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BULLETIN OF f*%u**a

THE BRITISH MUSEUM
(NATURAL HISTORY)

ZOOLOGY

VOL. 13

1965-1966

TRUSTEES OF

THE BRITISH MUSEUM (NATURAL HISTORY)

LONDON: 1967

DATES OF PUBLICATION OF THE PARTS

No. i ...... 6 August 1965

No. 2 . ." . . 14 September 1965

No. 3 . . . . . 14 September 1965

No. 4 . . . . . -15 October 1965

No. 5 ...... i October 1965

No. 6 ...... i October 1965

No. 7 . . . . . .22 October 1965

No. 8 . . . . . .3 December 1965

No. 9 ...... 2 March 1966

PRINTED IN GREAT BRITAIN

BY ADLARD & SON LIMITED

BARTHOLOMEW PRESS, DORKING

CONTENTS

ZOOLOGY VOLUME 13

PAGE

No. I. Primitive Cryptostigmatid mites from Rhododendron forests in the

Nepal Himalaya. By J. G. SHEALS i

No. 2. Japanese and other Ophiuroids from the collections of the Miinich

Museum. By AILSA M. CLARK (PI. i) 37

No. 3. The freshwater Gastropod molluscs of West Cameroon. By C. A.

WRIGHT (Pis. 1-3) 73

No. 4. Barbus (Pisces, Cyprinidae) of the Volta region. By A. J. & J. HOPSON 99

No. 5. Notes on the Cupuladriidae (Polyzoa, Anasca). By PATRICIA L.

COOK (Pis. 1-3) 151

No. 6. Polyzoa from West Africa : the Cupuladriidae (Cheilostomata,

Anasca). By PATRICIA L. COOK (Pis. 1-3) 189

No. 7. Descriptions of some Strongyles (Nematoda) from mammals in East
Nepal : with records of other parasitic nematodes. By WILLIAM G.
INGLIS & COLIN G. OGDEN 229

No. 8. Studies on the British Dermanyssidae (Acari : Mesostigmata), part i,

external morphology. By G. OWEN EVANS & W. M. TILL 247

No. 9. A review of the Muridae (order Rodentia) of Uganda. By M. J.

DELANY & B. R. NEAL 295

Index to Volume 13 357

PRIMITIVE CRYPTOSTIGMATID

MITES FROM RHODODENDRON

FORESTS IN THE NEPAL HIMALAYA

J. G. SHEALS

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. i

LONDON: 1965

PRIMITIVE CRYPTO STIGMATID

MITES FROM RHODODENDRON

FORESTS IN THE NEPAL HIMALAYA

J. G. SHEALS

British Museum (Natural History)

Pp. 1-35 ; 62 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. i

LONDON: 1965

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 13, No. i of the Zoological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

Trustees of the British Museum (Natural History) 1965

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued August, 1965 Price Eighteen Shillings

PRIMITIVE CRYPTO STIGMATID

MITES FROM RHODODENDRON

FORESTS IN THE NEPAL HIMALAYA

By J. G. SHEALS

INTRODUCTION

THIS report forms the first of a series on the soil- and litter-dwelling Acari collected
by the British Museum (Natural History) Expedition to East Nepal 1961-62. A full
account of this expedition with details of the localities examined is given by Sheals
& Inglis (1965) . In spite of the outstanding biogeographical interest of the Himalayan
region as a complex transitional zone, very little attention has been paid to the soil
and litter fauna of the area. In so far as the Acari are concerned, small collections
were made in 1954 by the British Museum (Natural History) Expedition to West
Nepal. Some of this material has been utilized in revisionary studies (Hyatt 1956,
Evans & Hyatt 1958, 1960), but apart from these no account of the Himalayan
free-living mites appears to have been published since the report on a collection of
Oribatidae from the Sikkim Himalaya by Pearce (1906).

The present paper is concerned with the taxonomy of the Oribatei Inferiores from
Rhododendron forest soil and litter in four localities in East Nepal : a large forest of
tree Rhododendrons (mainly R. arboreum) on the slopes of the Milke Danra ridge,
27° 19' N 87° 31' E, altitude 9,300-11,000'; an area of dwarf Rhododendron at
Topke Gola, 27° 38' N 87° 35' E, altitude 12,500-13,000'; an area of tree and bush
Rhododendron (mainly R. arboreum) at Selap, 27° 38' N 87° 49' E, altitude 10,100-
10,300' and thickets of bush Rhododendron (mainly R. arboreum) on the crest of the
Singalila ridge near Sandakphu, 27° 06' N 88° 01' E, altitude 12,000'. The higher
Cryptostigmata and other mites from these localities will be dealt with in later papers.

In the descriptions the setal nomenclature used in the later works of Grand jean
has been followed except in the case of the Nothroidea where it is convenient to
follow the system used by Sellnick & Forsslund (1955). The type material is
deposited in the British Museum (Natural History) .

Family ACARONYCHIDAE Grandjean, 1932
Himalacarus chimalae1 gen. et sp. nov.

ADULT. The prodorsum (figs, i & 2) is approximately i6ofjt, long and 140(1, wide
at the broadest point level with the sensilli. There is a prominent obliquely pro-
jecting rostral protuberance (the naso of Grandjean, 1958) below which a bilobed
eye such as Grandjean (op. cit.) has noted for other members of this family can
easily be discerned. The rostral setae (ro) are stout, prominent, quite smooth and
about 8o(jt in length while the lamellar setae (la) are barbed and about 95[z in length.
The interlamellar setae (in) and the exobothridial setae (xi) are barbed and about
I30|x in length, while the exobothridial setae (xs) are short (approx. 30(x long), smooth

xFrom the Nepali chimal — certain tree Rhododendrons

J. G. SHEALS

ps 2

FIG. i. Himalacarus chimalae gen. et sp. nov. Adult lateral.

and rather blunt. The sensillus (bo), i85-i90[x long, it quite smooth and tapers
gradually to a fine point. A rectangular design evidently formed by a thickening of
the cuticle can be discerned between the interlamellar setae ; this is almost identical
in form to the system of 'chitinous nervures' (false lamellae) described by Grand] can
(1952) for Stomacarus tristani.

The hysterosoma is about i75(J. in length and approximately 170^ wide at the
broadest point level with setae e2. The 'asthenique zone' between the posterior border
of the propodosoma and the median dorsal sclerite is very short. There are 17 pairs
of setae in the gastronotal series. Setae GI, cp, d2, ei, 62, f2 and hi are long, barbed,
dark brown in colour and arise from tubercles, those of setae et being particularly
prominent. An internal thickening such as Grandjean (1952, I952a, 1957) has
observed for Stomacarus, Archeonothms and Andacarus, respectively, can be seen
below the tubercle of e\. Difficulty was experienced in securing specimens with
completely undamaged setae but it would appear that the long barbed setae clf c2,
cp, d2 and e2 are about equal in length (i.e. about 170^) while the barbed setae f2 and
hj may be slightly longer than the total length of the idiosoma. Setae Ci and the

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL

short smooth c2 lie together on a median dorsal sclerite, while setae d2 and the
short smooth di lie together on paired sclerites. The remaining gastronotal setae,
with the exception of e2, are borne individually on small sclerites. Setae fj and those
of the ps and h series are short, smooth and simple, with the exception of setae ps2
which are shaped like spear-heads. Setae pst and h2 are rather blunt.

hllL

FIGS. 2-6. Himalacarus chimalae gen. et sp. nov. Fig. 2. Adult, dorsum. Fig. 3.
Adult, ano-genital region. Fig. 4. Adult, infracapitulum, Fig. 5. Adult, palp.
Fig. 6. Tritonymph, venter.

6 J. G. SHEALS

The ano-genital region is shown in fig. 3. There are 3 setae in the anal series
(AN) and 4 in the adanal (AD). All these setae lie individually on small sclerites.
There are 9 pairs of genital setae and 3 pairs of aggenitals (AG). The 10 pairs of
eugenital setae are all approximately similar in size and shape and it is suspected
that all the adults collected were males. The coxisternal region is essentially similar
to that of the tritonymph (fig. 6), the setal formula being (4-3-4-4).

FIGS. 7-10. Himalacarus chimalae gen. et sp. nov. Adult, legs.
Fig. 7. Leg I. Fig. 8. Leg II. Fig. 9. Leg III. Fig. 10. Leg IV.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 7

The infracapitulum is shown in fig. 4. There are four oral lips and in comparison
with other members of the family the lower lip is particularly large and easily seen.
The upper lip is also prominent and bears on its under surface a series of minute
denticles forming an inverted U-shaped pattern. The three pairs of adoral setae
are simple, the anterior pair (or^ being shorter than the other two. The chelicerae
have the characteristic 'ornithocephaline' outline common to other members of the
family and carry two setae dorsally. Their fixed digits bear three prominent teeth
and the movable digits only two ill-defined shallow projections. The five-segmented
palp (fig. 5) has the setal formula (0-2-1-3-18). The solenidion of the palp-tarsus
is borne on a prominence immediately distal to the lyriform fissure and of the 18
setae on this segment, 9 are eupathidia.

The legs (figs. 7-10) are relatively stout and robust, legs I and IV being markedly
longer than II and III. All the femora are divided. The setal formulae are : I (o-[4~6]-
5-6-31); II (i-[5-5]-5-7-27) ; III (2-[2-3]-3-6-26) ; IV (3-[3-3]-4-5-23), and the
formulae for the solenidia are: — 1(2-4-4) > H (J — 2 — 3) '> III (i-i-o) and IV (1-2-0).
On tarsus I the famulus, which is slender and ciliated, lies on a small prominence
between the lyriform fissure and the proximal solenidion. Of the 31 setae observed
on this segment at least 9 can be distinguished as eupathidia. As in Acaronychus,
only the pleural setae on tarsus II are eupathidial.

On all four legs the ambulacrum has two well-developed lateral claws. A minute
blunt central element can also be distinguished but this is evidently very much
less developed than in other Acaronychid genera.

TRITONYMPH. The number and disposition of dorsal setae are essentially similar
to those of the adult. Ventrally (fig. 6) the anal and coxisternal regions are similar
to those of the adult although in the figured specimen only 3 adanal setae were
present on one side. There are 7 genital and 2 aggenital setae.

The setal formulae for the legs are I (o-(4~6)-5-6-27), II (i-(3-5)-5~7-2i),
III (2-(i-3)-3-6-2i) ; IV (3-(i-3)~4-5-i7). The formulae for the solenidia are
the same as those for the adult.

OTHER STAGES. Unknown.

SYSTEMATIC POSITION

The Palaeacaroidea, now comprising n genera, were first recognised as a group
by Tragardh (1932) who considered they should constitute a separate sub-order of
the Acari — the Palaeacariformes. A few months later, however, Grandjean (1932)
failed to confirm Tragardh's observation of the presence of mandibulary stigmata
and concluded that the group should be incorporated with the Oribatei. In 1954,
Grandjean considered the group should be regarded provisionally as a super-family
— the Palaeacaroidea— comprising three distinct families, the Acaronychidae
(=Archeonothridae), Palaeacaridae and Ctenacaridae.

Himalacarus shows obvious affinites with the four genera, Acaronychus, Stomacams,

8 J. G. SHEALS

Andacarus and Archeonothms which comprise the family Acaronychidae in so much
as it possesses all the family attributes listed by Grandjean (1954), namely:

1. A 'naso' and its associated inferior protuberance.

2. A gastronotal series of 34 setae, the long dorsal setae being darkly
pigmented.

3. No large asthenique zone.

4. A short hysterosoma without large mediodorsal and pygidial shields.

5. Normal genital papillae and unmodified anterior genital setae.

6. Four oral lips.

7. The anterior adoral setae being much shorter than the other two pairs.

8. A large rounded palp tarsus with 16-19 setae of which 9 are eupathidia.

9. A mandible with an 'ornithocephaline' outline.

10. A first tarsus with a large erect famulus implanted close to the lyriform
fissure.

Grandjean further recognised two sub-families, the Archeonothrinae for the
3 genera Archeonothrus , Stomacarus and Andacarus; and the Acaronychinae for the
single genus Acaronychus. He separated the Archeonothrinae on the basis of their
non-claviform famulus and normal female eugenital setae and on the presence of
2 aggenital setae, false lamellae and small internal apophyses at the bases of setae e2.
The Acaronychinae were characterised by the absence of internal apophyses and
false lamellae and by the presence of a dilated famulus, 3 aggenital setae and, in
the female, swollen claw-like eugenital setae. However, Grandjean later (1957)
observed that the female of Stomacarus macfarlani, the species which was subsequently
made the type of Andacarus (Grandjean igsSa), had claw-like eugenital setae so
that this feature had to be abandoned as a sub-familial character.

Although having 3 aggenital setae, Himalacarus shows greater overall similarity
to the Archeonothrine genera than to Acaronychus, thus the number of aggenital
setae must also be eliminated from the list of sub-familial features, and, with the
reduction of the number of diagnostic sub-familial attributes to two, it might be
preferable to consider the family as a single relatively homogenous group. In table i,
the condition of the genera in relation to the main characters which have been used
in their classification is summarized.

Since the main part of this account was prepared, Balogh and Csiszar (1963) have
published a description of Stomacarus longicaudatus from South Argentina. Judging
by the figure of the dorsum this species appears to be very close to H . chimalae and
is probably congeneric with it. It can be noted that in S. longicaudatus the dorsal
setae di and i\ are comparatively long and barbed whilst in H . chimalae these setae
are shorter and smooth.

MATERIAL. H. chimalae occurred only in the samples from the Milke Danra forest
taken on 2nd December 1961. In all about 40 specimens were collected of which
15 were adult, apparently all males, the remainder being tritonymphs. The greater

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL

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io J. G. SHEALS

part of the material was collected in the shallow litter layer but a few specimens
were found in the underlying mineral soil.

Holotype (1964.5.27.1) and four paratypes (1964.5.27.2-5).

Family PARHYPOCHTHONIIDAE Grandjean, 1932
Parhypochthonius aphidinus Berlese

Parhypochthonius aphidinus Berlese, A. 1904. Redia 2: 25.

Parhypochthonius sp. Grandjean, F. 1934. Bull. Mus. Hist. nat. Paris (2) 6: 423.

The true identity of Berlese's aphidinus appears to be doubtful although it seems
most probable that the Parhypochthonius species described in detail by Grandjean
(op. cit.) is conspecific with this. A single specimen agreeing well with Grandj can's
description was recorded from Selap.

Family HYPOCHTHONIIDAE Berlese, 1910
Hypochthonius rufulus Koch

Hypochthonius rufulus Koch, C. L. 1836. Deutsch. Crust. Myr. Arach. Fasc. 3 t.ig.

This common species has a wide holarctic distribution having been recorded from
Europe and North America (van der Hammen, 1959) and from Japan (Aoki, 1959).
H. rufulus occurred in small numbers in the Topke Gola forest but was not recorded
from the other localities.

Family ENIOCHTHONIIDAE Grandjean, 1947
Eniochthonius minutissimus (Berlese)

Hypochthonius minutissimus Berlese, A. 1904. Redia 1: 252.
Hypochthonius pallidulus, Michael, A. D. 1888. British Oribatidae 2: 536.

The type locality is Florence and Michael found this species at the roots of ground
mosses in Epping Forest. E. minutissimus appears to be widely distributed in
Europe and in Nepal was found in all localities except Sandakphu. It was most
abundant in the Selap forest.

Family COSMOCHTHONIIDAE Grandjean, 1947
Cosmochthonius lanatus (Michael)

Hypochthonius lanatus Michael, A. D. 1885. /. roy. micr. Soc. (2) 5: 396.

The Nepalese material has been compared with specimens in the Michael collection.
Tarsi I are bidactylous while tarsi II-IV are tridactylous and as van der Hammen
(1952) has pointed out, Michael was in error when he figured the legs as being
monodactylous. Michael collected most of his material from an old thatched roof
in Cornwall and considered the species to be uncommon. However, it appears to be
widely distributed in soil and litter in Europe and in Nepal was found in all the 4
localities sampled.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL u

Family PHTHIRACARIDAE Perty, 1841
Phthiracarus robertsi sp. nov.

ADULT. The aspis (figs, n & 12) ranges in length from about 320-480^ and is
approximately 1.4 times as long as its greatest width immediately behind the
sensilli. All the prodorsal setae are weak. The sensillus is distinctly elbowed near the
base; proximally the basal portion is slender while at the elbow the sensillus thickens
abruptly before tapering gradually to a fine point. There are distinct lateral ridges
as well as a very pronounced broad median crest while behind the bothridium the
integument is raised into a series of longitudinal corrugations.

FIG. ii. Phthiracarus robertsi sp. nov. Lateral.

Along the line c3-h! the notogaster ranges in length from approximately 475-830(0,.
It is highly arched and, excluding the vestiges of fj and f2, carries 15 pairs of very
weak short setae. The fissures ia, ip and ips are well marked as also is the muscle
insertion point anterior to seta 62- The notogastral integument is distinctly pitted.
The ano-genital region is shown in fig. 13. There are five setae on each anal valve
and of these, two, evidently belonging to the anal series, are quite distinctly located
nearer the median border than the other three. All the anal and adanal setae are
very fine but equally well developed — cf. van der Hammen (1963 a) for P. laevigatus
(Koch). There are nine genital setae and a single aggenital seta. All the genital
setae are marginal and the anterior five setae are inserted more closely together than
the posterior four.

J. G. SHEALS

FIGS. 12-15. Phthiracarus robertsi sp. nov. Fig. 12. Aspis
Fig. 13. Ano-genital region. Fig. 14. Infracapitulum. Fig. 15. Chela.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL

FIGS. 16-19. Phthiracarus robertsi sp. nov. Legs, excluding trochanters.
Fig. 16. Leg I. Fig. 17. Leg II. Fig. 18. Leg III. Fig. 19. Leg IV.

I4 J. G. SHEALS

The infracapitulum is shown in fig. 14. The mentum is sub-trapezoidal in outline
and carries a single pair of long smooth setae while each gena carries two pairs of
similar setae. The three pairs of adoral setae are smooth and slightly sinuous. The
three-segmented palp has the formula (2-2-7) and °f the tarsal setae, three are
eupathidia. The chelicerae are shown in fig. 15. The fixed digit has three teeth and
the movable digit carries only a single tooth. Seta chb is located well down on the
anti-axial face, while seta cha is dorsal.

The legs are shown in figs. 16-19. The solenidia of the tarsi are all free, while on
each tibia the solenidion is coupled with a minute dorsal seta as also is the proximal
solenidion on genu I. The solenidion on tibia i extends beyond the tip of the tarsus
and the solenidion on genu i is about the same length as leg I. All the legs are mono-
dactyl and the formulae for the setae and solenidia are as follows: —
I (1-4-2-5-15) and (2-1-3) ; II (1-3-2-3-12) and (1—1—2) ;
III (2-2-1-2-10) and (i-i-o); IV (2-1-1-2-10) and (o-i-o).

This species is of some interest as the arrangement of setae on the anal valves
approaches the condition commonly associated with species of Steganacarus. This
species is dedicated to Lt. Colonel J. O. M. Roberts M.C., formerly Military Attache,
British Embassy, Kathmandu, in appreciation of his assistance to the East Nepal
Expedition.

FIG. 20. Phthiracarus cf. laevigatus Koch. Sensillus of Nepalese specimen.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 15

MATERIAL. Holotype (1964.6.19.1) and five paratypes (1964.6.19.2-6) Milke
Danra, 24.xi.6i. This species was not abundant and was recorded only from the
Milke Danra forest.

Phthiracarus cf. laevigatus (Koch)

Hoplophora laevigata Koch, C. L. 1841. Deutsch. Crust. Myr. Arach. Fasc. 38 t.i6.
Phthiracarus laevigatus, van der Hammen, L. 1963. Acarologia 5: 704.

Six specimens evidently referable to this species were taken at Sandakphu. The
material agrees reasonable well with van der Hammen' s redescription but the form
of the sensillus in the Nepalese specimens (fig. 20) appears to differ slightly. In the
Nepalese material the first adoral seta is feathered, a feature not mentioned by
van der Hammen.

Hoplophthiracarus nepalensis sp. nov.

ADULT. The aspis (figs. 21 & 22) ranges in length from about 215-290^ and is
approximately 1-2 times as long as its greatest width immediately behind the
sensilli. The rostral (ro), lamellar (la) and exobothridial (ex) setae are short and
simple, while the interlamellar setae (in) are stout, feathered apically, nearly erect,
and about three times as long as the rostrals. The sensilli are elbowed and taper
gradually to a fine point with slight apical feathering. The prodorsal integument is
coarsely pitted and behind the sensilli is raised into a series of low longitudinal ridges.

FIG. 21. Hoplophthiracarus nepalensis. sp. nov. Lateral.

J. G. SHEALS

Along the line 03-ps^ the notogaster ranges in length from about 330-500^, and,
excluding the vestigial fj and f2 there are 30 setae in the notogastral series. All the
notogastral setae are strong, nearly erect, weakly feathered apically and about equal
in length. Only the fissures ia and im could be discerned. The notogastral integument
is coarsely pitted, the margins of the pits being irregular. Ventrally (fig. 23) the anal
valves have prominent antero-median overlapping lobes, the right overlying the left.
The two anal setae are marginal and comparatively long while the three adanal
setae form an oblique row, the longest in the series being ad2, which is approximately
2-5 times as long as the anal setae. There are nine pairs of genital setae and a single
pair of aggenitals. The posterior four pairs of genital setae are comparatively long
and situated in a line somewhat remote from the margin, whilst the anterior five

FIGS. 22-25. Hoplophthiracarus nepalensis sp. nov. Fig. 22. Aspis.
Fig. 23. Ano-genital region. Fig. 24. Infracapitulum. Fig. 25. Chela.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL

FIGS. 26-29. Hoplophthiracarus nepalensis sp. nov. Legs, excluding trochanters.
Fig. 26. Leg I. Fig. 27. Leg II. Fig. 28. Leg III. Fig. 29. Leg IV.

i8 J. G. SHEALS

pairs are minute and marginal. There are three pairs of genital papillae, the anterior
papillae being rather small (cf. van der Hammen 1963, p. 314 for H . pavidus Berlese
which has only two pairs of genital papillae) . The ornamentation of the integument
of the genital and anal valves is similar to that of the notogaster.

The infracapitulum is shown in fig. 24. The mentum is subtrapezoidal in outline
and carries a pair of long simple setae (h) . The genae carry two pairs of setae (a & m) ;
setae m being about 1.5 — 2 times as long as setae a. There are three pairs of simple
adoral setae. The three-segmented palp has the setal formula (2-2-7) an(l a* least
three of the tarsal setae are eupathidial. The minute dorso-anterior seta appears to
be discrete and may be eupathidial. The chelicerae are shown in fig. 25.

The legs are shown in figs. 26-29. ^ is interesting to note that the solenidion on
genu I and the solenidia on tibiae I-III are coupled with minute dorsal setae, while
the solenidion on tibia IV is free. This condition also appears to obtain in H. pavidus
(Berlese), (van der Hammen 1963). All the legs are monodactyl and the formulae
for the setae and solenidia are as follows :

I (1-4-2-5-15) and (2-1-3) ; II (1-3-2-3-12) and (1-1-2) ;
III (2-2-1-2-10) and (i-i-o); IV (2-1-1-2-10) and (o-i-o).

H. nepalensis appears to be very similar to H. robustior Jacot which was taken from
second growth pine at Pensacola, Florida, U.S.A. Jacot (1933) did not figure the
venter of H. robustior but noted that the anal and genital plates were 'scrolled' and
not 'pock marked'. In H. nepalensis both the anal and genital valves are quite
clearly pitted in the same way as the notogaster.

MATERIAL. Holotype (1964.6.24.1) and five paratypes (1964.6.24.2-6) Sandakphu,
I2.xi.6i. This species was common at Sandakphu but was not found in the other
localities.

Steganacarus striculus (Koch)

Hoplophora stricula Koch, C. L. 1836. Deutsch. Crust. Myr. Arach. Fasc. 2 t.io.

Steganacarus striculus, Feider, Z. & Suciu, I. 1957. Stud. Cere, stiint. Biol. Agric. Acad. R.P.R.

8:33.

The Nepalese material conforms closely to the redescription of this species by
Feider and Suciu (op. cit.) and by Aoki (1958). In comparison with British material
however the Nepalese specimens have rather stouter notogastral setae and the apical
feathering of these setae is well marked i.e. as figured by Fieder & Suciu. 5. striculus
has a wide holarctic distribution and in Nepal was particularly abundant in the
Milke Danra and Topke Gola forests. It was prominent in Selap but only a few
specimens were taken at Sandakphu.

Family EUPHTHIRACARIDAE Jacot, 1930
Euphthiracarus inglisi sp. nov.

ADULT. The aspis (figs. 30 & 31) ranges in length from approximately 260-330^
and is about 1.3 times as long as its greatest width at the level of the sensilli. The
rostral setae (ro) are long and smooth and about 1.5 times the length of the lamellar

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 19

setae (la). Both the lamellar and the interlamellar setae (in) are slightly feathered
apically and the latter are about 1.6 times the length of the former. There is a very
pronounced scale below the bothridium, and the sensilli, which are elbowed, taper to a
comparatively blunt point with very slight apical feathering. The prodorsal integu-
ment is coarsely pitted.

FIG. 30. Euphthiracarus inglisi sp. nov. Lateral.

At the level of seta h2 the notogaster (fig. 30) ranges in length from 475 to 620^.
The 14 pairs of setae in the notogastral series are all equal in length, erect and slightly
feathered apically. The ornamentation of the notogastral integument, consisting of
very weak pits, is barely discernible. Ventrally (fig. 32), the anal portion of the
ano-genital valve carries six pairs of setae and a single pair of pores. Posteriorly the
ano-genital plate is very much produced dorsally so that the position of the posterior
setae is difficult to discern in the whole animal, but on dissection (fig. 34) the most
posterior pair are seen to be inserted near the notogastral border. Of the six pairs
of setae on the anal valves, two are marginal, quite smooth and distinctly longer
than the other four. The latter are removed from the margin and slightly feathered
apically as in the case of the notogastral setae. There are nine pairs of genital and
two pairs of aggenital setae. The anterior pair of aggenital setal are short and narrowly
lanceolate. The integument of the ano-genital region is distinctly pitted.

