectinate), usually present on actinal plates, sometimes on abactinal and marginal 

plates, occasionally lacking. 


interradial marginals more or less irregular in position, so that one plate of each series is 
often more or less mid-interradial but the armament of such odd superomarginals is not 
conspicuous. Inevitably, such intermediate taxa prompted him to amalgamate the 
Cheirasteridae with the Benthopectinidae. Apart from synonymizing Pararchaster with 
Benthopecten, he accepted the genera cited by Ludwig. 

Among the genera with no odd interradial marginals, Fisher distinguished Pontaster and 
Pectinaster with 'entire and swollen' papularia from Cheiraster and Luidiaster with 'flat and 
two-lobed' papular areas, separating Pontaster from Pectinaster on the presence of bivalved 
rather than fasciculate pedicellariae and Cheiraster from Luidiaster on the single rather than 
multiple subambulacral spines the only difference evident from a comparison of Ludwig's 
diagnoses. To supplement these criteria, Fisher proposed to use the proximal attachment of 
the dorsal arm muscles to one or two ambulacral plates and the two adjacent marginals in 
Luidiaster but to the dorsal body wall and the superomarginals in Cheiraster. However, in 
1919 he had to modify this after finding that the type-species of Cheiraster, C. gazellae 
Studer, does have tendons to the twelfth and thirteenth ambulacral plates, though 
inconspicuous ones, and these plates lack the enlarged crests of the comparable plates of L. 
hirsutus, the type species of Luidiaster, and even more L. dawsoni. I find that L. hirsutus 
additionally has the attenuated proximal ends of the muscles attached to the dorsal body wall 
just distal to the V-shaped area defined by the two lobes of the papular area, as well as having 
a tendon to a laterally swollen crest of an ambulacral plate (the sixth plate in a specimen with 
R 85 mm). Fisher himself (1940) found a much stouter link between the main part of the 
muscle and the dorsal body wall in the Discovery specimen of Pontaster planeta Sladen, a 
species later referred to Luidiaster (A.M.C., 1962) on account of the V-shaped flat papular 
areas and multiple subambulacral spines, and closely related to L. hirsutus. The muscle 
appears to me to extend nearly to the seventh ambulacral, ending opposite the fourth 
superomarginal, the distalmost papular pores being just proximal to this (as in L. hirsutus), 
although Fisher described it as ending level with the ninth ambulacral. He also noted a 
'rudimentary' tendon to the eighth ambulacral in planeta. In Cheiraster mirabilis (Perrier), 
which has abruptly enlarged fourth superomarginal plates, I find that there is a stout tendon 
to this marginal and the adjoining seventh ambulacral but no connection to the dorsal body 

Clearly, the muscle attachment shows too great a range in these various species for use as a 
character of generic weight. This leaves only the number of subambulacral spines to 
distinguish between Cheiraster and Luidiaster but I think that the alignment of the 
superomarginal plates is also of more than specific significance. Hitherto, this alignment has 
not been regarded as important, possibly because of the considerable range of form in the 
very variable species Pontaster tenuispinus, so that Ludwig suggested synonymy of Pontaster 
venustus with dorsally broad superomarginals with Pectinaster filholi where the marginals 
are almost entirely lateral. However, limitation of Pectinaster to species with laterally 
aligned superomarginals results in a much more natural grouping, since such species also 
have specialized compact papularia, those with dorsally broad superomarginals having 
much more diffuse pore areas potentially or actually bilobed in form with the abactinal 
plates within the areas little modified (compare Figs la & b). Verrill (1915 : 137) discounts 
the importance of the subambulacral spine number in ascribing the specimen he named 
enoplus to Cheiraster despite the two large subambulacral spines of the large holotype (R 
185 mm). Study of a good size range of C. mirabilis now shows that doubling of the 
subambulacral spine number is common in very large specimens, R > 120 mm, and that C. 
enoplus Verrill is a synonym, the holotype sharing the very distinctive abrupt enlargement of 
the fourth superomarginal plates and spines. 

With regard to the Atlantic species of Benthopectinidae, no less than 18 other nominal 
species which have been included in Pontaster, Pectinaster, Cheiraster and Luidiaster need 
to be taken into consideration, namely (in chronological order): 




Fig. 1 Half sections through the base of a ray bisecting the papularium or papular area, showing 
the armament of the second supero- and inferomarginal plates with their proximal and internal 
(hatched) faces, the fully developed spines of the third plates shown by discontinuous lines, (a) 
Pectinaster filholi Perrier, syntype of Pontaster forcipatus Sladen, Challenger st. 44, 45 or 50, 
BMNH no., R c.75 mm; (b) Cheiraster (Christopheraster) blakei sp. nov. Oregon 
st. 4294. R. c. 70 mm. [In (b) the vertical faces of the two marginals are not in the same plane; the 
position of the lobe of the papular area is indicated by the arrows.] 


Pontaster tenuispinus (Diiben & Koren, 1 846) NE Atlantic, Arctic 

Cheiraster echinulatus (Perrier, 1875) West Indies 

C. mirabilis (Perrier, 1881) West Indies 

Luidiaster hirsutus Studer, 1 883 Kerguelen, South Africa 

C. coronatus (Perrier, 1 884) West Indies 

Pontaster sepitus (Verrill, 1 885) NW Atlantic 

Pectinaster filholi Perrier, 1885 NE Atlantic 

Pontaster venustus Sladen, 1 889 Azores, Cape Verde area 

Pectinaster pristinus (Sladen, 1 889) SW Atlantic 

Luidiaster planeta (Sladen, 1 889) E & W Patagonia, Falklands 

Pectinaster vincenti (Perrier, 1 894) West Indies 

Pontaster perp lexus Perrier, 1894 West Indies 

Pectinaster oligoporus (Perrier, 1 894) West Indies 

P. dispar Verrill, 1915 West Indies 

P. gracilis Verrill, 1915 West Indies 

P. mixtus Verrill, 1915 West Indies 

C. planus Verrill, 1915 West Indies 

Luidiaster dubius H. L. Clark, 1 94 1 West Indies 

Of these, Pontaster venustus was synonymized with Pectinaster ftlholi by Ludwig (1910), 
followed by Macan (1938) but is now found to be referable to Cheiraster because of its 
irregular bilobed papular areas and dorsally broad superomarginal plates. The oval, well- 
defined papularia ofP.filholi are constructed of much deeper plates than the other abactinal 
ones, laterally grooved to accommodate the papulae (Fig. la). The same combination of 
less compact actually or potentially bilobed papular areas and relatively broad supero- 
marginals as in P. venustus is also shown by the types of Pontaster sepitus, Pectinaster 
vincenti, Pontaster perplexus, Pectinaster oligoporus, Pectinaster dispar, Pectinaster gracilis 
and Pectinaster mixtus, all of which have been examined. These encompass a size range of R 
26-c. 55 mm. 

Pectinaster pristinus is known only from a single small specimen, R c. 18 mm, from off the 
River Plate. This was in poor condition, the abactinal plates obscured by mud; when this was 
removed, no significant difference could be seen in the armament from specimens ofP.filholi 
of similar small size from the North Atlantic. 

Accordingly, both Pontaster and Pectinaster are reduced in the Atlantic to single species 
(indeed Pontaster is a monotypic genus). 

Following study of many benthopectinids from the U.S. National Museum and M.C.Z., 
Harvard collections, together with the Pillsbury and Gerda material of the University of 
Miami from the tropical Atlantic, I consider that most of the 1 6 remaining names listed 
above (which fall within the scope of Cheiraster and Luidiaster) are synonyms, only five 
being for valid species. This superfluity of names arose mainly from Perrier's confused 
treatment of the Blake collections (1881 and 1884 with afterthoughts in 1894), which was 
partly resolved but further complicated by Verrill's failure to take growth changes into 
account in his review of 1 9 1 5. 

One character of limited use is the R/r ratio, which tends to increase with the absolute size, 
so that species such as C. mirabilis, which may exceed R 120 mm, may achieve an R/r ratio 
of as high as 9/1, from usually 7-8/1 at Re. 100 mm or only c. 5/1 at R 20 mm. However, in 
C. sepitus with a maximum R of probably 70 mm, the ratio stays near 5/1 in specimens 
larger than R c. 30 mm. 

Since individual papulae do not appear to increase disproportionately in size with growth 
of the whole animal, their number must increase in order to maintain adequate respiratory 
function. In most of the species under discussion the first (primary) pore usually appears at R 
10-15 mm near the base of each arm distal to the primary radial plate, which at this small 
size is usually still distinguishable among the other disc plates; further pores, usually smaller, 
are added progressively around it, especially distally and laterally. Only Cheiraster mirabilis 
and C. coronatus (sensu H. L. Clark, 1941) differ from the rest in developing not one primary 



Fig. 2 (a and b) Cheiraster (Christophemster) mirabilis (Perrier). (a) Atlantis st. 3444, 3445, 3447 
or 3449, MCZ 3912(pt), R 25 mm, to show disc and proximal marginal spines; (b) Holotype, 
Blake st. 148, MCZ 10, R 87 mm, papular area (the proximal end lowermost); (c-f) Cheiraster 
(Christopheraster) blakei sp. nov. (c) Atlantis st. 3444, 3445, 3447 or 3449, MCZ 3912(pt), R 
21 mm, for comparison with (a); (d) Atlantis st. 2963F, MCZ 3915, R 112 mm, part of intact 
papular area (most of the spines unnaturally appressed); (e) Blake st. 19, MCZ 2179, R c. 
40 mm, second inferomarginal plate, adjoining actinal area and two adambulacral plates; (0 
Blake st. 295, MCZ 216, R c. 45 mm, denuded papular area showing the spine sockets on two 
midradial plates, the upper one probably the primary radial; (g, h) Cheiraster (Christopheraster) 
horridus Fisher, holotype, Albatross st. 4079 (Hawaiian Is), USNM 21 156, R 35 mm; (g) part of 
denuded papular area showing varied plate size and relatively small pores; (h) armanent of one 
large disc plate. 


pore but a pair of smaller pores offset one each side of the primary radial plate; additional 
similarly small pores appear laterally and distally but few develop in the proximal mid- 
radial area, even at a large size, so that the two lateral pore areas only become loosely linked 
(Fig. 2b & f)- At R > 100 mm, the total pore number in these two species exceeds 100 on 
each ray, far more than in the other species of Cheiraster with their larger papulae, so that for 
instance C. planus at R 100 mm has only c. 50 pores in each area. From the median primary 
pore these develop fairly regularly in this species in a curved double row on each side, 
forming a horseshoe-shape; only at R > 70 mm do the added pores form a more irregular and 
elongated area (Fig. 5). The early stages of this sequence are shown by the illustrated 
holotype and paratype of Pectinaster gracilis (Verrill, 1915, pi. 6, fig. 1 & pi. 15, fig. Ib); the 
smaller paratype (R probably c. 20 mm) has only three pores in a transverse line, the median 
one enlarged, while the holotype (pi. 6) at R 30 mm usually has just two additional pores 
placed distal to the lateral ones forming a very shallow U. Although not stated by Verrill, the 
holotype has most superomarginal spines distinctly longer and stouter than the corres- 
ponding inferomarginal spines a characteristic of C. planus as well as compact 
parapaxillae, uniformly short inferomarginal spinules giving a smooth contour to the plate 
(apart from the main and accessory spines), large single subambulacral spines and relatively 
blunt-angled furrow margins to the adambulacral plates. Consequently, P. gracilis should be 
treated as a synonym of C. planus. 

A similar sequence in the early development of pores is shown in Cheiraster echinulatus, 
Pectinaster vincenti and P. mixtus, also in Pontaster sepitus, P. venustus, P. perplexus, 
Pectinaster oligoporus and P. dispar (and almost certainly occurred in the holotype and only 
recorded specimen of Luidiaster dubius before it reached its final size of R c. 60 mm). In 
these, however, the later developed pores are more irregular in position than those of C. 
planus, forming at first a square or transverse patch (Fig. 4b) and then an elongated bilobed 
area, often asymmetrically so or with the distal pores of the two lobes approximating 
medially (as in the holotype of P. vincenti, Fig. 4g) to form an almost oval area. The 
maximum R in these nominal species does not appear to exceed 70 mm and the pore 
number 36. Most specimens taken are in the R 20-40 mm range. (Genital pores first appear 
in Cheiraster at R 10-15 mm close to the first superomarginal plates.) 

The holotypes of C. echinulatus, mixtus and vincenti have R 16 mm, 26 mm and 42 mm 
and the pore number in each area is 1-3, 3-5 and c. 12 respectively, an increase compatible 
with the size differences. All show the following characters: the abactinal plates are very 
mixed in size, polygonal, abutting fairly closely together, distinctly flattened in the two 
smaller specimens but somewhat convex in the larger one, especially on the papular areas, 
armed with numerous (up to c. 30) short 'spinules' of superficially granuliform shape 
(though with height/breadth >2/l in the largest) arising from most of the surface of the plate 
and encircling in a double ring on many plates a slender needle-like spinelet; the supero- 
marginal plates are broad in dorsal view, the two opposite ones together taking up c. 50% of 
the total arm breadth; the marginal spines are relatively slender, most inferomarginal ones 
longer than the corresponding superomarginal spines, while several of the inferomarginal 
spinules below the spine are more or less enlarged into accessory spinelets so as to give a 
rough profile to the body; the furrow margins of the proximal adambulacral plates form an 
acute angle prolonged into the furrow (the distal plates being truncated) and the sub- 
ambulacral spines are multiple (except perhaps in the small holotype of C. echinulatus 
where Perrier described a complete circle of 10-12 peripheral spines (including those 
projecting over the furrow, around a larger central spine but all the subambulacral spines are 
now [1980] lost or displaced). The coincidence of all these characters is convincing evidence 
that the three are conspecific, so that Cheiraster vincenti and Pectinaster mixtus become 
synonyms of C. echinulatus. The type localities of C. echinulatus and vincenti are near the 
neighbouring Lesser Antillean islands of Barbados and St Vincent; that of mixtus is off 
Havana, Cuba (Verrill's 'West of Florida' deriving from a misreading of the coordinates for 
Albatross st. 2341). Many other specimens from intermediate and western Caribbean 
localities to a confirmed depth of 567 metres show variation not only in the convexity of the 



f OOO 

e ooo 

d oooo 


abactinal plates and arrangement of the pores but also in the size of the pores, which reach a 
maximum number of 36 in a specimen with R 65 mm. The frequency of pedicellariae, as 
usual, is rather variable; there are some pectinate actinal ones is c. 50% of specimens but 
only c. 5% (including the holotypes of C. echinulatus and mixtus) have a few fasciculate 
inferomarginal pedicellariae adjoining the adambulacrals. The holotype of P. oligoporus 
from the Leeward Is. in 275 m is in very poor condition but its flattened abactinal plates and 
the prolonged furrow angles of the adambulacrals show that it too is conspecific with and a 
synonym of C. echinulatus. 

For the rest, the type localities of C. sepitus, venustus (with var. robusta), perplexus, dispar 
and dubius are scattered all over the North Atlantic, respectively: S of Cape Sable, Nova 
Scotia, 1570 metres, the Azores, 1650 metres (Cape Verde Islands, depth unknown), off Cap 
Blanc, W. Africa, c. 2325 metres, the Leeward Is, 1260 metres and S of Cuba, 485 metres. 
(For Pectinaster dispar Verrill cited the source as an Albatross station, locality unknown, but 
the labels now with the specimen indicate st. 2751, which is 1654'N, 63 12'W.) The 
relevant type specimens have R measurements and pore numbers as shown in table 2. 

Table 2 




v. robusta 









35+ (745-50) 









The figures for C. sepitus are from what may be called a 'topotype' in the USNM, no. 24525, since the 
two syntypes (from Albatross st. 2072) are now in a state of disintegration and were never properly 
described by Verrill. This specimen was named P. sepitus in Verrill's hand and was from a station 
(2075) with almost identical coordinates and depth. 

Although sharing with C. echinulatus the dorsally broad superomarginal plates taking up 
50% or more of the arm breadth, these all differ from that species in several features. The 
abactinal plates are more loosely joined and consistently markedly convex and parapaxilli- 
form, with more elongate and somewhat pointed spinules usually numbering only 5-15, 
clustered together on the central (highest) part of the plate, though again often surrounding a 
central spinelet several times longer; the superomarginal plates, though with a similar mean 
breadth to those of C. echinulatus, are somewhat longer and often more convex, numbering 
only c. 1 6 at R 20 mm or c. 27 at R 50 mm, compared with c. 20 and c. 30 at the same sizes in 
echinulatus; the marginal spines are shorter and much stouter basally, being conical in 
shape, though again the inferomarginal spines tend to be longer than the superomarginal 
ones and the upper inferomarginal spinules are likewise more or less elongated; the 
subambulacral spines are usually single, stout-based and conical (as usual in all these species 
similar in form to the marginal spines) and pedicellariae are rarely present at all, even on the 
actinal plates they were only found in less than 10% of specimens. Possibly in larger 
specimens, R > 50 mm, the arms remain relatively shorter than in C. echinulatus, R/r not 
much exceeding 5-0/1 where echinulatus may reach 6/1, but this remains to be seen from 

Fig. 3 Cheiraster (Barbadosaster) echinulatus (Perrier) (a) Holotype, Hassler, Barbados, Paris 
Museum (PM) 3481, R 16mm, disc and arm bases from above, the primary radial and 
interradial plates stippled, crystal bodies indicated on a few larger abactinal and marginal plates 
(the centre of the disc rubbed almost bare of spinules); (b, c) the same, part of a ventral 
interradius and the fourth and fifth adambulacrals with adjacent inferomarginal plates, most 
armament lost, the subambulacral spine drawn on the fifth plate appressed; (d-f) abactinal 
spinules of: (d) the holotype of C. echinulatus, (e) holotype of Pectinaster mixtus Verrill, R 
26 mm and (0 specimen from Gerda st. 628, R 66 mm. The scale equals 2 mm for (a), 1 mm for 
(b) and (c) and c. 0-5 mm for (dHO- 

104 A.M.CLARK 

further intact specimens. (Perrier's implication that there is more than one subambulacral 
spine in the holotype of perplexus is not borne out by re-examination. It may also be noted 
that he gave the pore number as 'une quinzaine' but removal of the spinelets in one area 
revealed the higher number of 2 1 . Yen-ill's estimate for the type of dispar was also too low). 

With all these features in common, I do not think that the wide spread of pore numbers 
from 2 1 at R 52 in the type of perplexus to only 8 in that of the geographically close venustus 
var. robusta at R 46 mm is significant. The 'topotype' of seprtus has as many as 1 5 at R 
<40 mm. Other similar specimens, with R 19-46 mm collected at five Pillsbury stations off 
Venezuela and two Discovery ones from the Azores, all of which I consider are conspecific 
with C. sepitus, have 0-12 pores in individual areas, often with a range of three or more in 
different radii of the same specimen, one with R 41 mm having from 6 to 12 pores in the five 
radii. Consequently, P. venustus, perplexus, dispar and L. dubius are now referred to the 
synonymy of C. sepitus, the oldest name. 

All the West Indian Pillsbury specimens now referred to C. sepitus are from depths greater 
than 1000 metres, as with the type locality of P. dispar, which casts doubt on the depth 
record of only 485 metres for the holotype of L. dubius. 

A further nomenclatural problem remains to be resolved concerning the two species 
generally known as C. mirabilis and C. coronatus, at least sensu H. L. Clark, 1941 . 

In his preliminary account of the Blake asteroids, Perrier (1881) described Archaster 
mirabilis (stations unspecified) firstly (paragraphs 1-4) from a specimen with R/r given as 
87/12 mm, only 6-10 spines around the anus and otherwise granulous abactinal plates; he 
then commented on another specimen with R/r 45/12 mm. In the full report (1884) the first 
specimen is redescribed at the beginning of the account of A. mirabilis, the station given as 
148, St Kitts, and the second specimen is separated off as holotype of Archaster insignis 
(since referred to Dytaster in the Astropectinidae), as noted by Verrill (1915). Other 
specimens from 46 different stations were also included under the name mirabilis but the 
one with R 87 mm from st. 148 must be the holotype. 

Also in 1884, Perrier described Archaster coronatus from a specimen with R 65 mm, R/r 
8/1, citing stations 2 and 19. The sample from st. 19 is in the MCZ, Harvard, labelled as 
syntypes but the largest specimen is badly broken and its R probably did not exceed 40 mm, 
so, as Verrill (1915) assumed, the holotype must be from st. 2, off Havana, Cuba, and in the 
Paris Museum. In 1894, Perrier inadmissably redescribed the holotype of mirabilis (R/r 
slightly modified to 85/10) as 'C. coronatus premier type', as Verrill (1915) noted when he 
repeated Perrier's 1884 description for coronatus, ranking it as a subspecies of mirabilis. H. 
L. Clark (1941) satisfactorily distinguished two West Indian species ofCheiraster with spines 
several mm long on the disc in the following terms: 

mirabilis coronatus 

1. Paxillae very low and closely covered 1. Paxillae more convex with fewer and 
by a group of minute spiniform more unequal granules, one (or 
granules, none of which are enlarged or sometimes more) of them elongated 
lengthened into spinules [spinelets in into a spinule. 

my terminology.] 

2. 3-10 relatively conspicuous spines 2. Spines up to 3-4 mm long but never 
3-8 mm long at the centre of the disc. forming a conspicuous isolated group. 

3. Fourth superomarginal spine up to 3. Fourth spine up to 5 mm long. 
10-12 mm long. 

After examining the Blake and Atlantis specimens in the MCZ and the Gerda and Pillsbury material I 

can add the following: 

1 a. Abactinal plates too low to be called 1 a. Armament of abactinal plates 

'paxilliform'; armed with up to consisting of up to c. 25 (mean c. 10) 

c. 35 (mean number c. \ 5) short distinctly elongated radiating spinules, 

pointed spinules in compact groups. the longer central spinelets graduated 

to the disc spines. 



2a. The big disc spines closely clustered 2a. 

in the centre, not extending to the 
primary radial plate on the arm base; 
usually 5-15 spines, rarely >20. 

3a. Fourth (rarely the fifth) superomarginal 3a. 

plate and spine abruptly enlarged, the 
corresponding inferomarginal plate 
reduced and often spineless. 

4. Marginal plates covered with fairly 4. 
uniform short pointed spinules, only 

a few on the lower face of the 
inferomarginals enlarged to spinelets; 
profile superficially smooth. 

5. Furrow margins of adambulacral plates 5. 
rounded or with obtuse angles. 

Big spines spaced, extending on to the 
papular areas in larger specimens and 
graduating to the central paxillar 
spinelets, usually > 10 even at R only 
30mm, up toe. 75atR 100mm. 
In larger specimens the fifth or sixth 
marginal spines the largest and 
graduated to the adjacent spines. 

Several more or less elongated 
spinules/spinelets at the upper end of 
the inferomarginals giving a shaggy 
appearance and rough profile. 

Furrow margins 90 or acute, angles 
prolonged into the furrow. 

Unfortunately, Perrier's 1884 description of C. coronatus described the abactinal armament 
as including c. 15 spines around the anus and the fourth superomarginal spine as much 
larger than the others (my italics). Although 1 5 disc spines is a high number for C. mirabilis 
at R 65 mm it is not unprecedented (see Table 4). Accordingly, it seems unavoidable that the 
name coronatus should be referred to the synonymy of mirabilis, as was done by Downey 
(1973), following Ludwig (1910), though evidently without appreciation of the existence of 
the two species recognized by H. L. Clark. 

A new name is therefore needed for the species with less conspicuous, more numerous and 
graduated disc spines and graduated superomarginal spines. Although H. L. Clark has the 
credit for distinguishing the species, knowing his antipathy to the use of personal names I 
propose the name C. blakei after the ship which first collected it. As the MCZ specimens 
from Blake station 19 are in poor condition, a large specimen from Gerda st. 918 (between 
Florida and the Bahamas), described on p. 113 has been selected as holotype. 

Turning to the species of Luidiaster (other than L. dubius disposed of above), the smallest 
specimen available of L. hirsutus from South Africa has R c. 35 mm (R/r 4*5/1) and c. 30 
pores/area but a primary median pore is distinguishable, as in Cheiraster sensu stricto. The 
pore areas are V-shaped with mostly two rows of pores in each limb whereas larger 
specimens, R c.80 mm, have 1 10-160 pores in a more irregular but still bilobed area. The 
type series of L. planeta from Chile, R 26-37 mm, shows 9-14 pores and the primary ones 
are distinct. These two southern species, L. hirsutus and L. planeta, both have the 
superomarginals inconspicuous in dorsal view, aligned mainly on the sides of the arms, 
unlike the species of Cheiraster. They also have two or more large subambulacral spines 
(though at R 35 mm in L. hirsutus the two spines are more or less markedly unequal) and H. 
L. Clark (1926) found single spines in his smallest specimen, R 16 mm). However, the West 
Indian Cheiraster echinulatus has two or even three subambulacral spines from a small size, 
R. c. 20 mm, while large specimens of C. mirabilis and C. blakei (R > 100 mm) often 
develop a second spine, though most individuals have only one on most plates. (Second 
spines usually appear first on the distal half of the arm and on the very first plate.) Although 
large specimens of C. mirabilis and blakei may have the superomarginal plates relatively 
narrow dorsally, they still form a small border to the paxillar area and in C. echinulatus they 
are consistently broad. 

I doubt whether the alignment of the superomarginal plates unsupported is of sufficient 
weight to justify a generic distinction for Luidiaster hirsutus so that Luidiaster is now treated 
as a subgenus of Cheiraster. Both names were introduced by Studer, 1883, who separated 
them, and even referred Luidiaster to the Astropectinidae, because the specimens of 
Cheiraster gazellae had the dorsal body wall sufficiently transparent to show the underlying 
muscles, which he took for gonads. However, he failed to name a type species for Luidiaster 
until 1 884, from which date only the name is valid. 

106 A.M.CLARK 

The main characters by which the seven Atlantic species of Cheiraster can be distin- 
guished are summarized in Table 3. (The number of adambulacral plates corresponding to 
the first ten inferomarginals does not seem to me to be significant in Cheiraster, though it 
may be useful for certain species of Benthopecten.) Synonymies and ranges for five of these 
species are given on pp. 1 12-8. 

Table 3 Tabular key to the Atlantic species of Cheiraster sensu lato. Brackets signify occasional or 
partial conditions. 

1 2 3 4 5 6 7 8 9 10 11 12 












c. 130 




L (0 











20/27/c. 35 

A c 










c. 5 


A c 






P 2,3 (1,4) 






A c 








c. 100 




A c 








c. 60 




A f 








c. 70 

*C. planeta. In the type series from S Chile, the furrow margins are acute but not in the Falkland Is specimen. 
fC. sepitus. Only five out of 60+ specimens examined have any actinal pedicellariae. 

1. R/ratR50+mm. 

2. Number of superomarginal plates at R 20, 40 and 60 mm (some obtained by extrapolation) 

3. Disc spines: 

L-few and relatively very large, rarely > 15, restricted to centre of disc; no central paxillar spinelets 

graduating to the spines: 
M-moderately large centrally but not more than ^ r, spaced and often extending on to the papular 

areas, graduating in size down to the central paxillar spinelets 
A-absent, only central paxillar spinelets on many disc and mid-radial arm plates, not > 1 5 mm long 

4. Proximal abactinal plates: 

c-convex, parapaxilliform, butting loosely together, spinules clustered on the raised area, mean no. 

10-12 or less 
f-flat or low convex, polygonal, butting more or less closely together, spinules covering most of the 

surface and more numerous, mean no. 1 5-20 

5. Papular areas: 

PB-initiating from a single large median primary pore, becoming irregularly bilobed, sometimes 

indistinctly so 
PC-initiating from a single large median primary pore, becoming crescentic (horseshoe-shaped), 

V-shaped or bilobed with long lobes 

D-double, initiating from a pair of pores, one each side of the mid-line, forming two diffuse lobes, 
merging proximally only in large specimens, R > 70 mm 

6. Largest marginal spines: 

a-Fourth superomarginal spine abruptly and conspicuously enlarged, fourth inferomarginal plate 

reduced and often lacking a spine 
e-Marginal spines fairly even in size or the fourth to seventh (according to size, only the fourth in 

smaller specimens) enlarged but graduated 

7. Marginal spines, relative length: 

I-most inferomarginal spines longer than the corresponding superomarginal ones 
S-most superomarginal spines longer (at least on the proximal half of the arm) 

8. Inferomarginal spinules at the upper end of the plate: 

g-graduated, several becoming more or less elongated, graduating to the accessory spine(s) or 

spinelets, profile rough 
u-fairly uniform, only some enlarged spinelets on the lower face of the plates, profile fairly smooth 

between the spines 


9. Furrow margins of adambulacral plates: 

P-with acute angles prolonged into the furrow, < 90 
R-rounded or with angles obtuse or right-angled 

10. Subambulacral spine number 

11. Pedicellariae: 

12. Maximum recorded R 

As a result of these comparisons between the Atlantic species, two further supraspecific 
groups can be recognized within the genus Cheiraster. 

Firstly, the essentially dual nature and different ontogeny of the papular areas in 
Cheiraster mirabilis and C. blakei compared with the other species of Cheiraster, in my 
opinion justifies at least a subgeneric distinction for these two species. The relatively small 
size of the papulae themselves, the marked size difference of some of the plates (including the 
primary ones) on the disc and arm bases and the development of spines several millimetres 
long on some of these plates provide supporting characters. At least two of the non- Atlantic 
species currently listed as Luidiaster can be placed in this subgenus, for which I propose the 
name Christopher aster. These are: Pontaster oxyacanthus Sladen, 1889, from Japan, which 
is very similar in abactinal armament to C. blakei and likewise has the small papular pores 
almost entirely offset from the midradii, and Cheiraster horridus Fisher, 1906, from the 
Hawaiian Islands, also with very small lateral papulae, a marked size difference of some of 
the abactinal plates and more or less enlarged spines on the larger plates (Fig. 2g & h). The 
holotype of oxyacanthus at R 73 mm has two large subambulacral spines on most plates, 
while that of horridus at R 35 mm has two subequal spines, so that the tendency to develop a 
second subambulacral spine, more or less accelerated in different species, may also be used to 
distinguish the subgenus. 

Secondly, the unusual combination in C. echinulatus of rather low, polygonal, closely- 
fitting abactinal plates armed over most of their surface w'th relatively numerous short 
spinules, dorsally broad superomarginals, unlike Cheiraster (Luidiaster), usually slender 
marginal spines, unlike the conical ones of Cheiraster (Cheiraster), and multiple sub- 
ambulacral spines in all but the smallest specimens (R <20 mm), also justifies a subgeneric 
distinction I believe. The name Barbadosaster (again reflecting the type locality of the type 
species) seems appropriate. In this case the subgenus appears to be monotypic. One non- 
Atlantic species of Cheiraster Pontaster teres Sladen, 1889, from the Banda Sea, East 
Indies, superficially resembles C. (Barbadosaster) echinulatus in the slender marginal spines 
and multiple subambulacrals but differs markedly in the pore areas. 


Archaster(pt.) Perrier, 1875 : 348 [1876 : 268] (A. echinulatus); 1884 : 256; Verrill, 1885a : 151. 
Cheiraster Studer, 1883 : 129; 1884 : 49-50; Perrier, 1885 (pt.) : 70; 1894 : 275; Fisher, 1906 : 1040; 

Ludwig, 1910:440, 454^57; Fisher, 1911:120, 123; Verrill, 1915:123-124; Fisher, 

1919: 190-191; H. L. Clark, 1941 : 26; Downey, 1973:40^1; Jangoux, 1978:95, Walenkamp, 

Luidiaster Studer, 1883 : 131 [nom. nud., no species named]; 1884 : 46^7; Ludwig, 1910 : 440^*41, 

451-454; Fisher, 1911 : 127; Verrill, 1915 : 149-150; Fisher, 1940 : 96; H. L. Clark, 1941 : 28. [Type 

species: Luidiaster hirsutus Studer, 1884.] 

Pontaster (pt.) Sladen, 1885 : 610; 1889 : 23-27; Verrill, 1894 : 247; 1895 : 131. 
Acantharchaster Verrill 1894 : 268-269. [Type species: Archaster dawsoni Verrill, 1880.] 
Pectinaster: Verrill, 1915:137-139; Downey, 1973:43; Walenkamp, 1979:26. [Non Pectinaster 

Perrier, 1885.] 

TYPE SPECIES. Cheiraster gazellae Studer, 1 883, by subsequent designation by Ludwig, 1910. 








.Q. - 





q . 





Fig. 4 Pore arrangements, showing presumed primary radial plates, (a-h) Cheiraster 
(Barbadosaster) echinulatus (Perrier). (a) Blake st. 45, MCZ 2181, R 27 mm; (b) Albatross st. 
2322, USNM 10109, R c. 30 mm; (c) Blake st. 36, MCZ 24, R c. 20 mm (with proximal 
superomarginals); (d-f) Atlantis st. 298 1C or 2982C, MCZ 3917, R 48, 47 and 51 mm; (g) 
holotype of C. vincenti Perrier, Blake st. 231, R c. 42 mm; (h) Atlantis st. 3480 or 3482, MCZ 
3924, R c. 55 mm. (i-r) Cheiraster (Cheiraster) sepitus (Verrill), (i) 'topotype', Albatross st. 
2075, USNM 24525, R c. 37 mm; 0) holotype of Pectinaster dispar Verrill, Albatross st. 2751, 
USNM 18468, R c. 50mm; (k) Albatross st. 2531, USNM 12062, R 35-40 mm (with 
proximal superomarginals); (1) holotype of Pontaster perplexus Perrier, Talisman 1883, st. 96, 
PM 3378, R 52 mm; (m, n) holotype of Luidiaster dubius H. L. Clark, Atlantis st. 3326, MCZ 
38 1 5, R c. 60 mm; (o, p) Princesse Alice st. 743 (Azores), MCZ 2689, R 22 mm; (q) same sample 
(named Pontaster venustus by Koehler, 1909), R c. 33 mm; (r) Talisman 1883, st. 93, MCZ 30, 
R c. 45 mm (named P. venustus by Perrier, 1894). 



(In fact, the spelling of the specific name in 1883 was Gazella, emended in 1884 to the 

A genus of Benthopectinidae without odd interradial marginal plates; abactinal plates 
usually parapaxilliform, more or less convex but flatter on the arms and sometimes all low, 
armed with several spinules, in most species many midradial and disc plates with a larger 
central spinelet, sometimes a spine > 2 mm long; papular pores limited to a bilobed area at 
the base of each arm, or sometimes a pair of lateral areas loosely integrated at their proximal 
ends medially, rarely the two lobes not well defined or asymmetrical, undeveloped in 
younger animals, R <c. 30 mm, only the plates in the median proximal part of the area 
somewhat deepened; superomarginal plates aligned partially on the dorsal surface and 
forming a more or less broad frame to the paxillar area (except in Cheiraster (Luidiaster) 
where they are inconspicuous), armed usually with a single conical or spike-like spine on the 
dorso-lateral convexity; inferomarginal plates sometimes with more than one large spine or 
graduated spinelets below the main spine; adambulacral plates with the convex or angular 
furrow margins armed with a fan of slender furrow spines, the arc usually continuous with a 
few similar but smaller proximal and distal spines, partly surrounding one, two, or several, 
more or less enlarged subambulacral spines, similar in shape and size to the marginal spines; 
fasciculate pedicellariae usually present, at least on some actinal plates, often also 
abactinally and on the lower faces of the inferomarginals adjoining the adambulacrals, rare 
in a few species (e.g. C. sepitus}, mostly bipectinate (i.e. arising from two adjacent plates) but 
the inferomarginal ones often lack an adambulacral component. Dorsal muscle bands of 
arms are attached proximally to one or two ambulacral plates, which may have modified 
ridges, the adjacent marginals and sometimes also to the dorsal body wall distal to the 
papular area. 

Subgenus CHEIRASTER Studer, 1883 

A subgenus with abactinal plates parapaxilliform convex and rounded, even those on the 
disc only abutting loosely together, most proximal and mid-radial ones armed with a central 
spinelet among the smaller spinules; papulae including a median proximal primary one, 
distinct from the rest (except in some larger specimens, R > 50 mm) by its larger size and 
position, which is at first distal to the primary radial plate though may become dissociated 
from it as other plates develop, the later developed pores at first forming a median patch and 
then extending into a distal lobe on each side; superomarginal plates forming a more or less 
broad frame to the paxillar area in dorsal view; marginal spines conspicuous, conical or 
spike-like, some species with a second but smaller inferomarginal spine; adambulacral plates 
predominantly with single subambulacral spines, similar in shape to the marginal ones, 
though the first plate and some distal plates may develop a second large spine. 

TYPE SPECIES. Cheiraster gazellae Studer, 1883. Philippines NW Australia: 360-470 

OTHER SPECIES INCLUDED. In chronological order: 

Archaster sepitus Verrill, 1885 

Pontaster pilosus Alcock, 1893 

C. inops Fisher, 1906 

C. snyderi Fisher, 1 906 

C. niasicus Ludwig, 1910 

C. ludwigi Fisher, 1913 

C. triplacanthus Fisher, 1913 

C.planus Verrill, 1915 

C. diomedeae Fisher, 1917 

C. weberi Doderlein, 1 92 1 

C. otagoensis McKnight, 1973 

North and Central Atlantic 

South of India 

Hawaiian Is to Sri Lanka 

Hawaiian Is-Nicobar Is 



Celebes & South-east Africa 

Gulf of Mexico-French Guiana 



South New Zealand 

485-3700 metres 
1080-1 3 10 metres 
430- 1250 metres 
420- 16 70 metres 
485-930 metres 
12 70 metres 
1050- 1280 metres 
550-715 metres 
770 metres 
470 metres 
1000- 12 80 metres 

(A specimen from offNatal with R c. 40 mm referred to C. triplacanthus (A. M. Clark, 1977), 

110 A.M.CLARK 

has distinctly enlarged primary papular pores; Fisher (1919) noted only that the pore areas in 
the holotype (R 49 mm) are 'double and joined only at the adcentral end'.) 

Subgenus LUIDI ASTER Studer, 1884 

A subgenus of Cheiraster with abactinal plates parapaxilliform, even those on the disc 
rounded and abutting loosely together or more or less spaced, most of the larger plates, even 
laterally on the arms, armed with a central spinelet among the smaller spinules; papulae 
including a median proximal primary one but this probably soon became indistinguishable 
as the number of pores increases and the areas spread, though are still bilobed; supero- 
marginal plates aligned mainly laterally, inconspicuous dorsally; marginal spines con- 
spicuous, conical or spike-like, usually at least two on most inferomarginal plates; 
adambulacral plates with two or more large subambulacral spines, the furrow angle variable. 

TYPE SPECIES. Luidiaster hirsutus Studer, 1884. Kerguelen, South Africa & Bouvet I.; 
240-550 metres. 

OTHER SPECIES INCLUDED. In chronological order: 

Archaster dawsoni Verrill, 1 880 Bering Sea-British Columbia 1 00-290 metres 

PontasterplanetaSladen, 1889 S. Chile, Argentina-Falklands 350-450 metres 

Cheiraster gerlachei Ludwig, 1 903 Antarctic 1 60-8 1 metres 

Pectinaster robustus A. H. Clark, 1917f Chile 2410 metres 

Luidiaster tuberculatus Djakonov, 1 929 Japan Sea 1 40-850 metres 

tSeep. 121. 

Subgenus BARBADOSASTER nov.* 

A subgenus of Cheiraster with most proximal abactinal plates polygonal in outline, fitting 
closely together and not markedly convex, sometimes almost flat, armed with relatively 
numerous short spinules, on the larger plates often forming two rings around a central 
spinelet; papulae including a median proximal primary one, usually distinct from the rest by 
its size and position, the later developed pores forming at first a median patch, then 
extending distally into a lobe on each side, though the two lobes may be unequal or ill- 
defined, hardly separated medially; superomarginal plates forming a broad frame to the 
paxillar area in dorsal view; marginal spines slender, not very stout basally, single, though 
some enlarged spinelets develop below the inferomarginal spine; most adambulacral plates 
with more than one subambulacral spine, except in very small specimens, R < 20 mm, the 
furrow angle of the proximal plates markedly acute. 

TYPE SPECIES. Archaster echinulatus Perrier, 1875. South Florida Caribbean; 150-570 
metres. Monotypic. 

Subgenus CHR1STOPHERASTER nov.* 

A subgenus of Cheiraster with the abactinal plates parapaxilliform (but in C. mirabilis only 
slightly convex), distal plates more or less spaced, the proximal plates mostly fitting fairly 
closely, very variable in size, the primary central, radial and interradial plates particularly 
enlarged, some or most of the larger plates armed with a central spine >2 mm long, often 
conspicuous, surrounded by small spinules, some other plates may have a central spinelet 

*Named after the type localities of the relevant type species: Barbados for C. (B.) echinulatus and St. Christopher's 
(Kitts) for C. (C.) mirabilis. 


but not in C. mirabilis; no primary midradial papular pore, the first-formed papulae a pair, 
each latero-distal to the primary radial plate, later pores developing laterally and distally, the 
two pore areas on each ray only loosely meeting proximally in larger specimens to form a 
single diffuse bilobed area with few, if any, midradial pores, the pores themselves relatively 
small and hardly visible with the naked eye, relatively numerous (c. 100 at R 100 mm on 
each ray); superomarginal plates partly aligned dorsally, forming a narrow frame to the 
abactinal area but becoming inconspicuous in the largest specimens (R > 120 mm); marginal 
spines large and spike-like; adambulacral plates with one or two large subambulacral spines, 
sometimes two only in the largest specimens, furrow angles variable in prominence. 

TYPE SPECIES. Archaster mirabilis Perrier, 1881. Florida Windward Is; 380-730 metres. 
OTHER SPECIES INCLUDED. In chronological order: 

Cheiraster blakei sp. nov. Florida-Northern Brazil 250-1030 metres 

Pontaster oxyacanthus Sladen, 1 889 Southern Japan 630 metres 

C. horridus Fisher, 1906 Hawaiian Is 260-330 metres 

This leaves a residue of the following species of uncertain affinities which have been referred 
to Cheiraster or Luidiaster. 

Cheiraster pedicellaris Studer, 1883. NE of Australia; 1000 metres. The holotype has R 
c. 30 mm and is probably a young specimen of C. gazellae according to Ludwig (1 9 10). 

Pontaster subtuberculatus Sladen, 1889. SE of Australia; 1740 metres. The holotype has R 
only c. 24 mm. Like Christopher aster it has no mid-radial papulae, the 17-21 pores 
forming two irregular longitudinal rows on each side, but the pores are relatively 
large and there is no sign of any enlarged spinelet or spine on any abactinal plates. The 
arms are unusually short for a Cheiraster and the single marginal spines are remarkably 
short and blunt. 

Pontaster teres Sladen, 1889. Indonesia; 260 metres. The holotype has R 42mm and 
superficially resembles C. (Barbadosaster) echinulatus, the superomarginals being 
relatively broad and the marginal spines unusually slender. However, denuding part of the 
paxillar area shows an unusual combination of larger convex plates with flattened inter- 
stitial ones between, fitting closely together except where the relatively large papular pores 
are developed, at the sides of the arm bases. The subambulacral spines are multiple. 

Pontaster trullipes Sladen, 1889. Philippines; 1920 metres. The holotype has R only 27 mm. 
The primary radial and interradial plates are enlarged, as in Christopheraster, but no large 
spines occur and the papulae form a median group, in which no primary one is distinct. 

Pontaster cribellum Alcock, 1893. Laccadive Sea; 2200 metres. The holotype has R 45 mm. 
It is said to resemble P. subtuberculatus in lacking spinopaxillae and having relatively 
broad superomarginals but the papularia are small and compact, with only 5 or 6 papulae 
in each and the marginal spines are sharp. 

Marcelaster antarcticus Koehler, 1907 (referred to Luidiaster by Fisher, 1940). South 
Orkney Is.; 3250 metres. R in the type series is up to only 33 mm. The proximal plates 
appear rather variable in size in the poor drawing and there are some relatively large disc 
spines but the papulae were not described at all so the position must remain uncertain. 

Cheiraster granulatus Ludwig, 1910. 'Indian Ocean'; depth not given. Ludwig's Valdivia 
report was never completed and the only information on this barely established species is 
in the key distinguishing it from C. subtuberculatus by the numbers of inferomarginal and 
furrow spines and the occurrence of actinal pedicellariae, which could all be correlated 
with a size difference, if it was appreciably larger than the small type of C. subtuberculatus. 

112 A.M.CLARK 

Cheiraster yodomiensis Goto, 1914 (referred to Luidiaster by Hayashi, 1973). Japan; 160 
metres. The holotype has R 1 70 mm and closely resembles a Nearchaster in the many disc 
spines, laterally aligned superomarginals and great extent of the papular pores along the 
sides of the paxillar areas. Also the interradial superomarginal spines are relatively large 
and possibly asymmetrical in some interradii, judging from the photograph. 

Cheiraster richardsoni Fell, 1958. Cook Strait, New Zealand; 730 metres. The holotype has 
R 3 1 mm. The superomarginals are broad dorsally and the subambulacral spines single. 
The papulae were undescribed. 

Cheiraster monopedicellaris McKnight, 1973. Northern New Zealand; 720-770 metres. The 
holotype has R 39 mm. The superomarginal plates are very broad but some abactinal 
plates have enlarged spinelets, unlike C. subtuberculatus. The 3-5 papulae are said to be 
in a more or less V-shaped arrangement. 

The complex synonymies of the North and Tropical Atlantic species are as follows: 

Cheiraster (Barbadosaster) echinulatus (Perrier) 
Figs 3a-f, 4a-h 

Archaster echinulatus Perrier, 1875 : 348 [1876 : 269]; 1884 : 263-264, pi. 10, fig. 4. 

Archaster mirabilis (pt) Perrier, 1 884 : 258-259 (sts 1 43 & 23 1 ). 

ICheiraster mirabilis: Perrier, 1 894 : 276-278, pi. 20, fig. 4. 

Cheiraster vincenti Perrier,.! 894 : 275-276. 

Cheiraster echinulatus: Perrier, 1894 : 278; Verrill, 1915 : 129-130 (non pp. 131-133, pi. 14, fig. 3, pi. 

19, fig. 2, pi. 25, fig. 1 ? = C. blakei); Downey, 1973 : 42, pi. 13, figsC, D. 
Pontaster oligoporus Perrier, 1894 : 293. 
Luidiaster vincenti: Ludwig, 1910: 452, 453. 
Pectinaster echinulatus: Ludwig, 1910 : 449. 
Pectinaster mixtus Verrill, 1915 : 140-145, pi. 6, fig. 2, pi. 15, fig. 2, pi. 17, fig. 1; Macan, 1938 : 349 

(listed); Downey, 1973 : 43^4, pi. 14, figsC, D. 

Pectinaster vincenti: Verrill, 1915 : 139-140; Macan, 1938 : 349 (listed). 
Pectinaster oligoporus: Verrill, 1915 : 147-148; Macan, 1938 : 349 (listed). 
Luidiaster mixtus: H. L. Clark, 1941 : 29-30; A. H. Clark, 1954 : 375 (listed). 

TYPE LOCALITY. Barbados, 183 metres. 

RANGE. Gulf of Mexico, Florida Strait, south to Nicaragua and Venezuela, east to the 
Leeward Islands; 150-570 (7586) metres. (The Atlantis specimen from 1000 fathoms E of 
Cuba said by H. L. Clark (1941) to show some anomalous features, is referable to C. sepitus, 
having convex spaced abactinal plates with relatively few spinules, stout conical marginal 
spines, single conical subambulacral spines and relatively few, c. 6, furrow spines in obtusely 
angled series.) 

Cheiraster (Christopheraster) mirabilis (Perrier) 
Fig. 2a, b 

Archaster mirabilis Perrier, 1881 : 27 (paragraphs 1-4 only); 1884:256-258, pi. 9, fig. 4 (non pp. 

258-259, sts 231 & 143, pi. 10, fig. 3 = C. echinulatus)^. 10, fig. 2 (or = C. blakei). 
Archaster coronatus Perrier, 1884 : 262-263 (non st. 19 = C. blakei). 
Cheiraster coronatus: Perrier, 1894 : 271-275; Ludwig, 1910 : 455,456. 
Cheiraster mirabilis (pt) Verrill, 1915: 124-127 (non pi. 14, fig. 5 = C. blakei; non C. mirabilis: 

Downey, 1973, Walenkamp, 1979 = C.planus). 
Cheiraster enoplus Verrill, 1915 : 135-137, pi. 18, fig. 1 (non C. enoplus: Downey, 1973 nee Luidiaster 

enoplus: H. L. Clark, 1941 = C blakei) 

TYPE LOCALITY. St. Kitts, Leeward Islands, 380 metres. 

RANGE. SE Florida to the Windward Islands (records from the Gulf of Mexico probably 
mistakes for other species); 380 (7265) 1470 (usually 400-700) metres; deeper records, e.g. 


Blake st. 196, \030fathoms, may be for C. sepitus. Uncertain identity precludes inclusion of 
lesser Blake station depths. 

Cheimster (Christopheraster) blakei sp. nov. 
Figs Ib, 2c-f 

Archaster coronatus (pt) Perrier, 1884 : 262 [st. 19 only]. 

lArchaster mirabilis (pt) Perrier, 1 884, pi. 10, fig. 2 (non A. mirabilis Perrier, 1 88 1 ). 

Cheiraster mirabilis (pt) Verrill, 1915, pi. 14, figs 5, 5a. 

Cheiraster echinulatus(pt)Verri\l 1915 : 131-133, pi. 14, fig. 3, pi. 19, fig. 2 [?pl. 25, fig. 1]. 

Cheiraster coronatus: H. L. Clark, 1941 : 26-27; A. H. Clark, 1954 : 375 (listed) (non C. coronatus: 

Perrier, 1894, Ludwig, 1910 = C. mirabilis). 

Luidiaster enoplus: H. L. Clark, 1 94 1 : 28-29 (non C. enoplus Verrill, 1 9 1 5 = C. mirabilis). 
Cheiraster enoplus: Downey, 1973 : 42-43, pi. 14, figs A, B. 

DESCRIPTION OF HOLOTYPE. R of the only entire original arm is 1 1 8 mm (two arms with 
regenerating tips are 95 mm or less); r is 12 mm; R/r = 9-8/1. Arm breadth at the fifth supero- 
marginals is 9'2 mm and the paxillar breadth here 6*0 mm, or 65%. The arms taper hardly at 
all for the first third of their length. 

There are about 35 spines more than 2 mm long on the disc based on enlarged abactinal 
plates including the primary radial and interradial plates and others. An additional 10-12 
smaller spines/spinelets arise in the papular area on the base of each ray, mostly on the row 
of plates between the two main irregular lines of pores each side of the midradial line. The 
longest central spines are 3 '6 mm long. Only a few other abactinal plates have an enlarged 
central spinelet among the small slender tapering spinules, length : basal breadth c. 2-5 : 1. 
These number usually 6-14, according to the plate size, mean c. 10, but the enlarged disc 
plates have up to 20 spinules around the central spine. All the abactinal plates are markedly 
convex and may be called parapaxilliform; they are rounded or slightly polygonal and only 
butt loosely together over most of the disc, while those on the arms are spaced from each 
other. The papulae number c. 75 on each ray; only one or two proximal ones (if any) are 
midradial, the rest being arranged in two distally diverging bands extending from alongside 
the primary radial plate to level with the distal ends of the third superomarginal plates. 

There are 55 superomarginals on the intact original arm, framing the paxillar area 
narrowly, each armed with a single spine of moderate size, the longest on the fifth and sixth 
plates up to 4-0 mm long, basal breadth c. 0*7 mm. The corresponding inferomarginal spines 
are similar or slightly longer, with an accessory spine about two-thirds as long below it, often 
also a third still smaller spine on the proximal plates besides several slender spinelets on the 
ventral face of the plate among the fine spinules which are of various sizes; the whole 
armament gives a hirsute or shaggy appearance to the plate as viewed from below. 

The actinal areas are small and triangular with four or five plates along the midinterradial 
line on each side of which the irregular adradial series numbers only about seven plates, 
those bearing the pedicellariae enlarged. The actinal armament is very sparse and mixed in 
size with only about two spinelets/spinules on most plates. 

Only 13^-15 adambulacral plates correspond to the first 10 inferomarginals. On each 
adambulacral no more than eight spines project over the furrow, their bases usually forming 
an angle of c. 90 but this may be blunted. The subambulacral spines are single, much larger 
and accompanied by only a few small spinules on the ventral face of the plate. The oral 
plates have 10 or 11 furrow spines, the apical one much enlarged and the second one 
intermediate. Probably only one suboral spine is enlarged but the plates are badly rubbed 
and most spines lost. 

Pectinate pedicellariae are developed on the adradial actinal plates (usually four in each 
interradius), on the ventral sides of the inferomarginals adjacent to the adambulacrals (for 
about three-quarters of the arm length) and abactinally (but only two are present near the 
centre of the disc). 

VARIATIONS. Some indications of variations and growth changes can be derived from Table 









C u 


U C 


03 u_ 

<u u 




5 R 
H <*> 



O 0) 


^ C/3 



5 6 
c )C 






O l*> 



o 2 



O O 

oo o .oooooor^t^ 


<N D. 1 ^- Tf 00 ^ OS 


.onaooo. ,c;v3tAoo 


















O o 
"v CX 


O <u 


X-. co 



<x> r ,\ 


o O 

>/^ >o o 


^ o 

t M r*^ I 




116 A.M.CLARK 

5. The disc is usually relatively larger than in the holotype and the arms more often taper 
evenly from the base, with the superomarginals forming a somewhat wider frame to the 
paxillar area, though never making up as much as 50% of the arm breadth. Often many 
abactinal paxillae have an enlarged central spinelet, graduating in size to the small spines on 
the papular areas. The relative number of papulae also varies; one specimen has well over 
100/ray even though R does not exceed 100 mm. In small specimens, the first one or two 
pores appear at R c. 16 mm (at the same time as the genital pores which are adjacent to the 
first superomarginal plates). 20 specimens with R 14-19 mm show a range in pore/ray 
number of 0-1 1 but with a mean of only 3. The marginal spines may be relatively larger than 
in the holotype, up to c. 6 mm long at R > 100 mm. In small specimens the fourth 
superomarginal spine is the largest but the fifth and sixth ones grow allometrically to equal 
and then exceed it. The number of adambulacral plates corresponding to the first ten 
inferomarginals is usually more than in the holotype, often 16 or up to 18 being noted. The 
bases of the furrow spines usually form a more pronounced angle projecting over the furrow. 
Finally, the occurrence of pedicellariae varies; some specimens may have more numerous 
abactinal ones; also at least one was observed with inter-superomarginal pedicellariae inter- 

AFFINITIES. Cheiraster (Christopher aster) blakei is closely related to C. (C.) mirabilis. Most 
of the differences are expressed in the tabular key (Table 3), the numerical tables here 
(Tables 4 and 5) and the comparison on pp. 104-5. These can be summed up as differences 
in armament, C. mirabilis having the enlarged abactinal and marginal spines relatively larger 
but fewer than C. blakei and the abactinal and inferomarginal spinules short and more 
nearly uniform. Additionally, the furrow spines tend to be more numerous in C. mirabilis, 
their arrangement on the furrow margin less angular and the contours of the abactinal plates 

TYPE LOCALITY. Gerda st. 918, 2603'N, 7805'W, between Florida and the Bahamas, 
804-822 metres. To be deposited in the U.S. National Museum. 

RANGE. Recorded from SE of Florida and the coasts of Cuba east and south to the Lesser 
Antilles and northern Brazil at least as far east as 44 W; 250 (?225)-1033 (but mostly 
500-800) metres. 

Cheiraster (Cheiraster) planus Verrill 
Fig. 5a, b 

Cheiraster planus Verrill, 1915 : 133-1 35, pi. 18, fig. 2. 

Pectinaster gracilis Verrill, 1915 : 145-147, pi. 6, fig. 1, pi. 14, fig. 4, pi. 15, fig. 1; Macan, 1938 : 349 

(listed); Jangoux, 1978 : 95-96, fig. 2 A, B. (IP. gracilis: Walenkamp, 1979 : 26-30, figs 9, 10, pi. 5, 

figs 1-4, pi. 6, figs 1-4). 
Cheiraster mirabilis: Downey, 1973 : 41-42, pi. 13, figs A, B; Walenkamp, 1979 : 23-26, figs 8, 10, pi. 

4, figs 2-4 (non C. mirabilis (Perrier, 1 88 1 )). 

TYPE LOCALITY. Unknown to Verrill but the holotype (USNM 18469) may have been from 
Albatross st. 2396 (Gulf of Mexico between the Mississippi delta and Cedar Keys) from 
which there is a sample (USNM 10564) including another specimen of C. planus (named 
mirabilis by Verrill but probably at an earlier date) and a note saying '17 specimens lent to 
A. E. Verrill'. 

RANGE. Western Gulf of Mexico and SW Caribbean, eastwards south of Jamaica to the 
Lesser Antilles and along the north coast of South America to French Guyana; 550 
(?408)-715 (7824) metres. A record from SW of Florida, Gerda st. 439, c. 24 N, 82+ W, 
needs confirmation since even the larger of the two specimens has R only 16 mm; the 
superomarginal spines appear to be larger than the inferomarginal ones, the abactinal plates 
are convex, the inferomarginals have fairly uniform spinules and the furrow angles of the 
adambulacrals are relatively shallow, as in C. planus as opposed to C. echinulatus. However, 
most of the distinctive subambulacral spines are lost. 



Fig. 5 (a, b) Pore areas of Cheiraster (Cheiraster) planus Verrill; (a) holotype, Albatross (st. ?), 
USNM 18469, R 92 mm; (b) Albatross st. 2396, USNM 10564, R c. 60 mm (same sample as 
holotype of Pectinaster gracilis Verrill); (c) Cheiraster (Cheiraster) sepitus (Verrill), ventral 
interradius partly denuded, same specimen as in fig. 4(k), the sockets of the missing 
subambulacral and suboral spines dotted; (d-g) Cheiraster (Luidiaster) robustus (A. H. Clark), 
holotype, Albatross st. 2789 (Chile), USNM 36945; (d) papular area and adjacent second and 
third superomarginals; (e) armament of four proximal abactinal plates; (f) ventral interradius, 
armament partly reconstructed, mostly lost; (g) fourth inferomarginal and adjacent adam- 
bulacral plates. 

Cheiraster (Cheiraster) sepitus (Verrill) 
Figs 4i-r, 5C 

Archaster sepitus Verrill, 1 885a : 151. 

Pontaster venustus Sladen, 1889:52-55, pi. 8, figs 5, 6, pi. 12, figs 5, 6; Perrier, 1894:287-288; 

1896:47;Koehler, 1909: 14-15, pi. l,fig. 13; Mortensen, 1927:71. 
Pontaster venustus var. robusta Sladen, 1 889 : 55. 
Pontaster sepitus: Verrill, 1894:247; 1895 : 137; 1915 : 1 30 (footnote). 
Pontaster perplexus Perrier, 1894 : 288-290, pi. 21, fig. 1, pi. 22, fig. 1; Mortensen, 1927 : 72. 
Pectinaster echinulatus (pt) Ludwig, 1910: 449 (sepitus a synonym). 
Pectinaster dispar Verrill, 1915 : 148, pi. 14, fig. 1; Macan, 1938 : 349 (listed). 
Pectinaster venustus: Koehler, 1924 : 186. 
Luidiaster dubius H. L. Clark, 1941 : 30, pi. 2, fig. 2. 
Luidiaster mixtus (pt) H. L. Clark, 1941 : 29-30 (specimen from 1000 fathoms). 

118 A.M.CLARK 

TYPE LOCALITY. S of Cape Sable, Nova Scotia; 1 570 metres. 

RANGE. South of Nova Scotia south to Cuba and the Lesser Antilles, east to the Azores, Bay 
of Biscay [? no details, Koehler, 1924], Portugal and south to the Cape Verde area; 485-3703 
metres (mostly 1000-2000 metres). 


Pectinaster Perrier, 1885:70; 1894:278-279; Ludwig, 1910:448-451 (pt); Fisher, 1911 : 122-123; 

1919 : 182; Macan, 1938 : 348-349 (pt). (non Pectinaster. Verrill, 1915, Downey, 1973). 
Pontaster (pt) Sladen, 1889 : 43-52. 

TYPE SPECIES. Pectinaster filholi Perrier, 1885, by subsequent designation by Ludwig, 1910. 
No type species was designated by Perrier in 1885 when he included two species, Archaster 
insignis Perrier, 1 884 and P. filholi which was evidently new (though not so stated), the genus 
being diagnosed only by the presence of fasciculate pedicellariae (though these are relatively 
simple in insignis having up to only four spinose 'valves'). In 1894 Perrier referred P. 
insignis to Dytaster Sladen, 1885, leaving P. filholi by elimination as the only eligible type 
species for Pectinaster, the formal designation awaiting Ludwig (1910: 449). 

DIAGNOSIS. A genus of Benthopectinidae with no odd interradial marginal plates (though the 
paired first marginals may be a little asymmetric in some interradii of some individuals), the 
interradial marginal spines relatively small; abactinal plates small paxilliform, at least the 
proximal ones elevated medially into a more or less well defined column bearing several 
spinules, on most plates of the disc and midradially on the arms surrounding a larger central 
spinelet, the spinules tending to form a continuous surface proximally in well-preserved (and 
probably in live) specimens; papulae restricted to an elongate oval papularium on to each 
arm base, formed of specialized markedly deepened abactinal plates, convex at the upper 
end and abutting together laterally with flanges to accommodate the papulae (see Fig. la), 
the surface of the papularium tending to bulge upwards (at least in preserved specimens 
when the dorsal body wall is contracted on to the ambulacra), the first-formed pores often 
forming transversely-aligned pairs, three or more pores developed by a size of R 20 mm; 
superomarginal plates mainly aligned laterally, only forming an inconspicuous rim to the 
paxillar area in dorsal view, each armed with an acute conical spine at the upper end; 
adambulacral plates with a single large conical subambulacral spine, only the first plate 
sometimes with a second spine; fasciculate pedicellariae usually present, often numerous, 
occurring on some abactinal, marginal, actinal and even a few adambulacral plates, usually 
oval in outline and formed of three or more modified spinelets based on either one or 
(pectinate pedicellariae) on two, rarely three, plates, the valves surrounding a deep hollow or 
bevelled notch in the plate or plates. Dorsal muscle bands of arms attached proximally to the 
wings of one or two ambulacral plates laterally and the adjacent marginal plates. 

REMARKS. This diagnosis emphasizing the highly modified compact oval papularia, 
paxilliform abactinal plates and laterally-aligned superomarginal plates not forming a wide 
border to the paxillar area in dorsal view, restores Pectinaster to the restricted concept of 
Perrier in 1894. As discussed under Cheiraster, the inclusion in Pectinaster of Archaster 
echinulatus Perrier, 1875, Pontaster cribellum Alcock, 1893, A. sepitus Verrill, 1885 and 
Pontaster venustus Sladen, 1889 (the last two as synonyms) by Ludwig (1910), followed by 
Verrill's addition in 1915 of Pectinaster gracilis, disparand mixtus (all new nominal species) 
and Pontaster oligoporus and Cheir -aster vincenti, both of Perrier, 1 894, broadened the genus 
to include species with less specialized papular areas and dorsally broad superomarginal 
plates. All these I consider have closer affinity with the type species ofCheiraster, C. gazellae 
Studer, 1883. The type material of most of them is relatively small so that the papulae have 
not yet formed bilobed areas, though the enlarged median proximal pore indicates that they 
are potentially bilobed. 


The elimination of these leaves Pectinaster in the Atlantic represented only by P. filholi 
and P. pristinus (Sladen), though the status ofP.forcipatus (Sladen) as distinct from P. filholi 
has been debated. 

In 1910, Ludwig referred Pontasterforcipatus Sladen, 1889, from off the eastern U.S.A., to 
the synonymy of Pectinaster filholi, type locality off Cap Blanc (with a paratype from the 
Azores area). He was followed in this by H. L. Clark (1923) but in 1962 I challenged this 
synonymy on the grounds of apparent differences in the numbers of actinal plates and their 
armament in the type series of P. forcipatus and P. forcipatus var. echinata (the latter from 
the Crozet Is area of the Southern Ocean), compared with Perrier's description and figures of 
P. filholi supported by a specimen (R only 33 mm) in the British Museum collections from 
SW of Ireland. However, after seeing further NE Atlantic specimens collected by the 
Discovery, in 1976 (Clark & Courtman-Stock) I came to agree with Ludwig and synonymize 
both forcipatus and echinatus with P. filholi, which name I used for specimens from South 
Africa, following H. L. Clark. Not only is there considerable variation in the numbers and 
shapes of the actinal plates in specimens from various parts of the Atlantic but also the 
records of Sibuet (1975) from the Gulf of Guinea and off Angola provide a zoogeographical 
link between filholi from NW Africa and echinatus from the Cape and Southern Ocean. 
Perrier's illustration of P, filholi (1894, pi. 20, fig. 3b) shows oral plates of exaggerated size at 
the expense of the actinal area, which appears exceptionally small, only c. 35% of the oral 
plate length, measured interradially. In four Discovery specimens from the Cap Blanc area 
(st. 9 1 33/5) an actinal area of each measures 6 1 , 68, 74 and 95% of the oral plate length. This 
compares with three specimens from W of Ireland with values of 48-92% (the minimum 
predictably in the smallest specimen with R only 40 mm), ten from the Gulf of Guinea off 
Ghana with 60-90%, six from off the U.S.A. with 65-109% and six from off South Africa and 
the Crozet Is area with 76-115%. The specimens with the relatively largest actinal areas, 
interradially longer than the oral plates, are mostly large, R > 70 mm (see Table 6). 

Sladen distinguished the variety echinata from the type material of P. forcipatus by the 
stouter marginal spines and larger central paxillar spinelets, the occurrence of which he 
thought was more widespread on the lateral paxillae. In the two largest syntypes of echinata 
(R c. 80 mm) the fourth superomarginal spine is 1*25 and 1-15 mm in basal breadth, the 
latter measurement being equalled in a South African specimen with R 72 mm. In North 
Atlantic specimens these spines are up to 0'9-1'Omm a small difference but just 
appreciable with the naked eye. Similarly, the longest central paxillar spinelets on the disc 
measure 1 '0 mm or more in the southern specimens as opposed to usually O'S-0'9 mm in the 
northern ones, though one syntype ofP.forcipatus has spinelets up to ! 1 mm. The extent of 
the central spinelets on the lateral paxillae varies also in P. mimicus from the Indo-West 
Pacific and P. agassizi from the East Pacific and I do not think it is a character of specific 
weight. Nevertheless there may be sufficient justification in the relative spinelet size for 
treating the southern specimens as subspecifically distinct from P. filholi but this should 
await comparison with material from the southern part of West Africa. 

One regional variation which may be significant is the relative number of pores in the 
papularia. In the available material this reaches a maximum of 20 in the american specimens 
and 21 in the NE Atlantic ones. The figured syntype(s) of P. filholi (Perrier, 1894, pi. 20, figs 
3a, 3d) shows 7 pores (only 6 in the description). A syntype in the MCZ, Harvard, with R 
only c. 45 mm, also from the Cap Blanc area, has 10 pores in one area examined. However, a 
Discovery specimen from this vicinity has as many as 2 1 pores in one area and 1 8 in another 
at R 65-70 mm. An even higher number is shown by some specimens of a large sample taken 
off Ghana by the Pillsbury (c. 4 N, 2^ W), of which 22 specimens with r > 8 mm (R 
probably 40+ mm) have 1 1-27 pores, mean 19. This compares with 6-20 pores and a mean 
of only 10 in 16 american specimens. Unfortunately the Pillsbury material is dried and in 
poor condition so other characters such as spinelet length cannot be ascertained in many 
specimens. Even in spirit-preserved specimens of Pectinaster, owing to frequent breakage of 
the arm tips, few accurate estimates of R/r or counts of marginals can be made. The mean 
R/r value for 16 of the type series of P. forcipatus is 5'9/l, compared with 5*5/1 in six 

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specimens from the NE Atlantic but this may not be significant since the latter are mostly 

As for Pontaster pristinus Sladen, based on a single small specimen from Challenger st. 
325, off the River Plate, this is clearly a Pectinaster sensu stricto, having relatively narrow 
superomarginals in dorsal view and compact, slightly bulging, papularia. Reexamination of 
the holotype throws considerable doubt on Sladen's arguments about its distinction from P. 
forcipatus. The specimen was coated with fine mud. When this was removed, the abactinal 
plates of the disc were found to have clusters of up to 7 spinules, usually 4 or 5, around the 
fine spinelets, the tips of which were seen by Sladen, the armament being in fact indistin- 
guishable from that of specimens of P. filholi of similar size (e.g. from Challenger st. 46, R 
c. 18 mm). (Sladen somehow misjudged R in pristinus as only 11 -5 mm but the complete 
arm remaining, though curled over, is at least 5 mm longer than this. Only two pores could 
be found in the papularium cleared and the abactinal spinelets of the disc are up to 0'6 mm 
long. The condition of the arm plates is poor but does not appear to be significantly different 
from small specimens of P. filholi and the madreporite is no more conspicuous than can be 
attributed to adpression of the armament in its vicinity. There are 4 or 5 furrow spines on the 
proximal adambulacral plates with a few similar spines in series with them proximal and 
distal to the large conical subambulacral spine. Three North American specimens with R 
18-21 mm, have 3, 4 or 6 pores in one papularium but this is probably not significant in 
view of the variation in pore number shown by other specimens of filholi. Nor do I consider 
the absence of pedicellariae in the single specimen of P. pristinus justifies a specific 
distinction, though pedicellariae are more or less numerous in most specimens from the NW 
and NE Atlantic examined. 

Outside the Atlantic, Pectinaster includes P. agassizi (Ludwig, 1905) from the East Pacific 
and P. mimicus (Sladen, 1889) from the Indian Ocean, East Indies and New Zealand. 
Despite its having enlarged spinelets on the lateral as well as midradial abactinal arm plates 
and only 12-14, rather than 15-18, proximal adambulacral plates matching the first ten 
inferomarginals, P. hylacanthus Fisher, 1913, from the Philippines and Indonesia, was 
reduced to a subspecies of P. mimicus by Doderlein, 192 1 . If these characters are of less than 
specific weight, then P. hispidus (Woodmason & Alcock, 1891) from the Indian Ocean is 
also conspecific with P. mimicus. This synonymy is supported by comparison of a paratype 
of P. hispidus with the holotype of P. mimicus. Further, Macan (1938) identified several 
specimens from the Gulf of Aden as P. agassizi or P. agassizi granuliferus subsp. nov. rather 
than P. mimicus. The new subspecies was said to differ in having abactinal central spine(let)s 
on the arms restricted to the midradial areas and more granuliform spinelets on the actinal 
plates (besides differences in the distribution of the pedicellariae not significant in my 
view), these two characters being also used by Macan to differentiate P. mimicus from the 
specimens he called P. agassizi. The fact that his largest specimens from the Gulf of Aden 
have R 1 19 and 158 mm could well account for the very high number of 40-50 pores in the 
papularia in apparent agreement with P. agassizi compared with up to only 20 in the few 
specimens attributed to P. mimicus in which pores have been counted (the types of P. 
mimicus with R 78 mm and of P. mimicus palawanensis, R c. 95 mm), both Fisher and 
Doderlein having apparently given little weight to relative pore number as taxonomically 
significant in Pectinaster. East Pacific material of P. agassizi has even more numerous 
pores already c. 55 at R only 65 mm from c. 25 at R 45 mm, so that the number is likely to 
be even greater when (if) R exceeds 100 mm. 

Of the other nominal species included in Macan's table for Pectinaster, Pontaster 
cribellum Alcock, 1893, was compared by Alcock rather with P. subtuberculatus Sladen, 
since referred to Cheiraster. Alcock's holotype had R only 45 mm and this could account for 
the still rounded rather than bilobed form of the papular areas. Finally, Pectinaster robustus 
A. H. Clark, 1917, from off Chile, has two large subambulacral spines and reexamination of 
the large holotype showed the pore areas (Fig. 5d) to be deeply bilobed distally and not 
restricted to specialized papularia. P. robustus is therefore now referred to the subgenus 
Luidiaster of Cheiraster. This action brings robustus into conflict as a junior secondary 



Table 7 Data for the Pillsbury specimens of Benthopecten. For the abactinal spinules per proximal 
plate '!+' signifies an enlarged spinelet or spine on some plates; for disc spines (which tally includes 
enlarged spinelets more than twice as long as the ordinary spinules), C, R and I in capitals or lower case 
signify large or small central, primary radial and primary interradial spines, the rest being irregular in 
position; for pore extent, the number is the superomarginal plate opposite the distalmost pores; the 
adambulacral count is of the proximal ones opposite the first 10 inferomarginals; brackets around 
entries signify occasional occurrence. 


Abactinal spinules 
proximal distal 

Disc spines 

Pore Adambs: 10 Furrow 
extent i.m. pis spines 

68 1 , Guyana 



lC,4R + c. 50 

4 19 
3 19* 



782, Colombia 



lc,2r + 3 








120, Florida/ 



lc,2r,2i + 2 
2R + 3 

4 16(15) 
2 21 
1 19 




3 14, Gulf of 




lC,2r + 8 

5 21 
4 21* 


34, Gulf of 

1 (2,3) 
1 (2,3) 



2-3 1\\ 



681, Guyana 




3(7 + 2)1 + 3 

3 16* 

5,6 (7) 

homonym with the species-group name robusta Sladen, 1889, originally described as a 
variety of Pontaster venustus but now referred to the synonymy of Cheiraster (Cheiraster) 
sepitus (Verrill, 1885). However, robustus (A. H. Clark) may well prove to be conspecific 
with C. (L.) planeta (Sladen, 1889), type locality off southern Chile, when specimens of 
intermediate size are available for comparison. 

Turning to benthopectinids with odd interradial marginal plates, Benthopecten itself is well 
represented in the Atlantic and also in need of review. 


Benthopecten Verrill, 1884:218 (footnote); 1895:130; Ludwig, 1910:464^66; Fisher, 

191 1 : 142-144; Macan, 1938 : 351-352; Downey, 1973 : 40. 
Pararchaster Sladen, 1885:610-611; 1889:4-7; Ludwig, 1910:462^64. (Type species, by 

monotypy, P. pedicifer Sladen, 1 885). 

TYPE SPECIES. Benthopecten spinosus Verrill, 1884, by monotypy. 

DIAGNOSIS. A genus of Benthopectinidae with an odd interradial marginal plate in each 
series, usually in all five interradii, the superomarginal one more or less inset on to the disc 
and bearing a conspicuous spike-like vertical spine (or occasionally a cluster of c. 3 small 
spines); abactinal plates thin, flat and scale-like, at least on the arms, but sometimes convex 
medially on the disc, never markedly paxilliform, each bearing one or more spinules, some 
of which may be elongated into spinelets or even spines more than 1*5 mm long, especially 





Oral f. 

Suboral Pedicellariae 
spines Abact. Act. i.m. 



681, Guyana 




4 (+) + + 
4 (+) + + 

100 + (?c. 150) 
40 + (760-70) 


782, Colombia 



4 + + + 

20 ++(770- 100) 





2,3 + + 

7? v. poor 


120, Florida/ 



5,6 + (+) (+) 
4,5(6) (+,1) (+,1) + 

50 + (?c. 70) 


3 14, Gulf of 



3 + + + 
5 + + + 

75++(7c. 130) 
100 + (?c. 150) 


34, Gulf of 



3 + + + 
4 + + + 

45 + (770-80) 
30 + (?c. 70) 


681, Guyana 


45++(?c. 110) 15 

on the primary plates of the disc; papulae scattered on the arm bases, usually extending 
further distally close to the marginal plates, so the areas become bilobed, sometimes covering 
much of the disc and a limited distance (usually <^R) along the arms; marginal plates with at 
least one long spine, the inferomarginals usually with an accessory spine (or two) below the 
main one but smaller; adambulacral plates with the furrow margin rounded or angular, 
armed with 3-8, sometimes more, radiating, slender furrow spines backed by one or two 
(rarely three) larger subambulacral spines usually aligned almost at right angles to the furrow 
when more than one occurs but in some species nearly parallel to it; some pectinate 
pedicellariae usually present, especially on the actinal plates, but often also between some 
abactinal or inferomarginal plates, especially proximally. 

REMARKS. Yen-ill's extraordinary omission in 1884 (though rectified in 18856:519) to 
mention the most obvious characteristic of Benthopecten the odd interradial supero- 
marginal plates with their conspicuous single spines led Sladen (1885 & 1889) to refer his 
several species to a new nominal genus Pararchaster. In 1894 Verrill referred Pararchaster to 
the synonymy of Benthopecten. Although Ludwig (1910) tried to retain two genera on the 
basis of the multiple abactinal spinelets of Pararchaster pedicifer Sladen, the type-species of 
Pararchaster, as opposed to the single ones of Benthopecten spinosus, Fisher (1911) found 
this distinction untenable at the generic level, following observations on the more abundant 
North Pacific material available to him. Study of the material of Benthopecten collected in 
the deep tropical Atlantic by the Pillsbury raises new doubts about even the specific 
significance of the frequency of the abactinal armament in many cases. These new specimens 

124 A.M.CLARK 

yield the data included in Table 7, from which it is evident that the very small samples 
collected at stations as wide apart as the Gulf of Guinea off West Africa, the Guyana Basin 
off northern South America (outside the Antillean chain) and the North American Basin 
between Florida and Bermuda, show a wide range within each sample in abactinal spinule 
frequency. The maximum spinule number is found in one large specimen from st. 68 1 off 
Guyana and another from st. 314 in the Gulf of Guinea, each sample including a second 
specimen with much less dense armament but otherwise not significantly different. St. 120 
also includes one more spinulose specimen out of three, though the difference is less marked. 
However, a third specimen was collected at st. 68 1 ; this is the single individual, segregated in 
Table 7, having the minimum of abactinal spinelets rarely more than one, which may be 
specifically distinct. Such a conclusion is supported by its primary interradial disc spines 
appearing to be better developed than the primary radial ones, coupled with there being 
rarely as many as 7 furrow spines (and that only on odd plates), where 6 or 7 is the usual 
number in the other specimens, and only a single (actinal) pedicellaria, where all the rest 
with R > 50 mm have several actinal and inferomarginal pedicellariae and usually also some 
abactinal ones. A review of the fairly abundant material of Benthopecten spinosus from the 
eastern continental slope of the U.S.A. has shown a consistent agreement in these same 
characters, except that the rare pedicellariae found were only on the inferomarginal and not 
the actinal plates. 

In comparison with other Atlantic species, a multispinulose condition with up to 10 
spinules on some abactinal disc plates is shown by the four syntypes of B. spinosissimus 
(Sladen) from off Ascension I (R 60-68 mm) and the holotype of B. fischeri (Perrier) from 
NW Africa (R c. 40 mm), the latter reexamined at the Paris Museum. 

In B. spinosissimus already at R < 70 mm the papulae extend distally to level with the 
sixth or seventh superomarginals and the marginals themselves are relatively short and 
numerous, numbering 47-53 in each series in the syntypes, giving a ratio of 1 -2-1 -4/1 for 
R/SM number. This compares with only about 30 marginals and a ratio of c. 2-2/1 at this 
size in B. spinosus and probably a similar number in B. simplex (Perrier), of which a 
specimen with an intact arm showing R 55 mm has only 25 superomarginals, also giving a 
ratio of 2 - 2/l . A further distinction of B. spinosissimus is the small number of furrow spines, 
only 3 or 4 on each plate, leaving little doubt of its specific distinction. 

With regard to B. fischeri, I found the holotype and only recorded specimen to have 4 or 5 
spinules on most proximal abactinal plates but additional ones on some plates with an 
enlarged spinelet or spine, apparently an unusually high frequency at this relatively small 
size. Although the jaws have a relatively rich armament of c. 10 furrow spines and up to 12 
suboral spines and spinelets on each oral plate, there are only 6 furrow spines on most 
adambulacral plates and the other characters agree with the bulk of the Pillsbury material. 
(The exception is the single Pillsbury specimen allied with B. spinosus, which is quite 
distinct in its very sparse armament.) B. fischeri was referred to the synonymy of B. spinosus 
by Grieg (1921 & 1 932) on its superficial resemblance. 

Grieg also followed Verrill (1895) in synonymizing Pararchaster armatus Sladen, 1889 
with B. spinosus, though Farran (1913) and Mortensen (1927) maintained armatus as a 
distinct species. Ludwig (1910) had instead referred armatus to the synonymy of Archaster 
simplex Perrier, 1881, the oldest name for any species of Benthopecten. The holotype (from 
the Gulf of Mexico NW of Cuba) has R only 18 mm and, until the Oregon and Alaminos 
took some rather larger specimens (R 48+ mm) in the Gulf of Mexico (described by Downey 
in 1973), little comparative information was available about B. simplex. Reexamination of 
these specimens and comparison with the type material of B. armatus from S and E of Cape 
Cod and various specimens from the vicinity of Portugal, SW Ireland and the Rockall 
Trough in the NE Atlantic reveals no differences which I consider to be significant (see Table 
10). The abactinal armament is multiple in all except immature specimens (numbering 
usually 1-5 spinules on the proximal plates), the disc spines (when developed) tend to 
enlarge first on the primary radial more than the interradial plates, the number of furrow 
spines averages 6 or 7 on most of the adambulacral plate series and the number of sub- 


ambulacral spines tends to increase from one to two with size, with occasional exceptions; 
lastly there is a relatively high incidence of pedicellariae, often in all three loci, in all but the 
smaller specimens (R <40 mm), again with some individual variation. 

The origin of some of the Pillsbury specimens from areas geographically adjacent to the 
Gulf of Mexico (the nearest being from st. 1178 east of Cuba) and their morphological 
resemblance to the rather smaller and mostly less well preserved Oregon and Alaminos 
specimens reinforces belief that all can be referred to B. simplex but for the exceptional 
specimen from st. 681 which is indistinguishable from B. spinosus. This serves both to 
extend the known size range of B, simplex to R c. 1 50 mm and also to extend the 
geographical range (not unexpectedly in view of the great depths) not only to both sides of 
the North Atlantic but also both sides of the tropical Atlantic, including the Gulf of Guinea, 
taking in the type locality of Benthopecten chardyi Sibuet, 1975. Despite several peculiarities 
of Sibuet's unique holotype (discussed further below), it seems to fall within the range of 
variation of B. simplex. 

CHARACTERS. Apart from the density of the abactinal armament (now considered to be of 
only limited use) the following characters have been adopted as of some taxonomic signifi- 
cance in distinguishing the species of Benthopecten: 

1 . The development of enlarged spines on the centre of the disc (by Ludwig) 

2. The relative size of any accessory spines on the inferomarginal plates (Sladen, Ludwig and 

3. The number of paired marginal plates in each series (Sladen) 

4. The numbers of furrow and subambulacral spines (Sladen et al.) 

5. The number of oral furrow spines (Ludwig) 

6. The number of adambulacral plates corresponding to the first ten paired inferomarginal 
plates (Ludwig and Macan, 1938) 

7. The occurrence of pedicellariae (Sladen et al.) 

8. The presence of more than one superomarginal spine (Sladen and Macan) 

9. The substitution of several small spines for the usual conspicuous odd interradial 
superomarginal spine (Perrier, 1894, forB.folini) 

10. The extent of papulae along the arm (Fisher and Macan) 

1 1 . The extent of the inferomarginal pedicellariae along the arm (Fisher and Macan) 

12. The number of actinal pedicellariae (Macan) 

Other characters which can be considered as possibly significant for some of the species 
but needing further comparisons for appreciation of their wider significance within the 
whole genus are: 

13. The occasional non-bifurcation of the papular areas distally 

14. The relative size of the primary radial and interradial disc spines, when developed 

15. The number of suboral spines 

16. The presence of more than one series of actinal plates and 

17. The relative size of the first paired superomarginal spines. 

Fortunately, since the arms are rarely preserved intact, the R/r ratio does not appear to be 
taxonomically significant in this genus, the relative disc size being apparently variable over a 
similar range in all the species. As in many long-armed asteroids, the R/r ratio tends to 
increase allometrically with absolute size. 

As with the characters based on the abundance of abactinal spinules, most of the others are 
subject to some degree of variation. 

No. 1 , the development of enlarged spines on the disc, depends to some extent on size and 
is also subject to variation. For instance, in large specimens of B. spinosus, R >90 mm, the 
disc spines range from few and inconspicuous, < 1'5 mm long, to a cluster of 20 or more, 
some exceeding 6 mm in length. An elaboration of this is character no. 14, the relative size of 
the primary radial and interradial disc spines, when developed, studied now only in B. 
spinosus and B. simplex, where a range of specimens is available. As detailed under the 
heading of B. simplex, on average the primary radial spines in that species are equal to or 

126 A.M.CLARK 

slightly larger than the primary interradial ones, whereas in B. spinosus the interradial spines 
are mostly better developed (not counting those of the madreporic interradius where the 
spine development is generally erratic). This character remains to be evaluated in other 

No. 2, the mean size of the accessory inferomarginal spine (if developed) relative to the 
main spine above it, appears to be somewhat variable and is of uncertain value because of 
paucity of information about its range, only holotypes having often been described. In some 
species such as B. spinosus it seems to be fairly consistently at least half as long as the main 
spine, whereas in B. simplex the reverse is true and it rarely exceeds half the spine length. 
(An exception again is the holotype of B. chardyi where the accessory spine is often large and 
may even equal the main spine on some plates.) Very spinose species such as B. spino- 
sissimus have several successive accessory spines on each plate, graduating in size. 

No. 3, the number of marginals relative to R, remains to be investigated for most species, 
marginal counts having rarely been cited by authors, partly because so few specimens are 
usually obtained with any arms intact. The development of additional marginal plates 
adjacent to the terminal does not keep pace with the elongation of the arms so that the ratio 
of R/SM number tends to increase with R. In 12 specimens of B. spinosus with R 
35-155 mm, the ratio ranges from 1-8/1 (in the smallest) to 3-4/1 (in the largest), whereas 
nine specimens with R 70-110 mm have a range of 2-3-3-0/1 and a mean of 2-6/1. The 
holotype of the closely related B. semisquamatus (Sladen), from Japan, does not differ 
significantly, having a ratio of 3-0/1 at R 166 mm. In contrast, as mentioned above, the four 
syntypes of B. spinosissimus with R 60-68 mm have a ratio of only 1-2-1-4/1, the marginals 
being relatively short and more numerous, 47-53, as opposed to only 32 in a specimen of B. 
spinosus with R 73 mm. 

No. 4, the numbers of furrow and subambulacral spines, may show a wide range along the 
furrow length of a single specimen, usually tending to increase beyond the proximal plates 
and to a greater extent in very large specimens. For instance, in the holotype of B. 
heteracanthus Macan from the Indian Ocean, R 210 mm, the number increases from 7 or 8 
proximally to 1 1 on many distal plates, while the holotype of B. polyctenius Fisher from 
Indonesia, at R 224 mm, has a range from 9-14. Two smaller specimens of B. polyctenius 
with R 120 and 80 mm have lower maxima of 1 1 and 9 respectively. In species of more 
restricted size range, the range in furrow spine number is probably restricted to c. 3 or 4. For 
instance, in B. spinosus, known to reach R 1 55 mm, the range is 4-6, predominantly 5, even 
in larger specimens, R >90 mm, while B. simplex with a similar size range usually has 6 or 
7, occasionally 5 or 8 spines. The holotype of B. chardyi Sibuet (now thought to be less than 
specifically distinct from B. simplex) has 5 or even 4 spines on about the first ten 
adambulacral plates but the number increases first to 6 then 7 beyond about the twentieth 
plate. As for the subambulacral spine number, up to a size of R 30 mm B. simplex has only 
single spines on nearly all the plates, then in most specimens a second spine becomes 
enlarged beyond the most proximal plates so that specimens with R >60 mm usually have 
two subambulacrals predominating throughout the series. (The holotype of B. chardyi with 
single spines on most plates could be interpreted as neotenous in this respect; two slightly 
smaller specimens of B. simplex also from the Gulf of Guinea have a small second spine on 
some plates.) 

No. 5, the number of oral furrow spines, would be expected to vary through the same sort 
of range as the mean number of adambulacral furrow spines. Certainly B. polyctenius shows 
a range of 8-14 according to Fisher (1919). B. spinosus has 5 or 6, occasionally 4, and B. 
simplex shows a range from 5 to 8, compared with 10 (or even 1 1) oral furrow spines in the 
holotype and only mature specimen of B. fischeri (Perrier) at R only c. 40 mm, which in 
other respects resembles the more spinulose specimens of B. simplex and has only c. 6 furrow 
spines on the adambulacral plates. Until more material from the type locality of B. fischeri is 
available, the significance of this character remains to be determined. 

No. 6, the number of adambulacral plates relative to the first ten paired inferomarginals, 
appears to vary over a range of 4 or 5, sometimes more, in a single species (see Tables 9, 10), 


judging from the data available. Unfortunately, even Ludwig, who initiated its use, failed to 
cite it for several species and not all subsequent authors have included counts in their 
descriptions. The usual number in B. spinosus is 1 7-20, while in B. simplex between 1 8 and 
21 is common but as many as 24 (or even 25) are found, for instance in the anomalous 
holotype of B. chardyi. Also in the holotype of B. violaceus (Alcock) from the Indian Ocean 
24 are supposedly found but this number is derived only from the drawing, not Alcock's 
description, and may not be correct. 

Nos 7, 1 1 and 12. The occurrence and frequency of pedicellariae are notoriously variable 
and rarely reliable in diagnosing most asteroid species; this is equally true of Benthopecten. 
Only a large sample (rarely provided) can give a good idea of the relative frequencies of 
abactinal, actinal and inferomarginal pedicellariae. Even in B. spinosus, in which 
pedicellariae have long been thought to be consistently rare or absent, an additional sample 
of ten specimens from the Atlantis II collections showed four with at least some infero- 
marginal pedicellariae. 

No. 8, the presence of accessory superomarginal spines in species such as B. spinosissimus 
is correlated with a generally higher frequency of spines on all the plates and is not in itself 

No. 9, the absence of conspicuous spines from the odd interradial superomarginal plates, 
as described in B.folini (Perrier, 1894), from off NW Africa, is not shown by all specimens of 
the type series so one must conclude that either two sympatric species were included or that 
the character is variable. If there are two species in the type series, the one with multiple 
diminutive spines on the odd plates also usually has two subambulacral spines. The other, 
with single large spines may have one subambulacral spine and is then indistinguishable 
from B. simplex chardyi. It is not certain which specimen in the Paris Museum collections 
Perrier used for his full description; the arms were broken and he could only cite r, which he 
gave as 12 mm. The largest specimen from off Cap Blanc (Talisman 1883 haul 96, P.M. no. 
3304) is almost certainly the one shown in Perrier's pi. 19, figs la and Ib of a general view, 
above and below; I estimate its r as lO'Q-10'5 mm (some shrinkage during nearly 100 years 
in spirit is not unlikely); R is 65 + 710-20 mm; all the odd superomarginals have small spines 
and there are two subequal subambulacral spines on most plates. Of the two specimens from 
haul 98, slightly further south (P.M. nos 33 1 7 and 33 1 8) one has had the abactinal body wall 
removed and is the specimen so mutilated shown in Perrier's pi. 20, fig. Ic. The second has 
R50 + 720-30 mm, r c. 1 1 mm and so could be the specimen from which pi. 20, figs la and 
Ib of details of parts of the disc were made, though there are some divergencies between the 
finer details of armament and Perrier's description. Accordingly, the largest specimen from 
off Cap Blanc is now designated as lectotype; its subambulacral armament was probably 
misrepresented as single in pi. 19, fig. la by Perrier's artist. However, of the four (not three) 
smaller and more broken specimens from this same haul 96, one has relatively large single 
odd superomarginal spines up to 6 mm long, R being 20 + ?c. 30 mm; its subambulacral 
spines are mostly single. The other three are more damaged interradially, the spines of the 
odd plates mostly lost; one has a pair of large spines still attached to one plate and signs of 
paired spines in at least two other interradii. From Talisman haul 97 (no. 3305) come three 
more damaged paratypes, at least one with three massive single odd superomarginal spines. 

Possibly these large spines are vulnerable to damage or loss from the attacks of predators 
and the anomalies in development are due to differential regeneration. In the holotype of B. 
pikei H. E. S. Clark, 1969, from New Zealand, in four out of five interradii the spines are 
relatively small and in 1970 she suggested that this might be attributable to injury when she 
described a variety australis ofB. pikei from two specimens with all the odd spines large. 

No. 10, the distal extent of the papulae along the arms, progresses during growth, usually 
from level with the first superomarginal to the third or fourth, while the distalmost pores are 
usually lateral rather than midradial (character no. 13). However, in B. acanthonotus Fisher, 
from off California, the distal extremity varies in three large specimens from level with the 
fifth superomarginal to the twentieth. In this species the papular areas are consistently 
bifurcating but the holotype and only described specimen of B. indicus Koehler, from off Sri 








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130 A.M.CLARK 

Lanka (R c. 120 mm) has the papulae extending to superomarginal 12 for the whole width of 
the arm. In a Pillsbury specimen from NE of Florida (R 100 mm), now referred to B. simplex, 
the papulae extend to the fourth superomarginals and the areas do not bifurcate. Also in the 
holotype of B. chardyi the distalmost pores are adradial rather than abradial and extend only 
to the third superomarginal though R is 90-100 mm. However, two slightly smaller Pillsbury 
specimens also from the Gulf of Guinea have the areas bifurcating; their papulae extend 
distally to the third (or even the distal ends of the second) superomarginals. Isolated cases of 
non-bifurcating areas have been recorded from at least one non-Atlantic species and proper 
scrutiny of more material may well show occasional instances of this, suggesting that 
character 1 3 has little taxonomic significance. Since in some benthopectinids the dorsal arm 
muscles are attached at least partially to the dorsal body wall near the base of the arm, as well 
as or rather than to the ambulacrals and marginals, this might have a limiting effect on the 
extent of the pores medially. 

No. 15, the number of suboral spines, is usually 3 or 4 (sometimes 5) on each plate in most 
species, accompanying spinelets being rare. Occasionally higher numbers are found. 
Reexamination of the holotype ofB.fischeri from NW Africa showed a total of c. 12 suboral 
spines and spinelets on each plate, four of them distinctly larger than the furrow spines. In 
contrast, the four similarly multispinose syntypes of B. spinosissimus from near Ascension 
Island have only 4 suboral spines on most plates, though there may be one or two additional 
spinelets on some plates. The oral furrow spines on the type ofB.fischeri are also unusually 
numerous, as discussed under character 5. 

No. 16, the number of series of actinal plates, is correlated with the relative disc size, as 
well as tending to increase with growth. Until an R of c. 100 mm, rarely more than one series 
is well developed. More larger specimens are needed to evaluate this character. 

No. 17, the relative size of the first paired superomarginal spines, shows a considerable 
range from very reduced in comparison with the fourth (usually the largest) spine to almost 
as large as the odd interradial spine. It can result in very different appearances of the 
interradii but preliminary observations suggest that it is too variable (even within one 
specimen) to be significant. 

The characters thought to be most useful are utilized in two tabular keys (Tables 8 and 9) 
to distinguish the Atlantic and non-Atlantic species of Benthopecten. Analysis of these tables 
shows only very minor differences in variable or largely untested characters between several 
of the nominal species or differences possibly attributable to growth changes; notably B. 
spinosus Verrill and B. semisquamatus (Sladen) are separated only by an insignificant 
difference in the number of adambulacrals relative to the inferomarginals. The occasional 
presence of inferomarginal pedicellariae in B. spinosus may well prove to be shared by some 
specimens of B. semisquamatus other than the syntypes, when more material is available for 
comparison. This close affinity was observed by Sladen, who treated an east American 
specimen, now regarded as conspecific with B. spinosus, as only a variety occidentalis of B. 
semisquamatus. Inter-oceanic distributions of abyssal species such as these are very likely. If 
the present decision to treat Atlantic specimens showing a considerable range of variation in 
density of the abactinal armament as a single species, B. simplex, is proved correct, then 
barriers to the conspecificity of many nominal species in the middle and lower parts of Table 
9 will be almost completely broken down and there should be a drastic reduction in the 
number of valid species of Benthopecten recognized. 

Since Benthopecten simplex is the first-named species of the genus and its limits both 
morphological and geographical are considerably extended by the Pillsbury and other 
specimens now considered to be referable to it, a full account of this species is given here. 

Benthopecten simplex (Perrier) 

Archaster simplex Perrier, 1881 : 28; 1884 : 264, pi. 1, fig. 8. 
Pararchaster armatus Sladen, 1889 : 19-22, pi. 1 , figs 5, 6, pi. 4, figs 5, 6. 
Pararchaster simplex: Perrier, 1894 : 253, 254-256. 


"Pararchasternov. sp. (P. Fischeri Perrieraff.)' Mortensen in Schmidt, 1904 : 24. 

Benthopecten simplex: Ludwig, 1910:461, 464, 465, 466; Fisher, 1911 : 143; Verrill, 1915: 1-22; 

Downey, 1 973 : 40, pi. 1 2, figs C, D. 

Benthopecten armatus: Farran, 1913 : 2-3; Mortensen, 1927 : 74-75, fig. 41. 
Benthopecten spinosus (pt) Grieg, 1921 (& 1932): 12-13, pi. 4, fig. 1 (Michael Sars sts 95 & 101 at 

least); H. L. Clark, 1941 : 26. (Non B. spinosus Verrill, 1884). 
Benthopecten chardyi Sibuet, 1975 : 283, 289-29 1 , fig. 3, pi. 1 , figs B, D 

R is probably up to c. 150 mm but rarely more than 80 mm; R/r is 7'7-7'9/l and R/no. of 
superomarginal plates c. 2*25/1 in three specimens with R 50-55 mm. 

DIAGNOSIS. A species of Benthopecten with multiple spinules on most proximal abactinal 
plates, often 2 or 3 but sometimes up to c. 10 when they are accompanied by a larger spinelet 
or spine, especially at R >c. 50 mm, those on the primary radial plates tending to be better 
developed than those of the primary interradial plates; papular areas usually distally 
bifurcate, the distalmost pores level with the second or third superomarginals in specimens 
with R 50-80 mm but extending to the fourth or even the fifth in those with R > 100 mm; 
odd interradial superomarginal plates with a single very large spine, other superomarginals 
also with single spines and a few small spinelets or spinules, inferomarginals with an 
accessory spine below the main one but usually less than half as long; 16-23, usually 18-21 
proximal adambulacral plates corresponding to the first ten paired inferomarginals (though 
up to 25 in the holotype of B. chardyi); actinal plates few, a second series only developed at R 
> 100 mm; adambulacral plates with 6 or 7 furrow spines (sometimes 5 or 8) and one or two 
subambulacral spines, usually two at R > 50 mm beyond the first ten or so plates, where 
there is more often only a single spine; oral plates with 5 or 6 (sometimes 4 or 7) furrow 
spines and 3 or 4 (5) suboral spines; actinal and some inferomarginal pedicellariae usually 
present at R > 50 mm and sometimes also some abactinal ones. 

VARIATIONS. As mentioned under the generic heading, the armament of the abactinal plates 
is very variable in B. simplex, if I am right to include most of the Pillsbury specimens. At R 
50-70 mm there are usually 2 or 3 (1-5) spinules on the proximal plates and single ones on 
the plates distal to the pore areas but occasional specimens of this size and larger ones may 
have some proximal plates enlarged with a bigger spinelet and up to 10 spinules. 

The degree of enlargement of spinelets and spines is also variable. Even at R c. 70 mm 
there may be only a single central spine appreciably larger than the spinules, though usually 
several others may exceed 1'5 mm in length at this body size, especially on some of the 
primary plates. 

The papular areas normally bifurcate distally but the largest specimen from Pillsbury st. 
120, NE of Florida, has the distalmost pores adradial. 

The odd interradial superomarginal spines are usually very conspicuous but on occasional 
specimens one or more of them may be reduced in size. 

The length of the accessory inferomarginal spine is usually less than half that of the main 
spine but it may be lacking altogether on most plates of a few individuals or conversely be 
nearly equal in length to the main spine on some plates, e.g. of the paratype of Pararchaster 
armatus with R 42 mm from Challenger st. 46 but not in the larger paratype from the same 

In larger specimens the single subambulacral spines of the proximal adambulacral plates 
often give way to two spines beyond, as in Downey's specimens from the Gulf of Mexico, 
shown by reexamination. 

Other variants are mentioned in the diagnosis. 

SYNONYMY. Until Downey (1973) described some Oregon and Alaminos specimens of B. 
simplex from the Gulf of Mexico, only the juvenile holotype (R 18 mm) had been described 
from the West Indian area and it was difficult to establish whether Sladen's type material of 
Pararchaster armatus from SE and E of Cape Cod (c. 42 N, 63 W and 40 N, 66 W) is 
synonymous with B. simplex, as proposed by Ludwig (19 10) or with B. spinosus Verrill, also 






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from the eastern U.S.A., as proposed by Verrill (1895). Comparison of the larger specimens 
of Downey and Sladen, together with a few further individuals from the NE Atlantic (off 
Portugal, the Bay of Biscay, SW Ireland and the Rockall Trough) yields the data shown in 
Table 10, which also includes for comparison the only available specimen of B. spinosus of 
comparable size. Table 7 of the Pillsbury specimens, all but one now referred to B. simplex, 
should also be consulted. 

Although the positions of the disc spines and enlarged spinelets are often irregular 
(especially adjoining the madreporic interradius), the contrast in the specimen of B. spinosus 
in Table 10 of four relatively large primary interradial spines with the three smaller primary 
radial spines (reflected also by most large specimens of B. spinosus) is very marked, as well as 
the absence of multiple abactinal spinules. Further, the presence of only 4 furrow spines on 
most adambulacral plates, as compared with the usual 6 or 7 of B. simplex, and the total 
absence of pedicellariae, combine to justify treating B. simplex as distinct from B. spinosus in 
my view and at the same time synonymizing B. armatus with B. simplex. In fact, two smaller 
american paratypes of P. armatus (R c. 40 and 30 mm) and the Portuguese paratype (R 
c. 32 mm) do have only 5 or sometimes 4 furrow spines but this is to be expected at this size 
and is probably only transitory. Also occasional larger specimens of B. simplex lack 
pedicellariae and conversely a few specimens of B. spinosus do have some inferomarginal 

With regard to the relationship of B. chardyi Sibuet with B. simplex, Sibuet had only a 
single specimen, from the Gulf of Guinea off Gabon (Jean Charcot st. CY 19, c. 2 S, 8 E, 
2584m.), R90-100mm, which has been reexamined. It has only a few inconspicuous 
spinelets enlarged near the anus, no disc spines, the papular areas do not bifurcate distally 
(not observed by Sibuet), the accessory inferomarginal spines are relatively large, sometimes 
almost equal in length to the main spine, there are as many as 24 or 25 adambulacrals 
corresponding to the first ten inferomarginals (not noted by Sibuet) and the subambulacral 
spines are almost without exception single; the furrow spines number 6 or 7 on most 
plates as usual in B. simplex the 4 or 5 recorded by Sibuet being limited to some of the 
proximal plates and transitional to the higher numbers. Apart from the last, the combination 
of these characters has not otherwise been found in B. simplex but each of them individually 
is liable to vary. Two (poorly preserved) specimens also from the Gulf of Guinea but to the 
NW off Ghana (Pillsbury st. 34, c. 4 N, 2{ W, c. 1970 m) with R c. 70 mm have a majority of 
the adambulacral plates with one rather than two subambulacral spines. However, one of 
them has two small disc spines developed, their papular areas are bifurcating, the accessory 
inferomarginal spines were probably relatively small and there are only 1\\ adambulacrals 
matching the first ten inferomarginals on the only arm of which sufficient remains to allow a 
count. Two much larger specimens again from the Gulf of Guinea but off western Nigeria 
(Pillsbury st. 314, c. 5 N, 4 E, 2300 m), R 7130-150 mm, both have only a single disc spine 
and a few enlarged spinelets, the pore areas bifurcate, the accessory inferomarginal spines are 
less than half as long as the main spine and there are only c. 2 1 adambulacrals corresponding 
to the first ten inferomarginals but the subambulacral spines are predominantly two. These 
few specimens indicate that in the Gulf of Guinea at least, the development of disc spines 
and a second subambulacral spine is relatively retarded. Whether or not this is sufficiently 
consistent to justify retention of B. chardyi as a distinct taxon remains to be seen from further 
material. At present it seems best to treat it as a subspecies of B. simplex. 

Similar doubts about the specific validity of B. folini must exist considering the inclusion 
in the type material of at least two specimens with conspicuous large odd superomarginal 
spines interradially, not the small multiple spines supposed by Perrier to be characteristic of 
the species. (Most of the remaining paratypes have lost these spines in preservation.) The 
larger of these two probably had R c. 50 mm and the single subambulacral spines would be 
expected in B. simplex at this size. It should also be noted that the holotype of B. chardyi has 
multiple small spines on one of its five odd superomarginal plates. 

Since the recognition of multispinulose specimens as conspecific with B. simplex, only the 
numerous oral furrow and suboral spines in the holotype of B. fischeri at R only c. 40 mm 

134 A.M.CLARK 

serve to distinguish it. In spite of this and the presence of two subambulacral spines on most 
proximal plates, I think it not unlikely that further material from the type locality off NW 
Africa will show the holotype to be only an aberrant B. simplex. 

RANGE. From the vicinity of Cape Cod (42^0 N, 63-66 W) south to the Gulf of Mexico, 
the Colombia and Guyana Basins and from south of Iceland (Thor st. 164) and the Rockall 
Trough (c. 57 N, 10 W) south to the Gulf of Guinea (c. 2{ S, 8 E); 1 1 75-2585 metres. 


Most of the observations on material of the U.S. National Museum and the Museum of 
Comparative Zoology, Harvard, were made during a visit in 1953/54 but I am indebted to 
Miss M. E. Downey and Professor R. M. Woollacott of those institutions for recent loans of 
specimens; also to Professor A. Guille, of the Paris Museum. 


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1917. Three new starfish and one new brittle-star from Chile. Proc. biol. Soc. Wash.SQ : 151-158. 

1954. Echinoderms (other than holothurians) of the Gulf of Mexico. Bull. US Fish Commn 

55 : 373-379. 

Clark, A. M. 1962. Asteroidea. Rep. B.A.N.Z. Antarctic Res. Exped. B9 : 1-104, 18 figs, 5 pis. 
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& Courtman-Stock, J. 1976. The echinoderms of southern Africa. British Museum (Nat. Hist.) 

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1970. Sea stars (Echinodermata : Asteroidea) from 'Eltanin' cruise 26, with a review of the New 

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1926. Echinoderms from the South African Fisheries and Marine Biological Survey. 2. Sea-stars 

(Asteroidea). Rep. Fish. mar. biol. Surv. Un. S. Afr. no. 4. Spec. Rep. no. 7 : 1-34, 7 pis. 

1941. Reports on the scientific results of the Atlantis Expeditions to the West Indies. The 

echinoderms (other than holothurians). Mem. Soc. cubana Hist. nat. 15 (1) : 1-54, 10 pis. 
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126: 1-158, 48 pis. 
Farran, G. P. 1913. The deep-water Asteroidea, Ophiuroidea and Echinoidea of the west coast of 

Ireland. Sclent. Invest. Fish. Brch Ire. 1912 (6) : 1-66, 17 figs, 2 pis. 
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1903 (3): 987-1 130, 49 pis. 

1910. New starfishes from the North Pacific. Zool. Anz. 35 : 545-553; 568-574. 

1911. Asteroidea of the North Pacific and adjacent waters. 1. Phanerozonia and Spinulosa. Bull. 

U.S. natn. Mus. 76 (1) : 1-419, 122 pis. 
1919. Starfishes of the Philippine Seas and adjacent waters. Bull. U.S. natn. Mus. 100 (3) : 1-546, 

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1940. Asteroidea. Discovery Rep. 20 : 69-306, 23 pis. 

Goto, S. 1914. A descriptive monograph of Japanese Asteroidea. J. Coll. Sci. imp. Univ. Tokyo 

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Biological Laboratory, Imperial Household of Japan, xi + 1 14 pp., 13 figs, 18 pis. 
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large des Guyanes et du Venezuela. Bull. zool. Mus. Univ. Amsterdam 6(13): 93-101 , 3 figs. 


Koehler, R. 1907. Asteries, Ophiures et Echinides recueillis dans les mers australes par la Scotia. Zool 

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1924. Les echinodermes de mers d'Europe. 1 .xiii + 360 pp., 9 pis. Paris. 

Ludwig, H. 1903. Seesterne. Result. Voy. Belgica (Zool.) 72 pp., 7 pis. 

1910. Notomyota, eine neue Ordnung des Seesterne. Sber.preuss. Akad. Wiss. 23 : 435^466. 

Macan, T. T. 1938. Asteroidea. Sclent. Rep. John Murray Exped. 4 (9) : 323^35, 12 figs, 6 pis. 
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Skr. Kommn Havunders., Kbh. no. 1 : 16-24. 

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Okutani, T. 1969. Synopsis of bathyal and abyssal megalo-invertebrates from Sagami Bay and the 

south off Boso Peninsula trawled by the R/V Soyo-Maru. Bull. Tokai reg. Fish. Res. Lab. no. 

57: 1-6 1,22 figs, 8 pis. 
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384pp. Paris. [Also published in: Archs Zool. exp. gen. 4(1875): 265-450; 5 (1876) : 1-104, 


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Manuscript accepted for publication 12 January 198 1 

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Zoology series Vol 41 No 4 26 November 1981 

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ISSN 0007-1498 Zoology series 

Vol 41 No 4 pp 137-234 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 26 November 1981 



Observations on clonal cultures of Euglyphidae (Rhizopoda, Protozoa). By C. G. 

Ogden 137 

The Nuttall and British Museum (Natural History) tick collections: lectotype 
designations for ticks (Acarina: Ixodoidea) described by Nuttall, Warburton, 
Cooper and Robinson. By J. E. Keirans&B. E. Brewster 153 

A revision of the spider genus Hispo (Araneae: Salticidae). By F. R. Wanless . . 1 79 
A revision of the spider genus Phaeacius (Araneae: Saliticidae). By F. R. Wanless . 1 99 
The protractor pectoralis muscle and the classification of teleost fishes. By P. H. 

Greenwood & G. V. Lauder Jnr. 


Observations of clonal cultures of Euglyphidae 
(Rhizopoda, Protozoa) 

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


The identification of small siliceous testate amoebae is often difficult due to three main 
factors, observations are made at the limits of the optical microscope, scarcity of specimens 
and inadequate descriptions. The introduction of scanning-electron microscopy for detailed 
surface examination has reduced the first of these obstacles, as this facility permits more 
accurate descriptions to be made of the shells of these animals. However, matching these 
more detailed descriptions with existing reports is often complicated as slight differences of 
shell structure have frequently prompted authors to propose new specific names, or new 
Varieties' or new 'forms'. The second factor can also be reduced by establishing clonal 
cultures in the laboratory; additionally such cultures enable one to study biology and 
variation in morphology. Previous reports (Hedley & Ogden, 1973, 1974; Hedley et al., 
1974) dealt with the biology of four species, namely Euglypha acanthopora (Ehrenberg, 
1841), E. rotunda Wailes, 191 1, E. strigosa (Ehrenberg, 1871) and Trinema lineare Penard, 

The present account describes the shell morphology and biology of four further species, 
based on clonal cultures, together with the redescription of a previous clone under a new 
specific name. 


The genera Euglypha and Assulina belong to the family Euglyphidae the classification 
adopted here is that proposed by Levine et al., 1980 and Loeblich and Tappan, 1964: 

Superclass RHIZOPODA Von Siebold, 1845 

Class FILOSEA Leidy, 1879 

Order GROMIIDA Claparede and Lachmann, 1859 

Superfamily EUGLYPHACEA Loeblich and Tappan, 1 96 1 

Family EUGLYPHIDAE Wallich, 1864 

shell hyaline, symmetrical, elongate, composed 
of rounded siliceous plates, aperture rounded 
or elongate; one nucleus. 

Euglypha acanthophora In identifying specimens from a clonal culture as E. acanthophora 
Hedley et al. (1974) accepted the reported variation attributed to this species, although with 
some reservation because the extent of the variations was not substantiated by our obser- 
vations of other Euglypha species in culture. These differences mainly concerned the shape 
and position of the siliceous spines and the shape of the body plates, which are illustrated in 
Leidy's superb figures (Leidy, 1879) of E. alveolata a species accepted by both Cash et al., 
(1915) and Penard (in Cash et al.) as a synonym of E. acanthophora showing that the 
siliceous spines project either distinctly from the sides of the body or discretely from the 
aboral extremity, and that there are distinct tooth-like projections on the posterior edge of 
the body plates. Penard (1902) proposed two varieties to accommodate specimens with 

Bull. Br. Mus. not. Hist. (Zool.) 41(4): 137-151 Issued 26 November 1 98 1 


138 C. G.OGDEN 

varying spines, and these were accepted by Cash et al., (1915) who also suggested that 
intermediate forms existed. The extent of these variations is given by Decloitre (1962) in a 
review of the genus Euglypha, which lists seven varieties of E. acanthophora, 

The limitation of our research was such that it was expedient at that time (Hedley et al., 
1974) to consider our clonal culture to be representative ofE. acanthophora. The problem of 
designated names now arises with the establishment of another distinct clonal culture within 
the heterogenous description of E. acanthophora. It appears that the most recent culture is 
more closely allied to the description of E. acanthophora, whilst the earlier culture (Hedley 
et al., 1974) fits the description of E. a. var. brevispina given by Cash et al, (1915). However, 
this latter description does not agree with the original (Penard, 1902) as discussed on p. 140, 
so to avoid confusion with these variety names it is proposed that a new specific name, E. 
cashii, be given to the earlier clonal culture. 

Materials and methods 

Information on the source of each species is given with the taxonomic description. 

The animals were obtained from crude cultures made by isolating small portions of each 
sample placed in a small plastic container and covered with a shallow layer of the culture 
liquid. Agnotobiotic cultures were kept at room temperature (18-20C), in similar 
containers, on a thin layer of agar agar (1 per cent in distilled water) with a sterilised wheat 
grain added prior to setting and covered by a shallow layer of culture medium. The culture 
medium was a 5 per cent solution of soil extract plus added nutrient salts (see Hedley & 
Ogden, 1973). Clonal cultures were established by isolating single active animals, and one 
clone has been used subsequently to produce the working cultures of each species. 
Subcultures made at 2-3 week intervals are adequate to maintain active animals. 

Live specimens were examined by optical microscopy using both phase-contrast and 
brightfield illumination. Specimens for scanning electron microscopy were cleaned by 
transferring them through several changes of distilled water. Then manipulated using a 
single-hair brush onto small drops of Araldite adhesive on a small cover slip, the cover slip 
having been previously cleaned and dried. The exception to this procedure were specimens 
of E. dickensii which proved to be fragile and collapsed using this technique. Alternatively 
they were fixed in 3 per cent glutaraldehyde in 1 M cacodylic acid buffer, rinsed in the 
buffer solution and several times in distilled water, then specimens in distilled water 
were micropipetted onto a clean cover slip and allowed to dry. Although some specimens 
still collapsed, the majority retained their natural shape. The prepared cover slips were 
fastened to aluminium stereoscan stubs with Araldite, prior to being coated evenly with a 
thin layer of conducting metal. The stubs were examined on a Cambridge Stereoscan 180 
operating at lOkV and the results recorded on Ilford HP5 film. 

Euglypha acanthophora (Ehrenberg, 1841) 

syn. Euglypha alveolata (Dujardin, 1 84 1 ) in Leidy, 1879, pi. 35, figs 3, 7, 15-18 
Euglypha brachiata Penard, 1902 (Non Leidy, 1878) 
Euglypha brachiata \ar.flexulosa Penard, 1902 

Euglypha brachiata var. brevispina Penard, 1 902 (Non Cash et al., 1915) 
Euglypha armata Wailes and Penard, 1911 
Euglypha crenulata var. minor Wailes, 1912 
Euglypha acanthophora var. dorsalis Schonborn, 1962 

This species was isolated from a sample of Sphagnum moss collected at Holmsley, near 
Burley, New Forest, Hampshire, in March 1979. 



Figs 1-5 Euglypha acanthophora: Figs 1 & 2, lateral views to illustrate the distribution of body 
and elongated plates x 1250 & x 970; Fig. 3, view of aperture with nine apertural plates x 1 170; 
Fig. 4, single apertural plate, note the slight thickening on the dentate margin x 7500; Fig. 5, 
terminal or aboral view showing the distribution of oval body plates x 1200. 

140 C. G. OGDEN 

Table 1 Range of measurements (in 

length breadth depth diameter of aperture 

Euglypha acanthophora 
Euglypha cashii 

a.* (53) 







b. (10) 







Euglypha compressa 












Euglypha dickensii 












Assulina muscorum 












* measurements from Hedley el ai, 1 974; bracketed figures indicate the number of specimens measured, 
a. Clone from Hertfordshire, b. clone from Wales. 

DESCRIPTION. The shell is elliptical or ovoid, circular in cross section, composed of about 
one hundred and thirty body plates and up to six elongated plates or spines (Table 1 , Figs 1 & 
2). The aperture is terminal, circular and surrounded by between eight and eleven, usually 
evenly spaced, denticulate apertural plates (Fig. 3). Each apertural plate is roughly oval, 
between 8'1-10'4 urn in length and 6'3-7 - 8 um in width, often both the denticular and the 
opposite margin are pointed (Fig. 4). The dentate edge is barely thickened on the inner 
surface around the median tooth and the five smaller lateral teeth positioned on each side 
(Fig. 4). Denticular margins are also seen on the second circle of plates surrounding the 
aperture. The body plates are arranged in alternate, longitudinal rows, and range from 
7'5-9'8 urn in length and 6'3-8 - 1 urn in width. Usually the posterior margin of these roughly 
oval plates has a median rounded projection, this appears to be more pronounced when the 
plate has an angular shape (Fig. 6). Body plates appear to vary in shape depending on their 
position, the pointed posterior margin of the plate becoming more pronounced in progres- 
sion down the length of the body (Fig. 1), only to be rounded in some instances at the aboral 
extremity (Fig. 5). The elongated plates are dagger shaped, ranging in length from 18-30 urn, 
and usually project from the sides of the body at an acute angle (Figs 1 & 2). They may be 
positioned anywhere in the posterior half of the shell, but are only rarely seen projecting 
from the extremity. The elongated portion of these plates is thin, hence their brightness when 
bombarded by electrons in the scanning electron microscope (Figs 7 & 8), and vary markedly 
in breadth which in turn alters the degree of tapering. It would appear that narrow and long 
spines are more flexible (Fig. 7), whilst the broader and shorter spines are more robust (Fig. 

DISCUSSION. Both Cash and Penard described E. acanthophora with long thin spines and E. 
acanthophora var. brevispina with short stout spines. There is good agreement between the 
diagrams of E. acanthophora given by Cash (1915: pi. 35, fig. 2), Leidy (1879 pi. 35 figs 3, 7, 
15-18) and Penard (1902 p. 505 and in Wailes & Penard, 1911: pi. 3, fig. 16), but not 
between the diagrams for E. a. var. brevispina, those of Cash and Leidy (1915: pi. 33, figs 3, 
5; 1879 pi. 35, figs 1, 2, 4) being quite different from the original Penard figures (1902 p. 505 
figs 6-10). Whereas the spines shown by the former authors are pointed, Penard's are blunt 
and appear to have broken or ragged edges. As Penard's two varieties brevispina and 
flexulosa differ only in having as their names imply either truncate or flexous spines, 
features which are considered to be artifacts from observations on clonal cultures reported 
here, they are listed as synonyms of E. acanthophora. 

Although the apertural plates in the above mentioned descriptions are similar, there are 
differences in the distribution and shape of the body plates. For example, Leidy (1879) noted 
that there were two rows of denticulate plates surrounding the aperture and that the body 
plates had small projections from their posterior margins. Both Cash and Penard agree with 



Figs 6-8 Euglypha acanthophora: Fig. 6, area of shell surface showing typical body plates x 3300; 
Fig. 7, 'dagger-shaped' elongated plate, a narrow and flexible type x2500; Fig. 8, a stout and 
robust type of elongated plate x 2700. 

the two rows of denticulate plates, but neither commented on the body plates-except for a 
reference in the generic discussion (Cash et al, 1915: p. 5) naming just three species with 
'scutiform' plates : Zs. scutigera Penard, 1911; E. aspera Penard, 1899 and E. crenulata 
Wailes, 1911 nevertheless they synonymize Leidy's description with E. acanthophora. 
Wailes (1912) in describing a new variety, E. crenulata var. minor, noted that it was only 
distinguished from E. armata Wailes & Penard, 1911 (a synonym of E. acanthophora 
proposed by Wailes in Cash et al., 1 9 1 5) by the shape of the body plates. 

Of the eleven varieties listed by Decloitre (1962, 1976), two have already been mentioned, 
E. a. var. brevispina and E. a. \ar.flexulosa; four appear to be similar to E. acanthophora as 
described here, these are E. a. var. cylindracea (Playfair, 1918), E. a. var. dorsalis Schonborn, 
1962, E. a. var. fantastica Decloitre, 1965 and E. a. var. longispina Decloitre, 1969; two, E. 
a. var. equeis Decloitre, 1956 and E. a. var. heterospina Decloitre, 1949, have discrete 
arrangements of spines and may represent distinct species; and the remaining three are 
considered later on p. 143. The four similar varieties differ in size (cylindracea), slight 
deflection of the aperture (dorsalis), the shape of the spines (fantastica), and the number of 
spines (longispina). Specimens with misaligned apertures are not uncommon in clonal 
culture, and E. a. var. dorsalis is therefore considered to be a synonym of E. acanthophora. 
The differences of the three remaining varieties may also be artificial, but it is not possible to 
comment further on these. 

Euglypha cashii nom. nov. 

syn. Euglypha acanthophora var brevispina in Cash et al., 1915 
Euglypha alveolata in Leidy, 1879: pi. 35, figs 1, 2, 4 
Euglypha alveolata var. cirrata Wailes, 1912 
Euglypha acanthophora var. deflanderi Decloitre, 1956 
Euglypha acanthophora var. elegans Stepanek, 1963 
Euglypha acanthophora in Medley et al., 1974 
Euglypha acanthophora in Ogden & Hedley, 1980 

Two clones of this species have been isolated, the first from sewage sludge from the Maple 
Lodge Works of the Colne Valley Sewage Board, Hertfordshire, in December, 1972 and the 
second from a sample of Sphagnum moss from Myndd Hiraethog, Denbigh, Clwyd, North 
Wales, in August, 1980. 



Figs 9-13 Euglypha cashii: Fig. 9, lateral view showing the distribution of body and elongated 
plates x 1250; Fig. 10, aperture with ten evenly distributed apertural plates x 1100; Fig. 11, 
detail of apertural plates x 4400; Fig. 12, elongated plates projecting from aboral 
extremity x 1 800; Fig. 13, elongated plates following the curvature of the shell x 2400. 