The infracapitulum is shown in fig. 33. The sub-triangular mentum carries a
single pair of long setae (h). The genae carry two pairs of simple setae (a & m), seta
m being rather longer than a, and there are three pairs of simple adoral setae. The
three-segmented palp has the setal formula (2-2-8) and at least four of the tarsal

J. G. SHEALS

FIGS. 31-35. Euphthiracarus inglisi sp. nov. Fig. 31. Aspis. Fig. 32. Ano-genital
region. Fig. 33. Infracapitulum. Fig. 34. Anal valve — dissected. Fig. 35. Chela.

setae are eupathidial. The minute dorso-anterior seta is discrete and has the appear-
ance of being eupathidial. The chelicerae are shown in fig. 35. Seta cha is dorsal
and seta chb is inserted well down on the antiaxial face. Each digit has four teeth.
The legs are shown in figs. 36-39. A noticeable feature is that on all the tibiae the
dorsal seta is extremely long and on legs II-IV exceeds the tibial solenidia in length.
All the tarsi are tri-heterodactyl and the formulae for the setae and solenidia are as
follows :—

I (1-3-4-5-16) and (2-1-3) ; II (1-3-3-5-14) and (1-1-2) ;

III (2-2-2-2-11) and (i-i-o); IV (2-1-1-2-10) and (i-i-o).

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL

FIGS. 36-39. Euphthiracarus inglisi sp. nov. Legs, excluding trochanters.
Fig. 36. Leg I. Fig. 37. Leg II. Fig. 38. Leg III. Fig. 39. Leg IV.

22 J. G. SHEALS

E. inglisi is very similar to E. flavus (Ewing). However, an examination of a
specimen of the latter collected in Illinois, revealed that, in contrast to E. inglisi,
the notogastral integument is very heavily ornamented and the rostral setae are
very much shorter than those of E. inglisi. Moreover, in E. flavus the distal third of
the sensillus is distinctly feathered whilst in E. inglisi the feathering is slight and
confined to the tip. This species is dedicated to my colleague on the East Nepal
Expedition, Dr. W. G. Inglis.

MATERIAL. Holotype (1964.7.2.1) and five paratypes (1964.7.2.2-6), Sandakphu,
I2.xi.6i. E. inglisi was common at Sandakphu but was not found in the other
localities.

Rhysotritia ardua (Koch)

Hoplophora ardua Koch, C. L. 1841. Deutsch. Crust. Myr. Arach. Fasc. 32 t.i5.

Pseudotritia loricata, Feider, Z. & Suciu, I. 1957. Stud. Cere, stiint. Biol. Agric. Acad. R.P.R.

8: 40.
Rhysotritia ardua, Markel, K. & Meyer, I. 1959. Zool. Am. 163: 341.

This species was extremely common in the Milke Danra and Selap forests. It
occurred in small numbers at Topke Gola but was not found at Sandakphu. The
Nepalese material has been compared with topotypic material from the Regensberg
area kindly provided on loan by Dr. L. van der Hammen. The Nepalese specimens
are generally larger and more heavily sclerotized than the topotypic material. Thus,

FIG. 40. Mesotritia maerkeli sp. nov. Lateral.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL

the length of the aspis in the Nepal material ranged from about 285-310(0. whilst in
the six topotypes examined the length of the aspis ranged from about 215-250^.
No other morphological differences could be detected.

cha

FIGS. 41-44- Mesotritia maerkeli sp. nov. Fig. 41. Aspis.
Fig. 42. Ano-genital region. Fig. 43. Infracapitulum. Fig. 44. Chela.

24 J. G. SHEALS

Mesotritia maerkeli sp. nov.

ADULT. The aspis (figs. 40 & 41) ranges in length from approximately 300-360^
and is about 1.2 times as long as its greatest width at the level of the sensilli. The
rostral setae which are inserted well behind the lamellar setae are long and
smooth and about 1.6 times the length of the latter. Both the lamellar and inter-
lamellar setae (in) are smooth. There is a very pronounced scale below the bothridium
while the sensilli are short — about the same length as the interlamellar setae — and
expanded sub-apically before tapering abruptly to a fine point.

At the level of seta h1; the notogaster ranges in length from approximately
550-600^. The 14 pairs of setae in the notogastral series are all approximately
equal in length, slender and rather weak. The notogastral integument has no
distinct ornamentation. Ventrally (fig. 42) the anal valve carries a single pair of
setae and there are three pairs of setae in the adanal series. The adanal pore (iad)
is located well in front of the anal seta. The genital valve is completely separated

FIGS. 45-46. Mesotritia maerkeli sp. nov. Legs.
Fig. 45. Leg I. Fig. 46. Leg II.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 25

from the aggenital plate by a distinct suture while the oblique fissure separating the
aggenital from the adanal elements characteristic of Oribotritia is lacking. There are
six pairs of genital and two pairs of aggenital setae.

The infracapitulum is shown in fig. 43. The sub-triangular mentum carries a
single pair of long simple setae (h). The genae carry two pairs of simple setae
(a & m) seta a being about 1.7 times the length of seta m. Only two pairs of adoral
setae could be discerned. The three-segmented palp has the setal formula (2-2-8)
and four of the tarsal setae are eupathidial. The chelicerae are shown in fig. 44.
Seta chb is situated well down on the anti-axial face and both the fixed and movable
digits have two teeth.

The legs are shown in figs. 45-48. On leg I the famulus is forked and on all legs
the solendia of the tibiae and genua are coupled with dorsal setae, the dorsal setae
on tibia I being particularly long. All the tarsi are triheterodactyl and the formulae
for the setae and solenidia are as follows :—

I (1-3-5-5-17) and (2-1-3) I II (1-4-4-4-12) and (1-1-2) ;

\ *_/ *_/«_/ / / \ »j/ * \ i i i / \ /"

III (2-3-2-2-11) and (i-i-o); IV (2-3-2-2-10) and (o-i-o).

FIGS. 47-48. Mesotritia maerkeli sp. nov. Legs.
Fig. 47. Leg III. Fig. 48. Leg IV.

26 J. G. SHEALS

I am indebted to Professor Konrad Markel, to whom this species is dedicated,
for information on the genus Mesostritia. Professor Markel is currently preparing a
revision of the Euphthiracaridae and for the present it is sufficient to note that while
M . maerkeli has affinities with M. testacea Forsslund (Forsslund & Markel 1963) it
can be readily separated from the latter by reference to the position of the rostral
setae and adanal pore.

MATERIAL. Holotype (1964.7.20.1) and one paratype (1964.7.20.2), Selap, 4.ii.62.
Only four specimens were taken and all were found in the Selap Forest.

Oribotritia berlesei (Michael)

Phthiracarus berlesei Michael, A. D. 1898. Das Tierreich Lief 3: 81.

Oribotritia decumana, Willmann, C. 1931. Die Tierwelt Deutschlands Teil 22: 193.

Oribotritia berlesei, Grandjean, F. 1933. Bull. Mus. Hist. nat. Paris 5: 309.

Eight specimens were taken from the Topke Gola forest, four from Selap and one
from the Milke Danra forest.

Family EPILOHMANNIIDAE Oudemans, 1923
Epilohmannia cylindrica (Berlese)

Lohmannia cylindrica Berlese, A. 1904. Redia 2: 23.

Lesseria szanisloi Oudemans, A. C. 1917. Arch. Naturgesch. 88A. (6): 79.

A single specimen agreeing well with Oudemans' description was recorded from
Selap.

Family EULOHMANNIIDAE Grandjean, 1931
Eulohmannia ribagai Berlese

Lohmannia (Eulohmannia) ribagai Berlese, A. 1910. Redia 6: 223.

Arthronothrus biunguiculatus Tragardh I. 1910. Naturwissenschaftliche Unlersuchungen des
Sarekgebirges in Schwedisch-Lappland 4: 544.

Generally regarded as a relict, this species has a wide holarctic distribution
(Balogh 1961). In Nepal a small number of specimens was taken at Topke Gola
and a single specimen was recorded from Selap.

Family NOTHRIDAE Berlese, 1896
Nothrus springs my t hi sp. nov.

ADULT. The prodorsum (fig. 49) is approximately 35O[j. long and 350^ wide at the
broadest point level with the interlamellar setae (in). The median rostral incision
extends almost to the base of the rostral setae (ro) and the latter are short and
simple while the lamellar (la) and interlamellar setae are narrowly spatulate and
serrated apically. The exobothridial setae (ex) are simple. The sensilli are approxi-
mately 2iO[A in length and slightly expanded in one plane subapically. A cluster of
pseudotracheae can be discerned below the bothridia. The integument of the pro-
dorsum is pitted and, centrally, the raised margins of the pits form a regular polygonal
network.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL

FIGS. 49-52. Nothrus springsmythi sp. nov. Adult. Fig. 49. Dorsum.
Fig. 50. Venter. Fig. 51. Chela. Fig. 52. Infracapitulum.

J. G. SHEALS

The notogaster (fig. 49) is approximately 720^ in length and about 620^ wide at
the broadest point near the level of setae e2. There are 16 pairs of notogastral setae,
those in the c, d, e and f series being narrowly spatulate. Seta ct is about 2.5 times
the length of c2 and the insertion point of the latter is slightly closer to GI than to c3.
The setae of the pn series are broadly spatulate and seta kj is about 1.9 times the
length of pn^ All the setae of the pn and k series have a very prominent core of
active chitin with ramifying branches near the tip. The integument of the central
part of the notogaster is irregularly pitted, the margins forming a sub-circular
pattern.

FIGS. 53-56. Nothrus springsmythi. Adult, legs excluding trochanters.
Fig. 53. Leg I. Fig. 54. Leg II. Fig. 55. Leg III. Fig. 56. Leg IV.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 29

Ventrally (fig. 50) there are two setae in the anal (an) and three setae in the
adanal (ad) series. The latter are very narrowly spatulate. Owing to the granular
nature of the integument of the epimera the setal bases are difficult to discern.
However there appears to be considerable variation in the chaetotaxy of the inter-
coxal region and the arrangement is not always bilaterally symmetrical. The
observed variation in epimera was as follows: I (6-7), II (3-4), III (5) and IV (5-7).
This condition is evidently neotrichous, and, as Grand] ean (1934) has noted for
Nothrus silvestris (Nicolet), the 'abnormal' condition first becomes apparent in the
protonymph (3-2-3-0), the formula for the larva (2-1-2) being normal (vide infra).

The infracapitulum is shown in fig. 52. The arrangement is 'stenarthric' (Grand-
jean, iQ57a) and the rutella are basically of the Camisia type with large overlapping
ventral lobes completely covering the adoral papillae. There are two pairs of long
simple adoral setae and the four pairs of setae on the genae are arranged in a straight
line. The five-segmented palp has the formula (0-1-1-3-9) and four of the tarsal
setae are eupathidia.

The chelicera is shown in anti-axial view in fig. 51. It resembles the chelicera of
Camisia figured by Grandjean (1947) although seta cha is longer and more posteriorly
situated. The organ of Tragardh is discernible only with difficulty.

The legs are shown in figs. 53-56. Legs I and IV are much longer than II and III
and the integument of trochanters II and IV as well as that of all the femora is
sculptured with a distinct polygonal pattern. All the tarsi are tri-heterodactyl and
the formulae for the setae and solenidia are as follows : —

I (1-9-5-6-27) and (1-2-3) ; II (1-9-5-5-24) and (i-i-i) ;
III (3-5-5-5-25) and (i-i-o); IV (2-5-5-5-22) and (i-i-o).

LARVA. The prodorsum (fig. 57) is approximately 165(0. long. The lamellar setae
(la) are spatulate while the remaining prodorsal setae are simple. The sensilli are
represented by short stumps. Along the mid-line the notogaster is approximately
26o[j, in length and the integument is tuberculate. With the exception of setae c2
and ki, all the dorsal setae are broadly spatulate. Setae c2 are simple but somewhat
blunt and less than one third of the length of setae q. Setae kx are narrowly spatulate
and about twice the length of setae f2. Ventrally (fig. 58) seta pn: is spear-shaped and
approximately the same length as k^ The structure of the ano-genital region is
normal for the genus and the intercoxal formula is (2-1-2). The setal and solenidial
formulae for the legs are as follows :

I (0-2-3-4-14) and (i-i-i) ; II (0-2-3-3-13) and (i-i-i) ;
III (0-2-2-3-12) and (i-i-o).

PROTONYMPH. The prodorsum (fig. 59) is approximately 200(j, in length and its
integument is ornamented with a polygonal pattern. The lamellar (la) and inter-
lamellar setae (in) are broadly spatulate, while the sensilli, approximately 260^ in
length, taper to a fine point after bending sharply near the base. Along the mid-line
the notogaster is approximately 380^ in length, its integument is tuberculate although
this feature is not so well marked as in the larva. With the exception of seta k^
all the dorsal setae are broadly spatulate. Seta c2 is about half as long as seta ct

J. G. SHEALS

FIGS. 57-58. Nothrus springsmythi sp. nov. Larva.
Fig. 57. Dorsum. Fig. 58. Venter.

and seta kj is about six times the length of seta pn^ Ventrally (fig. 60) setae pnj and
pn3 are broadly spatulate, while the two pseudanal setae (op) are sub-lanceolate.
There are no setae in the anal or adanal series and each crescent-shaped genital
shield carries a single seta. The intercoxal formula is (3-2-3-0) and the setal and
solenidial formulae for the legs are as follows :

I (0-3-5-5-15) and (1-1-2); II (0-3-5-5-13) and (i-i-i);
III (1-2-4-4-13) and (i-i-o); IV (0-0-0-0-7) and (i-i-o).

DEUTONYMPH. Along the mid line the prodorsum and notogaster are approximately
200 and 450(j, in length, respectively. The arrangement and form of the dorsal
setae is essentially similar to that of the protonymph. Setae lq are about i8o[j. long
and 4-5 times as long as setae pnt. Ventrally (fig. 61) the pseudanal setae (op) have

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 31

become broadly spatulate, while three pairs of simple adanal and four pairs of genital
setae have appeared. The setal and solenidial formulae for the legs are as follows:
1 (I-7-5-5-I9) and (1-2-2) ; II (1-7-5-5-17) and (i-i-i) ;
III (3-3-5-5-17) and (i-i-o); IV (1-2-4-4-13) and (i-i-o).

TRITON YMPH. Along the mid-line the prodorsum and notogaster are approximately
230 and 650^ in length, respectively. The arrangement of dorsal setae is essentially
the same as in the adult except that the spatulate setae are rather broader. Setae
ki are about 240^ long and about 3 times the length of setae pn^ Ventrally (fig. 62),
the anal region differs from that of the deutonymph by the addition of 2 pairs of
anal setae and a pair of anal pores. There are seven pairs of genital setae. The setal
and solenidial formulae for the legs are as follows :

I (1-8-5-6-23) and (1-2-3); H (1-7-5-5-17) and (i-i-i) ; III (3-3-5-5-17)
and (i-i-o) and IV (1-2-4-4-13) and (i-i-o).

Nothrus springsmythi appears to have affinities with Nothrus biciliatus C. L. Koch.
However, European specimens of the latter kindly provided by Dr. K.-H. Forsslund
had an overall length of 790-800 [z, whilst N. springsmythi is much larger ranging in
length from 1070-1 130^. Moreover, in N. springsmythi setae kt are much more
slender although the form of these setae, as well as that of setae pn1; seems to vary
considerably. An interesting difference can also be seen in the form of the setae on
the tarsi. In N. biciliatus the three proximal latero-dorsal setae of the tarsi are
markedly spatulate with a sickle-like curve, whilst in N. springsmythi these setae
are much more slender and of the same type as the proximal ventral setae.

N. springsmythi also resembles two recently described South American species
viz. Nothrus oblongus Hammer 1961, and Nothrus macedi Beck 1962. It differs from
the latter in lacking the tubercle at the base of seta pn2, by having a smooth sensillus,
comparatively smooth distal setae on the tarsi and also in the length, relative to the
basal distance of setae f^ In comparison with N. oblongus, N. springsmythi is larger
and stouter. In the former species the first adanal seta is described as being similar
to the dorsal hysterosomal setae (narrowly spatulate) whilst in N. springsmythi all
the adanal setae are very narrowly lanceolate. This species is dedicated to Major
T. Le M. Spring Smyth in appreciation of his services to the East Nepal Expedition.

MATERIAL. Holotype adult (1964.6.15.1) and five paratypes comprising an adult
and all four postembryonic developmental stages (1964.6.15.2-6), Selap, 4.11.62.
N. springsmythi was extremely common in the Milke Danra and Selap forests. It
was particularly abundant in the latter locality and in one sample constituted 24%
of the Cryptostigmatid population. It was less abundant at Topke Gola and was
not found at Sandakphu.

Nothrus palustris Koch

Nothrus palustris Koch, C. L. 1839. Deutsch. Crust. Myr. Arach. Fasc. 29 1.13.
Nothrus palustris, Sellnick, M. & Forsslund, K.-H. 1955. Ark. Zool. 8: 498.

This species dominated the community at Topke Gola. It occurred in small
numbers in the Milke Danra and Selap forests and a single tritonymph was recorded
from Sandakphu.

J. G. SHEALS

FIGS. 59-62. Nothrus springsmythi sp. nov. Fig. 59. Protonymph, dorsum. Fig. 60.
Protonymph, venter. Fig. 61. Deutonymph, venter. Fig. 62. Tritonymph, venter.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 33

Family CAMISIIDAE Oudemans, 1900
Camisia lapponica (Tragardh)

Nothrus lapponicus lapponicus Tragardh, I. 1910. N aturwissenschaftliche Untersuchungen des

Sarekgebirges in Schwedisch-Lappland 4: 526.
Camisia lapponica, Sellnick, M. & Forsslund, K.-H. 1955. Ark. Zool. 8: 489.

According to Sellnick & Forsslund (op. cit.) this species has a wide holarctic
distribution. It occurred in small numbers at Sandakphu.

Platynothrus peltifer (Koch)

Nothrus peltifer Koch, C. L. 1839. Deutsch. Crust. Myr. Arach. Fasc. 29 t.g.
Platynothrus peltifer, Sellnick, M. & Forsslund, K.-H. 1955. Ark. Zool. 8: 515.

This species, which has a wide holarctic distribution, was found in all four localities,
but was most prominent in the Selap and Milke Danra forests. P. peltifer was
amongst the species recorded from the Sikkim Himalaya by Pearce (1906).

Family TRHYPOCHTHONIIDAE Willmann, 1931
Trhypochthonius tectorum (Berlese)

Hypochthonius tectorum Berlese, A. 1896. Acar. Myr. et Scorp. Ital. rep. Fasc. 78 8.
Trhypochthonius tectorum, Willmann, C. 1931. Die Tierwelt Deutschlands Teil 22: 103.

The type material appears to have been collected from moss on roofs in Florence
(van der Hammen, 1959). This species is widely distributed in Europe and the
collections of the British Museum (Nat. Hist.) include three specimens from Pearce's
Sikkim-Himalaya collection. This species was particularly common at Topke Gola
but occurred in small numbers in all the localities sampled.

Family MALACONOTHRIDAE Berlese, 1916
Malaconothrus monodactylus (Michael)

Nothrus monodactylus Michael, A. D. 1888. British Oribatidae 2: 528.

This species was common in the Milke Danra and Topke Gola forests and occurred
in small numbers at Selap.

Family NANHERMANNIIDAE Sellnick, 1928
Nanhermannia nana auct.

Nanhermannia nana, Willmann, C. 1931. Die Tierwelt Deutschlands Teil 22: 96.
Nanhermannia nana, Strenzke, K. 1953. Zool. Anz. 150: 73.

Although rather larger, ranging in length from approximately 580-600^, the
Nepalese specimens agree well with Strenzke's description. Van der Hammen
(1959) has shown that Nicolet's nanus has been widely misinterpreted and is not
identical with nana sensu Willmann, but he considered that the latter should not
be renamed until detailed redescriptions of Banks' species had been made. This
species was plentiful in the Milke Danra and Topke Gola forests.

34 J. G. SHEALS

SUMMARY

Twenty two species of Oribatei Inferiores were found in the soil and litter of
Rhododendron forests in four localities in East Nepal. The following six species
are considered to be new.

1. Himalacarus chimalae gen. et sp. nov. (p. 3)

2. Phthiracarus robertsi sp. nov. (p. n)

3. Hoplophthiracarus nepalensis sp. nov. (p. 15)

4. Euphthiracarus inglisi sp. nov. (p. 18)

5. Mesotritia maerkeli sp. nov. (p. 24)

6. Nothrus springsmythi sp. nov. (p. 26)

ACKNOWLEDGMENTS

I am indebted by my colleague Mr. D. Macfarlane for advice on a number of
problems and to Mrs. E. Hyatt for assistance with sorting the samples.

REFERENCES

AOKI, J. 1959. Die Moosmilben (Oribatei) aus Siidjapan. Bull, biogeogr. Soc. Japan 21: 1-22.
BALOGH, J. 1961. Identification keys of World Oribatid (Acari) families and genera. A eta zool.

Budapest 7: 243-344.
— & CSISZAR, J. 1963. The zoological results of Gy Topal's collectings in South Argentina.

Ann. hist.-nat. Mus. hung. Zool. 55: 463-485.
BECK, L. 1962. Beitrage zur Kenntnis der neotropischen Oribatidenfauna 2. Nothridae,

Camisiidae, Heterobelbidae. Senck. biol. 43: 385-407.
EVANS, G. O. & Hyatt, K. H. 1958. The genera Podocinum Berl. and Podocinella gen. nov.

(Acarina: Mesostigmata) Ann. Mag. nat. Hist. (12) 10: 913-932.

1960. A revision of the Platyseiinae (Mesostigmata: Aceosejidae) based on material in the

collections of the British Museum (Natural History). Bull. Brit. Mus. (nat. Hist.), Zool. 6:
25-101.

GRANDJEAN, F. 1932. Au sujet des Palaeacariformes Tragardh. Bull. Mus. Hist. nat. Paris (2)
4: 411-426.

1934. Les poils des epimeres chez les Oribates (Acariens). Bull. Mus. Hist. nat. Paris (2)

6: 504-512.

1947. L'origine de la pince mandibulaire chez les Acariens actinochitineux. Arch. Sci.

phys. nat. 29: 3O5~355-

1952. Observations sur les Palaeacaroides (Acariens) (ire serie). Bull. Mus. Hist. nat.

Paris (2) 24: 360-367.

I952a. Observations sur les Palaeacaroides (Acariens) (36 serie). Bull. Mus. Hist. nat.

Paris (2) 24: 547-554-

1954. Etude sur les Palaeacaroides. Mem. Mus. Hist. nat. Paris (NS) 7A: 179-274.

I954a. Essai de classification des Oribates (Acariens). Bull. Soc. zool. Fr. 78: 421-446.

1957. Observations sur les Palaeacaroides (46 serie). Bull. Mus. Hist. nat. Paris (2) 29:

213-220.

I957a. L'infracapitulum et la manducation chez les Oribates et d'autres Acariens. Ann.

Sci. nat. (n) 19: 234-279.

1958. Au sujet de naso et de son oeil infere chez les Oribates et les Endeostigmata

(Acariens). Bull. Mus. Hist. nat. Paris (2) 30: 427-435.

I958a. Observations sur les Palaeacaroides (56 serie). Bull. Mus. Hist. nat. Paris (2)

30: 76-83.

PRIMITIVE CRYPTOSTIGMATID MITES FROM NEPAL 35

HAMMEN, L. van der. 1952. The Oribatei (Acari) of the Netherlands. Zool. Verh. Leiden 17:
I-I39-

- 1959. Berlese's primitive Oribatid mites. Zool. Verh. Leiden 40: 1-93.

- 1963. The Oribatid family Phthiracaridae. I. Introduction and redescription of
Hoplophthiracarus pavidus Berlese. Acarologia 5: 306-317.

- ig63a. The Oribatid family Phthiracaridae. II. Re-description of Phthiracarus laevigatus
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HAMMER, M. 1961. Investigations on the Oribatid fauna of the Andes Mountains. II. Peru.

Biol. Skr. 13 No. i.
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SHEALS, J. G. & INGLIS, W. G. 1965. The British Museum (Natural History) Expedition to

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TRAGARDH, I. 1932. Palaeacariformes a new sub-order of Acari. Ark. Zool. 24B: 1-6.

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JAPANESE AND OTHER OPHIUROIDS

FROM THE COLLECTIONS OF

THE MUNICH MUSEUM

AILSA M. CLARK

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 2

LONDON: 1965

JAPANESE AND OTHER OPHIUROIDS FROM

THE COLLECTIONS OF THE

MUNICH MUSEUM

AILSA M. CLARK

British Museum (Natural History)

Pp. 37-71 ; i Plate ; 6 Text-figures

BULLETIN OF

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ZOOLOGY Vo1- 13 No. 2

LONDON: 1965

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In 1965 a separate supplementary series of longer
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JAPANESE AND OTHER OPHIUROIDS FROM

THE COLLECTIONS OF THE

MUNICH MUSEUM

By
AILSA M. CLARK

THIS paper deals with some ophiuroids from the collections of the Zoologische
Staatssammlung des Bayerischen Staates in Munich entrusted to me by Dr. H.
Fechter, to whom I am deeply indebted, not only for the opportunity of studying
such interesting material but also for permission to retain a proportion of the dupli-
cate specimens for the British Museum collections.

Most of the material came from southern Japan (collected by Doderlein, Doflein
and Haberer) or from the colder waters of the northern Japan Sea (Schmidt, Brash-
nikow and Domaschnew) but a number of other localities, ranging from the Mediter-
ranean and West Indies to the Indian Ocean and western as well as eastern Pacific,
are involved, including some specimens from the " Albatross " collections originating
with the United States National Museum — the greater part of which has already been
worked up by Koehler, Doderlein and others. Unfortunately some of the specimens
were labelled only with collectors numbers or dates and in the case of the Japan Sea
collections the details were often given only in almost illegible Cyrillic script so that,
in some instances, only an approximate transliterated version can be given where
the name is untraceable in the atlases available. I am indebted to Mrs. H. Sabo for
help in deciphering the labels.

The most interesting family represented is the Amphiuridae, which has been the
subject of a recent revision by Fell (1962), involving extensive dismemberment of the
major genera Amphmra, Amphiodia, Amphipholis and Amphioplus. I am not
convinced that the resultant groups form natural genera and accordingly am retain-
ing the customary combinations of names in this paper.

A number of the species represented evoke no special comment ; accordingly the
records of these are simply listed at the end of the paper.