DESCRIPTION. The shell is elliptical or ovoid, circular in cross section, composed of about 
two hundred body plates and up to six elongated plates (Table 1 , Fig. 9). The aperture is 
terminal, circular and surrounded by between ten and thirteen evenly spaced denticulate 
apertural plates (Fig. 10). Each apertural plate is roughly oval, varying from 8-1-12-5 um in 
length and 7-2-1 1-5 um in width. The dentate edge is slightly thickened and carries a 
median tooth with either four or five smaller lateral teeth on each side (Fig. 1 1). Specimens 
are sometimes seen with the average number of plates surrounding the aperture but with 
either one or two being dentate body plates, similar to those shown in Fig. la of Hedley et al. 
(1974), instead of dentate apertural plates. The body plates are arranged in alternate 
longitudinal rows and range from 10-9-12 um in length and 7-9-9-1 um in width. Each oval 
body plate has a median rounded projection on the posterior margin, these projections are 
not as pronounced as in E. acanthophora (compare Figs 1 & 9). The elongated plates project 
slightly from the aboral extremity, or closely follow the outline of the shell in the aboral 
region, they range from 19-6-29-2 um in length and 6- 1-7-6 um in width (Figs 12 & 13). 

DISCUSSION. The description of this species is in good agreement with that given by Cash et 
al. (1 9 1 5: pi. 33, figs 3 & 5) for E. a. var. brevispina. As there is some disparity between these 
two descriptions and the original description of the variety brevispina given by Penard (1902) 
(see also p. 140), in addition to the variety name brevispina being used with other species of 
Euglypha, it was considered that a new specific name would eliminate future confusion of 
these species. E. cashii is distinguished from E. acanthophora by size, shape of body and 
elongate plates, but mainly by the positioning of the elongated plates. 

The three varieties of E. acanthophora listed by Decloitre (1962, 1976) which are similar 
to E. cashii are E. a. var. cirrata: E. a. var. deflanderi and E. a. var. elegans, they apparently 
differ only in the deflection of the posterior spines or elongated plates to the shell body. Such 
features have been seen in clonal culture as a normal variation, and these varieties are there- 
fore considered to be synonyms of E. cashii. Another species, E. tiscia Gal, 1969, listed by 
Decloitre (1976), although similar to E. cashii is considered to represent a valid species 
because of its larger dimensions. 

ETYMOLOGY. This species is named after Mr James Cash, who contributed so much to the 
taxonomy of testate amoebae at the beginning of this century. 

Euglypha compressa Carter, 1 864 

This species was isolated from a sample of Sphagnum moss, collected at Subberthwaite, 
Broughton in Furness, Cumbria, in July, 1978. 

DESCRIPTION. The shell is ovoid, laterally compressed and composed of about two hundred 
body plates and thirty spines (Table 1, Fig. 14). The aperture is terminal, circular or 
elliptical and surrounded by between eleven and fifteen, evenly spaced, denticulate apertural 
plates (Fig. 15). Each apertural plate is oval, between 6-2-7-1 um in length and 4-6-5-4 um 
in width. The dentate edge is distally thickened around the large median tooth, but this 
thickening tapers evenly outwards to the three smaller teeth positioned on each side and 
equates with the normal shell thickness close to the last tooth (Fig. 16). The body plates are 
arranged in sequence and range in size from 7'4-8'2 um in length and 3-4-4-4 um in width. 
They are roughly oval and often characterised by their hexagonal appearance, this latter 
feature is variable but the squared nature of the narrow margins is usually distinctive (Fig. 
17). The spines are positioned along the lateral margins from about the mid-body position to 
the aboral extremity (Fig. 15). They are sometimes in pairs and usually alternate when 
viewed laterally, one pointing upwards and the next downwards along the length of the body 
(Figs 14 & 15). Each spine has a narrow base from which it tapers to its widest point about 
a quarter of the total spine length, then it tapers gradually over the remaining threequarters 
to the terminal point (Fig. 1 8). 

DISCUSSION. The only recorded variety of this species is E. compressa f. glabra Cash et al., 



Figs 14-18 Euglypha compressa: Fig. 14, lateral view showing the distribution of body plates and 
spines x 1250; Fig. 15, latero-apertural view to illustrate the circular aperture and the 
positioning of the spines x910; Fig. 16, detail of single apertural plate, note the thickening 
around the dentate margin x9500; Fig. 17, arrangement of typical body plates x2350; Fig. 18, 
illustration of spine projecting from organic cement matrix between two adjacent body plates 
x 4400. 


1915, a form without spines. However, variation in the shape and size of the siliceous spines 
was also reported by Cash et al. for E. compressa, and such differences were considered to be 
a normal feature. This may not now prove to be the case, if, as the information derived from 
clonal cultures in the present work shows that differences in spine positioning is specific, 
then it would appear to follow that structural differences of spine construction may also be 
specific. The spines described here are similar to those reported by de Graaf (1956) and 
Brown (1910), but they are different from those described by Ogden & Hedley (1980). In the 
latter report the spines are stout at the base, concave, and taper evenly to a point (see 
PI. 78D, Ogden & Hedley, 1980). Both types of spine are here considered to represent E. 
compressa, which is distinguished by the compressed shell and lateral position of the spines, 
but further work on clonal cultures may establish spine shape as a specific character. 

Euglypha dickensii sp. nov. 

This species was isolated from a sample of damp moss taken from underneath sweet chestnut 
trees at Cobham woods, Rochester, Kent in February, 1974. 

DESCRIPTION. The shell is ovoid, laterally compressed and composed of about two hundred 
and eighty elongate body plates (Table 1, Figs 19 & 20). The aperture is terminal, oval or 
circular, and surrounded by between eleven and fifteen, evenly spaced, denticulate apertural 
plates (Fig. 21). In a few specimens additional apertural plates are seen in the second circle of 
plates around the aperture. Each apertural plate is roughly circular, from 5-4-6-4 um in 
length and 3'8-4'4 um in width. The dentate edge has a large, thick, distinctly curved, 
median process with a terminal pointed tooth, this is flanked on each side by a medium 
outward facing tooth and one or two smaller teeth (Figs 22 & 24). The denticular thickening 
equates with the normal shell thickness at the position of the small teeth. The body plates are 
elongate, ranging from 5*1-6-2 um in length and 1-6-2-5 um in width, and are arranged in 
alternate longitudinal rows. Around the mid-body region there are some randomly 
distributed pointed body plates (Fig. 23). These pointed plates vary from normal plates with 
a small sharp spike, about 6*9 um long, to tapered spines about twice the length, 9- 1 um, of a 
normal body plate. Although there is variation in the dimensions of these pointed body 
plates, their presence in a mid-body position is a reasonably stable feature. 

DISCUSSION. The species described here is similar to three species previously reported from 
soil samples, namely E. capsiosa Couteaux, 1978, E. cuspidata Bonnet & Thomas, 1960 
and E. simplex Decloitre, 1965. It has similar dimensions to E. simplex but differs in the 
shape of the apertural plates, which in the latter species have a distinct diamond-shape (see 
Couteaux et al, 1 979). Both E. capsiosa and E. cuspidata are smaller species with fewer body 
and apertural plates, and again differ in the shape of the apertural plates. E. dickensii is 
distinct in size, dentation of the apertural plates and the presence of pointed body plates in 
the mid-body region. 

ETYMOLOGY. As this species was found in the countryside frequented by Charles Dickens, 
the nineteenth century author, and subsequently featured in many of his novels, it is named 
in his honour. 

Assulina muscorum Greef, 1888 

This species was isolated from a sample of dry moss on soil, collected at Rolestone Farm, 
Banwell, Somerset in February, 1974. 

DESCRIPTION. The shell is ovoid, laterally compressed and composed of about two hundred 
shell plates (Figs 25 & 26). The aperture is terminal and surrounded by between ten and 
fourteen shell plates arranged in a rather irregular manner, most with their minor axis 
bordering the opening but often a few have their major axis (Figs 27 & 28). The opening is 
edged with a thin band of organic cement, this band is frequently thickened on the tips of 
some plates to form tooth-like projections (Fig. 28). The shell plates are oval, ranging from 
5'8-6'8 um in length and 2-5-3-1 um in width. They are usually arranged in alternate, 



Figs 19-24 Euglypha dickensii: Fig. 19, lateral view showing the distribution of body plates 
x 1970; Fig. 20, view to illustrate the lateral compression x 1300; Fig 21, aperture with fourteen 
apertural plates x 2400; Fig. 22, view showing the overlapping of the apertural plates and the 
thickness of the median tooth x 7800; Fig. 23, portion of shell surface with two pointed body 
plates x 4600; Fig. 24, circular apertural plate with typical dentate margin x 1 1000. 




Figs 25-28 Assulina muscorum: Fig. 25, lateral view to illustrate the distribution of shell plates 
x2150; Fig. 26, view to show lateral compression and arrangement of plates x 1400; Fig. 27, 
apertural view x 2100, Fig. 28, detail of 'tooth-like' projections of organic cement around the 
aperture x 4900. 

148 C. G. OGDEN 

longitudinal rows, with their major axis parallel to the major axis of the body. However, in 
some instances the axes of the plates are not parallel with the body and in these cases the 
general pattern is altered. The aboral extremity is also subject to irregular arrangements of 
shell plates. 

DISCUSSION. The shell of A. muscorum is reported as being usually brown but occasionally 
colourless in the wild, whereas in culture it is mainly colourless. Nevertheless, live animals 
have a distinct band, probably the 'pigment zone', at the mid-body region which is often so 
large that it may tend to give colour to the shell. Variation in structure of the shell appears to 
be limited to the formation of an extra large individual, usually confined to less than three 
per cent of the population. Such specimens have more than the normal compliment of shell 
plates but the arrangement is the same. Similar large specimens have been reported (Hedley 
& Ogden, 1973) in clonal cultures of Euglypha rotunda. 


The formation of a daughter-cell by simple division has been observed for all the described 
species and follows the same pattern in each, the only variation being the additional arrange- 
ment of elongated plates or spines in E. acanthophora, E. cashii and E. compressa. The 
sequence of events in E. compressa are described. 

Euglypha compressa 

The onset of division begins with the protrusion of a short thick cytoplasmic extension from 
the parent aperture. The apertural plates are the first to be passed from the storage area 
adjacent to the nucleus, via the peripheral cytoplasm to become arranged in a circle around 
this cytoplasmic extension. The remainder of the plates follow the same route and are 
arranged in sequence, in distinct rows radiating from the apertural plates (Fig. 29a). Each 
plate is added on the inside of the previous plate so that there is a considerable overlap. This 
excessive overlapping of the plates in the region of attachment is very noticeable in the early 
stages of shell construction. All the body plates are therefore in position, but not correctly 
spaced, well in advance of the shell attaining its full size. When the daughter shell has 
attained a size about two-thirds that of the parent, vacuolar activity is seen in the anterior 
third of the parental cytoplasm. This activity proceeds into the daughter cytoplasm as it 
increases in volume, at the same time the zone of pigment granules begin to extend towards 
the aperture of the parent. The spines are the last elements to be passed from the parent to 
the daughter (Figs 29b & c), where they are arranged centrally in the cytoplasm parallel 
to the shell walls. The pair of mid-body spines are the first to be pushed by cytoplasmic 

Fig. 29 Diagram illustrating division in Euglypha compressa: formation of daughter shell (a-e) 
followed by nuclear division (f-1). 

a. body plates (bp) pass via the peripheral cytoplasm to extremity of cytoplasmic extension of 
parent, 10-15 minutes; b. as last plates pass into forming daughter shell, siliceous spines (ss) 
begin moving from parent, 20 minutes; c. spines move into daughter shell, cytoplasmic vacuoles 
(v) appear close to parent aperture and granules of pigment zone (pz) begin to move, 25 minutes; 
d. spines arranged for positioning in daughter, and granules in parent move towards aperture, 27 
minutes; e. daughter shell full size with some spines in position and cytoplasm filled with 
cytoplasmic vacuoles, granules begin to extend from parent into daughter, 30 minutes; 
f. daughter shell complete, granules in position between the opposed apertures and nucleus (n) in 
parent has small polar extensions, 40 minutes; g. elongation of parent nucleus, 55 minutes; h. 
granules mainly in daughter and nucleus now 'diamond-shape', 60 minutes; i. elongated nucleus 
becoming indistinct behind moving granules, 70 minutes; j. arrows depict pathways of rapidly 
moving granules, 75 minutes; k. movement of granules has slowed down and zones beginning to 
reform, faint nucleii appear, 90 minutes; 1. contractile vacuoles (cv) and nucleii apparent, pig- 
ment zones almost reformed, 100 minutes. The times given are based on an estimated starting 



150 C. G.OGDEN 

movement out between adjacent, lateral body plates, they appear to be positioned on each 
side simultaneously. This pair are followed in succession by other pairs of spines in a 
progressive sequence towards the aboral region. The last spines to be positioned are those 
that project from the terminal region. These spines are manoeuvred in the cytoplasm (Fig. 
29e) until they are parallel to the aboral shell surface, they are then pushed through and 
cytoplasmic movement continues until the spines are at right angles to the shell surface. 
Throughout the time taken to position the spines those that project from the shell are in 
continuous movement, backwards and forwards like oarsmen in a rowing-boat except that 
their movements are not synchronized, and there are smaller movements still visible 
amongst the body plates. Vacuolar activity seems to be concentrated mainly in the daughter 
with the pigment granules being pushed from the parent into the peripheral cytoplasm of the 
daughter. A halo effect is most noticeable around the parent nucleus at this time. 

A few seconds after the last aboral spines are in position, movement of all the shell 
elements slows down, until the spines are projecting slightly posteriorly when viewed from 
above. All movement has stopped after a further two minutes from the final positioning of 
the last aboral spines, and the daughter shell is complete. At about the same time the parent 
nucleus begins to elongate, initially there are two small polar extensions (Fig. 290, but this 
changes into a 'diamond-shape' as most of the granules are passed into the daughter cyto- 
plasm (Fig. 29g). Slightly later, movement of the granules in the daughter cytoplasm seem to 
suggest that when they reach the aboral extremity they are reflected back via the peripheral 
cytoplasm to the parent, meanwhile the nucleus has become more elongate (Fig. 29h). Rapid 
movement of the granules now obscures the changing nucleus (Fig. 29i), and a regular 
pathway of constantly moving granules is formed between the parent and daughter cyto- 
plasm (Fig. 29j). When the rapid movement ceases, after about five minutes it is replaced by 
a slow regular movement with no apparent directional flow, cytoplasm of both cells looking 
homogenous. After a further five minutes the apertural region of both are relatively free of 
granules, contractile vacuoles are apparent in the anterior region of each and there is some 
movement of granules (Fig. 29k). A little while later nuclei are apparent in the posterior 
region of both cells as the pigment granules are concentrated into the mid-body region (Fig. 
291). Just prior to separation the cytoplasm in each shell is withdrawn slightly so that the 
cytoplasmic connection is severed, the animals move apart independently shortly after this 

The approximate time taken to produce the daughter shell is forty-five minutes, whilst the 
total time for division is one hundred minutes. 

Doubling time 

Estimations on the length of time required to double the population (doubling time) were 
made on cultures established and maintained under similar conditions. Growth curves 
calculated from records of daily counts of individuals were made on three replicate cultures 
of four species, but only one culture was available for E. acanthophora. The results are given 
in Table 2. 

Table 2 Estimates of doubling time (in days). 

Euglypha acanthophora 2*7 

Euglypha cashii 2-3-2-8 

Euglypha compressa 4-0-4-4 

Euglypha dickensii 2-6-3- 1 

Assulina muscorum 2-3-2-9 


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Limnologica 1 : 83-91. 

StCpanek, M. 1 963. Die Rhizopoden aus Katanga (Kongo-Afrika). Annls. Mus. r. Afr. cent. 1 17 : 9-9 1 . 
Wailes, G. H. 1912. Freshwater Rhizopoda and Heliozoa from the States of New York, New Jersey and 

Georgia, U.S.A.; with supplemental note on Seychelles species. J. Linn. Soc. Zool. 32 : 121-161. 
& Penard, E. 19 1 1 . Clare Island Survey. Part 65. Rhizopoda. Proc. R. Ir. Acad. 31 : 1-64. 

Manuscript accepted for publication 12 December 1980 

The Nuttall and British Museum (Natural History) 
tick collections: lectotype designations for ticks 
(Acarina: Ixodoidea) described by Nuttall, 
Warburton, Cooper and Robinson 

James E. Keirans 

Rocky Mountain Laboratories, Hamilton, Montana 59840, U.S.A. 
Bernice E. Brewster 

Arachnida Section, Zoology Department, British Museum (Natural History), London 
SW7 5BD 


A monographic revision of the Ixodoidea was begun by Nuttall et al. (1908) with subsequent 
volumes published by Nuttall & Warburton (1911, 1915) and Robinson (1926). A further 
volume was produced by Arthur (1960). These works were based largely on material 
gathered by Nuttall and his associates and the Entomological Research Committee for 
Tropical Africa, forerunner of the present day Commonwealth Institute of Entomology. 
During his lifetime, Professor Nuttall presented a considerable number of specimens to his 
correspondents. In 1939, the remainder of the Nuttall Collection was donated by the 
Molteno Institute to the British Museum (Natural History) and at that time it was the largest 
tick collection in the world. Currently, the Nuttall collection is kept separately from the 
main tick collection of the British Museum (Natural History), and types of Ixodoidea 
designated by Nuttall et al. may be found in both collections. 

Prior to the formulation of the International Code of Zoological Nomenclature, type 
specimens were frequently labelled only as 'TYPE' or 'COTYPE' with the result that many 
of the taxa described by Nuttall et al. were in need of lectotype designations. Consequently 
these studies were undertaken and form the basis of the present report. At the same time an 
attempt was made to trace the type material of all taxa described by Nuttall and his 
associates; appendices are therefore provided listing holotypes, previously selected 
lectotypes and other types together with their location, where known. 

The following list of lectotypes is in alphabetical order by specific name within the genera 
Argas and Ornithodoros in the family Argasidae and within the genera Amblyomma, 
Aponomma, Dermacentor, Haemaphysalis, Ixodes, Rhipicentor, and Rhipicephalus in the 
family Ixodidae. First the species binomen, or in a few cases the trinomen as originally 
described, is given followed by author(s), date, page, and figure(s). Any currently recognized 
synonyms and the Nuttall tick catalogue number precede a statement detailing all pertinent 
information relating to the original collection. Next, the current status of the type material is 
given, including selection of a lectotype and paralectotypes and an updating of the host and 
locality data. This is followed by subsequent published information relating to the species in 
question. Finally, there is a brief statement which summarizes hosts and distribution of each 
tick species. 

During the course of the studies reported below, each lectotype and each paralectotypic 
series was isolated within the vial or jar containing specimens of the taxon; the nomenclative 
status of this material was clearly indicated on an additional included label. 

Bull. Br. Mm. not. Hist. (Zool.) 41(4): 1 53-1 78- Issued 26 November 198 1 




Argas reflexus indicus Warburton, 1910: 396. 

NUTTALL 1649. The Indian Museum collection contained three specimens of this form. One 
was found in a book in the Entomological room, and another on a wall of the Museum 
building. The third specimen was found in a box sent from the Museum to Kurseong, E. 
Himalayas. The birds building in the roof of the Museum were identified as Passer 
domesticus and Cypselus affinis. The type species has not been recorded from India.' 

TYPES. Lectotype 9 Argas reflexus indicus (N1649) found in a book, Entomological room, 
Indian Museum, Calcutta (22'32'N, 8822'E), India, 15.111.1910, C. A. Paiva. [Date and 
collector information taken from vial label]. 

REMARKS. A second collection (N1243) consisting of 1 cf, 1 9, 1 N A. reflexus indicus, found 
in basin of water, Entomological room, Indian Museum, Calcutta, 13. IV. 19 10, F. H. 
Gravely, has a blue label with the word TYPE. We have not accorded these specimens 
paralectotype status because the specimen numbers and locality data do not conform to 
Warburton 's published record. 

Argas reflexus is not known to occur east of 1500' longitude and without additional 
collections, the exact taxonomic status of A. r. indicus remains moot. 

HOSTS & DISTRIBUTION. Argas reflexus indicus is an avian parasite known only from India. 

Ornithodoros gurneyi Warburton, 1926 : 5 5, fig. 1 

NUTTALL 3562. 'Described from two specimens taken in an old shed at Tibboburra, N.S.W., 
24. XI. 1922, by Mr W. B. Gurney and sent with other ticks by the Government 
Entomologist of N.S.W. in 1924 to the Imperial Bureau of Entomology, where it was 
numbered 1378.' 

TYPES. Lectotype cf Ornithodoros gurneyi (N3562) taken in an old shed, Pindera Selection, 
Tibboburra, New South Wales, Australia; 1 N paralectotype Ornithodoros gurneyi, date as 
above, deposited in the BM(NH) 97 collection (1925.5.9.4). 

HOSTS & DISTRIBUTION. Ornithodoros gurneyi is primarily a parasite of large macropodids 
(kangaroos and wallaroos) but will also attack man and dogs in most areas of Australia. 

Ornithodoros piriformis Warburton, 1981 : 285, fig. 2 

NUTTALL 3335. 'Described from 15 specimens taken at Mahabaleshwar, Satara District, at a 
height of 4200 ft, (?host)by F. H. G., 13-16.IV.12. Indian Mus. SPA./10.' 

TYPES. Lectotype 9 and 2 9, 1 cf paralectotypes Ornithodoros piriformis (N3335) host 
unknown, Robber's Cave, Mahabaleshwar (1755'N, 7340'E), Maharashtra, India, at a 
height of 1280 m, 13-16.IV.1912, F. H. G. 

REMARKS. The locality Robber's Cave is taken from NuttalPs catalogue and on the vial label 
with the ticks. Two slides, one with hypostome palp and leg I, the other with hypostome, are 
both missing. 

HOSTS & DISTRIBUTION. Ornithodoros piriformis has been collected from Miniopterus 
schreibersii from the type locality. Robber's Cave contains several other bat species 
including Rousettus leschenaulti and probably Rhinolophus affinis. This tick is known only 
from India. 


Amblyomma atrogenatum Nuttall & Warburton, 1908 : 409, figs 30-32 
Junior subjective synonym of Amblyomma clypeolatum Neumann, 1 899. 


NUTTALL 43. 'Description based on 6 d 1 from Testudo elegans, received from India at the 
Zoological Gardens, London, 15. VIII. removed 2. X.I 905. (Preserved in spirit.)' 

TYPES. Lectotype d 1 and 4 d 1 paralectotypes Amblyomma atrogenatum (N43) ex Testudo 
elegans received from India at Zoological Gardens, London, 15.VHI.1905. Ticks removed 

REMARKS. The 4 rf paralectotype total includes 1 d pinned specimen. An empty token tube 
in the Nuttall collection indicates a slide mount of 1 cf which is now missing. 

HOSTS & DISTRIBUTION. Amblyomma clypeolatum is a tortoise parasite in India and Sri 

Amblyomma caelatumm Cooper & Robinson, 1908 : 460, figs 6-9 
Junior subjective synonym of Amblyomma geoemydae (Cantor, 1 847). 

NUTTALL 2938. Original description 'based on two d"s and one gorged 9. Collected by Mr F. 
F. Laidlaw, Skeat Expedition, 1899, Kwala Aring, Java. Host, a tortoise (Geoemyda 
spinosd). Preserved in alcohol.' 

TYPES. Lectotype 3 and 1 d 1 , 1 $ paralectotypes Amblyomma caelaturum (N2938), ex 
Heosemys spinosa, Kwala Aring, Malaysia (locality not verified) collected by Dr N. 
Annandale, 1899. Gift of Mr F. F. Laidlaw. Correction of locality and collector: Robinson 

REMARKS. The name A. caelaturum was published in July, 1908. The name A. maylanum 
Neumann was published in March of the same year. Anastos (1950 : 107) sank the former as 
a junior synonym of A. malayanum on the grounds of priority. After examination of all 
relevant types Kohls (1957 : 87) concluded that both A. caelaturum and A. malayanum were 
junior synonyms of A. geoemydae (Cantor, 1 847). 

HOSTS & DISTRIBUTION. Amblyomma geoemydae parasitizes reptiles, especially tortoises and 
monitor lizards in Indonesia, Malaysia, Thailand, Ryukyu Islands, Japan, Taiwan, and the 

Amblyomma cooperi Nuttall & Warburton, 1908 : 410, figs 33-36 

Junior synonyms: Amblyomma lutzi Aragao, 1908:112; Amblyomma ypsilophorum Schulze, 
1941 :93. 

NUTTALL 70. Original description, 'based on 1 d 1 and 5 9 collected from "Carpinchio" 
(Hydrochoerus capybard) at Puerto Cooper, Alto, Paraguay, S. America, by W. F. Cooper in 
1904. (Dry specimens.)' 

TYPES. Lectotype d 1 and 1 d, 6 cf paralectotypes Amblyomma cooperi, (N70), ex 
Hydrochoerus hydrochaeris Puerto Cooper (2303'S, 5743'W), Alto, Paraguay, 1904, W. F. 
Cooper. [Included in the above total is one slide-mounted 9]. 

REMARKS. Robinson (1926 : 75) indicated that more than 1 rf was present in the original 
collection but gave no total. 

HOSTS & DISTRIBUTION. Amblyomma cooperi is a parasite of capybaras and tapirs in 
Paraguay, Bolivia, Brazil, Uruguay, and Argentina. 

Amblyomma darlingi Nuttall, 1912 : 50, figs 1^ 
Junior synonym of Amblyomma oblongoguttatum Koch, 1844. 

NUTTALL 1285-1287. 'Described from 5 d 1 , 9 9, 40 N, and 23 larvae found on a deer 
(Odocoleus sp.), Corozal, Panama Canal Zone, 14.VIII.1910, by A. H. Jennings, of Ancon, 
and 2 N, found on the nape and head of a turkey buzzard (Catharista atratus), Empire, Canal 


Zone, IV. 191 1, by S. T. Darling, M. D., Chief of Laboratory, Isthmian Canal Commission, 
Ancon, Canal Zone. Named in honour of Dr Darling in view of his distinguished service to 
parasitology. The types, for which I am indebted to Dr Darling, are in Cambridge 

TYPES. Lectotype rf and 2 <?, 1 9 paralectotypes Amblyomma darlingi (N1285) ex Odocoileus 
virginianus, Corozal (08'59'N, 7935'W), Panama, 14.VIII.1910, A. H. Jennings [above 
paralectotype total includes 1 cT, 1 9 which were pinned and are now in the BM(NH) 
collection] 33N, 20L paralectotypes Amblyomma darlingi (N1286) same collection data as 
N1285; 2N paralectotypes Amblyomma darlingi (N1287) ex Coragyps atratus, perhaps from 
Empire Known Distance Range (09'04'N, 7940'W), Panama, IV. 19 1 1 , Dr S. T. Darling. 

HOSTS & DISTRIBUTION. Amblyomma oblongoguttatum occurs on a wide variety of 
mammalian hosts including deer, peccary, horse, cattle, dog, goat, armadillo, raccoon, etc., 
and ranges from the State of Sinaloa, Mexico, southward through Central America to 
Venezuela and Brazil. 

Amblyomma fiebrigi Robinson, 1912: 482, figs 3, 4 
Junior subjective synonym of Amblyomma pacae Aragao, 1911. 

NUTTALL 2939. 'Described from 1 cf and 1 9, collected at San Bernardino, Paraguay, S. 
America, by Dr Karl Fiebrig. Host not specified.' 

TYPES. Lectotype cf and 1 9 paralectotype Amblyomma fiebrigi (N2939), host unknown, San 
Bernardino (2516'S, 5719'W), Paraguay, Dr Karl Fiebrig. 

REMARKS. There is no date given for this collection in the original description but both 
Nuttall's tick catalogue and the vial label with the ticks bear the date 1911. 

HOSTS & DISTRIBUTION. Amblyomma pacae is most often collected, although not in large 
numbers, from Agouti paca but it has also been recorded from the genera Dasyprocta and 
Tamandua. This tick has been collected in British Honduras, Panama, Colombia, 
Venezuela, Brazil and Paraguay. 

Amblyomma laticaudae Warburton, 1933 : 561, figs 5, 6 

BM(NH) 1933.3.15.24-26. 'Described from 1 cf, 1 9 (gorged) and 1 nymph taken from a 
sea-snake, Laticauda colubrina. The ticks were adhering to the skin, shed in captivity, at the 
Raffles Museum, Singapore, and were sent to Cambridge by the Assistant Curator, Mr 
Norman Smedley, who adds the following note: "The large 9 and others were of the same 
grey as the snake and usually attached to the grey parts of the body, although I observed one 
on a black stripe. Laticauda colubrina belongs to that group of the sea-snakes possessing 
broad ventrals, and spends quite a considerable part of its time out of water, but even so the 
survival of the ticks while the snake is submerged seems remarkable." Types in the British 
Museum (Nat. Hist.).' 

TYPES. Lectotype cf and 1 9, 1 N paralectotypes Amblyomma laticaudae BM(NH) 
1933.3.15.24-26 ex Laticauda colubrina, Raffles Museum, Singapore (0117'N, 10351'E), 
Republic of Singapore, VI. 1930, sent by Mr Norman Smedley. 

REMARKS. Collection date of VI. 1930 taken from vial label. 

HOSTS & DISTRIBUTION. Amblyomma laticaudae has been collected only from Laticauda 
colubrina, a venomous sea snake which is widely distributed throughout the tropical Indo- 
Pacific Islands, southern Japan to the Bay of Bengal. 

Amblyomma robinsoni Warburton, 1927 : 408, pi. xxvii, figs 1,2 

NUTTALL 3617. 'Described from 8 cf and 3 9 taken from Varanus komodoensis and 
contained in a tube labelled "Komodo VII, 1923, Museum Buitenzorg," Java.' 


TYPES. Lectotype cf and 2 cf, 1 9 paralectotypes Amblyomma robinsoni (N3617), ex Varanus 
komodoensis, Komodo Island (0835'S, 11928'E), Lesser Sunda Islands, Indonesia, 
VII. 1923, ex Museum, Buitenzorg, Java. 

HOSTS & DISTRIBUTION. Amblyomma robinsoni is apparently restricted to the giant monitor, 
Varanus komodoensis, on Komodo Island. 

Amblyomma uncatum Nuttall & Warburton, 1908 : 412, figs 37^0 
Junior subjective synonym of Amblyomma nodosum Neumann, 1899. 

NUTTALL 36. 'Description based on 5 cf from Tamandua tetradactyla, S. America (Zool. 
Society's Gardens, London, 1905. Dry specimens).' 

TYPES. Lectotype cf and 6 cf paralectoypes Amblyomma uncatum (N36) [Lectotype and one 
paralectotype in alcohol, 2 paralectotypes pinned, 3 paralectotypes mounted on slides] ex 
Tamandua tetradactyla, South America, 1905. 

REMARKS. The host (an anteater), from which the type series was collected, arrived at the 
Zoological Society's Gardens, London, 10.XII.1905 and died 22.XII.1905 (Robinson 
1926 : 199). The date on the vial label is 22.XII.1905. There are also present two additional 
slides, one with the capitulum and leg IV of a cf, and the second with chelicerae and 
hypostome of a cf. Evidently, Nuttall & Warburton based their description on 5 of the 7 
specimens comprising the collection. 

HOSTS & DISTRIBUTION. A. nodosum adults are specific parasites of anteaters and the species 
has been reported from Costa Rica, Panama, Guatemala, Colombia, Bolivia, Nicaragua, 
Brazil and Trinidad. 

Amblyomma v-notatum Nuttall, 1910 : 412, fig. 5 
Junior subjective synonym of Amblyomma geayi Neumann, 1 899. 

NUTTALL 1 149. 'Described from (Nl 149) 4 9*5 found on a sloth (Bradypus tridactylus), at 
Manaos, Brazil, by Mr T. P. Beddoes, 1903.' 

TYPES. Lectotype 9 and 2 9 paralectotypes Amblyomma v-notatum (N1149) ex Bradypus 
tridactylus Manaus (0308'S, 60'01'W), Brazil, 1903, T. P. Beddoes. 

HOSTS & DISTRIBUTION. Amblyomma geayi is parasitic on both B. tridactylus, the three- 
toed sloth and Choloepus spp. two-toed sloths in Panama, Colombia, Guyana, Surinam, 
Brazil and Peru. 

Amblyomma variegatum nocens Robinson, 1912 : 480, fig. 2 
Junior subjective synonym of Amblyomma pomposum Donitz, 1909. 

NUTTALL 2941. 'Rhodesia, South Africa. Host not specified. Described from 2 cf's collected 
by Mr E. M. Jarvis, F.R.C.V.S., of the Dept. of Agriculture, Southern Rhodesia.' 

TYPES. Lectotype cf Amblyomma variegatum nocens (N2941), Zimbabwe, Mr E. M. Jarvis. 

REMARKS: There is no date given for this collection in the original description but both 
Nuttall's tick catalogue and the vial label with the tick bear the date 1911. Robinson 
(1926 : 100) stated that this collection was '?off cattle'. The second cf specimen of the type 
series could not be located. 

HOSTS & DISTRIBUTION Amblyomma pomposum is most often collected from domestic 
cattle but is found on many other Artiodactyla. It is distributed in highland areas from 
Uganda and Zaire southward to Zambia. 


Amblyomma versicolor Nuttall & Warburton, 1908 : 407, figs 27-29 
Junior synonym of Amblyomma cajennense (Fabricius, 1 787). 

NUTTALL 212. 'Description based on 1 d 1 and 1 9 from a horse, Tolosa, Oaxaca, Mexico, 
collected by Mr A. Laughton, 1907. (Preserved in 70% spirit.)' 

TYPES. Lectotype rf and 1 9 paralectotype Amblyomma versicolor (N212) ex Equus caballus, 
Tolosa (2231'N, 10 1'22'W), Mexico, I.X.I 907, Mr A. Laughton. 

HOSTS & DISTRIBUTION. A. cajennense is a common species occurring in abundance on 
domesticated animals and less frequently from wild hosts. It readily attacks man and has 
been found naturally infected with the etiologic agent of Rocky Mountain spotted fever, 
Rickettsia rickettsii. The tick ranges from southern Texas in the United States southward to 
Argentina and the Caribbean islands. 

Aponomma draconis Warburton, 1933 : 546, fig. 7 
Junior subjective synonym of Aponomma komodoense Oudemans, 1928. 

NUTTALL 3875. 'Described from 3 dtf and 6 <$<3 (in poor condition) taken from a "dragon", 
Varanus komodoensis, brought from Komodo Island, Flores, Dutch East Indies, and for 
some years past in the Zoological Gardens, London. Communicated by Dr P. A. Buxton, 
April, 193 1 . Types in Cambridge (N. 3875).' 

TYPES. Lectotype cf and 2 rf, 6 9 paralectotypes Aponomma draconis (N3875) ex Varanus 
komodoensis, brought from Komodo Island (0836'S, 11930'E) Indonesia, and for some 
years past in the Zoological Gardens, London, England, IV. 193 1 . Dr P. A. Buxton. 

HOSTS & DISTRIBUTION. Aponomma komodoense has been collected on Komodo and 
nearby Flores Island, Indonesia, on Varanus komodoensis. One record of this tick from 
Varanus salvator in Djakarta Zoo was probably a case of accidental parasitism. 

Dermacentor imitans Warburton, 1933 : 559, figs 3, 4 

NUTTALL 3874. 'Described from 12 d and 1 9 taken by Mr W. J. Hamilton, Jun., on a peccary 
(Pecari angulatus) at Turrialba, Costa Rica, on August 1st, 1927, and sent to Cambridge by 
Prof. R. Matheson. In the same tube were specimens of Amblyomma mantiquirense. Types 
in Cambridge (N.3874).' 

TYPES. Lectotype <3 and 9 rf, 1 9 paralectotypes Dermacentor imitans (N 3874), ex Tayassu 
tajacu, Turrialba (0954'N,83'41'W), Costa Rica, 1.VIII.1927, W. J. Hamilton, Jr. 

HOSTS & DISTRIBUTION. Known also from Panama, Guatemala, and Mexico, Dermacentor 
imitans has been recorded from Mazama americana as well as from its common host, the 

Haemaphysalis aciculifer Warburton, 1913 : 125, figs 4, 5 

NUTTALL 1981. 'Described from 1 rf and 1 9 taken, in company with Rhipicephalus 
appendiculatus, from Cobus thomasi (an antelope), on the N.E. shore of Lake Edward, 
Uganda, X. 191 1, by S. A. Neave. Types in Entomol. Research Committee's Collection, No. 
463 a.' 

TYPES. Lectotype d 1 [designated as holotype by Hoogstraal & El Kammah, 1972:965], 
paralectotype 9 [designated as allotype by Hoogstraal & El Kammah, 1972 : 965] (N 1981) ex 
Kobus kob thomasi, N.E. shore of Lake Edward (approx. 0025'S, 2930'E), Uganda, X. 191 1 , 
S. A. Neave [ex Entomological Research Committee 463a] BM NH 1913.8.12: 7-8. 

HOSTS & DISTRIBUTION. Haemaphysalis aciculifer adults are parasitic on antelopes, 
domestic cattle, sheep, goats, Cape buffalo, hares and occasionally carnivores that prey upon 


antelopes and other game. This tick has been recorded from Cameroon, Uganda, Ethiopia, 
Sudan, Kenya, Tanzania, Zimbabwe, and South Africa. Hoogstraal & El Kammah (1972) 
gave detailed host and distributional data for H. aciculifer. 

Haemaphysalis bancrofti Nuttall & Warburton, 1915: 487, figs 427-430 

Junior synonyms: Haemaphysalis novaeguineae Krijgsman & Ponto, 1932:17; Haemaphysalis 
meraukensis Taylor 1944 : 187; Haemaphysalis krijgsmani Kohls 1948 : 157. 

NUTTALL 2100, 2114, 2115, 2689, 2691. 'Described from specimens derived from 
Queensland, Australia: (N. 2100), 2 d and 5 9 from Macropus dorsalis, Burnett District, 
III. 19 13; (N. 2114), 1 9 and 1 N, found crawling on collector's trousers, at Brigaton Scrub, 
Burnett; (N. 21 15), 1 rf, from Macropus giganteus, Burnett District, III. 19 13; (N. 2689), 2 9, 
with eggs and larvae, the 95 taken from Macropus dorsalis, Eidsvold, XII. 19 13; (N. 2691), 6 
d , 3 9 and 2 N host (?) found at Eidsvold. All collected by Dr T. L. Bancroft, of Eidsvold, after 
whom the species is named. 

Types in Cambridge: (N. 2 100, c? 9 ; N. 269 1 , N; N. 2689, larvae). 

In the following collections (by Nuttall number), Professor Nuttall placed blue type 
labels N2 100, N2689 (ii) and N2691 (ii). Nuttall 2100 was divided, one vial containing 1 d, 
19 the other vial containing 1 rf, 4 9. In the following collections (by Nuttall number), 
Professor Nuttall placed blue cotype labels N2 1 14 (i), (ii), N2 1 1 5, N2689, N269 1 . 

TYPES. Lectotype d (in vial containing 1 cT, 1 9) and 1 d, 5 9 paralectotypes Haemaphysalis 
bancrofti (N2100) [1 9 vulva on slide] ex Macropus dorsalis, Burnett District, Queensland, 
Australia, III. 19 13, Dr T. L. Bancroft; paralectotypes 1 9 (N21 14ii), 1 N (N21 14i) crawling 
on collector's trousers, Brigaton Scrub, Burnett, Queensland, Australia, III. 19 13, Dr T. L. 
Bancroft [leg IV of N21 14(i) is mounted on a slide] 1 d (N21 15) ex Macropus giganteus, 
Burnett District, Queensland, Australia, III. 19 13, Dr T. L. Bancroft; 2 9 (N2689), 223 L 
(including 7 L mounted on slides) (N2689H) ex Macropus dorsalis, Eidsvold, Queensland, 
Australia, XII. 19 13, Dr T. L. Bancroft; 2 N (N269H), 6 c?, 3 9 (N2691ii), no host cited, 
Eidsvold, Queensland, Australia, 1913, Dr T. L. Bancroft. 

REMARKS. Roberts (1963) cited the original collection data and redescribed the d 1 , 9 and 

HOSTS & DISTRIBUTION. Haemaphysalis bancrofti is found in the coastal and subcoastal 
areas of Queensland and northern New South Wales, Australia, and from Kangaroo Island, 
South Australia (Roberts, 1970). It is also present in New Guinea. Hosts include a wide range 
of marsupial mammals as well as most of man's domesticated animals. 

Haemaphysalis calcarata houyi Nuttall & Warburton, 1915: 444 

NUTTALL 2996. 'Described from 7 d , 1 9 and 2 N from Bate, New Cameroon, collected by Dr 
Houy. Types in the Berlin Museum (No. 279), co-types in Cambridge (N2996).' 

TYPES. Lectotype c? and 2 d 1 , 1 9, 1 N paralectotypes Haemaphysalis houyi (N2996), host not 
stated, Bate Cameroun [If the coordinates given by Hoogstraal (1955) are correct, the type 
locality would now be called Bade, Central African Republic], no date stated, Dr Houy. 

REMARKS. Hoogstraal (1954:307) raised the taxon to specific rank, then (1955:365) 
reviewed and illustrated H. houyi and from information supplied to him from J. Bequaert 
gave the type locality at approximately 0640'N, 1 7WE. 

HOSTS & DISTRIBUTION. Haemaphysalis houyi is known primarily from Euxerus erythropus 
in a narrow belt through Kenya, Uganda and Sudan westward to Senegal. 


Haemaphysalis calvus Nuttall & Warburton, 1915: 445, figs 378, 379 

NUTTALL 2120a. 'Described from 3 d and 1 9 (N2120a) taken from buffalo, Sekong River, 
Sandakan, British North Borneo, 19. IV. 19 13, Dr H. F. Conyngham coll.' 

TYPES. Lectotype <5 and 2 rf, 1 9 paralectotypes (N2120a) ex buffalo, Sekong River, 
Sandakan (05 e 50'N, 1 18'05'E), Sabah, Malaysia, 19.IV.1913, Dr H. F. Conyngham [9 vulva 
mounted on a slide]. 

HOSTS & DISTRIBUTION. Haemaphysalis calvus in addition to the original collection has been 
recorded from Cervus unicolor, a tiger, and a bear in Malaysia and Thailand. 

Haemaphysalis campanulata Warburton, 1908 : 5 1 3, figs 5, 6 

NUTTALL 897. Original description 'based on numerous specimens taken from Chinese dogs 
in Mongolia by Major M. L. Hearn, and received by us from Colonel Skinner in 1906.' 

TYPES. Lectotype d and 91 of, 59 paralectotypes Haemaphysalis campanulata (N897) ex 
Chinese dogs, Mongolia, rec'd 14. X.I 906, Major M. L. Hearn. 

The above paralectotype total includes 3 cf , 1 9 which were previously pinned and now are 
retained in a separate vial and 1 <?, 1 9 mounted on a slide. 

REMARKS. Nuttall & Warburton (1915) stated, Types in Cambridge (N897); we have 
presented cotypes (d 1 9) to the British Museum and London School of Tropical Medicine, also 
(cf) to the Museums in Paris, Berlin, Toulouse (Neumann coll.), and Washington, D.C. (U.S. 
Dept. Agric.).' [Those USDA specimens consist of 2 cf, 1 9 and are now housed at the Rocky 
Mountain Laboratories (RML 109832).] 

HOSTS & DISTRIBUTION. This species is known from China, Mongolia, Manchuria, Japan 
and Korea. Hosts include cows, horses, dogs, rats and man. 

Haemaphysalis crassa Warburton, 1908 : 516, fig. 8 
Junior subjective synonym of Haemaphysalis punctata Canestrini & Fanzago, 1878. 

BM(NH) 1908.1.14.103-105. 'Description based on two gorged 9 at the British Museum, in a 
tube labelled Haemaphysalis papuana.' 

TYPES. Lectotype 9 and 19 paralectotype Haemaphysalis crassa BM(NH) 
1908.1 .14.103-105, ex. cattle, Zurnabad (4030'N, 461 5'E), Azerbaijan SSR, USSR. 

REMARKS. One of the five vial labels with these ticks bears the following information: 
''Haemaphysalis papuana Thorell 9, Surnabat Antirinderpest Station, from cattle, Dr E. 
Dschunkovskii.' Nuttall & Warburton (1915:384) spell the locality Surnabad and the 
collector's name Dschunkowsky. 

HOSTS & DISTRIBUTION. Haemaphysalis punctata is a parasite on most of man's domesti- 
cated animals as well as many species of ground nesting birds. This tick is found throughout 
central and southern Europe, northern Africa and into the Asiatic areas of Turkey and the 

Haemaphysalis hoodi Warburton & Nuttall, 1909 : 62, figs 7, 8 
Junior synonym: Haemaphysalis africana Howard, 1909 : 219. 

NUTTALL 424. 'Described from numerous specimens of both sexes sent by Dr P. Hood, from 
Bathurst, Gambia, where it occurred on fowls in May, 1908.' 

TYPES. Lectotype cf and 14 cf, 34 9 paralectotypes Haemaphysalis hoodi (N424) ex fowls, 
Bathurst, now Banjul (1 328'N, 1 639'W), Gambia, V. 1 908, Dr P. Hood. 

REMARKS. Nuttall & Warburton (1915) p. 486 stated 'Types in Cambridge (N424, adults and 
N, N1364 N and L); cotypes (cf 9 from our collection) at Toulouse.' Specimens in collection 


(N1364) mentioned by Nuttall & Warburton (1915) are the types of Haemaphysalis 

HOSTS & DISTRIBUTION. Distributed throughout the Ethiopian faunal region but perhaps 
found more in West rather than East Africa, it is exclusively parasitic on birds with a 
preference for ground dwellers. 

Haemaphysalis hoodi orientalis Nuttall & Warburton, 1915: 486 
Junior synonym: Haemaphysalis zambeziae Santos Dias 1953 : 1 . 

NUTTALL 2847, 2848. 'Described from 6 cf and 3 9 taken from Procavia manningi 
Wroughton, S.W. Shore of Lake Nyasa, British Central Africa, III. 19 10, S. A. Neave coll. 
Types in the Imperial Bureau of Entomology coll., London (Nos. 66a and 69); cotypes in 
Cambridge (N2847 and N2848, 1 cf, 2 9).' 

TYPES. Lectotype cf and 1 cf, 1 9 paralectotypes Haemaphysalis hoodi orientalis (N2847) 
(Imperial Bureau of Entomology 66a) ex Heterohyrax brucei manningi, S.W. Shore, Lake 
Malawi, Malawi, III. 1910, S. A. Neave [9 vulva mounted on a slide]; BM(NH) 1914.6.15: 
1-3; paralectotypes 3 cf, 2 9 (N2848) (Imperial Bureau of Entomology 69), collection data as 
for (N2847) [1 cf, 1 9 hypostome mounted on a slide]. 

REMARKS. Hoogstraal (1956 : 161) elevated H. orientalis to specific rank. 

HOSTS & DISTRIBUTION. Haemaphysalis orientalis is a parasite of hyraxes in Malawi, 
Tanzania, Zambia and Mozambique. 

Haemaphysalis howletti Warburton, 1913 : 123, figs 2, 3 

NUTTALL 1979. 'Described from 1 cf and 1 9 taken by F. M. Hewlett from a hill pony at 
Rawalpindi, India (no date). The types are in the Quick Laboratory, Cambridge (N1979).' 

TYPES. Lectotype cf and 1 9 paralectotype Haemaphysalis howletti ex hill pony, Rawalpindi 
(3336'N, 7304'E), Pakistan, date not stated on vial label or in NuttalFs catalogue, F. M. 
Hewlett. [9 vulva mounted on a slide.] 

REMARKS. Nuttall & Warburton (1915 : 494) stated the collection date 1912. Dhanda (1964) 
redescribed the adults and described the immature stages. Nymphs and larvae were collected 
mostly on rodents and birds. The only adult collected was from a bandicoot. All were from 
the vicinity of Poona, India. 

HOSTS & DISTRIBUTION. Haemaphysalis howletti is known from western India, northern 
Pakistan, and southern Nepal on the above-mentioned hosts. 

Haemaphysalis humerosa Warburton & Nuttall, 1909 : 60, figs 4, 5 

NUTTALL 669. 'Described from 4 cf and 1 9 taken from Perameles macrura, Barrow Island, 
N.W. Australia. Presented by Hon. N. C. Rothschild.' 

TYPES. Lectotype cf and 1 cf, 1 9 paralectotypes Haemaphysalis humerosa (N669) 
(Rothschild 178) ex Isoodon macrourus, Barrow Island (2048'S, 1523'E), Australia. [9 
vulva mounted on a slide.] 

REMARKS. An additional 1 cf of this collection was sent to Dr L. O. Howard, U.S. Department 
of Agriculture, Washington, D.C., 21.V.1909. Nuttall & Warburton (1915) indicated 3 cf , 1 9 
and 1 N present in (N669). Roberts (1963) noted that the tube label states 3 cf 1 9 off 
Perameles macrura, W. Australia. Nuttall has added the note 'one missing from 20. X.I 934'. 

HOSTS & DISTRIBUTION. Haemaphysalis humerosa is collected most frequently from the 
bandicoot and, in addition to Australia, has been found in New Guinea. 


Haemaphy sails inermis aponommoides Warburton, 1913: 128, fig. 8 

NUTTALL 1566. 'Described from numerous 9*5 taken by Colonel F. Raymond F.R.C.V.S., 
from a Himalayan Zebu at Belgachia, Calcutta, III. 19 12, in company with Boophilus 
australis. Types in the Quick Laboratory, Cambridge (Nl 566).' 

TYPES. Lectotype cf and 19 9 paralectotypes Haemaphysalis inermis aponommoides 
(N1566), ex Himalayan Zebu cattle, Belgachia, Calcutta (2232'N, 8822'E), India, III. 19 12, 
Colonel F. Raymond. 

REMARKS. Nuttall & Warburton (1915 : 367) repeated the original description and figures 
and (p. 549) listed 23 9 in the collections of BM(NH), Berlin Museum, and Toulouse. 
Hoogstraal (1961 : 317) redescribed the female and Hoogstraal (1962 : 195) described the <S 
and raised aponommoides to specific rank. The cattle undoubtedly were imported from the 
north to Calcutta, a city far from this tick's distributional range. 

HOSTS & DISTRIBUTION. Haemaphysalis aponommoides is known from Nepal, India, Sikkim 
and China. Adult H. aponommoides primarily parasitize artiodactyls but have been found 
on man, dog, flying squirrel, wildcat and black bear. Immatures have been collected on 
pheasant and 2 shrew and 4 rodent species (Hoogstraal & Mitchell, 1971). 

Haemaphysalis japonica [spelled japonnica] Warburton, 1908 : 512, figs 3, 4 

NUTTALL 1247. 'Description based on numerous specimens in a tube labelled H.flava at the 
British Museum . . . Habitat, Japan.' 

TYPES. Lectotype cf and 1 9 paralectotype Haemaphysalis japonica (N1247) ex Capricornis 
crispus Hondo (3227'N, 13012'E), Kyushu, Japan, received IV. 1911 [i.e., received at 
Cambridge University from Mr A. S. Hirst, BM(NH)], The Duke of Bedford's collector, 6 cf, 
paralectotypes data as above, deposited in the BM(NH) 1 908. 1 1.7.1-11 (part). 

REMARKS. Nuttall & Warburton (1915 : 403) gave the collection data for Warburton's type 

HOSTS & DISTRIBUTION. Haemaphysalis japonica is found on dogs, deer, serow, cattle and 
horses in Japan, Korea, China and Southern Primor, U.S.S.R. 

Haemaphysalis japonica douglasi Nuttall & Warburton, 1915: 403 
Junior subjective synonym of Haemaphysalis japonica Warburton, 1908. 

NUTTALL 1248. 'Described from 4 cf taken from roe-deer, at Ten-an-fu, Shiensi, Northern 
China, 19.V.1909, by Captain H. E. M. Douglas. Types in the British Museum and co-types 
(N. 1248 2 cf) in Cambridge.' 

TYPES. Lectotype cf Haemaphysalis japonica douglasi (N1248) ex Capreolus capreolus, 
Yen-An (3638'N, 10927'E), Shensi, Peoples Republic of China, 19. V.I 909, Captain H. E. 
M. Douglas. 7 cf paralectotypes, data as above, deposited in the BM(NH) 19 1 1 .7.7.28-36. 

HOSTS & DISTRIBUTION. See Haemaphysalis japonica. 

Haemaphysalis koningsbergeri Warburton & Nuttall, 1909 : 65, figs 11,12 

NUTTALL 496. 'Described from 2 cf and 2 9 sent by Dr J. C. Koningsberger of the Zoological 
Museum at Buitenzorg, Java, in 1908. Its host was Felis pardus.' 

TYPES. Lectotype cf and 2 9 paralectotypes Haemaphysalis koningsbergeri (N496) ex 
Pantherapardus, Java, Indonesia, received X. 1908. [9 vulva mounted on a slide]. 

REMARKS. Nuttall & Warburton (1915 : 467) reduced H. koningsbergeri to a synonym of//. 


HOSTS & DISTRIBUTION. Haemaphysalis koningsbergeri is a parasite of small and medium- 
size carnivores and found in Vietnam, southern Thailand, Malaysia, Borneo, and on Java 
and Sumatra in Indonesia. 

Haemaphysalis longipalpis Warburton, 1910: 399, figs 4, 5 
Junior subjective synonym of Haemaphysalis aculeata Lavarra, 1904. 

NUTTALL 1107. 'Described from 10 d 1 and 1 9 taken by Mr C. C. Dobell from Tragulus 
meminna, the Mouse Deer, at Colombo, Ceylon, 3. VIII. 1909.' 

TYPES. Lectotype cf and 7 cf, 1 9 paralectotypes Haemaphysalis longipalpis (Nil 07) ex 
Tragulus meminna, Colombo (0656'N, 7951'E), Sri Lanka, 3.VHI.1909, C. C. Dobell [9 
vulva mounted on a slide.] 

HOSTS & DISTRIBUTION. Haemaphysalis aculeata is an uncommon species. Adults are most 
often collected from the Mouse Deer, immatures have been recorded from Macaca radiata 
and Paradoxurus zeylonensis. This tick has been found in Sri Lanka, India and Indonesia. 

Haemaphysalis mjobergi Warburton, 1926 : 57, fig. 3 

NUTTALL 3560. 'Described from two specimens in a tube labelled "Rissa equina, Borneo", in 
a small collection of ticks sent by Dr E. Mjoberg from Sarawak, 7.IV.24.' 

TYPES. Lectotype c? Haemaphysalis mjoebergi (N3560) ex Cervus unicolor (probably), Mt 
Poi, Sarawak, Malaysia, sent 7. IV. 1924 by Dr. E. Mjoberg. 

REMARKS. Kohls (1961 : 305) noted that 'Rissa' is probably a lapsus for Rusa, a name used 
for the sambar or red deer, Cervus unicolor. He also stated that the second specimen was 
probably returned to the Sarawak Museum but that it could not be located and may have 
been lost during the Japanese occupation. The collecting locality of Mt Poi was found in 
Nuttall's catalogue. 

HOSTS & DISTRIBUTION. Haemaphysalis mjoebergi has only been collected from Cervus 
unicolor (probably) in Kalimantan, Indonesia, and from grass and fallen leaves at Poi, 
southwest Sarawak, Malaysia. 

Haemaphysalis silacea Robinson, 1912 : 478, fig. 1, a-f 

NUTTALL 1629, 2944. 'Described from 4 9 found on oxen allowed to run on "starvation 
camp" from which stock had been excluded for two years, Gonubie Park, East London, 
South Africa.' 

TYPES. Lectotype 9 and 1 9 paralectotype Haemaphysalis silacea (N1629), ex cattle, 
"Starvation Camp", Gonubie Park (3255'S, 2759'E), Republic of South Africa, 1911, 
received from W. F. Cooper [paralectotype 9 vulva mounted on a slide]; paralectotypes 2 9 
(N2944), collection data as for (N1629) except date given as 24.11.191 1, and collector R. J. 

REMARKS. Nuttall & Warburton (1915:418) added the collector's name, Mr R. J. Davys, 
and the deposition of the types Cambridge (N1629, N2944). Hoogstraal (1963) reviewed 
and updated the existing knowledge on H. silacea. 

HOSTS & DISTRIBUTION. Haemaphysalis silacea is known only from South Africa but is no 
longer found in the type locality. Adults are found most often on cattle but have a 
comparatively wide host range including sheep, goat, Kudu, hare, mongoose, bushbuck, etc. 

Haemaphysalis \vellingtoni Nuttall & Warburton, 1908 : 397, figs 9-1 1 

NUTTALL 22 1 . 'Description based on 1 d 1 and 2 9 from fowl, Sarawak, Borneo, collected by Dr 
A.R.Wellington, 1907.' 


TYPES. Lectotype d and 10 9, 2 N paralectotypes Haemaphysalis Wellington! (N221) ex 
fowls, Kuching (01'33'N, 11020'E), Sarawak, 1.1907, Dr A. R. Wellington [1 9 vulva 
mounted on a slide.] 

REMARKS. Nuttall & Warburton (1915:481) stated that 'our description is based on 2 rf, 1 1 9 
and 2 N taken from domestic fowl at Sarawak, Borneo, collected by Dr A. R. Wellington, 

HOSTS & DISTRIBUTION. Haemaphysalis Wellington! ranges from Sri Lanka and India through 
Burma, Thailand, Vietnam, Malaysia, to Borneo and Indonesia. It is a parasite of ground 
feeding birds and, occasionally, small mammals. 

Ixodes ampullaceus Warburton, 1933 : 559, fig. 2 
Junior subjective synonym of Ixodes ugandanus Neumann, 1906. 

NUTTALL 3873. 'Described from 2 9 taken by Mr Rupert L. L. Hart on the "edible rat", 
Cricetomys ganibicus, in Uganda.' 

TYPES. Lectotype 9 Ixodes ampullaceus (N3873) ex Cricetomys gambianus, Kingdom of 
Ankole (0030'S, 3030'E), Uganda, sent 1 1 .III. 1 93 1 , Mr Rupert L. L. Hart. 

REMARKS. Nuttall's catalogue lists only 1 9 as being present in this collection rather than the 
2 9 cited by Warburton in the original collection. The locality of Ankole and the date were 
also taken from his catalogue. 

HOSTS & DISTRIBUTION. Ixodes ugandanus is primarily a rodent parasite especially of cane 
rats in Cameroon, Nigeria, Uganda, Tanzania, Kenya (2 records), Zimbabwe, Mozambique, 
and South Africa. 

Ixodes arvicolae Warburton, 1926 : 56, fig. 2 

NUTTALL 3594, 3595. 'Described from four specimens taken by Miss G. E. Pickford and Mr 
G. E. Hutchinson on water-voles, Arvicola amphibius amphibius, at Quy, near Cambridge, 
in May 1925. One specimen was in company with /. tenuirostris. Type at Cambridge.' 

TYPES. Lectotype 9 Ixodes arvicolae (N3595) ex Arvicola terrestris, Quy (5213'N, 0013'E), 
England, 23.V.1925, Miss G. E. Pickford & Mr G. E. Hutchinson; 2 9 paralectotypes Ixodes 
arvicolae (N3594) from nest of Arvicola terrestris, Quy (52'13'N, 0013'E), England, 
21-23.V.1925, MissG. E. Pickford & Mr G. E. Hutchinson. 

REMARKS. In Nuttall's catalogue, N3594 is labelled 3 9 cotypes [one of which is now missing] 
and N3595 is labelled type. Ixodes arvicolae is possibly a junior synonym of Ixodes 
apronophorus Schulze, 1924. 

HOSTS & DISTRIBUTION. In addition to the type host, /. arvicolae has been collected on the 
coypu, Myocastor coypus. The tick is known from Cambridgeshire and Kent, England. 

Ixodes caledonicus Nuttall, 1910 : 408, figs 1-3 

NUTTALL 961, 1142, 1200. 'Described from 1 9 (N961) on rocks below a dove's nest, 
Fastcastle, Scotland, 6.IX.1909 by Dr J. H. Ashworth (Edinburgh); 1 N and 3 L (Nl 142) 
found on a young pigeon, from a dovecot [sic], at Duniface [sic], Stirlingshire, Scotland, 
18. IV. 19 10 and 1 9 and 4 N (N1200) from the same source, 9. VIII. 19 10, communicated by 
Mr Williams Evans (Edinburgh).' 

TYPES. Lectotype 9 Ixodes caledonicus (N961) rocks below dove's nest, Fastcastle (5556'N, 
0214'W), Scotland 6.IX.1909, Dr J. H. Ashworth [Nuttall & Warburton (1911) state J. F. 
Cormack and J. Waterston as collectors]; paralectotypes 3 L (Nil 42), 2 N (N1200) ex 
Columba livia, Dunipace (560 1 'N, 0355'W), Stirlingshire, Scotland 1 8.1 V. 1 9 1 0, W. Evans. 


REMARKS. Nuttall & Warburton (1911 : 346) indicated that 1 9, N, L were deposited in 
Cambridge and 1 9, N, L were deposited in Edinburgh. The specimens deposited in 
Edinburgh are now missing. 

HOSTS & DISTRIBUTION. Ixodes caledonicus is an avian ectoparasite and ranges from 
Scotland to Norway, Denmark, and Germany. 

Ixodes cavipalpus Nuttall & Warburton, 1908 : 394, figs 1-5 

NUTTALL 245. 'Description based on 2 d 1 and 7 9 from a baboon (Cynocephalus babuin), 
Kansanshi, N.W. Rhodesia, collected by Dr A. Yale Massey.' 

TYPES. Lectotype d 1 and 3 9 paralectotypes (in alcohol) 1 d 1 , 1 9 (each mounted on a slide) 
Ixodes cavipalpus (N245), ex Papio cynocephalus, Kansanshi (1205'S, 2625'E), Zambia, 
sent 22.1.1907 to London School of Hygiene and Tropical Medicine, Dr A. Yale Massey [A 
slide is present containing 1 d 1 , 1 9 spiracular plate.] 

REMARKS. Massey's (1908) mention of the name Ixodes cavipalpus was a nomen nudum. 
Nuttall & Warburton (19 1 1) add the collection date 1. 1907. 

HOSTS & DISTRIBUTION. Ixodes cavipalpus is found on a wide range of hosts including 
primates, carnivores, and lagomorphs but is found most often on the Artiodactyla. This tick 
ranges from Cameroun in West Africa, eastward through Zaire, Uganda, Kenya, and 
southward through Tanzania, Malawi, Zambia, Angola, Zimbabwe, Mozambique into the 
northern Transvaal of South Africa. 

Ixodes dentatus spinipalpis Hadwen & Nuttall in Nuttall, 1916:301, figs 6-8 

NUTTALL 3 1 80, 3181: 'My description is based on (N. 3 1 80) 1 9 found on Lepus americanus, 
1. IX. 19 10, (N. 3181) 3 N found on L. americanus, Sciurus hudsonius douglasii and upon a 
child respectively, besides 4 larvae (17 were originally collected) from Lepus americanus, 
5. III. 19 10, all taken at Mount Lehman, British Columbia, Canada, by Dr S. Hadwen. Types 
in Cambridge.' 

TYPES. Lectotype 9 Ixodes dentatus spinipalpis (N3180) ex Lepus americanus, Mount 
Lehman (4907'N, 12223'W) British Columbia, Canada, 1. IX. 19 10, DrSemour A. Hadwen; 
2 N, 4 L paralectotypes Ixodes dentatus spinipalpis [1 N ex Homo sapiens; 1 N ex Lepus 
americanus; 4 L (1 L mounted on a slide) ex Lepus americanus], Mount Lehman (4907'N, 
12223'W), British Columbia, Canada, 5.III.1910, Dr Semour A. Hadwen. 

REMARKS. The nymph from Tamiasciurus hudsonicus could not be located and is not 
mentioned in Nuttall's catalogue of ticks. Cooley & Kohls (1942 : 1734) raised /. spinipalpis 
to specific rank. 

HOSTS & DISTRIBUTION. Adults and immatures of Ixodes spinipalpis are found on numerous 
species of lagomorphs and rodents. Immatures are also found on birds. The tick is found in 
the Pacific and Rocky Mountain areas of the U.S.A. and in British Columbia and Alberta, 

Ixodes eichhorni Nuttall, 1916 : 295, figs 1, 2 

NUTTALL 2672, 2675. 'Described from a 9 found on the collector's person at Rook Island, 
Friedrich Wilhelmshaven, German New Guinea, IX. 19 13 and several unfed N's found on a 
Kingfisher, Manus, Admiralty Islands, IX-X.1913 by Mr A. F. Eichhorn after whom the 
species is named and to whom I am indebted for the specimens. Considering the part of the 
world they come from and the remarkable agreement in structure between the 9 and N I 
attribute them to one species although they were collected from different hosts. The only 
species hitherto known to possess circular anal grooves is /. rasus Neumann, 1899, from 


Africa, which differs from /. eichhorni in numerous respects, the coxae being unarmed and 
trenchant, etc. The types (N. 2672, 2675) are in Cambridge.' 

TYPES. Lectotype 9 Ixodes eichhorni (N2672) ex Homo sapiens, Rook Island, now known as 
Umboi Island (0536'S, 148WE), Papua New Guinea, IX. 19 13, Mr A. F. Eichhorn; 2 N 
paralectotypes Ixodes eichhorni (N2675), ex Kingfisher, Manaus Island (0205'S, 147WE), 
Admiralty Islands. 

HOSTS & DISTRIBUTION. Ixodes eichhorni is a parasite of birds in Papua New Guinea, New 
Hebrides, Admiralty Islands, and Western Samoa. 

Ixodes fecialis aegrifossus Warburton & Nuttall, 1909 : 250. 
Junior subjective synonym of Ixodes fecialis Warburton & Nuttall, 1909. 

NUTTALL 339. 'Described from three mutilated 95 taken from an Opossum, Tamborine 
Mountain, Logan River, S. E. Queensland, April, 1907, and labelled "Scrub-ticks," sent by 
Professor R. T. Hewlett (London) who received them from the Bacteriological Institute, 

TYPES. Lectotype 9 and 1 9 paralectotype Ixodes fecialis aegrifossus (N339) ex Opossum, 
Tamberine Mountain, Logan River (2743'S, 15318'E), Queensland, Australia, IV. 1907, 
sent by Professor R. T. Hewlett (London) who received them from the Bacteriological 
Institute, Brisbane, Australia. 

HOSTS & DISTRIBUTION. Ixodes fecialis is found on a wide range of both small and large 
marsupials as well as mice, rats, and occasionally man. It is found in Victoria, Tasmania, 
New South Wales, Queensland and Western Australia. 

Ixodes gigas Warburton, 1910: 397, figs 1 , 2 
Junior subjective synonym of Ixodes acutitarsus (Karsch, 1 880). 

NUTTALL 1 104. 'Described from two specimens in the India Museum, Calcutta (no. 5992/10 
and no. ?) taken at Punkabani, Darjiling District, E. Himalayas (no host recorded).' 

TYPES. Lectotype d 1 Ixodes gigas (N1104), host unknown, Punkabani, Darjeeling District 
(2702'N, 8816'E), India; 1 d paralectotype with same data deposited in the Indian Museum, 
Calcutta (59927 10). 

REMARKS. In Nuttall's catalogue of ticks the collecting locality is spelled Punkabari. 

HOSTS & DISTRIBUTION. Ixodes acutitarsus will parasitize most medium to large sized wild 
and domestic animals and often man and has been recorded from India, Nepal, Peoples 
Republic of China (Tibet), Burma, Japan, and Taiwan. 

Ixodes kelloggi Nuttall & Warburton, 1908 : 396, figs 6-8 
Junior subjective synonym of Ixodes brunneus Koch, 1 844. 

NUTTALL 278, 279, 280. 'Description based on 3 9, one from a thrush (from Palo Alto, Cal. 
1895), one from Cyancitta [sic] stelleri frontalis (R. C. Snodgrass, coll. 1895, California), and 
a third from a bird (probably California), sent by Prof. V. L. Kellogg of Stanford University 
in 1907. (In alcohol.)' 

TYPES. Lectotype 9 Ixodes kelloggi (N280) ex Thrush, Palo Alto (37*2 1'N, 12207'W), 
California, U.S.A., 14. XI. 1895; 1 9 [slide mounted capitulum] paralectotype Ixodes kelloggi 
(N279) ex Cyanocitta stelleri, California, U.S.A. 1 5.XII. 1 895, R. E. Snodgrass. 

REMARKS. Exact dates for (N280) and (N279) are taken from Nuttall's catalogue. The 
specimen from a bird, probably California (N278) is missing. 


HOSTS & DISTRIBUTION. Ixodes brunneus is exclusively a parasite of birds usually infesting 
migratory species. With the exception of one record from Venezuela, valid records of /. 
brunneus have only been from the United States. For a listing of the 21 states from which it 
has been collected see Keirans & Clifford (1978 : 54). 

Ixodes kempiNuttall, 1913 : 131 fig. 1 

Junior subjective synonym of Ixodes granulatus Supino, 1897. 

NUTTALL 2066. 'Described from eight partly gorged 95, found on Sciurus erythraceus 
intermedius, at Kobo (400 feet elevation), India, Abor Expedition, by S. W. Kemp, 
29. III. 19 12. (Indian Museum, Calcutta, No. 1247/17.)' 

TYPES. Lectotype 9 and 2 9 paralectotypes Ixodes kempi (N2066) ex Callosciurus erythraeus 
intermedius, Kobo (2747'N, 9523 / E), India (altitude 122 m), 24.111.1912, S. W. Kemp. 

REMARKS. The date of 24. III. 19 12 is in Nuttall's catalogue and on the vial label with the 

HOSTS & DISTRIBUTION. Ixodes granulatus is primarily a rodent parasite in Nepal, India, 
Vietnam, Thailand, Burma, Cambodia, Indonesia, Philippines, Malaysia, Japan, Korea, 
China, and Taiwan. 

Ixodes loricatus spinosus Nuttall, 1 9 1 : 4 1 1 , fig. 4 

[Preoccupied by spinosus Neumann, 1 899] 
Junior subjective synonym of Ixodes luciae Senevet, 1940. 

NUTTALL 647: 'Described from (N647) 3 9*5 taken from a large Opossum, Tabasco de la 
Frontera, Mexico, in the month of May (ex Hon. N. C. Rothschild's collection).' 

TYPES. Lectotype 9 and 1 9 paralectotype Ixodes loricatus spinosus (N647) ex Didelphis sp., 
Frontera (1832'N, 9238'W), Tabasco, Mexico, May, ex Hon. N. C. Rothschild's collection. 

HOSTS & DISTRIBUTION. Adults of Ixodes luciae prefer opposums as hosts but immatures are 
more often found on rodents. The tick is found from southern Mexico to Peru, Bolivia and 
Brazil with one record from Argentina. 

Ixodesoldi Nuttall, 1913 : 135, fig. 3 

NUTTALL 2065. 'Described from 14 slightly engorged 9's found on a bush-cat, at Komatendu, 
Sierra Leone, West Africa, by J. J. Simpson, 13. VIII. 19 12 (Imperial Bureau of Entomology, 
No. 604).' 

TYPES. Lectotype 9 and 5 9 paralectotypes Ixodes oldi (N2065) ex bush cat (probably a West 
African species ofGenetta], Komatendu, Sierra Leone (locality not verified), 13. VIII. 1912, J. 
J. Simpson. 

REMARKS. Nuttall (1913) states 14 9 in the original collection. In his tick catalogue, however, 
he recorded 7 9 in collection (N2065). Arthur (1958:47) and (1965: 138) discusses a 
'holotype 9' deposited in the BM(NH) bearing the above original data. However, as can be 
seen from Nuttall's statement, he did not select a holotype. 

HOSTS & DISTRIBUTION. Ixodes oldi is primarily a parasite of carnivores, especially genets, 
although it is also found on the insectivore, Crocidura sp. It ranges from Sierra Leone in the 
west, eastward to Tanzania and south to the Republic of South Africa. 

Ixodes percavatus rothschildiNutla\\ & Warburton, 191 1 : 221 

NUTTALL 634. 'Described from (N634) 2 9's and 3 N taken from a puffin (ex Rothschild 
Coll., no locality recorded).' 


TYPES. Lectotype 9 and 1 9, 2 N paralectotypes Ixodes rothschildi (N634) (Rothschild 1 59) ex 
Fratercula arctica, no other data. 

REMARKS. Zumpt (1952 : 17) and independently Arthur (1953 : 222) gave specific rank to /. 
rothschildi. Arthur (1963 : 134) referred to 2 9, 3 N in the type series. One nymph is now 

HOSTS & DISTRIBUTION. Ixodes rothschildi is a parasite of marine birds in the coastal areas of 
Great Britain, Ireland, and France. 

Ixodes rid no ides Nuttall, 1913: 1 36, fig. 4 

[Preoccupied by ricinoides De Geer, 1 778] 

Ixodes nuttallianus Schulze, 1930 nom. nov. 

NUTTALL 1401, 1402. 'Described from five partially fed 98; one 9 found on a swamp deer, 
Wen-chwan-hsien, near Si-ho-hsien, China, and four 95 found on musk deer (same locality). 
The specimens, obtained by purchase from Mr T. V. Sherrin, Taxidermist, Hampton, 
Middlesex, are in Cambridge. (N. 1401, 1402).' 

TYPES. Lectotype 9 Ixodes ricinoides (N1401) ex Hydropotes inermis, Wen-Ch'uan 
(4459'N, 8104'E), Sinkiang Province, Peoples Republic of China 1.VII.1911, purchased 
from Mr T. V. Sherrin; 3 9 paralectotypes Ixodes ricinoides (N1402) ex Mochus moschiferus, 
Wen-Ch'uan (4459'N, 8 1 04'E), Sinkiang Province, Peoples Republic of China, 1 .VII. 1911, 
purchased from Mr T. V. Sherrin. 

REMARKS. Collection dates were taken from Nuttall's Catalogue of Ticks. 

HOSTS & DISTRIBUTION. In addition to the members of the Cervidae mentioned above, /. 
nuttallianus has also been collected from the barking deer, Muntiacus muntjak. It has been 
found also on Capricornis sumatraensis, Naemorhedus gpral, dog, goat, cattle and zhum 
[hybrid cow-yak]. Ixodes nuttallianus is distributed from Sinkiang Province, China, through 
Tibet and Nepal. For a detailed discussion of /. nuttallianus see Clifford etal (1971). 

Ixodes ricinus gibbosus Nuttall, 1916 : 300, fig. 5 

NUTTALL 2553. 'Described from (N2553) 5 d 1 , 6 9 found by Mr W. H. J. van Heemstra on 
Capra hircus, Smyrna, Asiatic Turkey, XI. 19 13, in company with Hyalomma aegyptium 
and Haemaphysalis cinnabarina var. punctata. Types in Cambridge.' 

TYPES. Lectotype c? and 4 cf, 6 9 paralectotypes Ixodes ricinus gibbosus (N2553) ex domestic 
goat, Izmir (3825'N, 2709'E), Turkey, XI. 1 9 1 3, Mr W. H. J. van Heemstra. 

REMARKS. Saratsiotis (1970 : 661) raised I. gibbosus to specific rank. 

HOSTS & DISTRIBUTION. Ixodes gibbosus infests, cattle, sheep, goats, horses and camels. 
Ixodes gibbosus replaces /. ricinus in the drier biotopes of Italy, Greece, Yugoslavia, 
Albania, Bulgaria, Cyprus, Turkey, and Israel. 

Ixodes victoriensis Nuttall, 1916, 297, fig. 3 

NUTTALL 3194. 'Described from 2 gorged 9's from wombat (doubtless Phascolomys 
mitchelli), Victoria, Australia, X.I 892, received from Dr Georgina Sweet . . . Types in the 
collection of Dr Sweet (No. 236a), Melbourne, Australia, and (N. 3 194) in Cambridge.' 

TYPES. Lectotype 9 Ixodes victoriensis (N3194) from wombat (doubtless Vombatus ursinus) 
Victoria, Australia, X. 1 892, received from Dr Georgina Sweet. 

REMARKS. The 9 specimen in the collection of Dr Sweet (No. 236a) is now missing. 


HOSTS & DISTRIBUTION. Ixodes victoriensis has been collected only from wombats in 
Victoria, Australia. 

Rhipicentor bicornis Nuttall & Warburton, 1908 : 399, figs 12-16 

NUTTALL 320f. 'Description based on 3 cf and 1 9 collected by Dr Old in North Nyassa, 
British Central Africa. Host not stated (preserved in 5% formalin, then in spirit).' 

TYPES. Lectotype d and 2 c? [1 d in alcohol, 1 d mounted on two slides, i.e., capitulum, body] 
paralectotypes Rhipicentor bicornis (N320f), host unknown, Malawi, III. 1907, Dr J. E. S. 

REMARKS. The 1 9 Rhipicentor bicornis is missing. 

HOSTS & DISTRIBUTION. Rhipicentor bicornis has been collected on cattle, goats, jackal, 
eland, dogs, lions, and several other hosts in Zaire, Ruanda, Burundi, Angola, Zambia, 
Malawi, Zimbabwe, and Namibia. 

Rhipicephalus coriaceus Nuttall & Warburton, 1908 : 402, figs 17-20 
Junior subjective synonym of Rhipicephalus supertritus Neumann, 1907. 

NUTTALL 320e. 'Description based on 2 cf and 6 9 collected in North Nyassa, British Central 
Africa, by Dr Old, 1907; 1 rf (the larger) from Benguella, W. Africa, collected by Dr F. C. 
Wellman, 1907. Hosts not stated. (Preserved in 4% formalin, transferred to spirit.)' 

TYPES. Lectotype 9 and 1 cf, 2 9 [1 9 mounted on a slide] paralectotypes Rhipicephalus 
coriaceus (N320e) host unknown, Malawi, III. 1907, Dr J. E. S. Old; 1 cf paralectotype 
Rhipicephalus coriaceus (N320e), Distrito de Benguela (12'00'S, 15WE), Angola, 1907, Dr 
F. Creighton Wellman. 

REMARKS. The paralectotype 9 in alcohol has had its genital aperture removed and mounted 
on a slide. A photograph of this structure appeared in Feldman-Muhsam (1956 : pi. Ill, fig. 

HOSTS & DISTRIBUTION. Rhipicephalus supertritus adults have been collected on Carnivora- 
Panthera leo, Perissodactyla-D/cmw bicornis and Equus burchelli and many species of 
Artiodactyla. The tick ranges from Sudan, Zaire, Tanzania, Angola, Zambia, and Malawi to 

Rhipicephalus evertsi albigeniculatus Nuttall & Warburton, 1916: 327 
Junior subjective synonym of Rhipicephalus evertsi mimeticus Donitz, 1910. 

NUTTALL 1619, 1650c, 1694b: 'Received by us from the Lower Congo, only differs from the 
above [reddish yellow legs] in having banded legs.' 

TYPES. Lectotype d 1 and 15 cf, 8 9 paralectotypes Rhipicephalus evertsi albigeniculatus 
(N1650c) ex cattle, Mateba Island (0554'S, 1250'E), Zaire, 23-30.IV. 19 12. Received from 
Professor Meuleman (Brussels); 5 d 1 , 3 9 paralectotypes Rhipicephalus evertsi albigeniculatus 
(N1694b) ex cattle Mateba Island (0554'S, 1250'E), Zaire, received 22. VI. 19 12; 2 d 
paralectotypes Rhipicephalus evertsi albigeniculatus (N1619) ex cattle, Kasonga either 
(0528'S, 211 1'E) or (0542'S, 18'48'E), Zaire, Drs Dutton & Todd. 

HOSTS & DISTRIBUTION. Rhipicephalus e. mimeticus parasitizes cattle, horses, sheep, goats, 
and a few of the larger wild hosts such as zebras and antelope. It occurs in drier areas of 
southern Africa and can be found together with R. e. evertsi in the savannahs of northern 
Angola and southwestern Zaire. 


Rhipicephalus longiceps Warburton, 1912: 11, figs 6, 7 

NUTTALL 351, 393. 'Described from 1 8 cfs and 3 95 (No. 351) from "Klipspringer Bok" taken 
by Dr F. C. Wellman in 1907 in the Benguella Hinterland, Angola, long. E.1505' lat. 12 C 44', 
altitude 1360 metres, and 19 cfs and 2 $s. (No. 393) in a mixed collection of ticks taken by the 
same collector in the same district during 1908 but with no host recorded. Types in 

TYPES. Lectotype cf and 17 cf, 2 9 (1 cf abnormal) paralectotypes Rhipicephalus longiceps 
(N351), ex Oreotragus oreotragus, Benguela (1235'S, 1325'E), Angola, received 
3. XII. 1907, sent by Dr F. Creighton Wellman [9 vulva mounted on a slide]; paralectotypes 
15 cf, 2 9 (N393), host not stated, Benguela (1235'S, 1325'E), Angola, 3.V.1908, Dr F. 
Creighton Wellman. 

REMARKS. The one abnormal cf from N35 1 was described and figured by Nuttall (1914: 253). 
Santos Dias (1958) examined the type-material of this species and stated that it consisted of: 
12 cfcf and 1 9 (Types), from a 'Klipspringer bok' (Oreotragus sp.), Benguela, by Dr F. C. 
Wellman, 30/12/1907 No. 351, ex-Nuttall collection; 12 cfcf and 1 9 (Cotypes), by the same 
collector (3/5/1908), at the same locality and host No. 393, ex-Nuttall collection. 

HOSTS & DISTRIBUTION. Rhipicephalus longiceps is a rare tick known only from Angola and 
Southwest Africa (Namibia) and parasitic upon kudu and klipspringer bok. 

Rhipicephalus masseyi Nuttall & Warburton, 1908 : 404, figs 2 1-26 

NUTTALL 246. 'Description based on 31 cf and 21 9 from Bos coffer, Kansanshi, N.W. 
Rhodesia, collected by Dr A. Yale Massey, 1907 (preserved in spirit).' 

TYPES. Lectotype cf and a total of 21 cf, 13 9 paralectotypes [19 cf, 10 9 in alcohol, 1 cf, 2 9, 
chelicerae mounted on slides plus 1 cf , 1 9 in alcohol deposited in the BM(NH) collection 
1906.11.6.11-12] Rhipicephalus masseyi (N246) ex Syncerus coffer, Kansanshi (1205'S, 
2625'E), Zambia, 1907, Dr A. Yale Massey [1 9 vulva mounted on a slide]. 

HOSTS & DISTRIBUTION. Adult Rhipicephalus masseyi have been collected from Atilax 
paludinosus, Phacochoerus aethiopicus, Potamochoerus porcus, Tragelaphus angasi, T. 
scriptus, and Syncerus coffer in Rwanda, Zaire, Tanzania (southern), Angola, Zambia and 

Rhipicephalus neavei Warburton, 1912:7, figs 2, 3 
Junior subjective synonym of Rhipicephalus kochi Donitz, 1905. 

NUTTALL 1414. 'The specimens which appeared most characteristic and were selected as 
types were taken by Mr S. A. Neave from an eland near the mouth of the Tasangazi R., 
Luangwe Valley, N.E. Rhodesia [E.R.C. No. 168]. Types at British Museum and 

TYPES. Lectotype cf and 3 cf, 4 9 paralectotypes Rhipicephalus neavei (N1414) ex 
Taurotragus oryx near mouth of the Lusangazi River (1325'S, 3134'E), Luangwa Valley, 
Eastern Province, Zambia, 1. IX. 19 10, Mr S. A. Neave; 14 cf, 6 9 paralectotypes, collection 
data as above, deposited in the BM(NH) collection (Reg. No. 1 9 1 1 . 1 2.5. 1 6-40). 

REMARKS. Santos Dias (1958 : 478) cited this collection as containing 1 cf, 1 9. 

HOSTS & DISTRIBUTION. Adults of Rhipicephalus kochi are found on medium to large-size 
mammals in central and eastern Africa and as far south as Zambia. 

Rhipicephalus neavei var. punctatus Warburton 1912: 10, figs 4, 5 
NUTTALL 1411. 'Described from 13 cfs and 8 95 from Kudu, near Fort Mlangeni, Central 


Angoniland, Nyasaland (Neave, V.I 9 10, E.R.C. No. 132), 1 9 from Impala aepiciros 
melampur (sic) on N.-W. shore of L. Nyasa (Neave, VII. 19 10, E.R.C. No. 127), and 1 9 from 
reed-buck, Valley of Rukuru R., N. Nyasaland (Neave, 26.VI.1910, E.R.C. No. 
1 58) ... Types at British Museum and Cambridge.' 

TYPES. Lectotype cf and 1 cf, 1 9 paralectotypes Rhipicephalus neavei punctatus (N141 1) ex 
Tragelaphus strepsiceros, near Fort Mlangeni (1441'S, 3432'E), Ncheu District, Malawi, 
V.I 9 10, S. A. Neave; 11 cf, 4 9 paralectotypes, collection data as above, deposited in the 
BM(NH) collection (Reg. No. 191 

REMARKS. Santos Dias (195 1 : 373) raised Rhipicephalus punctatus to specific status. 

HOSTS & DISTRIBUTION. Host predilections and distributional data for Rhipicephalus 
punctatus in Africa are very poorly known because so few collections of this species have 
been accurately identified. 

Rhipicephalus sculptus Warburton, 1912: 1 3, figs 8, 9 

NUTTALL 1409, 1448. 'Described from 11 cfs and 5 95 (E.R.C. No. 230a) [This is Nuttall 
1448] from roan antelope, Mpalali R., Marimba, Nyasaland (Old, 1.1911) . . . 1 cf (E.R.C. 
No. 227a) [This collection is now missing] from the same locality and host . . . 3 cf and 1 9 
(No. 115b) [This is Nuttall 1409] from zebra, S. Rukura Valley, N. Nyasaland (Neave, 
VI. 19 10) . . . Types at the British Museum and Cambridge.' 

TYPES. Lectotype cf and 1 9 paralectotype Rhipicephalus sculptus (N1409), ex Equus 
burchelli, locality possibly South Rukuru River (1044'S, 3414'E), Malawi, VI. 19 10, S. A. 
Neave; paralectotypes 2 cf, data as above, deposited in the BM(NH) collection. Reg. no. 
191; paralectotypes 3 cf, 2 9 (N1448) ex Hippotragus equinus, locality possibly 
Mpala River, now known as Mipala River (1555'S, 3516'E), Malawi, 1.1.191 1, DrJ. E. S. 
Old [9 vulva mounted on a slide]. 

REMARKS. Santos Dias (1958 : 479) indicated 1 cf 1 9 present in N1409 and 3 cf 2 9 present in 
N1448. Although he did not mention the latter Nuttall number, he listed all relevant 
collection data. 

HOSTS & DISTRIBUTION. Rhipicephalus sculptus is a rare species known only from the zebra 
and a few members of the Artiodactyla in eastern and southern Africa. 

Rhipicephalus simpsoni Nuttall, 1910:413, figs 6, 7 

NUTTALL 1214. 'Described from 5 cf's and 11 9*5 found on a large rodent, Oshogbo, S. 
Nigeria, W. Africa, 28.11.1910, J. J. Simpson (N1214). (41b, Entomological Research 
Committee for Tropical Africa).' 

TYPES. Lectotype cf and 1 cf, 1 9 paralectotypes Rhipicephalus simpsoni (N1214), ex 
Thryonomys sp. (probably), Oshogbo (0746'N, 0434'E), Nigeria, 28.11.1910, J. J. Simpson 
[9 vulva mounted on a slide]. Paralectotypes 1 cf, 4 9 Rhipicephalus simpsoni, data as above, 
deposited in the BM(NH) collection Reg. no. 191 

REMARKS. Santos Dias (1958) indicated 2 cf, 1 9 R. simpsoni present in Nuttall 1214 when he 
examined the Nuttall collection in 1955. 

HOSTS & DISTRIBUTION. Rhipicephalus simpsoni is a specific parasite of cane rats, 
Thryonomys spp. and is widely distributed in the Ethiopian faunal region. 


We are grateful to Mr K. H. Hyatt, Arachnida Section, Zoology Department, British 
Museum (Natural History) for his cooperation during this study. Mr Donald Macfarlane, 


Commonwealth Institute of Entomology, generously gave of his time and provided us with 
his expertise in matters nomenclatural. We also thank the following individuals in assisting 
in locating or confirming the presence or absence of types of species described by Nuttall et 
al. and now under their care: Dr S. K. Gupta, Zoological Survey of India, Calcutta; Professor 
Ph. Dorchies, Ecole Nationale Veterinaire, Toulouse; Dr M. Moritz, Museum fur 
Naturkunde, Berlin; Dr J. A. Campbell, Department of Zoology, University of Edinburgh; 
Dr A. Neboiss, Department of Entomology, National Museurn of Victoria; and Dr G. 
Pretzmann, Naturhistorisches Museum, Vienna. This work is part of a research project on 
Professor G. H. F. Nuttall and the Nuttall Tick Catalogue conducted while the senior author 
was on a Visiting Scientist assignment to the British Museum (Natural History) 1977-1978 
from Department of Health and Human Services, Public Health Service, National Institutes 
of Health, National Institute of Allergy and Infectious Diseases, Epidemiology Branch, 
Rocky Mountain Laboratories, Hamilton, Montana 59840, U.S.A. 


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Saratsiotis, A. 1970. Etude morphologique et observations biologiques sur Ixodes gibbosus Nuttall, 

\9\6.Annls Parasit. hum.comp.45 : 661-675. 
Schulze, P. 1924. Ixodes apronophorus n. sp. eine neue deutsche Zecke von Arvicola amphibius L. 

Zool. Am. 59: 28 ; 1-284. 

1930. Uber linige Verwandte von Ixodes ricinus L. aus Ostaisen. Zool. Am. 90 : 294-303. 

1941. Neues Uber brasilianische Amblyommen. Zool. Anz. 134 : 93-104. 

Senevet, G. 1940. Quelques Ixodides de la Guyane franchise. Especes nouvelles d 1 'Ixodes et 

$ Amblyomma. VI Congr. Int. Ent. (Madrid, Sept. 6-12, 1935) 2 : 891-898. 

Supino, F. 1897. Nuovi Ixodes della Birmania (Nota preventiva). Alii Soc. ven-trent. Sci. 3 : 230-238. 
Taylor, F. H. 1944. Arachnid notes. Proc. Linn. Soc. N.S. W. 69 : 185-187. 
Trapido, H. 1965. Notes on critical Asian Haemaphysalis specimens in European museum collections 

with designations of lectotypes and a neotype. Proc. ent. Soc. Wash. 67 : 1 52-165. 
Warburton, C. 1908. On some new and obscure species of the genus Haemaphysalis of the Ixodidae. 

Proc. Camb. phil. Soc. math. phys. Sci. 14 : 508-5 19. 
1910. On two collections of Indian ticks. Parasitology, Cambridge^ : 395-407. 

1912. Notes on the genus Rhipicephalus, with the description of new species, and the 

consideration of some species hitherto described. Parasitology, Cambridge 5 : 1-20. 

1913. On four new species and two new varieties of the ixodid genus Haemaphysalis. 

Parasitology, Cambridge 6 : 121-130. 

1918. Notes on Ticks. Being descriptions of two new species of Ornithodorus and of the hitherto 

unknown female ofHyalomma monstrosum. Parasitology, Cambridge 10 : 284-287. 

1926. On three new species of ticks (Arachnida, Ixodoidea), Ornithodoros gurneyi, Ixodes 

arvicolaeand Haemaphysalis mjobergi. Parasitology, Cambridge 18 : 55-58. 

1927. On five new species of ticks (Arachnida Ixodoidea), Ornithodorus nattereri, Ixodes theodori, 

Haemaphysalis toxopei, Amblyomma robinsoni and A. dammermani, with a note on the ornate 
nymph of A. latum. Parasitology, Cambridge 19 : 405-410. 



1933. On five new species of ticks (Arachnida Ixodoidea). Ixodes petauristae, I. ampullaceus, 
Dermacentor imitans, Amblyomma laticaudae and Aponomma draconis, with notes on three 
previously described species, Ornithodoros franchinii Tonelli-Rondelli, Haemaphysalis cooleyi 
Bedford and Rhipicephalus maculatus Neumann. Parasitology, Cambridge 24 : 558-568. 

& Nuttall, G. H. F. 1909. On new species of Ixodidae, with a note on abnormalities observed in 

ticks. Parasitology, Cambridge 2 : 57-70. 
Zumpt, F. 1952. The ticks of sea birds. Aust. Nat. Antarctic Res. Rep. Series B\ : 12-20. 

Manuscript accepted for publication 10 December 1980. 

Appendix I 

Holotypes and previously selected lectotypes of tick species described by Nuttall, Warburton, Cooper, 

and Robinson 



Number Original Name 

Senior Subjective 

Type Status 



Amblyomma annandalei 
1106 Warburton, 1910:403, 

fig. 8 
*None Amblyomma crassum 

Robinson, 1926: 177, 

fig. 83 
36 1 5 Amblyomma dammermani 

Warburton, 1927:409, 

pi. XXVII, fig. 3 
2940 Amblyomma longirostrum 

Cooper & Robinson, 

1908: 457, figs 1-5 
565 Amblyomma quasicyprium 

Robinson, 1926:237, 

fig. 117 
2943 Aponomma quadratum 

Cooper & Robinson, 

1908: 468, figs 22-24 
2942 Aponomma simplex 

Cooper & Robinson, 

1908: 466, figs 18-21 
5 1 5a Haemaphysalis bispinosa 


Warburton & Nuttall, 
1909: 69, fig. 16 

1108 Haemaphysalis cuspidata 

Warburton, 1910:401, 
figs 6, 7 

693a Haemaphysalis dentipalpis 
Warburton & Nuttall, 
1909: 67, fig. 14 

635 Haemaphysalis doenitzi 

Warburton & Nuttall, 
1909: 64, figs 9, 10 

A. supinoi Neumann, 1905 

A. cyprium Neumann, 1899 

A. humerale C. L. Koch 

A. cyprium Neumann, 1 899 

A. varanensis Supino, 1897 

A.fimbriatum C. L. Koch, 

H. asiaticus (Supino, 

Holotype 9 BM(NH) 
Nuttall Coll. 

Holotype 9 BM(NH) 
reg.No. 1926.7.27.1 

Holotype 9 BM(NH) 
Nuttall Coll. 

Holotype 9 
Depository unknown 

Holotype 9 
Depository unknown 

Holotype rf BM(NH) 
Nuttall Coll. 

Holotype <? BM(NH) 
Nuttall Coll. 

Lectotype 9 
Hoogstraal & Trapido 

BM(NH) Nuttall 

Lectotype rf Trapido 
Nuttall Coll. 
Lectotype cf 
Hoogstraal & Trapido, 
Nuttall Coll. 
Lectotype 9 
Hoogstraal & Wassef 
Nuttall Coll. 



Appendix I continued 



Number Original Name 

Senior Subjective 

Type Status 



1997 Haemaphysalis kinneari 

Warburton, 1913: 127, 

fig. 6 

760, 76 1 , Haemaphysalis 
762 montgomeryi 

Nuttall, 1912:57, 

figs 7, 8 
540c Haemaphysalis proximo 

Warburton & Nuttall, 

1909: 61, fig. 6 
1 647 Haemaphysalis spiniceps 

Warburton & Nuttall, 

1909: 68, fig. 15 
3614 Haemaphysalis toxopei 

Warburton, 1927:407, 

fig. 3 

1 648 Haemaphysalis vidua 

Warburton & Nuttall, 

1909: 66, fig. 13 
1400 Haemaphysalis warburtoni - 

Nuttall, 1912:55, 

figs 5,6 

1219 Hyalomma monstrosum 

Nuttall & Warburton, 

1908: 4 14, figs 4 1^5 
*None Ixodes daveyi v 

Nuttall, 1913: 133, fig. 2 
650 Ixodes fee ialis 

Warburton & Nuttall, 

1909: 58, figs 1,2 
*None Ixodes nairobiensis 

Nuttall, 1916:299, 

fig. 4 
3872 Ixodes petauristae 

Warburton, 1933 : 558, 

fig. 1 
3581 Ixodes theodori 

Warburton, 1927:405, 

fig. 2 
1 1 03 Rhipicephalus breviceps 

Warburton, 1910:398, 

fig. 3 

H. cornigera Neumann, 1897 

H. cornigera Neumann, 1897 

/. acuminatus Neumann, 

R. sanguineus (Latreille, 

Holotype 9 BM(NH) 
Nuttall Coll. 

Lectotype <t (N760) 
Hoogstraal et al. 
Nuttall Coll. 
Holotype 9 BM(NH) 
Nuttall Coll. 

Holotype cf BM(NH) 
Nuttall Coll. 

Lectotype cf 

Hoogstraal (1964) 

BM(NH) Nuttall 


Holotype cf BM(NH) 

Nuttall Coll. 

Lectotype cf 
Hoogstraal (1966) 
BM(NH) Nuttall 

Holotype cf 
Depository unknown 

Holotype 9 BM(NH) 
reg. No. 1913.8.12.6 
Holotype 9 BM(NH) 
Nuttall Coll. 

Holotype 9 BM(NH) 
reg. No. 1915.10.14.1 

Holotype 9 BM(NH) 
Nuttall Coll. 

Holotype 9 BM(NH) 
Nuttall Coll. 

Holotype 9 BM(NH) 
Nuttall Coll. 

Specimens donated to the British Museum (Natural History) by Nuttall et al. or described from material in the 
Museum collections. 

Appendix II 


Types of tick species described by Nuttall, Warburton, Cooper, and Robinson which are not contained 
in the collections of the British Museum (Natural History) 





Cooper & Robinson, 1908 
Aponomma gervaisi lucasi 

Warburton, 1910 
Haemaphysalis aborensis 

Warburton, 1913 
Haemaphysalis cornigera anomala 

Warburton, 1913 
Haemaphysalis leachi indica 

Warburton, 1910 
Haemaphysalis turturis 

Nuttall & Warburton, 1915 
Ixodes neumanni 

Nuttall & Warburton, 1911 
Ornithodoros asperus 

Warburton, 1918 
Ornithodoros nattereri 

Warburton, 1927 
Rhipicentor nuttalli 

Cooper & Robinson, 1908 
Rhipicephalus phthirioides 

Cooper & Robinson, 1907 

Unknown 'Description based on 3 9's.' 

ZSI, Calcutta 

ZSI, Calcutta 

ZSI, Calcutta 

ZSI, Calcutta 

MNHU, Berlin - 

ENV, Toulouse - 

MNHU, Berlin - 

NM, Wien 

Unknown 'Description based on a single d 1 . 1 

Unknown 'Description based on two dried 

ticks ... a male and a female . . 

ZSI, Calcutta = Zoological Survey of India, Calcutta (formerly Indian Museum, Calcutta). 
MNHU, Berlin = Museum fur Naturkunde der Humboldt-Universitat, Berlin, D.D.R. 
ENV, Toulouse = L'Ecole Nationale Veterinaire de Toulouse, France. 
NM, Wien = Naturhistorisches Museum, Wien, Austria. 

A revision of the spider genus Hispo (Araneae : 

F. R. Wanless 

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


Unlike most genera of the spider family Salticidae, Hispo Simon, 1886 can be easily 
recognized by the characteristic form of the carapace and the presence of a constriction 
behind the posterior median eyes. The genus is represented in both the Oriental and 
Ethiopian regions and at present includes 1 1 known species. Several of these were originally 
described in the genera Astaenochestes Simon, 1900 or Pseudomarengo Caporiacco, 1947. 
The later genus was revised by Roewer (1965), however, they are here considered to be 
synonymous with Hispo. 

In this paper, the species are divided into two species groups which may be readily 
separated by the presence or absence of a fovea. The species included in both groups are 
morphologically similar and in most cases they can be distinguished from one another by the 
structure of the genitalia, particularly the form of the palpal tibial apophyses in males. The 
presence or absence of white hairs on the clypeus and the occurrence of furrows or striae on 
the anterior surface of the chelicerae provide useful characters for separating otherwise 
similar species. The degree of variation cannot be adequatey demonstrated for most species 
as the genus is not well represented in museum collections. The biology is virtually 
unknown, some species live in woodland or forest litter and could be considered ant-like, but 
there is no behavioural evidence to suggest a mimetic relationship with ants. 

The measurements were made in the manner described by Wanless (1978a). 

Genus HISPO Simon 
Hispo Simon, 1886 : 393. Type species Hispo cingulata Simon, by monotypy. Simon, 1901 : 449, 451, 

452; 1903a : 1050. Petrunkevitch, 1928 : 186. Roewer, 1954 : 985. Bonnet, 1957 : 2219. Wanless 

Astaenorchestes Simon, 1900:397. Type species Astaenorchestes frenatus Simon, by original 

designation and monotypy. Simon, 1901 : 450-452. Petrunkevitch, 1928 : 185. Roewer, 1954 : 985. 

Bonnet, 1955 : 766. Syn. n. 
Pseudomarengo Caporiacco, 1947 : 228. Type species Pseudomarengo inermis Caporiacco, by original 

designation. Roewer, 1954 : 951; 1965 : 32. Syn. n. 

Although Roewer (1954) and Bonnet (1957) give the year 1885 as the publication date for 
the genus Hispo, the date of publication given on the title page is 'avril 1886'. The 
confusion arose because the part concerned was issued for the year 1885. 

DEFINITION. Small to large spiders ranging from about 3*5 to 8 mm in length. Sexual 
dimorphism not marked. Distinctive colour markings sometimes present; not hirsute. 
Carapace: very low to moderately high; constricted behind posterior median eyes and 
usually with two shallow depressions behind anterior medians; fovea present or absent; 
microsculpturing variable, cuticle sometimes iridescent. Eyes: anteriors contiguous, apices 
level or slightly procurved; posterior median eyes relatively small, closer to anterior laterals 
than to posterior laterals; posterior row usually narrower than anterior row with a space 
between the posterior lateral eyes and the lateral margins of the carapace; quadrangle length 
between 39 and 50 per cent of carapace length. Clypeus: very low to low. Chelicerae: small to 
medium, more or less vertical, anterior surface irregularly strigose or smooth, sometimes 

Bull. Br. Mus. nat. Hist. (Zool.)41(4): 179-198 Issued 26 November 1981 


180 F. R. WANLESS 

with a 'scooped-out' furrow on each chelicera; promargin with 4-6 contiguous teeth, 
retromarginal teeth similar, but minute more numerous (4-10) and forming a serrated ridge. 
Maxillae: parallel to subparallel, blades more or less rounded. Labium: tongue-shaped, 
about as long as broad or broader than long. Sternum: elongate scutiform, sometimes 
attenuate anteriorly. Pedicel: short. Abdomen: elongate; scuta absent; spinnerets subequal in 
length, anteriors and posteriors unequally robust, medians slender; colulus apparently 
lacking; traecheal system (Fig. 5C) (one species examined), arising from transverse slit 
immediately in front of anterior spinnerets, apparently simply branched and limited to 
abdomen. Legs: moderately long and slender, I and sometimes II enlarged or elongate; 
fringes usually lacking, spines weak to moderately strong, usually more numerous on 
posterior legs; claws generally pectinate (difficult to see in most species), tufts present, 
usually a stiff curved setae lying between claws on legs I-II; scopulae rarely present. Female 
palp: moderately long and slender. Male palp (Fig. 1A, B): with branched tibial apophysis, 
occasionally with small ventral apophysis; cymbium sometimes depressed and sclerotized 
basally; embolus (e) usually robust, long and curved originating centrally or subapically; 
conductor (c) a modified part of the embolic shaft, rarely present (Fig. 1 1 A, B); tegulum (t) 
pleated, and folded distally, the resulting lobe, when present, resembling an apophysis (1) 
(Fig. 7F); median apophysis (m) small, slender with a distal hook, often difficult to see in 
lateral view; subtegulum (st) not pleated, seminal reservoir dark, usually indistinct. Epigyne 
Fig. 5G, J): opening paired, indistinct with sclerotized margins; introductory ducts (0 paired, 
proximally translucent with glandular 'appendixes' (c), distally dark and coiled around the 
tube connecting the primary and secondary spermathecae; spermathecae dumb-bell-shaped, 
primary spermathecae (p) small with leaf-like fertilization ducts; secondary spermathecae (s) 
small to large and globular, on distal margin a lobe-like structure (r) which resembles the 
stretcher found in some linyphiid spiders. 

REMARKS. The epigyne structure of Hispo bipartita Simon clearly differs from that 
described above, but the vulva has not been examined as only one adult female is available 
for study. In any event the generic placement of//, bipartita will have to be reviewed when 
the related genera Massagris Simon, and Tomocyrba Simon are revised. 

AFFINITIES. Hispo is closly related to the genera Massagris and Tomocyrba, both of which 
have foveae and a constriction behind the posterior median eyes which are positioned much 
closer to the anterior laterals than to the posterior laterals. The male palps of Hispo although 
distinctive, show morphological similarities to these genera in the form of the tegulum and 
median apophysis. It is possible that the bipartita-group of Hispo may have to be transferred 
into Massagris as the genitalia would appear to be rather similar. 

It will also be necessary to re-examine some of the amber salticids described by 
Petrunkevitch (1942, 1958). The fossil genus Gorgopsina Petrunkevitch, is evidently close to 
Tomocyrba and the type species of Gorgopsina, G.frenata (Koch & Berendt) may well prove 
to be synonymous with recent Tomocyrba. 

Wanless (19786) suggested that the simple tracheal system found in species of Portia 
Karsch indicated a relationship with lyssomanid spiders. However, as Hispo does not appear 
to have any close affinities with known lyssomanid spiders, while possessing a simple 
tracheal system (Fig. 5C), this proposition will need to be reconsidered when more genera 
have been examined. 

DIAGNOSIS. The cingulata-group of species of Hispo can be easily distinguished from the 
genera Massagris and Tomocyrba by the absence of a fovea. The bipartita-group can be 
distinguished by the very low carapace. 

BIOGEOGRAPHY. The present distribution of Hispo can suggest historically different patterns 
of dispersal depending in which period of geological time the genus evolved and the present 
species came into being. As most species have been described from Madagascar it seems 
plausable to suggest that Hispo evolved in the Malagasy region. If this happened before the 
disruption of Gondwanaland then vicariance may be responsible for the occurrence of only 



Fig. 1 Hispo striolata Simon, rf, expanded palp: A, lateral view; B, ventral view. 

one species in Africa (H. inermis (Caporiacco)) and two species in the Seychelles (H. 
alboclypea Wanless and H. striolata Simon). It is worth noting that although the Seychellois 
species are closely related to each other, H. alboclypea is evidently closer to H. inermis, 
while H. striolata, is closer to H. pullata sp. n. from Madagascar. These four species 
comprise the striolata-subgroup which is hopefully of monophyletic origin. 

If on the other hand these species reached Africa and the Seychelles after the disruption of 
Gondwanaland we should not postulate a relict distribution but rather a recent one which 
may have occurred as a result of aerial dispersal, i.e. ballooning or some other mechanism 
such as rafting. Platnick (1976) has suggested that vicariance is responsible for most inter- 
continental distribution patterns in spiders and that the role of ballooning, at least for 
medium-sized and large ground-dwelling species has been greatly exaggerated. Some 
salticids are known to balloon as adults, and no doubt, ballooning plays an important role in 
their distribution particularly at a regional level. Howevever, Platnick is probably correct in 
his basic hypothesis and we may be sure that some distribution patterns in the Salticidae are 
the result of vicariance. 

If vicariance is responsible for the present distribution patterns in Hispo then species can 
be expected to occur in the Comoro Islands and India. Unfortunately the occurrence of H. 
bipartita in India and Sri Lanka does not fulfil the prediction for as mentioned above, this 
species may have to be transferred into Massagris, and its distribution reconsidered when 
that genus is revised. 

List of species in the genus Hispo Simon, 1886 

Hispo alboclypea Wanless (1981) 

H. alboguttata Simon, 1903 

H. bipartita Simon, 1903 

H. cingulata Simon, 1886 

H.frenata (Simon, 1900) 

H. inermis (Caporiacco, 1947) 

H. macfarlanei sp. n. 

H. pullata sp. n. 

H. striolata Simon, 1897 

H. sulcata sp. n. 

H. tennis sp. n. 

182 F. R. WANLESS 

Key to species of Hispo 

1 Fovea lacking; sternum truncate anteriorly (Figs 2C, 10D) . . . cingulata -group 2 

- Fovea present; sternum attenuate anteriorly (Fig. 12B). . . . bipartita-group 15 

2 Males 3 

Females 11 

3 Palpal conductor present (Fig. 1 1 A, B) (Madagascar) . . . cingulata Simon (p. 193) 

- Palpal conductor absent 4 

4 Tibial apophysis with backward pointed spur (Fig. 9J, K) (Madagascar) 
macfarlanei sp. n. (p. 191) 

- Tibial apophysis without backward pointed spur 5 

5 Clypeus clothed in white hairs 6 

Clypeus not white haired 7 

6 Embolus large and robust (Fig. 2 E) (Madagascar) pullatasp. n. (p. 183) 

- Embolus small and slender (Fig. 4A) (Seychelles) .... alboclypea Wanless(p. 185) 

7 Embolus very robust (Fig. 7F); dorsal prong of tibial apophysis with small prolateral spur 

(Fig. 7G) (Madagascar) tenuis sp. n. (p. 1 89) 

Embolus otherwise; dorsal prong of tibial apophysis without a prolateral spur ... 8 

8 Anterior surface of chelicera with a distinct furrow (Fig. 6E,8F) (Madagascar) ... 9 
Furrow lacking; anterior surface of chelicerae irregularly strigose .10 

9 Lateral prong of tibial apophysis with a small spur (Fig. 8B, C) . . sulcata sp. n. (p. 1 90) 

- Lateral prong of tibial apophysis without a spur (Fig. 6C) . . . frenata (Simon) (p. 188) 

10 Legs I swollen (Fig. 5E); embolus slender (Fig. 5H) (Africa) . inermis (Caporiacco) (p. 1 86) 
Legs I elongate (Fig. 3C); embolus robust (Fig. 3 A) (Seychelles) striolata Simon (p. 1 84) 

11 Clypeus clothed in white hairs -12 

Clypeus not white haired 14 

1 2 Epigynal opening with median scape-like projection (Fig. 9C) (Madagascar) 
macfarlanei sp. n. (p. 191) 

- Epigynal opening without median scape-like projection .13 

13 Epigynal openings subcontiguous, transversely elongate, lateral margins extending beyond 

secondary spermathecae (Figs 10F, 1 1C, D) (Madagascar) . . cingulata Simon (p. 193) 

Epigynal openings separate, obliquely elongate, lateral margins not extending beyond secondary 

spermathecae (Fig. 4B) (Seychelles) alboclypea Wanless(p. 185) 

14 Carapace with depressions behind anterior median eyes; epigynal openings subcontiguous 

(Fig. 3D) (Seychelles) striolata Simon (p. 1 84) 

Carapace without depressions behind anterior median eyes; epigynal openings separate 
(Fig. 5 F, I) (Africa) inermis (Caporiacco) (p. 186) 

15 Abdomen with discoidal white and black markings (Fig. 13 A) [species only known from a 

subadult 9] (Sumatra) alboguttataSimon(p. 196) 

Discoidal markings lacking; epigyne with large openings and tangled fertilization ducts 
near posterior margin (Fig. 1 2F) (India, Sri Lanka) .... bipartita Simon (p. 1 95) 

The cingulata-group 

The cingulata-group is comprised of nine known species, one from Africa, two from the 
Seychelles and six from Madagascar, which can be readily distinguished from species of the 
bipartita-group by the absence of a fovea. 

Within this group two subgroups based on the structure of the male palp can be 
recognized. In the striolata-subgroup (H. striolata Simon, H. alboclypea Wanless, H. inermis 
(Caporiacco) and H. pullatta sp. n.), the dorsal prong of the palpal tibial apophysis is 
relatively small and unmodified. In the cingulata-subgroup (H. cingulata Simon, H.frenata 
(Simon), H. tenuis sp. n., H. sulcata sp. n., and H. macfarlanei sp. n.), the tibial apophysis is 
relatively large and often provided with spurs. H. cingulata is further distinguished by having 
a modified embolic shaft. 



Fig. 2 


Hispo pullata sp. n., holotype d: A, dorsal view; B, carapace, lateral view; C, sternum; D, 
palp, lateral view; E, palp, ventral view. 

H'spo pullata sp. n. 
(Fig. 2A-E) 

DIAGNOSIS. Hispo pullata is closely related to H. striolata Simon, but can be separated by the 
presence of a fringe of white hairs on the clypeus. 

FEMALE. Unknown. 

MALE HOLOTYPE. Carapace (Fig. 2A, B): finely rugose to finely punctate-reticulate within 
quadrangle; brownish orange, weakly iridescent. Eyes: with black surrounds; anteriors 
contiguous, apices slightly procurved, sparsely fringed in light brown hairs. Clypeus: fringed 
in white hairs. Chelicerae: with fine irregular striae; brownish orange, weakly iridescent; 
teeth not examined. Maxillae and labium: brownish orange. Sternum (Fig. 2C): light orange, 
shiny. Abdomen: yellow-brown lightly tinged black with indistinct whitish markings clothed 
in white hairs; anal tubercle whitish. Legs: legs I heaviest, dark brown; other legs light brown 
to yellow-brown. Spination of legs I: metatarsi with 1 proventral spine, tibiae with 1 distal 
proventral, femora with 3 dorsal and 1 distal prolateral spine; other leg spines weak, but 
more numerous especially on legs III-IV. Palp (Fig. 2D, E): similar to that of//, striolata, but 
the embolus is slightly more robust and the ventral apophysis of the tibia a little more 

Dimensions (mm): total length 3*84; carapace length 1*76, breadth 1-16, height 0*72; 
abdomen length 2-12; eyes, anterior row 1-12, middle row 0'88, posterior row 0'92; 
quadrangle length 0'86. Ratios: AM : AL : PM : PL :: 10-5 : 4-5 : 1 : 4, AL-PM-PL :: 5-9. 

184 F. R. WANLESS 

VARIATION, d total length 3'3 to 4- 1 mm, carapace length 1 -56 1 -88 mm (4 specimens). 
DISTRIBUTION. Madagascar. 

MATERIAL EXAMINED. Madagascar: Beanana, holotype cf, ii. 1970 (A. Lambillon, MT. 
142.600) (MRAC, Tervuren). Antongil, paratypes 3 cfd 1 , (A. Mocqueries) (MNHN, Paris. 

Hispo striolata Simon 

Hispo striolata Simon, 1897 : 387, <?, 9. Lectotype cf, Seychelles (MNHN, Paris, 1 1409) [Examined]. 
Simon, 1901 : 450, 451. Hirst, 1911 : 382. Roewer, 1954:985. Bonnet, 1957:2220. Proszynski, 
1971:417. Wanless, (1981) [Lectotype designated]. 

DIAGNOSIS. Hispo striolata is closely allied to H. pullata sp. n., but can be distinguished by 
the absence of a fringe of white hairs on the clypeus. The female of//, pullata is unknown. 

REMARKS. This species has recently been redescribed in a faunistic study on Salticidae from 

Fig. 3. Hispo striolata Simon, cf: A, palp, ventral view; B, palp, lateral view; C, leg I. 9: D, 
epigyne; E, vulva, ventral view; F, vulva, dorsal view. 



Fig. 4 Hispo alboclypea Wanless, cf; A. palp, ventral view; C, palp, lateral view. 9: B, epigyne; D, 

vulva, ventral view; E, vulva, dorsal view. 

the Seychelle Islands (Wanless, 1981) but, for completeness, I have provided figures of the 

DISTRIBUTION. Seychelle Islands: Mahe, Silhouette, Praslin. 

MATERIAL EXAMINED. 9 dtf, 3 99, including the lectotype; for data see Wanless (1981). 

Hispo alboclypea Wanless 
(Fig. 4A-E) 

Hispo alboclypea Wanless (1981), cf, 9. Holotype cf, allotype 9, Seychelles, Silhouette, (BMNH. 
1952. 12. 17.2 10-2 14) [Examined]. 

DIAGNOSIS. H. alboclypea is closely related to H. inermis (Caporiacco) but can be readily 
distinguished by the presence of a fringe of white hairs on the clypeus and the structure of the 
genitalia (Fig. 4A-E). 

186 F. R. WANLESS 

REMARKS. This species has only recently been described (Wanless, 1981) and as in the case 
of//, striolata, only figures of the genitalia are provided. 

DISTRIBUTION. Seychelle Islands: Mahe, Silhouette. 

MATERIAL EXAMINED. 4 dtf, 299, including the holotype; for data see Wanless (1981). 

Hispo inermis (Caporiacco) comb. nov. 
(Fig. 5A-J) 

Pseudomarengo inermis Caporiacco, 1947 : 228, rf. Holotype cf, Kenya, Lac Dijpe (TM, Budapest) 

[Examined]. Roewer, 1954 : 951 : 1965 : 33. 
Pseudomarengo rufescens Caporiacco, 1947:229, 9. Holotype 9, Kenya, Moshi (TM, Budapest) 

[Examined]. Roewer, 1954 : 951; 1965.: 33 [=P. inermis]. 

DIAGNOSIS. Hispo inermis is closely related to H. alboclypea Wanless, but can be readily 
separated by the absence of a fringe of white hairs on the clypeus and the structure of the 
genitalia (Fig. 5D, F-J). 

MALE FROM ANGOLA. Carapace (Fig. 5A, B): punctured-reticulate, weakly iridescent 
especially within quadrangle; depressions behind AM hardly apparent; dark orange-brown 
with faint sooty markings; thinly clothed in fine pale brown hairs. Eyes: with black 
surrounds; anteriors contiguous, apices slightly procurved, sparsely fringed in whitish hairs. 
Clypeus: edged black with scattered pale brown hairs. Chelicerae: with fine irregular striae; 
dark brown, shiny; teeth not examined. Maxillae: orange-brown, inner distal margins 
whitish. Labium: slightly broader than long; orange-brown. Sternum: elongate scutiform; 
orange tinged black, shiny. Abdomen: mottled yellow-brown and black with vague 
transverse bands of whitish lanceolate hairs; spinnerets blackish. Legs: legs I (Fig. 5E) and II 
robust; legs I dark orange-brown with tarsi yellow-brown, shiny with a violet tinge under 
some angles of illumination, venter of tibiae and patellae scantily fringed in brown hairs; 
other legs brownish yellow lightly tinged black. Spines few and weak, apparently lacking on 
legs I-I. Palp (Fig. 5D, H): ventral tibial setae relatively stout; embolus slender partly hidden 
by the subtegulum in ventral view. 

Dimensions (mm): total length 3'08; carapace length 1-56, breadth 1*03, height O47; 
abdomen 1*56; eyes, anterior row 0'84, middle row 0'8, posterior row 0'82; quadrangle 
length 0-64. Ratios: AM : AL : PM : PL :: 8 : 3-5 : 1 : 3-5, AL-PM-PL :: 3-2-6-5. 

FEMALE FROM ANGOLA. Similar to c? except for the following. Carapace: finely punctate- 
reticulate, weakly iridescent especially in eye region; dark orange, thinly clothed in fine pale 
orange hairs. Chelicerae: small, light orange, shiny; promargin with 3 teeth, retromargin with 
5 or 6 forming a serrated ridge. Sternum: pale yellow-orange with darker margins. Abdomen: 
whitish yellow clothed in fine pale brown hairs. Legs: more or less as in rf, but pale yellow 
orange with scanty scopulae on tarsi IV (rf similar, but scopulae less noticeable). Epigyne 
(Fig. 5G, I, J): in this specimen the broken off embolus tip (e) can be seen in the left opening. 

Dimensions (mm): total length 4*56; carapace length 1-88, breadth 1'26, height 0-60; 
abdomen length 2*84; eyes, anterior row 1*04, middle row 0*94, posterior row 1-04; 
quadrangle length 0*8. Ratios: AM : AL : PM : PL :: 9-2 : 4-2 : 1 : 4, AL-PM-PL :: 4-5-8-5. 

VARIATION. <J total length 3-0 to 4-2 mm, carapace length 1-36-1-88 mm (8 specimens). 9 
total length 3-9 to 5-2 mm, carapace length 1-64-1-88 mm (9 specimens). Depth of colour 
varies from yellow-brown to brown-black, but this could be the effects of preservation. The 
epigynes are sometimes plugged and the number of coils visible through the integument 
varies; there are usually three on each side (Fig. 5F), but evidently the coils can overlap so 
that in some specimens only two are apparent (Fig. 51). 

DISTRIBUTION. Angola, Botswana, Kenya, Zaire. 

MATERIAL EXAMINED. Angola: Dundo, R. Luachimo, 1 d, 25.11.1948 (A. de Barros 
Machado, Ang. 400.26); Dundo, in house, 1 <?, v.1960 (A. de Barros Machado, Ang. 14938); 



Fig. 5 Hispo inermis (Caporiacco), d 1 : A, dorsal view; B, carapace, lateral view; D, palp, lateral 
view; E, leg I; H, palp, ventral view. 9: C, tracheal system, schematic; F, epigyne; G, vulva, 
ventral view; I, epigyne, another specimen; J, vulva, dorsal view. 

Dundo, forest gallery nr. museum, 2 dtf, 8 99, iii.1948 (A. de Barros Machado, Ang. 414.5); 
Tchivinguiro, forest litter, 1 cf, 27. ix. 1949 (A. de Barros Machado, Ang. 1877.21). 
Botswana: Maqwee, Mopane woodland, pitfall trap, 1 cf, 12.xii.1975 (A. Russel- Smith) 
(BMNH). Kenya: Lac Djipe, holotype <t [Pseudomarengo inermis], x.1904 (K. Kittenberger, 
1228/1905) (TM, Budapest); Moshi, holotype 9 [P. rufescens], ix.1903 (K. Kittenberger, 
1228/1906) (TM, Budapest). Zaire: lac Tanganika, He de Mboko, humus, 1 9, 6.H.1957 (N. 
Leleup, MT. 91383); Kivu, Terr. Uvira, massif de 1'Itombwejtete de source riv. 
Kalimabenge, foret de montagne avec bambous, 2900 m, 1 9, xii.1958 (N. Leleup, MT. 
1 13 194); Kwango, Terr. Feshi, foret de tete de la source Sengi, dans humus, 1 9, iii.1959 (N. 



Leleup, MT. 1 13785); Tshuapa, Bamania, 1 <f, xii.1954 (P. Hulstaert, MT. 84034) (MRAC. 

Hispo frenata (Simon) comb. nov. 
(Fig. 6A-G) 

Astaenorchestes frenatus Simon, 1900:397, d. LECTOTYPE d' (here designated) Madagascar 
(MNHN, Paris, 13127) [Examined]. Simon, 1901:450^52. Roewer, 1954:985. Bonnet, 
1955 : 766. Proszyriski, 1971 : 379. 

DIAGNOSIS. Hispo frenata is closely related to H. tennis sp. n., H. sulcata sp. n., and H. 
macfarlanei sp. n., but can be separated by the absence of spurs on the palpal tibial 
apophyses (Fig. 6C). 

FEMALE. Unknown. 

MALE LECTOTYPE (in poor condition). Carapace (Fig. 6A, D): brown-black, an iridescent 
sheen within quadrangle; scattered white hairs below lateral eyes (badly rubbed). Eyes: with 
black surrounds; anteriors more or less contiguous, apices slightly procurved, sparsely 
fringed in white hairs. Clypeus: rubbed, but probably sparsely fringed in light brown hairs. 
Chelicerae (Fig. 6E): smooth with deep cross furrows; brown-black, rather shiny; promargin 
with 4 teeth, retromargin with 6 or 7 forming a serrated ridge. Maxillae and labium: brown- 
black. Sternum (Fig. 6F): grey-yellow, shiny. Abdomen: moderately elongate; brown tinged 
black, original pattern lost, but irregular spots of whitish hair present. Legs: legs I longest, 
brown-black with yellow-brown tarsi, thinly clothed in fine pale orange hairs; other legs 
yellow-brown tinged black. Spination of legs I: tibiae with 2 distal ventral spines; 

Fig. 6 Hispo frenata (Simon), lectotype d; A, carapace, dorsal view; B, cheliceral teeth; C, palp, 
lateral view; D, carapace, lateral view; E, chelicera, showing furrow; F, sternum; G, palp, ventral 



Fig. 7 Hispo tennis sp. n., holotype d: A, dorsal view; B, carapace, lateral view; C, sternum; D, 
cheliceral teeth; E, palp, lateral view; F, palp, ventral view; G, tibial apophysis, lateral aspect, 
viewed slightly from below. 

other leg spines few and moderately weak. Palp (Fig. 6C, G): additional spurs on the tibial 
apophyses are lacking in this species. 