Family OPHIOMYXIDAE
Ophiobyrsa intorta (Koehler)

Ophiobyrsella intorta Koehler, 1922, 27-29, pi. iv, figs. 2-5, pi. xcii, fig. 4.

MATERIAL. " Albatross " station 5215, 12° 31' 30" N., 123° 35' 24" E. (Philip-
pines), 1,105 metres, 3 specimens ; station 5219, 13° 21' oo" N., 122° 18' 45" E.,
970 metres, 2 specimens.

4o A. M. CLARK

The type of Ophiobyrsella intorta was also taken in the Philippines by the
" Albatross ". The present specimens agree with Koehler's photographs and descrip-
tion of the type except that there is no row of median teeth up the apex of each jaw
but simply multi-serial thorny-tipped tooth papillae throughout. The papillae
altogether number about 20 on each jaw, including one or two almost superficial
papillae lateral to the apex which might be termed oral papillae. The apical papillae
deeper in the mouth are longer than the superficial ones. Koehler notes " Together
with the tooth papillae the mouth (oral) papillae form a little bundle of short, slender
and spinulose spines continuous with the teeth which are fairly regular in their
arrangement and which have exactly the same form as these papillae." [My italics],
If the " teeth " are as narrow and spiniform as the papillae in the type it would be
surprising if they formed only a single vertical row not flanked by other papillae. In
view of the otherwise close correspondence between Koehler's description and figures
and the present specimens, I think the only difference is one of terminology and
accordingly am referring intorta to the genus Ophiobyrsa, characterized by the presence
of few oral papillae, many tooth papillae and no proper teeth.

It should be noted that the type species of Ophiobyrsa is 0. rudis Lyman, not 0.
hystricis Lyman as stated by Koehler.

In comparison with the holotype of 0. rudis the present specimens differ notably in
the much smaller oral shields with the adorals meeting proximal to them, the fewer
and smaller disc platelets and spinelets and the absence of spinelets on the fragmented
dorsal arm plates. Ophiobyrsa synaptacantha H. L. Clark, from south-west Japan
differs in the separated adorals though it has similarly fragmented and spineless
dorsal arm plates.

Ophiobyrsa acanthinobrachia H. L. Clark

Ophiobyrsa acanthinobrachia H. L. Clark, 1911, 269-270, fig. 134 ; Matsumoto, 1917, 24.

MATERIAL. Doflein, no. 588, Yogashima, 150 metres, 31. x. 1904, i specimen.

Murakami (1944) has described a species Ophiobyrsa strictima, from unknown
locality, in his paper on the ophiuroids of Ogasawara and Yaeyama, the unique type
having the disc diameter 23 mm. He notes that it is closely related to 0. acan-
thinobrachia, which it resembles in having spinelets on the dorsal arm plates. The
differences between them given by Murakami are that strictima has oval rather than
rounded-triangular oral shields, narrower adoral shields, more compact disc scaling
and the tooth papillae arranged in three rather than two vertical rows.

The type of 0. acanthinobrachia is also large with the disc diameter 20 mm., whereas
in the present specimen it is only 7 mm., the arms being about 45 mm. long, giving a
ratio of i : 6-4 compared with i : 14 in the type of 0. acanthinobrachia. The disc
scales are slightly spaced and the tooth papillae are arranged in two vertical rows so
I am referring the specimen to H. L. Clark's species. Besides the tooth papillae,
which number about six in each vertical row with the longest ones highest in the oral
slit, there is an irregular cluster of smaller papillae at the superficial end of the two
rows as well as one slightly larger papilla lateral to the apex of the jaw each side,

JAPANESE AND OTHER OPHIUROIDS 41

which might be termed an oral rather than a tooth papilla. On the distal side of
the superficial second oral tentacle there is an erect papilla or scale not mentioned by
H. L. Clark, though his figure shows an oral papilla just proximal to the tentacle pore.
The arm spines number up to five, as might be expected at this size, the type having
seven proximally.

Family OPHIAGANTHIDAE
Ophiacantha acanthinotata H. L. Clark

Ophiacantha acanthinotataH. L. Clark, 1911, 203-204, fig. 94 ; Matsumoto, 1917, 117 ; Murakami,

I942. 3-

MATERIAL. Doderlein, Enoshima 22, i specimen.

The disc diameter is 6 mm. and the basal arm segments have not more than nine
arm spines each side in comparison with the type, in which there were ten or eleven
spines proximally at a disc diameter of 9 mm. The two more distal oral papillae in
each series of three are both rather paddle-shaped, the third one not the widest, unlike
that of the type. Another slight difference is that a wedge-shaped area at the distal
end of each radial shield is left bare.

Ophiacantha rhachophora H. L. Clark

Ophiacantha rhachophora H. L. Clark, 1911, 201-202, fig. 92 ; Matsumoto, 1917, 119-120, fig. 30 ;
Murakami, 1942, 5-6 ; Djakonov, 1954, 4I-42' %• IO-

MATERIAL. Doderlein : Enoshima 22, i specimen ; Enoshima 25, 14 specimens ;
Yogashima, 7 specimens ; Yogashima, 2.xi.i88i, i specimen ; 7.xi.i88i. Ill, 250
fathoms (457 metres), 2 specimens; 2.xi.i88i. Ill, i specimen; no details, 2
specimens.

The number of oral papillae is variable in these specimens. As Matsumoto has
shown, the distalmost and thorniest one arises vertically from the adoral shield. One
or two of the other papillae are bifurcate dorso-ventrally at the tip but none are really
thorny. In one specimen with disc diameter 5 mm. there are only three oral papillae
each side of the apical one, including the clavate, thorny, adoral shield papilla, but
usually the number is five, though six may be found in some series as on one side of
the jaw figured by Matsumoto. The oral shields usually have a more prolonged and
acute proximal angle than is shown in H. L. Clark's figure of the type.

The thorny tentacle scales and distalmost oral papilla as well as arm spines make
this species easily recognizable from the sympatric Ophiacantha pentagona.

Ophialcaea congesta (Koehler)

Ophiacantha congesta Koehler, 1904, 103-104, pi. xxiv, figs, i, 2.
Ophialcaea congesta : H. L. Clark, 1915, 217.
Ophialcoea congesta : Koehler, 1922, 85, pi. xv, figs. 6, 7.

MATERIAL. "Albatross" station 5119, 13° 45' N.. 120° 30^' E. (Philippines),
721 metres, i specimen ; station 5618, o° 37' N., 127° 15' E. (Moluccas), 763 metres, i
specimen.

42 A. M. CLARK

As Koehler (1922) noted, only the basal dorsal arm spines are enlarged. In the
case of the specimen from " Albatross " station 5618 this applies to the uppermost
spines of the first three free segments ; on the second and third segments these spines
measure 1-9 mm. in length, whereas on the fourth segment tqe corresponding spine of
each side is only 1-3 mm. long. The length of the basal segments is about i-o mm.
These enlarged spines also differ in being distinctly clavate in shape. The disc
diameter is c. 9-5 mm. but the disc is rather distorted. The arm length is c. 50 mm.
There are four arm spines basally but the number increases to five beyond the thir-
teenth segment or thereabouts. The dorsal arm plates of at least the proximal half
of the arm each have a row of minute thorns along the distal edge. Neither of these
last two features was observed by Koehler in his material. This specimen also differs
from his description in the apical oral papilla (or lowest tooth), which is much wider
than the lateral papillae, not similar to them. The disc granules are conical (or at
least appear so through the thin skin). The dorsal arm plates have the distal edge
convex, unlike those of the type, which Koehler described as having parallel sides,
but much like the specimen he figured in 1922.

Because of the thorns on the dorsal arm plates, this species runs down to Ophiogema
in Fell's generic key (1960) but differs markedly from the type species, Ophiogema
punctata Koehler, 1922, in having a coat of conical granules (or they could be called
very short spinules) on the disc rather than " a rather stout pointed spine " on each
disc plate ; the ventral arm plates are also different, the dorsal arm plates have thorns
rather than spinelets and in Ophiogema punctata the arm spines are more needle-like
and numerous, seven in number, none of them are clavate and they are " finely
echinulated " not smooth. With so many differences I have no doubt that the two
species are generically distinct. The rib-like concealed radial shields give Ophialcaea
congesta the appearance of a Euryalid rather than an Ophiacanthid.

The specimen from " Albatross " station 5119 is similar in size but only the first
two free arm segments, especially the second, have the uppermost spine enlarged.
The thorns on the dorsal arm plates are also less distinct.

Family OPHIAGTIDAE
Ophiactis macrolepidota Marktanner-Turneretscher

Ophiactis macrolepidota Marktanner-Turneretscher, 1887, 298, pi. xii, figs. 12, 13 ; Doderlein,
1898, 484, pi. xxxvii, fig. i ; Matsumoto, 1917, 155-156, fig. 37 ; Murakami, 1942, 8 ; 1943,
167.

MATERIAL. Doderlein : Yogashima, 2 specimens ; Kashiyama, i specimen.

Whereas the largest specimen of Ophiactis pteropoma in the present collection has
only three arm spines though the disc diameter is 4 mm., all three of these specimens,
which are smaller, have four spines on the second free arm segment, though the fol-
lowing segments have only three ; also the second from lowest spine differs in being
somewhat squared at the tip.

I agree with Matsumoto and Murakami that such Japanese specimens are almost
certainly conspecific with Doderlein's from Amboina but there is some doubt whether

JAPANESE AND OTHER OPHIUROIDS 43

this is also true of the holotype of 0. macrolepidota. Not only was it supposed to come
from " Sidney " (presumably Sydney, N.S.W.), a temperate locality (though a few
tropical species do have their southern limit there) and one from which there are no
later records, but also it was described as having the ventral side of the disc (rather
improbably for an Ophiactis} covered with skin only. If Marktanner's description
was wrong in this respect it is possible that his species is now known by another
name, there being several nominal species of Ophiactis found in that part of Australia.

Family AMPHIURIDAE
Amphiura iridoides Matsumoto

Amphiura iridoides Matsumoto, 1917, 205-207, fig. 56.
M onamphiur a iridoides : Fell, 1962, u.

MATERIAL. Doderlein : nr. Yogashima, i specimen ; Yogashima, 200 metres, i
specimen ; Enoshima 22 and 24, 5 specimens ; no details, i specimen.

One of the two from Yogashima particularly approaches Amphiura iris Lyman in
having the distal oral papilla thicker than is usual in iridoides, while on some jaws
the papilla appears to have a double apex, being preserved in the erect position, since
its thin cross-section is slightly bowed ; when appressed the papillae appear leaf-
shaped. There are six arm spines on one or two basal segments although the disc
diameter is only about 3-5 mm., whereas Matsumoto's holotype of A. iridoides with
disc diameter 4 mm. has only five arm spines. The type of Amphiura iris at a disc
diameter of 5 mm. has only four spines although Matsumoto's largest specimen of iris,
with diameter 5-5 mm., did have five spines. The Yogashima specimen also differs
from A. iris in having the disc scaling very smooth and the dorsal arm plates not
" humped ", agreeing in these characters with A. iridoides. The radial shields are
nearly three times as long as broad and distinctly divergent, whereas in the type
specimen of A. iris they are relatively larger and are parallel in alignment.

Amphiura iris Lyman
(Text-fig, i)

Amphiura iris Lyman, 1879, 23, pi. xi, figs. 302-304 ; 1882, 132, pi. xvi, figs. 4-6 ; Matsumoto,

1917, 204-205.
M onamphiur a iris : Fell, 1962, n.

MATERIAL. Enoshima, 230 fathoms (420 metres), i specimen.

The disc diameter is 5 mm. and the arm length 40 mm., a ratio of i : 8, whereas
Matsumoto's specimen measured 5-5/25 mm. or i : 4-5. Unfortunately the holotype
has all the arms badly broken and no estimate of their length can be made. The
Enoshima specimen has the two infradental oral papillae of each jaw spaced from
one another with a third papilla or tooth at the same level between them ; on one
jaw, however, the infradental papillae are placed asymmetrically. In the holotype

A. M. CLARK

FIG. I. Amphiura iris Lyman. Holotype, B.M. no. 82.12.23.104. (a) Dorsal and (b)
ventral views of parts of the disc and oral area. In (a) the scales between the radial shields
are very indistinct and some imagination has been used in showing their limits ; never-
the less, they are certainly more numerous than shown in Lyman's figure, where only one
row is drawn. In (b) one of the four distal oral papillae included is displaced and seen
edge-on.

of A. iris the infradental papillae are closer together on three jaws, the small lowest
tooth being just above and partially between them ; a fourth jaw is damaged but the
fifth has a cluster of terminal papillae almost like an Ophiopsila, though these too are
damaged. In other respects the Enoshima specimen agrees with the type, that is in
the disc scaling, long parallel radial shields, shapes of the arm plates (the dorsal ones
slightly humped), the form of the distal oral papilla and the large single tentacle scale.

Amphiura trachydisca H. L. Clark

Amphiura trachydisca H. L. Clark, 1911, 149-150, fig. 60 ; Matsumoto, 1917, 201 ; Murakami,
1942, 19.

MATERIAL. Enoshima, 230 fathoms (420 metres), i specimen.

H. L. Clark's figure is rather misleading since the disc scales of this species do not so
much bear tubercles as have their free edges thickened and erected. This is true not
only of this specimen but also of the type, which I have examined in the U.S. National
Museum.

Amphiura arcystata H. L. Clark

Amphiura acrystata (lapsus for arcystata) H. L. Clark, 1911, 145-148, fig. 58 ; Matsumoto, 1917,

203 ; Murakami, 1942, n.

Amphiura arcystata H. L. Clark, 1915, 224 ; Djakonov, 1954, 74-
Hemilepis arcystata : Fell, 1962, 10.

MATERIAL. Doderlein, no details, i specimen.

In this specimen the disc is almost completely scaled above, though interradially
the scales are obscured ; the ventral side is quite bare. Beyond the base of the arm
the middle spines become square at the tips and even slightly bihamulate but the tip
is never very broad and the general impression given by most of the spines is that they

JAPANESE AND OTHER OPHIUROIDS 45

are tapering. The dorsal arm plates are ovate, as in the holotype, not rhombic as in
the two other specimens figured by H. L. Clark.

Fell (1962) referred Amphiura arcystata to Hemikpis, which he characterized as
having the disc scaled above but partly or wholly naked below. This move was
evidently a compromise measure between the three specimens figured by H. L. Clark
(1911, fig. 58, p. 146) under the name of arcystata, two of which (including the holo-
type) have more or less extensive skin on the dorsal side interradially and centrally,
though the ventral side is quite naked, while the third specimen has a complete scale-
covering on both sides of the disc. In fact the type of arcystata conforms more closely
to the diagnosis of Fell's Amphinephthys, with type species Amphiura crossota
Murakami from the Caroline Islands, which similarly has scales only around the radial
shields. Matsumoto (1917) recorded some additional Japanese specimens as arcystata,
noting that the larger ones have discs of the " Ophionephthys-type " (i.e. with scales
only around the radial shields), implying that the smaller ones have more extensive
scaling, as with H. L. Clark's " Albatross " specimens. Djakonov (1954) also found
that some small specimens from the north-west Pacific have fine granuliform scaling
on both sides of the disc in contrast to the more usual nearly naked condition.

Unless the station number was incorrect (as H. L. Clark suggested might be
possible), the holotype of arcystata, together with the 25 " topotypes " from the same
station, was from Californian waters, though, with a single exception of doubtful
identity, all the other " Albatross " specimens were from Japan. However, in
support of the correctness of the " Albatross " station given for the type there are
further records from the eastern Pacific, notably by May, Nielsen and Ziesenhenne.
Although all the specimens of the two latter authors appear to have had mainly naked
discs, May noted that specimens from Monterey Bay usually show this condition but
sometimes have the disc completely scaled above.

If H. L. Clark, Matsumoto, Djakanov and May are right in considering specimens
with more fully scaled discs as conspecific with others with reduced scaling, it seems
to me inadvisable to set up generic distinctions based on the extent of the disc scaling
alone, as Fell has done. Though very useful in reducing the large number of species
of Amphiura to more manageable groups, the resultant subdivision is I think too
artificial for these groups to be acceptable as genera (or even subgenera) without
further limitation of characters. The second character used by Fell to delimit his
new genera, namely the number of tentacle scales, is also of doubtful value at the
generic level, in my view, but I hope to be able to examine this problem in the detail
which it deserves before too long.

Amphiura sp. cf. euopla H. L. Clark

see Amphiura euopla H. L. Clark, 1911, 144-145, fig. 57 ; Matsumoto, 1917. 201-202, fig. 55 ;

Murakami, 1944, 265-266.
Hemilepis euopla : Fell, 1962, 10.

MATERIAL. Doderlein, Kagoshima, i specimen.

This specimen has the disc diameter 5 mm. and differs from H. L. Clark's descrip-

46 A. M. CLARK

tion of the type of euopla (d.d. 10 mm.) in having the disc finely scaled ventrally as well
as dorsally, the radial shields just contiguous, the primary plates distinct, the oral
shields with no distal lobes, the distal oral papilla more pointed and the second and
third of the seven arm spines conspicuously bihamulate. However, Matsumoto and
Murakami both refer specimens to euopla with disc diameter less than 6 mm. having
the discs scaled ventrally, the primaries distinct, shorter oral shields and the middle
arm spines not just blunt but thorny-tipped (some appearing almost hatchet-shaped
in Matsumoto's figure while he uses the term "spur-shaped" to describe them).
The difference in the radial shields may be illusory since I found on examination of the
holotype that the scales only extend between the two radial shields of each pair for
the proximal two- thirds of their length, beyond which they gape apart from each
other, probably unnaturally. The artist has used his imagination in drawing scales
between the distal ends of the shields, which I believe were originally contiguous as
they are in this specimen. Even so, there are still so many differences between the
type and this specimen, together with Matsumoto's smaller ones, that I am not con-
vinced that they are conspecific. In addition to the characters already mentioned,
H. L. Clark notes that a young specimen of euopla with disc diameter less than 4 mm.
has only four or five arm spines, whereas this one at diameter 5 mm. has as many as
seven spines.

In running down this specimen in Matsumoto's key to the Japanese species of
Amphiura I found the key rather misleading. Both A. euopla and arcystata come
within the section with two tentacle scales, a single distal oral papilla and five to
seven arm spines, the last distinction being coupled with the nakedness of the disc at
least ventrally, despite the fact that H. L. Clark has referred fully scaled specimens to
arcystata and Matsumoto himself proceeds to do the same for euopla. Then the two
species are supposedly distinguished from each other by four characters, according to
Matsumoto. However, the first of these four is the thickness of the disc, said to be
thick in euopla and thin in arcystata, and I think that the thickness is too liable to be
influenced by abundance of food, seasonal conditions and preservation to allow its use
as a specific character. The second point is the extent of the scaling on the disc ;
this is given as less extensive in arcystata but, in view of the variation in this character
observed by both H. L. Clark and Matsumoto, this does not seem to provide a valid
distinction. Thirdly, there is the shape of the radial shields, described as " short " in
euopla. but " long and rather narrow " in arcystata ; although the type of euopla does
have fairly short shields, in the larger, and to a lesser extent also the smaller, specimen
figured by Matsumoto under the name of euopla, I would describe the shields as long
and narrow (if the proportions shown are true) . Also in the third specimen figured
by H. L. Clark under the name of arcystata (the one with the disc fully scaled), the
radial shields are much shorter than in the type and other specimen figured. Finally
the shape of the arm spines is given as " spur-shaped and rough at the tip " in euopla
as opposed to " conical, not rough " in arcystata ; this may be a genuine difference
but it should be pointed out that the single specimen in the present collection referred
by me to arcystata does have the tips of the middle arm spines beyond the base of the
arms slightly squared-off and there may even be a small thorn proximally and

JAPANESE AND OTHER OPHIUROIDS 47

distally, though this is quite inconspicuous in comparison with the much widened
bihamulate tip of the second and third spines in the specimen from Kagoshima which
I believe is conspecific with the smaller ones Matsumoto referred to A . euopla, if not
with the type of euopla.

Another Japanese species with which the Kagoshima specimen has some affinity
is A. pachybactra Murakami, 1942, from the Izu Peninsula. The latter similarly has
the disc completely scaled, two tentacle scales and seven arm spines (at a disc diameter
of 8 mm.) but the primary plates are not distinct (though this may well be expected at
such a relatively large size) , all the arm spines are square-tipped and the second to fifth
of them are thorny-tipped without being distinctly bihamulate, the distal oral papilla
is blunter (in this specimen it is spiniform), the dorsal arm plates are narrower and
the oral shields have a flat distal side, their shape being broadly pentagonal. In fact,
A. pachybactra is very similar to A. ambigua Koehler, 1905, of which a large specimen
with disc diameter n mm. from Indo-China (identified by Mortensen) is in the British
Museum collections. I have compared this specimen with the one from Kagoshima
and find that it too has the middle arm spines modified with bihamulate tips, but these
tips are much smaller, barely squared-off on the second and third spines and thus
more like the corresponding spines in the specimen from the Miinich collection which
I have referred to arcystata. However, A . ambigua also has the disc fully scaled, two
tentacle scales and divergent radial shields just contiguous distally, though it differs
in having long distal lobes on the oral shields and the distal oral papilla shorter and
blunter. In the specimen of A . ambigua figured by Koehler in 1922 (pi. 69, figs. 5
and 6) the oral shields appear to have a rather shorter distal lobe but it is still not as
short as the lobe in this specimen, where the length : breadth ratio of the whole
shield is 14-5 : 14 and the distal side has only a very obtuse angle.

Murakami compared A. pachybactra instead with A. rapida Koehler, 1930, which
is a synonym of A. poecila H. L. Clark, 1915, according to H. L. Clark in 1946, the
types of both originating in the waters of southern Australia. A. poecila has similar
oral shields, distal oral papillae, tentacle scales, fully-scaled disc with primaries
distinct and fan-shaped dorsal arm plates proximally, like the Kagoshima specimen
but it differs in having the radial shields quite separate.

It is suprising that Murakami (1944) referred specimens with the disc fully-scaled
to A . euopla rather than to his A . pachybactra without commenting on their inevitable
resemblance to the latter.

Amphiura sp. juv. aff. koreae Duncan
seeAmphiura koreae Duncan, 1879, 466, pi. x, figs. 18, 19 ; Matsumoto, 1917, 198-199, fig. 53.

MATERIAL. Doderlein, Kagoshima, 2 specimens.

Even the larger specimen has the disc diameter only 2-6 mm. It has two short
distal oral papillae like A . koreae but differs from Duncan's type specimen in having
the proximal end of the oral tentacle scale more nearly superficial and the two papillae
of each infradental pair often spaced from each other with a conical tooth in between.
Also the type of koreae has the inner distal oral papilla arising from the side of the

48 A. M. CLARK

oral plate rather than from the adoral shield as it does here. However, Liitken and
Mortensen (1899) and Matsumoto (1917) emphasize that the oral structure is very
variable in the types of A . diomediae, which is generally considered to be a synonym
of koreae.

I believe that Amphilepis diastata Murakami, 1942, may prove to be another
synonym of Amphiura koreae. Although Murakami describes it as having only two
oral papillae (i.e. one infradental and one distal) his figure shows two distal papillae,
the inner one arising from the distal end of the oral plate where it joins the adoral
shield.

The Kagoshima specimen also has some resemblance to Amphiura confinis Koehler,
1904, from the East Indies, especially in the superficial position of the oral tentacle
scale. The same character allies it with the specimens named Amphiura concolor by
Lyman from " Challenger " station 191 in the Aru Islands, which also have two distal
oral papillae on the adoral shield and the infradental papillae often spaced, conical
and with a cusped tooth between them. The " Challenger " specimens are much
bigger, even the smallest of them having the disc diameter 6 mm. and their consecutive
dorsal arm plates are quite separate from each other, also the radial shields are not
contiguous at all distally ; otherwise they are very like this specimen. I do not
believe that they are conspecific with the type specimen of A . concolor, described by
Lyman also in the " Challenger " report, since the type has rounded, closely-placed
infradental papillae, the dorsal arm plates just contiguous, the disc scales thicker, the
oral and adoral shields of different shapes and more numerous arm spines. Indeed,
I believe that the Aru Island specimens should be referred to A . confinis, though they
differ slightly in having the disc scales thinner and the disc as a result smoother than
in the types of confinis.

Amphiura inepta Djakonov

Amphiura inepta Djakanov, 1954, 77~79» £§?• 23-

Amphiura carchara : Djakonov, 1954, 80-8 1, fig. 25. [Non A. carchara H. L. Clark, 1911.]

Monamphiura inepta : Fell, 1962, n.

MATERIAL. Brashnikow, no, 18, June, 1899, (northern Japan Sea) 2 specimens.

The larger specimen has the disc diameter 6 mm. and there are four spines on each
side for the first 17 free arm segments on the only arm remaining attached. The
smaller specimen has the disc diameter c. 4-5 mm. and has four spines on about 12
proximal segments. The discs of both have the ventral scales spaced from each other
in transparent skin, some enlarged plates dorsally and centrally but not forming a
regular rosette, the oral shields very blunt proximally, the adorals barely or not quite
meeting and the dorsal arm plates oval or wide fan-shaped with a very obtuse proximal
angle.

The type specimen of A. inepta has the disc diameter 14 mm., while in the type of
A. carchara H. L. Clark it is 8 mm. Even at this relatively large size the latter has
no more than three arm spines, also its primary rosette is not mentioned by H. L.
Clark and so is probably indistinct, the disc skin is completely lacking in scales on the
ventral side, the adorals are separate interradially and the orals have a proximal

JAPANESE AND OTHER OPHIUROIDS 49

angle. The specimen from the northern Okhotsk Sea figured by Djakonov under the
name of A. car char a has the disc diameter 5-2 mm. but already has four arm spines
proximally and also agrees with the present material in the shape of the oral and
adoral shields and in the spaced scales of the ventral side of the disc. Djakonov
himself recognized ah1 these differences from the type of A . carchara but still did not
compare the specimen with A. inepta. Although in his key he included inepta
among the species with two tentacle scales proximally, rarely only one, the figure he
gives of the species shows only one scale throughout ; the two Brashnikow specimens
similarly have no more than one scale. Djakonov's material of A. inepta came from
La Perouse Strait (between Hokkaido and Sakhalin), southern Sakhalin and the
north-east Okhotsk Sea.

Amphiura digitula (H. L. Clark)

(Text-fig. 2)

Amphiodia digitula H. L. Clark, 1911, 162-164, fig. 70.

Amphiura digitula : Matsumoto, 1917, 199-200, fig. 54 ; Djakonov, 1954, 71-

Amphiura leptopholida H. L. Clark, 1915, 226-227, pi- lv> fig8- II> I2 [Possibly recognisable as a

distinct form of digitula.]
Diamphiodia digitula : Fell, 1962, 14.

MATERIAL. Doderlein : Enoshima 22, i badly damaged specimen ; Enoshima
24, 3 specimens without discs; (PDoflein), Kachiyama, 5 specimens with discs, 6
without ; Doderlein : Enoshima, 4 specimens of forma leptopholida.

The present material together with Matsumoto's observations and my drawings of
the types of some of H. L. Clark's nominal species from Japan, made in 1953 in the
U.S., suggest that Amphiura digitula tends to intergrade with Amphioplus ancistrotus
(H. L. Clark). This is not so surprising as it may seem at first sight since the two
genera are closely related and species of Amphiura with two distal oral papillae, such
as A . digitula, need only the development of a single intermediate papilla to bridge
the gap to Amphioplus. The species of both genera possess a first oral tentacle scale
higher in the oral slit each side of the jaw, which is completely absent in Amphiodia,
where the oral papillae are only numerically, not morphologically, intermediate
between those of the type species of Amphioplus with four papillae and of Amphiura
with two.

H. L. Clark in 1911 referred both digitula and ancistrotus to Amphiodia but when
Matsumoto studied the homologies of the oral papillae of Amphiurids he recognized
that, although digitula has two distal oral papillae, these are based on the adoral
shield rather than the oral plate and an oral tentacle scale (additional papilla in
Matsumoto's terminology) is present ; accordingly the species should be referred to
Amphiura. There are several comparable species of Amphiura, notably A. koreae,
in which there are two distal oral papillae, similarly arising from the edge of the adoral
shield (though the inner of the two may be based about the point of junction of the
adoral shield and the oral plate) . Even if Fell's Diamphiodia proves to be sufficiently
natural for recognition as a genus distinct from Amphiodia, there is no question that

50 A. M. CLARK

digitula could be referred to it since the type species of Diamphiodia has oral papillae
similar to Amphiodia. Matsumoto also referred Amphiodia ancistrota to Amphioplus
because of its total of four (sometimes even five) oral papillae together with an oral
tentacle scale.

Both these species have rather unusual paired digits outside the radial shields,
which prompted the name digitula ; also both have two tentacle scales of moderate
size, somewhat similar arm plates and radial shields which are divergent and more or
less narrow (the width is exaggerated in H. L. Clark's figure 69 (1911) of A. ancistrotus
in comparison with the type specimen). The type of Amphiura digitula is distin-
guished particularly by the spiniform shape of the inner one of the two distal oral
papillae and by the diastema between this papilla and the infradental one (charac-
teristic of the genus Amphiura as opposed to Amphioplus) also by the fine, almost
granuliform, scaling of the disc ventrally and marginally, contrasting with the
smoother and larger dorsal scales. In the type of Amphioplus ancistrotus there are
four rounded oral papillae in continuous series and the ventral disc scales are smooth
and not extremely fine.

Between or close to one or other of these two extremes come two other Japanese
amphiurids described by H. L. Clark in 1915 as the types of new species. One of
these was referred to Amphiura and the other to Amphioplus (incidentally supporting
Matsumoto's generic dispositions), so that Dr. Clark failed to compare them with
either digitula or ancistrotus since he had left these both in Amphiodia in 1911.

FIG. 2. Amphiura digitula forma leptopholida H. L. Clark. Holotype of A. leptopholida,
M.C.Z. no. 1365. (a) Dorsal and (b) ventral partial views of disc and oral area.

The first of these nominal species (fig. 2), which he named Amphiura leptopholida,
has oral papillae almost identical with those of the type of digitula, the inner of the two
distal ones being spiniform ; also it has pairs of digits outside the radial shields. The

JAPANESE AND OTHER OPHIUROIDS 51

only differences are that the ventral disc scales are not very small and granuliform
but smooth and somewhat larger and the second from lowest of the four arm spines
has a very slight terminal hook rather than a simple tip. The four specimens in the
Munich collection from " Enoshima " differ from the complete specimens labelled
"Enoshima 22 " or " 24 " and the rest from Kachiyama (all of which are immediately
recognizable as digitula) in having the ventral disc scales smooth, not granuliform,
and the second arm spine distinctly hooked on the segments beyond the base of the
arm. Since the disc is easily lost in this species, judging from the proportion of
incomplete specimens in the present collection, it is possible that the difference in
disc scaling at least is correlated with regeneration. Three of the four Enoshima
specimens exceed by 2 or 3 mm. the 6 mm. disc diameter of the types of both A.
digitula and leptopholida, so the presence of five rather than four arm spines basally
is only to be expected ; the fourth specimen, disc diameter 5 mm., has four spines
basally.

The differences being of such small magnitude, I believe that the types of digitula
and Uptopholida are conspecific, nevertheless I think it worthwhile to distinguish
specimens with smooth disc scales and hooked arm spines (if these characters prove
to be consistently linked) as forma leptopholida, though such infrasubspecific taxa
have no status in taxonomy, according to the Code of the International Commission.

The second nominal species of H. L. Clark, 1915, is Amphioplus lobatod.es. Apart
from minor differences in the proportions of the oral shields and lesser curvature of
the second arm spine, there seems to me no reason why it should not be referred to
the synonymy of A . ancistrotus.

The Munich collection also includes three Amphiurids which, like Amphiura
leptopholida , present an intermediate condition between A. digitula and Amphioplus
ancistrotus. These are described under the heading of the latter species.

Amphioplus ancistrotus (H. L. Clark)
(Text-figs. 3 and 4)

Amphiodia ancistrota H. L. Clark, 1911, 161-162, fig. 69.

Amphioplus ancistrotus: Matsumoto, 1917, 171-172, fig. 43; Chang, 1948, 54-55, fig. n, pi.

viii, fig. 7 ; Djakonov, 1949, 54, fig. 69 ; 1954, 6l >' Fell> I9&2, 17.
Amphioplus lobatodes H. L. Clark, 1915, 254-255, pi. vii, figs. 12, 13.

MATERIAL. No details, presumably southern Japan, 2 specimens ; Haberer,
Sagami Bay, 7. vii. 1904, 1 specimen of a distinct form ; Doderlein, Enoshima 24, 2
specimens of the same form.

As mentioned under the heading of Amphiura digitula, I consider Amphioplus
lobatodes to be a synonym of A . ancistrotus. The drawing of the type, which I made
in 1954 when visiting the United States, shows no digits distal to the radial shields in
the radius depicted. Unfortunately I omitted to notice whether this was also true of
the other radii. H. L. Clark's brief description, largely comparative with A . lobatus,
makes no mention of this feature. If digits are really absent, then it may be possible
to distinguish lobatodes from ancistrotus but I doubt this.

52 A. M. CLARK

In the two specimens with no detailed locality the oral papillae are all short and
rounded. Although H. L. Clark's figure of the holotype shows the third (from inner-
most) papilla as conical and somewhat elongated, I found on examination of the
holotype that the three inner papillae are all rounded, though subequal, while the
third one is not erect (fig. 3#) ; in fact they are just as described by Dr. Clark, accor-
dingly I have no doubt that these two specimens are conspecific with the type.

imm

FIG. 3. Amphioplus ancistrotus (H. L. Clark), (a) Holotype of Amphiodia ancistrota H. L.
Clark, U.S.N.M. no. 25601, oral papillae, (b) to (e) holotype of Amphioplus lobatodes H. L.
Clark, M.C.Z. no. 1480 ; (b) dorsal and (c) ventral partial views of disc and oral areas,
(d) segment with twenty-second dorsal arm plate, (e) arm spines of twelfth free segment.
In (c) the ventral scaling is in reality barely distinct, especially proximally where it is very
tenuous and broken in most interradii.

The other three specimens in the Munich collection, however, have the third oral
papilla, and to some extent the second also, erect and spiniform. Matsumoto's
figures 436 and c both show these papillae as elongated and acute, but in Chang's
drawing all the papillae are short and rounded as in the type of Amphioplus lobatodes
(fig. 3b-e). (It should be noted that Chang's specimen has the disc naked ventrally
so the identification may be incorrect, however the disc diameter was only 3 mm. and
the small size might contribute to this deficiency.) In most other characters, namely
the form of the arm plates, tentacle scales, hooked second arm spine and particularly
the paired digits outside the radial shields, these three specimens agree with the first
two mentioned but their oral papillae provide such a sharp contrast that I consider it
worthwhile to designate this form with the two middle oral papillae erect and spini-
form as forma anisopapilla of Amphiura ancistrotus. Although such infrasubspecific
names are not recognized by the International Commission on Zoological Nomen-
clature and have no status in taxonomy, I think they still have their uses for the sake
of comparison. The shape of the oral shields, the extent of contact of the adorals
and the proportions of the radial shields all appear to be variable in A . ancistrotus and
it is interesting that some of the shapes exhibited approximate to those found in
Amphiura digitula. (It may be noted here that H. L. Clark's figure of the holotype
of Amphioplus ancistrotus shows the radial shields shorter than they really are and
therefore relatively too wide.) There are two minor differences between the specimen
of forma anisopapilla figured and the type of ancistrotus, namely in the former the
disc scales are particularly indistinct and the dorsal arm plates appear thinner and

JAPANESE AND OTHER OPHIUROIDS

more transparent. However, I doubt whether these differences are significant.

The two specimens of unknown locality, which may be designated as forma ancis-
trotus, have disc diameters of 9 mm. and 6 mm. In both of them the second from
lowest arm spine is hooked but the larger specimen has five arm spines on the first
four or five free segments, while the smaller one has no more than four basally.

FIG. 4. (a) to (d) Amphioplus ancistrotus forma anisopapilla nov. Enoshima 24. (a) Dorsal
and (b) ventral partial views of disc and oral area, (c) segment with fourteenth dorsal arm
plate, (d) segment with twelfth ventral arm plate, (e) and (f) Amphioplus conductus
Koehler, holotype, U.S.N.M. no. 41161, (e) a pair of radial shields with hyaline processes
distal to them, (f) the second from lowest arm spine of a middle arm segment. In (a) the
dorsal disc scales are very indistinct, also the dorsal arm plate is semi-transparent and the
underlying ossicles are indicated below it. In (b) three of the third oral papillae shown
are more or less fore-shortened, whereas the fourth has become appressed.

The larger Enoshima specimen of forma anisopapilla (fig. ^a-c) has the disc 7 mm.
in diameter. The radial shields are 1-3-1 -45 mm. long and 0-4 mm. in maximum
width ; the proximal angle is acute. The smaller specimen from the same locality
has the disc 3 mm. across and differs in having the oral shields longer and more acute
proximally. The Sagami Bay specimen has the disc diameter just over 7 mm. and,
unlike the other two, has the primary rosette just distinct ; its radial shields are
smaller, with length : breadth 0-8 : 0-2 mm. and only the third oral papilla is spiniform.

There are a few species of Amphioplus which similarly have the middle oral papillae
spiniform, but all of them have only a single tentacle scale. They include A. dispar
(Koehler), 1897, from the Indian Ocean, A. aciculatus from off the Congo and A.
acutus from the Antarctic, both species of Mortensen, 1936 (Discovery Report), also
A. gastracantha and notacantha (Liitken and Mortensen), 1899, from the East Pacific,
though the two last-named differ further in having a few disc spinelets so that they
may not be referable to Amphioplus at all but to Amphiacantha.

Amphioplus ancistrotus is closely comparable with A. diacritus Murakami (1943,
p. 225), the type of which has the disc diameter as much as 10 mm. The plates of

A. M. CLARK

the primary rosette in A . diacritus are small but distinct, the radial shields appear to
be longer and narrower than in ancistrotus and not so divergent, also the arm spines
number only three even at this large size, the middle one the longest but evidently
not hooked at the tip (though, as all the arms are said to be broken at the base, since
specialized spines only develop further out on the arm, in most Amphiurid species
that have them, it is possible that all the segments with hooks have been lost).
A. diacritus is possibly more closely related to A. rhadinobrachius H. L. Clark,
another Japanese species with only three arm spines. It is noteworthy that in
Murakami's figure of A . diacritus there are paired processes distal to the radial shields
though none are mentioned in his description.

A few other species of Amphiurids from the western Pacific have similar processes.
One such is Amphiacantha acanthina (H. L. Clark), also from Japan, which has distal
angles to the dorsal arm plates like Amphioplus ancistrotus but differs in having only
three arm spines as well as in the generic character of possessing spines on the disc.
A second species is Amphioplus conductus Koehler from the Philippines, in the
holotype of which I found paired hyaline bifurcating digits distal to the radial shields
(fig. 4^) not observed by Koehler. The only significant differences I can see between
conductus and ancistrotus are that, at a disc diameter of 6 mm., A. conductus already
has six arm spines on some basal segments and the second of these is not hooked but
bifurcated ; also the dorsal arm plates lack the marked distal angle.

Amphioplus asterictus H. L. Clark

(Text-fig. 5)
Amphioplus asterictus H. L. Clark, 1915, 252, pi. vii, fig. 9-11.

MATERIAL. Haberer, Sagami Bay, 7. vii. 1904, i specimen lacking the disc.

3mm

FIG. 5. Amphioplus asterictus H. L. Clark. Holotype, M.C.Z. no. 1486. (a) Disc and one
arm base viewed dorsally, (b) two jaws and part of the disc viewed ventrally, (c) spines
of two proximal segments. In (a) the primary disc scales are slightly hollowed, not
thickened.

JAPANESE AND OTHER OPHIUROIDS 55

Since H. L. Clark published only photographs of the type, I give here a drawing of
it which I was able to make at the Museum of Comparative Zoology, thanks to the
kindness of Dr. Elisabeth Deichmann.

Amphioplus japonicus forma parvus (Matsumoto)

Amphioplus relictus (part), Koehler, 1922, 180, pi. Ixxi, figs. 7, 8 ; Fell, 1962, 17. [Non A.

relictus (Koehler), 1898].
Ophiophragmus japonicus var. parvus Matsumoto, 1941, 334-336, figs. 3, 4.

MATERIAL. Doderlein: Tango, 40 fathoms (73 metres), I specimen; Tango 4, 7
specimens ; Kagoshima, 6 specimens ; no details, 7 specimens.

The generic position of Ophiophragmus japonicus has been the subject of some
controversy. Matsumoto (1915) included the species in Ophiophragmus because of
the erect marginal scales, despite the fact that it has four oral papillae. Since the
possession of only three papillae is partly diagnostic of Ophiophragmus, H. L. Clark
(1918, p. 271) referred japonicus to Amphioplus noting that the erect " fence " of
marginal scales is not homologous with the articulated marginal papillae or spinelets
found in the other species of Ophiophragmus, including the type, 0. wurdemanni
(Lyman), 1860 (of which no good figures existed up to that time, Koehler (1914)
having confused the issue by publishing photographs of Amphiodia limbata under
the name of Ophiophragmus " wundermanni "). Likewise H. L. Clark referred
Ophiophragmus affinis Duncan, 1887, to Amphioplus [where the name became invalid
as a homonym of A. affinis (Studer)], noting that it is probably identical with A.
relictus (Koehler), 1898. Matsumoto had considered Duncan's affinis, which came
from the Bay of Bengal, to be a close relation of japonicus together with Amphi-
pholis andreae Liitken, 1872, from Java, Amphiura praestans Koehler, 1905, from
the Flores Sea and Amphiodia periercta H. L. Clark, 1911, from Alaska and Oregon,
U.S. [The last-named of these is a true Amphiodia but the others are currently
referred to Amphioplus.']

In 1922 Koehler also referred affinis Duncan, as well as his own Amphiura relictus,
to Amphioplus on the grounds of differences in the internal structure in comparison
with Ophiophragmus wurdemanni.

In 1941 Matsumoto noted that the additional oral papilla above the main series of
superficial papillae (alternatively called the first oral tentacle scale) is absent in
japonicus unlike the type species of Amphioplus, A. tumidus (Lyman). Accordingly
he disagreed with H. L. Clark's transfer of the species to Amphioplus and retained it in
Ophiophragmus. Having removed the three distal oral papillae of one series in a
specimen from this collection, I can confirm that Matsumoto 's observation is correct.
However, in the closely related species Amphioplus hastatus (Ljungman) the oral
tentacle scale may not be visible in some specimens, even when the overlying papillae
are displaced, though its development is clearly variable since in others a small scale
or papilla can be distinguished. A. hastatus may also have spinous projections on
the uppermost row of ventral scales similar to those occurring in japonicus, as
Mortensen (1940, Echinoderms from the Iranian Gulf) has shown.

A. M. CLARK

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JAPANESE AND OTHER OPHIUROIDS 57

Accordingly, together with Fell (1962), I agree with H. L. Clark tha.t japonicus is
better referred to Amphioplus, although I believe that, together with A. hastatus,
depressus, relictus, andreae, laevis, praestans, megapomus, miyadii and some others, it
may be possible to distinguish it as forming part of a subgenus marked off from
Amphioplus tumidus, the type species of the genus, by the reduction of the oral ten-
tacle scale coupled with linear arrangement of the oral papillae, contiguity of the
radial shields and enlargement of the two tentacle scales of each arm pore.

The present specimens are referred to Matsumoto's forma parvus, of A. japonicus
because none of them have the row of enlarged outermost dorsal disc scales shown by
Matsumoto in his figures of japonicus itself. I have not seen any specimens of the
latter and am unable to assess whether or not the forma is worth retaining.

The table given here shows some measurements of 21 specimens of A. japonicus
forma parvus. It indicates that the relative size of the radial shields decreases as
growth proceeds, from a maximum of just under half the disc radius at about 3 mm.
disc diameter to a minimum of only a quarter when the diameter is over 8 mm.
Since this ratio has been used to distinguish between related species belonging to this
section of the genus Amphioplus it is clearly advisable that the total size of the disc
should be taken into account when making such comparisons.

When describing parvus in 1941, Matsumoto commented that it is allied to Amphio-
plus megapomus H. L. Clark. Having examined the discless type specimen of
megapomus I cannot agree with this. The dorsal arm plates of megapomus have a
median distal peak, so appearing trilobed, and the arm spines are much more acute
than in japonicus. In 1915, H. L. Clark noted that complete specimens of A.
megapomus have relatively wide radial shields as in A. japonicus. Despite this, I
believe that A. megapomus is more likely to prove conspecific with A. miyadii
Murakami, 1943, also of Japanese origin, since it agrees in the oral structure and in
the distinctive trilobed dorsal arm plates. Regeneration of the disc in those species
of Amphioplus which are particularly liable to shed it, such as A. integer, results at
first in abnormally short radial shields, though their relative length increases as
growth proceeds. If the specimens with discs studied by Clark in 1915 were regenera-
ting, the shields would probably be abnormally short. However, another possibility
is that these intact specimens were not conspecific with the types of megapomus since
Clark noted that their arm plates were not identical. A . miyadii is certainly more
closely related to A . laevis (Lyman) and praestans (Koehler) with similarly elongated
radial shields, than it is to A . japonicus.

In contrast to the type of forma parvus, the majority of the present specimens
have the six primary plates distinct, if not by larger size then by a slight hollowing of
their surface. In the smallest specimens the primaries are partly contiguous with
each other, having interstitial scales only at their corners (as in Koehler's photograph
of the small type of A. hastatus (1927), Ark. Zool., 19, pi. 3, fig. 2) but in larger speci-
mens the primaries are more or less widely separated.

In 1922 Koehler referred sixty-four specimens from Kagoshima, Japan, as well as
a number of Philippine specimens to Amphioplus relictus, noting that the relative
width of the oral shields is variable and that some specimens (of which he figures one

58 A.M. CLARK

from Kagoshima) have a distinct marginal row of erect disc scales. I have found
that some of the specimens earlier (1905) identified as A. relictus by Koehler, at least
those from " Siboga " stations 51 and 71 (Makassar and Molo Strait in the East
Indies), have a distinct enlarged row of marginal disc scales contrasting with the
uppermost row of smaller ventral scales, which are erect and project to form a scal-
loped edge to the disc, as in some specimens of A. japonicus. Nevertheless, I think
that Koehler's Kagoshima specimens are more likely to be referable to A . jpaonicus
than relictus since the ventral view of one (1922, pi. 71, fig. 7) suggests that the distal
edge of the oral shields is simply convex as in all the present specimens, without
having a distinct, more or less constricted, distal lobe, as occurs even in Koehler's
Manila specimen (pi. 71, fig. 6) where the shields are particularly wide in comparison
with those of most Philippine and East Indian examples of the species, in which the
distal lobe is usually as long as the proximal angle. [In the Manila specimen the
shape is very like that found in one of the syntypes of A . andreae, seen by me at the
Museum of Comparative Zoology, and it is possible that it should have been referred to
andreae and not to relictus.']

As these remarks indicate, the distinctions between Amphioplus hastatus, depressus,
relictus, andreae and japonicus are very subtle and in need of further consideration
when a true assessment of variation and growth changes can be made together with
re-examination of the types.

Amphiodia craterodmeta H. L. Clark

Amphiodia craterodmeta H. L. Clark, 1911, 155-157, fig. 65 ; Matsumoto, 1917, 182 ; Djakonov,

1938, 463 ; 1949, 54, fig. 72 ; 1954, 62.
Diamphiodia craterodmeta : Fell, 1962, 14.

MATERIAL. Domaschnew, nos. 53 and 55, 20. vi. 1900 and 27.vii.i9oo, 4 speci-
mens ; Brashnikow, no. 6, July, 1899, 2 specimens ; Schmidt, nos. 9 and 10, Mauka,
south-west Sakhalin, 46-47 sagenes (c. 100 metres), S.vi.igoi, 19 specimens. (All
N. Japan Sea).

Of the two specimens from Brashnikow's number 6, the larger has the disc diameter
8 mm. and the smaller 4-5 mm. The former has no distinct primary disc scales, the
oral shields are very small with the distal lobe both sunken and constricted, also the
infradental papillae are widely spaced and head the series of three oral papillae each
side and the disc plates are all extremely well-defined. The smaller specimen from
no. 6 differs in having the primaries much larger than the other scales.

In comparison with Amphiodia fissa, this species differs in having narrower (but
still mainly contiguous) radial shields, entire dorsal arm plates, smaller oral shields
and larger adorals, the primary disc scales not so different from the other scales, four
arm spines proximally and the ventral tentacle scales longer.

Amphiodia fissa (Liitken)
(Text-fig. 6)

Amphipholis fissa Liitken, 1869, 12-13 (3O-31)-

JAPANESE AND OTHER OPHIUROIDS

Amphiodia rossica Djakonov, 1935, 4^5, figs. 7 and 8 ; 1938, 465-466, fig. 8 ; 1949, 55, fig. 71 ;

1954, 64, fig. 1 8.
Diamphiodia fissa and rossica : Fell, 1962, 14.

MATERIAL. Schmidt, nos. 3 and 5, Mauka, south-west Sakhalin, 20 and 15 sagenes
(42 and 31 metres), 3.vi.i9Oi and 4.11.1901, 5 specimens ; no. 21, Posjet Bay, near
the bay of Minanosok, 12 sagenes (26 metres), 6 . iv . 1900, 4 specimens ; no. 81, entrance
to the bay of Tshogu-tschien-dogu, n specimens.

FIG. 6. Amphiodia fissa (Liitken). Specimen from Peter the Great Bay, M.C.Z. no. 4622,
labelled as A . rossica Djakonov. (a) Dorsal and (b) ventral views of parts of the disc and
oral area, (c) segment with fifteenth (split) dorsal arm plate, (d) lateral view of an arm
base showing the individually projecting disc scales ; the middle arm spines fore-short-
ened.

NOMENCLATURE. Liitken described this species in Danish from material collected
on the Amur coast (i.e. probably in the vicinity of Vladivostok), giving also a short
latin diagnosis but no figures. The very unusual feature for an Amphiurid of the
subdivided dorsal arm plates, which gave the species its name, taken in conjunction
with the peculiar erect marginal scales of the disc, leaves me in no doubt that
Amphiodia rossica Djakonov (also from the Amur region), which shares these charac-
ters, was based on material of this same species. In his original description of
Amphiodia rossica (1935) Djakonov made no mention of Amphipholis fissa but in
1954 he commented on the possible identity of the two ; however, with no published
figure of the type of A. fissa in existence he evidently thought that sufficient doubt
exists to justify the continued use of the name he had himself established.

Since Liitken's name Amphipholis fissa has remained in oblivion for well over 50
years, I submitted to the International Commission on Zoological Nomenclature a
petition to obtain the suppression of the name A. fissa as a nomen oblitum. This case

60 A. M. CLARK

is suspended in view of the controversy regarding nomina oblita in general raised at
the last colloquium on nomenclature.

REMARKS. The smallest of the five specimens from Mauka has the disc diameter
only 3'5 mm- and the proximal tentacle scale on the lateral plate is reduced or absent
on all but the basal pores, though the scale on the ventral arm plate is quite large.
The dorsal arm plates are nearly all split and fairly regularly so, also the marginal disc
scales are erect. In another small specimen with disc diameter 3 mm. the ventral
scale is also absent on segments beyond the tenth and sometimes as far proximally
as the fifth. The larger specimens have the disc diameter about 10 mm.

Yet another small specimen, with disc diameter only 2-75 mm., has the dorsal
arm plates fan-shaped and not split. It also has the plates of the primary rosette in
direct contact with each other, whereas in specimens with disc diameter 5 mm. or
more the primaries are all separated. The splitting of the dorsal arm plates seems
to start with the distal plates and progress proximally, but even when the plates are
not split the arms appear distinctly carinate. One specimen with disc diameter
4-5 mm. has the first five to seven plates entire ; it also has three arms regenerating,
each with a soft, worm-like tip, not yet distinctly segmented. There are two
tentacle scales only to about the twentieth segment, then one.

Other variable characters include the erection of the marginal disc scales, which
may be lacking, also the development of the distal lobe of the oral shields.

Since Djakonov's figures are unnatural with regard to the mouth parts and not
quite in agreement with the specimens I have seen, it seems worthwhile to give here a
figure of an oral angle drawn from a specimen presented by Djakonov to the Museum
of Comparative Zoology, Harvard.

Family OPHIOTRICHIDAE
Ophiothrix sp. ? eusteira H. L. Clark

Ophiothrix eusteira H. L. Clark, 1911, 265-267, fig. 132 ; Matsumoto, 1917, 222-223, fig- 60.