Dimensions (mm): total length 4*32; carapace length 2*04, breadth 1*52, height 0'84; 
abdomen length 2-2; eyes, anterior row 1'36, middle row 1'08, posterior row MO; 
quadrangle length 1-0. Ratios: AM : AL : PM : PL :: 12 : 5-5:1: 4, AL-PM-PL :: 5-5-1 1. 

DISTRIBUTION. Madagascar. 

MATERIAL EXAMINED. Madagascar: Imerina, Lectotype 

(Camboue) (MNHN, Paris. 

Hispo tenuis sp. n. 

(Fig. 7A-F) 

DIAGNOSIS. Hispo tenuis is closely related to H. frenata (Simon), H. sulcata sp. n., and //. 

190 F. R. WANLESS 

macfarlanei sp. n., but may be distinguished by the more robust embolus (Fig. 7F) and 
presence of a prolateral spur on the dorsal prong of the tibial apophysis (Fig. 7G). 

FEMALE. Unknown. 

MALE HOLOTYPE. Carapace (Fig. 7A, B): finely rugose to finely punctate-reticulate within 
eye region; brownish orange, weakly iridescent. Eyes: with black surrounds; anteriors 
contiguous, apices slightly procurved, fringed in whitish hairs'. Clypeus: with scattered long 
pale brown hairs. Chelicerae: finely rugose; dark brown, weakly iridescent; promargin with 4 
teeth, retromargin with 6 or 7 forming a serrated ridge. Maxillae and labium: brownish 
orange, inner distal margin of maxillae whitish. Sternum (Fig. 7C): brownish orange lightly 
tinged black. Abdomen: mottled yellow-brown and black with whitish markings; anal 
tubercle whitish. Legs: legs I longest, brownish orange to light brown; other legs 
yellow-brown to whitish yellow with blackish femora. Spination of legs I: metatarsi with 1 
prolateral spine; other leg spines few and weak. Palp (Fig. 7E-G): embolus robust, prolateral 
spur on tibial apophysis can only be seen when palp is viewed slightly from below (Fig. 7G). 

Dimensions (mm): total length 4*08; carapace length 1-68, breadth 1*05, height O6; 
abdomen length 2- 12; eyes, anterior row 0*96, middle row 0-78, posterior row 0*84; 
quadrangle length 0-8. Ratios: AM : AL : PM : PL :: 9 : 4 : 1 : 3-5, AL-PM-PL :: 5-9. 

VARIATION. 3 total length 3'6 to 4-08 mm, carapace length 1-48-1-68 mm (3 specimens). 
DISTRIBUTION. Madagascar. 

MATERIAL EXAMINED. Sri Lanka: Kandy, lectotype juvenile, (E. Simon) (MNHN Paris 
paratypes2d-cf,x.l971 (B. Ranson, MT. 1 42.876) (MRAC, Tervuren). 

Hispo sulcata sp. n. 
(Fig. 8A-I) 

DIAGNOSIS. Hispo sulcata is closely related to H. frenata (Simon), H. tenuis sp. n., and H. 
macfarlanei sp. n., but can be distinguished by the presence of a spur on the lateral prong of 
the tibial apophysis (Fig. 8B, C). 

FEMALE. Unknown. 

MALE HOLOTYPE. Carapace (Fig. 8H): finely rugose to aciculate in eye region; dark brown, 
quadrangle paler, shiny with a greenish iridescent tinge on thoracic part under some angles 
of illumination. Eyes: with black surrounds; anteriors more or less contiguous, apices level, 
sparsely fringed in whitish hairs. Clypeus: sparsely fringed in fine whitish hairs. Chelicerae 
(Fig. 8F, G): smooth with deep cross furrows; orange-brown, shiny; promargin with 5 teeth, 
retromargin with 6 or 7 forming a serrated ridge. Maxillae (Fig. 8E, H): blades more or less 
rounded, but with a broad groove laterally; orange-brown. Labium: orange-brown. Sternum 
(Fig. 8D): yellow-brown, shiny. Abdomen: as in H. pullata, but whitish markings less 
distinct. Legs: legs I longest, dark brownish orange; other legs yellow-brown to orange- 
brown. Spination of legs I: tibiae with 1 distal ventral spine, femora with 2 dorsal and 1 distal 
proventral; other spines weak, but more numerous especially on legs III-IV. Palp (Fig. 
8A-C): the spur on the lateral prong of the tibial apophysis is best seen when viewed from 
slightly below. 

Dimensions (mm): total length 4-9; carapace length 2-1, breadth 1-64, height O23; 
abdomen length 2-68; eyes, anterior row 1-4, middle row 1-12, posterior row 1-16; 
quadrangle length 1-04. Ratios: AM : AL : PM : PL :: 12 : 6 : 1 : 5, AL-PM-PL :: 6-12. 

DISTRIBUTION. Madagascar. 

MATERIAL EXAMINED. Madagascar: holotype rf, no other data, (MCZ, Harvard). 



Fig. 8 Hispo sulcata sp. n., holotype d". A, palp, ventral view; B, tibial apophysis, lateral aspect, 
viewed from below; C, palp, lateral view; D, sternum; E, maxillae and labium; F, chelicera, 
showing furrow; G, cheliceral teeth; H, carapace, lateral view; I, leg I. 

Hispo mac far land sp. n. 
(Fig. 9A-M) 

DIAGNOSIS. Hispo macfarlanei is closely related to H.frenata (Simon), H. tennis sp. n., and 
H. sulcata sp. n., but may be distinguished by the backward pointed spur on the dorsal prong 
of the tibial apophysis (Fig. 9J, K) and the presence of a small epigynal scape (Fig. 9C). It is 
not known if the scape is diagnostic for females, as females offrenata, tenuis and sulcata are 

MALE HOLOTYPE. Carapace (Fig. 9A, B): finely rugose, shiny in eye region; dark reddish 
brown with light orange guanin within quadrangle and yellow-brown patch behind posterior 
lateral eyes; below posterior eyes and on thoracic sides narrow stripes of short white hairs. 
Eyes: with black surrounds; anteriors contiguous, apices level, sparsely fringed in fine shiny 
hairs. Clypeus: thinly clothed in fine shiny hairs. Chelicerae (Fig. 9F, I): smooth with shallow 



cross furrows; dark reddish brown, shiny; promargin with 4 contiguous teeth, retromargin 
with 9 or 10 forming a serrated ridge. Maxillae and labium: orange-brown. Sternum (Fig. 
9H): yellow with darker margins, shiny. Abdomen: pale yellow to yellow-brown with 
blackish markings and spots of short white hairs; anal tubercle sparsely fringed in white 
hairs. Legs: legs I elongate (Fig. 9D), orange-brown, thinly clothed in light orange hairs with 
white haired patches on femora and upper parts of trochanters and coxae; other legs light 


Fig. 9 Hispo macfarlanei sp. n., holotype d: A, dorsal view; B, carapace, lateral view; D, leg I; F, 
chelicera, showing furrow; H, sternum; I, cheliceral teeth; J, tibial apophysis, lateral aspect, 
viewed slightly from below; K, palp, lateral view; L, palp, ventral view. Paratype 9: C, epigyne; 
E, leg I; G, vulva, ventral view; M. vulva, dorsal view. 


orange-brown with yellowish trochanters and coxae, thinly clothed in fine pale yellowish 
hairs and with scattered white hairs on femora. Spination of legs I-II: femora with 3 dorsal 
spines; other leg spines moderately strong and numerous. Palp (Fig. 9J-L): tibial apophysis 
distinctive, unlikely to be confused with other known species of Hispo. 

Dimensions (mm): total length 6'56; carapace length 2*84, breadth 2*16, height 1-24; 
abdomen length 3'68; eyes, anterior row 1*18, middle row 1-56 posterior row 1'58; 
quadrangle length 1-4. Ratios: AM : AL : PM : PL :: 17 : 7-5 : 1-4 : 6'5, AL-PM-PL :: 8-17. 

FEMALE PARATYPE. Similar to cf except for the following. Clypeus: densely clothed in white 
hairs. Chelicerae: more bulbous anteriorly, furrow hardly apparent; dark orange, shiny; 
sparsely clothed in fine light brownish hairs. Legs: legs I (Fig. 9E) and II slightly more robust 
than in the cf; legs I dark orange; other legs pale yellow-orange to dark orange. Epigyne (Fig. 
9C, G, M): the small finger-like scape which arises from the anterior margin of the indistinct 
opening is surrounded by a dark red-brown substance, (?plugged) in the two epigynes 
available for study. Unfortunately it has not been possible to determine the precise nature of 
the scape, furthermore the spermathecae, which are blackish, may have altered during 

Dimensions (mm): total length 7 - 92; carapace length 2*84, breadth 2*08, height 1'24; 
abdomen length 4 - 6; eyes, anterior row 1'87, middle row l - 48, posterior row 1'52; 
quadrangle length 1-4. Ratios: AM : AL : PM : PL :: 17 : 7-5 : 1-3 : 7, AL-PM-PL :: 8-16. 

VARIATION, d total length 4*88 to 6*84 mm, carapace length 2-44-2-84 mm (8 specimens). A 
second 9 measures 6*36 mm total length, 2-72 mm carapace length. 

DISTRIBUTION. Madagascar. 

MATERIAL EXAMINED. Madagascar: Antongil, holotype cf, paratypes 7 cfcf, 2 99, (A. 
Mocqueries) (MNHN, Paris, 19541). 

ETYMOLOGY. This species is named after Mr D. Macfarlane of the Commonwealth Institute 
of Entomology, London. 

Hispo cingulata Simon 

Hispo cingulata Simon, 1886 : 394, 9. LECTOTYPE $ (here designated) Madagascar, (MNHN, Paris, 
5.487) [Examined]. Simon, 1901:450-452. Petrunkevitch, 1928:186. Bonnet, 1957:2220. 
Proszyriski, 1971 : 41 7. 

H. cingulatus: Roewer, 1954 : 985 [unjustified emendation]. 

DIAGNOSIS. Hispo cingulata is a distinctive species readily separated from other known 
species of Hispo by the presence of a 'conductor' (c) at the base of the embolus (Fig. 1 1 A, B), 
and the structure of the epigyne (Figs 10F, 1 1C, D). 

MALE FROM ANTONGIL. Carapace: finely rugose within eye region especially in depressions 
behind anterior median eyes; dark reddish, weakly iridescent; below posterior lateral eyes a 
narrow white haired stripe. Eyes: with black surrounds; anteriors contiguous, apices level, 
sparsely fringed in fine brown hairs. Clypeus: white hiared. Chelicerae: irregularly strigose 
with shallow vertical furrows; dark reddish, weakly iridescent; promargin with 4 contiguous 
teeth, retromargin with 8-10 forming a serrated ridge. Maxillae and labium: orange-brown. 
Sternum: elongate scutiform; orange-brown, shiny; slightly depressed opposite coxae I. 
Abdomen: yellow-brown tinged black with transverse bands of short white hairs; anal 
tubercle white haired. Legs: legs I-II robust, dark orange-brown clothed in stiff fine brownish 
hairs; other legs orange-brown lightly tinged black, slightly less hirsute. Spination of legs I-II: 
metatarsi with 1 proventral spine, tibiae with 2 ventral spines and 1 prolateral, femora with 3 
dorsals and 2 prolaterals; other leg spines more numerous. Palp (Fig. 1 1 A, B): the conductor 
(c) appears to be diagnostic for this species. 
Dimensions (mm): total length 7*52; carapace length 2-4, breadth 2-2, height 1-2; abdomen 




Fig. 10 Hispo cingulata Simon, lectotype 9: A, dorsal view; B, cheliceral teeth; C, carapace, 

lateral view; D, sternum; E, leg I; f, eipgyne. 

length 4-0; eyes, anterior row 1'84, middle row 1*44, posterior row 1'56; quadrangle length 
1-52. Ratios: AM : AL : PM : PL :: 17 : 7-5 : 2 : 7, AL-PM-PL :: 8-18. 

FEMALE LECTOTYPE (Fig. 10A-F): similar to rf except for the following. Cheliceme (Fig. 10B): 
slightly more bulbous anteriorly, furrow not apparent, irregularly strigose; dark reddish, 
shiny. Epigyne (Fig. 10F): vulva of another specimen (Fig. 1 1C, D). 

Dimensions (mm): total length 8'5; carapace length 3*32, breadth 2'02, height 1-16; 
abdomen length 4-8; eyes, anterior row 1-7, middle row 1-36, posterior row 1-46; quadrangle 
length 1-46. Ratios: AM : AL : PM : PL :: 16 : 7 : 1-5 : 5-5, AL-PM-PL :: 8-18. 

VARIATION, d 1 total length 5'84 to 7*52 mm, carapace length 2 - 4-3 - 36 mm (4 specimens). A 9 
in poor condition measures about 5'8 mm total length, 3'08 mm carapace length. The 
vertical cheliceral furrow is less evident in the smaller male and it is probably an allometric 
growth character. 

DISTRIBUTION. Madagascar. 

MATERIAL EXAMINED. Madagascar: lectotype 9, (MNHN, Paris. 5.487); Antongil, 2 dtf, (A. 
Mocqueries) (MNHN, Paris. 19541); Ste Marie Island, 2 dtf, 1 9, (A. Mocqueries) (MNHN, 
Paris. 19944). 

The bipartita-group 

This group is comprised of two species characterized by the presence of a fovea. H. bipartita 
Simon, is only known from the female, while H. alboguttata Simon is known from one 
subadult specimen. 


Hispo bipartita Simon 
(Fig. 12A-F) 

Hispo bipartita Simon, 19030 : 1050. LECTOTYPE juvenile, (here designated) Sri Lanka (MNHN, 
Paris, 20406) [Examined]. Bonnet, 1957 : 2219 [nomen nudum}. Proszyriski, 1971 :417. 

REMARKS. Bonnet, 1957 was mistaken in regarding the name H. bipartita as a nomen 
nudum. Simon (1903a) provided the briefest diagnosis and the name is therefore available. 
Furthermore an adult female in the collections of the British Museum (Natural History) is 
considered to be conspecific with the lectotype, a juvenile specimen. 

DIAGNOSIS. H. bipartita seems to be closely related to H. alboguttata Simon, which is only 
known from a single subadult female. They are separated by their colour markings (Figs 
1 2 A, 1 3 A), although this may or may not hold good for adults. 

MALE. Unknown. 

FEMALE FROM INDIA. Carapace (Fig. 12 A, D): finely rugose in eye region to very finely 
papillate on thoracic part; orange-brown, sparsely clothed in short white lanceolate hairs. 
Eyes: with black surrounds; anteriors contiguous, apices slightly procurved, fringed in pale 
brown hairs with tufts outside the anterior lateral eyes. Clypeus: sparsely fringed in long 
whitish hairs. Chelicerae (Fig. 12C): irregularly strigose; lateral condyle prominent; 
orange-brown, weakly iridescent; promargin with 4 contiguous teeth, retromargin with 5 or 6 
forming a serrated ridge. Maxillae and labium: pale orange to light yellow. Sternum (Fig. 
12B): pale orange with darker margins. Abdomen: mottled yellow-brown and black with 
oblique fringes of short white hairs. Legs: legs I (Fig. 12E) and II enlarged; legs I pale orange, 
thinly clothed in pale orange hairs that arise from small circular pits (best seen on the tibiae); 
other legs yellow-brown to pale yellow. Spination of legs I: femora with 3 dorsal, almost 
hair-like spines; other leg spines few and weak. Epigyne (Fig. 1 2F): vulva not examined. 

Dimensions (mm): total length 5-6; carapace length 2*4, breadth 1-64, height O56; 
abdomen length 3*12; eyes, anterior row 1-2, middle row 0*96, posterior row 1-2; quadrangle 
length 1-0. Ratios: AM : AL : PM : PL :: 10-5 : 5 : 2 : 5, AL-PM-PL :: 4-13. 

Fig. 11 Hispo cingulata Simon, cf: A, palp, ventral view; B, palp, lateral view. 9: vulva, ventral 

view; D, vulva, dorsal view. 



Fig. 12 ///s/?o bipartita Simon, 9: A, dorsal view; B, sternum; C, cheliceral teeth; D, carapace, 

lateral view; E, leg I; F, epigyne. 

VARIATION. The juvenile has the markings of the adult with a tuft of hairs outside of the 
anterior lateral eyes; the fovea, although present, is very small. 

DISTRIBUTION. India, Sri Lanka. 

MATERIAL EXAMINED. Sri Lanka: Kandy, lectotype juvenile, (E. Simon) (MNHN, Paris, 
20406). India: Madras, 1 ?,(N. S. Jambunathan) (BMNH. 1923.12.21.76). 

Hispo alboguttata Simon 
(Fig. 13A-B) 

Hispo alboguttata Simon, [June] 19030 : 1050. LECTOTYPE subadult 9 (here designated) Sumatra 
(MNHN, Paris. 22180) [Examined]. Simon, [September] 19036 : 306. Roewer, 1954 : 985. Bonnet, 
1 957 : 22 1 9. Proszynski, 1971:417. 

REMARKS. Although this species is only known from a subadult female the colour markings 
would appear to be distinctive and adults should be recognizable. The specimen is therefore 
figured and briefly described below. 

DIAGNOSIS. H. alboguttata can be distinguished from H. bipartita by the discoidal colour 
markings (Fig. 13 A). 



MALE. Unknown. 

FEMALE LECTOTYPE (subadult). Carapace (Fig. 13 A, B): orange with black markings in eye 
region; very sparsely clothed in fine orange and whitish hairs. Eyes: with black surrounds, 
anteriors contiguous, apices slightly procurved, fringed in whitish hairs with light orange 
hair tufts outside anterior lateral eyes. Clypeus: sparsely fringed in long whitish hairs. 
Chelicerae: pale yellow. Maxillae, labium & sternum: pale yellow, shiny. Abdomen: yellow- 
brown with pattern of discoidal white hair tufts and black markings; spinnerets yellow- 
brown, tinged black and thinly clothed in golden hairs. Legs: legs I-II enlarged; pale yellow, 
but legs I tinged black. Spines few and weak. 
Dimensions (mm): total length 7*05; carapace length 3*04. 


MATERIAL EXAMINED. Sumatra: Palembang, Foret du Nirou, lectotype subadult 9, (M. J. 
Bouchard) (MNHN, Paris. 22 1 80). 

Species Incertae Sedis 

Astaenorchestes continentalis Caporiacco 

Astaenorchestes continentalis Caporiacco, 1949:462, 9 juv. Kenya, Nairobi, 1944. Roewer, 

This species was originally described from a juvenile female. I have been unable to examine 
the type and the species cannot be positively identified from the original description. 
However, the description is consistent with Hispo and the specimen was probably a juvenile 
of//, inermis which Caporiacco himself first described from Kenya. 


Fig. 13 Hispo alboguttata Simon, lectotype, subadult 9: A, dorsal view; B, carapace, lateral view. 

198 F. R. WANLESS 


I wish to thank the following colleagues who kindly made types and other material available 
for study: Professor P. L. G. Benoit, Musee Royal d'Afrique Centrale, Tervuren, Belgium 
(MRAC, Tervuren); M. M. Hubert, Museum National d'Histoire Naturelle, Paris, France 
(MNHN, Paris); Professor H. W. Levi, Museum of Comparative Zoology, Harvard, U.S.A. 
(MCZ, Harvard); Dr S. Mahunka, Termeszettudomanyi Muzeum Allattara, Budapest, 
Hungary (TM, Budapest). 

I am also grateful to Mr D. Macfarlane of the Commonwealth Institute of Entomology, 
London, for reading the manuscript and for linguistic help. 


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de Belgique, Bruxelles. 
1965. Die Lyssomanidae und Salticidae Pluridentati der Athiopischen Region (Araneae). Annls 

Mus. r. Afr. cent. No. 1 39-86 pp. 
Simon, E. 1886. Etudes Arachnologiques 18e Memoire (1) XXVI Materiaux pour servira la faunedes 

arachnides du Senegal. Annls Soc. ent. Fr. (6) 5 : 345-396. 
1897. Etudes Arachnolgiques 29e Memoire (1) XLVI Arachnides recueillis en 1895 par M. le Dr 

A. Brauer (de 1'Universite de Marburg) aux iles Sechelles. Annls Soc. ent. Fr. 66 : 370-388. 
1900. Descriptions d'arachnides nouveaux de la famille des Attidae. Annls Soc. ent. Belg. 


1901 . Histoire Naturelle des Araignees, 2 (3). Roret, Libraire Encyclopedique : Paris. 

1903a. Histoire Naturelle des Araignees, 2 (4). Roret, Libraire Encyclopedique : Paris. 
19036. Arachnides recueillis a Sumatra par M. J. Bouchard. Annls Soc. ent. Fr. 72 : 301-310. 

Wanless, F. R. 1978a. A revision of the spider genera Belippo and Myrmarachne (Araneae : Salticidae) 

in the Ethiopian region. Bull. Br. Mus. nat. Hist. (Zool.,) 33 (1 ) : 1 39 pp. 
19786. A revision of the spider genus Portia (Araneae : Salticidae) Bull. Br. Mus. nat. Hist. (Zool.) 

34 (3): 83-124. 
1981. Contributions a 1'etude de la faune terrestre des iles granitiques de 1'archipel des Sechelles 

(Mission P.L.G. Benoit J. J. Van Mol 1972) Salticidae (Araneae). Annals Mus. r. Afr. cent. 8, Zool. 

No. 235. 

A revision of the spider genus Phaeacius ( Araneae : 

F. R. Wanless 

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


Phaeacius Simon 1900 is a small genus of oriental salticids comprised of five known species 
three of which are described here as new. They are characterized by the presence of a long 
slender secondary conductor in males (Fig. 5C) and a specialized patch of spatulate setae, of 
unknown function, on the underside of coxae IV in females. It is unfortunate that the biology 
is unknown as it would be of interest to compare the behaviour of these salticids with that of 
Portia Karsch, a closely related genus of detritus mimics, some species of which build webs 
of their own as an aid to capture prey or invade the webs of other spiders and feed on the 
occupants (Wanless, 1978&). In spite of their large size (total length 7'5 to 11 -5 mm) 
Phaeacius species are rare in museum collections and in the majority of cases only the types 
were available for study. 

The measurements were made in the manner described by Wanless (19780), but for the leg 
spination the system adopted is that used by Platnick and Shadab (1975). 

Genus PHAEACIUS Simon 

Phaeacius Simon, 1900:32. Type species Phaeacius fimbriatus Simon, by original designation. 

Simon, 1901:406-408. Waterhouse, 1902:280. Petrunkevitch, 1928:182. Roewer, 1954:935. 

Bonnet, 1958:3493. 
Cocalus: Thorell, 1892 : 353 [in part]. 

DEFINITION. Medium to large spiders ranging from about 7'5 to 1 1*5 mm in length. Sexual 
dimorphism not marked. Most species hirsute, with similar colour patterns consisting of 
white marginal carapace bands and leaf-like abdominal markings composed of setae which 
are easily rubbed; abdominal markings may also include granules of subcutaneous guanin 
(Fig. 9A) possibly an artifact of preservation. Carapace: of medium height, longer than 
broad, widest between coxae II-III, lateral margins with distinct membraneous border 
(arrowed in Fig. 5E); fovea long and sulciform, situated just behind posterior median eyes; 
eye region finely punctate and weakly iridescent (evident only in rubbed specimens). Eyes: 
set on well developed tubercles, with black surrounds that are normally covered in hairs; 
arranged in three transverse rows comprised of anterior medians and anterior laterals, 
posterior medians, and posterior laterals; anteriors densely fringed in hairs, more or less 
subcontiguous with apices slightly procurved in frontal view; anterior medians largest, 
anterior laterals more than half diameter of anterior medians; posterior medians relatively 
large, positioned slightly closer and on or slightly inside optical axis of anterior laterals, 
posterior laterals almost as large as anterior laterals, set slightly closer together and well 
inside lateral margins of carapace; quadrangle formed by posterior median and posterior 
lateral eyes broader than long and widest posteriorly; entire quadrangle (measured from base 
of anterior medians to posterior margin of posterior laterals) occupying between 37 and 44 
per cent of carapace length. Clypeus: between 25 and 36 per cent of diameter of anterior 
median eyes; clothed in hairs; ventrally a sometimes rather broad segmental membrane 
bearing a median subtriangular sclerite. Chelicerae: robust; more or less parallel, slightly 
inclined anteriorly; basal region usually clothed in hairs, short in central area, long on lateral 

Bull. Br. Mus. nat. Hist. (Zool.) 41(4): 1 99-2 1 2 Issued 26 November 1 98 1 




Fig. 1 Phaeacius lancearius (Thorell), d 1 palp, x 32. Note distorted secondary conductor. 

margins; promargin with three teeth, retromargin with three to five. Maxillae: moderately 
long, subparallel. Labium: tongue-shaped, moderately long, slightly greater than half 
maxillae length. Sternum: elongate scutiform; margins not deeply excavated. Coxae: 
subequally robust or coxae IV slightly larger; in females coxae IV with spatulate setae on 
ventral surface. Pedicel: short. Abdomen: elongate ovoid with two pairs of impressed spots, 
the anterior pair the least conspicuous; spinnerets moderately long, posteriors longest with 
apical conical segment bearing moderately elongate spiggots, anteriors robust, medians and 
posteriors relatively slender; tracheal system not examined (insufficient material), spiricle an 
indistinct transverse slit near base of anterior spinnerets; former position of colulus indicated 
by hair tuft; anal tubercle a well developed pointed cone. Legs: moderately long and robust; 
spines strong and numerous; ornate fringes lacking, but female tarsi I-II and metatarsi I-II 
with fine ventral setae; claws pectinate scopulae lacking. Female palps: moderately long and 
robust with terminal claw. Male palps: large, usually dark; moderately complex and 
morphologically similar; densely clothed in hairs (not shown in figures). Tibiae with massive 
retrolateral apophysis, ventral apophysis and sometimes an intermediate apophysis; 
cymbium with small basal protuberance and characteristic profile in dorsal view (Figs 4E; 
8B); embolus (e) robust, long and curved with conspicuous pars pendula (p); primary 
conductor lacking; secondary conductor (sc) translucent long, slender and curved, arising 
from membraneous region which appears to separate the embolic base from the tegulum 
(Fig. 4A), also arising from the membraneous region, below the embolic base, a short 
apophysis (a) (Figs 2E; 6B); tegulum (t) with pronounced tegular furrow (tf), peripheral 
seminal ducts (sd) and retrolateral striae (s) (Figs 1 ; 2 A-F). Epigynes: large and dark; orifice 
paired, ovoid to furrowed, opening of looped introductory ducts lying alongside inner 
margins. Internal structure not examined (insufficient material). 

The secondary conductor clearly arises from a membraneous region of the tegulum which 
lies adjacent to the embolic base (Fig. 2C, D). This region seems to be homologous with a 



Fig. 2 Phaeacius lancearius (Thorell), 3 palp: A, ventral and retrolateral tibial apophyses, x 32; 
B, tegulum with striae, x 80; C, embolus secondary conductor and tegular furrow, x 55; D, origin 
of secondary conductor, x 160; E, apophysis protruding from below embolic base, x 580; F, tip 
of embolus, x 700. 

202 F. R. WANLESS 

similar area found in some species of Portia and referred to as a tripartite membrane in 
Wanless (19786). The dorsal and prolateral elements of the tripartite membrane are not 
evident in Phaeacius. 

AFFINITIES. Phaeacius seems on the basis of the elongate fovea, procurved anterior eye row 
and male palpal structure to be most closely related to the schultzii-group of species in the 
genus Portia Karsch (see Wanless, 19786). The kenti-group of Portia would now appear to 
require a new genus. However, for the present I am reluctant to propose formal changes 
because morphological characters and relationships between other genera in the subfamily 
Boethinae are largely unknown. 

List of species in the genus Phaeacius Simon, 1900 

Phaeacius canalis sp. n. 
P. fimbriatus Simon, 1900 
P. lancearius (Thorell, 1895) 
P. malayensis sp. n. 
P. saxicola sp. n. 

REMARKS. Simon (1901) transferred Cocalus ramipalpis Thorell, from Sumatra, into 
Phaeacius, but to judge from his illustration (Fig. 435, g) the species probably belongs in 
Boethus and is therefore not included in the present revision. 

Key to species of Phaeacius 

Males (males of saxicola are unknown) 

1 Dorsal prong of retrolateral tibial apophysis long (Fig. 8C) (Phillipines) . canalis sp. n. (p. 209) 
Dorsal prong of retrolateral tibial apophysis short (Figs 4C; 5 F; 6 A) 3 

2 Intermediate tibial apophysis present (Fig. 6B) (Singapore) . . . malayensis sp. n. (p. 205) 

- Intermediate tibial apophysis lacking 3 

3 Retrolateral tibial apophysis relatively short and broad, median notch moderately conspicuous 

(Figs 4C; 5 B) (Java) fimbriatus Simon (p. 202) 

- Retrolateral tibial apophysis relatively long and narrow, median notch inconspicuous (Fig. 

5 A, F) (Burma) lancearius (Thorell) (p. 205) 

Females (females of lancearius are unknown) 

1 Epigynal openings subovoid (Fig. 9C) (Nepal) saxicola sp. n. (p. 210) 

Epigynal openings relatively narrow or furrowed 2 

2 Epigynal openings furrowed (Fig. 8G) (Phillipines) canalis sp. n. (p. 209) 

Epigynal openings relatively narrow (Figs 4D; 7C) 3 

3 Anterior margin of epigynal orifice level with anterior loop of seminal duct (Fig. 7C) (Sumatra) 
' . . malayensis sp. n.(p. 205) 

- Anterior margin of epigynal orifice posterior to anterior loop of seminal duct (Fig. 4D) (Java) 
fimbriatus Simon (p. 202) 

Phaeacius fimbriatus Simon 

Phaeacius fimbriatus Simon, 1900:32, 9. lectotype 9 (here designated) Java (MNHN, Paris) 
[examined]. Simon 1901:406^08. Reimoser, 1925:90. Petrunkevitch, 1928:182. Roewer, 
1954 : 935. Bonnet, 1958 : 3493. 

DIAGNOSIS. P. fimbriatus seems to be most closely related to P. lancearius, but may be 
distinguished by the form of the retrolateral tibial apophysis (Figs 4A, C; 5B) which is shorter 
and somewhat broader basally. Also, the median notch (arrowed in Fig. 5B) is more 



) P canalis 

^ . s 

(A) P fimbriatus 
P lancearius 
P malayensis 

M P saxicola 

Fig. 3 Distribution ofPhaeacius spp. 

pronounced. In the lectotype the embolus is clearly shorter than that of P. lancearius, but the 
tip appears to have broken off. Unfortunately the right palp is missing and no other 
specimens are available for comparison. 

REMARKS. A female labelled '20032 Phaeac. fimbriatus E.S. Java: Gedi (Fr)' corresponds 
with the data given in the original description, but was accompanied by an apparently 
conspecific male not mentioned by Simon. As no other specimens can be found in the Simon 
collection (MNHN, Paris) the single female is presumed to be the type specimen. The male is 
described below. 

FEMALE LECTOTYPE. Carapace (Fig. 4B, G): orange-brown with sooty markings; eye region 
finely punctate and weakly iridescent under some angles of illumination; badly rubbed, but 
patches of recumbent short whitish hairs present. Eyes: with black surrounds; fringed in 
whitish hairs especially AM. Clypeus: clothed in brownish orange hairs. Chelicerae: orange, 
shiny; basal region clothed in whitish hairs; promargin with three teeth, retromargin with 
four. Maxillae: light orange grading to whitish yellow on inner distal margins. Labium: pale 
orange tipped whitish yellow. Sternum: pale yellow with light orange margins; clothed in 
light yellow hairs. Coxae: pale yellow to yellow-brown. Abdomen: with two pairs of 
impressed spots; light brownish grey with numerous granules of yellowish subcutaneous 
guanin and lateral dark brown markings forming a foliated pattern more or less as in P. 
saxicola sp. n. (Fig. 9A); basal region with rather scanty clumps of white and dark brown 
hairs, otherwise rubbed; spinnerets pale yellow-brown. Legs: yellow-brown with vague 
orange-brown markings; spines numerous and strong. Spination of legs I: metatarsi v 1-0-0, 
p 1-1-0, r i_i_i ; tibiae v 1^-1 , p 1-0-1 , r 1-0-1 , d 1-1-0; patellae p 1-0-0, r 1-0-0; femora 
p 0-1-1, d 0-2-3. Epigyne (Fig. 4D): dark reddish brown; clothed in light orange-brown 

Dimensions (mm): total length 8 -72; carapace length 3'88, breadth 3*16, height 1*76; 
abdomen length 4-64; eyes, anterior row 2*16, middle row 1-92, posterior row 2*02; 



Fig. 4 Phaeacius fimbriatus Simon, cf: A, palp, ventral view; C, palp, ectal view; E, palpal 
cymbium, dorsal view; G, carapace, lateral view. 9 lectotype: B, carapace lateral view; D, 
epigyne; F, cheliceral teeth; H, carapace, dorsal view. 

quadrangle length 1-6. Ratios: AM : AL : PM : PL : 16 : 9 : 6 : 8'4; AL-PM-PL : 1 1-5-14; 
AM:CL(clypeus): 16:4. 

MALE (formerly undescribed) FROM JAVA. Similar to 9 except for the following: Carapace 
(Fig. 4G): with uneven marginal band of white hairs. Chelicerae: not quite as robust as in 9; 
basal region densely clothed in white hairs noticeably long on lateral margins. Abdomen: 
more or less as in 9, but guanin granules not evident. Legs: similar to 9, but darker markings 
forming indistinct annuli on tibiae and metatarsi. Spination of legs I: metatarsi v 2-0-0, p 


1_1_0 9 r 1-1-2, d 0-1-0; tibiae v 2-3-2, p 1-1-0, r 0-1-1 , d 1-1-1; patellae p 1-0-0, r 1-0-0; 
femora p 0-1-1 , d 0-2-1 , r 0-0-1 . Palp (Figs 4A, C; 5B): rather dark and thickly clothed in 
orange-brown hairs. 

Dimensions (mm): total length 8' 1 ; carapace length 3'8, breadth 3'24, height 2'0; abdomen 
length 4*32; eyes, anterior row 2-1, middle row 1*82, posterior row 1'96; quadrangle length 
1-52. Ratios'. AM : AL : PM : PL : 15 : 9 : 6 : 8; AL-PM-PL : 1 1-13; AM : CL : 15 : 5. 


MATERIAL EXAMINED. Java: Mons Gede, lectotype 9, 1 cT, (Fruhstorfer) (MNHN, Paris. 

Phaeacius lancearius (Thorell) 

Cocalus lancearius Thorell, 1895 : 357, <f. LECTOTYPE rf, (BMNH); PARALECTOTYPE d, (NR, 

Stockholm) (here designated), Burma [examined]. Bonnet, 1956 : 1 173. 
Phaeacius lancearius (Thorell): Roewer, 1954 : 935. 

DIAGNOSIS. P. lancearius seems to be most closely related to P. fimbriatus, but may be 
separated by the longer and somewhat narrower retrolateral tibial apophysis and the less 
pronounced median notch (arrowed in Fig. 5 A). It is possible that intermediate forms may 
occur, but for the moment P. lancearius is regarded as a good species. 

FEMALE. Unknown. 

MALE LECTOTYPE. Carapace (Fig. 5D, E): greyish black with finely punctate orange-brown 
eye region; margins with broad uneven white haired bands, otherwise badly rubbed. Eyes: 
with black surrounds, irregularly fringed in whitish and light brown hairs (mostly rubbed). 
Clypeus: clothed in light brown hairs. Chelicerae: orange-brown, basal area clothed in 
whitish hairs, longest on outer lateral margins; promargin with three teeth, retromargin with 
four. Maxillae: dark brown grading to yellow-brown on inner distal margins. Labium: dark 
brown tipped yellowish brown. Sternum: yellow-brown with thin brownish margins, shiny; 
clothed in white hairs. Coxae: greyish, shiny. Abdomen: with two pairs of impressed spots; 
light greyish with granules of yellow guanin; sparsely and irregularly clothed in recumbent 
light brownish hairs; spinnerets greyish yellow. Legs: greyish brown to orange-brown; spines 
numerous and robust. Spination of legs I: metatarsi v 2-0-0, p 1-1-0, r 1-1-2, d 0-1-0; tibiae 
v 2-3-2, p 0-2-0, r 0-2-0, d 1-1-1; patellae p 1-0-0; femora p 0-1-1, d 0-1^. Palp (Figs 1, 
2 A-F; 5A, C, F): very dark. 

Dimensions (mm): total length about 7'76; carapace length about 3'7, breadth 3*08, height 
1-6; abdomen length about 3'8; eyes, anterior row 1*95, middle row 1*68, posterior row 1-8; 
quadrangle length 1-48. Ratios: AM : AL : PM : PL : 14:9 : 5: 5:7; AL-PM-PL : 1 1-13; 
AM:CL: 14:5. 

VARIATION. cT total length 7*76 to 8'72 mm, carapace length 3'72^-Q mm (3 specimens). 
The paralectotype and the male from India are generally paler in colour, the carapace being 
orange to dark orange with a dense covering of mixed white and light orange hairs. The palps 
are also paler and densely clothed in light brownish hairs. 

DISTRIBUTION. Burma; India. 

MATERIAL EXAMINED. Burma: Tharrawaddy, lectotype <?, (E. W. Oates) (BMNH. 
1895.9.21.1065); Tonghoo, paralectotype rf, (E. W. Oates) (NR, Stockholm, 1607). India: 
Southern India, 1 d 1 , no other data (BMNH). 

Phaeacius malayensis sp. n. 


DIAGNOSIS. P. malayensis is closely related to both P. fimbriatus and P. lancearius, but may 



Fig. 5 (A, C-F) Phaeacius lancearius (Thorell) <f lectotype: A, palpal tibia, ectal view; C, palp, 
ventral view; D, carapace, dorsal view; E, carapace, lateral view; F, palp, ectal view. (B) P. 
fimbriatus Simon, d: palpal tibia, ectal view. 

be distinguished by the presence of a pronounced apophysis lying between the ventral and 
retrolateral tibial apophyses (Fig. 6B) in males, and by having the anterior margin of the 
epigynal orifice more or less level with the anterior loop of the seminal duct (Fig. 7C) in 

MALE HOLOTYPE. Carapace (Fig. 6D, E): light orange to orange-brown; finely punctuate with 
iridescent sheen in eye region; sparsely and irregularly clothed in recumbent white hairs 
(rubbed). Eyes: with black surrounds; irregularly clothed in whitish and pale golden hairs. 
Clypeus: fringed in light brown and whitish hairs. Chelicerae: pale orange; basal region white 
haired; promargin with three teeth, retromargin with four. Maxillae: light orange and 




mottled grey to whitish on inner distal margins. Labium: orange-brown tipped greyish white. 
Sternum: pale yellow with light orange margins, shiny; clothed in pale yellow hairs. Coxae: 
pale yellowish, shiny. Abdomen: rubbed; pale yellow with granules of whitish guanin and 
blackish lateral mottling (suggesting a foliated pattern as in other species of Phaeacius). 
Legs: yellowish orange; spines strong and numerous. Spination of legs I: metatarsi v 2-0-0, p 
1_1_0 9 r 1-1-2, d 0-1-0; tibiae v 2-3-2, p 1-0-1, d 2-2-1; patellae p 1-0-0, r 1-0-0; femora 
p 0-1-1 , d 0-2-2, r 0-0-1 . Palp (Fig. 6A, B): densely clothed in light brown and whitish hairs 
which tend to obscure the apophyses that lies between the ventral and retrolateral 

Dimensions (mm): total length 8'0; carapace length 3*8, breadth 3'24, height 1-8; abdomen 
length 4-12; eyes, anterior row 2-16, middle row 1'92 posterior row 2*0; quadrangle length 
1-68. Ratios: AM : AL: PM : PL: 16'5 : 9'5 : 6'5 : 9; AL-PM-PL : H-13'5; AM : CL 

Fig. 6 


Phaeacius malayensis sp. n., d 1 holotype: A, palp, ectal view; B, palp, ventral view; C, 
cheliceral teeth; D, carapace, dorsal view; E, carapace, lateral view. 



FEMALE PARATYPE. Carapace (Fig. 7A): yellow-brown to light amber with sooty markings; 
clothed in short recumbent light brown and white hairs with narrow white haired marginal 
bands. Eyes: with black surrounds; densely fringed in white and pale amber hairs. Clypeus: 
fringed in long whitish hairs. Chelicerae: orange-brown; clothed in long white hairs; 
promargin with three teeth, retromargin with four. Maxillae: orange-brown with whitish 
inner distal margins. Labium: orange-brown tipped whitish. Sternum (Fig. 7B): pale yellow 
with dark margins, shiny; clothed in fine pale yellowish hairs. Abdomen: greyish yellow; 
sparsely clothed in light orange-brown hairs forming a leaf-like pattern with scanty tufts of 
long white hairs posteriorly. Legs: light yellow grading to yellow-brown distally with 
indistinct annuli on metatarsi; spines numerous and robust. Spination of legs I: metatarsi v 
2-0-0, p 1-1-0, r 1-1-1 ; tibiae v 2-3-2, p 1-0-1 , r 1-0-1 , d 1-1-0; patellae p 0-1-0, r 0-1-0; 
femora p 0-1-1 , d 0-1-4. Epigyne (Fig. 7C): large and dark, clothed in fine pale orange hairs. 
Dimensions (mm): total length 8*96; carapace length 3*8, breadth 3' 16, height 1'68; 
abdomen length 5*04; eyes, anterior row 2-12, middle row 1*8, posterior row 1-96; 
quadrangle length 1-6. Ratios: AM : AL : PM : PL : 15 : 9 : 6 : 9; AL-PM-PL : 12-14; 
AM:CL: 15:3. 
DISTRIBUTION. Malaysia, Singapore; Sumatra. 

MATERIAL EXAMINED. Malaysia: Singapore, holotype rf, 1898 (H. N. Ridley) (BMNH, 
1980.4.9.1). Sumatra: Lembang Anai, paratype 9, xii.1974 (W. S. Bristowe) (BMNH. 

Fig. 7 Phaeacius malayensis sp. n., 9 paratype: A, dorsal view; B, sternum; C, epigyne. 


Phaeacius canalis sp. n. 

(Fig. 6A-F) 

DIAGNOSIS. P. canalis is a fairly distinctive species readily separated from other species of 
Phaeacius by the elongate dorsal prong of the retrolateral tibial apophysis (Fig. 6B) in males, 
and the grooved epigynal openings (Fig. 6F) in females. 

HOLOTYPE MALE. Carapace (Fig. 6C, E): pale to dark amber with vague radiating markings; 
irregularly clothed in recumbent clear whitish and pale amber hairs (rubbed), from clypeus 
to posterior thoracic margin a wide uneven white haired band. Eyes: with blackish 
surrounds; densely fringed in whitish and pale amber hairs. Clypeus: clothed in pale amber 
hairs. Chelicerae: dark amber, shiny; basal half densely clothed in white hairs, longest on 
lateral margins; promargin with three teeth, retromargin with four. Maxillae: orange-brown 
grading to whitish yellow on inner distal margins. Labium: orange-brown tipped whitish 

Fig. 8 Phaeacius canalis sp. n., cf holotype: A, palp, ventral view; B, palpal cymbium, dorsal 
view; C, palp, ectal view; D, carapace, dorsal view; E, sternum; F, carapace, lateral view. 9 
paratype: G, epigyne. 

210 F. R. WANLESS 

yellow. Sternum (Fig. 6D): pale yellow-brown, shiny; clothed in whitish yellow hairs. Coxae: 
pale yellow-brown. Abdomen: with two pairs of impressed spots; pale yellow-brown; clothed 
in yellow-brown and orange-brown hairs with scanty white tufts posteriorly; pattern 
essentially as in P. malayensis sp. n. (Fig. 7A). Legs: yellowish orange with obscure dark 
orange annuli; spines strong and numerous. Spination of legs I: metatarsi v 2-0-0, p 1-1-0, r 
1-1-1, d 0-1-0; tibiae v 3-2-2, p 1-0-1, r 1-1-0, d 1-1-1; patellae p 1-0-0, r 1-0-0; femora 
p 0-1-1, d 0-1-3, r 0-0-1. Palp (Fig. 6A, B): densely clothed in whitish and dark amber 
hairs; secondary conductor relatively short. 

Dimensions (mm): total length 9*2; carapace length 4*6, breadth 3*72, height 2'0; abdomen 
length 2-68; eyes, anterior row 2*22, middle row 2*0, posterior row 2*12; quadrangle length 
1-7. Ratios: AM : AL : PM : PL: 17: 10 : 7 : 9; AL-PM-PL : 13-15-5; AM : CL : 17-6. 

FEMALE PARATYPE. Similar to c? except for the following: Carapace: darker; marginal band 
narrower and less clearly defined. Clypeus: clothed in light orange-brown hairs with dense 
covering of long white hairs in lower space between AM. Chelicerae: dark orange, shiny; 
basal region clothed in white hairs; promargin with three teeth, retromargin with five. Legs: 
spination of legs I: metatarsi v 2-0-0, p 1-1-1, r 1-1-0; tibiae v 3-2-2, p 1-1-0, r 1-1-0, d 
1-1-0; patellae p 1-0-0, r 1-0-0; femora p 0-1-1 , d 0-1-4. Epigyne (Fig. 6F): large and dark, 
clothed in long whitish hairs; lateral openings plugged. 

Dimensions (mm): total length about 8'9; carapace length 3-76, breadth 3-04, height 1*86; 
abdomen length 4-8; eyes, anterior row 2*16, middle row 1*84, posterior row 1*96; 
quadrangle length 1-64. Ratios: AM : AL : PM : PL : 15'5 : 8'5; AL-PM-PL : 12-14; 
AM:CL: 15'5:5. 

DISTRIBUTION. Philippines. 

MATERIAL EXAMINED. Philippines: Albay, S.E. Luzon, holotype rf xi-xii.1894 (G. J. 
Whitehead) (BMNH, 1895.1.17.920); Cape Engano, Paratype 9, (G. J. Whitehead) (BMNH, 

Phaeacius saxicola sp. n. 
(Fig. 9A-C) 

DIAGNOSIS. P. saxicola is a moderately distinctive species, readily separated from others in 
the genus by the appearance of the epigyne (Fig. 9C). 

MALE. Unknown. 

FEMALE HOLOTYPE. Carapace (Fig. 9A, B): dark orange-brown with sooty markings, also an 
inverted light orange-brown leaf-shaped patch on thoracic region; clothed in dull whitish 
and light brown hairs with white haired marginal bands and lateral dark brown haired spots. 
Eyes: with black surrounds; densely fringed in whitish and light orange-brown hairs. 
Clypeus: clothed in dull white and orange-brown hairs. Chelicerae: orange to brownish 
orange, shiny; basal region clothed in white hairs; pro and retromargins with three teeth. 
Maxillae: dark orange-brown with inner distal margins light yellowish. Labium: dark 
orange-brown tipped light yellow. Sternum: light yellow-brown with irregular reddish 
orange margins; clothed in creamy white hairs. Coxae: light yellow-brown. Abdomen: light 
grey with granules of yellowish guanin; rubbed, but with leaf-life pattern outlined in brown- 
black hairs. Legs: pale yellowish orange grading to orange-brown distally with indistinct 
annuli on metatarsi and tibiae; spines strong and numerous. Spination of legs I: metatarsi v 
2-0-0, p 1-1-0, r 1-1-1 ; tibiae v 2-3-2, p 1-1-0, r 1-1-0, d 1-1-0; patellae p 0-1-0, r 0-1-0; 
femora p 0-1-1, d 0-1-4. Epigyne (Fig. 9C): large and dark; covered in whitish and light 
yellow-brown hairs. 

Dimensions (mm): total length 11-3; carapace length 4*0, breadth 3'44, height 1-76; 
abdomen length 7-28; eyes, anterior row 2-16, middle row 1-89, posterior row 2'04; 



Fig. 9 Phaeacius saxicola sp. n., 9 holotype: A, dorsal view; B, carapace, lateral view; C, epigyne. 

quadrangle length 1-74. Ratios: AM : AL : PM : PL : 15-5 : 9'5 : 6 : 9; AL-PM-PL : 13-14; 
AM:CL: 15-5:4. 


MATERIAL EXAMINED. Nepal: Maewa Khola, Sanghu, holotype 9, 6,000ft, on rock face, 
10.x. 1961 (K.H.Hyatt, British Museum Nepal Expedition, 42b) (BMNH. 1980.4.15.1). 


I wish to thank M. M. Hubert, Museum national d'Histoire naturelle, Paris, France (MNHN, 
Paris) and Professor T. Kronestedt, Naturhistoriska Riksmuseet, Stockholm, Sweden (NR, 
Stockholm) for providing specimens for study. Also Mr D. Macfarlane (CIE, London) for 
reading the manuscript. 


Bonnet, P. 1956. Bibliographia Araneorum. 2 (2) : 919-1925. Imprimerie Douladoure, Toulouse. 
1958. Bibliographia Araneorum. 2 (4) : 3027-4230. Imprimerie Douladoure, Toulouse. 

212 F. R. WANLESS 

Petrunkevitch, A. 1928. Systema Aranearum. Trans. Conn. Acad. Arts Sci. 29 : 270 pp. 

Platnick, N. I. & Shadab, M. U. 1975. A revision of the spider genus Gnaphosa (Araneae : 

Gnaphosidae) in America. Bull. Am. Mus. nat. Hist. 155 : 3-66. 
Reimoser, E. 1925. Fauna sumatrensis. Supplta ent. 11 : 89-94. 
Roewer, C. F. 1954. Katalogder Araneae. 1 Abt. B: 924-1290. Institut Royal des Sciences naturelle de 

Belgique, Bruxelles. 
Simon, E. 1900. Etudes arachnologiques 30e Memoire (1) XLVII Descriptions d' especes nouvelles de 

la famille des Attidae. Annls Soc. ent. Fr. 69 : 27-6 1 . 

1901. Histoire Naturelle des Araignees, 2 (3) : 381-668. Paris, Roret: Libraire Encyclopedique. 

Thorell, T. 1892. Studi sui ragni Malesi e Papuani. (4) 2. Mus. civ. Stor. nat. Giacomo Doria. 31 : 490 


1 895. Descriptive catalogue of the spiders of Burma. 406 pp. British Museum (natural History). 

Wanless, F. R. 1978a. A revision of the spider genera Belippo and Myrmarachne (Araneae : Salticidae) 

in the Ethiopian region. Bull. Br. Mus. nat. Hist. (Zool.) 33 (1) : 139 pp. 
1978ft. A revision of the spider genus Portia (Araneae : Salticidae). Bull. Br. Mus. nat. Hist. (Zool.) 

34 (3): 83-1 24. 
Waterhou'se, C. O. 1 902. Index zoologicus 42 1 pp. London. 

Manuscript accepted for publication 2 October 1980 

The protractor pectoralis muscle and the 
classification of teleost fishes 

Peter Humphry Greenwood 

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

George V. Lauder 1 

Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, 02 1 38 


The protractor pectoralis muscle is currently thought to be one of the shared derived features 
characterizing certain neoteleostean fishes. Further investigation shows, however, that a 
protractor pectoralis muscle occurs in many taxa throughout the Teleostei, the Dipnoi, the 
Palaeopterygii and other lower actinopterygians, and, probably, in the Elasmobranchio- 
morphi as well. 

Since the protractor pectoralis is apparently homologous with the trapezius muscle of 
tetrapods it therefore would appear to be a derived feature of the Gnathostomata as a whole 
and not just a synapomorphy of a group (the Eurypterygii) within the Teleostei. 

In his paper on the interrelationships of the higher Euteleostei, Rosen (1973) considered 
the presence of a protractor pectoralis muscle to be one of the synapomorphies 
characterizing the section Eurypterygii of his subdivision Neoteleostei. Rosen also included 
in the Neoteleostei, as the primitive sister group of the Eurypterygii, the section 
Stenopterygii (=Stomiatiformes). Although lacking a protractor pectoralis muscle, the 
Stomiatiformes share other derived features with their apomorph sister taxa, viz. the 
Aulopiformes, Myctophiformes, Protacanthopterygii and Acanthopterygii (Rosen, 

According to Rosen (who based his conclusions involving the protractor pectoralis largely 
on the then unpublished work of Winterbottom) this muscle does not occur in any other 
group of teleostean fishes. When that work was published the following year, Winterbottom 
(1974 : 269) repeated the claim that '. . . the protractor pectoralis appears to be confined to 
the neoteleosts', despite the fact that he described and illustrated the muscle in three non- 
neoteleostean taxa, the ostariophysans Brycon, Cyprinus and Diplomystus. 

This lapsus is probably explained by Winterbottom's definition of the concepts 
'neoteleosts' and 'non-neoteleosts' (Winterbottom, 1974 : 227). He uses the term neoteleost 
*. . . in the sense proposed by Rosen & Patterson (1969 : 460)' but uses its antithesis, the 
non-neoteleosts, '. . . to designate Divisions I and II, and the salmoniforms (less mycto- 
phoids) of Division III of the Greenwood et al. (1966) classification'. Thus the Ostariophysi 
and the Gonorynchiformes of Greenwood et al. were left in limbo, neither neoteleosts nor 
non-neoteleosts, and were overlooked. 

A further complication is introduced by Winterbottom's report in some clupeomorphs 
(sensu Greenwood, Rosen, Weitzman & Myers, 1966) of a muscle which, in his opinion 
(1974 : 269) '. . . seems to be analogous to the protractor pectoralis', and with which '. . . the 
muscle in the neoteleosts would appear to be a homologous structure, and indicative of 
common ancestry'. 

These incongruities in recently published accounts, coupled with the fact that a muscle 
apparently identical to the protractor pectoralis in neoteleosts has been described in 

'Present address: Department of Anatomy, University of Chicago, 1025 East 57th Street, Chicago, Illinois 60637, 

Bull. Br. Mus. nat. Hist. (Zool.) 41(4): 2 1 3-234 Issued 26 November 1 98 1 


chondrichthyans (=Elasmobranchiomorphi), dipnoans and non-teleostean actinopterygians 
(see Allis, 1917; Edgeworth, 191 1 & 1935), led us to review and reconsider the value of this 
muscle as a character in reconstructing gnathostome phylogenies, especially those of 
teleostean fishes. 

As a first step we have reviewed the literature dealing with the protractor pectoralis (ie 
essentially the cucullaris and cephaloclavicularis muscles of Edgeworth and the trapezius of 
Allis) in lower gnathostomes, and have compared these findings with our dissections 
covering a wider range of taxa than any previously examined (see Table 1). 

Allis (1917 : 343-350, and 402) gives a detailed summary of the situation at that time with 
respect to gnathostome fishes, and includes a critique of Edge worth's earlier (1911) paper, 
the only wide-ranging review of cranial musculature in vertebrates then available. Later, 
Edgeworth (1935) expanded his earlier essay. Apart from these three works, and 
Winterbottom's (1974) recent contribution, there are no other papers dealing with the prime 
question we wished to review, namely the homology of the protractor pectoralis both within 
and outside the teleostean fishes. 

Allis (1917) concludes that amongst gnathostome fishes the protractor pectoralis (his 
trapezius) is always innervated by the vagus, and that it is derived, embryologically, from the 
muscle plate of the last branchial arch. Thus, at least implicitly, Allis would consider the 
muscle to be homolgous in these animals. Edgeworth (1911; 1935) reaches a similar 
conclusion regarding the muscle's homology, and extends it even beyond the limits of the 
gnathostome fishes. According to Edgeworth (1935 : 151) the protractor pectoralis (which he 
calls cucullaris) 'In Dipnoi, Teleostomi, Amphibia, Reptilia and Mammalia ... is developed 
as a backgrowth of the most caudal branchial muscle plate', and more specifically, (p. 151), 
'The similar primary innervation of the Cucullaris from the most caudal fibres of the Vagus 
shows that the muscle has had a continuous phyletic history and is to be regarded as a 
homologous structure whatever its source'. 

Ontogenetically, Edgeworth considered that in Dipnoi and other teleostomes the 
protractor pectoralis is derived from the posterior margin of the levator arcuum 
branchialium anlage (which would be topologically equivalent to Allis' muscle plate of the 
last branchial arch). However, in the Elasmobranchiomorphi (except the rays, Batoidei), he 
believed that the muscle is formed from the dorsal ends of all the embryonic branchial 
muscle plates (ie the embryonic branchial constrictors since in these animals no levator 
muscles are differentiated; see Edgeworth, 1935 : 232-233). In the Batoidei, according to 
Edgeworth, the protractor develops from the dorsal end of the fifth muscle plate alone, a 
condition which he thought was '. . . probably secondary to that in the Selachii and without 
any genetic relationship to that of the Teleostomi' (see Edgeworth: 1 52, and also 140 & 142). 
In other words, the batoid condition was derived from the selachian one which, in turn, 
Edgeworth considered to be derived from that basic to the gnathostomes, and characterized 
by the failure of the levator muscles to develop from the branchial muscle anlage 
(Edgeworth, 1911 : 193; 1935 : 152). 

Allis (1917 : 346-7) contested Edgeworth's general conclusions (first published in 1911) 
regarding the derivation of the protractor pectoralis in elasmobranchiomorphs. In his 
opinion (based on personal observations and the literature available) the muscle in 
selachians '. . . is simply a differentiation of the constrictor superficialis of the ultimate 
branchial arch'. Allis took further support for his views from Dohrn's (1884; 1885) failure to 
find that the muscle in selachians was developed from the dorsal ends of all the branchial 
muscle plates. 

This discrepancy between the viewpoints of Allis and Edgeworth has, as far as we can 
determine, never been resolved. It is of interest to note that in Edgeworth's 1935 account of 
the muscle and its development he entirely ignores Allis' comments. In part, their difference 
in viewpoints may be attributable to the fact that Edgeworth's approach was primarily 
embryological whilst Allis' first-hand information was derived from the dissection of 
post-embryonic fishes. Further and more critical work needs to be done before the problem 
can be resolved. 


Until this uncertainty has been eliminated we must have some reservations about 
accepting the homology of the protractor pectoralis in elasmobranchiomorphs with the 
muscle identified as the protractor pectoralis in lower Teleostomi. 

As far as the teleostome fishes are concerned, the results of both Allis' and Edgeworth's 
reviews strongly indicate the homology of the muscle throughout the group, at least with 
respect to those members of the Dipnoi and Actinopterygii which they had studied or which 
had been reported upon by others. 

Winterbottom (1974 : 269) has clarified the confusion surrounding Edgeworth's (1935 : 
131 & 143) account of the muscle in teleosts, especially the confusion stemming from 
his use of two names for a morphologically identical muscle. From our own experience we 
would endorse fully the conclusions reached by Winterbottom, and we therefore also treat 
Edgeworth's cucullaris and cephaloclavicularis muscles as synonymous, and thus, in turn, as 
a synonym of Winterbottom's protractor pectoralis muscle. We have also been able to check 
and clarify certain claims for the absence of the muscle, made by Allis (1917) and Vetter 
( 1 878). These are discussed below (p. 228). 

None of our anatomical investigations on post-larval specimens would seem to refute the 
hypothesis that the protractor pectoralis muscle is homologous throughout the lower 
gnathostomes (with, of course, the reservations noted above regarding that muscle in the 
Elasmobranchiomorphi). We are very conscious that our research has only involved an 
investigation of post-embryonic material; the absence of detailed ontogenetic studies is a 
serious drawback in any attempt to investigate homologies. 

Since Winterbottom (1974 : 269) expressed some uncertainty about the homology of the 
muscle he called the protractor pectoralis in the Clupeomorpha, that problem will be 
considered now. 

Our observations (Table 1), based on 21 non-engraulid clupeoid taxa (the muscle 
apparently is absent in engraulids), lead us to believe that Winterbottom's difficulty stems 
partly from his mis-identification of a muscle as being entirely the fourth levator externus, 
and partly from the extreme postero-lateral displacement of the origin for the protractor 
pectoralis and consequently its very close association with the cranial insertion point of some 
epaxial body muscles (see Fig. 3). 

We agree with Winterbottom in identifying the thin, strap-like muscle, running from the 
pterotic to the shoulder girdle, as the protractor pectoralis (see Winterbottom, 1974, fig. 24). 
But, we would identify the usually thin, sheet-like but somewhat expanded muscle lying 
ventral to it as the levator posterior muscle and not, as he does, a muscle composed entirely 
of the expanded 4th levator externus; we have not found, even in Clupea harengus, such an 
expansive and continuous muscle sheet lying between the protractor pectoralis and the main 
levator muscle mass as is shown in Winterbottom's figure 24. In other words, we believe that 
Winterbottom included the posterior levator, the 4th levator externus, and some non- 
muscular tissue lying above and between these muscles, in the 'muscle' he identified as the 
4th levator externus. 

In the clupeoids we examined, the protractor pectoralis and the levator posterior share a 
common site of origin and often a common tendon of attachment to that site. Both muscles 
are closely apposed to the surface of the epaxial body muscles which lie dorsal and medial to 
them, and both are often difficult to locate. 