MATERIAL. Doderlein, y.xi.iSSi. Ill, 170 fathoms (311 metres), i specimen.

The single specimen has the disc diameter 7-5 mm. The radial shields are large,
c. 2-4 mm. long, and completely bare. The disc scales are clearly visible below the
sparse covering of small, spaced, tapering stumps, though some have about three
long thorns remaining on their tips. In the middle of the disc are a few short spines
only, agreeing more with Matsumoto's figured specimen than with the holotype,
which has the central spines very long. There is more resemblance to the specimen
figured by Koehler (1922, pi. 45, fig. 6) under the name of Ophiothrix koreana, which
I think would be better referred to 0. eusteira in view of my observations (below)
about the types of 0. koreana.

This specimen is only doubtfully identified as Ophiothrix eusteira because it already
has as many as nine arm spines on the second free segment, whereas the holotype,
with disc diameter 1-5 mm. greater, has only six or seven spines, according to H. L.
Clark.

JAPANESE AND OTHER OPHIUROIDS 61

Ophiothrix koreana Duncan
(pi. I, figs. 3 and 4.)

Ophiothrix koreana Duncan, 1879, 473-476, pi. xi, figs. 28-32 ; H. L. Clark, 1911, 257-262,
figs. 127-128; Matsumoto, 1917, 220 ; Murakami, 1942, 20 ; 1943, 232-233 ; 1944, 267 ;
Djakonov, 1954, 88.

MATERIAL. Doderlein : Yogashima, 3 specimens; Yogashima, 2.xi.i88i, 3
specimens; Yogashima 2 & 3.xi.i88i, I specimen; Enoshima 18, 183 metres,
8.xi.i88i.II, i specimen ; Tango, 73 metres, 2 specimens.

Duncan's figures of the syntypes of Ophiothrix koreana were not very good so I give
here photographs of the upper surfaces of two of them. These show that, unlike
the specimens figured under the name of koreana by H. L. Clark, which subsequent
writers have taken as models of the species, their radial shields carry a number of
thorny, mostly trifid, stumps at their distal ends as well as proximally, though the
stumps appear more sparse on the shields than on the scales. The largest syntype,
with disc diameter 8 mm., has about 30 stumps on each radial shield including four or
five at the distal tip, while the smallest specimen, disc diameter 4-5 mm., has about 12
stumps on each shield. This smallest syntype, as well as one other out of the six,
has some thorny disc spines in addition to the stumps but the remaining four have no
spines and superficially resemble H. L. Clark's figure of Ophiothrix hylodes (1911,
p. 263, fig. 130), which Matsumoto and subsequently Clark himself referred to the
synonymy of Ophiothrix marenzelleri Koehler, 1904^. The type of hylodes has the
disc stumps (or " stout, blunt, rough spines " in Dr. Clark's terminology) with
irregular thorns along their lengths and at their tips rather than the trifid form found
in the types of 0. koreana and also in the specimens identified as 0. marenzelleri by
Koehler (1922). However, some of the specimens of the present collection also have
" stumpy spines " of this kind, so the form of the stumps cannot be considered as
diagnostic. H. L. Clark did not mention 0. marenzelleri in his work of 1911, but
commented that 0. hylodes might be " only an extreme variety of koreana ". Mat-
sumoto repeated this supposition and noted that " the sublittoral form (of marenzel-
leri) approaches 0. koreana more closely than the littoral form ", this sublittoral form
sometimes having a few long spines on the disc as well as the close covering of thorny
stumps or tubercles. Matsumoto did not comment on the density of tubercles on the
radial shields as being any less in the littoral form than in the sublittoral but Koehler
(19040, p. 104) noted that in the types of 0. marenzelleri the radial shields are almost
as closely covered with stumps as the rest of the disc.

In the present collection I have identified as 0. marenzelleri specimens in which the
stumps appear equally dense on the radial shields as on the rest of the disc and those
with the radial shields distinctly barer I have named 0. koreana. However, so close
are the types of 0. koreana to the form generally accepted as 0. marenzelleri that I
think the latter name might well be considered a synonym. Judging from the
variation of Ophiothrix fragilis in the north Atlantic it is quite possible that the
Japanese specimens hitherto designated as koreana, marenzelleri, hylodes and possibly
even eusteira H. L. Clark, 1911, all represent a single very variable species. Alter-
natively the specimens with almost or completely bare radial shields such as those

62 A. M. CLARK

figured under the name of 0. koreana by H. L. Clark in 1911 might be referred to
0. eusteira. A much larger collection than that available to me is necessary before a
proper assessment of this problem can be made, so notorious is the genus Ophiothrix
for its variability.

Ophiothrix stabilis Koehler

Ophiothrix stabilis Koehler, 19040, 84-86, figs. 46-49 ; Matsumoto, 1917, 224 ; Murakami, 1944,

268.
Ophiothrix ciliaris : H. L. Clark, 1911, 257. [Non O. ciliaris (Lamarck), 1816.]

MATERIAL. Schmidt, 1901, Nagasaki, 3 specimens.

These three specimens were taken with one which I have referred to Ophiothrix
marenzelleri. They are immediately distinguishable from it by the disc stumps
which are shorter and better-termed tubercles than stumps ; they also have a coronet
of usually five to seven divergent points. Some disc spines are also present as in the
holotype 0. stabilis but there is a difference in the shape of the ventral arm plates.
None of these have the distal edge markedly convex as Koehler describes it in the
holotype (though his figure may not be reliable since it shows the ventral-most arm
spines with their hooks directed inwards instead of outwards or proximally, when
aligned perpendicularly, as in other species of Ophiothrix and in the three specimens
of 0. stabilis in the present collection).

Since Matsumoto used the convex or concave shape of the distal edge of the ventral
arm plates as an important character in his key to the Japanese species of Ophiothrix,
the identification of these three specimens as 0. stabilis invalidates this particular
dichotomy of his key. This dichotomy was already untrue for 0. eusteira, included
with stabilis as having convex edges to these plates, whereas only two proximal
plates are so depicted in H. L. Clark's figure of eusteira and none at all in Matsumoto's
own figure, where all the plates appear incurved in the middle of the distal side.

Ophiothrix panchyendyta H. L. Clark

Ophiothrix panchyendyta H. L. Clark, 1911, 264-265, fig. 131 ; Matsumoto, 1917, 219.

MATERIAL. Okinose Bay, 600 metres, i specimen.

The disc spines appear longer than shown in H. L. Clark's figure of the type. The
disc diameter of this specimen is 10-5 mm. and the majority of the spines towards
the centre of the disc are 1-7-2-0 mm. long. The length of the corresponding spines
was not specified in the type. There are hardly any short stumps on the disc besides
the spines. Close to the proximal interradial edge of each radial shield is a row of
four to eight short, tapering thorny-tipped spines or stumps, but superficially the
radial shields appear relatively large and very bare. There is no spine in the middle
of the first dorsal arm plate, unlike the type of 0. panchyendyta, but the arm plates
are similarly conspicuously granular in surface texture. Although H. L. Clark sus-
pected that neither of these last two characters might prove to be diagnostic, the
latter at least holds good for this specimen.

JAPANESE AND OTHER OPHIUROIDS 63

Ophiopsammium rugosum Koehler
Ophiopsammium rugosum Koehler, 1905, 116-117, pi- x"i» ngs- 4~6 ; 1930, 197.

MATERIAL. Sprater, Gulf of Siam, 1910/4500, clinging to a pennatulid, 10 speci-
mens.

The outlines of the radial shields are visible through the skin and granules. The
disc spines are very variable in occurrence, even in different interradii of the same
specimen ; there may be just two widely separated spines right on the edge of an
interradius or about 10 conical spines more or less inset or some shorter conical
tubercles (all these with truncated tips). One specimen has two to four very short,
thick, pointed tubercles in each dorsal interradius level with the tangent to the distal
ends of the radial shields, then at the very edge of the disc or slightly towards the
ventral side there are two separate clusters each of about four short conical spines.
Another specimen has about 35 very large low granules in the central part of the disc
and central interradial areas, not projecting out of the skin.

The colour of the disc is pinkish dorsally and dark-red ventrally. There are also
patches of intense black spots dorsally of varying extent and conspicuousness in the
different specimens. One has particularly large spots in asymmetrical areas of the
disc and arms, so that one pair of radial shields and one adjacent shield are covered
with big spots and one pair with only fine spots, while the rest are parti-coloured.
On the arms the spotted patches extend over about four segments. The tube feet
are dark red.

These specimens appear to have been epizooic on the pennatulid, since they are well
entwined within its branches. Possibly the peculiarities of the genus are correlated
with this habit.

Family OPHIODERMATIDAE
Ophioconis permixta Koehler

Ophioconis permixta Koehler, 1905, 14-15, pi. ii, figs. 4 & 7 (5 in caption).
Ophiurodon permixtus : Matsumoto, 1915, 84 ; 1917, 315 ; Koehler, 1922, 352.

MATERIAL. Xenia reef, Dar-es-Salaam, Tanganyika, 3 specimens.

The use of the original generic name for this species results from a comparison
between the three present specimens and two examples in the British Museum
collections of Ophioconis forbesi (Heller), 1863, the type species of Ophioconis Liitken,
1869, one specimen from the Adriatic originating with Dr. Heller himself and the
other from La Ciotat in the south of France. Unfortunately I have no material of
Ophioconis grandisquama Koehler, 1904, the type species of Ophiurodon Matsumoto,
1915 but there are several examples in the British Museum collection of Ophioconis
cupidum Koehler, which Matsumoto referred to Ophiurodon together with Ophioconis
cincta Brock and 0. permixta Koehler, all of which I consider to be congeneric with
Ophioconis forbesi.

Matsumoto's restricted Ophioconis included only 0. forbesi and 0. brevispina
Ludwig, 1880, both from the Mediterranean and neither known to Matsumoto at

64 A. M. CLARK

first hand. He distinguished Ophioconis from Ophiurodon, Ophiuroconis , Ophiuro-
chaeta, Ophiolimna and Ophiarachna by the arm spines, which he said are " very
short, lying flat on the arm, hyaline ". In fact the dorsal arm spines of the first two
free segments on both specimens of Ophioconis forbesi seen by me are equal to or even
slightly longer than the corresponding dorsal arm plates and in the better-preserved
(French) specimen (pi. I, figs, i & 2) most of the spines are erect, only those of the
distal parts of the arms having become flattened against the arms in preservation.
In Ophioconis brevispina and in 0. vivipara Mortensen, 1925, from Morocco, the arm
spines are much shorter, only about half as long as the corresponding segments near
the bases of the arms, judging from the figures, but it is 0. forbesi which is the type
species and criterion of the genus.

The three specimens of 0. permixta in the Munich collection are all small, with the
disc diameter little more than 3 mm. but their arm spines have very similar propor-
tions and alignment to those of 0. forbesi and, apart from the scattering of spinelets
among the granules of the disc, there is little difference between them, certainly not
enough to warrant a generic separation. Hertz (1927) has described a subspecies
nueva of Ophiurodon grandisquama based on four specimens from Madeira. She
maintains that these are very similar to a Japanese specimen of 0. grandisquamawhich
she has studied. Certainly the arm spines appear from her photographs to be rela-
tively longer than those of Ophioconis forbesi , since most of the spines of the proximal
half of the arm exceed or at least equal the segments in length. Nevertheless I can
see no difference of sufficient magnitude to justify generic separation. If Hertz is
right in asserting that nueva is closely related to grandisquama then Ophiurodon must
be considered as a synonym of Ophioconis. However, this needs confirmation from a
study of Japanese specimens. Certainly I do not think that the presence of disc
spinelets in permixta warrants a generic distinction from Ophioconis forbesi, par-
ticularly as H. L. Clark (1938) has found the occurrence of such spinelets in the
related species Ophioconis cincta Brock to be very variable. In eleven specimens
from northern Australia he found that five had a distinct marginal fringe of disc
spinelets, as in the type of cincta, while five others had these marginal spinelets poorly
developed or absent and the last specimen had both marginal and dorsal spinelets,
as in 0. permixta. This last observation casts doubt on the validity of permixta as
distinct from cincta, but much better sampling of these rather inconspicuous ophiuroids
is needed before a true appreciation of the specific limits can be reached. Ophiurodon
cupidum should also be referred to Ophioconis ; having no disc spinelets it is the
Indo-Pacific counterpart of the Mediterranean Ophioconis forbesi , the main difference
being only of colour pattern.

In the small specimens of 0. permixta the second tentacle scale is present for the
first six to twelve segments, on one arm even to the seventeenth.

This record from East Africa provides a considerable extension of range for
Ophioconis permixta, which was previously known only from the East Indian area.

Pectinura anchista H. L. Clark

Pectinum anchista H. L. Clark, 1911, 23-25, fig. i ; Matsumoto, 1917, 322.

JAPANESE AND OTHER OPHIUROIDS 65

MATERIAL. Haberer, no. 4168 (pt.), Sagami Bay, between Ito and Hatsushima
Island, c. 150 metres, March 1903, 2 specimens.

Matsumoto thinks that Pectinura anchista may be a synonym of P. cylindrica
(Hutton), 1872, from New Zealand. On morphological grounds it seems to me to be
equally likely that anchista is synonymous with P. aequalis (Lyman), 1880 and on
zoogeographical grounds more probable since the " Challenger " collected aequalis off
New Guinea and it has since been recorded by Koehler (1904 and 1922) from the
vicinity of Celebes and from the Philippines, while Murakami (1944) records it from
Yaeyama, Japan. Murakami did not state the size of his specimens but Lyman's
type of P. aequalis has the disc diameter 25 mm. and Koehler 's Philippine specimens
are equally large or larger still. The presence of 10 arm spines in Koehler's specimens,
whereas these from Sagami Bay have only seven proximally, may be attributable to
the much smaller size of the latter, the disc diameter being only u or 12 mm. The
holotype of P. anchista also has seven arm spines ; its disc diameter is 14 mm. I
think that a direct comparison between Japanese and East Indian specimens of
similar size will show the two to be indistinguishable.

Family OPHIURIDAE
Aspidophiura uniumbonata Murakami
Aspidophiura uniumbonata Murakami, 1942, 21-22, fig. 8.

MATERIAL. No details, presumably southern Japan, i specimen.

The disc diameter is 4-5 mm. and the longest stump of arm remaining attached is
only 3-5 mm. long. There is a suggestion of a boss on the central disc plate as in the
type of Aspidophiura uniumbonata. Two of the five primary radials are irregularly
subdivided but the other three are regular and about equal in size to the radial shields.
The oral shields are wider than in either A . watasei or A.forbesi but agree with those of
the type of uniumbonata. The only difference from the type is that the uppermost
arm spine of at least the first and second free segments is just longer than the
segment and the middle arm spine too is relatively longer, almost equal to the
segment in length. In the two other species the spines are shorter.

A number of other species were included in the collection for which no particular
comment was elicited. They are as follows :

Southern Japan

Astrodendrum sagamimtm (Doderlein)

Haberer, no. 4118, Fukuura, Sagami Bay, c. 150 metres, 1-2 . iii . 1903, 2 specimens.

66 A. M. CLARK

Ophiacantha pentagona Koehler

Doflein, no. 320, Sagami Bay, 180 metres, 25 .x. 1904, 10 specimens ; Doderlein :
Enoshima 14, i specimen; Enoshima 22, 2 specimens; 2.xi.i88i, IV, i
specimen.

Ophiopholis mirabilis (Duncan)

Doflein: Yogashima (Misaki), 150 metres, 31. x. 1904, 4 specimens; Sagami
Bay, towards Boshu, 120 metres, i.x.1904, i specimen; Haberer, Fukuura,
Sagami Bay, March, 1903, i specimen ; (Haberer?), no. 4356, Sagami Bay
7 . vii . 1904, i specimen.

Ophiopholis sp.? brachyactis H. L. Clark

Doderlein, Enoshima 18, 8.xi.i88i, II, i specimen.

Ophiactis pteropoma H. L. Clark

Doderlein : Enoshima 25, 2 specimens ; 2.xi.i88i, III, i specimen.

Amphipholis sobrina Matsumoto

Doderlein, Enoshima 22, i specimen ; Tango, 40 fathoms (73 metres), i speci-
men.

Amphiacantha acanthina (H. L. Clark)
Doflein, station 16, no. 22b, 3 specimens.

Ophiothrix marenzelleri Koehler

Schmidt, Nagasaki, 1901, i specimen ; Haberer, no. 4356, Sagami Bay, 7. vii.
1904, i specimen; Doflein, Yogashima (Misaki), 150 metres, 31. x. 1904, I
specimen; (Doflein?), Kachiyama, i specimen; Doderlein, 2.xi.i88i, III,
2 specimens.

Ophiomastix mixta Liitken

Schmidt, Nagasaki, i.iii.igoi, 3 specimens.

Ophiarachnella gorgonia (Miiller and Troschel)

Schmidt, Nagasaki, February-March, 1901, i specimen.

Ophioplocus japonicus H. L. Clark

Schmidt, Nagasaki, i.iii.igoi, i specimen.

Ophiura kinbergi (Ljungman)

Schmidt, Nagasaki, i specimen; Doflein, Tzushi (?), 130 metres, n.xi.i9O4,
2 specimens; Doderlein, Tagawa (?), 7.xi.i88i, II, i specimen ; (Doderlein?),
Tagawa, i specimen.

JAPANESE AND OTHER OPHIUROIDS 67

Ophiozonetta projecta (Koehler)

Doderlein, Yogashima, i specimen.

Ophiozonella longispina (H. L. Clark)

Haberer : no. 4168 (pt.), Sagami Bay, between Iso and Hatsushima Island,
c. 150 metres, March, 1903, 20 specimens ; Fukuura, Sagami Bay, March, 1903,
i specimen ; Doderlein : Enoshima 22, i specimen ; 2 .xi. 1881, III, i specimen.

Stegophiura sladeni (Duncan)

Haberer : no. 4356 (pt.), Sagami Bay, 7.vii.i904, i specimen ; no. 4168 (pt.),
Sagami Bay, between Ito and Hatsushima Island, c. 150 metres, March, 1903,
i specimen ; Sagami Bay, 1900, 2 specimens.

Stegophiura vivipara Matsumoto

Doflein, no. 582, Uraga Channel, Sagami Gulf, 150 metres, 22.x. 1904, i speci-
men ; (Doderlein?), Yogashima, 2 and 3.xi.i88i, 4 specimens.

Ophioleuce charischema (H. L. Clark)

Doderlein : Enoshima 22, i specimen ; Yogashima 2 and 3.x.i88i, i specimen ;
no details, i specimen.

Amur-Sakhalin (Localities probably east of Vladivostok, unless Sakhalin included).
Ophiacantha adiaphora H. L. Clark

Schmidt, no. 64, 19. v. 1900, i specimen.

Ophiacantha bidentata (Retzius)

Schmidt : no. 57, i specimen ; no. 48, off Cape Povorotny, 230-196 metres ;
1 8 specimens.

Ophiopholis aculeata (Linnaeus)

Schmidt : no. 68, 24^.1900, i specimen ; no. 46, Strelok Channel, i specimen ;
Brashnikow : no. 7, June 1899, I specimen ; no. 14, June, 1899, i specimen ;
no. 49, August, 1899, 4^ specimens ; nos. 19 and 14, 1899/1901, n specimens ;
Domaschnew, no. 59, July, 1900, i specimen.

Amphiura lepidevaspis Djakonov

Schmidt, no. 10 (pt.), Mauka, S.W. Sakhalin, 46-47 sagenes (c. 100 metres),
8 . vi . 1 901 , i specimen .

Amphiodia craterodmeta H. L. Clark

Domaschnew, no. 55, 20 . vi . 1900, 3 specimens ; no. 53, 27 . vii . 1900, i specimen ;
Brashnikow, no. 6 (pt.), July, 1899, 2 specimens ; Schmidt, nos. 9 (pt.) and 10
(pt.), Mauka, S.W. Sakhalin, 46-47 sagenes (c. 100 metres), S.vi.igoi, 19
specimens.

68 A. M. CLARK

Amphioplus macraspis (H. L. Clark)

Schmidt, no. 81 (pt.), 1900, 2 discless specimens.

Stegophiura nodosa (Lutken)

Schmidt, no. 47, 95-100 (? sagenes), 9. v. 1900, I specimen.

Ophiura leptoctenia H. L. Clark

Schmidt : nos. 8, 9, Mauka, S.W. Sakhalin, 46-47 sagenes (c. 100 metres),
S.vi.igoi, 5 specimens ; no. 48 (pt.) off Cape Povorotny, 230-196 metres, 1900,
9 specimens ; no. 58, same locality, 200-132 metres, 3 specimens.

Ophiura quadrispina H. L. Clark

Schmidt, no. 48 (pt.), off Cape Povorotny, 230-196 metres, 1900, i specimen.

Ophiura sarsi Liitken

Schmidt : nos. 9, 10 (pts.), Mauka, S.W. Sakhalin, 46-47 sagenes (c. 100 metres),
S.vi.igoi, 16 specimens; no. n, "east facing Patrok Bay (?)," 22.iii.i9OO,
i specimen ; no. 30 (locality illegible), 17.^.1900, I specimen; no. 48 (pt.), off
Cape Povorotny, 230-196 metres, 1900, 2 specimens ; no. 69, between Askold
and (?) Skriplev Islands, 55 sagenes (c. 115 metres), 24^.1900, 2 specimens:
Brashnikow : no. 6 (pt.), July, 1899, 3 specimens ; no. 35, July, 1899, 2 speci-
mens.

Ophiura sarsi vadicola Djakonov

Schmidt : no. 18, off Cape Povorotny, 3.^.1900, n specimens ; no. 24, Aniva
Bay, S. Sakhalin, 13-14 sagenes (27-29 metres), 28.viii.i9Oi, i specimen ; no.
46, Strelok Channel, 48-42 (? sagenes), 8. v. 1900, i specimen; no. 69 (pt.),
between Askold and (?) Skriplev Islands, 55 sagenes (c. 115 metres), 24^.1900,
i specimen ; Brashnikow, no. 35, i6.vii.i899, 7 specimens.

Ophiura maculata (Ludwig)
Brashnikow : no. 15, June, 1899, 2 specimens ; no. 27, July, 1899, i specimen.

Miscellaneous Localities

Aster onyx longifissus Doderlein

Albatross st. 2892 (Santa Barbara Channel, S. California) 35

st. 2979 (Anacapa Island, S. California) 30

st. 3198 (off central California) 23

st. 2891 (Point Conception, Oregon) 26

Aster onyx loveni Muller and Troschel

Albatross st. 2923 (off San Diego, California) i

st. 3787 (off Punta Gorda, N. California) i

st. 5637 (Molucca Islands) i

JAPANESE AND OTHER OPHIUROIDS 69

Asteroschema (or Ophiocreas) spp.

(Doderlein, 1927, regards Ophiocreas as hardly distinct from
Asteroschema even at the subgeneric level, but Mortensen and
Fell since consider Ophiocreas to be a distinct genus. With-
out the disc no distinction is possible) .

Albatross st. 5621 (Molucca Islands) i arm

(Doderlein recorded Asteroschema (Ophiocreas) gilolense from
this station) .

Albatross st. 5634 (Molucca Islands) i arm

(Doderlein recorded A. (0.) ambonesicum from this station).

Amphiura chiajei Forbes

Villefranche. Dr. Neresheimer i

Ophictais savignyi Miiller and Troschel

Ceylon. Haeckel i

Ophiactis simplex (Le Conte)

Academy Bay, Santa Cruz, Galapagos Islands

Foerster, 15. v. 1959 2

Ophiothrix sp. prob. fragilis (Abildgaard)

44° 48' N : 13° 45' E (northern Adriatic)

36 metres, 26. ix. 1895 i

Ophiothrix angulata (Say)

Kingston Harbour, Jamaica. Dr. Heitz, 30.xii.i9O3 c. 50

Macr ophiothrix hirsuta (Miiller and Troschel)

Ras Mohammed, Red Sea. Dr. Hofer, 1892 i

Ophiothela danae Verrill
Thursday I. Regenerating 3 arms and half the disc i

Ophiocoma scolopendrina (Lamarck)

Red Sea, Schadwan I. (? spelling), southside. H. Haas,
2y.iii.i956 5

Mombasa, E. Africa, May, 1960, Papp 5

Washington Museum, no. 46985, Philippines, E. A. Mearns, 1912

2 (badly damaged)

Ophioderma cinereum Miiller and Troschel
New Port, Curacao. Dr. Hellmich, 24.1937 i

7o A. M. CLARK

Ophioderma longicauda (Retzius)
Tunis. Pet. Muller, 13. ix. 1960 i

Bathypectinura conspicua (Koehler)

Albatross st. 5215 3

st. 5219 2

Ophionereis albomaculata E. A. Smith

Academy Bay, Santa Cruz, Galapagos Islands, Foerster, 15 . v. 1959 5

OphiuroUpis gelida (Koehler)

German Siidpolar-Expedition : 25 . vi . 1902 4

17. iv. 1902, 385 m. (one infested
with sponge lophon) 9

12 . viii . 1902 (largest with lophon) 5
14. iv. 1902 2

REFERENCES
CHANG, F.-Y. 1948. Echinoderms of Tsingtao. Contr. Inst. Zool. Acad. Peiping, 4, 33-104,

24 figs. 1 1 pis.
CLARK, H. L. 1911. North Pacific Ophiurans in the collection of the United States National

Museum. Bull. U.S. not. Mus., 75, 1-302, 144 figs.
— 1915. Catalogue of recent Ophiurans. Mem. Mus. comp. Zool. Haw., 25 (4), 165-376,

20 pis.
— 1918. Brittle-stars, new and old. Bull. Mus. comp. Zool. Harv., 62 (6), 265-338, 8 pis.

1938. Echinoderms from Australia. Mem. Mus. comp. Zool. Harv., 55, viii + 596, 63 figs.

28 pis.

- 1946. The Echinoderm Fauna of Australia. Publ. Carneg. Instn., 566, 1-567.
DJAKONOV, A. M. 1935. New Ophiurans of the genus Amphiodia from the Sea of Japan.

Dokl. Akad. nauk. SSSR., n.s. 3, 379-380.

- 1938. The Echinodermata of Siaukhu Bay (Japan Sea). Reports of the Japan Sea
Hydrobiological Expedition of the Zoological Institute, Academy of Sciences of the U.S.S.R.
in 1934, pt- T> 425-498, 10 figs. [In Russian].