The clupeoid fourth levator externus muscle is much thicker and more nearly spindle- 
shaped than are the protractor pectoralis and posterior levator muscles. Its origin is 
contiguous with those of the other levators and is thus well separated from the origin of the 
posterior levator. It inserts on the dorso-medial angle of the enlarged 4th epibranchial. In 
contrast, the posterior levator inserts broadly along and behind almost the entire dorsal 
margin of that epibranchial. 

As in many other teleosts, the origin of the presumed protractor pectoralis in clupeoids is 
closely associated with the origin of the levator posterior, the muscle is well separated from 
the origins of the levatores externi and interni muscles, and it inserts onto the cleithrum. We 
therefore consider that, within the terms of reference available to us and other workers, the 


muscle in clupeomorphs is homologous with the muscle identified as a protractor pectoralis 
in other teleosts. 

Clearly, the presence of a protractor pectoralis muscle cannot be taken as a synapomorphy 
for the eurypterygian neoteleosts (see Rosen, 1973), a conclusion which is supported by 
evidence from many other taxa, and which will be discussed further below (p. 232). 
Parenthetically we note that the interrelationships of the protractor pectoralis and the 
levator posterior muscles have been the cause of confusion on previous occasions, as is 
shown by Winterbottom's (1974:269) lucid unravelling of that problem in Edgeworth's 
account of the so-called cucullaris and cephaloclavicularis muscles (Edgeworth, 
1935: 131-143). 

Materials and methods 

The comparative anatomy of the protractor pectoralis muscle was examined in over 250 taxa 
of primitive gnathostomes. In Table I we list the species examined, the presence or absence 
of the protractor pectoralis in each taxon, an indication of whether further comments on the 
anatomy will be given in the text (see pp. 222-232), and the register number(s) for the 
specimen(s) examined. All specimens are from the British Museum (Natural History) fish 
collection unless otherwise noted: the abbreviation 'MCZ' before a number indicates a 
specimen in the collection of the Museum of Comparative Zoology, Harvard University, and 
'I.O.S.' refers to the Institute of Oceanographic Sciences' Discovery collection station 
number. A figure in parenthesis after the register numbers indicates the number of specimens 
which were examined from that particular lot. 

The protractor pectoralis muscle is often extremely thin, and great care must be exercised 
when dissecting to prevent accidental removal of the muscle. In particular, removal of gill 
filaments from the arches, and the superficial fascia covering the dorsal branchial muscles 
laterally, should be done with caution. Portions of the pectoral girdle, especially the post- 
temporal, supracleithrum, and dorsal aspect of the cleithrum, were often removed to enable 
obliquus superioris muscle fibres to be distinguished from those of the protractor pectoralis. 

The muscle nomenclature in this paper follows that of Winterbottom (1974) because of its 
general acceptance by investigators of teleostean morphology. We realize, however, that in 
studies of other basal gnathostome groups use of the term 'trapezius' has become common 
for a muscle which we consider to be homologous with the protractor pectoralis. 

Protractor pectoralis Annotations 

in text Register 

Taxon Present Absent (pp. 222-232) number 


Centrophorus sp. (embryo) + 1973.7.12:18-21 

Hexanchus sp. (embryo) + 1973.7.12:1-3 

Scyliorhinus caniculus + 196 1 . 10. 10 : 1-1 1 


Latimeria chalumnae + BMNH uncatalogued 


Neoceratodusforsteri + + 1959.8.11:12 

Protopterus aethiopicus + + 1957.6.11:1-4 



Acipenser schrencki + + 1925.8.6:3 


Calamoichthys calabaricus + + 1894.7.30:14-15 

Polypterus bichir + + 1928.7.3:1 

Polypterus ornatipinnis + + BMNH uncatalogued 

Polypterus senegalus + 1969.3.17:1-3 



Table 1 cont. 


Protractor pectoral is 
Present Absent 


in text 

(pp. 222-232) 



Lepisosteus oculatus 
Lepisosteus osseus 





Arapaima gigas 
Brienomyrus taverni 
Gnathonemus longibarbis 
Gymnarchus niloticus 
Heterotis niloticus 
Hiodon tergisus 
Hyperopisus bebe 
Mormyrops deliciosus 
Mormyrops engystoma 
Mormyrus kannume 
Mormyrus macrophthalmus 
Notopterus kapirat 
Pantodon buchholzi 
Papyrocranus afer 
Papyrocranus afer 
Petrocephalus catostoma 
Xenomystus ni'gri 


Albula vulpes 
Albula vulpes 
Anguilla anguilla 
Elops hawaiensis 
Elops machnata 
Halosaurus guentheri 
Megalops atlanticus 
Nemichthys scolopaceus 
Notacanthus bonaparti 


Alosa pseudoharengus 
Alosa pseudoharengus 
Anchoa hepsetus 
Anchoa spinifer 
Anchoa spinifer 
Brevoortia tyrannus 
Chirocentrus dorab 
Clupea harengus 
Cynothrissa mento 
Denticeps clupeoides 
Dorosoma cepedianum 
Dussumieria acuta 
Dussumieria hasseltii 
Engraulis edentulus 

MCZ 34650 
MCZ uncatalogued 

MCZ uncatalogued 

BMNH uncatalogued 
BMNH uncatalogued 
MCZ 50425 
1976.5.21 : 1-4 
BMNH uncatalogued 
1971.9.28: 15-18 
BMNH uncatalogued 
BMNH uncatalogued 
1977.11.8: 16-18 
BMNH uncatalogued 

MCZ 18064 
1962.6.29: 11^2 
1962.4.3 : 1-25 
1962.3.26: 1-8 
1966.10.14: 1-2 
BMNH uncatalogued 
1968.3.2: 1 
1973.10.29: 161-183 


MCZ unc 













: 1-79 

: 1-24 , 


Table 1 cont. 



Protractor pectoral is 
Present Absent 

in text 
(pp. 222-232) 


Engraulis japonica 
Etrumeus teres 
Gilchristella sp. 
Jenkinsia stolifera 
Nematalosa come 
Neopisthopterus sp. 
Odontognathus panamensis 
Odontognathus panamensis 
Opisthopterus dovi 
Pellonula afzeliusi 
Sardinella jussieu 
Sprattus sprattus 
Stolephorus commersonii 
Stolephorus indicus 
Stolephorus heterolobus 



Chanos chanos 
Chanos chanos 
Gonorynchus gonorynchus 
Phractolaemus ansorgii 


Abramis brama 
Aspius vorax 
Barbus barbus 
Barbus intermedium australis 
Barilius bendelisis 
Capoeta capoeta 
Carassius auratus 
Catostomus commersonii 
Cobitis caspia romanica 
Cyprinus carpio 
Cyprinus carpio 
Gyrinocheilus aymonieri 
Noemacheilus barbatulus 
Notemigonus crysoleucas 
Opsariichthys uncirostris 
Opsariichthys uncirostris 
Opsaridium ubangensis 
Oxygaster anomalura 
Rutilus rutilus 
Tinea tinea 
Alestes nurse 
Alestes rutilus 
Brycon dentex 
Erythrinus erythrinus 
Hoplias malabaricus 

1969.4.22: 1667-1676 


1973.2.9: 1-30 



1974.7.11 : 559-578 


1974.7.11 : 79-123 

1974.6.26: 139-147 



1939.2.21 :5-7 




MCZ uncatalogued 
BMNH uncatalogued 

1974.9.5: 1-31 

1920.1.22: 127-146 

1864.4.11 :41^2 







1977.7.19: 1-4 

MCZ uncatalogued 


1969.6.12: 13-24 

MCZ uncatalogued 





BMNH uncatalogued 


BMNH uncatalogued 
MCZ uncatalogued 
1972.10.17: 1398-1411 
1974.5.22: 154-173 



Table 1 cont. 


Protractor pectoral is 
Present Absent 


in text 

(pp. 222-232) 



Apteronotus albifrons + 

Gymnotus anguillaris + 

Hypopomus artedi + 

Arius heudeloti + 

Diplomystes papillosus + 

Mystus cavisus + 




1971.9.28: 118-121 
MCZ 8290 


Alepocephalus agassizii 
Aplochiton zebra 
Argentina sphyraena 
Argentina sphyraena 
Bathylagus sp. 
Coregonus albula 
Dallia pectoralis 
Dallia pectoralis 
Esox americanus 
Esox niger 
Galaxias auratus 
Galaxias brevipinnis 
Galaxias maculatus 
Galaxias vulgaris 
Galaxias waitei 
Galaxias weedoni 
Novumbra hubbsi 
Opisthoproctus soleatus 
Osmerus eperlanus 
Osmerus mordax 
Plecoglossus altivelis 
Plecoglossus altivelis 
Retropinna retropinna 
Retropinna retropinna 
Salmo trutta 
Thymallus thymallus 
Umbra krameri 
Umbra limi 
Umbra pygmaea 

1977.6.23: 1-6 
1912.12.20: 18-19 
1971.7.21 : 22-24 
1906.12.5: 1-3 
MCZ uncatalogued 
1883.12.14: 172 
MCZ uncatalogued 
1972.1.27: 15-18 
1972.1.27: 10-14 
1965.10.19: 17-23 
1934.12.19: 1 
1979.11.26: 11-223 
1963.10.28: 16-25 
1923.2.26: 121 
1930.2.5: 1 
1936.11.13: 1-2 
1883.12.14: 172 
MCZ 33 124 


Astronesthes lucifer 
Astronesthes niger 
Chauliodus sloani 
Diplophos taenia 
Gonostoma elongatum 
Photichthys argenteus 
Sternoptyx diaphana 
Stomias boa 

1922.6.7: 14-23 
MCZ 52868 
1972.2.22: 17-19 
MCZ uncatalogued 
MCZ 42 184 
BMNH uncatalogued 


Table 1 cont. 



Protractor pectoral is 
Present Absent 

in text 
(pp. 222-232) 



A ulopusfila mentosus 
Bathysaurus agassizi 
Chlorophthalmus agassizi 
Evermanella atrata 
Evermanella balbo 
Evermanella indica 
Evermanella normalops 
Harpadon macrochir 
Paralepis elongata 
Saurida undosquamis 


Diaphus fragilis 
Electrona antarctica 
Gymnoscopelus aphya 
Lampadena speculigera 
Neoscopelus microchir 
Notoscopelus kroyeri 
Scopelopsis multipunctatus 
Scopelus humboldti 


Amblyopsis spelaea 
Antennarius altipinnis 
Aphredoderus sayanus 
Carapus acus 
Coelorhynchus occa 
Gadus ogac 
Gobiesox papillifer 
Lophius budegassa 
Lycodonus mirabilis 
Malacocephalus laevis 
Muraenolepis microps 
Ophiodon rochei 
Percopsis omiscomaycus 
Porichthys notatus 
Porichthys porosissimus 



Anableps anableps 
Aphanius dispar 
Atherina duodecimalis 
Cyprinodon pecosensis 
Exocoetus obtusirostris 
Fundulus heteroclitus 
Lamprichthys tanganicanus 
Melanotaenia nigrans 
Menidia menidia 
Neostethus lankesteri 

1953.11.1 : 10-13 
MCZ uncatalogued (2) 
MCZ 40530 
I.O.S. 5420 
I.O.S. 7824, 24 
I.O.S. 4947 
I.O.S. 7089, 24 
MCZ 44232 
MCZ 43 140 
MCZ 561 11 

MCZ uncatalogued 
1970.8.11: 1-2 
1970.8.11 : 99-1 12 
1962.1.3: 11-12 
MCZ 55532 

1898.12.29: 141-149 
1952.11.25: 1-4 
1974.6.27: 1-6 
MCZ 52937 
MCZ 44836 
MCZ 38301 
MCZ 44993 
1937.7.12: 11-17 
MCZ 54922 
MCZ uncatalogued 
1961.9.4: 171-173 

1973.9.10: 154-194 
1978.8.1 : 303-502 
MCZ 42538 

1955.12.20: 1449-1465 
1975.3.20: 119-186 
MCZ uncatalogued 



Table 1 cont. 


Protractor pectoral is 
Present Absent 

in text 
(pp. 222-232) 


Oryzias latipes + 

Parexocoetus brachypterus + 

Poecilia reticulata + 

Potamorrhaphis guianensis + 

Scomberesox saurus + 

Ammodytes hexapterus + 

Anabas testudineus + 

Archamia zosterophora + 

Ariomma indicus + 

Aulostomus chinensis + 

Bothuspodas + 

Brachydeuterus auritus + 

Callionymus lyra + 

Capros aper + 

Caranx malabaricus + 

Centropyge bispinosus + 

Congiopodus perorianus + 

Cottus gobio + 

Cyclopterus lumpus + 

Dactylopterus volitans + 

Epinephelus aeneus + 

Eupomacentrus fasciolatus + 

Genes poeti + 

Gobius niger + 

Grammistes sexlineatus + 

Gymnocephalus cernua + 

Holocentrus spinifer + 

Holocentrus suborbitalis + 

Hoplichthys acanthopleurus + 

Hoplolatilus starcki + 

Kutajlammeo sammara + 

Kyphosus cuierasceus + 

Lates microlepis + 

Lepomis auritus + 

Liparis liparis + 

Lutjanus synagris + 

Macrorhamphosus gracilis + 
Mastacembelus albomaculatus + 

Monocirrhus polyacanthus + 

Monopterus albus + 

Mugil cephalus + 

Nandus nebulosus + 

Notothenia larseni + 

Ophioblennius steindachneri + 

Ostichthys murdjan + 

Pachypopsfourcroi + 

Parapercis cephalopunctata + 

Pelates quadrilineatus + 

Phanerodonfurcatus + 

Platycephalus mulleri + 

Pleuronectes platessa + 

1923.2.26: 160-169 
1967.2.1 : 53-57 
1972.7.27: 1014-1019 
BMNH uncatalogued 

1974.5.25: 1548-1560 
1962.9.18: 109-117 
1962.6.1 : 15-29 
BMNH uncatalogued 
1936.8.26: 1100-1104 
1968.12.31 :2 
1967.2.1 : 308-3 11 

1967.2.1 : 73-76 
1971.2.16: 1072-1081 
1951.1.16: 145-147 
1961.4.19: 106-115 
1960.3.15: 170-172 
MCZ 43537 
1939.5.24: 1684-1695 
BMNH uncatalogued 
1973.1.22: 105-112 
MCZ 492 12 

MCZ 460 17 

1976.4.2 : 77-83 












1979.10.16: 16-29 
1971.2.16: 1992-1993 


Table 1 cont. 



Protractor pectoralis Annotations 

in text Register 

Present Absent (pp. 222-232) number 

Polymixia nobilis 
Pomacanthus arctifrons 
Psettodes erumei 
Scomber japonicus 
Scorpaenodes insularis 
Sebastes crameri 
Solea solea 

Sphyraena chrysotaenia 
Stephanoberyx monae 
Synbranchus marmoratus 
Syngnathus acus 
Tautogolabrus sp. 
Thalassoma purpureum 
Trachipterus taenia 
Trichogaster pectoralis 
Uranoscopus scaber 
Zaniolepis latipinnis 

1862.4.22: 17-18 


1933.7.31 : 1-2 

1967.2.1 :41^4 



1971.2.16: 118-127 





BMNH uncatalogued 


1891.8.31 : 27-35 





1971.7.21 : 86-90 

Comments on the protractor pectoralis muscle in certain taxa 


In selachians the protractor pectoralis (=trapezius or cucullaris of authors) originates from 
the fascia covering the epaxial muscles laterally (Fig. 1). The fibres extend postero ventral ly 
to insert on the pectoral girdle. Medial to the protractor pectoralis, another muscle, which 
appears to be an epaxialis derivative, extends anteriorly from the dorsal aspect of the 
pectoral girdle to merge with epaxial muscle fibres. The fibres of this muscle lie at 90 to 
those of the protractor pectoralis. 

Anterior to the protractor pectoralis a muscle runs posteroventrally from the epaxialis 
fascia to insert on the posterodorsal aspect of the last gill arch (Fig. 1 ). We consider this 
muscle to represent an anterior division of the protractor pectoralis, as did Allis (1917) and 
Edgeworth(1935: 141). 

The protractor pectoralis lies medial to the branchial constrictor muscles although both 
share a common origin from the epaxial muscle fascia. Levator arcuum branchialum 
muscles are absent in selachians. 


Millot & Anthony (1958 : 63, fig. 29) do not describe a protractor pectoralis muscle (or a 
muscle with the anatomical relationships of the protractor) in adult Latimeria chalumnae. 
We have dissected and made observations on a 32 cm foetus, and also find no trace of the 


Neoceratodus forsteri. The protractor pectoralis is a well-developed and distally expansive 
muscle (Fig. 2). Its narrow origin lies immediately posterior to the common origin of the 3rd 


and 4th levator extern! muscles; it has a wide, musculose insertion onto the cleithrum, its 
antero-posterior orientation being almost in the sagittal plane. 

From its apparent anterior face some fibres attach dorsally to the posterior aspect of the 
5th gill arch, immediately above the origin of the subarcualis rectus of Wiley (1979 : fig. 4). 
Unlike Wiley, we cannot identify a separate pharyngoclavicularis internus muscle associated 
with the 5th gill arch. Possibly the muscle he illustrates is what, in our specimen, appears to 
be the anterior portion of the protractor pectoralis. In other words, the two muscles in this 
specimen are so closely contiguous that they give the appearance of a single muscle. The 
fibre orientation of this anterior muscle (if it be separate) is like that in the protractor 
pectoralis itself, and its attachment to the cleithrum is medial and ventral to the attachment 
area of the pharyngoclavicularis muscles from arches 1-4. 

Protopterus aethiopicus. A protractor muscle, originating from the posterior margin of the 
cartilaginous skull broadens out from its narrow point of origin to insert, tendinously, on the 
cleithrum. The muscle, as compared with that in Neoceratodus, is narrow and instead of 
having a simple inverted fan-shape has a slightly concave and twisted anterior margin which, 
again unlike Neoceratodus, is aligned almost at right angles to the sagittal plane; posteriorly a 
short length of the muscle lies in that plane. Continuous with the muscle's posterior margin 
are several much more expansive muscle bundles. These are apparently derived from the 
hypaxial body musculature; dorsally the muscles attach to the horizontal septum, and 
posteriorly they continue beyond the girdle but give off fibres which insert on the anterior 
face of the cleithrum as the main muscle mass passes below that bone. Careful dissection 
shows that the apparent continuity of fibres from the branchial protractor pectoralis with 
those of the hypaxial pseudo-protractor is in fact false; the two muscles are separated by a 
very narrow hiatus obscured by the rather dense tissue overlying them. 


Acipenser schrencki. The very well-developed, broad and thick protractor pectoralis 
originates along the posterior transverse region of the skull and inserts on the cleithrum. Its 
origin is clearly separated from that of the muscle plate representing the levators of the 3rd 
and 4th gill arches. 

Sewertzoffs (1928) embryological studies of the cranial muscles in Acipenser ruthenus 
clearly demonstrate the common origin of the branchial levators, and the derivation of the 
protractor pectoralis, from the posterior part of the same plate. 

The muscle's ontogenetic history in other chondrosteans is less clear (see Edgeworth, 
1935 : 142) and would repay further investigations on both embryological and adult 
material. According to Edgeworth (loc. cit.) the muscle is absent in A. sturio '. . . possibly by 
atrophy during developmental stages owing to the fixation of the pectoral girdle'. 


Polypterus ornatipinnis. The protractor in this species in a thin, largely tendinous muscle 
which shares its origin with the 3rd and 4th levatores externi, and inserts on the cleithrum 
near its point of maximum curvature. 

Calamoichthys calabaricus. In this species, unlike P. ornatipinnis, the protractor is large 
and noticeably broader distally than proximally. As in P. ornatipinnis it has a common 
origin with the 3rd and 4th levatores externi but inserts onto the cleithrum at a point slightly 
dorsal to the bone's region of maximum curvature. 


Lepisosteus osseus lacks both protractor pectoralis and levator posterior muscles. Just dorsal 
to the origin of the fourth levator externus, the obliquus superioris inserts tendinously on the 
skull. In Lepisosteus oculatus some lateral fibres of the obliquus superioris insert on a medial 
flange of the supracleithrum, but no protractor pectoralis is present. 



Amia calva possesses a well-developed protractor pectoralis (=fifth levator externus of Allis, 
1897). The protractor pectoralis originates from the otic region of the skull and inserts via a 
long tendon onto the cleithrum. 


The osteoglossomorph fishes lack both a protractor pectoralis (Table 1) and a levator 
posterior muscle. Winterbottom (1974 : 252) noted that the levator posterior is confined to 
the neoteleosts (in his usage of the category) with the exception of Hiodon. We find that 
Hiodon lacks the levator posterior and possesses a posteriorly displaced origin of the fourth 
levator externus due to the large swimbladder extension in the otic region. 


Some comments on the protractor pectoralis of clupeomorph fishes have been made already 
(p. 2 15). 

In none of the clupeid and chirocentrid species we have examined is the muscle well- 
developed; usually it is mainly tendinous, flat and narrow, and invariably it is closely applied 
to the ventro-anterior face of the overlying body muscles. Often part of the protractor 
pectoralis inserts on the supracleithrum. 

A protractor pectoralis muscle is absent in all the engraulid species examined, and 
apparently, in the sole extant representative of the Denticipitoidei, Denticeps clupeoides. 
However, because the only specimens of Denticeps available are small (ca 50 mm SL), a 
narrow, thin muscle could easily be overlooked. The muscle is present in a large proportion 
of the Clupeidae (sensu law) we examined (Fig. 3). 

A levator posterior muscle occurs in the majority of clupeomorph taxa we dissected. Its 
origin is shared with, or is very close to, that of the protractor pectoralis. In general it too is a 
thin, narrow and partly tendinous muscle, and is always closely applied to the ventro- 
anterior face of the body musculature (Fig. 3). 

No levator posterior was found in the engraulid Anchoa spinifer (although the muscle is 
present in A. heterolobus). It also appears to be absent in the chirocentrid Chirocentrus 
dorab, unless, atypically, it is closely associated with the 4th external levator muscle which, 
in this species, seems to be composed of two very closely contiguous parts. We are uncertain 
about the condition in the denticipitoids because of the small size of the available specimens. 

The presence of a levator posterior in many clupeomorph fishes contradicts 
Winterbottom's (1974:252) statement about the occurrence of that muscle within the 
teleosts, namely, that apart from its occurrence in the osteoglossomorph Hiodon, it is a 
neoteleostean feature (see also above). It also contradicts his suggestions about a 
possible origin for the posterior levator from a condition like that supposedly occurring in 
the clupeomorphs (Winterbottom, loc. cit.). As noted already (p. 215), and in contradistinc- 
tion to Winterbottom's observations, none of the clupeoids we examined showed any 
continuity of fibres between the 4th external levator and '. . . a thin sheet of muscle whose 
origin extends to the posterolateral tip of the pterotic' (the condition which Winterbottom 
considered to be the typical clupeomorph one). Our interpretation of the clupeomorph 
condition is that part of Winterbottom's 'thin sheet of muscle . . .' is the levator posterior 
muscle, but that the area of tissue between it and the 4th levator externus is devoid of muscle 
fibres, and consequently the two muscles are distinct from one another (see also p. 2 1 5). 


A protractor pectoralis is present in some or all taxa of every otophysan subdivision (see 
Table 1 ; also Winterbottom, 1 974; figs 20, 2 1 & 22), and apparently is absent only in certain 



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cypriniform taxa. It is well-developed in the three members of the anotophysi we dissected, 
particularly in Chanos chanos where it is a thick, deep muscle proximally, and has a broad, 
tendinous insertion onto the upper half of the cleithrum; in Gonorynchus gonorynchus it is 
narrow and strap-like, inserting directly onto the girdle, while in Phractolaemus ansorgii it is 
broad, thin and partly tendinous. 

Amongst those otophysans with a protractor pectoralis, the muscle, although intra- 
specifically constant, shows varying degrees of development; also, in some species it inserts 
directly onto the cleithrum, whereas in others it inserts onto the membrane extending from 
the girdle to form the posterior wall of the branchial chamber. When the protractor has a 
'membrane insertion' it is closely applied to the anterior face of the body musculature which 
delimits the posterior boundary of the branchial chamber. In general, the protractor 
pectoralis is narrow and strap-like, and often has a near-vertical orientation. 

Vetter (1878) claims that the protractor pectoralis (his trapezius), is absent in Cyprinus 
carpio and Barbus barbus. The muscle certainly is present in C. carpio (see Winterbottom, 
1974, fig. 22; personal observations), but may have been overlooked by Vetter because of its 
slenderness and its rather tendinous nature. A short, fine, protractor pectoralis is also present 
in a specimen of Barbus barbus we dissected, although the muscle is absent in another 
member of that genus, B. intermedius, from Kenya, east Africa. 

A levator posterior muscle appears to be absent in some anotophysans (Chanos chanos), 
but is present in Gonorynchus gonorynchus. Its absence in Chanos could be correlated with 
the development of an expansive and complex suprabranchial organ in that genus. The 
muscle is seemingly also absent in Phractolaemus ansorgii, but the condition of our material 
does not permit a definite conclusion on that point. 

All the otophysan taxa examined have a levator posterior muscle; it is particularly well- 
developed in members of the Cypriniformes (see also Winterbottom, 1974; 252-253, figs 
20-22), but in other groups it is often a slender strap-like muscle. 

The occurrence of a levator posterior muscle in the Ostariophysi further negates 
Winterbottom's claim that its presence is a neoteleostean feature (see above [Clupeo- 
morpha], and also p. 2 1 3 regarding that author's handling of the category Ostariophysi). 

Brousseau (19760 & b) has described parts of the branchial musculature in six 
ostariophysan taxa. It is clear from his descriptions and figures that, depending on the species 
involved, he has either misidentified the protractor pectoralis, or confused and compounded 
it with the levator posterior muscle. 


The distribution of the protractor pectoralis in this group is exceedingly irregular, with, in 
addition, the muscle occurring in relatively few taxa. As an example of its irregular 
occurrence, we may note its presence in some but not in other Galaxias species, and its 
presence in Dallia pectoralis (Fig. 4) but not in Umbra limi, U. krameri or Novumbra 

None of the protacanthopterygian species we examined has a levator posterior muscle, and 
we can find no reference in the literature to its occurrence in these fishes (see also 
Winterbottom, 1974:252-253). 


Lauder & Lanyon (1980: fig. 2) identified as a protractor pectoralis in Lepomis macrochirus 
a muscle inserting on the supracleithrum and posttemporal, and extending anteriorly to 
originate on the posterodorsal aspect of the skull. This muscle is apparently a derivative of 
the epaxialis and is not homologous with the protractor pectoralis as defined in this paper. 
The true protractor pectoralis in Lepomis (Fig. 5) originates dorsally from the pterotic 
adjacent to the origin of the levator posterior, and extends posteroventrally to fan out and 
insert in the connective tissue between the last gill arch and the cleithrum. No fibres of the 
protractor pectoralis contact the cleithrum. 

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Fig. 5 Lepomis auritus (lateral view, left side; gill filaments removed). Cl: cleithrum. Lm 1-4: 
levator muscles of lst-4th gill arches. Lp: levator posterior muscle. Mccl: membrane overlying 
cleithrum and distal part of pharyngocleithralis externus muscle. Phce & Phci: pharyngo- 
cleithralis internus and externus muscles, respectively. Pp: protractor pectoralis muscle. Rcc: 
rectus communis muscle. Scl: supracleithrum. Drawn from specimen BMNH 1973.1.22: 

Many other acanthopterygians also possess a thin protractor pectoralis muscle which does 
not insert directly on the cleithrum (eg. Zeus, Nandus, Trachipterus, Macrorhamphosus, 
Sebastes). Another common condition in the Acanthopterygii is the presence of a protractor 
pectoralis inserting either directly onto the cleithrum (eg. most atherinomorphs, cichlids, 
Platycephalus, Liparis, Hoplolatilus), or inserting onto both the connective tissue posterior 
to the 4th ceratobranchial and the cleithrum (eg. Lates, Epinephalus, Grammistes, Genes). 

In both the synbranchiform taxa examined, the protractor pectoralis is, however, a well- 
developed, strap-like muscle running horizontally from the skull to the pectoral girdle. In 
Synbranchus marmoratus (Fig. 6) its origin is shared with that of the branchial levator 
muscles, and it inserts onto the small, moveable supracleithrum. An insertion onto the 
cleithrum itself is precluded by the position of the gill arches, which are so positioned that 


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the path of the protractor is directed so as to lie above the dorsal extremity of the cleithrum. 
The origin of the protractor in Monopterus albus lies posterior to and well-separated from 
that of the levators, and its orientation is such that it inserts onto the dorsal tip of the 
cleithrum. The dorsoposterior margin of the protractor pectoralis is closely applied to 
the ventral margin of the supracleithrum, but no fibres from the muscle are inserted onto the 

The muscle in Lepomis identified by Lauder & Lanyon (1980, fig. 2) as a protractor 
pectoralis (see above), is of uncertain homology. It appears to be similar to the levator 
pectoralis of Winterbottom (1974: 270, and fig. 25) which has been found in batrachoids, 
lophiids, and tetraodontiforms. We have also found a similar muscle to the levator pectoralis 
in scorpaeni forms and nandids, and it may have a much wider distribution than is presently 

Summary and conclusions 

Since our interest in the protractor pectoralis was stimulated initially by irregularities in its 
presumed phylogenetically based pattern of distribution within the Teleostei (see p. 213), 
our principal concern is with the effects additional data have on the phylogenetic 
conclusions reached previously (Rosen, 1973; Winterbottom, 1974). 

Clearly, because of its occurrence in some or even many members of the Clupeomorpha, 
Ostariophysi and Protacanthopterygii, the protractor pectoralis is not, as was once thought, a 
feature restricted to the Neoteleostei (see pp. 217-219). Indeed, its overall pattern of 
occurrence extends beyond the current limits of the Euteleostei (of which the Neoteleostei 
are a major subdivision; Rosen, 1973; Patterson & Rosen, 1977) to include the Teleostei 
(sensu Patterson & Rosen) as a whole. 

Taking the argument further: if one accepts the supposed homology of the muscle 
throughout the gnathostomes, the protractor pectoralis occurs in some or all members of the 
Neopterygii, Palaeopterygii, Dipnoi and, very probably, the Elasmobranchiomorphi as well 
(see pp. 214; 216-221). Amongst the major groups making up the lower gnathostomes, a 
protractor pectoralis is absent only in the extant Actinistia, a group represented by the single 
taxon Latimeria chalumnae. 

The protractor pectoralis, therefore, would seem to be a synapomorphy of the 
Gnathostomata and thus, within the various lineages of that group, its presence must be 
treated as a plesiomorph character and not as a synapomorphy of the Teleostei in part. 

Its distribution among the lower gnathostomes has a somewhat patchy pattern (see 
Table 1), a pattern whose patchiness is, in some respects, even more marked and more 
puzzling within the Teleostei. 

A protractor pectoralis is not developed in the two lineages which are generally thought to 
be the most primitive amongst living teleosts, namely the Osteoglossomorpha and the 
Elopomorpha. Patterson & Rosen (1977) for example, consider the Osteoglossomorpha to 
be the plesiomorph sister group of all other living teleosts combined (ie the Elopocephala), 
and the Elopomorpha to be the plesiomorph sister group of the other Elopocephala (ie the 
Clupeocephala). The muscle is also absent in one 'higher' group, the Stomiatiformes, which, 
on the basis of its having certain derived characters, Rosen (1973: 505) identified as the 
plesiomorph sister group of all other lineages he brought together as the Neoteleostei (see 
also p. 213). Within those other lineages (ie the Aulopiformes, Myctophiformes, 
Paracanthopterygii and Acanthopterygii) a protractor pectoralis has been found in virtually 
all taxa placed in the supposedly 'higher' categories Paracanthopterygii and Acanthop- 
terygii, 1 in the majority of Aulopiformes, and in many Myctophiformes as well (See Table I; 
also Winterbottom, 1974). 

The exceptional taxon is Stephanoberyx monae; no protractor was found in the two specimens we examined. 


Among lineages which lie, cladistically speaking, between the plesiomorph Osteoglosso- 
morpha + Elopomorpha on the one hand and the apomorphic Neoteleostei (sensu Rosen) on 
the other, the muscle occurs in most members of the Clupeomorpha and in many 
Ostariophysi (including the plesiomorphic Anotophysi), but is found in only a few members 
of the Protacanthopterygii. 

An intriguing aspect of this pattern is the contrast between the absence of a protractor 
pectoralis (a derived condition) in the two most primitive lineages, and its presence (the 
primitive condition) in the more derived ones. Equally intriguing is the way the muscle, in 
taxa belonging to the 'intermediate' groups, may be absent in some species of a genus but not 
in others (for example in the protacanthopterygian genus Galaxias], or in some but not all 
members of seemingly closely related taxa (for example, again amongst the protacanthop- 
terygians, its presence in Dallia, but its absence in Umbra and Novumbrd). Similar patterns 
of presence or absence can be found within the Clupeomorpha and the Ostariophysi (see 
Table 1). 

We can offer no explanation for these patterns, and must conclude that the presence or 
absence of a protractor pectoralis muscle is a feature of little value as an indicator of phyletic 
relationship except at a high level of universality. In other words, it is a synapomorphy of the 

Within the Teleostei we are impressed by the constancy of its presence in the 
Paracanthopterygii and Acanthopterygii, and by the constancy of its absence in some other 
groups (for example, the Osteoglossomorpha). This pattern would suggest to us that in these 
lineages the ontogenetic canalization leading either to the development or to the suppression 
of a protractor pectoralis was fixed very early in the history of each lineage. In contrast, the 
irregular patterns seen in other lineages (for example the Ostariophysi and Protacanthop- 
terygii) would seem to indicate the retention of a flexible linkage between the pathway of 
protractor pectoralis ontogeny and other elements of the total ontogenetic pattern. What 
significance, if any, this may have in tracing phyletic histories remains obscure. 

All our conclusions (and speculations) are, of course, dependent on the hypothesis that the 
muscle is an homologous (ie synapomorphic) feature within the Gnathostomata. Our 
investigations provide no refutation of that hypothesis, but we are of the opinion that the 
only critical test will come from comparative embryological and ontogenetic studies on a 
larger scale than has been carried out so far. 


We are most grateful for the assistance given to us by our colleagues in the Museum of 
Comparative Zoology and the British Museum (Natural History), especially Karel Liem, 
William and Sara Fink, Keith Banister and Gordon Howes (who also drew the figures). 

This investigation began when the senior author was a Agassiz Visiting Professor at the 
Museum of Comparative Zoology. He is much indebted to that Museum for providing 
financial assistance for his visit, and for providing such congenial working conditions when 
he was there. The junior author gratefully acknowledges the Society of Fellows (Harvard 
University) for a Junior Fellowship, and a Penrose Fund award (grant 8636) from the 
American Philosophical Society. 


Allis, E. P. 1897. The cranial muscles and cranial and first spinal nerves in Amia calva. J. Morph. 

12 : 487-808. 
1917. The homologies of the muscles related to the visceral arches of the gnathostome fishes. Q. Jl. 

Microsc. Sci. 62 : 303^06. 
Brousseau, R. A. \916a. The pectoral anatomy of selected Ostariophysi. I. The Characiniformes. 

J. Morph. 148:89-135. 
\916b. The pectoral anatomy of selected Ostariophysi. II. The Cypriniformes and Siluriformes. 

J. Morph. 150:79-115. 


Dohrn, A. 1884. Studien zur Urgeschichte des Wirbeltierkorpers IV. Mitt. zool. Stn Neapel 
5: 102-151. 

1885. Studien zur Urgeschichte des Wirbeltierkorpers VII. Mitt. zool. Stn Neapel 6 : 

Edgeworth, F. H. 1911. On the morphology of the cranial muscles in some vertebrates. Q. Jl. Microsc. 
Sci.56: 167-316. 

- 1935. The cranial muscles of vertebrates. Cambridge. 

Greenwood, P. H., Rosen, D. E., Weitzman, S H. & Myers, G. S. 1966. Phyletic studies of teleostean 

fishes, with a provisional classification of living forms. Bull. Am. Mus. nat. Hist. 131 : 339-456. 
Lauder, G. V. & Lanyon, L. E. 1980. Functional anatomy of feeding in the bluegill sunfish, 

Lepomis macrochirus: in vivo measurement of bone strain. J. exp. Biol. 84 : 33-55. 
Millot, J. &. Anthony, J. 1 958. Anatomie de Latimeria chalumnae. 1 : 1 1 8 pp. C.N.R.S., Paris. 
Patterson, C. & Rosen, D. E. 1977. Review of ichthyodectiform and other Mesozoic teleost fishes and 

the theory and practice of classifying fossils. Bull. Am. Mus. nat. Hist. 158 : 8 1-1 72. 
Rosen, D. E. 1973. Interrelationships of higher euteleostean fishes. Zool. J. Linn. Soc. 53, Supp. 

1 -.397-513. 

- & Patterson, C. 1969. The structure and relationships of paracanthopterygian fishes. Bull. Am. 
Mus. nat. Hist. 141 : 357-474. 

Sewertzoff, A. N. 1928. The head skeleton and muscles of Acipenser ruthenus. Acta zool, Stockh. 

9: 193-319. 
Vetter, B. 1878. Untersuchungen zur vergleichenden Anatomie der Kiemen-und Kiefermusculatur der 

Fische, II. Jena Z Naturw. 12 : 431-550. 
Wiley, E. O. 1979. Ventral gill arch muscles and the interrelationships of gnathostomes, with a new 

classification of the Vertebrata. Zool. J. Linn. Soc. 67 : 149-179. 
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nat. Sci. Philad. 125 : 225-317. 

Manuscript accepted for publication 9 January 198 1 

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the research carried out "behind the scenes". 

Titles to be published in Volume 41 

Anatomy and phytogeny of the Chinese Major Carps 
Ctenopharyngodon Steind., 1866 and Hypophthalmichthys Blkr., 
1860. By Gordon Howes 

Morphology and systematics of some interior-walled cheilostome 
Bryozoa. By P. L. Cook and P. J. Chimonides 

Notes on Atlantic and other Asteroidea. 1. Family Benthopectinidae. 

By Ailsa M. Clark 


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Bulletin of the 

British Museum (Natural Histo 


Zoology series Vol 41 No 5 17 December 1981 

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ISSN 0007-1498 Zoology series 

Vol 41 No 5 pp 235-294 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 17 December 1981 



The ostracod genus Loxoconcha from Abu Dhabi lagoon, Persian Gulf. 
By R. H. Bate & A. Gurney 

A revision of the spider genus Cocalus (Araneae: Salticidae). By F. R. Wanless 
The Phthiracarus species of C. L. Koch. By B. W. Kamill 

The status of Lamingtona lophorhina McKean & Calaby, 1968 
(Chiroptera: Vespertilionidae). By J. E. Hill & K. F. Koopman . 

The status of Hipposideros galeritus Cantor, 1 846 and Hipposideros cervinus 
(Gould, 1 854) (Chiroptera: Hipposideridae). By P. D. Jenkins & J. E. Hill 




The ostracod genus Loxoconcha Sars from 
Abu Dhabi lagoon and the neighbouring near-shore 
shelf, Persian Gulf 

Raymond H. Bate & Ann Gurney 

Departments of Palaeontology and Zoology, British Museum (Natural History), Cromwell 
Road, London SW7 5BD 


The present study on the genus Loxoconcha from the region of Abu Dhabi lagoon (Lat. 
24 32' N, Long. 54 21' E), follows a number of ecological and taxonomic papers (see Bate 
1971, 1973; Bate & Sheppard, 1980; and Gurney 1979a & b) that have been based on 
material collected by Dr Graham Evans of Imperial College, London, during the period 
1961-65. This material, collected during an investigation of the Recent sediments of the 
Trucial coast, has been supplemented, for the ostracod studies, with samples collected by 
Professor John Murray (Exeter) in 1969. 

Elsewhere in the Gulf, Sabeekah Razzaq (1979) has undertaken a preliminary study of the 
benthic microfauna of Kuwait while Paik (1976 & 1977) has examined the ostracod faunas 
on the Iranian side of the Gulf as well as from the Gulf of Oman. Other papers relevant to our 
study are those of Hartmann 1964, on ostracods from the Red Sea; Jain 1978, on the fauna of 
Mandvi Beach, west coast of India and Bhatia 1979, on part of the fauna from off Karwar, 
west coast of India. 

Four species of Loxoconcha (Loxoconcha) and one of the new subgenus Epakroconcha are 
described here; it is the purpose of this paper not only to place these on record but to discuss 
the relevance of the ornamental differences (ranging from coarsely reticulate to finely pitted) 
that are exhibited between the species. 

Systematic descriptions 

Genus LOXOCONCHA Sars, 1866 

REMARKS. In Bate et al. (in press), the genus Loxoconcha was divided into two subgenera: 
Loxoconcha and Loxocorniculum. Here a third subgenus Epakroconcha is recognized. Of 
the five species described from Abu Dhabi lagoon and the Persian Gulf, four belong to the 
subgenus Loxoconcha, being forms having a clear cut dimorphism of subquadrate females 
and subrectangular males. The fifth species differs by having a subtrigonal outline for both 
males and females and is accordingly placed in the separate subgenus Epakroconcha. 

Subgenus LOXOCONCHA Sars, 1866 

REMARKS. The four species described here differ ornamentally by having either a reticulate 
or a pitted shell surface. We have grouped the species according to this and, as will be 
discussed under environment, we show that this has a bearing on their distribution either 
within the lagoon or outside on the nearshore shelf. 

Bull. Br. Mus. nat. Hist. (Zool.) 41 (5): 235-25 1 Issued 1 7 December 1 98 1 


236 R. H. BATE& A. GURNEY 

Reticulate species 

Loxoconcha (Loxoconcha) multiornata sp. nov. 

Loxoconcha ornatovalvae Hartmann; Bate 1971 : 245, 246, 248, 250, pi. 1, figs. Ik & 2k, pi. 2, fig. 3k, 

pi. 3, figs. 2k & 3k. 
Loxoconcha sp. A, Jain 1978 : 126, fig. 5 A. 

HOLOTYPE. BM(NH) no. 1980.236, cf carapace, sample 65 14, central lagoon terrace. 
PARATYPES. BM(NH)nos 1980.237-243. 

TYPE LOCALITY. Sample 6514 Central lagoon terrace, shallow marine, tidal; salinity 

DIAGNOSIS. Species of Loxoconcha having very coarsely reticulate ornamentation (as 
illustrated) with some reticulae developed into ridges; dorsal ridge of carapace characteristic- 
ally producing pear-shaped pattern. 

DESCRIPTION. Species dimorphic, the males more elongate than the females. Ornamentally 
the reticulae are very coarsely developed and may even be produced as low ridges. The eye 
node is developed as a swelling situated on a low antero-dorsal ridge. Sieve plate normal 
pores are situated within the reticulae. Ventrally, the edge of the right valve, where it over- 
laps the left, is marked with short ridges while the left valve in the posterior part of the 
carapace possesses four rowlock-type projections (Fig. ID). Hinge typically gongylodont 
with coarsely dentate/loculate median element. Muscle scars with three oval adductor scars 
in a vertical row with a fourth situated almost in front of the lowermost scar. Frontal scar 
V-shaped. Duplicature broad with small antero-ventral and narrow postero-ventral vesti- 
bule. Marginal canals straight, widely spaced and few in number: 9 anteriorly. Outside the 
selvage a broad flange extends around the anterior and along the ventral margin. 

DIMENSIONS. Holotype: 1980.236 cf carapace, length 0'46 mm; height 0'26 mm; width 
0-26 mm. Paratypes: 1980.237 d 1 RV., length 0'47 mm; height 0*26 mm. 1980.238 9 
carapace, length 0'38 mm; height 0'24 mm; width 0'23 mm. 1980.239 9 carapace, length 
0-40 mm; height 0'25 mm; width 0'23 mm. 1980.240 cf carapace, length 0*50 mm; height 
0-26 mm; width 0'28 mm. 1980.241 9 RV., length 0'40 mm; height 0'24 mm. 1980.242 rf 
RV., length 0*44 mm; height 0*25 mm. 

REMARKS. Loxoconcha (Loxoconcha) multiornata sp. nov. is morphologically very close to 
the Red Sea species, Loxoconcha ornatovalvae Hartmann 1964 and was, in fact, identified as 
such by one of us (Bate, 1971). There are, however, significant differences that make 
separation of the two species possible: firstly in dorsal view, the curved dorsal ridge of L. 
multiornata produces a pear-shaped pattern when both valves are together (Fig. IE, F) 
whereas in L. ornatovalvae (Hartmann, 1964, pi. 20, fig. 93) there is a looped ridge pattern 
extending from the eye node back along the dorsal margin. The eye node of the Red Sea 
species is separate from the antero-dorsal ridge, whereas in L. multiornata it is a swollen part 
of the ridge. The differences in the lateral ornamentation are best explained by illustration 
(see Fig. 2). 

Loxoconcha (L.) multiornata has been recorded by Jain 1978 as Loxoconcha sp. A from 
the Recent sediments of Mandvi Beach, west coast of India and is thus known to occur within 
the Indian Ocean area. As yet it has. not been recorded from the African coast. If, as appears 
possible, the Red Sea Loxoconcha ornatovalvae has developed from the Indian Ocean/ 
Persian Gulf L. multiornata, then we are going some way to implying that species give rise to 
similarly ornamented species and this could lead us to a subdivision of the subgenus 
Loxoconcha on basic external morphology. By grouping the five species described in this 
paper on similar ornamentation we are partly suggesting this but we would not claim that the 
groupings here necessarily relate to all the species described here. 



Fig. 1 Loxoconcha (Loxoconcha) multiornata sp. nov.: A, X 125, holotype, cf left side, BM(NH) 
1980.236; B, X 125, paratype 9 RV., specimen lost; C, X 125, paratype cf RV., BM(NH) 
1980.237; D, G, enlargement of ventral margin X 580 and ventral view of carapace X 125, 9 
paratype BM(NH) 1980.239; H, X 1- 12K, sieve plate, specimen lost; E, X 125, paratype <s dorsal 
carapace, BM(NH) 1980.240; F, X 125, paratype 9 dorsal carapace, BM(NH) 1980.239; I, J, X 
345, paratype 9 terminal hinge elements RV, BM(NH) 1 980.24 1 . 

238 R. H. BATE& A. GURNEY 


Fig. 2 Comparison of ornamentation between Loxoconcha (Loxoconcha) multiornata sp. nov. 
[A] and Loxoconcha (Loxoconcha) ornatovalvae Hartmann [B]. 

DISTRIBUTION. Loxoconcha (L.) multiornata is essentially a nearshore shelf species but does 
occur sporadically in the inner, central and outer lagoon areas (Fig. 1 1 ). 

Loxoconcha (Loxoconcha) gurneyi sp. nov. 

Loxoconcha sp. A. (pars), Bate 1971 : 245, 246, 248, pi. 1, figs 11, 21, pi. 2, figs 21, 31 & pi. 3, figs 11, 21 

Loxoconcha sp. B., Jain 1978 : 126, fig. 5B. 

DERIVATION OF NAME. Named in honour of Dr Robert Gurney, 1879-1950, Zoologist, well 
known for his work on Crustacea. 

HOLOTYPE. BM(NH) no. 1980.245, d LV. sample 62 107, nearshore shelf. 
PARATYPES. BM(NH)nos 198.246-251 &431. 

TYPE LOCALITY. Sample 62107, nearshore shelf; shallow marine; salinity 43'75%o; surface 
water temperature 23'0-24-4C. 

DIAGNOSIS. Species of Loxoconcha with neat, honeycomb reticulation. 

DESCRIPTION. Carapace quadrate (9) to rectangular (rf) with straight dorsal margin; sinuous 
ventral margin; broadly rounded anterior and posteroventral margins; posterodorsal margin 
short and concave. Eye node oval, situated below anterior cardinal angle. Entire shell surface 
covered by neat reticulation of somewhat rounded, 5-sided, pits; the pits becoming 4-sided 
around the posterior margin and along the dorsal and ventral surfaces. Ventral margin of 
right valve ornamented by short ridges situated perpendicular to valve edge (Fig. 3B). 
Normal pore canals open, through a sieve plate, on the murae of the reticulation. Hinge, well 
developed gongylodont. Muscle scars with a vertical row of 4 adductors and an oval frontal 
scar (Fig. 3G). Duplicature broad around anterior (with a narrow vestibule) and in the 
posteroventral region; eight long, straight and widely spaced anterior canals. A distinct 
flange extends around the entire free margin. Copulatory appendage with small, blunt- 
ended, lappet (Fig. 4). 

DIMENSIONS. Holotype: 1980.245, 9 carapace, length 0'50 mm; height 0*29 mm; width 
0-27 mm. Paratypes: 1980.246, rf LV., length 0'53 mm; height 0'28 mm. 1980.247, cf RV., 
length 0-52 mm; height 0'28 mm. 1980.248, rf RV., length 0'49 mm; height 0*27 mm. 
1980.249, 9 carapace, length 0'49 mm; height 0'31 mm; width 0'25 mm. 1980.250, <S LV., 
length 0-53 mm; height 0'29 mm. 

REMARKS. Loxoconcha (L.) gurneyi sp. nov. has previously been figured as Loxoconcha sp. 
A. by Bate from the Persian Gulf and as Loxoconcha sp. B. by Jain from Mandvi Beach, west 
coast of India. Loxoconcha gruendeli Jain 1978 from Mandvi beach is very close to this 
species but differs by having pits rather than reticulae in the dorsal region and by having a 
smooth posterior border; in L. (L.) gurneyi short ridges cross the posterior border. Like L. 
(L.) multiornata, a series of short ridges occur along the free outer margin of the ventral edge 
of the right valve (Fig. 3B, D). A number of individual valves completely lack a duplicature 




Fig. 3 Loxoconcha (Loxoconcha) gurneyi sp. nov.: A, E, LV. X 100 and sieve plate X 600, d 
holotype BM(NH) 1980.250; B, D, enlarged view posteroventral margin X 200 and ventral 
surface X 100 d carapace, specimen lost; C, X 100, paratype d dorsal carapace, specimen lost; F, 
X 100 paratype 9 left carapace, BM(NH) 1980.249; G, X 100, holotype 9 right carapace, 
BM(NH) 1980.245; H, X 100, paratype d RV., BM(NH) 1980.248; I, J, X 250, paratype -.d 
terminal hinge elements RV., BM(NH) 1980.246; K, X 125, paratype d internal RV., BM(NH) 

240 R. H. BATE& A. GURNEY 

Fig. 4 Copulatory appendage of Loxoconcha (Loxoconcha) gurneyi sp. nov. d paratype BM(NH) 

1 980.43 1 . Scale bar represents 0" 1 mm. 

(e.g. BM(NH) no. 1980.250) this is considered to be due to the animal dying immediately 
after moulting and before calcification of the duplicature part of the inner lamella. 

DISTRIBUTION. Loxoconcha (L.) gurneyi sp. nov. is a nearshore shelf species in the Persian 
Gulf its presence in Abu Dhabi lagoon (Fig. 1 1) was almost certainly the result of being 
taken into the lagoon by tidal currents. 

Pitted species 

Loxoconcha (Loxoconcha) indica Jain, 1978 

Loxoconcha sp. C., Bate 1971 : 246, 250, pi. 3, figs. 2n, 3n. 
Loxoconcha megapora indica n. subsp. Jain 1978 : 126, figs. 4L 1-2; 6P. 

TYPE LOCALITY. Mandvi beach, west coast of India. 

DIAGNOSIS (emended). Small species of Loxoconcha having densely pitted surface orna- 
mentation. Posteroventral margin of carapace broadly convex; posterior end broadly 
rounded with short, weakly concave, posterodorsal margin. Carapace convex in dorsal view; 
eye nodes distinct. 

DESCRIPTION. Carapace dimorphic, males rectangular, females quadrate; both sexes with 
broadly rounded anterior and posterior ends, posterodorsal margin being short and insignifi- 
cantly concave. Eye node distinct. Shell surface covered by a dense, finely pitted 
ornamentation, smooth patches medianly and dorsally marking the location of muscles on 
the inside of the shell. Periphery of carapace with very fine network of striae. Normal pores 
open to exterior by means of a sieve plate. Left valve larger than right. Hinge gongylodont, 
muscle scars a subvertical row of 4 oval scars with a crescent-shaped anterodorsal frontal scar 
and two, large, rounded, anteroventral mandibular scars. Duplicature of moderate width 
with a distinct vestibule extending around the anterior end; anterior marginal canals long, 
straight and widely spaced; approximately 9-10 anteriorly. 

DIMENSIONS. 1980.252, cf carapace, length 0*49 mm; height 0'26 mm; width 0*23 mm. 
1980.253, 9 LV., length 0'42 mm; height 0'26 mm. 1980.254, d carapace, length 0'48 mm; 
height 0-26 mm; width 0*23 mm. 1980.255, 9 RV., length 0'41 mm; height 0-26 mm. 
1980.256, 9 carapace, length 0*43 mm; height 0'27 mm; width 0'22 mm. 

REMARKS. Loxoconcha (L.) indica Jain, 1978 was first described as a subspecies of L. 
megapora Benson & Maddocks, 1964 by Jain from Mandvi beach, west coast of India. L. 
megapora, a smooth species found in the Knysna estuary, South Africa has a different dorsal 
outline (being parallel-sided to slightly concave) to Jain's material and the two are not 
considered to be conspecific. Jain's subspecies is, therefore, raised to specific status. 



Fig. 5 Loxoconcha (Loxoconcha) indica Jain. A, H, rf left carapace X 125 and sieve plate X 750, 
BM(NH) 1980.252; B, F, 9 LV. X 125 and internal view X 100, BM(NH) 1980.253; C, X 125, rf 
right carapace, BM(NH) 1980.254; D, G, external view X 125 and internal view, X 100, 9 RV. 
BM(NH) 1980.255; E, X 125, dorsal view, 9 carapace BM(NH) 1980.256. Loxoconcha 
(Loxoconcha) amvgdalanux sp. nov. I, X 125, paratype dorsal carapace -2 instar, BM(NH) 
1980.257; J, X 1 18, holotyperf RV. BM(NH) 1980.258. 

242 R. H. BATE & A. GURNEY 

Loxoconcha sp. C. recorded by Bate, 1971 from Abu Dhabi lagoon, Persian Gulf is 
considered to be conspecific with Jain's L. indica even though the postero- ventral margin of 
the Persian Gulf material is more broadly convex; the geographical range of the species is 
thus extended. Laterally L. indica has a superficial resemblance to L. matagordensis Swain, 
1955 from San Antonio Bay, Texas but lacks the posterodorsal Loxocorniculum projection, 
anterior marginal ridges and oblique anterodorsal slope of matagordensis. Ornamentally, 
there are many species, including L. rhomboidea the type species, that possess the type of 
ornamentation exhibited here; carapace outline differences as well as the male copulatory 
appendage being used to distinguish between them. Unfortunately although females with 
appendages occur for L. indica no male was available for dissection. Loxoconcha spec, 
illustrated by Hartmann (1980, pi. 10, figs. 14, 15) from Pt. Welshpool, Australia is compar- 
able in shape with L. indica although lacking the straight dorsal margin of the latter and 
having a much more finely pitted ornamentation. 

DISTRIBUTION. Mandvi Beach, west coast India and in the Persian Gulf where it occurs on 
the shallow terraces of Abu Dhabi lagoon having a salinity tolerance of 44-80-55'30%oand 
a surface water temperature of 20-36'2C; so far only found allochthonously on the near- 
shore shelf. 

Loxoconcha (Loxoconcha) amygdalanux sp. nov. 
(Figs 5, 1, J; 6, A-K; 7; 8, A-C & 1 1) 

Loxoconcha sp. B., Bate 1971 : 245, 246, 248, pi. 1, fig. 2 m; pi. 2, fig. 3 m. 
Loxoconcha sp. A., Paik 1977 : 56, 58, pi. 6, figs. 1 12-1 14. 

DERIVATION OF NAME. Latin amygdala, almond + nux, a nut. 

HOLOTYPE. BM(NH) no. 1980.258, rf RV., sample 6278, back reef, nearshore shelf. 

PARATYPES. BM(NH) nos 1980.257 & 259-263, 269 & 430. 

TYPE LOCALITY. Sample 6278 nearshore reef; salinity gradient 43'05-44-45%o; surface 
temperature 23'0-33'6C. 

DIAGNOSIS. Coarsely pitted species of Loxoconcha in which pits are larger towards centre of 
valve. Dimorphism distinct: females subquadrate, males rectangular. Posteroventral border 
broadly convex in female, less so in male. Male with distinct postero-ventrolateral carapace 
indentation. Copulatory appendage shovel-shaped with small, triangular, terminally pointed 

DESCRIPTION. Carapace quadrate with broadly rounded anterior margin and deep, convex 
posteroventral margin and straight to slightly convex dorsal margin in the female. Male very 
elongate with rounded anterior and deep depression in posterior ventrolateral surface. Eye 
nodes in both dimorphs distinct. Ornamentation of large pits more evenly sized in the 
female. In the male the pits are very much larger towards the centre of each valve. Normal 
pore sieve plates situated on broad murae between pits. Left valve only slightly larger than 
right, almost equi valve. In ventral view female carapace has appearance of an almond, from 
which the species name has been taken. Internally the hinge is well developed, gongylodont. 
Muscle scars a curved row of four adductors, an anterocentral crescent-shaped frontal scar 
and two oval anteroventral mandibular scars. Duplicature broad with narrow anterior and 
posteroventral vestibules. 9 anterior marginal canals, long, straight and widely spaced. 
Copulatory appendage shovel-shaped with small, triangular, terminally pointed lappets. 

DIMENSIONS. Holotype: 1980.258, 3 RV., length 0'57 mm; height 0'30 mm. Paratypes: 
1 980.257, -2 instar, carapace, length 0-38 mm; height 0*22 mm; width 0- 19 mm. 1 980.259, 3 
LV., length 0'58 mm; height 0'29 mm. 1980.260, -2 instar, carapace, length 0*40 mm; height 
0-26 mm; width 0'22 mm. 1980.261, -2 instar, LV., length 0'34 mm; height 0-19mm. 
1980.262, 9 RV., length 0'49 mm; height 0'30 mm. 1980.263, 9 RV., length 0'43 mm; height 




Fig. 6 Loxoconcha (Loxoconcha) amygdalanux sp. nov.: A, X 122, paratype rf LV., BM(NH) 
1980.259; B, D, sieve plate X 1-Q5K and external view X 125 -2 instar LV, paratype BM(NH) 
1980.261; C, X 127, paratype ventral carapace -2 instar, BM(NH) 1980.260; E, X 260 muscle 
scars paratype 9 RV, BM(NH) 1980.262; F, J, K, internal view X 123 and terminal hinge 
elements X 265, female RV., paratype BM(NH) 1980.263; G, H, I, internal view X 122 and 
terminal hinge elements X 350, -2 instar LV., paratype BM(NH) 1 980.264. 



Fig. 7 Copulatory appendage of Loxoconcha (Loxoconcha) amygdalanux sp. nov. cf paratype, 
BM(NH) 1 980.430. Scale bar represents 0- 1 mm. 

0-27 mm. 1980.264, -2 instar, LV., length 0*40 mm; height 0'22 mm. 1980.265, -2 instar, 
RV., length 0'38 mm; height 0-22 mm. 1980.266, -1 instar, RV., length 0'41 mm; height 
0-26 mm. 1 980.267, -2 instar, LV., length 0'37 mm; height 0'24 mm. 

REMARKS. Loxoconcha (L.) amygdalanux sp. nov. has been recorded from the Gulf of Oman 
and the Persian Gulf by Paik (1977) who records this species from his faunal zones A to C, 
open marine conditions ranging in depth from 7 to 208 metres. The coarse ornamentation, 
particularly towards the centre of the carapace, in association with carapace outline clearly 
sets this species apart from all others described from the Gulf/Indian Ocean region. Neither 
has any comparable species been described by Hartmann from the Red Sea nor Australia. 

DISTRIBUTION. Nearshore shelf and outer lagoon channels, Abu Dhabi lagoon; Central 
Basin Persian Gulf and Biaban Shelf, Gulf of Oman. 

Submenus EPAKROCONCHA nov. 

DERIVATION OF NAME. Greek epakros, pointed at the end + concha, Latin, shell. 
GENDER. Feminine. 
TYPE SPECIES. Epakroconcha batei sp. nov. 

DIAGNOSIS. Carapace subtrigonal in lateral outline of both dimorphs with left valve dorsal 
margin umbonate; right valve dorsal margin almost straight. Left valve larger than right. Eye 
node distinct. Normal pore canals with sieve plate. Shell surface ornamented. Hinge gongy- 
lodont. Muscle scars with 4 oval adductors, a crescent-shaped frontal and an oval antero- 
ventral mandibular. Anterior and posteroventral duplicature broad with vestibula and 
straight marginal canals, few in number. First antenna long, slender, 6 jointed with 4 
terminal bristles. 2nd antenna with two terminal claws and long spinneret bristle. Maxilla 
with small comb of spines on inner (1st) endite and having approx. 16 (one aberrant) bristles 
on respiratory plate. Walking legs slender. Copulatory appendage boot-shaped with small 
terminally rounded lappet. 

REMARKS. Epakroconcha subgen. nov. differs from other loxoconchid genera by having a 
subtrigonal shell outline in both dimorphs; the male being only slightly more elongate than 
the female. 

Loxoconcha (Epakroconcha) batei sp. nov. 
(Figs 8, D-H; 9, A-C; 10 & 1 1) 

DERIVATION OF NAME. Named in honour of Charles Spence Bate 1818-1889 for his work on 
Recent Crustacea. 

HOLOTYPE. BM(NH) no. 1 980.27 1 , d carapace, sample 1 094, central lagoon terrace. 