1949. Definitions of Echinoderms from far-eastern seas. Bull. Pacif. Inst. Fish. Oceanogr.

30, 1-130, 22 pis. [In Russian].
1954. Ophiuroids of the Soviet Union. Tabl. anal. Faune URSS, 55, 1-136, 47 figs, i pi.

[In Russian].
DODERLEIN, L. 1898. Ueber einige epizoisch-lebende Ophiuroidea. In Semon, Zoologische

Forschungsreisen in Australien und dem Malayischen Archipel. Denkschr. med.-naturw.

Ges. Jena, 8, 483-488, pi. xxxvii.
DUNCAN, P. M. 1879. On some Ophiuroidea from the Korean Seas. /. Linn. Soc. (Zool.),

14, 445-482, pis. ix-xi.

- 1887. On the Ophiuridae of the Mergui Archipelago, collected for the Trustees of the
Indian Museum by Dr. John Anderson. /. Linn. Soc. (Zool.), 21, 85-106, figs. 28-40, pis.
viii, ix & xi.

FELL, H. B. 1960. Synoptic keys to the genera of Ophiuroidea. Zool. Publ. Viet. Univ. N.Z.,

no. 26, 1-44, 6 figs.
1962. A revision of the major genera of Amphiurid Ophiuroidea. Trans, roy. Soc. N.Z.

(Zool.), 2, 1-26, i pi.

JAPANESE AND OTHER OPHIUROIDS 71

HERTZ, M. 1927. Die Ophiuroiden der Deutschen Tief see-Expedition, i. Wiss. Ergebn.

Valdivia 22 (3), 59-122, 7 figs. pis. vi-ix.
KOEHLER, R. 1897. Echinodermes recueillis par 1' Investigator dans 1'Ocean Indien. i.

Les Ophiures de Mer Profonde. Ann. Sci. Nat., (8), 4, 277-372.

1898. Echinodermes recueillis par 1'Investigator dans 1'Ocean Indien. 2. Les Ophiures

littorales. Bull. sci. Fr. Belg., 31, 54-124, pis. ii-v.

1904. Ophiures de 1'Expedition du Siboga. i . Ophiures de mer profonde. Siboga Exped.,

45a, 1-167, 36 pis.

i9O4a. Ophiures nouvelles ou peu connues. Mem. Soc. Zool. Fr., 17, 54-119, 98 figs.

1905. Ophiures de 1'Expedition du Siboga. 2. Ophiures littorales. Siboga Exped., 45b,

1-142, 18 pis.

1914. A contribution to the study of Ophiurans of the United States National Museum.

Bull. U.S. nat. Mus., 84, 1-173, 18 pis.

1922. Ophiurans of the Philippine Seas and adjacent waters. Bull. U.S. nat. Mus.,

100 (3), 1-486, 103 pis.

- 1930. Papers from Dr. Th. Mortensen's Pacific Expedition, 1914-16. LIV. Ophiures
recueillis par le Docteur Th. Mortensen dans 1'Archipel Malais. Vidensk. Medd. dansk.
naturh. Foren., 89, 1-295, 2O pls-

LJUNGMAN, A. 1866. Ophiuroidea viventia hue usque cognita enumerat. Ofvers. Vetensk-

Akad. Fork. Stockh., 23, 303-336.
LUTKEN, C. F. 1869. Additamenta ad Historiam Ophiuridarum. c. Beskrivende og kritiske

Bidrag til Kundskab om Slangestjernerne. K. danske Vidensk. Selsk., 5 (8), 24-109, 3 figs.

- 1872. Ophiuridarum novarum vel minus cognitarum descriptiones nonnullae. Overs,
danske Vidensk. Selsk. Fork., 77, 75-158, 2 pis.

LUTKEN, C. F. & MORTENSEN, T. 1 899. Reports on an exploration off the west coasts of Mexico,

Central and South America, and off the Galapagos Islands. XXV. The Ophiuridae. Mem.

Mus. comp. Zool. Harv., 23 (2), 97-208, 23 pis.
LYMAN, T. 1879. Ophiuridae and Astrophytidae of the " Challenger " Expedition. II. Bull.

Mus. comp. Zool. Harv., 6 (2), 17-83, 8 pis.

1882. Ophiuroidea. Rep. Sci. Res. Challenger, Zool. 5, 1-386, 46 pis.

MARKTANNER-TURNERETSCHER, G. 1887. Beschreibung neuer Ophiuriden und Bemerkungen

zu bekannten. Ann. naturh. (Mus.) Hofmus. Wien, 2, 291-316, pis. xii, xiii.
MATSUMOTO, H. 1915. A new classification of Ophiuroidea : with descriptions of new genera

and species. Proc. A cad. nat. Sci. Philad., 67, 43-92.
1917. A Monograph of Japanese Ophiuroidea, arranged according to a new classification.

J. Coll. Sci. Tokyo, 38 (2), 1-408, 100 figs. 7 pis.

1941. Report of the biological survey of Mutsu Bay and vicinities. Sci. Rep. Tohuku

Univ., (4) 16, 331-344, 10 figs. pis. xxi-xxiii.

MURAKAMI, S. 1942. Ophiurans of Izu, Japan. /. Dep. Agric. Kyushu Univ., 7 (i), 1-36,

12 fififS.

I943. Report on the Ophiurans of Palao, Caroline Islands. Report on the Ophiurans of

Yaeyama, Ryukyu. Ophiurans from some Gulfs and Bays of Nippon. /. Dep. Agric
Kyushu Univ., 7 (4-6), 159-204, figs. 1-17 ; 205-222, figs. 1-2 ; 223-234, figs. 1-2.

I944. Report on the Ophiurans from off Ogasawara Islands and from off the Yaeyama

group, Nippon. Note on the Ophiurans of Amakusa, Kyusyu. /. Dep. Agric. Kyushu
Univ., 7 (7-8), 235-257, figs. 1-14 ; 259-280, figs. 1-5, pi. i.

PLATE i

FIGS, i, 2. Ophioconis forbesi (Heller). B.M. reg. no. 94. u . 19.2, from La Ciotat, S. of France.
X3-

FIGS. 3, 4. Two syntypes of Ophiothrix koreana Duncan, B.M. reg. no. 80.1.3.13, from the
Korean Straits, 42 metres, both X3.

Bull. EM. (N.H.) Zool. 13, 2

PLATE i

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THE FRESHWATER GASTROPOD
MOLLUSCS OF WEST CAMEROON

C. A. WRIGHT

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 3

LONDON: 1965

THE FRESHWATER GASTROPOD MOLLUSCS
OF WEST CAMEROON

C. A. WRIGHT

British Museum (Natural History)

Pp. 73-98 ; Plates 1-3 ; 21 Text-figs

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. 3

LONDON: 1965

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In 1965 a separate supplementary series of longer
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THE FRESHWATER GASTROPOD MOLLUSCS
OF WEST CAMEROON

By C. A. WRIGHT

SCHISTOSOMIASIS, the disease of man caused by blood-flukes of the genus Schistosoma,
is commonly associated with the savannah and semi-arid regions of Africa and the
Middle East. In 1953 a focus of infection with S. haematobium was reported by
Zahra in two crater lakes in the rain-forest area of West Cameroon. In 1957 Mandahl-
Barth (1957!)) reported two species of Bulinus (one of them new) from one of these
crater lake foci and, in reply to a request made to Dr. B. O. L. Duke of the Helminth-
iasis Research Unit at Kumba, I received a preserved sample of bulinid snails from
the second of the two infected lakes reported by Zahra. In order to obtain living
material of both snails and schistosomes from this unusual focus and to investigate
the possible wider distribution of the disease in West Cameroon a short visit was
made to the territory in October and November 1963. This paper reports the mala-
cological results of the expedition.

I am greatly indebted to Dr. S. P. Tchoungui, Federal Minister of Health of the
Republique Federale du Cameroun and to Colonel R. Aretas, Conseiller Technique
at the Federal Ministry of Health for their encouragement and support. It is a
pleasure also to acknowledge the facilities made available by Dr. G. G. Dibue,
Director of Medical Services, West Cameroon and Dr. B. O. L. Duke, Director of the
Helminthiasis Research Unit at Kumba. Special thanks are due to Mr. and Mrs.
Peter Moore of the Helminthiasis Research Unit for their kindness and hospitality ;
it is largely due to their help that a series of minor disasters were overcome and the
objectives of the expedition were achieved. Both in the field and subsequently
in the laboratory I was assisted by Mr. M. S. Bennett. This work was made possible
by grant No. AI-O365O-03 from the U.S. Public Health Service.

TOPOGRAPHY OF WEST CAMEROON

The territory now designated West Cameroon was formerly the south-western part
of the British Trusteeship territory lying between Nigeria and the French-adminis-
tered Cameroun. In a number of scientific papers the same area has been referred to
by different authors as West Cameroon, the Southern Cameroons and the North-
western Cameroons. The region extends about 450 miles north-eastward from roughly
4°N., 9°E. on the coast of the Gulf of Guinea to about g°N., I3°E. The country is
mountainous with dense rain-forest in the south-western part and open grassland
in the hills to the north-east ; its geography and geology have been very fully des-
cribed by Geze (1943). The mountains are partly tectonic in origin but the majority
are volcanic, forming a chain of which the Gulf of Guinea islands (Fernando-Poo,
Principe, Sao Thome and Annobon) are a south-western extension. Overlying the

3§

76 C. A. WRIGHT

pre-Cambrian bedrock of gneisses and gneissic granites are a sedimentary series of
Cretaceous age which form a band running roughly parallel to the coast and there is
another band in the Mamfe region. Overlying the Cretaceous deposits in the south-
western part of the country is a thick sheet of basalt of volcanic origin. Geze
quotes evidence suggesting that volcanic activity in Cameroon began in the Creta-
ceous period but Reyment (1954) does not support this view and considers that the
activity has been entirely post-Cretaceous. There are some sedimentary grits and
sandstones of Tertiary age in the coastal area and Quaternary deposits are confined
to the mangrove swamps which surround much of the coast. Volcanic activity
resulting in extensive layers of tuffs in some areas has continued into recent times and
the last eruption of Mount Cameroon occurred in 1954.

The crater lakes which were one of the major objectives of the present expedition
are considered by Geze to be the result of violent gaseous explosions of relatively
recent origin. This opinion is based on the absence of igneous material associated
with the craters, other than that of the older basalt sheet through which the eruptions
occurred or demonstrably younger elements. These younger igneous elements in-
clude the island in Lake Barombi Kotto (the remains of a small volcano which
appeared after the lake was formed) and the basalt stream on the north-east side of
Lake Barombi Mbo. Evidence of the relatively recent origin of the craters is provided
by the generally steep slope of their sides but in some this incline is less abrupt,
suggesting a considerable variation in their age. Typically the lakes are roughly
circular in outline and the internal slopes of the craters are thickly forested with
dense vegetation extending down to and overhanging the water's edge. Hydro-
graphic information is available for only the three main lakes in Kumba division,
Barombi Mbo, Barombi Kotto and Soden ; the first and last of these are steep-sided
and deep but the volcanic island in Barombi Kotto has filled a good deal of the lake,
making it relatively shallow and creating a more gently sloping shore, particularly
around the island. On the north-west side of Barombi Mbo the entering stream has
created a small, swampy delta. The extension of this silt deposition has formed a
shelf extending out into the lake about one hundred yards. The water depth over
this shelf probably does not exceed ten feet and there is considerable growth of
aquatic vegetation. Access to Lake Soden is difficult and the only point which was
visited had a very steep slope and exceedingly dense, overhanging marginal vegeta-
tion. The fringing forest appears to be uninterrupted and it is not known if there
are any shallow areas. Lake Ejaghem near Mamfe has less steep banks and forest
clearing has resulted in some gaps in the marginal vegetation ; this lake and the
three preceding ones have endemic fish populations which are utilized to varying
extents by the local people. In Bamenda Division there are a number of lakes
whose origin is uncertain but which have many of the characteristics of the craters
in Kumba Division. Only three of these were visited, Lakes Bambuluwe, Bafeng
and Wum ; the first two are small, steep-sided and with dense marginal vegetation
while Lake Wum is open and has some areas with a gently sloping margin with
aquatic vegetation. Fish have been introduced into Lake Wum and appear to be
thriving but there is no natural fish population in either Bambuluwe or Bafeng and

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 77

information from inhabitants of the village at Oku indicated that there are no fish in
Lake Oku either.

HISTORICAL

Cameroon has attracted a good deal of malacological attention as a result of its
rich and interesting fauna of terrestrial species but, for a territory so well-endowed
with rivers, streams and lakes, the records of freshwater species are poor, von
Martens (1877) described a large West African collection made by Professor Bucholz
and this included six species of fresh- and brackish-water prosobranchs from the
coastal region, mostly in the neighbourhood of Victoria. In 1891 the same author
reported on a collection made by Preuss in the area of the " Barombi-Station " and
mentioned the two large species of freshwater prosobranchs still found in Lake
Barombi Mbo and its tributary stream. d'Ailly's (1896) account based upon ex-
cellently documented collections made by the Swedish surveyor Dusen included
nine species of prosobranchs and de Rochebrune (1898) gave a list of species received
from Sjostedt, one of Dusen's colleagues, and mentioned two of the prosobranchs
present in d'Ailly's material. O. Boettger (1905) published a list of Cameroons
species based on material received from several collectors but here again the only
freshwater gastropods noted were prosobranchs. Two small collections made by
Dyke were reported upon by Spence (1925 and 1928) and included three species of
prosobranchs, two of which were described as new. The first basommatophoran
recorded was a planorbid, found on the northern slopes of Mount Cameroon and
described by C. R. Boettger as Australorbis camerunensis (1941). A Swiss expedition
in north-eastern Cameroon collected two freshwater prosobranchs not included in any
of the earlier accounts (Forcart, 1951) and the Danish expedition to the French
Cameroons in 1949-50 brought back only terrestrial species (N0stvik, 1956). Man-
dahl-Barth (1957^ described Bulinus camerunensis and recorded B. truncatus rohlfsi
from specimens collected by Mr. P. J. Moore in Lake Barombi Kotto. These two
species together with Boettger's planorbid are the only basommatophoran pul-
monates recorded from the area. The present account adds seven more planorbids,
one lymnaeid and one ancylid to the list as well as an additional prosobranch not
previously recorded.

Family PLANORBIDAE
Biomphalaria camerunensis (Boettger)

Australorbis camerunensis C. R. Boettger, 1941 : 121.

Biomphalaria camerunensis camerunensis Mandahl-Barth, igsya : 1142.

MATERIAL : Bambalang, N'dop Plain, Bamenda Division, about 70 specimens
collected from shallow water with thick mud bottom and scarcely any vegetation,
heavily shaded by thickets of low-growing palm trees. 6th November, 1963.

Shell (PI. I, figs. 16-23) : yellowish-brown, discoidal, flattened above with the
centre slightly depressed ; Whorls weakly angled beneath, umbilicus wide ; well-
developed spiral lines give marked granular microsculpture even on later whorls ;

?8 C. A. WRIGHT

larger shells irregularly rugose and patchily eroded. Nearly all shells up to 12-5 mm.
diameter have apertural lamellae (PI. I, fig. 23). The largest specimen collected
had a maximum diameter of 17-3 mm., umbilicus 7-2 mm. and height 5-4 mm.
The mean ratio maximum diameter/umbilicus diameter for 30 specimens of all sizes
is 2-67 and for the ratio maximum diameter /height the value is 2-75.

Anatomy : male copulatory organ large, penis-sheath longer than preputium
(Text-fig. 4), mean ratio penis sheath /preputium for a series of adult specimens is
1-42 (range 1-07-1-86), penis slightly shorter than the sheath ; prostate long with
30 or more primary diverticula, all with secondary and many with tertiary branches.
Albumen gland relatively small, oviduct and muciparous gland (the translucent
white proximal part of the uterus) short but the distal, yellow oothecal gland is long ;
vagina long, without marked dilatation, receptaculum seminis club-shaped (Text-fig.
4), vesicle about equal in length to duct. The pericardia of all specimens dissected
were packed with metacercariae of an unidentified trematode and the gonad and part
of the digestive gland of others were found to be completely destroyed by rediae of
probably the same species of parasite.

Radula (Text-fig. 10) : teeth small, tendency towards arrow-head shape of some
lateral mesocones but majority triangular, ecto- and endocones roughly equal ;
six to eight tricuspid laterals in each half-row, about the same number of intermedi-
ates, division of the endocone starts at about tooth 15, most marginal ectocones
undivided but some have outer edge serrated.

Biomphalaria camerunensis has so far been known only from the type series of
shells collected at Mongonge on the northern slope of Cameroon Mountain. Mandahl-
Barth (ig57a) described a sub-species B. c. manzadica from a few localities in the
Lower Congo and, on shell characters my material is closer to this sub-species than
the nominate form in that the umbilicus diameter is greater than the height. How-
ever, the illustration of the type specimen of B. camerunensis shows that it differs
from any of my specimens in that the aperture and terminal part of the body whorl
are deflected downward, a character which gives a misleading increase in the height
of the shell in very large specimens. All other members of the Biomphalaria sudanica
species group have the umbilicus diameter greater than the shell height as in my
material and it seems unwise to set B. camerunensis apart because this proportion
is reversed in a single, large, possibly aberrant specimen. Anatomically the present
material differs from all other members of the B. sudanica group (with the exception
of B. sudanica rugosa Mandahl-Barth, 1960) in that the penis sheath is consistently
longer than the preputium. The radula teeth conform to the general characters of
the sudanica group. Three shells (B.M. (N.H.) coll. no. 1936-6-2-24-26) from the
Batouri District of East Cameroon which compare well with my material were labelled
" Planorbis salinarum " by Connolly in 1936 and there is no doubt that they bear a
close resemblance to that species. However, the umbilical diameters and heights
of the Cameroons shells are proportionately slightly higher than in the type series of
B. salinarum. It is probable that snails reported from Yaounde in East Cameroon
as B. sudanica by Gaud (1955) are also B. camerunensis.

The presence of apertural lamellae does not appear to have been noted before in

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 79

members of the B. sudanica species group. Smith (1881) originally described the
form tanganyicensis (now considered by Mandahl-Barth (i957a) to be a sub-species
of B. sudanica) as a member of the genus Segmentina but there is no trace of lamellae
in any of Smith's specimens. The lamellae found in B. camerunensis do not differ
from those reported in B. pfeifferi and in various Puerto Rican species by Richards
(1963) who suggested that they may serve as supporting structures for the delicate
aperture during aestivation. The unusually shallow and shaded habitat from which
this sample was collected is most probably subject to desiccation in the dry season.
The specimens brought alive into the laboratory showed a persistent tendency to
crawl out of the water. Compared with Sudanese and Kenyan strains of B. sudanica
kept in this laboratory B. camerunensis is an exceptionally slow-growing species and
it has so far proved refractory to infection with strains of Schistosoma mansoni
from Egypt and Liberia.

Biomphalaria pfeifferi (Krauss)

Planorbis pfeifferi Krauss, 1848 : 83, pi. 5, fig. 7.

MATERIAL : Lake Wum, Bamenda Division, on sparse aquatic vegetation in 2-3
feet of water. Six specimens, yth November, 1963.

Shell (PL I, figs. 14-16) : light yellowish-brown, whorls without marked angulation,
well-developed spiral lines intersect fine growth lines to give a strong granular
microsculpture. The dimensions of the largest specimen in the sample were, maxi-
mum diameter 6-4 mm., umbilical diameter 1-8 mm., height 2-8 mm. The mean
ratio for the whole sample of maximum diameter/umbilical diameter was 3-66 and
of maximum diameter /height was 2-14.

Anatomy : the state of preservation of the sample was poor. The two largest
specimens (6-4 and 5-5 mm. diameter) were dissected and found to be fully mature.
The penis sheath is shorter than the preputium, the prostate has about 15 primary
diverticula, most of them with secondary branches. In contrast to B. camerunensis
the oviduct is strongly convoluted and the muciparous gland encloses more than half
the uterus. The receptaculum is club-shaped and the vesicle is about equal in
length to its duct. The pericardia of both specimens contained many trematode
metacercariae.

Radula : the teeth are larger than in B. camerunensis, there are seven tricuspid
laterals in each half-row, their mesocones are broad and spatulate and the marginal
ectocones are divided.

Biomphalaria pfeifferi has an almost universal distribution in the Ethiopian region
and its presence in Cameroon is to be expected.

Anisus coretus (de Blainville)

Le Coret, Coretus Adanson, 1757 : 7-10, pi. I, fig. 3.

Planorbis coretus Adanson, de Blainville, 1826 : 230.

non Planorbis coretus Adanson, Dautzenberg, 1890.

Coretus adansonii Gray, 1850 : 119, pi. 309, fig. 4.

Planorbis (Gyraulus) gibbonsi Nelson. Binder, 1957 : I2O> n§- I7-

3§§

C. A. WRIGHT

\ A ( \

FIGS. 1-3. Anisus coretus. i, Whole genital system. 2, Tip of male
copulatory organ. 3, Prostate.

FIG. 4. Biomphalaria camerunensis. Male copulatory organs and receptacula.

MATERIAL : Lake Barombi Kotto, Kumba Division. Ten specimens and
laboratory-bred material. I5th November, 1963.

Pamol Estate, Lobe, Kumba Division, in ditches, about 70 specimens. i5th
November, 1963.

Shell (PL II, figs. 4-6) : small, discoidal, flattened above, slightly concave beneath,
whorls 3-3^, rounded without angulation, sutures relatively deep, aperture rounded.
The shells are glossy with fine growth lines, many of those from Lobe have a very
fine spiral sculpture on the underside giving a wavy appearance to the growth lines.
The largest specimen seen was laboratory bred and measured 2-7 mm. maximum
diameter. The mean dimensions of five adult specimens from Lobe are 2-4 mm.
maximum diameter, 1-24 mm. umbilicus diameter and 0-68 mm. height.

Anatomy (Text-fig, i) : the male copulatory organ is small with the penis sheath
and preputium about equal in length and the preputium slightly greater in diameter

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 81

than the proximal dilatation of the sheath. The penis is usually a little longer than
the sheath, there is a small claw-like stylet at its tip and the opening of the vas
deferens is sub-terminal (Text-fig. 2) . The prostate (Text-fig. 3) consists of a glandular
part of the male duct with a single diverticulum. Examination by phase-contrast
microscopy shows that the wall of the sperm-duct consists of irregularly rounded
cells, the glandular prostatic part and the diverticulum are finely and densely
granular and the vas deferens appears to be composed of connective tissue with sparse
oval cells with well-defined nuclei. The seminal vesicle on the hermaphrodite duct
is coiled and has small projections on its surface. The vagina is short and proximally
dilated, the uterus is long, narrow and straight and the albumen gland is relatively
large. The receptaculum seminis is ovoid and slightly shorter than its slender duct.

Radula (Text-fig. 16) : teeth small and few in number, there are about eight
tricuspid laterals, a single intermediate in which the ectocone is divided and a single
five-cusped marginal in each half-row.

Anatomically this material does not differ significantly from Binder's (1958)
description of Gyraulus gibbonsi from the Ivory Coast. The terminal stylet of the
penis is similar in both forms and differs from the smooth cap-like structure of
Anisus misellus (Morelet) from Angola (Wright, 1963). The single prostatic diverti-
culum of the Ivory Coast material is very much longer than in specimens from the
Cameroons but this is probably a variable character in view of the occurrence of
occasional individuals with a single diverticulum in A . misellus, a species which usually
has a more normal prostate with several diverticula. Binder shows the seminal
vesicle to be convoluted but smooth in his specimens and in this respect the Cameroon
form resembles A. misellus which also has small projections from the surface of the
vesicle.

Binder referred his specimens to Gyraulus gibbonsi on the basis of an illustration by
Pilsbry & Bequaert (1927) . G. gibbonsi was described by Nelson (1878) from Zanzibar
but his illustration does not agree with his description in that it shows a shell com-
pletely flattened beneath and with a sharp basal angle while the description says that
the aperture is rounded. The whereabouts of the type-specimen are unknown but
Pilsbry & Bequaert examined material from Zanzibar and found that it agreed with
specimens from the Congo with a rounded aperture. I have also examined topotype
shell material and have found that it resembles the form from the Cameroons except
that it lacks any trace of spiral sculpture and reaches a much greater size at the 3^
whorl stage than does the West African form. There is, however, a very striking
resemblance between my specimens and the description and photographs of Adanson's
(1757) Le Coret, Coretus, published by Fischer-Piette (1942). Fischer-Piette pointed
out that Adanson's species had never been rediscovered and that Dautzenberg's
(1890) re-description of it from the type-locality at Podor, Senegal, certainly did not
agree with the original specimens recovered later. Gray's (1850) species Coretus
adansonii was based only on Adanson's original description as was the earlier
Planorbis coretus of de Blainville (1828). Adanson emphasized the small size of this
snail and reported copulation between individuals just over 3 mm. in diameter.

The relationships of A. coretus to other species are not yet readily determined.

82 C. A. WRIGHT

A. misellus is obviously close and it is possible that some of the many species of
small planorbids indifferently described by Germain (1907, 1909, 1911, 1917) from
West Africa and the Lake Chad region are no more than local variations, de
Azevedo et al, (1961) have recently described the anatomy of A. natalensis from
Mozambique as having a single prostatic diverticulum, no penial stylet and a terminal
opening to the vas deferens. These characters are in contrast to the normal prostate
and sclerotized terminal part of the penis in A . natalensis from Ethiopia and Trans-
vaal (Wright & Brown, 1962).

Gyraulus costulatus (Krauss)

Planorbis costulatus Krauss, 1848 : 83, pi. 5, fig. 8.

MATERIAL : Lake Wum, Bamenda Division. On dead leaves near the lake margin.
3 specimens. 7th November, 1963.

Shell (PL II, figs. 7-9) : small, discoidal, pale yellowish brown, strongly and
regularly ribbed with a well-marked equatorial carination. The largest specimen
was broken but probably had 3-3 1 whorls ; an individual with 3 whorls measured
3-2 mm. diameter, 0-8 mm. umbilical diameter and 1-08 mm. high.

Anatomy : one specimen was aphallic but the anatomy of the other two corres-
ponded well with that described for this species from Angola (Wright, 1963). The
penis papilla noted by de Azevedo et al. (1961) was observed but the Cameroon
specimens had the penis relatively shorter with the stylet not reaching to the end of
the sheath.