Fig. 8 Loxoconcha (Loxoconcha) amygdalanux sp. nov.: A, X 126, RV., -2 instar, paratype 
BM(NH) 1980.265; B, X 118, RV., -1 instar, paratype BM(NH) 1980.266; C, X 122, LV., -2 
instar, paratype BM(NH) 1980.267. Loxoconcha (Epakroconcha) batei subgen. et sp. nov.: D, X 
124, paratype rf right carapace, BM(NH) 1980.268; E, X 137, paratype 9 left carapace, BM(NH) 
1980.269; F, H, internal view X 1 32 and hinge X 146, 9 RV., paratype BM(NH) 1980.270; G, X 
2K, holotype rf, sieve plate, BM(NH) 1 980.27 1 . 




Fig. 9 Loxoconcha (Epakroconcha) batei subgen. et sp. nov.: A, X 127, holotype <f right 
carapace, BM(NH) 1980.271; B, X 121, paratype <t left carapace, BM(NH) 1980.272; C, X 122, 
paratype 9 dorsal carapace, BM(NH) 1980.273. D, X 125, paratype 9 right carapace, BM(NH) 
1980.428. E,X 1 -8K, sieve plate, paratype 9 right carapace, BM(NH) 1980.428. 

Fig. 10 Loxoconcha (Epakroconcha) batei subgen. et sp. nov. Appendages and copulatory organ. 
Scale A 0- 1 mm for appendages; scale B 0- 1 mm for copulatory organ. A, F, 1 st antenna and 
copulatory organ, d paratype, BM(NH) 1980.433; B, C, D, 2nd antenna, mandible and maxilla, 
9 paratype, BM(NH) 1980.432; E, 1st, 2nd & 3rd thoracic appendages, 9 paratype, BM(NH) 



Fig. 10 Loxoconcha (Epakroconcha) batei subgen. et sp. nov. Caption p. 246. 

248 R. H. BATE& A. GURNEY 

PARATYPES. BM(NH)nos 1980.268-270,272-273,428-9 & 432-3. 

TYPE LOCALITY. Sample 1094, weed sample preserved in alcohol, inner lagoon terrace. 
Shallow water marine, tidal; salinity gradient 49'70-55'3%o; surface temperature of 

DIAGNOSIS. Species of Epakroconcha having coarsely pitted shell ornamentation, the pits 
larger towards centre of each valve. Periphery of carapace with fine network of striae. 
Posteroventral border of right valve with five small marginal denticles; denticles smaller and 
sometimes fewer in left valve. Small tubercle present on ventrolateral part of shell just below 
mid point, more clearly seen in dorsal or ventral view. Approximately 8 straight anterior 
marginal canals; 3 posterior canals. Copulatory appendage boot-shaped with small rounded- 
triangular lappet. 

DESCRIPTION. Carapace subtrigonal in lateral outline; dorsal margin umbonate in the left 
valve. Anterior margin of both valves rounded; posterior with deep, convex posteroventral 
margin and short concave, posterodorsal margin. Left valve over-reaches right along dorsal 
margin. Males slightly more elongate than females. Eye node distinct; shell surface coarsely 
ornamented with large pits that decrease in size towards valve margins in which region a 
reticulation of low ridges (peripherally striae) are superimposed. A low tubercle is developed 
below mid-point on each valve, more clearly observed in dorsal or ventral view. Postero- 
ventral margin with five, small, marginal denticles. Normal pores with slightly recessed sieve 
plate each bearing a central setal pore: setae long and slender. Hinge gongylodont. Muscle 
scars with an oval to crescent-shaped frontal scar situated medially in front of 4 oval 
adductor scars, mandibular scar elongate-oval, situated well below adductors. Duplicature 
broad, with broad anterior and posterior vestibula; marginal canals short, straight, approxi- 
mately 8 anteriorly, 3 posteriorly and 6 posteroventrally. 1st antenna long, slender, 6 jointed 
with 4 slender terminal bristles and 4 bristles at distal end of penultimate podomere. 2nd 
antenna with two strong terminal claws and long spinneret bristle; terminal two podomeres 
fused. Mandible with strong masticatory process. Maxilla with blade-like respiratory process 
having 16 bristles (or less) of which one is aberrant. Thoracic limbs slender with strong 
terminal claws. Copulatory appendage boot-shaped with small, terminally rounded lappet. 

DIMENSIONS. Holotype: 1980.271 cf carapace, length 0'52 mm; height 0'32 mm; width 
0'23 mm. Paratypes: 1980.268 rf carapace, length 0'51 mm; height 0*31 mm; width 
0-24 mm. 1980.269, 9 carapace, length 0'49 mm; height 0'34 mm; width 0*25 mm. 
1980.270, 9 RV., length 0'50 mm; height 0'32 mm. 1980.272 d 1 carapace, length 0'52 mm; 
height 0-31 mm; width 0*26 mm. 1980.273 9 carapace, length 0'50 mm; height 0'33 mm; 
width 0'26 mm. 1980.428 9 carapace, length 0*52 mm; height - 35 mm; width 0'25 mm. 
1 980.429 9 RV., length 0'49 mm; height 0'29 mm. 

REMARKS. Loxoconcha (Epakroconcha) batei sp. nov. is readily separable from previously 
described species of the genus on its subtrigonal lateral outline and spinose posteroventral 

DISTRIBUTION. Inner lagoon and lagoon channels, Abu Dhabi lagoon. Transported as dead 
valves onto the nearshore shelf. 


It is popularly regarded that the strength of the carapace ornamentation of an ostracod is a 
direct reflection on the nature of the environment in which the species lives. A coarse 
ornament reflects an adaptation to survival within a high energy environment and a poorly 
ornamented shell is suitable for quieter low energy environments. This does not necessarily 
hold true for burrowing forms nor for those that are able to shelter within dense patches of 
weed or among growing coral communities. Also many genera are totally smooth for all their 




< o a 

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d - 3 




m ? S 




'3 -S 


S .2 1 



a i r 




250 R. H. BATE& A. GURNEY 

species; strength of their shell possibly being effected by their shape, for example, the rather 
rotund genus Xestoleberis. Here, all the species described live on weed or in algae on lagoon 
terraces, within lagoon channels or on the more open nearshore shelf environment, and as 
such are subjected to the considerable tidal influences of the region. For details of the 
oceanography of the region the reader is referred to Evans et al. 1973 from which the 
following information has been taken: 

Tides -maximum rise of 2- 5 m in front of the islands to 1 m at the back of the 

lagoon except during periods of prolonged onshore winds. 

Water depth -up to 7 m deep in the lagoon channels and rarely exceeding 2 m on the 
lagoon terraces; often becoming dry at low tide. 

Salinity -nearshore waters range from 42'7%o to 44'5%o: lagoon waters range from 

53'6%o to 66'9%o. Higher values have been recorded from tidal pools but no 
species of Loxoconcha or of Epakroconcha have been obtained from these 
high salinity (77'4%o) pools. 

Temperature -nearshore waters have a surface temperature of 23-24C while lagoon waters 
range from 22-36C. 

Loxoconcha (Loxoconcha) multiornata sp. nov. (Fig. 11). This species of Loxoconcha is 
the most coarsely ornate of those present in our material. It is both a common nearshore shelf 
species and at the same time a common inhabitant of the more quiet higher salinity waters of 
the lagoon where it occurs both on the terraces and within the lagoon channels. This species 
thus shows no special preference for a particular environment. We did not, however, find 
any live specimens of L. (L.) multiornata and because of this there is a possibility that at least 
part of its distribution is due to postmortem dispersal. We consider that this would only be 
partially responsible for the distribution of the species as such a wide dispersal has not been 
observed for the other species. 

Loxoconcha (Loxoconcha) gurneyi sp. nov. (Fig. 1 1). This species falls into the pattern of 
distribution expected of an ornate ostracod. The distribution of L. (L.) gurneyi, either as live 
specimens or dead valves, is restricted to the nearshore shelf environment. The presence of 
individuals just inside the mouth of the lagoon as well as on the tidal delta is to be expected 
through simple dispersal in a particularly high energy environment of both living and dead 
material. L. (Loxoconcha) gurneyi has also been recorded from beach sediments off the west 
coast of India by Jain (1978) see synonymy. 

Loxoconcha (Loxoconcha) indica Jain (Fig. 11). A finely pitted species, L. (L.) indica is 
considered to be restricted solely to a lagoon environment. Live specimens have only been 
obtained from within Abu Dhabi lagoon where it lives on the outer and central lagoon 
terraces. It is not a common ostracod and the presence of a small number of dead valves on 
the nearshore shelf significantly opposite the lagoon entrance is considered to be due to 
postmortem dispersal. The original description of this species from beach sand from Mandvi, 
west coast India, is regarded as not being the true habitat for live specimens. 

Loxoconcha (Loxoconcha) amygdalanux sp. nov. (Fig. 11). This species, like L. (L.) 
indica, is pitted but rather coarsely so. Indeed the intensity of the ornamentation is reflected 
by the dispersal of the species being equally dominant both on the nearshore shelf and within 
the channels and terraces of the outer and inner lagoon. The species has also been recorded 
by Paik (1977) from the Central Basin, Persian Gulf and from the Gulf of Oman. L. (L.) 
amygdalanux is thus considered to be tolerant of both high and low energy environments as 
well as tolerating a range of salinity and temperature gradients. 


Loxoconcha (Epakroconcha) batei sub. gen. et sp. nov. (Fig. 11). The recorded distribu- 
tion here represents our total knowledge of this species. Although live specimens were 
restricted to sample 1094, the type locality, the distribution of the species is not considered to 
be far removed from that shown here. The presence of dead valves outside the lagoon and 
possibly of those just inside the entrance is considered to be due to postmortem dispersal. 
The species is regarded as being restricted to the inner and central lagoon terraces, inhabiting 
a shallow water environment where a thin layer of filamentous algae often covers the 


Apart from Loxoconcha (L.) multiornata the species of Loxoconcha exhibit the pattern of 
distribution that relates the degree of ornamentation to the energy level of the environment: 
the finest ornamented form (L. (L.) indica) being restricted to a lagoon environment and the 
more coarsely ornamented species (L. (L.) gurneyi) being restricted to the nearshore shelf, 
more open sea environment. L. (L.) amygdalanux, intermediate in the strength of its 
ornamentation, has a distribution equally shared between the outer part of the lagoon and 
the shelf environment outside. L. (Epakroconcha) batei, although rather coarsely orna- 
mented, is the only species currently placed in this subgenus and in the absence of other 
species we cannot draw any conclusions from its restriction to a lagoon environment. 


Bate, R. H. 1971. The distribution of Recent Ostracoda in the Abu Dhabi Lagoon, Persian Gulf. Bull. 

Centre Rech. Pau-SNPA suppl. 5 : 239-256, 3 pis. 
Bate, R. H. 1973 in Evans, G. et al. The oceanography, ecology, sedimentology and geomorphology of 

parts of the Trucial Coast Barrier Island complex, Persian Gulf. In: Purser, B. H. (Ed.) The Persian 

Gulf, 411 pp., Springer- Verlag. 
Bate, R. H. & Sheppard, L. M. 1980. On Xestoleberis postangulata Bate & Sheppard sp. nov. 

Stereo- Atlas Ostracod Shells 7 : 1 1 - 1 06 . 
Bate, R. H., Whittaker, J. E. & Mayes, C. (in press). Marine Ostracoda of the Galapagos Islands and 

Ecuador. J. Linn. Soc. zool. 
Benson, R. H. & Coleman, G. L. 1963. Recent marine ostracodes from the eastern Gulf of Mexico. 

Univ. Kansas Paleont. Contr. No. 2 : 1-52, pis. 1-8. 
Benson, R. H. & Maddocks, R. 1964. Recent ostracodes of Knysna Estuary, Cape Province, Union of 

South Africa. Univ. Kansas Paleont. Contr. No. 5 : 1-39, pis. 1-6. 
Bhatia, S. B. & Kumar, S. 1979. Recent Ostracoda from off Karwar, west coast of India, 173-178, 3 

pis. In: VII International Symposium on Ostracodes, Beograd. 
Evans, G. et al. 1973. The oceanography, ecology, sedimentology and geomorphology of parts of the 

Trucial Coast Barrier Island Complex, Persian Gulf, pp. 233-269. In: Purser, B. H. (Ed.) The 

Persian Gulf, 41 1 pp., Springer- Verlag. 
Gurney, A. 1979a. On Paijenborchellina alata Gurney sp. nov. Stereo- Atlas of Ostracod Shells 

6 : 27-30. 

19796. On Paijenborchellina venosa Gurney sp. nov. Stereo-Atlas oj 'Ostracod Shells 6 : 107-1 12. 

Hartmann, G. 1 964. Zur Kenntnis der Ostracoden des Roten Meeres. Kieler Meeresforsch 20 : 35-1 27. 
1980. Die Ostracoden der Ordnung Podocopida G. W. Miiller 1894 der warmtem perierten und 

subtropisch-tropischen Kiistenabschnitte der Siid und Sudostkuste Australiens. Mitt. hamb. zool. 

Mus.Inst.ll: 1 1 1-204, 15 pis. 
Jain, S. P. 1978. Recent Ostracoda from Mandvi Beach, West Coast of India. Bull. Ind. Geol. Assoc. 

Paik, K. H. 1976. Rezente Ostracoden aus Oberflachen-sedimenten des Persischen Golfs und des Golfs 

von Oman. Diss. Univ. Kiel, 196 pp. . 

Paik, K. H. 1977. Regionale Untersuchungen zur Verteilung der Ostracoden im Persischen Golf und in 

Golf von Oman. 'Meteor' Forsch.-Ergebnisse, No. 28 : 37-76. 
Razzaq, S. 1979. A preliminary investigation of the marine benthic microfauna of Kuwait. Kuwait 

Institute for Scientific Research, 91 pp., 3 pis. 
Swain, F. M. 1955. Ostracoda of San Antonio Bay, Texas,/ Paleont. 29 : 561-646, pis. 59-64. 

Manuscript accepted for publication 27 February 198 1 

A revision of the spider genus Cocalus (Araneae: 

F. R. Wanless 

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


Cocalus Koch 1846 is a small genus of oriental and Australian Salticidae comprised of four 
known species, one of which is described here as new. Although the type species, Cocalus 
concolor Koch, is only known from a single adult male which has lost both palps, it is 
thought the low elevation on the carapace, shared by all species is sufficiently characteristic 
to unite the species here included in Cocalus. In the only known female (C. murinus Simon) 
the epigyne is dark with a median orifice and is not especially characteristic when compared 
with epigynes of related genera. However, the male palps (in the two species, C. gibbosus 
sp.n., and C. limbatus Thorell, for which these are known) are distinctive, the sinuous finger- 
like protuberance of the tibiae being, as far as I am aware, unique within the Salticidae. 

Opisthoncus Koch from Australia is the only other genus known to the author to have a 
similar elevation on the carapace. It does not, however, appear to be closely related to 
Cocalus as the eye patterns are different, the elevation is situated between the posterior 
lateral eyes and the palps are less complex. The biology of Cocalus is unknown. 

The standard abbreviations and measurements are those used by Wanless (1978a), but for 
the leg spination the system adopted is that used by Platnick and Shadab (1975). 

Genus COCALUS Koch 

Cocalus Koch, 1846: 180, 230. Type species Cocalus concolor Koch, by subsequent designation 
(Simon, 1901:408). Koch, 1850:48. Marschall, 1873:393. Scudder, 1882:72. Peckham & 
Peckham, 1885:266, 288. Simon, 1901:405, 407, 408. Sherborn, 1922:1375; 1932:345. 
Petrunkevitch, 1928 : 181. Neave, 1939, 1 : 778. Roewer, 1954 : 934. Bonnet, 1956 : 1 173. 

Cocala: Simon, 1 864 : 327, [lapsus calami]. Waterhouse, 1912:61. Neave, 19391: 778. 

DEFINITION. Medium to large spiders ranging from about 1'5 to 8'6 mm in length. Sexual 
dimorphism not marked. Carapace: characteristic of genus, profile generally as in Figs 2 A, 
B; 3A, B; moderately high, longer than broad, widest between coxae II-III, lateral margins 
usually with membraneous margin; fovea moderately long, situated just behind posterior 
lateral eyes; positioned centrally within posterior ocular quadrangle a small bump (arrowed 
in Fig. 2 A, B, E). Eyes: set on moderately well developed tubercles, with black surrounds that 
are normally covered in hairs; arranged in three transverse rows comprised of anterior 
medians and anterior laterals, posterior medians, and posterior laterals; anteriors 
subcontiguous with apices strongly procurved in frontal view and recurved in dorsal view; 
anterior medians largest, anterior laterals about half diameter of anterior medians; posterior 
medians relatively large, positioned more or less midway between anterior laterals and 
posterior laterals and outside optical axis of anterior laterals; posterior laterals about as large 
as anterior laterals, set close to lateral margins of carapace when viewed from above; 
quadrangle formed by posterior median and posterior lateral eyes broader than long and 
widest posteriorly; entire quadrangle (measured from between bases of anterior medians to 
posterior margins of posterior lateral eyes) occupying between 45 and 47 per cent of carapace 
length. Clypeus: between 32 and 64 per cent of diameter of anterior median eyes, usually 
clothed in hairs. Chelicerae: robust, more or less parallel, slightly inclined anteriorly; 

Bull. Br. Mus. nat. Hist. (Zool.) 41 (5): 253-26 1 Issued 1 7 December 198 1 




promargin with three teeth, retromargin with four or five. Maxillae and labium: more or less 
as in Fig. 3D; bases usually obscured by coxae I. Sternum: elongate scutiform, generally as in 
Fig. 3E. Coxae: robust; coxae IV largest, others subequal. Pedicel: short. Abdomen: long and 
tapered; spinnerets long, posteriors robust and almost twice as long as robust anteriors, 
medians slender and slightly shorter than anteriors; anal tubercle a well developed cone; 
tracheal system not examined; spiricle an indistinct transverse slit near base of anterior 
spinnerets; former position of colulus apparently indicated by patch of hairs. Legs: 
moderately long and robust; spines strong and numerous; legs I sometimes with scanty 
ventral fringes on metatarsi and tibiae, also minute ventral setae on tarsi and metatarsi (c.f. 
Portia Karsch); claws pectinate, tufts present, scopulae absent. Female palps: moderately 
long and robust with terminal claw. Male palps: large, moderately complex and morpho- 
logically similar; densely clothed in hairs (not always shown in figures). Tibiae with truncate 
ventral apophysis and large retrolateral apophysis containing a sinuous finger-like protuber- 
ance of the tibiae (Fig. 4C); cymbium with basal retrolateral lobe; embolus (e) robust, arising 
from distal end of tegulum and curving inwards towards the alveolar cavity; near embolic 
base an indistinct sclerite (a), also arising from embolic base a translucent lobe (1); tegulum (t) 
ovoid with tegular furrow (0, retrolateral striae and dark peripheral seminal ducts (Fig. 40). 
Epigyne: large and dark, median orifice plugged in specimen examined. Internal structure 
not examined. 

REMARKS. Platnick (pers. comm.) suggested that the tegular furrow of Phaeacius (see 
Wanless, 1981) and related genera may contain pores, as the furrow and striae resembled 
distribution channels which may spread a secretion over the surface of the tegulum (Fig. 1 A, 
B). A re-examination of the male palp of Phaeacius fimbriatus Simon, confirmed Platnick's 
suggestion by showing the presence of a single minute pore in the wall of the furrow (position 
arrowed in Fig. 1 A). The pore, which seems disproportionately small in relation to the size 
of the furrow, cannot be resolved with the stereoscopic microscope. Unfortunately there is 
insufficient material to confirm the presence of pores in Cocalus. 

AFFINITIES AND DIAGNOSIS. Cocalus belongs to a group of genera in which the male palps are 
characterized by the presence of a tegular furrow and truncate or obtuse apophysis of the 
venter of the tibiae. It is readily distinguished from other genera of this group (i.e. Boethus 
Thorell, Brettus Thorell, Codeta Simon, Cyrba Simon, Phaeacius Simon and Portia Karsch) 
by the presence of an elevation in the posterior ocular quadrangle (Fig. 2 A, B, E). 

Fig. 1 Phaeacius lancearius (Thorell), rf palp: A, apical region, x 60, showing tegulum (t), tegular 
furrow (0 and striae (s); location of pore indicated by arrow. B, pore in wall of tegular furrow, 
x 1200. 



Fig. 2 Cocalus concolor Koch, holotype cT; A, carapace, lateral; B, carapace, dorsal; C, abdomen, 
dorsal; D, chelicera, inner view; E, carapace, facies; F, abdomen, lateral. 

List of species in the genus Cocalus Koch, 1846 

Cocalus concolor Koch, 1 846 
C. gibbosus sp. n. 
C. limbatus Thorell, 1878 
C. murinus Simon, 1899 

Summary of taxonomic changes effecting species listed in the catalogues of Roewer (1954) 
and Bonnet (1956) as belonging in Cocalus. 

Cocalus africanus Thorell, junior synonym of Portia africana (Simon) (Wanless 1 9786) 

256 F. R. WANLESS 

Cocalus lancearius Thorell, transferred into Phaeacius Simon (Roewer 1954), redescribed 

Cocalus longipes Thorell, Cocalus macellus Thorell and Cocalus protervus Thorell all 

belong in Cocalodes Pocock, comb. n. 
Cocalus salax Thorell, belongs in Codeta Simon, comb. n. 

The synonyms listed by Bonnet (1956) are accepted, but for one species, Phaeacius 
ramipalpis (Thorell) which evidently belongs in Boethus (see Wanless 1981). Cocalodes, 
Codeta and Boethus are presently being revised by the author. 

Cocalus concolor Koch 
(Fig. 2A-F) 

Cocalus concolor Koch, 1846:180, rf. Holotype cf, Indonesia, Bintang Island (MNHU, Berlin) 
[examined]. Koch, 1850:49. Thorell, 1892:475. Simon, 1896:351; 1901 :407, 408. Rainbow, 
1898 : 352. Petrunkevitch, 1928 : 181. Roewer, 1954 : 935, Bonnet, 1956 : 1173. Proszyriski, 
1971 : 390. Wanless, 19786:85. 

Cocala concolor: Simon, 1864 : 327. [lapsus calami]. 

DIAGNOSIS. The holotype of this species, a male in poor condition, has no palps and a 
diagnosis cannot be given at the present time. 

FEMALE. Unknown, but C. murinus Simon from Sumatra may belong here. 

MALE HOLOTYPE. In poor condition with carapace and legs translucent. Carapace (Fig. 
2A, B): brown, irregularly clothed in recumbent white hairs; punctate with iridescent violet 
sheen in eye region. Eyes (Fig. 2A, B, E). Clypeus: with light brownish hairs below AM, 
densely white haired below AL. Chelicerae: dark brown with long light brown and black 
hairs; teeth as in Fig. 2D. Maxillae, labium and sternum: more or less as in C. murinus (Fig. 
3D, E); brownish. Abdomen (Fig. 2C, F): dark brown. Legs: detached, some missing; 
brownish. Palps: missing. 

Dimensions (mm): total length c. 7*5; carapace length 3'0, breadth 2*12, height 1*82; 
abdomen length 4'48; eyes, anterior row 1-9, middle row 1-90, posterior row 1*96; quad- 
rangle length 1-42. Ratios: AM : AL : PM : PL :: 14 : 7-5 : 5 : 7; AL-PM-PL :: 12-1 1-5; 
AM:CL:: 14:8. 

DISTRIBUTION. Indonesia, Bintang Island. 

MATERIAL EXAMINED. Holotype cf, data given in synonymy. 

REMARKS. Rainbow (1898) records this species from British New Guinea, but gives no details 
as to the precise locality, neither does he indicate the sex of the specimen concerned. For the 
present it seems best to regard this record with caution and not formally include it in the 
distribution list. 

Cocalus murinus Simon 
(Fig. 3A-E) 

Cocalus murinus Simon, 1899 : 102,9. LECTOTYPE 9 (here designated) Indonesia, Sumatra (MNHN, 
Paris) [examined]. Simon, 1901 : 406, 407. Roewer, 1954:934. Bonnet, 1956: 1173. Proszyriski, 
1971 :391. 

DIAGNOSIS. Other females of Cocalus are unknown and a diagnosis cannot be given at the 
present time. Previous studies on Portia and Phaeacius suggest that the epigynes of Cocalus 
may be difficult to distinguish from one another. 

MALE. Unknown, but possibly synonymous with the type species C. concolor from Bintang 



^ i 


Fig. 3 Cocalus murinus Simon, lectotype 9: A, dorsal view; B, carapace, lateral; C, epigyne; D, 

maxillae and labium; E, sternum. 

258 F. R. WANLESS 

FEMALE LECTOTYPE. In fair condition. Carapace (Fig. 3A, B): amber with vague markings 
radiating from foveal region; irregularly clothed in short recumbent white hairs (rubbed). 
Eyes: with black surrounds, fringed in white hairs. Clypeus: clothed in long white hairs. 
Chelicerae: amber, shiny with long white hairs basally; promargin with three teeth, retro- 
margin with five. Maxillae and labium (Fig. 3D): orange-brown with inner maxillae margins 
and labial tip yellowish. Sternum (Fig. 3E): yellow-brown tinged black, shiny; clothed in 
long coarse white hairs and fine pale brown ones. Abdomen: rubbe<^; dull whitish yellow with 
sooty lateral markings, pattern indistinct; spinnerets yellow-brown. Legs: pale amber to light 
yellowish orange; spination of legs I: metatarsi v 2-0-0, p 0-1-0, d 2-1-1, r 0-1-0; tibiae v 
2-2-2, p 1-0-1, d 1-1-1, r 1-0-1; patellae p 1-0-0, r 1-0-0; femora d 0-2^. Palp: light 
yellowish orange to pale amber. Epigyne (Fig. 3C): dark amber clothed in creamy hairs, 
orifice indistinct and evidently plugged. 

Dimensions (mm): total length 8'56; carapace, length 3'36, breadth 2'52, height l - 88; 
abdomen length 5*6; eyes, anterior row 1-96, middle row 2'02, posterior row 2'16; quad- 
rangle length 1-54. Ratios: AM : AL : PM : PL :: 15-5 : 8 : 5 : 8; AL-PM-PL :: 12-12-5; 
AM : CL :: 1 5*5 : c. 5 (too hairy for accurate measurement). 

DISTRIBUTION. Indonesia, Sumatra. 

MATERIAL EXAMINED. Lectotype 9, Sumatra (J.-L. Weyers) (MNHN, Paris, 16290). 

Cocalus gibbosus sp. n. 
(Fig. 4A-D) 

DIAGNOSIS. Similar to C. limbatus, but distinguished by the truncate dorsal prong of the 
retrolateral tibial apophysis (Fig. 4C). 

FEMALE. Unknown. 

MALE HOLOTYPE. In good condition. Carapace (Fig. 4A, B): dark orange-brown; clothed in 
short recumbent white hairs. Eyes: with black surrounds except AM; fringed in whitish hairs. 
Clypeus: covered in dark grey hairs below AM with whitish hairs below AL. Chelicerae: dark 
orange-brown, sparsely covered in fine black hairs; promargin with three teeth, retromargin 
with four. Maxillae: brownish with dull white inner margins. Labium: dark brown tipped 
dull white. Sternum: brownish with black hairs. Abdomen: dorsally orange-brown to greyish 
brown, clothed in short recumbent white hairs (rubbed); pattern indistinct; venter dark 
greyish with vague longitudinal rows of lighter spots, clothed in short black hairs. Legs: 
generally orange-brown with whitish, and black hairs; spination of legs I: metatarsi v 2-0-0, 
p 1-1-0, d 0-1-1, r 1-1-0; tibiae v 2-2-2, p 1-1-0, d 1-1-1, r 2-1-0; patellae p 1-0-0, r 
1-0-0; femora p 0-0-1 , d 0-2-3. Palp (Fig. 4C, D): similar to that of C. limbatus, clothed in 
white hairs. 

Dimensions (mm): total length 8*16; carapace, length 3*0, breadth 2*28, height 1*88; 
abdomen length 5*04; eyes, anterior row 1-78, middle row 1*88, posterior row 2*0; quad- 
rangle length 1-38. Ratios: AM : AL : PM : PL :: 14 : 7 : 4-5 : 7; AL-PM-PL :: 14 : 9. 


MATERIAL EXAMINED. Holotype <f, Australia, Lockerbie, N. Queensland (R. Raven) 
30.i-l.ii.1975 Queensland Museum, reg. no. QM S846. 

Cocalus limbatus Thorell 
(Fig. 5A-D) 

Cocalus limbatus Thorell, 1878 : 283, 311, rf. Holotype rf, Amboina (MCSN, Genova) [examined]. 
Thorell, 1881 : 492, 706. Simon, 1901 : 407. Roewer, 1954 : 934. Bonnet, 1956 : 1 173. Proszyhski, 
1971 :391. 



Fig. 4 Cocalus gibbosus sp. n., holotype rf: A, dorsal view; B, carapace, lateral; C, palp, retro- 
lateral; D, palp, ventral. Abbreviations: a, sclerite; e, embolus; 1, lobe; f tegular furrow; t 

DIAGNOSIS. Similar to C. gibbosus, but distinguished by the obtuse dorsal prong of the 
retrolateral tibial apophysis (Fig. 5C). 

FEMALE. Unknown. 

MALE HOLOTYPE. In fair condition. Carapace (Fig. 5A, B): amber to dark amber, irregularly 
clothed in whitish pubescent hairs (rubbed). Eyes: with black surrounds except AM; fringed 
in whitish hairs. Clypeus: fringed in light brownish hairs below AM with white hairs below 
AL. Chelicerae: dark amber; thinly covered in pale amber hairs; teeth not examined. 
Maxillae: brownish amber with inner distal margins yellowish. Labium: dark amber. 



Sternum: partly obscured by coxae, apparently similar in form to that of C. murinus; dark 
amber with pale amber hairs. Abdomen: dull brownish amber lightly tinged black (original 
markings bleached); a tuft of pale amber hairs anteriorly, otherwise irregularly covered in 
dull whitish hairs. Legs: generally amber; spination of legs I: metatarsi v 1-0-0, p 1-0-0, d 
2-1-1, r 0-1-0; tibiae v 2-2-2, d 2-2-1, r 1-0-0; patellae p 1-0-0, r 1-0-0; femora d 0-2-4. 
Palp (Fig. 5C, D): the protuberance resting within the 'bowl' of the retrolateral tibial 
apophysis is slightly shorter than that found in C. gibbosus. 

Dimensions (mm): total length 7'44; carapace, length 2*94, breadth 2-28, height 1*8; 
abdomen length 4*52; eyes, anterior row 1'92, middle row 1-96, posterior row 2*06; quad- 
rangle length 1-4. Ratios: AM : AL : PM : PL :: 16 : 8 : 5 : 7; AL-PM-PL :: 1 1-1 1; 
AM:CL:: 16:6. 

DISTRIBUTION. Indonesia, Amboina. 

MATERIAL EXAMINED. Holotype d 1 , data given in synonymy. 

Fig. 5 Cocalus limbatus Thorell, holotype cf; A, dorsal view; B, carapace, lateral; C, palp, retro- 
lateral; D, palp, ventral. 



I wish to thank the following colleagues for providing specimens for study. Dr G. Arbocco, 
Museo Civico di Storia Naturale, Geneva (MCSN, Geneva); M. M. Hubert, Museum 
national d'Histoire naturelle, Paris (MNHN, Paris); Dr M. Moritz, Zoologisches Museum, 
Berlin (ZM, Berlin); Dr Valerie Todd Davies, Queensland Museum, Brisbane (QM, 
Brisbane). Also Mr D. Macfarlane (CIE, London) for reading the manuscript. 


Bonnet, P. 1956. Bibliographia Araneorum. 2 (2) : 919-1925. Imprimerie Douladoure, Toulouse. 
Koch, C. L. 1846. Die Arachniden. 13 : 234 pp. Lotzbeck, Niirnberg. 

1850. Ubersicht des Arachnidensystems. Heft. 5, 104 pp. Lotzbeck, Niirnberg. 

Marschall, A. de 1873. Nomencl. Zool. 482 pp. C. R. Societatis Zoologico-Botanicae, Vindobonae. 

Neave, S. A. 1939. Nomencl. Zool. Vol I A-C: 957 pp. Zoological Society, London. 

Peckham, G. W. & Peckham, E. G. 1885. Genera of the family Attidae: with a partial synonymy. 

Trans Wis. Acad. Sci. Arts Lett. 6 : 255-342. 

Petrunkevitch, A. 1928. Systema Aranearum. Trans. Conn. Acad. Arts Sci. 29 : 270 pp. 
Platnick, N. I & Shadab, M. U. 1975. A revision of the spider genus Gnaphosa (Aranea: Gnaphosidae) 

in America. Bull. Am. Mus. nat. Hist. 155 : 3-66. 
Prdszyriski, J. 1971. Catalogue of Salticidae (Aranei) specimens kept in major collections of the world. 

Annlszool. Warsz. 28 : 367-J-519. 
Rainbow, W. J. 1898. Contribution to a knowledge of the arachnidan fauna of British New Guinea. 

Proc. Linn. Soc. N.S. W. 23 : 328-356. 
Roewer, C. R. 1954. Katalog der Araneae. 2 Abt. B: 924-1290. Institut Royal des Sciences Naturelle de 

Belgique, Bruxelles. 
Scudder, S. H. 1882-1884. Nomencl. Zool. I Supplemental List. 376 + 340 pp. II Universal Index. 

Bull. U.S. natn. Mus. No. 19. 
Sherborn, C. D. 1925. Index Animalium. Pt. VI: 1197-1452. British Museum (Natural History), 

Simon, E. 1 864. Histoire Naturelle des Araignees (Araneides), 540 pp. Roret: Libraire Encyclopedique, 

1896. Liste der Arachniden der Semon'schen Sammlung in Australien und dem Malayischen 

Archipel. In Semon, Zoologische Forschungsreisen in Australien und dem Malayischen Archipel. 

Denkschr. med.-naturw. GesJenaS : 341-352. 
1899. Contribution a la faune de Sumatra. Arachnides recuellis par M. J. L. Weyers, a Sumatra 

(Deuxieme memoire). Annls Soc. ent. Belg. 43 : 78-125. 

1 90 1 . Histoire Naturelle des Araignees. 2(3): 38 1-668. Roret: Libraire Encyclopedique, Paris. 

Thorell, T. 1878. Studi sui ragni Malesi e Papuani. Part II. Ragni di Amboina raccolti da Prof. O. 

Beccari. Mus. civ. Stor. nat. Giacomo Doria 13:317 pp. 
1881. Studi sui ragni Malesi e Papuani. Part III. Ragni dell Austro-Malesia e del Capo York, 

conservati nel Museo Civico di Storia Naturale di Geneva. Mus. civ. Stor. nat. Giacomo Doria 

17 : 720 pp. 

1892. Studi sui ragni Malesi e Papuani. Part IV, 2. Mus. civ. Stor. nat. Giacomo Doria. 

31 : 490 pp. 
Wanless, F. R. 1978a. A revision of the spider genera Belippo and Myrmarachne (Araneae: Salticidae) 

in the Ethiopian region. Bull. Br. Mus. nat. Hist. (Zool.) 33 (1) : 139 pp. 
19786. A revision of the spider genus Portia (Araneae: Salticidae). Bull. Br. Mus. nat. Hist. (Zool.) 

34 (3): 83-124. 

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(Zool.) 41(4): 199-212. 
Waterhouse, C. O. 1 9 1 2. Index zoologicus Noll: 324 pp. London. 

Manuscript accepted for publication 3 March 198 1 

The Phthiracarus species of C. L. Koch 

B. W. Kamill 

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


Koch (1841) described a number of species of Hoplophora from Regensburg, Germany, of 
which nine are currently classified in Phthiracams: crinita, ferruginea, globosa, laevigata, 
lentula, longula, lucida, straminea and testudinea (Parry, 1979). Apart from the last 
mentioned species, which appears from Koch's original figure to have a greater affinity with 
the Euphthiracaroidea (but see Jacot, 1936), all are undoubtedly members of the genus 
Phthiracams. These species are, however, extremely difficult to reidentify since Koch's very 
brief descriptions are based almost entirely on body shape and colour, two characters now 
known to be uniform in many species of this genus. Only globosa and laevigata can be 
characterized by body shape alone: in comparison with the other Phthiracams species 
collected at Regensburg, H. globosa was said to be very 'globular' while the illustration of//. 
laevigata shows the notogaster to be 'angled' anteriorly. 

Several redescriptions of Koch's species have been published, including those of Jacot 
(1936), Feider and Suciu (1957) and van der Hammen (1963). Jacot collected at more than 
20 localities in the Regensburg area, recognizing six of Koch's species (laevigata, testudinea, 
crinita, lentula, straminea and ferruginea) and one new Phthiracams species, P. boresetosus. 
Localities were chosen based not only on Koch's habitat descriptions but on Fiirnrohr's 
detailed lists of the Regensburg flora published in 1839. Jacot treated the redescriptions in 
considerable detail, identifying each species on the basis of body shape and size, and setal 
form and length. However, he neglected to include details of the leg chaetotaxy which are 
now regarded as being essential for the separation of Phthiracarus species. Although Jacot's 
interpretations of crinitus and lentulus correspond with those of the present study, his 
specimens have not been considered for neotype designation, since all are entire, uncleared 
and mounted together with one or more other species in Canada Balsam. In this condition, 
Jacot's material would not easily withstand dismounting, dissection and remounting. 
Further collecting at Regensburg by van der Hammen in 1959 and 1961 resulted in his 
description of P. laevigatus and the designation of a neotype. A number of other species were 
tentatively identified by van der Hammen (pers. comm.) as representing Koch's seven other 
species this material has been examined as part of the present study. Material collected by 
the late Dr Max Sellnick at Regensburg, and made available to the author through the 
courtesy of Dr Gisela Rack, Hamburg, has also been examined in an attempt to determine 
the identities of Koch's Phthiracarus species. Unfortunately, it has not been possible to 
borrow any of the specimens studied by Feider and Suciu. 

Family PHTHIRACARIDAE Perty, 1841 

Phthiracarus crinitus (C. L. Koch) 
(Figs 1-7) 

Hoplophora crinita Koch, 1841* : Heft. 32, t. 8. Regensburg [type series presumed lost]. NEOTYPE 
(here designated), Schweighauser Forest, Regensburg (ZM, Hamburg, A30/80). 

Phthiracarus crinitus: Jacot, 1936 : 172 [in part]. Topotypes, Dechbetten Forest, Regensburg (MHN, 
Geneva, 3 119h) [examined]; Willmann, 1931 : 130. 

The dating for the various Hefte follows Sherborn ( 1 923). 

Bull. Br. Mm. not. Hist. (Zool.) 41 (5): 263-274 Issued 1 7 December 1 98 1 




Figs 1-6 Phthiracarus crinitus: (1 ) aspis, lateral; (2) notogaster, lateral; (3) notogaster, dorsal; (4) 
sensillus; (5) aspis, dorsal; (6) ano-genital region. 



ADULT: Large and strongly sclerotized. The aspis (Figs 1, 4-5) ranges in length from 
255-330 jum with a maximum width of 220-270 /zm. All the dorsal setae are long, simple 
and procumbent. The interlamellar setae (//) which are inserted just posterior to the 
bothridia, are about 1*5 times the length of the lamellars (la) and extend to the rostral bases. 
Setae (ro) almost reach the anterior aspal margin. The sensilli are 60-100 /um in length and 
expanded basally the distal portion is slender, serrated and blunt terminally. The exoboth- 
ridial setae (ex) are short. The notogaster (Figs 2-3), 480-550 /zm long (measured along a line 
through <:,-/,) and with a greatest depth of 330-390 /um, is elongate in lateral aspect. All the 
notogastral setae are long (greater than the distance c r d { ) and simple, the c and d series being 
erect while those in the e, h and ps series are recurved. Seta c, is situated on the posterior 
margin of the collar and setae c 2 _ 3 submarginally. Vestigial /| is located just dorsal to the seta 
/i, while the fissures ip and ips are situated between setae h 2 and /z 3 and between setae ps 3 and 
ps 4 respectively. On each anal plate (Fig. 6) there are three setae of more or less equal length 
(an } _ 2 and ad 3 ). The nine genital setae are arranged in two rows, an anterior marginal row of 
five setae (g,_ 5 ) and a posterior submarginal row of four (g 6 - 9 ). A single aggenital seta ag is 
located antiaxially in the genital furrow. The genital papillae are typically phthiracaroid in 
form, the anterior pair being considerably smaller than the two posterior pairs. The 
chelicerae are 1 80-2 10 //m long with about 20 sharply pointed spines on the paraxial surface 

Figs 7-8 Leg I, trochanter to tibia, dorsal aspect: (7) Phthiracarus crinitus; (8) Phthiracarus 


266 B. W. KAMILL 

of the principal segment and about 25 conical spines antiaxially. The leg chaetotaxy is of the 
'complete type' (see Parry, 1979) with four setae on femur I and a single seta on genu IV. All 
the solenidia are long and more or less straight. On tarsus I the seta coupled with solenidion 
a> 2 is short and barely discernible. Setae (tc) and (w) on this segment, and (?c), (w), (p) and s on 
tarsi II-IV are ribbon-like, hooked distally and covered with whorls of spicules in the middle 
third. The four setae on femur I (Fig. 7), d, (v) and /', are all located at about the same level on 
the segment. Seta d which is the shortest of the four (about two-thirds as long as /'), is 
thickened, bluntly serrated and somewhat curved. 

MATERIAL: Three specimens from a sample (no. 59015) of moss and bilberries (Vaccinium 
myrtillus), Schweighauser Forest, Unterkaulhausen, Regensburg, 19.vii.1959 (M. Sellnick), 
deposited in the collections of the Zoological Museum of the University of Hamburg. One of 
these specimens (A30/80) is hereby designated as the neotype. Other material was examined 
from rotten wood and litter beside a brooklet (sample no. 61 R45), Donaustaufer 
Forest, Regensburg, (L. van der Hammen) (RNH, Leiden). P. crinitus was 
not widespread in the Regensburg material, being present only in small numbers in the 
above two samples. 

REMARKS: Koch recorded crinita predominantly in mosses, Regensburg. Although the 
original description of the species was rather incomplete, crinita appeared to be character- 
ized by unusually long notogastral setae a feature which was noted by Jacot (1936) in his 
description of specimens which he considered to be conspecific with H. crinita. Jacot's 
specimens are for the most part conspecific and in good condition. However, as mentioned 
above, they have not been considered for neotype designation. A neotype was selected from 
amongst Sellnick's spirit specimens which were found to be conspecific with the mite 
described by Jacot. 

Phthiracarus ferrugineus (C. L. Koch) 
(Figs 9-1 3) 

Hoplophoraferruginea Koch, 1841 : Heft. 32, t. 10. Regensburg [type series presumed lost]. NEOTYPE 
(here designated), Taimering, Regensburg (RNH, Leiden, P2005-7). 
[Phthiracarus ferrugineus: Jacot, 1936 : 179. Misidentification, see under P. longulus.} 

ADULT: Rather small and weakly sclerotized. The aspis (Figs 10-12) is about 225 /zm in 
length with a greatest width of about 160 /zm. The rostrals (ro) are short and do not reach the 
anterior aspal margin. Setae (it) are twice the length of setae (Id) and extend to the rostral 
bases. The sensilli are long (70-80 //m), slender and distally serrated. The exobothridial setae 
(ex) are short. The notogaster (Fig. 9), about 350 /zm in length, has a. maximum depth of 
about 270 jum and bears moderately long (more or less equal to the distance c, -/,), simple 
setae which are erect to recurved. Setae c, and c 3 are inserted close to the posterior collar 
margin and seta c 2 submarginally. Vestigial f { is located one-third of the distance between 
setae h\ and ps } . Only the fissures ia and im appear to be present. On each anal plate there 
are three setae (setae ad t _ 2 vestigial) of which an,_ 2 are the longest. The chelicerae are 
135-140 //m long with 9 to 10 sharply pointed spines on the paraxial surface of the principal 
segment and 6 to 10 conical spines antiaxially. The leg chaetotaxy is of the 'complete type' 
with rather short and straight solenidia. On femur I (Fig. 1 3) seta d is short (about half as long 
as /'), thickened and, as in crinitus, this seta is curved and bluntly serrated. In the three 
available specimens, the setal arrangement on this segment shows considerable variation. In 
the neotype (Fig. 13) seta d is located on a level with seta /' and anterior to the ventral setae, 
while in a second specimen d is located anterior to the lateral seta, and in a third specimen, 
posterior to the lateral seta. 

MATERIAL: Three specimens from rotten material in a moist hayfield, Taimering, Regens- 
burg, (L. van der Hammen) (sample no. 61 R34), deposited in the collections of 
the Rijksmuseum van Natuurlijke Historic, Leiden. One of these specimens (P2005-7) is 
hereby designated as the neotype. 






Figs 9-13 Phthiracarus ferrugineus: (9) notogaster, lateral; (10) aspis, lateral; (1 1) aspis, dorsal; 
(12) sensillus; (13) leg I, trochanter to tibia, dorsal aspect. 

268 B. W. KAMILL 

REMARKS: Koch recorded ferruginea in mosses on trees. He described the species as being 
very small with long, fine notogastral setae and from his figure it is evident that the species 
possessed rather long and prominent sensilli. 

The smallest of the Regensburg species examined in the present study, P. ferrugineus 
(notogastral length 350 um) is somewhat unusual in its possession of four setae on femur I 
and a single seta on genu IV, two features which are associated with larger species 
(notogastral length 500-1000 um) of the genus. Smaller species with a notogastral length of 
less than 500 um are generally characterized by a 'reduced' form of leg chaetotaxy. 

Although rather smaller, P. ferrugineus bears considerable resemblance to P. membranifer 
Parry (notogastral length 31 0-500 /zm) recorded from the fermentation layer under Sitka 
spruce, Tintern Forest, Monmouthshire. In comparison with the latter, P. ferrugineus differs 
only in having a 'complete' form of leg chaetotaxy. 

Phthiracarus globosus (C. L. Koch) 
(Figs 8, 14-18) 

Hoplophora globosa Koch, 1841 : Heft. 32, t. 12. Regensburg [type series presumed lost]. NEOTYPE 

(here designated), Burgweinting, Regensburg (ZM, Hamburg, A3 1/80). 
Phthiracarus globosus: Willmann, 1931 : 193; Feider &Suciu, 1957 : 5; Sellnick, 1960: 131. 
Phthiracarus globus Parry, 1979:341. Holotype, Rydal Water, Westmorland, England (BMNH, 

London, 1976.2.18.3.) [examined]. Syn. nov. 

ADULT: Large and strongly sclerotized. The aspis (Figs 15-17) ranges in length from 
240-3 10 //m with a maximum width of 1 80-250 um. All the dorsal setae are very long, 
simple and procumbent. Setae (if) and (la) are more or less equal in length and both pairs of 
setae reach the rostral bases. The sensilli are 40-55 um long, lanceolate and serrated 
marginally. The exobothridial setae (ex) are moderately long. The notogaster (Figs 14, 18), 
500-640 um long and ^ith a maximum depth of 300-460 um, is rather globular in 
comparison with other species of the genus. All the setae are long (greater than the distance 
c,-J,), simple and almost erect. Seta c, is inserted on the posterior margin of the collar and 
setae c 2 _ 3 submarginally. Vestigial /J is located adjacent and just dorsal to the seta /z,, towards 
the mid-dorsal line. The fissures ip and ips are absent. On each anal plate there are five long 
setae; an t _ 2 and ad 3 being more or less equal in length and somewhat shorter than setae ad t _ 2 . 
The chelicerae are 170-1 90 um long. The principal segment carries 12-26 sharply pointed 
spines on the paraxial surface and 22-25 conical spines antiaxially. The leg chaetotaxy is of 
the 'complete type' and on femur I (Fig. 8) the dorsal seta is long (equal in length to /'), 
slightly curved and covered in whorls of blunt serrations. 

MATERIAL: Two specimens from litter under 'broom (Genista? [= Sarothamnus] growing 
under willows, Burgweinting, Regensburg, 16.viii.1959 (M. Sellnick), deposited in the 
collections of the Zoological Museum of the University of Hamburg. One of these specimens 
(A3 1/80) from sample no. 59059 is hereby designated as the neotype. A further ten 
specimens were examined from rotten wood and litter beside a brooklet (sample no. 6 1 R45), 
Donaustaufer Forest, Regensburg, (L. van der Hammen) (RNH, Leiden). P. 
globosus was always recorded in small numbers in the Regensburg samples examined. 

REMARKS: P. globosus is a very distinctive species being characterized by the 'globular' form 
of the notogaster. In comparison with German specimens, it was found that the British 
material examined was considerably larger (notogastral length 570-850 um) and more 
heavily sclerotized. It is interesting that Jacot did not regard globosus as a Phthiracarus 
species, but rather as the male of Hoplophora decumana ( = Oribotritia decumana). 

Phthiracarus laevigatus (C. L. Koch) 
Hoplophora laevigata Koch, 1844 : Heft. 38, t. 16. Regensburg [type series presumed lost]. 






Figs 14-18 Phthiracarus globosus: (14) notogaster, lateral; (15) aspis, dorsal; (16) sensillus; (17) 

aspis, lateral; ( 1 8) notogaster, dorsal. 

270 B. W. KAMILL 

Phthiracarus laevigatus: Jacot, 1936: 167; van der Hammen, 1963:704. NEOTYPE (designated), 
Donaustaufer Forest, Regensburg (RNH, Leiden) [examined]. 

This species has been adequately redescribed by van der Hammen (1963). 

Phthiracarus lentulus (C. L. Koch) 
(Figs 19-24) 

Hoplophora lentula Koch, 1841 : Heft. 32, t. 16. Regensburg [type series presumed lost]. NEOTYPE 

(here designated), Taimering, Regensburg (RNH, Leiden, P2001^4). 
Phthiracarus lentulus: Jacot, 1936 : 175 [in part]. Topotypes, Unter-Isling/Burgweinting, Regensburg 

(MHN, Geneva, 3135h) [examined]. [Phthiracarus lentulus: Feider & Suciu, 1957 : 5. Misidentifi- 


ADULT: Medium-sized. The aspis (Figs 21-23) ranges in length from 270-3 10 //m with a 
maximum width of 220-240 /urn. Setae (it) and (la) are about equal in length and both pairs of 
setae extend half the distance il-ro. The rostral setae (ro) reach the anterior aspal margin. The 
sensilli are 70-90 //m in length and slender; proximally the sensillar margin is smooth while 
the distal part is serrated. The exobothridial setae (ex) are moderately long. The notogaster 
(Figs 19-20) ranges in length from 500-580 urn with a greatest depth of 400-440 urn. All the 
setae are short (less than c,-J,), stout and more or less erect. Seta c 3 is located on the posterior 
collar margin and setae c,_ 2 submarginally. Vestigial f } is located anterior to the seta h { while 
the fissures ip and ips are absent. On each anal plate there are three setae (ad t _ 2 vestigial); 
setae tf,_ 2 being slightly longer than ad v The chelicerae are 1 90-200 /zm long with 18-26 
sharply pointed spines on the paraxial surface of the principal segment and 17-35 conical 
spines antiaxially. The leg chaetotaxy is of the 'complete type', the solenidia being long and 
almost straight. On tarsus I the distal seta coupled with solenidion a) 2 is long and divided into 
two parts by a longitudinal constriction. On femur I (Fig. 24) setae d, (v) and /' are all located 
at about the same level on the segment. Seta d, which is almost as long as /', is thickened, 
curved and bluntly serrated. 

MATERIAL: Four specimens from moist wood, moss and litter, Taimering, Regensburg, (L. van der Hammen), deposited in the collections of the Rijksmuseum van 
Natuurlijke Historic, Leiden. One of these specimens (P2001-4) is hereby designated as the 

REMARKS: Koch recorded lentula predominantly in moss in woods and described the species 
as being of medium size with short notogastral setae. It is evident that the sensilli were 
probably either short or very fine, since these were not included by Koch in his original 
figure. The sensilli are indeed very fine in lentulus (Fig. 23), a feature which was also noted 
by Jacot in his redescription of this species. 

Phthiracarus longulus (C. L. Koch) 
(Figs 25-31) 

Hoplophora longula Koch, 1841 : Heft. 32, t. 17. Schweighauser Forest, Regensburg [type series 

presumed lost]. NEOTYPE (here designated), Donaustaufer Forest, Regensburg (RNH, Leiden, 

Phthiracarus ferrugineus (Koch) sensu Jacot, 1936: 179 [in part]. Topotypes, Dechbetten Forest, 

Regensburg (MHN, Geneva, 3 1 1 9h) [examined]. 
Phthiracarus tardus Forsslund, 1956:216. Holotype, Vasterbotten, Sweden (paratype, BMNH, 

London, 1 964.7. 1 3.78) [examined]. Syn. nov. 

ADULT: Small and weakly sclerotized. The aspis (Figs 27, 29-30) ranges in length from 
220-255 //m with a maximum width of 1 70-190 /^m. The interlamellar setae (it) which are 
inserted on a level with the bothridia, are twice as long as the lamellars (la) and extend almost 
to the rostral bases (ro). The sensilli are lanceolate, serrated marginally, and range in length 
from 30-40 um. The exobothridial setae (ex) are short. The notogaster (Figs 26, 28) is some- 
what elongate and ranges in length from 430-490 //m with a maximum depth of 







Figs 19-23 Phthiracarus lentulus: (19) notogaster, lateral; (20) notogaster, dorsal; (21) aspis, 

lateral; (22) aspis, dorsal; (23) sensillus. 




Figs 24-25 Leg I, trochanter to tibia, posterolateral aspect: (24) Phthiracarus lentulus; (25) 

Phthiracarus longulus. 

300-320 //m. All the setae are erect and more or less equal to the distance c,-d,. Setae c,_ 3 
form a row immediately behind the posterior collar margin while vestigial f } is located just 
dorsal to the seta /?, and towards the mid-dorsal line. The fissures ip and ips are absent. On 
each anal plate (Fig. 31) there are three rather short setae, ad^ being the shortest. The 
chelicerae are 1 50-200 /zm long with 4-14 sharply pointed spines on the paraxial surface of 
the principal segment and 6-10 conical spines antiaxially. The leg chaetotaxy is of the 
'reduced type' (see Parry, 1979) with three setae on femur I (V absent) and no setae on genu 
IV (/' absent). Tarsus II bears 12 setae (subunguinial seta present), a feature which is 
generally associated with 'complete chaetotaxy' species. All the solenidia are moderately 
long and straight. Solenidion w 2 on tarsus I is coupled with a minute distal seta. On femur I 
(Fig. 25) the dorsal seta is short, serrated, curved distally and located posterior to the setae /' 
and v'. 

MATERIAL: Eight specimens from rotten wood and litter beside a brooklet, Donaustaufer 
Forest, Regensburg, (L. van der Hammen), deposited in the collections of the 
Rijksmuseum van Natuurlijke Historic, Leiden. One of these specimens (P2012-15) from 
sample no. 6 1 R45 is hereby designated as the neotype. 

REMARKS: Koch recorded longula in moss in woods, a species he described as being small 
with long, fine notogastral setae and rather short, round-ended sensilli. 

Of the species here described, P. longulus appears to be the most abundant in all the 
Regensburg samples. The species has been recorded only rarely in the British Isles but in 
Sweden the author has found it to be extremely widespread. 






Figs 26-31 Phthiracarus longulus: (26) notogaster, lateral; (27) aspis, lateral; (28) notogaster, 
dorsal; (29) sensillus; (30) aspis, dorsal; (3 1 ) ano-genital region. 

274 B. W. KAMILL 


Of the eight Phthiracarus species described by C. L. Koch, six are capable of being 
reidentified with reasonable certainty. The two remaining species (lucida and straminea) 
cannot in the author's opinion be separated by any morphological characters mentioned in 
the original descriptions, although lucida was collected from a 'swampy' meadow, while 
straminea was recorded from moss in woods. However, Phthiracarus species are not 
generally considered to be habitat specific and it seems likely therefore, that Koch was in fact 
concerned with only one species, different specimens of which exhibited minor differences in 


The present study was based largely on specimens kindly sent by Dr L. van der Hammen, 
Rijksmuseum van Natuurlijke Historic (RNH), Leiden. Specimens from the Jacot and 
Sellnick Collections were examined through the courtesy of Dr B. Hauser, Museum 
d'Histoire Naturelle (MHN), Geneva, and Dr G. Rack, Zoologisches Institut und 
Zoologisches Museum (ZM), Hamburg. 


Feider, Z. & Suciu, I. 1957. Contributii la cunoaterea oribatidelor (Acari) din R.P.R. familia 

Phthiracaridae Perty 1 84 1 . Studii Cere, stiint. lasi (Biol.) 8 : 23^6. 
Forsslund, K-H. 1956. Schwedische Oribatei (Acari). III. Ent. Tidskr. 11 : 210-218. 
Furnrohr, A. E. 1839. Naturhistorische Topographic von Regensburg, Bd II. Regensburg: Manx. 
Hammen, L. van der, 1963. The oribatid family Phthiracaridae. II. Redescription of Phthiracarus 

laevigatus (C. L. Koch). Acarologia 5 : 704-7 1 5. 

Jacot, A. P. 1936. Les Phthiracaridae de Karl Ludwig Koch. Revue suisse Zool. 42 : 161-187. 
Koch, C. L. 1835-1844. Deutschlands Crustaceen, Myriapoden und Arachniden. Regensburg. 
Parry, B. W. 1979. A revision of the British species of the genus Phthiracarus Perty, 1841 

(Cryptostigmata: Euptyctima). Bull. Br. Mus. nat. Hist. (Zool.) 35 (5) : 323-363. 
Sellnick, M. 1960. Formenkreis: Hornmilben, Oribatei. Tierwelt Mitteleur. 3 : 45-132. 
Sherborn, C. D. 1923. On the dates of C. L. Koch, 'Deutschlands Crustaceen, Myriapoden und 

Arachniden,' 1835^4. Ann. Mag. nat. Hist. 9(11): 568. 
Willmann, C. 193 1 . Moosmilben oder Oribatiden (Cryptostigmata). Tierwelt Dtl. 22 : 79-200. 

Manuscript accepted for publication 28 April 1 98 1 

The status of Lamingtona lophorhina McKean & 
Calaby, 1968 (Chiroptera: Vespertilionidae) 

J. E. Hill 

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

K. F. Koopman 

Department of Mammalogy, The American Museum of Natural History, Central Park West 
at 79th Street, New York, New York 10024, U.S.A. 


The genus Lamingtona and its sole species L. lophorhina were described by McKean & 
Calaby (1968) on the basis of six specimens from Mount Lamington in Northern Province, 
eastern Papua New Guinea, at 8 55' S, 148 10' E. Although showing Lamingtona to have a 
clear relationship to Nyctophilus, the original account is insufficiently detailed to permit a 
proper assessment of its status or of possible relationships of L. lophorhina to any of the 
several currently recognized species of this predominantly Austro-Papuan genus. The 
original material of L. lophorhina is housed in the South Australian Museum, Adelaide and 
in the collections of the Commonwealth Scientific and Industrial Research Organisation 
(Division of Wildlife Research), Canberra. Through the courtesy of Dr J. H. Calaby we have 
been able to examine the two paratypes from Canberra and to compare them directly with 
most of the described forms of Nyctophilus, with especial attention to those already known 
from New Guinea. 

Systematic Section 

McKean & Calaby (1968) compared Lamingtona primarily with Nyctophilus, although only 
two species (timoriensis and geojfroyi) of the latter are actually mentioned in their account. 
The principal diagnostic character that they use in separating Lamingtona from Nyctophilus 
and from its close relative Pharotis is the absence in Lamingtona of any band of integument 
connecting the ears across the forehead and they also remark that the ears of Lamingtona are 
smaller and the tragus not broadened basally as it is in these genera. In Nyctophilus it has 
been possible to examine specimens in alcohol of geojfroyi, timoriensis, gouldi, bifax, 
walkeri, microdon, arnhemensis and microtis (the last including bicolor), the species that 
tentatively we recognize in this genus, and in Pharotis similar exampks of its sole represent- 
ative, P. imogene. The ears are connected by a moderate or high band (which incidentally is 
not an extension of their inner or medial margins, but a separate structure arising from the 
outer or medial surface of each conch) in Pharotis imogene and in all of the species of 
Nyctophilus except arnhemensis and microtis. In arnhemensis the band is low and in 
microtis only slightly developed and barely discernible or absent in the midline, a point 
made clear in the original description of this species by Thomas (1 888) who says 'band across 
the forehead nearly or quite obsolete in the centre'. On occasion, therefore, the ears are not 
joined at all. In Pharotis imogene and in several of the species of Nyctophilus the ears are 
large or sometimes very large, but those of N. microtis are relatively small, as its name 
implies, although Thomas (1915) found them to be rather larger than he thought originally 
(1888). The ears of walkeri and arnhemensis are of similar size or are only a little larger than 

Bull. Br. Mus. nat. Hist. (Zool.) 41(5): 275-278 Issued 1 7 December 1 98 1 



those of microtis, which in turn resemble the ears of Lamingtona in size. The tragus is 
broadened basally in all except Nyctophilus microtis, although the extent of such widening 
may be slight. In microtis, however, this broadening of the tragus is often very poorly marked 
and may be virtually absent. 