Radula (Text-fig. 18) : teeth small, 9-10 tricuspid laterals and about 6 five cusped
marginals in each half-row. The cusps of the laterals are a little broader than those
in Angolan specimens.

This species was probably more abundant than the number of specimens collected
suggests but the samples obtained from Lake Wum were, for various reasons, in-
adequate. The largest of the three specimens was grossly infected with rediae of an
unidentified trematode. This was probably the same species of parasite as the
metacercariae found in the pericardia of the other two individuals and the Biom-
phalaria pfeifferi and SegmentorUs angustus from the same locality.

Segmentorbis angustus (Jickeli)

Segmentina angusta Jickeli, 187/1 : 22O> pi- 7> fi§- 24-
Segmentorbis angustus (Jickeli), Mandahl-Barth, 1954 : 9^-

MATERIAL : Lake Wum, Bamenda Division. On dead leaves and emergent grasses.
About 50 specimens, also laboratory-bred material. 7th November, 1963.

Shell (PI. II, figs. 1-3) : lenticular, flattened beneath, relatively sharp basal angle,
translucent, pale yellowish-brown, wild specimens often thickly crusted with dark
brown deposits, umbilicus narrow and very deep. There are usually two, rarely
three sets of lamellae in the body whorl, the basal is well-developed, straight, about
half the width of the whorl, the inner curves strongly over the parietal wall, the

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 83

outer is usually weak, often sub-divided and sometimes absent while the dorsal is
rarely present (Text-figs. 5 and 6). The shell is smooth and shiny with a faint trace
of spiral sculpture on the underside, visible only under high magnification.

Anatomy (Text-fig. 7) : penis-sheath about two-thirds the length and a quarter the
width of the preputium, with a single, large, thin-walled flagellum attached to the
proximal end of the sheath. There is a separate prostatic duct from which arise
about fifteen unbranched diverticula. The seminal vesicle is widely dilated and
convoluted with small surface protruberances. There is a slight dorsal dilatation
of the vagina, the uterus is long and straight and the receptaculum seminis is clavate,
about equal in length to its duct. Several specimens had heavy infections of trema-
tode metacercariae in their pericardia.

Radula (Text-fig. 17) : the teeth are similar to those of Anisus coretus but they are
smaller and more numerous. The first 7-9 laterals are tricuspid, the endocone is
subdivided in the next six or seven and from about tooth fifteen outward the ectocone
is also divided giving six to eight typical 5-6 cusped marginals.

5. angustus was originally described from Ethiopia and is known from many
places in East and South Africa. A single small specimen was recorded from
Kikondja in the Congo by Pilsbry & Bequaert (1927). Their illustration of the shell
shows a greater number of basal lamellae (4) than are found in specimens of com-
parable size in the present material and the outer lamellae are even more degenerate
in the Congo specimen. Mandahl-Barth (1954) notes that there is usually only a
single set of septa but occasionally up to five sets are seen in this species in Uganda.
He also mentions specimens of Segmentorbis with even larger numbers of septa in
which the other shell characters do not differ from normal S. angustus. The form of
the receptaculum seminis in the Cameroon specimens differs from that described by
Mandahl-Barth (1954) for Uganda material and by de Azevedo et al. (1961) for
specimens from Mozambique. Both of these authors show the vesicle to be small
and spherical with a diameter only about a quarter the length of the duct. Even
young specimens from Lake Wum have an elongate-ovoid receptaculum about equal
in length to its duct and this form is similar to that in Ethiopian specimens (Brown,
1965).

Segmentorbis kanisaensis (Preston)

Segmentina kanisaensis Preston, 1914 : 265, pi. 18, figs. 17-19.
Segmentorbis (Carinorbis) kanisaensis ; Mandahl-Barth, 1954 : 98, fig. 45.

MATERIAL : Lake Barombi Mbo, Kumba Division. One specimen.

Shell : lenticular, flattened beneath, sharp basal angle ; translucent, yellowish-
brown, fine growth lines and slight spiral microsculpture on the underside ; a single
set of well-developed lamellae (Text-figs. 8 and 9), the basal about two-thirds the
width of the whorl, the inner strong, curved, the outer long, oblique and the dorsal
a pronounced dot-like structure. Dimensions of the single specimen were diameter
2 mm., umbilicus 0-5 mm., height 0-6 mm.

Anatomy : the specimen was immature but dissection showed the characteristic
lack of a flagellum on the male copulatory organ.

C. A. WRIGHT

FIGS. 5-7. Segmentorbis angustus. 5, Dorsal view of shell. 6, Ventral
view of shell. 7, Whole genital system.

FIGS. 8, 9. Segmentorbis kanisaensis. 8, Dorsal view of shell. 9, Ventral view of shell.

Segmentorbis snails were relatively common on dead leaves in Lake Barombi Mbo
near the mouth of the entering stream on the north-west shore. Unfortunately all

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 85

of the wild specimens were lost and the single individual reported here was later
found in a tank in the laboratory containing leaf-litter from the lake. This species
is common in low-lying areas of West Africa from the Gambia to Angola and is
also found in East and South Africa.

Bulinus rohlfsi (Clessin)

Physa rohlfsi Clessin, 1886 : 349, pi. 49, fig. 7.

Bulinus truncatus rohlfsi ; Mandahl-Barth, I957(b) : 29, pi. 20, fig. 18.

MATERIAL : Lake Barombi Mbo, Kumba Division. About 100 specimens received
from Dr. B. O. L. Duke, 8th January, 1957 ; 20 specimens and laboratory-bred
material, I4th November, 1963.

Lake Barombi Kotto, Kumba Division. About 40 specimens and laboratory-bred
material, 3ist October and I5th November, 1963.

Shell (PL I, figs. 7-10) : light yellowish-brown, spire short, no marked ribbing or
other micro-sculpture ; columella more or less straight, columellar margin reflexed,
closing the umbilicus. Adult specimens from Lake Barombi Kotto usually have the
aperture margin reflexed giving to the shell a bell-mouth appearance ; this distortion
occasionally occurs in younger individuals and, if followed by a period of normal
growth, results in the formation of a marked ridge on the body whorl. Similar
reflexion of the aperture margin has not been seen in wild specimens from Barombi
Mbo but it occasionally occurs in laboratory colonies from the Mbo stock. The
maximum size in the Kotto sample is length 11-2 mm., width 8-8 mm. and aperture
length 8-9 mm. From Mbo the largest specimen is 7-7 mm. long, 5-1 mm. wide and
has an aperture length of 5-1 mm. This disparity in maximum size between the two
populations has been maintained in laboratory colonies bred from the stocks. The
colony from Kotto is also remarkable for the number of distorted shell-forms which
it produces. The mean ratio shell-length/aperture length for adult specimens of both
populations is 1-2.

Anatomy : all specimens dissected from both populations were aphallic ; in some
there is a trace of sperm-duct and a vestigial prostate but in the majority there is no
development of the male system. Specimens from Mbo have the uterine glands fully
developed and apparently functional at shell-length 4-0 mm. while those from Kotto
are less well-developed at 6 mm. The mantle in specimens from Kotto is pale grey
with black spots and patches while in the Mbo samples the markings vary, as they do
in the same species from Angola, from almost unmarked to strongly patterned.

Radula (Text-fig. 13) : teeth smaller than in B. truncatus, 3-5 tricuspid laterals,
all with slightly arrowhead-shaped mesocones flanked by fine interstitial cusps,
13-18 marginals in which the mesocone is not so clearly dominant as it is in the next
species.

Bulinus rohlfsi is the oldest name of a considerable number of species and varieties
described from Lake Chad. Mandahl-Barth (i957b) treats rohlfsi as a sub-species of
B. truncatus and there is no doubt about the close relationship of the two species.
However, the sub-species concept has little real meaning in the Basommatophora

86 C. A. WRIGHT

and I prefer to regard rohlfsi as a distinct species. The large number of names given
to this species in Lake Chad is some indication of the great variability of its shell
form. The distinctions observed here between the populations from the two Kumba
lakes are further evidence of this polymorphism. In both lakes B. rohlfsi is acting
as intermediate host for Schistosoma haematobium.

Bulinus tropicus (Krauss)

Physa tropica Krauss, 1848 :

Bulinus (Bulinus) hemprichii depressus Haas, 1936 : 28, pi. I, fig. 15.

Bulinus tropicus tropicus (in part) ; Mandahl-Barth, I95yb : 19, pi. 10, fig. 4.

MATERIAL : Babungo, N'dop Plain, Bamenda Division. On aquatic plants in a
slow-flowing stream about i mile east of the village. 20 specimens, also laboratory
bred material. 6th November, 1963.

Shell (PI. I, figs, ii and 12) : thin, translucent, yellowish-brown, spire flat, aper-
ture wide, columella slightly twisted ; early whorls ribbed, body-whorl smooth with
fine growth-lines. Dimensions of the largest specimen were — length 8-1 mm.,
width 6-9 mm. and aperture length 7-4 mm. In the laboratory shell-lengths in
excess of 12 mm. have been achieved by mature individuals. The mean ratio
shell-length/aperture-length for a series of juveniles is 1-03, a clear indication of the
flatness of the spire. However, laboratory-bred specimens develop a prominent
spire even in the first generation.

Anatomy : penis sheath about twice the length of the preputium, its proximal
dilatation is a little wider than the distal part ; epiphallus short and without dilata-
tion. The largest wild specimen (8-1 mm.) was protandrously mature with apparent-
ly functional copulatory organ and prostate but poorly differentiated uterine glands.
The remainder of the sample were all juvenile but in all there were at least traces of
rudimentary male genitalia and no potentially aphallic individuals were seen. The
mantle is light grey with sparse, well-defined black spots and patches.

Radula (Text-fig. 12) : teeth larger than in B. rohlfsi, 6-7 tricuspid laterals with
sub-division of the endocone occurring in the seventh or eighth tooth ; lateral
mesocones slightly arrow-head shaped, usually flanked by lanceolate interstitial
cusps, inner edge of the endocones frequently corrugated. In the 20-22 marginals
the mesocones remain undivided and are very prominent.

Wild specimens of this form closely resemble Bulinus depressus, described as a
sub-species of " B. hemprichii " from a canal near Lake Bangweulu in Zambia. In
an earlier paper (Wright, 1957) I suggested that B. depressus was no more than a
juvenile of B. globosus but, in a private communication, Dr. Mandahl-Barth drew my
attention to certain features of depressus which indicate that it is definitely not related
to the B. africanus complex and he (i957b) included it in the synonomy of B. tropicus
tropicus. Examination of the immature wild material from Babungo left me in
some doubt as to whether it should be assigned to the tropicus or truncatus species
groups. However, electrophoresis of the egg-proteins of the one living specimen
brought back to London demonstrated without doubt that this form belongs to the

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 87

B. tropicus complex (Wright & Ross, in press). This record is probably near to the
extreme north-west limits of the range of this species group for I am unable to agree
with Mandahl-Barth on the inclusion of B. guernei from West Africa in the tropicus
group ; I have examined B. guernei from three of the four localities mentioned by
Mandahl-Barth (ig^jb) and I have found a high proportion of aphallic individuals
and arrowhead shaped lateral mesocones, Smithers (1956) has also reported wild-
caught specimens infected with Schistosoma haematobium. These three characters
all indicate an affinity with the truncatus group and no other species with tropicus
characteristics has been reported from West Africa. It is possible that the B.
strigosus reported by Gaud (1955) from Bangui is B. tropicus.

Bulinus forskali (Ehrenberg)
Isidora Forskalii Ehrenberg, 1831 : 20.

MATERIAL : Ditches and streams on Pamol Estate, Lobe, Kumba Division.
36 specimens. 25th October, 1963.

Shell (PI. I, figs. 5 and 6) : strongly turretted with well-marked shoulders and ribs
on all whorls, distinct small spines on the shoulder of the third whorl where the ribs
intersect the angle. Mean dimensions in millimeters of 14 adults (maxima in
brackets) were length 5-4 (7-2), width 2-7 (3-2) and aperture length 2-7 (3-0).

Anatomy : male copulatory organ small and slender, penis sheath /preputium ratio
about 3/2.

Radula (Text-fig. 15) : similar to that described for this species from Angola but
few malformations present in this population ; 5 or 6 tricuspid laterals and up to
eighteen marginals.

B. forskali is probably the most widely distributed and variable species of fresh-
water snail in Africa and its presence in Cameroon was to be expected. It was recorded
by Zahra (1953) from the stream flowing into the lake past Barombi Mbo village but
it was not found there during the present visit despite repeated examination of the
area.

Bulinus camerunensis Mandahl-Barth

Bulinus camerunensis Mandahl-Barth, iQ57b : 31, pi. 22, fig. 21.

MATERIAL : Lake Barombi Kotto, Kumba Division. About 150 specimens,
also laboratory-bred material. 3ist October and i5th November, 1963.

Shell (PL I, figs. 1-4) : high-spired, translucent, yellowish-white, most specimens
show some quite marked ribbing and in a few there is a weak shoulder on the second
and third whorls ; about half the shells examined show irregular spiral lines giving
patches of reticulate sculpture. The mean dimensions in millimeters (maxima in
brackets) for a sample of 50 wild adults are length 5-4 (6-9), breadth 2-9 (3-5) and aper-
ture length 3-2 (4-0). Larger specimens, up to 11-5 mm. high have been bred in the
laboratory. The mean value and range of the ratio shell-length/aperture-length is
1-65 (1-47-1-94). In only three out of the 50 wild snails measured was the breadth

88 C. A. WRIGHT

of the shell greater than the aperture length. This is in contrast to the dimensions
given in the original description.

Anatomy : male copulatory organ small and slender, sheath usually longer than
the preputium and its proximal dilatation about equal in diameter to the distal part.
The relative lengths of the two parts of the copulatory organ vary widely, the mean
ratio of sheath-length to preputium is 1-43 but it ranges from about i-o-2-o. The
prostate is relatively small in all of the fully adult specimens examined. Mantle
markings in wild specimens are irregular and somewhat diffuse black patches on a
grey ground but laboratory-bred individuals have the mantle almost uniformly black.

Radula (Text-fig. 14) : teeth similar to those of B. forskali but slightly smaller and
nearly all of the lateral mesocones are flanked by fine interstitial cusps. There are
five or six laterals as in B. forskali but the number of marginals is greater (up to
twenty-four) in B. camerunensis.

In his original description of this species Mandahl-Barth mentioned the possibility
that B. camerunensis was no more than a local form of B. forskali but that it differed
from that species by its smaller radula teeth. The contrast in shell-characters
between B. camerunensis and the form of B. forskali cc-llected a few miles away at
Lobe is well-marked and the present species shows greater similarity to B. beccarii
(Paladilhe) from Aden (Wright, I963a) than to any other member of the forskali
group. In the hope of finding further foci of this species six other crater lakes were
visited but without success. Repeated attempts to infect B. camerunensis with the
West Cameroon strain of Schistosoma haematobium have failed.

Family LYMNAEIDAE
Lymnaea natalensis Krauss

Limnaeus natalensis Krauss, 1848 : 85, pi. 5, fig. 15.

MATERIAL : Babungo, N'dop Plain, Bamenda Division. On aquatic vegetation
in a slow-flowing stream about I mile east of the village. 12 specimens. 6th Novem-
ber, 1963.

Lake Wum, Bamenda Division. 6 specimens, yth November, 1963.

Shell (PI. I, fig. 13) : thin, shiny, elongate-ovoid, spire short and sharply pointed ;
the only ornament is fine growth lines. Mean dimensions in millimeters (maxima
in brackets) of four adult specimens from Babungo are length 8-4 (9'i), breadth
4-6 (5-6) and aperture length 6-4 (7-0).

Anatomy : penis-sheath approximately twice the length of the preputium,
receptaculum seminis duct long with the receptaculum lying on the left side of the
body. Two of the four adult specimens from Babungo had the accessory genital
glands completely destroyed by an unidentified larval trematode but the other two
had the characteristic distal dilatation of the prostate.

Radula (Text-fig. 19) : eight to ten tricuspid laterals in which the endocones and
mesocones are fused together for over half their length ; fifteen to twenty marginals,
transition from the laterals occurring by sub-division of the ectocone which almost

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON

FIGS. 10-21. Radula Teeth. 10. Biomphalaria camerunensis. n. Biomphalaria pfeifferi.
12. Bulinus tropicus. 13. Bulinus vohlfsi. 14. Bulinus camerunensis. 15. Bulinus
forskali. 16. Anisus coretus. 17. Segmentorbis angustus. 18. Gyraulus costulatus.
19. Lymnaea natalensis. 20. Lanistes libycus. 21. Potadoma freethi .

go C. A. WRIGHT

disappears in the outer teeth which have only four or five cusps derived from the
endo- and mesocones.

L. natalensis is widely distributed throughout Africa. The only feature of interest
in the present material is the relatively small size at which maturity is reached.

Family ANCYLIDAE
FERRISSIA sp.

MATERIAL : Ditches on Pamol Estate, Lobe, Kumba Division. 5 specimens.
i5th November, 1963.

Shell (PI. II, figs. 10 and n) : small, rectangularly ovoid, light yellow, apex
striate, displaced to the right, smoothly rounded. The margin of all five specimens
was of soft periostracum causing considerable variation in shell shape. The mean
dimensions in millimeters (maxima in brackets) of four entire specimens are length
178 (1-92), width 1-2 (1-2), height 0-57 (0-64).

No preserved specimens were available for anatomical study. Small ancylids
were also found in Lake Wum and at Babungo, both in Bamenda Division but the
material from both localities was lost. The small amount of material and lack of
anatomical data do not permit an assessment of the affinities of this species but the
proportions of the shell are similar to those of F. eburnensis Binder, 1957, from the
Ivory Coast. Population variations in the outline of the shell from uniformly
oval to rectangularly ovoid were noted by Binder.

Family AMPULLARIIDAE
Lanistes libycus (Morelet)

Ampullaria libyca Morelet, 1848 : 28, pi. 3, fig. 9.
Ampullaria bernardiana Morelet, 1860 : 190.

MATERIAL : Lake Barombi Mbo, Kumba Division. On stones and sandy bottom
near canoe beach on north-east side of the lake. 40 specimens. 24th October, 1963.

Ditches on Pamol Estate, Lobe, Kumba Division. 12 specimens. I5th Novem-
ber, 1963.

Shell (PI. Ill, figs. 9 and 10) : large, globose, low-spired, apex often eroded ;
whorls with well-marked shoulder angle, distinctly flattened below the suture,
moderate carination around the umbilicus. Specimens from Lake Barombi Mbo
have the shell smooth with strong spiral bands of colour which appear brownish-red
externally and purple internally, those from Lobe have the colour bands less well-
marked but have strong spiral ridges of periostracum which persist on adult shells ;
these ridges are present on some juveniles from the lake population. The largest
specimen collected (from the lake) measured 27 mm. in height, 28 mm. in width and
had an aperture length of 18 mm.

Radula (Text-fig. 20) : central tooth massive, median cusp flanked usually by two,
sometimes three, smaller cusps all of which may be fused into a single transversely
elongate blade ; lateral four-cusped and the two marginals bi-cuspid.

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 91

The type-locality for L. libycus is in Gaboon and the species has a wide distribution
in West Africa. It has been reported from many places in the Cameroons by previous
workers. The treatment of L. bernardianus as a variety of L. libycus was retained by
d'Ailly (1896) because the large number of samples which he examined from Cameroon
showed no intermediates between the two forms. However, the characteristic
spiral ridges of bernardianus are confined to the periostracum and are easily rubbed
off. This distinguishing character is therefore dependent on the nature of the
substratum in the habitat. Binder (1957) has put forward a strong claim for the
retention of the name libycus despite the recently demonstrated prior claim of
L. intortus (Lamarck). In view of the confusion likely to be created by the use of
Lamarck's name I am in entire agreement with Binder's argument.

Family BITHYNIIDAE
GABBIA sp.

MATERIAL : Lake Wum, Bamenda Division, i specimen, yth November, 1963.

Shell (PL II, fig. 12) : pale yellowish-brown with three rounded whorls, a very
faint spiral microsculpture intersects the fine growth lines but the general appearance
of the shell is smooth and shiny. The umbilicus is narrowly open and the spiral
nucleus of the operculum occupies about half its total width. The single specimen
measures 2-32 mm. in height and width and has an aperture height 1-52 mm. and
aperture width of 1-36 mm.

The genus Gabbia is mainly East and Central African in distribution, the only
West African records being Frauenfeld's B. africana and Binder's (1955) Bithynia
(Gabbia) tournieri from the Ivory Coast. Several species have been recorded from
the Lake Chad region but the affinities of the present specimen appear to be nearest
to G. parva Mandahl-Barth, 1954.

Family THIARIIDAE
Potadoma freethi (Gray)

M elania freethi Gray, 1831 : n.

Melania nigritina Morelet, 1848 : 345; von Martens, 1877 : 270; von Martens, 1891 : 33;

d'Ailly, 1896 : 121.

Melania (Nigrilella) nigritina ; Boettger, 1905 : 181.
Thiara (Melanoides) dykei Spence, 1925 : 249.
Potadoma graptoconus Pilsbry & Bequaert, 1927 : 276, fig. 46a and pi. XX, figs. 10 and n.

MATERIAL : Stream on Bai Rubber Estate, Kumba Division. 10 specimens.
25th October, 1963.

Stream on Pamol Estate, Lobe, Kumba Division. 10 specimens. 25th October,
1963.

Stream near Barombi Kotto Stranger Town, Kumba Division. 18 specimens.
I5th November 1963.

92 C. A. WRIGHT

Stream through Barombi Mbo village, Kumba Division. 20 specimens. i6th
November 1964.

Shell (PI. Ill, figs. 1-8) : large, solid, turreted, adults usually black and decollate,
whorls flattened, suture shallow, aperture pear-shaped, pointed above, columella
gently curved outward resulting in a slight expansion of the aperture at the junction
of the basal and columellar margins. There is a well-marked spiral sculpture and
on the later whorls fine growth lines are more accentuated and intersect the spiral
lines to give a granular pattern. Young specimens have a marked basal angle on
the body whorl (PI. HI, figs. 7 and 8) but this almost disappears in the adults. The
usual black colour of adult shells results from a fine covering of silt which can be
removed by ultrasonic cleaning to show the basic greenish-brown colour of the young
shell with at least one dark purple-brown band. Specimens from habitats with
sandy bottoms are often found in this clean condition.

Radula (Text-fig. 21) : central tooth complex with five small cusps on its anterior
edge. The single lateral on each side is assymetrically four-cusped and the two
marginals are tricuspid.

The genus Potadoma is typically West African and more or less confined to the
forested areas. The type locality for P. freethi is the island of Fernando Poo off
the Cameroon coast, Melania nigritina was described from Gaboon, P. dykei from
near Victoria in Cameroon and P. graptoconus from the Congo. All authors are in
agreement concerning the identity of P. freethi and Morelet's nigritina. P. dykei
was described without reference to freethi ; a specimen of dykei identified by Spence
(PI. Ill, figs. 2 and 5) is in the collection of the British Museum (Natural History) and
apart from being slightly more obese and having more well-marked cords around the
columella it is not possible to separate it from large specimens of freethi. P. grapto-
conus was distinguished by Pilsbry & Bequaert because of its well-marked spiral
sculpture without the granulation said to be characteristic of freethi. However,
examination of numbers of specimens from West Cameroon indicates that this
granulation is not always present and is somewhat irregular on the holotype of
freethi (PI. Ill, fig. 4).

P. freethi was always found in gently moving water and although it is present in
the stream flowing into Lake Barombi Mbo live specimens were not found in the lake
itself. This species is common as a sub-fossil in the area of Lake Barombi Kotto
and near Bekondo on the route to Lake Soden.

DISCUSSION

The collections made during this expedition fall readily into two groups, those from
the low-lying forested area of Kumba Division and those from open savannah
country at 4-5,000 feet in Bamenda Division. No species has been found which is
common to both areas and although the search in Mamfe Division which intervenes
between Kumba and Bamenda was superficial no freshwater snails of any kind were
found. This suggests that an effective barrier exists between the two areas. The
broad differences between the two parts of the snail fauna are those typical of altitude

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 93

and correspond to the differences between the coastal plain and southern plateau
regions of Angola (Wright, 1963!)). The low-lying areas are characterized by an
abundance of prosobranchs, the presence of Anisus and members of the Bulinus
forskali group and B. rohlfsi while the highlands have Biomphalaria, Gyraulus
costulatus and Bulinus tropicus. The presence of Segmentorbis kanisaensis in Lake
Barombi Mbo is in keeping with its occurrence in the coastal plain of Angola and at
low altitudes in the Ivory Coast and Gambia while S. angustus in Lake Wum corres-
ponds with its distribution in higher crater lakes in Uganda. If the affinities of the
species of Gabbia in the same lake have been correctly assessed it also points to a
similarity with the Uganda fauna. Geze (1943) mentions the occurrence of a number
of East African elements in the insect fauna of the mountains in Cameroon and
attributes this to a more extensive temperate connection with the east along the
highland chain during the pluvial periods. Sram (1955) has even gone so far as to
suggest that there is evidence of glaciation on Mount Cameroon and Mannengouba
during this period. The type locality of Biomphalaria camerunensis on Mount
Cameroon suggests that this is a relic population now isolated from the more general
highland areas by the intervening tropical rain forest.

In Kumba Division the difference between the snail faunas of lakes Barombi Mbo
and Barombi Kotto is interesting. The only species which they have in common is
Bulinus rohlfsi, the two populations of which have distinctive characteristics.
Trewavas (1962) has recently shown that the two lakes have no species of fish in
common and that all but one of eight species in Barombi Mbo are probably endemic
while Barombi Kotto has an endemic genus, two endemic species and one endemic
sub-species out of a total of seven species. Bulinus camerunensis appears to be
endemic to Barombi Kotto but the other snail species in both lakes have wide distribu-
tions in West Africa. The effluent from Barombi Kotto passes into the Meme
River near the mouth of which lies Lobe. Anisus coretus is common to both the
lake and the streams in the Lobe area while B. camerunensis in the lake, although
closely related to B. forskali, is clearly distinct from the local form of that species
at Lobe. Differentiation of the two populations of B. rohlfsi may be the result of the
" founder principle " and does not necessarily imply any prolonged separation.