According to McKean & Calaby (1968) Lamingtona lophorhina lacks an ear band. 
Examination of the two paratypes (99 CM 2090, 2091) shows this to be correct in the sense 
that the ears are not joined near the bases of their inner or medial margins by an obvious 
band of integument. However, although in both paratypes the area between and in front of 
the ears is densely pilose, careful investigation reveals the presence of a low integumentary 
ridge in the dry skins, extending from a point just behind the inner margin of each ear 
towards the midline. These ridges do not quite reach the median line but diminish in height 
towards the centre of the head. The condition exactly resembles that found in Nyctophilus 
microtis in which the ears are supported medially by a thick, tapered integumentary ridge 
whose outer and highest parts are integral with the medial faces of the ears. As in the 
paratypes of Lamingtona lophorhina the integumentary ridges taper to the midline, which 
on occasion they do not quite reach, and at best the ears are joined medianly only by a low 
elevation. So far as can be determined from the dry skins the tragus in one paratype (CM 
209 1 ) is more or less spatulate and little widened basally but in the other example (CM 2090) 
the tragus is slightly widened in its lower part: in both the tragus has a bluntly rounded tip, 
inclined anteriorly and is closely similar to the tragus of Nyctophilus microtis. We can see no 
reason that supports the recognition of Lamingtona as a genus distinct from Nyctophilus on 
account of the characters originally proposed. Miller (1907) in defining Nyctophilus noted 
'Ears . . . usually joined across forehead by a conspicuous band of membrane, but this 
obsolete at middle in one species . . .'. [Since Miller stated that he had examined timoriensis, 
microtis and walkeri, the 'one species' was presumably microtis.} Moreover, the extent of 
joining of the ears (and the shape of the tragus) is known to vary within other bat genera and 
scarcely justifies even subgeneric recognition, especially since in Nyctophilus the relatively 
rudimentary integumentary ridges of microtis and lophorhina are linked by the low band of 
arnhemensis to the higher, more obvious connecting bands of species of similar size such as 
bifax, walkeri and microdon. There appear to be no significant cranial features to support the 
subgeneric separation of microtis and lophorhina (and possibly arnhemensis) from the 
remaining species currently recognized in Nyctophilus. 

In discussing the affinities of Lamingtona McKean & Calaby (1968) remarked that it 
'probably belongs in the subfamily Nyctophilinae but because of the ear characteristics it 
falls outside that subfamily as currently defined'. This claim deserves examination, 
especially in view of the close correspondence between the ear of L. lophorhina and that of 
Nyctophilus microtis, even to the shape and size of the small internal lobe near the base of its 
posterior or outer margin. Although Tate (1941) did indeed mention 'large united ears' as 
one of several features of the subfamily, the original definition by Miller (1907) characterized 
it externally by the form and structure of the muzzle and noseleaf, without reference to the 
ears. Miller remarked that the subfamily 'Differs from the Vespertilioninae in the abruptly 
truncate muzzle, on the anterior face of which the nostrils open forward beneath a distinct 
horseshoe-shaped ridge or small noseleaf. Incidentally, though the New World genus 
Antrozous (usually, though perhaps erroneously placed in the Nyctophilinae) has an ear 
band basically similar to that of Nyctophilus, it is interrupted in the middle, as on occasion is 
the rudimentary band of N. microtis, and therefore does not (contra Tate) join the two ears 
together. There are evidently no grounds for excluding lophorhina and microtis from the 
Nyctophilinae unless it is proposed radically to redefine this alleged subfamily. As pointed 
out by McKean & Calaby, lophorhina has the small noseleaf and grooved muzzle character- 
istic of Nyctophilus: in fact, so far as can be determined from the dry skins of the paratypes, it 
is very similar in rhinarial structure to microtis, with an apparently truncate muzzle. 

The two paratypes (CM 2090, 2091) are dorsally a rather uniform reddish brown, 
ventrally a little paler and more ochreous: the wing membranes are blackish brown, the tail 
membrane brownish. Both have every indication of prolonged immersion in alcohol (from 


1929 to 1967 according to McKean & Calaby, 1968): the dorsal pelage is blackish brown at 
the base but otherwise bleached reddish brown, the ventral pelage similarly coloured but 
with the tips of the hairs more ochreous or buffy brown. In general the pelage is dense and 
long both above and below, almost shaggy. It is noticeably long and thick on the head as far 
anteriorly as a line just anterior to the eyes: the muzzle and the anterior part of the underside 
of the chin are sparsely covered with shorter hairs. A similar distribution of fur on the head is 
found in Nyctophilus microtis. 

So far as can be determined, the narial foliations of lophorhina consist of a raised ridge 
surmounting the nostrils, the second low ridge lying behind it divided medianly. A similar 
structure (type 2 of Thomas, 1915) characterizes Nyctophilus microtis. The ears closely 
resemble those of this species, and are moderate and full, with a rounded tip, the posterior 
margin of the ear evidently concave above the antitragal lobe. The basal third to one half of 
the medial surface of the conch is sparsely pilose as in N. microtis. As already discussed, the 
interaural band in the paratypes exactly resembles the rudimentary band found in that 
species, and both lophorhina and microtis have an essentially similar tragus. 

The skull of lophorhina closely resembles that of Nyctophilus microtis in both size and 
structure. The sagittal crest in the two paratypes examined (one, CM 209 1 with slightly worn 
teeth), however, is less developed frontally than in adults of N. microtis. The upper surface of 
the rostrum also provides a point of variation: in CM 2090 the rostrum has a very shallow, 
almost imperceptible median depression, while CM 2091 has a relatively deep, well-defined 
longitudinal sulcus. Mr Peter Aitken of the South Australian Museum has examined the 
holotype (M6404) and two of the remaining paratypes (M6402, M6403): the final paratype 
(M6401) is in alcohol with the skull in situ. He reports that M6402 has a relatively shallower, 
more saucer-shaped depression on the top of the rostrum than M6403 and M6404, which 
each have a very distinct, deep longitudinal rostral depression forming a more obvious 

There is a similar range of variation among the specimens representing Nyctophilus 
microtis and those referred to this species that we have examined, all from Papua New 
Guinea. The young adult holotype of bicolor ( = N. microtis bicolor) (BM(NH) 5.1 1.28.2) 
from an unknown locality near the coast on the Aroa River, c. 9 05' S, 146 48' E has a 
dorsally flattened rostrum with no more than the slightest suggestion of any median 
depression, while another immature (AMNH 108531) from Sogeri, 9 30' S, 147 33' E, the 
type locality of microtis itself, has such a depression only faintly indicated. The holotype of 
microtis (BM(NH) has a very shallow rostral depression much like that of CM 
2090. A further specimen (BM(NH) from Sogeri has a moderate longitudinal 
rostral depression similar to that of CM 209 1 but shallower, while an example (BM(NH) from Kokoda, 8 53' S, 147 44' E has a large, very deep median sulcus that is 
more greatly developed than the sulcus of this paratype of lophorhina. This last locality, 
incidentally, is but little distant from Mount Lamington at 8 55' S, 148 10' E, whence the 
original series of lophorhina was obtained. The collection of the American Museum of 
Natural History also includes a further six specimens from a variety of localities in Papua 
New Guinea that reflect this variation in the degree of development of the rostral depression. 
From the limited material available to us it does not seem to be correlated with geography, 
sex or with age (judged by tooth wear) once full maturity is achieved: the only specimens in 
which the rostral depression is virtually absent are clearly immature or young. 

In size lophorhina almost exactly resembles Nyctophilus microtis. Ranges of selected 
measurements of lophorhina (from McKean & Calaby, 1968), followed by those of adults of 
N. microtis in the British Museum (Natural History) and in the American Museum of 
Natural History (number of specimens examined in parentheses): length of forearm 39-40 
(6), 37-5^0-5 (8); length of ear 15-19-5 (6), 15-19 (7); condylobasal length 13-8-14-7 (5), 
13-5-14-5 (5); length of maxillary toothrow (c-m 3 ) 5-3-6-0 (5), 5-5-6-4 (8). 

Although small, the sample of specimens that we have examined leaves little doubt that 
lophorhina and microtis are conspecific, if indeed not synonymous, and also closely related 
to bicolor. However, the close proximity in eastern Papua of the type localities of these three 

278 J. E. HILL & K. F. KOOPMAN 

forms and the paucity of material from this area makes any further evaluation of their 
taxonomic validity difficult if not impossible. The differences between such specimens as we 
have seen are small and indicate that at best only subspecific separation can be justified. For 
the present, therefore, we are inclined to accept only a single species, Nyctophilus microtis, 
with the provisional recognition of three subspecies, N. m. microtis Thomas, 1888, N. m. 
bicolor Thomas, 1915 and N. m. lophorhina McKean & Calaby, 1968. 


A review of the characters ascribed to the vespertilionid genus Lamingtona McKean & 
Calaby, 1968 from New Guinea shows that it cannot be separated generically from the 
predominantly Austro-Papuan genus Nyctophilus and that there is no justification for its 
retention as a valid subgenus within Nyctophilus. Examination of paratypes of its type 
species L. lophorhina McKean & Calaby, 1968 together with further information from the 
original series indicates that it is conspecific and possibly synonymous with Nyctophilus 
microtis Thomas, 1888. Three subspecies of N. microtis (N. m. microtis, N. m. bicolor 
Thomas, 1915, N. m. lophorhina McKean & Calaby, 1968) are tentatively recognized in 
eastern Papua but lack of adequate material prevents any proper assessment of their valdity. 


Our thanks are due to Dr J. H. Calaby, of the Division of Wildlife Research, Commonwealth 
Scientific and Industrial Research Organisation, Canberra, Australia who made possible an 
examination of part of the original series of Lamingtona lophorhina, and to Mr Peter Aitken, 
Senior Curator at the South Australian Museum, Adelaide, Australia who examined the 
holotype and other relevant paratypes on our behalf. 


McKean, J. L. & Calaby, J. H. 1968. A new genus and two new species of bats from New Guinea. 

Mammalia 32 : 372-378, 3 figs, 2 tabs. 
Miller, G. S. 1907. The families and genera of bats. Bull. U.S. natn. Mus. 57 : i-xviii, 1-282, 49 figs, 14 

Tate, G. H. H. 1941. Results of the Archbold Expeditions. No. 40. Notes on vespertilionid bats. Bull. 

Am. Mus. nat. Hist. 78 : 567-597, 4 figs. 

Thomas, O. 1 888. Description of a new bat of the genus Nyctophilus. Ann. Mag. nat. Hist. (6), 2 : 226. 
1915. Notes on the genus Nyctophilus. Ann. Mag. nat. Hist. (8), 15 : 494^99. 

Manuscript accepted for publication 7 May 198 1 

The status of Hipposideros galeritus Cantor, 1846 
and Hipposideros cervinus (Gould, 1854) 
(Chiroptera: Hipposideridae). 

Paulina D. Jenkins & J. . Hill 

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


Considerable taxonomic confusion has surrounded the small leaf-nosed bats of Indo- 
Australia allied to Hipposideros galeritus Cantor, 1 846 from Penang Island, and numerous 
names have been proposed since that date for specimens or populations from this extensive 
region. Suggested classifications have varied from the recognition of three or more species to 
the definition of no more than one, the grouping of the various named forms changing 
accordingly. However it has long been apparent that the Bornean population of these bats 
seemed critical to any consideration of the groups of taxa as a whole, but for many years the 
relative lack of specimens from a variety of Bornean localities made critical assessment 
difficult. Moreover, this difficulty is compounded by much confusion in the literature. 

This study was initiated through the interest of the Earl of Cranbrook (formerly Lord 
Medway) who obtained specimens in Sarawak while with the Royal Geographical Society 
Expedition to Mount Mulu, 1977-1978. These although superficially referable to H. 
galeritus, on close examination seemed to represent two very similar but nevertheless 
distinct taxa. They have led to a further examination of specimens ascribed to H. galeritus or 
to its nominal allies in the collections of the British Museum (Natural History) and as a result 
it has proved possible to attempt some clarification of the status and relationships of the 
majority of the named forms previously associated in one way or another with this species. 
Representative material of a minority of described taxa or populations is either lacking or 
unavailable and to this extent our conclusions must remain provisional, although even so it 
has been possible to advance tentative opinions. At the same time we have endeavoured to 
indicate through detailed synonymies the wide variety of conflicting usages and differing 
opinions that exist in the literature of this relatively small group of bats, of which 
representatives are likely to be found in the majority of collections from Indo- Australia. 

Material and Methods 

One hundred and thirty-three specimens have been examined in the course of this study, all 
in the British Museum (Natural History) [BM(NH)]. For the most part they are identified 
by their registration numbers but a minority of specimens collected recently in southeastern 
Sulawesi has yet to be accessed and will be reported more fully in an account of the 
collection of which they form a part. By the courtesy of Dr C. Smeenk, we have also 
been able to borrow a series of critical specimens from the Rijksmuseum van Natuurlijke 
Historic, Leiden (RNH) which have proved very valuable. In addition, Professor J. D. Smith 
of California State University has generously provided measurements of a series of 
specimens that he obtained recently on the islands of the Bismarck Archipelago, off 
northeastern New Guinea. Qualitative assessments of external, cranial and dental characters 
are summarized in the systematic accounts. Measurements are in millimetres and were 

Bull. Br. Mus. nat. Hist. (Zool.) 41(5): 279-294 Issued 1 7 December 1 98 1 



made entirely by one of us (P.D.J.) with dial calipers. In the text those relevant in diagnosis 
appear with the following abbreviations: 

Length of forearm FA 

Length of 2nd phalanx of third and fifth digits III 2 , V 2 

Length of tail TL 

Length of skull from canine to occipital condyle (condylocanine length) CCL 

Greatest width across zygomata ZW 

Greatest mastoid width MW 

Greatest diameter of tympanic annulus TD 

Crown width across third molars M 3 -M 3 

Length of upper toothrow from canine to third molar inclusive C-M 3 

Width of third molar M 3 

Where series of specimens have been available the range of each measurement is given, 
preceeded in parentheses by the number examined and followed by the arithmetic mean, 
also in parentheses. Measurements of holotypes appear separately but are also included in 
the appropriate series. 


Although Hipposideros galeritus Cantor, 1846 and H. cervinus (Gould, 1854) are in general 
use as the earliest identifiable names in this small complex of bats, it is possible that both 
may be antedated by Rhinolophus crumeniferus Lesueur & Petit, 1807 (pi. 35), from Timor 
Island. The status of this name has been reviewed and discussed by Tate (1941 : 367, 
footnote, 382, 387), Laurie & Hill (1954 : 56), Oey & Feen (1958 : 230), Hill (1963 : 23) and 
Goodwin (1979 : 79). It is based solely on a coloured plate and no holotype appears to exist. 
There is very little doubt that the bats depicted belong to the galeritus complex and although 
the plate is insufficiently detailed, they are probably referable either to galeritus or to 
cervinus. There are no subsequent records of bats referable to either of these from Timor and 
for the present the name is best left unallocated. 

The taxonomic history of this small but involved grouping of bats thus begins with the 
description of Hipposideros galeritus Cantor, 1846 from Penang Island, Malaya, followed in 
quick succession by the recognition of cervinus Gould, 1854 from northern Australia, 
labuanensis Tomes, 1859 from Borneo, longicauda Peters, 1861 from Java and, rather later, 
of brachyotus Dobson, 1874 from India. Even at this early stage, considerable differences of 
opinion are apparent, Peters (1871) for example separating galeritus subgenerically from 
labuanensis, cervinus and longicauda but Dobson (1876, 1878) synonymizing labuanensis, 
longicauda and brachyotus into galeritus which, like cervinus, he considered a valid species. 
This author noted variations in the size of the noseleaf, tail length and wing insertion on the 
hindlimb in H. galeritus as he understood it, considering specimens of labuanensis and 
brachyotus intermediate between longicauda at one extreme and galeritus at the other. 
However, this variation is at least partially due to the fact that his H. galeritus is a composite 
of//, galeritus and H. cervinus as we define them, although not unexpectedly some of the 
features to which Dobson drew attention retain diagnostic significance. His association of 
labuanensis with galeritus, rather than with cervinus, has persisted until the present day and 
has led to many of the difficulties encountered in classifying this group of bats, since the 
majority of subsequent comparisons have been made against members of the Bornean 
population that labuanensis represents, rather than galeritus of which few specimens are 

Five further names have been proposed since Dobson's studies: batchianus Matschie, 
1901 from Batjan Island in the Molucca Islands, schneideri Thomas, 1904 from Sumatra, the 
posthumously published misorensis Peters, 1906 probably from Misor Island ( = Schouten 
Island) off northeast New Guinea, insolens Lyon, 1911 from Borneo and finally celebensis 
Sody, 1936 from Sulawesi. Among these the account by Lyon of insolens marks an 


important point since this author found insolens and 'galeritus' to occur sympatrically and 
thus demonstrated for the first time the existence of two distinct species in the Bornean 
population. Most later authors have endeavoured to recognize this in different ways but have 
been handicapped by the unavailability of type material, by a lack of specimens from critical 
areas or by reliance on the concept of labuanensis as representative of H. galeritus. On this 
last point for example, it seems clear from the measurements that they cite, that 'galeritus' as 
understood by Lyon (191 1), by Chasen (1931) when reporting specimens from Sabah or by 
Sody (1936) when using Bornean specimens for comparison is in fact the taxon that we 
recognize as H. cervinus labuanensis. Hipposideros galeritus and its allies have been listed, 
reviewed and discussed either entirely or in part by Chasen (1940), Tate (1941), Laurie & 
Hill (1954) and Hill (1963). 

Chasen (1940) admitted weaknesses in his arrangement of the Malaysian members of the 
complex but correctly maintained two separate taxa in Borneo. He recognized two species, 
H. longicauda with insolens as its subspecies and H. galeritus with subspecies labuanensis 
and schneideri. Although he had at one time examined the holotype of galeritus, Chasen had 
no access to it when writing and took for comparison 'galeritus' from Pahang which may 
well have been H. cervinus labuanensis as we understand it, leading him to associate 
galeritus with labuanensis and schneideri, rather than with insolens and longicauda which 
we have found to be its allies. 

The first attempt at a cohesive review of the entire complex was made by Tate (1941) who 
presented much useful information and many useful characters for distinguishing the taxa. 
Unfortunately it is difficult to extract a coherent view of their classification from this rather 
confusing paper. The caption to the distribution map (p. 365) for the galeritus group 
provides one opinion with the recognition of four species: H. galeritus with subspecies 
schneideri, H. longicauda, H. labuanensis and H. cervinus with subspecies insolens, 
celebensis and batchianus. The text however (p. 367) offers a different view, Tate observing 
that for the sake of consistency with the then prevailing view that galeritus extends through 
Sumatra, Java and Borneo to Celebes, it must be further extended to include cervinus of New 
Guinea and Australia. He concludes that brachyotus, labuanensis, insolens, celebensis, 
batchianus and cervinus would be more or less valid subspecies of galeritus, with schneideri 
and longicauda as derived species. Later (p. 369) he reaffirms that cervinus, batchianus, 
celebensis and insolens are no doubt conspecific but remarks that labuanensis and schneideri 
are good species. Finally (p. 391) labuanensis, celebensis and cervinus (also on p. 392) are 
considered subspecies of galeritus but insolens and schneideri distinct species. Moreover 
(p. 391) he reports both galeritus and schneideri from North Pagi Island in the Mentawei 

Tate (p. 367) also suggested that the holotype of H. galeritus consisted of a mismatched 
skin and skull, although apparently he only saw a photograph of the skull and had not 
examined the skin but instead relied upon Cantor's description. He considered that the skin 
had distinct affinities with cervinus but that the skull appeared to be that of a bat of the 
bicolor group. As a result he designated the skin as the type of Cantor's name. This opinion 
was accepted by Laurie & Hill (1954) and more particularly by Hill (1963) who concluded 
that the skull agreed closely with that of the cranially similar species H. ater, a member of his 
bicolor subgroup, otherwise readily distinguished from H. galeritus by the absence of lateral 
supplementary leaflets. Subsequent examination has shown clear cranial differences between 
H. ater and H. galeritus and with the advantage of additional specimens from Malaya and 
Thailand, that agree closely both externally and cranially with the holotype of galeritus, we 
have been able to establish that these assumptions are erroneous and that it is very unlikely 
that any mismatch has occurred. 

Laurie & Hill (1954) endeavoured to reconcile the varying opinions advanced by Tate 
with the specimens then available in the British Museum (Natural History) although 
concerned chiefly with New Guinea and Sulawesi. These authors recognized two species, H. 
galeritus extending eastwards perhaps to Sulawesi, H. cervinus from Borneo to the Solomon 
Islands, with subspecies insolens, celebensis and batchianus. They considered the eastern 



cervinus to be slightly less specialized than the western galeritus and its allies and, although 
differing in detail, their concept has elements in common with that advanced in this paper. 

Hill (1963) in reviewing the complex as a whole, adopted much of the treatment by Laurie 
& Hill and carried it further in uniting galeritus and cervinus into one species, galeritus, of 
which all other forms were made subspecies; insolens becoming a synonym of labuanensis, 
their differences being thought to reflect individual variation. This arrangement has been 
widely accepted but proves to be incorrect. 

Systematic Section 

The review by Hill (1963, p. 19) of the galeritus subgroup and (p. 52) of//, galeritus as this 
author understood it provides an account of the major external, cranial and dental features of 
bats here attributed to H. galeritus or H. cervinus but since it is based on a composite of both 
further definition is needed. 

Hipposideros galeritus Cantor, 1 846 

DIAGNOSIS AND DESCRIPTION. Antitragus subangular, tall, nearly one third of ear length in 
height; antitragal projection absent (a small antitragal fold towards rear of antitragus may 
suggest an antitragal projection but disappears when ear is flattened); internarial septum 
linear, slightly swollen (Fig. 1); upper margins of narial lappets slightly lobed; intermediate 
leaf simple, slightly inflated medianly, expanded laterally to form a small lappet on either 
side, nearly as wide or wider than posterior leaf; wing elements relatively long; tail long; 
generally five caudal vertebrae (exceptionally four or six); tail membrane large and extensive. 
Skull with long braincase and short toothrow relative to condylocanine length (Fig. 2); 
rostrum inflated, in profile swollen above interorbital region which is very constricted and 
shortened anteroposteriorly, with an angular outline when viewed from above; zygomata 
slender with prominent, short, abruptly rising jugal process; greatest zygomatic width 
generally less or subequal to mastoid width; anterorbital foramen subtriangular to suboval; 



Fig. 1 Frontal view of noseleaf of: (a) Hipposideros galeritus insolens; (b) Hipposideros cervinus 




braincase globose, frontal and parietal regions divided by a shallow depression; interparietal 
moderately swollen; mastoid well developed; post-palatal spicule obsolescent or absent; 
anterior sphenoidal ridges equal in length to posterior ridges to enclose a diamond-shaped 
sphenoidal depression; width of cochleae exceeding their distance apart, each one third 
concealed by a large tympanic annulus whose greatest diameter (2 -70-3 '00 [2 -8 3]) is a little 
less than or approximately equal to the distance between the annuli across the sphenoidal 



Fig. 2 Lateral view of the skull of: (a) Hipposideros galeritus galeritus; (b) Hipposideros cervinus 





$3v-&v.i .-: 

>S * . 'V . 




>^?^ :.-^ 

-^/iSfl* ! ; * 



Fig. 3 Ventral view of external structure of right auditory region of skull of: (a) Hipposideros 
galeritus insolens; (b) Hipposideros cervinus labuanensis. 

depression (except in H. g. longicaudus see discussion of this subspecies), annulus acutely 
angled to longitudinal axis of skull (Fig. 3); upper incisors bicuspid, outer lobes 
sometimes small; M 3 with well developed third commissure, fourth commissure small but 
evident; crown area of inner lower incisors less than crown area of outer pair. 

DISTRIBUTION. India and Sri Lanka to Borneo and Java. 

Hipposideros galeritus galeritus Cantor, 1 846 

Hipposideros galeritus Cantor, 1846:183 (Pinang [ = Penang I.], Malaya); Lekagul & McNeely, 
1977 : 167 (Satun Province, Thailand); Tate, 1941 : 367 (discussion, lectotype designated). 

Rhinolophus (Phyllorrhina) galeritus, Wagner, 1855 : 659 (Pinang [ = Penang I.], Malaya). 

Phyllorrhina galerita, Fitzinger, 1870 : 885 (Malay Peninsula; Pinang [ = Penang I.]). 

Phyllorhina galerita, Peters, 1871 : 316 (Pinang [ = Penang I.]); Dobson, 1876 : 69, 1878 : 141 (in part, 
Pinang [ = Penang I.]). 

Hipposideros galeritus galeritus, Chasen, 1940 : 46 (in part, Malay States); Tate, 1941 : 365 (mapped), 
367 (type material discussed), 391 (N. Pagi, Mentawei Is.); Laurie & Hill, 1954 : 56 (in part, Penang, 
Malay States); Hill, 1963 : 53 (in part, Malay Peninsula); Medway, 1969 : 29, 1978 : 29 (in part, 
Penang I.). 

Hipposideros galeritus group, Tate, 1941 : 358, 360, 364 (discussed). 

[Hipposideros] galeritus, Tate, 1941:359 (listed), 369 (discussed), 384 (type material, further 

HOLOTYPE. Adult cf BM(NH) 79. 1 1 .2 1 .85. Slightly damaged skin, skull in good condition but 
left premaxilla and incisor missing, left zygoma broken, occipital region slightly damaged. 
Penang Island, Malaya. Originally in Indian Museum, Calcutta. 


OTHER MATERIAL. MALAYA: 9 BM(NH) 61.1718 (in alcohol, skull extracted) Bukit Kutu, 
Selangor, 1061 m; THAILAND c? 9 BM(NH) 78.2351-2 (skins and skulls) Wang Bla Chan, 
Muang, Satun Province. 

DIAGNOSIS. As species diagnosis and description; relatively small. 

Measurements of holotype: FA 46'3; III 2 19'2; V 2 9'5; CCL 14-0; ZW 7'9; MW 8'9; TD 
2-9; M 3 -M 3 5-6; C-M 3 5-3; M 3 1 -2. 

Measurements of the series examined: FA (4) 39-(M8'l (43*95); III 2 (4) 19-0-2O7 (19-80); 
V 2 (4) 9-5-10-7 (10-15); TL (3) 36-0, 40'0, doubtfully 49-0; CCL (4) 14-0-14-5 (14-30); ZW (4) 
7-9-8-3 (8-15); MW (4) 8'3-8'9 (8'68); TD (4) 2-80-2-90 (2-88); M 3 -M 3 (4) 5'5-5'7 (5-61); 
C-M 3 (4) 5-3-5-6 (5-46); M 3 (4) 1-20-1-35 (1-26). 

REMARKS. Dobson (1876, 1878) evidently saw the holotype of Hipposideros galeritus, 
probably at the Indian Museum, Calcutta (it was presented to the British Museum (Natural 
History) in 1879) although (1876; 198) he does not list it from that collection. Since then it 
appears to have been commented upon only by Tate ( 1 94 1 ) and Hill ( 1 963). 

Tate (1941) considered that the skin and skull of the holotype of H. galeritus were 
mismatched, although he had not examined the skin and apparently only had a photograph 
of the skull, and stated that the skin alone represented Cantor's type. However, the skull 
associated with this skin has an inflated rostrum, slender zygomata and U-shaped palation, 
all features distinguishing it from the bicolor subgroup, to which Tate and subsequently Hill 
(1963) thought it to belong and it is very similar to the skulls of other specimens of//. 
galeritus as understood in this paper from India, Sri Lanka, Thailand, Malaya and 
Borneo. In all, the zygomatic width is less than or subequal to the mastoid width, a feature 
otherwise characteristic of//, bicolor and its close allies. Tate also thought that the skin had 
affinities with H. cervinus. However, it has been possible to examine the ears and noseleaf in 
some detail after thorough soaking and there is clearly a tall, angular antitragus lacking any 
posterior antitragal projection, a prominent internarial septum and well developed, broad 
intermediate leaf, features which are characteristic of our concept of galeritus but not found 
in cervinus. Therefore there seems little to substantiate the contention that a mismatch of 
skin and skull has occurred and Tate's designation (p. 367) of the skin as the lectotype of H. 
galeritus should be discounted. 

Lekagul & McNeely (1977) provided useful photographs of the noseleaf and skull, with 
notes on ecology and behaviour. However, those remarks extrapolated from Medway (1958) 
probably refer to //. cervinus (H. c. labuanensis) as understood here, rather than to H. 

DISTRIBUTION. MALAYA: Selangor; Penang Island. THAILAND: Satun Province. 

Hipposideros galeritus brachyotus (Dobson, 1874) 
Phyllorhina brachyota Dobson, 1874 : 237 (Central India). 
Phyllorhina galerita, Dobson, 1876 : 69 (in part, Central India; Deccan). 
Hipposideros galeritus, Blanford, 1888:287 (Indian Peninsula; Ceylon [=Sri Lanka]); Brosset, 

1962 : 618 (Western and Central India: Chikalda; Ellora; Kanheri; Bombay; Bedsar Caves; Badomi). 
Hipposideros brachyotus, Wroughton, 1913 : 36 (Kanara); 1915 : 85, 101 (Bihar; Ceylon [ = Sri Lanka]; 

Ryley, 1914 : 689 (Gujerat); Phillips, 1923fl : 274, 19236 : 155, 1924 : 25, 1935 : 94 (Ceylon [ = Sri 

[Hipposideros] brachyotus, Tate, 1941 : 358 (listed), 364 (mentioned), 369 (discussed), 381 (listed, 

Hipposideros galeritus brachyotus, Tate, 1941 : 367 (discussion); Hill, 1963 : 54 (Ceylon [ = Sri Lanka]; 

Mysore; Bombay; Bengal). 

HOLOTYPE. Adult rf BM(NH) In alcohol in fair condition, wing membranes 
damaged, skull extracted, in good condition, right upper incisor, left upper canine and left 
anterior upper premolar (PM 2 ) missing. Central India. 

OTHER MATERIAL. INDIA: BM(NH) (skin and skull) Lingasoogar, N.N.W. of 


Bellary; <fg BM(NH) (in alcohol, skulls extracted) Konkan; <S BM(NH) (skin and skull) Honawar, Kanara; 3 BM(NH) (skin and skull) Gaya, 
Bihar; 9 BM(NH) (skin and skull) Danta, Gujerat; SRI LANKA: rf BM(NH) (skin and skull) Matugama; rf $ BM(NH) (skins and skulls) 
Kala-Oya; BM(NH) 66.5525 (skin and skull) Dammeria, Passara, Uva. 

DIAGNOSIS. Similar in structure and size to H. g. galeritus but the material available is 
insufficient to assess its subspecific validity. 

Measurements of holotype: FA 45-6; IIP 18'6; V 2 12-2; TL 35-6; CCL 14*2; ZW 8'6; MW 
8-7; TD 2-70; M 3 -M 3 5-8; C-M 3 5-4; M 3 1'45. 

Measurements of the series examined: FA (10) 44-0^8-4 (46*05); IIP (10) 18-3-21-6 
(1 7-09); V 2 (10) 9-3-12-2 (10'97); TL (10) 30'0-37'5 (34-08); CCL (10) 14- 1-1 5-4 (14-67); ZW 
(10) 8-4-9-1 (8-76); MW (10) 8-5-9-1 (8'79); TD (10) 2'70-3-QO (2-83); M 3 -M 3 (10) 5-6-6-2 
(5-88); C-M 3 (10) 5-3-5-9 (5-67); M 3 (10) 1-30-1-50 (1-43). 

REMARKS. This taxon has a relatively simple historical background, different authors 
considering it to be either specifically distinct or a subspecies of//, galeritus. Phillips (19230, 
19236, 1924, 1935) provided notes on its behaviour, reproduction and ecology in Sri Lanka, 
Brosset (1962) gave similar comments on Indian populations and included a frontal 
photograph of the head. Both maintained that these bats have been observed only as isolates 
or in very small family groups, Brosset considering that they avoided other species. This is an 
interesting point that may reflect the fact that, in contrast to //. cervinus, H. galeritus is 
poorly represented in trapping samples and is comparatively rare in museum collections. 

DISTRIBUTION. INDIA: Gujerat; Mysore; Bihar. SRI LANKA. 

Hipposideros galeritus insolens Lyon, 1911 

Hipposideros insolens Lyon, 1911 : 129 (Upper Pasir River, southeastern Borneo); Chasen, 1931 : 112 

(Baturong Caves, N. Borneo); Tate, 1941 : 391 (Perboewa, N.W. Borneo). 
Hipposideros longicauda insolens, Chasen, 1940 : 46 (Borneo). 
[Hipposideros] insolens, Tate, 1941 : 359 (listed), 368 (discussed), 384 (holotype). 
Hipposideros cervinus insolens, Tate, 1941 : 365 (mapped); Laurie & Hill, 1954 : 56 (discussed). 
Hipposideros galeritus insolens, Tate, 1941 : 367 (discussed), 369 (systematic status). 
Hipposideros galeritus labuanensis, Davis, 1962 : 39 (in part?, Sandakan, Borneo); Hill, 1963 : 54 (in 

part, Borneo); Medway, 1965 : 57, 1977 : 51 (in part, Borneo). 

HOLOTYPE. Adult cf USNM 154389. In alcohol, skull extracted, both in good condition 
(Poole & Schantz, 1942). Near Lowatsi, on the Upper Pasir River, southeastern Borneo. Not 

OTHER MATERIAL. BORNEO: BM(NH) (should be in alcohol but not found, skull 
only) Baram, Sarawak; 3d 1 49 BM(NH) 51.120-126 (in alcohol, skulls extracted) Long Lama 
Caves, Baram River, Sarawak; c?9 BM(NH) 59.150-1 (skins and skulls) Gua Bungoh, Bau, 
Sarawak; 39 BM(NH) 75.1896-8 (in alcohol) Niah Cave, Gunong Subis, 4th Division 
Sarawak; 9 BM(NH) 78.123 (in alcohol) Gunong Mulu, Sarawak, 500m; 3d 1 4 9 BM(NH) 
78.2495-2501 (in alcohol) inland from Sangkulirang, mouth of River Baai, E. Kalimantan, 
059'N, 117'58'E. 

DIAGNOSIS. Larger than H. g. galeritus or //. g. brachyotus with more robust skull; 
interorbital region longer and less angular in shape; upper incisor occasionally with reduced 
outer lobe. 

Measurements of the series examined: FA (19) 45-7-52-2 (48'94); IIP (19) 17-1-21-9 
(19-46); V 2 (19) 10-1-13-2 (10-81); TL (19) 28-4-37-6 (33-81); CCL (4) 14-5-1 5-7 (14-90); ZW 
(7) 9-0-9-3 (9-10); MW (6) 8-8-9-2 (8-98); TD (7) 2*75-2-90 (2-85); M 3 -M 3 (9) 6-1-6-6 
(6-37); C-M 3 (7) 5-9-6:3 (6-19); M 3 (10) 1-50-1-70 (1-62). 

REMARKS. Lyon (1911) distinguished H. insolens from H. galeritus (clearly = //. cervinus 


labuanensis as understood here, from his measurements) by its distinctly longer forearm, tail 
and tibia, although he considered the skulls and dentition of the two taxa to be indistinguish- 
able. Chasen (1931) also considered insolens a distinct species but later (1940) evidently 
revised this view and listed it as a subspecies of H. longicauda from Java. Despite the 
confusion of Tate's (1941) treatment of insolens, this author made an important contribution 
in distinguishing it from labuanensis and listing a number of cranial characters useful in 
separating the two taxa. This distinction was maintained by Laurie & Hill (1954) who 
regarded insolens as a subspecies of H. cervinus and labuanensis as a subspecies of H. 

More recently, Davis (1962) synonymized insolens and labuanensis but the measurements 
that he gave suggest that his sample consisted only of the latter and included no specimens 
referable to insolens. This view was followed by Hill (1963) who also synonymized insolens 
and labuanensis and further suggested that insolens, as understood by Lyon and Tate, might 
well refer to large individuals of labuanensis, the differences observed by Tate reflecting 
individual variation. However, although these features may be more variable than Tate 
thought, in combination with others listed above but not noted by Tate, they establish two 
species in Borneo, one represented by H. galeritus insolens, the other by H. cervinus 

DISTRIBUTION. BORNEO: Sarawak; Brunei; E. & S.E. Kalimantan. 

Hipposideros galeritus longicaudus (Peters, 1861) 

Phyllorhina longicauda Peters, 1861 : 708; 1871 : 321 (Surakarta, Java); 1906: pi. 5L, fig. 3 (head 


Phyllorhina galerita, Dobson, 1876 : 69; 1878 : 141 (in part, Java). 
Hipposideros galerita longicauda, Sody, 1930 : 270 (C. Java). 
Hipposideros longicauda longicauda, Chasen 1940 : 46 (Java) 

[Hipposideros] longicauda, Tate, 1941 : 359 (listed), 369 (systematic status), 385 (holotype, remarks). 
Hipposideros longicauda, Tate, 1941 : 365 (mapped), 367 (systematic status). 
Hipposideros galeritus longicauda, Hill, 1963 : 56 (Java). 

HOLOTYPE. Attempts to locate the holotype in the Museum fur Naturkunde der Humboldt- 
Universitat in Berlin (where it might be expected to be housed), the Forschungsinstitut und 
Natur-Museum Senckenberg, Frankfurt and the Staatliches Museum fur Naturkunde, 
Stuttgart, have failed. 

OTHER MATERIAL. JAVA: c? 9 RNH 15103-7, 15168-9, 15258, 15319, 15330 (skins and 
skulls) Panganderan, Central Java; 3$ RNH 1 5 1 70, 1 5259 Toenggilis, West Java. 

DIAGNOSIS. This taxon is poorly known with few specimens or records. As species 
diagnosis but averaging slightly smaller than H. g. galeritus and H. g. brachyotus, 
considerably smaller than H. g. insolens and differing from all other subspecies in having 
small tympanic annuli (2-4-2-5[2'46]). The most noticeable feature is the long tail of the 
holotype, although Sody (1930) reported four specimens which have rather shorter tails. 
According to this author the skulls of these Javan examples are strikingly larger than those of 
the Sulawesian specimens that he subsequently described (1936) as celebensis, although the 
series that we have examined does not support this. 

Measurements of the holotype: FA 48 (Peters, 1861), 48-3 (Dobson, 1876); TL 44 (Peters, 
1861), 40-6 (Dobson, 1876), TL of four additional specimens 32-36 (Sody, 1930). 

Measurements of the series examined (skin measurements taken from dried skins): FA (II) 
45-3^8-0 (46-50); IIP (II) 16'5-19'2 (17-76); V 2 (II) 9-5-11-4 (10*64); TL not measured 
because vertebrae have been removed from dried skins; CCL (II) 13-6-14-4 (13*93); 
ZW (II) 8-1-8-6 (8-29); MW (II) 8'3-8'8 (8'53); TD (II) 2-4-2-5 (2-46); M 3 -M 3 (10) 
5-4-5-8 (5-63); C-M 3 (II) 5'2-56 (5*45); M 3 (10) 1-3-1-45 (1'39). 


REMARKS. Peters (1861) and Dobson (1876) seem to have been the only authors who have 
made a direct examination of the original material. There are, however, excellent drawings 
of the head among the posthumously published drawings of Peters (1906) which, with the 
original account, establish longicaudus as a member of the galeritus/cervinus complex. It has 
a tall, subangular antitragus and large noseleaf, with prominent internarial septum and broad 
intermediate leaf, all features which with the long tail are characteristic of H. galeritus. A 
small projection illustrated towards the rear of the antitragus may be an antitragal projection 
similar to that of H. cervinus but may equally result from a fold in the ear membrane, 
disappearing when the ear is flattened. Such a fold is found in some individuals of H. 
galeritus. The illustration by Peters may not be based on the specimen measured in his 
original account, since he appears to have had more than one example. The frontal pit in 
males is described by Peters (1861) yet Dobson (1876) stated that he had examined the adult 
female type, indicating the existence of at least two specimens. Chasen (1940) and Tate 
(1941) regarded H. longicauda as a distinct species; Tate considered that it agreed with the 
galeritus group but was probably atypical. 

DISTRIBUTION. JAVA: central and west. 

Hipposideros cervinus (Gould, 1 854) 

DIAGNOSIS AND DESCRIPTION. As in H. galeritus but antitragus rounded, short, approxi- 
mately one-eighth of ear length in height; an acutely pointed antitragal projection; inter- 
narial septum low, narrow, not swollen (Fig. 1); narial lappets well developed, not lobed; 
intermediate leaf slightly inflated dorsally, slightly expanded laterally, narrower than either 
anterior or posterior leaves; forearm, distal phalanges and tail not especially lengthened; 
usually four caudal vertebrae, sometimes five, rarely six. 

Skull with short braincase and long toothrow relative to condylocanine length (Fig. 2); 
rostrum only slightly inflated, in profile normally level with interorbital region, with a 
rounded outline when viewed from above; zygomata with rather long, gradually sloping jugal 
process; zygomatic width greater than mastoid width; anterorbital foramen suboval to 
ellipsoidal; interparietal only slightly swollen; mastoid uninflated; a blunt median post- 
palatal spicule sometimes present; anterior sphenoidal ridges longer than posterior ridges to 
enclose a pyriform sphenoidal depression; cochleae only slightly wider than their distance 
apart; tympanic annulus overlaying less than one quarter of the cochlea; greatest diameter of 
annulus (2-05-2-65 [2-36]) one half to three quarters of the distance between the annuli 
across the sphenoidal depression; annulus shallowly angled to longitudinal axis of skull (Fig. 
3); upper incisors small, at most only weakly bicuspid, with rounded inner lobe, outer lobe 
when present minute; M 3 slightly reduced, third commissure shortened; fourth commissure 
absent; crown area of inner lower incisor equal to or slightly less than crown area of outer 

DISTRIBUTION. Malaya, Sumatra and Philippine Islands east to New Hebrides. 

Hipposideros cervinus cervinus (Gould, 1 854) 

Rhinolophus ? cervinus Gould, 1 854; pi. 34 (Cape York and Albany I., Australia). 
Rhinolophus (Phyllorrhina) cervinus, Wagner, 1 855 : 66 1 (Cape York and Albany I., Australia). 
Phyllorrhina cervina, Fitzinger, 1870 : 873 (Cape York and Albany I., Australia); Peters, 1871 : 321 (in 

part, Cape York and Albany I., Australia; Am I.). 
Phyllorhina cervina, Dobson, 1878 : 142 (Cape York and Albany I., Australia; Aru Islands; 

Waigiou Island; New Guinea). 

Hipposideros galerita galerita, Sody, 1930 : 267 (Mampoegrotto, S. Celebes [ = Sulawesi]). 
Hipposideros cervinus, Sanborn, 1931 : 24 (New Guinea: Marienberg; Santa Cruz Is.: Fenvaloa; New 

Hebrides: Espiritu Santo, Efate, Malo); Sanborn & Nicholson, 1950 : 331 (New Hebrides: Espiritu 



Hipposideros celebensis Sody, 1936 : 47 (Mampoe Cave, S. Celebes [ = Sulawesi]). 

(?) Hipposideros galeritus galeritus. Shamel, 1940 : 354 (Peling [ = Peleng] I.; Gimpoe or Bada, Celebes 

[ = Sulawesi]). 
[Hipposideros] cervinus, Tate, 1941 : 359 (listed), 368 (discussed, S. New Guinea: Fly R.; Aru I.; New 

Hebrides); 381 (holotype). 
Hipposideros cervinus cervinus, Tate, 1941 : 365 (mapped), 369 (systematic status); Laurie & Hill, 

1954:56 (New Guinea; Japen [ = Jobi] I.; Kei Is.; Aru Is.; Trobriand Is.; Kiriwina; Bismarck 

Archipelago: New Ireland; Solomon Is., Guadalcanar, Bougainville); Hill, 1956 : 78 (Solomon Is.: 

Rennell); McKean, 1972 : 26 (New Guinea: Papua). 
Hipposideros cervinus celebensis, Tate, 1941:365 (mapped); Laurie & Hill, 1954:57 (Celebes 

[ = Sulawesi]). 
Hipposideros galeritus cervinus, Tate, 1941:367 (discussed), 391 (New Guinea: W. Irian), 392 

(variation); Hill, 1963 : 57 (New Guinea: Waigeo I.; Japen [ = Jobi] I.; Kei Is.; Aru Is.; N. Australia; 

Bismarck Archipelago: New Ireland; Trobriand Is.: Kiriwina; Solomon Is.: Bougainville, 

Guadalcanar, Fauro, Russell, Rennell; Santa Cruz Is.: Vanikoro; New Hebrides: Espiritu Santo, 

Efate, Fenvaloa); Hill, 1971 : 574 (Solomon Is.: San Jorge I.). 

[Hipposideros] celebensis, Tate, 1941 : 359 (listed), 368 (S. Celebes [ = Sulawesi]), 38 1 (holotype), 
Hipposideros galeritus celebensis, Tate, 1941 : 367 (discussed), 369 (systematic status), 391 (S.Celebes 

[ = Sulawesi]); Hill, 1963 : 56 (Celebes [ = Sulawesi]). 
Hipposideros galeritus labuanensis, Laurie & Hill, 1954:56 (in part, ? extending to Celebes 

[ = Sulawesi]). 
Hipposideros galeritus, Koopman, 1979:9 (Karkar I., Crown I., off N.E. coast of New Guinea; 

Bismarck Archipelago: New Britain). 

HOLOTYPE. Adult 9 BM(NH) 55.1 1.7.13. Skin in good condition, rostrum and mandible of 
skull. Caves on Albany Island, Cape York, N. Australia. 

and skull), 9 BM(NH) (in alcohol) Cowal Creek, 9 BM(NH) unregistered (in 
alcohol) Cape York; NEW HEBRIDES: Id 1 39 BM(NH) 73.1361^4 (in alcohol, skulls 
extracted) Amok, Malekula Is., ^9 BM(NH) unregistered (in alcohol, skulls extracted) 
Espiritu Santo; SANTA CRUZ Is.: <? BM(NH) 55.359 (skull) Vanikoro; SOLOMON 
ISLANDS: 2rf 1 9 BM(NH) (in alcohol, skulls extracted) Fauro I, rf BM(NH) (in alcohol, skull extracted) Aola, Guadalcanar, rf 9 BM(NH) (skins 
and skulls) Talena, Russell I., 9 BM(NH) 54.883 (skin and skull) Rennell I., 1 IcM 39 BM(NH) 
67.2063-86 (in alcohol, skulls extracted) Talise, San Jorge I.; TROBRIAND ISLANDS: 9 
BM(NH) 1 (in alcohol, skull extracted) Kiriwina; BISMARCK ARCHIPELAGO: 
BM(NH) (skin and skull) New Ireland; NEW GUINEA: BM(NH),5 
(skins and skulls), BM(NH) (skins and skulls) Jobi I., 4cf 29 BM(NH) (in alcohol, skulls extracted) Haveri, BM(NH) (skull) Arfak Mts., 8cf 
89 BM(NH) 33. 6.1.10-25 (in alcohol, skulls extracted), BM(NH) 69.325 (skin and skull) 
Javavere Caves, Papua, <?9 BM(NH) 69.326-7 (skins and skulls) E. of Port Moresby, Papua, 
6rf 89 BM(NH) 80.618-631 (in alcohol) Siboma, Morobe Province; ARU ISLAND: + \6<? 
+ 109 BM(NH) (in alcohol and skins and skulls) Wasi; KEI ISLAND: ^9 
BM(NH) (in alcohol, skulls extracted); SULAWESI: 2rf 39 BM(NH) 
unregistered (in alcohol, skulls extracted), paratypes of H. celebensis tf<? RNH 
29297, 29303 (skins and skulls) Mampoe Grot, 20 km from Watoe Pone. 

DIAGNOSIS. As species diagnosis and description. Variable in size but generally relatively 
small. Specimens from Sulawesi, Aru and Kei Islands have a blunt post-palatal spicule 
which is lacking in those from New Guinea, Cape York, Solomon Islands and New Hebrides. 

Measurements of holotype: M 3 -M 3 6-3; C-M 3 6*0; M 3 1-7. 

Measurements of the series examined: FA (37) 40-4-48-2 (45-70); IIP (37) 14-6-18-8 
(16-81); V 2 (37) 8-7-1 1-1 (10-02); TL (37) 22-8-31-0 (27-16); CCL (25) 13-8-15-6 (14-44); ZW 

(26) 8-7-10-0 (9-18); MW (25) 8-2-4-2 (8-64); TD (24) 2-15-2-60 (2-29); M 3 -M 3 (27) 5-7-7-0 
(6-27); C-M 3 (27) 5-5-6-3 (5-91); M 3 (27) 1-4-1-7 (1-51). 

REMARKS. Dobson (1878) distinguished H. cervinus from H. galeritus, with which he 


synonymized labuanensis, here regarded as a subspecies of cervinus. Among more recent 
authors, Tate (1941) provided comments on a number of specimens from New Guinea and 
remarked that two colour phases occurred, one red the other grey, and that there was some 
geographical variation in size. This author variously considered cervinus a valid species or a 
subspecies of//, galeritus. Laurie & Hill (1954) attempted to formalize the uncertainties of 
Tate's taxonomic treatment of cervinus, considering it a distinct species closely allied to //. 

Initially Sody (1930) referred Sulawesian specimens to H. galeritus galeritus but after 
comparison with specimens he thought represented Bornean galeritus, he decided (1936) 
that the Sulawesian specimens were specifically distinct, naming them H. celebensis. It is 
impossible to determine the identity of his comparative material from the measurements 
cited and it may have consisted of H. cervinus labuanensis, H. galeritus insolens or even a 
mixture of both. No morphological characters are described in either account. Some of the 
measurements given by Sody for celebensis, such as the length of the tail, the zygomatic and 
mastoid widths and the length of the upper toothrow are apparently comparable with those 
used in this study. However our measurements of two of the paratypes of celebensis conflict 
with those quoted by Sody, for instance the upper toothrow length which we find greater 
than given by Sody while the zygomatic and braincase breadths are less than his values. Since 
the two paratypes unquestionably belong to //. cervinus, agreeing closely with specimens of 
this species obtained recently in Sulawesi, it seems reasonable to assume that the entire series 
of celebensis, including the holotype, also belong to H. cervinus. 

Shamel (1940) recorded specimens (in the National Museum of Natural History, 
Washington) from Sulawesi and Peleng Island as //. galeritus galeritus but gave no 
diagnostic information, although Tate later observed (1941 : 368) that the Peleng Island 
specimens exactly resembled celebensis. Tate stated that a series of specimens (in the 
American Museum of Natural History, New York) from S. Sulawesi, which he also referred 
to celebensis, agreed in most respects with insolens although smaller in size but approached 
labuanensis in a shorter M 3 with an incomplete cusp pattern. Tate allocated celebensis 
variously to //. galeritus (p. 367,369, 391) or to//, cervinus (p. 365) and it is impossible from 
these remarks to decide to which species these specimens from Peleng Island and Sulawesi 
belong or if they are a mixture of both species. Only further examination will determine their 
status. Laurie & Hill (1954) treated celebensis as a subspecies of//, cervinus but suggested 
that the specimens recorded by Shamel seemed most likely to belong to labuanensis (which 
they considered to be a subspecies of galeritus), evidently assuming that two species occur on 
Sulawesi. Our investigations suggest that only one species, //. cervinus cervinus, occurs there. 

DISTRIBUTION. The majority of islands and island groups from Sulawesi to the New Hebrides 
and North Australia. 

Hipposideros cervinus labuanensis (Tomes, 1 859) 

Phyllorrhina labuanensis Tomes, 1859 : 537 (Labuan I. and Sarawak, Borneo). 

Rhinophylla labuanensis, Gray, 1 866 : 82 (no locality given). 

Phyllorhina labuanensis, Peters, 1871 : 321 (Labuan I. and Sarawak, Borneo). 

Phyllorhina galerita, Dobson, 1876 : 69, 1878 : 141 (in part, Labuan I., Borneo). 

Hipposideros galeritus, Thomas, 1895 : 108 (Engano I., W. of Sumatra); Lyon, 191 1 : 129 (Panebangan 
I., Pangkallahan River, upper Pasir River, Borneo). 

Phyllorhina speoris, Jentink, 1897 : 52 (C. Borneo). 

Hipposideros schneidersi (misprint = schneideri) Thomas, 1904:722 (Sukaranda, Sumatra); Tate, 
1941 : 367, 369 (systematic status), 391 (N. Pagi I., Mentawei Is.). 

Hipposideros galeritus galeritus. Chasen, 1931 : 1 1 1 (N. Borneo); Chasen, 1940 : 46 (in part, Malay 
States; Rhio [ = Riouw] Archipelago; Banka I.; S. Natuna Is.); ? Sanborn, 1952; 104 (Mindanao I., 
Philippine Is.); Laurie & Hill, 1954:56 (in part, Malay States); Davis, 1961 : 90 (Malaya); Hill, 
1963 : 53 (in part, Malay Peninsula; Riau [ = Riouw] Archipelago; Banka; S. Natuna Is.); Medway, 
1 969 : 29, 1 978 : 29 (in part, Malaya; Singapore I.). 

Hipposideros galeritus labuanensis, Chasen, 1940 : 47 (Borneo); Tate, 1941 : 367 (systematic status), 


391 (N.W. Borneo); Laurie & Hill, 1954:56 (in part, Borneo); Davis, 1962:39 (Borneo); Hill, 

1963 : 54 (in part, Borneo; Labuan I.; Mindanao I. Philippine Is.); Medway, 1965 : 57, 1977 : 51 (in 

part, Borneo). 
Hipposideros galeritus schneideri, Chasen, 1940 : 47 (Sumatra; ? Engano I.); Tate, 1941 : 365 (mapped); 

Hill, 1963 : 54 (Sumatra; Engano I.; Sipora I.; N. Pagi, Mentawei Is.). 
[Hipposideros] labuanensis, Tate, 1941 : 359 (listed), 368 (discussed), 385 (holotype). 
[Hipposideros] schneideri, Tate, 1941 : 359 (listed), 368 (discussed; Sumatra, Pagi), 389 (holotype). 
Hipposideros labuanensis, Tate, 1941 : 365 (mapped), 369 (systematic status). 

HOLOTYPE. Adult BM(NH) Skin in good condition, skull lacking premaxillae, 
cranium damaged. Labuan Island, Borneo. 

OTHER MATERIAL. BORNEO: 4rf 39 BM(NH) (in alcohol, skulls extracted), dtf 99 
BM(NH), BM(NH), BM(NH), BM(NH) 61.1046-9 (skins and 
skulls) Sarawak, rfcf 99 BM(NH), BM(NH), BM(NH), BM(NH) (in alcohol and skins and skulls) Baram, Sarawak, 23 BM(NH) 95.1 1.5.2-3, 6d* 
BM(NH), 2rf 19 BM(NH) 59.157-9, 3d 1 69 BM(NH) 62.21 18-26 (in alcohol and 
skins and skulls) Miri, Baram River, Sarawak, 8rf 199 BM(NH) 51.88-1 14 (in alcohol and 
skins and skulls) Long Lama Caves, Baram River, Sarawak, 29 BM(NH) (in 
alcohol, skulls extracted) Mt. Pisang, rf BM(NH), 6<f BM(NH), 2cf 
BM(NH) 58.26-7, 2 9 BM(NH) 59.155-6, 9 BM(NH) 69.232, rf 9 BM(NH) 75.1899-1900 (in 
alcohol and skins and skulls) Niah Cave, Sarawak, 9 BM(NH) (skin and skull) Mt. 
Dulit, BM(NH), d 1 9 BM(NH) 78.119-120 (in alcohol, skulls extracted) 
Gunong Mulu, 2rf BM(NH) 57.471-2 (in alcohol) Gomanton Caves, 9 BM(NH) 64.38 (in 
alcohol) Kuching, Sarawak, cf 9 BM(NH) 78.121-2 (in alcohol) Sungei Buloh, BM(NH) 
unregistered (in alcohol) Bidi Caves, Sarawak. MALAYA: BM(NH) (skin and 
skull) Singapore, HOrf 259 BM(NH) 60.1706-1840 (in alcohol, skulls extracted) Bukit 
Lagong Forest Reserve, Kepong, d 1 BM(NH) 61.1717 (in alcohol) Pulau Kaban, E. coast of 
Johore, rf BM(NH) 62.714 (in alcohol, skull extracted) Ulu Gombok, Selangor, d 9 BM(NH) 
67.1602-3 (in alcohol and skin and skull) Gunong Benom, Pahang. RIOUW 
ARCHIPELAGO: 2<5 1 9 BM(NH) (skins and skulls) Karimon I. SUMATRA: cf 
BM(NH) (in alcohol, skull extracted) Upper Langkat, Holotype of H. schneideri 9 
BM(NH) (in alcohol) Soekaranda, 9 BM(NH) (in alcohol) Lebong Tandai, 
Benkalen Reserve, 9 BM(NH) 94. 1 .7.5 (in alcohol, skull extracted) Engano Island, 9 BM(NH) 
95.1 .9.5 (in alcohol) Simatobe, Sipora. 

DIAGNOSIS. Larger on average than H. c. cervinus. Specimens from Malaya and Borneo have 
a blunt post-palatal spicule which is lacking in those from Sumatra. 

Measurements of holotype: M 3 -M 3 7'0; C-M 3 6*7; M 3 1-75. 

Measurements of the series examined: FA (32) 43-7-51-8 (47-81); III 2 (32) 14-4-18-3 
(16-58); V 2 (31) 7-8-10-5 (9-43); TL (32) 18-5-26-8 (22-7); CCL (10) 15-2-16-2 (15-78); ZW 
(14) 9-3-10-4 (9-91); MW (1 1) 9-0-9-5 (9-21); TD (12) 2-40-2-65 (2-50); M 3 -M 3 (15) 6-5-7-2 
(6-83); C-M 3 (16) 6- 1-6-7 (6-45); M 3 (16) 1-60-1-80 (1-75). 

REMARKS. Much of the confusion between H. galeritus and H. cervinus originates from 
Dobson (1876) who erroneously synonymized labuanensis with galeritus and who suggested 
(1878) that labuanensis was an intermediate form between galeritus at one extreme and 
longicauda at the other. There is every indication in the accounts by Lyon (1911), Chasen 
(1931, 1940) and Davis (1962) that the specimens or galeritus' used in their discussions and 
comparisons of this species with insolens were actually representatives of H. cervinus 
labuanensis. Consequently, the specimens recorded by Sanborn (1952) from the Philippine 
Islands, should also be referred to H. c. labuanensis or represent a closely related taxon, since 
Sanborn stated that his specimens agreed with (unspecified) descriptions of galeritus and 
with Lyon's (1911) measurements. 

Tate (1941) generally regarded schneideri as a distinct species (pp. 367, 369, 391) although 
he mapped it (p. 365) as a subspecies of//, galeritus, while Chasen (1940), Laurie & Hill 


(1954) and Hill (1963) considered it a subspecies of//, galeritus. The Sumatran specimens 
examined are very similar to H. cervinus labuanensis, to which they are referred. Thomas 
(1904) separated schneideri from 'galeritus'' because of small differences in the premolar 
dentition but these appear to be the result of individual variation and are within the range of 
H. cervinus. 

Medway (1958) provided a useful account of roosting and flight behaviour in Borneo 
which probably applies to H. c. labuanensis, although it is possible that his remarks refer in 
part to H. galeritus insolens since the two sometimes occur in the same caves. However, 
insolens is much less common in collections and may be solitary or live in small groups (c.f. 
H. g. brachyotus) and if so is unlikely to form part of the swarms described by Medway. 

DISTRIBUTION. Malay Peninsula and Sumatra to Borneo and possibly the Philippine Islands. 

Hipposideros cervinus batchianus Matschie, 1901 

Hipposideros batchianus Matschie, 1901 : 273 (Batjan I.) 

[Hipposideros] batchianus, Tate, 1941 : 359 (listed), 381 (holotype). 

Hipposideros galeritus batchianus, Tate, 1941 : 367, 369 (systematic status), 392 (Batjan); Hill, 

1963 : 56 (Batchian I. [ = Batjan I.]). 
Hipposideros cervinus batchianus, Tate, 1941 : 365 (mapped), Laurie & Hill, 1954 : 57 (Batchian I. 

[ = Batjan I.]). 

HOLOTYPE. In the Museum fur Naturkunde der Humboldt-Universitat zu Berlin. Adult 9, 
number 1 5 628, skin and skull. Batjan Island. Not seen. 


DIAGNOSIS. Measurements given by Matschie suggest that H. c. batchianus is similar in size 
to H. c. cervinus. 

Measurements of the holotype taken from Matschie (1901): FA 47'5; IIP 12; V 2 8*5; TL25; 
ZW 10; Temporal breadth from the auditory aperture 8'2; C-M 3 6'4. 


Hipposideros cervinus misorensis (Peters, 1906) 

Phyllorhina cervina var. misorensis Peters, 1906: pi. 5L, fig. 4 (probably Misor I. [ = Schouten I.], New 

HOLOTYPE. Based solely on the plate by Peters, its original specimen probably in the 
Museum fur Naturkunde der Humboldt-Universitat zu Berlin, but not traceable February 


REMARKS. The status of this taxon cannot be evaluated from the plate upon which it is based, 
beyond remarking that the noseleaf closely resembles that of H. cervinus, with which 
misorensis is provisionally associated. 

Summary of taxonomic changes 

The following taxa have been discussed and are listed below with their synonyms in brackets: 
Hipposideros galeritus galeritus, H. g. brachyotus, H. g. insolens, H. g. longicaudus, H. 
cervinus cervinus (celebensis), H. c. labuanensis (schneideri), H. c. batchianus, H. c. 


We are grateful to the Earl of Cranbrook whose dissatisfaction with the currently accepted 
classification of Bornean ''galeritus'' prompted the initial investigations. Relevant material 


has recently been received from Mr B. H. Gaskell of 'Operation Drake' who provided 
specimens from Papua New Guinea and Sulawesi. We are indebted also to Professor J. D. 
Smith, Department of Biology, California State University, who generously provided 
collectors, notes and measurements of specimens from the Bismarck Archipelago. We are 
grateful to Dr C. Smeenk, Rijksmuseum vanNatuurlijke Historic, Netherlands for the loan of 
material relevant to this study and to Dr F. Dieterlen, Staatliches Museum fur Naturkunde 
Stuttgart, Dr H. Hackethal, Museum fur Naturkunde der Humboldt-Universitat zu Berlin 
and Dr D. Kock, Forschungsinstitut und Natur-Museum Senckenberg, Frankfurt for 
providing information about specimens in their collections. 


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Manuscript accepted for publication 27 January 1 98 1 

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