SCHISTOSOMIASIS IN WEST CAMEROON

Zahra's (1953) original report has remained virtually the only source of information
on schistosomiasis in West Cameroon, de Azevedo (1958) included these data in a
general report covering what was at that time the whole British Cameroons but did
not add anything further. One of the objectives of the present expedition was to
confirm the identity of the snails responsible for transmission of the parasite in the
known foci and to investigate their wider distribution in the territory.

Wild specimens of Bulinus rohlfsi from Barombi Kotto were found to be shedding
cercariae of Schistosoma haematobium (confirmed by exposure of hamsters with
subsequent recovery of adult worms). Laboratory bred snails of this species from
both of the Barombi lakes have been successfully infected by miracidia hatched
from urine samples from Barombi Mbo. Bulinus camerunensis from Barombi Kotto

94 C. A. WRIGHT

was not found naturally infected nor has it been successfully exposed to infection in
the laboratory. Laboratory bred specimens of Bulinus tropicus from the N'dop
plain were not susceptible to miracidia from Barombi Mbo. This is to be expected
because so far no member of the B. tropicus complex has been demonstrated to act
as an intermediate host for 5. haematobium. B. rohlfsi has not been found in any
of the other places investigated in West Cameroon so there is little chance of the
disease spreading unless the snails become established in other areas.

The habitat for B. rohlfsi in both lakes is limited to shallow water with reasonable
sun-exposure and the snails are found on dead leaves rather than on aquatic vegeta-
tion. It is remarkable that the focus in Lake Barombi Mbo has remained confined
to the small area on the north-west side of the lake near to the beaches where fishing
canoes from the village are drawn up since there is ample opportunity for distribution
of the snails both on the canoes and on fish traps which are set daily in all suitable
places around the lake margin. It is probable that shading of the water's edge by
fringing vegetation is the most important factor in preventing the colonization of
new areas and any major clearance of the marginal bush should be viewed with
concern. It is probable that transmission in both lakes is partly seasonal ; to judge
from conditions in October and November it seems likely that maximum snail
population densities occur in January-February and, allowing for development of
the larval schistosomes there is probably a peak of transmission in February-March.

An interesting aspect of these lake foci is that schistosomiasis is almost confined
to members of the Barombi tribe who fish both lakes. The Barombis appear to be
the only local people who do not have a profound superstitious fear of the lakes and
contact with the water by people of other tribes is relatively infrequent. Lakes such
as Soden and Ejaghem are known to contain fish but this is not a sufficient attraction
to draw people to them regularly. On Lake Soden there are not even any canoes
and only poor tracks to the water's edge. The few cases of urinary schistosomiasis
encountered outside the Kumba area are usually from Nigeria or the East but some
are found in people who have lived in Barombi Kotto Stranger Town and visited the
lake for washing.

Dissemination of Bulinus rohlfsi from the Barombi lakes might occur if develop-
ment of fisheries in other lakes is undertaken using Barombi fish for stocking. There
is also a slight possibility that B. rohlfsi from Barombi Kotto might gain access via
the Meme river to the Lobe area where bush clearance has provided potentially
good snail habitats in the streams and ditches. It seems that the West Cameroon
foci of urinary schistosomiasis are effectively closed with little opportunity for
spread and not much chance of importation of new strains. The strain in the
Barombi foci is probably derived from the B. rohlfsi-borne form in Nigeria (Cowper,
1963). Snail control by molluscicides or habitat modification would not be practical
in the lakes since the danger of damaging the fish population would be too great.
The confined nature of the foci will probably lend themselves to control by drug
treatment when suitable compounds are available.

There is little information available concerning Schistosoma mansoni in West
Cameroon. It is perhaps significant that Gaud's (1955) distribution map of this

FRESHWATER GASTROPOD MOLLUSCS OF W. CAMEROON 95

parasite shows the edge of a lightly endemic zone in the east coinciding exactly with
the border between East and West Cameroon in the area of the N'dop plain.
Enquiries in the area near Bambalang revealed that dysentery accompanied by
passing of blood is common in the area but time did not permit a survey to be carried
out. Biomphalaria camerunensis is a possible host for S. mansoni but limited
infection experiments with laboratory-bred specimens have so far failed. Both an
Egyptian and a Liberian strain of parasite have been tested. There is at present
no evidence for the existence of 5. mansoni in the Wum area despite the presence of
the potential intermediate host Biomphalaria pfeifferi in Lake Wum.

In conclusion brief mention must be made of two other trematode diseases, para-
gonimiasis (lung-fluke) in man and liver-fluke in cattle. Zahra (1952) reported a
widespread incidence of infection with Pamgonimus in Victoria and Kumba Divisions.
This distribution coincides with that of the snail intermediate host, Potadoma
freethi, which is common in small streams throughout the area. The incidence of
Paragonimus is said to be particularly high among the Bakossi people in the region
of Mount Kupe up to a height of 5,000 feet. No published reports of the occurrence
of Fasciola gigantica in cattle in West Cameroon have been seen. Inquiries through
the sanitary inspectors responsible for meat inspection suggest that the few cases
seen are in cattle brought from outside the territory. However, around Banso and
Ndu (6-7,000 feet) the parasite is scarcely ever seen while in the N'dop plain where
the intermediate host Lymnaea natalensis is found it is said to be of slightly more
frequent occurrence. Most of the grazing area in Bamenda Division appears to be
above the altitude limit for L. natalensis and examination of apparently ideal habitats
near Oku failed to yield any snails.

ALPHABETICAL LIST OF COLLECTING PLACES

This list includes the major places visited but not a large number of minor streams
and pools in which nothing was found.

1. Babungo, N'dop Plain, Bamenda Division. 6°O2' N., io°i2'E. Altitude
about 4,000 feet. Slow-moving, weed-choked stream about one mile east of the
village. 6th November, 1963. Bulinus tropicus, Lymnaea natalensis and unidenti-
fied ancylid.

2. Bafeng (Lake), near falls on Mencham River, about 12 miles south of Wum on
Bafut road, Bamenda Division. 6°2o' N., io°02' E. Altitude unknown, probably
between 5,000 and 6,000 feet. Small, very steep-sided crater lake with dense bush
within the crater but surrounded by open grassland. 7th November, 1963. No
snails found.

3. Bambalang, N'dop Plain, Bamenda Division. 5°52' N., io°25' E. Altitude
about 4,000 feet. Stagnant, shallow, marshy pools, heavily shaded by palm scrub.
Thick, black mud bottom and mixed emergent vegetation. 6th November, 1963.
Biomphalaria camerunensis.

4. Bambuluwe (Lake), south of Bamenda town, Bamenda Division. 5°5o' N.,
io°io' E. Altitude probably over 6,000 feet. Moderate-sized crater lake, slightly

96 C. A. WRIGHT

irregular in shape, dense surrounding bush except near outlet of effluent stream where
grassland runs down to lake shore. Few leeches and insect larvae seen. 4th Novem-
ber, 1963. No snails found.

5. Barombi Kotto (Lake), Kumba Division. 4°2g' N., 9°2o' E. Altitude about
350 feet. Large crater lake with less steep margins than most and a volcanic island
in the centre on which stands the village. No snails found on the lake margin at
canoe beach but numerous specimens collected from dead leaves and debris on
gently sloping beaches of the island. 3ist October and I5th November, 1963.
Bulinus rohlfsi, B. camerunensis and Anisus coretus.

6. Barombi Mbo (Lake), Kumba Division. 4°42'N., g°28' E. Altitude about
1,000 feet. Large crater lake, about two miles in diameter, very deep except on
north-west side where entering stream has deposited a shelf of silt and gravel.
Snails confined to this part of the lake. Visited six times in October and November,
1963. Bulinus rohlfsi, Segmentorbis kanisaensis, Lanistes libycus and Potadoma
freethi (in entering stream).

7. Ejaghem (Lake), Mamfe Division. 5°45' N., 8°56' E. Altitude about
600 feet. Moderately large lake, shallow sloping surround with thick bush which
has been cleared in patches. Natural(?) fish population but no snails found. 9th
November, 1963.

8. Lobe, Pamol Estate, Kumba Division. 4°37' N., 9°oi' E. Altitude, near
sea level. Oil palm plantation on which most of the bush has been cleared and some
water-management of small streams undertaken. 25th October and I5th November,
1963. Bulinus forskali, Anisus coretus, Ferrissia sp., Lanistes libycus and Potadoma
freethi.

9. Soden (Lake), Kumba Division. 4°45' N., 9°i6' E. Altitude, 1,500 feet.
Moderately large, isolated crater lake with very steep, densely forested internal slope,
difficult of access, very little human contact, no canoes, natural fish population.
28th October, 1963. No snails found.

10. Tiko, Victoria Division. 4°5' N., 9°2o' E. Altitude, near sea level. Heavily
polluted stream flowing through the edge of the town, visited at several points after
hearing reports of urinary schistosomiasis in the area. Reports later proved to be
without foundation. i2th November, 1963. No snails found.

11. Wum (Lake), Bamenda Division. 6°28' N., io°02' E. Altitude about
5,000 feet. Moderate sized lake, gently sloping bank with patches of shallow mar-
ginal water and some aquatic vegetation. Lake open, surrounded by savannah,
margins unshaded, yth November, 1963. Biomphalaria pfeifferi, Gymulus costu-
latus, Segmentorbis angustus, Lymnaea natalensis, Gabbia sp. and unidentified ancylid.

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PLATE i

FIGS. 1-3. Bulinus camerunensis, Lake Barombi Kotto, x 8.

FIG. 4. Bulinus camerunensis, laboratory bred, x 8.

FIGS. 5, 6. Bulinus forskali, Lobe, x 8.

FIGS. 7, 8. Bulinus rohlfsi, Lake Barombi Mbo, x 2-5.

FIGS. 9, 10. Bulinus rohlfsi, Lake Barombi Kotto, x 2-5.

FIGS, n, 12. Bulinus tropicus, Babungo, N'dop plain, x 2-5.

FIG. 13. Lymnaea natalensis, Lake Wum, X 2-5.

FIGS. 14-16. Biomphalaria pfeifferi, Lake Wum, x 2-5.

FIGS. 17-23. Biomphalaria camerunensis, Bambalang, N'dop Plain, x 2-5.

Bull. B.M. (N.H.) Zoo/. 13, 3

PLATE i

PLATE 2

FIGS. 1-3. Scgmentorbis angustus, Lake \Yurn, x 14.
FIGS. 4-6. Anisus coretus, Lobe, x 14.
FIGS. 7-9. Gyraiilus costulatus, Lake Wum, x 14.
FIGS. 10, ii. Ferrissia sp., Lobe, x 14.
FIG. 12. Gabbia sp. Lake Wum, x 14.

Bull. B.M. (N.H.) Zoo/. 13, 3

PLATE 2

PLATE 3

Figs. 1-3 & 7-10 x 1-6, Figs. 4-6 x 6-5.
FIG. i. Potadoma freethi, Fernando Poo, (Holotype).
FIG. 2. Potadoma freethi, (P. dykei Spence).
FIG. 3. Potadoma freethi, Barombi Mbo.
FIG. 4. Microsculpture of specimen in Fig. i.
FIG. 5. Microsculpture of specimen in Fig. 2.
FIG. 6. Microsculpture of specimen in Fig. 3.
FIGS. 7, 8. Potadoma freethi, juveniles, Lobe.
FIG. 9. Lanistes libycns, Lake Barombi Mbo.
FIG. 10. Lanistes libvcus, Lobe.

Bull. B.M. (N.H.) Zoo/. 13, 3

PLATE 3

PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING

T /§& ' '•>>,

) I i?-."

BARBUS (PISCES, CYPRINIDAEJ^y^
OF THE VOLTA REGION

A. J. & J. HOPSON

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 4

LONDON: 1965

BARBUS (PISCES, CYPRINIDAE) OF THE
VOLTA REGION

A. J. & J. HOPSON

Fisheries Division, Accra, Ghanaf

f Present address: Federal Fisheries Service, Lake Chad Research Station, P.O. Box 227, Maiduguri,
Northern Nigeria.

Pp. 99-149; 1 8 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. 4

LONDON: 1965

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 13, No. 4 of the Zoological series.
The abbreviated titles of periodicals cited follow those of
the World List of Scientific Periodicals.

Trustees of the British Museum (Natural History) 1965

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued October, 1965 Price £i 2s.

BARBUS (PISCES, CYPRINIDAE) OF THE
VOLTA REGION

By A. J. & J. HOPSON

CONTENTS Page

SYNOPSIS . . . . . . . . . . . 101

INTRODUCTION ........... 101

CEPHALIC PIT-LINES . . . . . . . . . .103

Barbus ablabes (Bleeker) . . . . . . . . .105

Barbus spurrelli Blgr. . . . . . . . . .108

Barbus macrops Blgr. . . . . . . . . .110

Barbus par ablabes Blanc & Daget . . . . . . .114

Barbus hypsolepis Daget . . . .. . . . .117

Barbus nigeriensis Blgr. . . . . . . . . .119

Barbus lawrae Hopson . . . . . . . . .121

Barbus subinensis Hopson . . . . . . . . .124

Barbus trispilus (Bleeker) . . . . . . . . .126

Barbus sublineatus Daget . . . . . . . . .128

Barbus macinensis Daget . . . . . . . . .130

Barbus voltae Hopson . . . . . . . . .132

Barbus atakorensis Blanc & Daget . . . . . . .135

Barbus punctitaeniatiis Daget . . . . . . . .137

Barbus bawkuensis Hopson . . . . . . . . .139

Barbus pobeguini Pellegrin . . . . . . . . .141

Barbus stigmatopygus Blgr. ........ 143

Barbus leonensis Blgr. . . . . . . . . .145

KEY TO THE VOLTAIC SPECIES OF Barbus . . . . . .147

ACKNOWLEDGEMENTS .......... 148

REFERENCES ........... 149

SYNOPSIS

Eighteen species of Barbus, collected recently in Ghana, are described and figured. Fifteen
of the species were obtained from the Volta basin. The synonymy of West African Beirabarbus
is revised and four species of this subgenus are shown to occur in Ghana. Barbus nigeriensis is
redescribed using the holotype augmented with new material. A key is given for voltaic species
of Barbus.

INTRODUCTION

Barbus is one of the most specifically numerous genera of freshwater fish in Africa
and well over 200 species have been described. The genus is well represented in
West Africa and fifteen species were identified from a collection of fish which we made
in the Volta basin, chiefly in northern Ghana, during 1961. Most of the species are
widespread and common and Barbus forms an important and characteristic element
in the fish fauna of all habitats. The present investigations deal chiefly with voltaic

ZOOL. 13, 4 4

102 A. J. & J. HOPSON

species but observations are also made on three species of Barbus collected from the
Prah basin in the forest region of southern Ghana :

B. ablabes (Bleeker)
B. trispilus (Bleeker)
B. subinensis Hopson

Environmental conditions in the rivers of northern Ghana are comparable with those
described by Blanc & Daget (1957) for Haute Volta and by Holden (1963) for the
Sokoto River in Northern Nigeria which are indeed typical of the whole savannah
region of West Africa. Flooding occurs during a short rainy season lasting from
mid-May to mid-September followed by a rapid fall in water level with the onset of
the dry season. By the end of December the output of the Black Volta in northern
Ghana is greatly diminished and the White Volta is reduced to a mere trickle linking
a chain of sandy pools. Smaller tributaries dry out completely before the onset of
the rains in May. A more stable environment is found in permanent, well-vegetated
oxbow lakes which are particularly characteristic of the valley of the White Volta.
Relatively stable conditions are also found in dams constructed across seasonal
streams for agricultural purposes ; these are now common in Ghana.

Previous observations on Barbits in Ghana have dealt almost exclusively with
species occuning outside the Volta basin (Boulenger, 1911, 1916 ; Trewavas in
Irvine, 1947). A detailed account of voltaic species was made only during the last
decade by Blanc & Daget (op. cit.} who worked on material from Haute Volta. They
recorded nine species of Barbus from the Volta basin :

B. macrops Blgr. (= B. ablabes, sensu Blanc & Daget, 1957)

B. parablabes Blanc & Daget

B. voltae Hopson (=B. nigeriensis, sensu Blanc & Daget, op. cit.)

B. macinensis Daget

B. punctitaeniatus Daget

B. atakorensis Blanc & Daget

B. leonensis Blgr.

B. stigmatopygus Blgr.

Most of these are widely distributed elsewhere in the savannah region of West Africa
and all, with the exception of B. parablabes and B. atakorensis, were shown by Blanc
& Daget to be common to the Niger basin.

Six additional species were recorded from the Volta basin during the present survey :

B. spurrelli Blgr.
B. hypsolepis Daget
B. nigeriensis Blgr.
B. lawrae Hopson
B. bawkuensis Hopson
B. pobeguini Pellegrin

Of these, B. hypsolepis, B. nigeriensis and B. pobeguini also occur in the Niger basin.
In the course of the present investigations extensive use has been made of material

BARBUS OF THE VOLT A REGION 103

in the British Museum (N.H.) and in the Musee National d'Histoire naturelle, Paris.
Comparisons were made with type material whenever possible. Most of our obser-
vations agree with Daget (1954) and Blanc & Daget (1957). We differ mainly in the
synonymy of the subgenus Beirabarbus and in the diagnosis of B. nigeriensis.

Four species of Beirabarbus are shown to occur in Ghana : B. ablabes,: B. spunelli,
B. macrops and B. parablabes. Barbus deserti of Daget (1954) and B. ablabes of
Blanc & Daget (1957) are now regarded as misidentified examples of B. macrops,
which proves to be widely distributed through the savannah regions of West Africa
and the Chad basin. Barbus ablabes and B. deserti, with which B. macrops has been
confused, are apparently restricted in distribution, B. ablabes to the forest rivers
between south west Ghana and Liberia and B. deserti to the north central Sahara.

A redescription has been prepared for B. nigeriensis, based on the holotype, on
material from Western Nigeria and on specimens recently collected in northern
Ghana. The species described by Blanc & Daget (1957) as B. nigeriensis was
misidentified. It has been shown recently to be a new species, B. voltae (Hopson,

1965)-

Methods : the snout length was measured between the verticals to the tip and the
anterior margin of the eye ; the head length was taken to the posterior extremity
of the bony operculum. Ledges of skin forming the dorsal margin of each orbit are
included in the interorbital width. Perforated scales overlapping the base of the
caudal fin are included in the lateral line count. Other data were obtained in
accordance with standard practice.

All specimens were fixed initially in 5% formalin for three to four months. The
material was then carefully washed and transferred to an 80% solution of alcohol for
permanent preservation. It was found that this procedure satisfactorily exposes the
underlying melanophores, the pattern of which is often of taxonomic importance.

The relationship of lateral line to horizontal myoseptum often provided characters
of diagnostic value. Marked differences were noted among the Barbus under
consideration in the degree to which the lateral line dipped below the myoseptum.
In species of the subgenus Beirabarbus and in B. punctitaeniatus for example the dip
is slight whereas in B. nigeriensis and B. lawrae it is more pronounced. The maximum
distance between horizontal myoseptum and lateral line has been expressed in terms
of the depth of adjacent scale rows. The point at which the myoseptum and lateral
line converge also varies interspecifically. In Beirabarbus the point is usually above
the base of the anal fin whereas in B. perince and in B. werneri they converge near the
tip of the caudal peduncle.

CEPHALIC PIT-LINES

Herre (1932) first drew attention to cephalic pit-lines in Barbus palustris (== B.
radiatus, vide Greenwood, 1963) and considered them to be of sufficient importance
to be the criterion for establishing a new genus, Beirabarbus. Apparently unaware
of Metre's work, Schultz (1942) described a new genus Mannicthys (M. lucileae
Schultz = B. ma-crops Boulenger), on an identical basis. Johnels (1954) later showed
that cephalic pit-lines were present in at least six West African Barbus and doubted

io4 A- J- & J- HOPSON

the advisability of using them as generic or subgeneric characters. Recently,
Greenwood (1962) has given a detailed discussion on the occurrence of pit-lines in
African Barbus. He upholds Herre's taxon Beirabarbus at subgeneric level and
recognizes two distinct patterns of pit-line distribution.

(a) Pit-lines of the Beirabarbus type where the individual pits are small and very
numerous, grouped tightly into lines like beads and raised above the surface as
ridges. These lines sometimes branch and are present in a characteristic pattern on
the side of the snout, the cheek, the operculum and on the dorsal surface of the head.
On the basis of these characters, Greenwood referred the following species to the
subgenus Beirabarbus :

B. ablabes (Bleeker) B. deserti Pellegrin

B. radiatus Peters B. aurantiacus Blgr.

B. jae Blgr. B. macrops Blgr.

B. aspilus Blgr. B. spurrelli Blgr.
B. callipterus Blgr.

(b) The pits are relatively larger than in Beirabarbus and much fewer in number.
They are sometimes arranged in lines but with the pits always well separated from
one another. The lines are never raised above the surface in ridges and are usually
visible only on the cheek and operculum. Greenwood found this type of pit in fourteen
species of Barbus :

B. nigeriensis Blgr. B. trispilomimus Blgr.

B. kessleri (Steindachner) B. pumilus Blgr.

B. trispilus (Bleeker) B. anema Blgr.

B. congicus Blgr. B. svenssoni Johnels

B. pseudognathodon Blgr. B. collarti Poll

B. pleiwopholis Blgr. B. leonensis Blgr.

B. urostigma Blgr. B. cercops Whitehead

Of the above species, only B. kessleri has a serrated last simple dorsal ray and all
with the exception of B. cercops are West African in distribution.

Our own observations agree with Greenwood's. Cephalic pit-lines were noted in
all of the eighteen species of Barbus described in the present work. Of these, four
are referable to Beirabarbus :

B. ablabes

B. spurrelli

B. macrops

B. par ablabes Blanc & Daget

The remaining fourteen species all had weakly developed pit-lines corresponding
to the description under (b) above. They are as follows :

B. hypsolepis Daget B. voltae Hopson

B. nigeriensis Blgr. B. atakorensis Blanc & Daget

B. lawrae Hopson B. punctitaeniatus Daget

BARBUS OF THE VOLTA REGION 105

B. subinensis Hopson B. bawkuensis Hopson

B. trispiliis (Bleeker) B. pobeguini Pellegrin

B. siiblineatus Daget B. stigmatopygus Blgr.

B. macinemis Daget B. leonensis Blgr.

Cephalic pit-lines were always most clearly seen in formalin-fixed specimens.

Barbus (Beirabarbus) ablabes (Bleeker, 1863)
(Text-fig, i)

Puntius (Barbodes) ablabes Bleeker, 1863, Nat. Verh. Wet. Haarlem, 23 : 114, pi. 23, fig. i.
Barbus ablabes : Boulenger, 1911, Cat. Afr. Fish. 2 : 156, fig. 133.

Barbus ablabes : Pellegrin, 1923, Les poissons des eaux donees de I'Afrique occidentale : 134, Paris.
[non Barbus albabes : Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 132, fig. 13,
see p. no below].

SYNTYPES : two fish S.L. 64 and 66 mm. from Dabo Crom, Guinea in the Leyden
Museum.

DESCRIPTION based on 30 fish, 31-53 mm. S.L. from the R. Weiwei, Kumasi, south
Ghana supplemented with notes on the following material in the British Museum
(N.H.) : 10 fish, 44-70 mm. S.L. from the Kotchwah River, south Ghana (reg. no.
99.12.22.46-55) ; 4 fish, 36-48 mm. S.L. from the R. Atesu, south Ghana (reg. no.
1903.4.24.94-97) ; 2 fish, 58-62 mm. S.L. from near Bunsu, south Ghana (reg. no.
1938.12.15.40-41).

Body moderately compressed. S.L. 3-1 to 3-7 times the maximum depth and
3-1 to 3-6 times the length of the head. Predorsal profile convex with a slight
nuchal hump. Eyes inclined slightly upwards, the diameter 2-9 to 3-4 times in the
length of the head and 1-05 to 1-2 times in the interorbital width. Snout bluntly
pointed, 0-6 to 0-9 times the diameter of the eye. Mouth moderate, subterminal.
Anterior barbel overlapping the base of the posterior barbel, 0-4 to 0-6 times the
diameter of the eye. Posterior barbel extending well past the vertical to the centre
of the eye, 0-6 to 0-9 times the eye diameter. Pit-lines of the Beirabarbus type
present on the side of the snout, the cheek, the operculum and on the dorsal surface
of the head, between the eyes. Pectoral fin 0-6 to 0-8 times the length of the head,
the tip just overlapping the pelvic origin in the four smallest individuals (31-37 mm.
S.L.). Last simple dorsal ray smooth, flexible, 0-8 to i-o times the length of the
head. Tip of dorsal fin slightly rounded, distal margin concave. Dorsal fin rays
III 7 (f.i) or III 8 (f.29). Anal fin rays III 5. The pelvic origin lies between the
verticals to the third simple and the first branched dorsal rays. Caudal peduncle
i-i to 1-4 times as long as deep. The lateral line dips to a maximum of the depth of
half a scale row below the horizontal myoseptum in the anterior third of the body ;
the two converge at a point in the verticals to the last anal fin rays. Lateral line
scales 22 to 27 (modal number 24). Three and a half scales between the lateral line
and the dorsal origin, 3! scales between the lateral line and the mid-ventral line
immediately in front of the pelvic origin and 2\ scales between the lateral line and the
pelvic origin. Eleven (f.i) or 12 (f.29) scales round the caudal peduncle.

io6

A. J . & J. HO PS OX

FIG. i. Barbus ablabes ; N.B. in this and all following figures the caudal fin is simplified.

Summary of morphometric data based on 30 fish, 31-53 mm. S.L. ; measurements
are expressed as percentages of S.L.

Maximum depth .
Length head
Diameter eye
Interorbital width
Length snout
Length anterior barbel .
Length posterior barbel .
Length pectoral fin
Length dorsal fin .
Length caudal peduncle
Depth caudal peduncle .

COLORATION : in living specimens, greenish-brown above, silvery on the sides,
whitish below. A conspicuous dark mid-lateral band runs from the snout to the
posterior end of the body ; the band frequently shows a brilliant green iridescence.
Dorsal and caudal fins are tinged basally with yellow which becomes more intense
towards the distal margin to merge with a terminal band of orange-red. Anal fin
orange-red with a colourless free margin. Paired fins yellow on the anterior rays ;
the pelvic fins are also tinged with orange in brightly coloured individuals. Iris,
orange-red above. In formalin fixed specimens the dorsal surface is densely pig-
mented with fine melanophores free from a narrow zone near the margin of each
scale ; the pigment-free zone becomes wider on more lateral scales. Dorsal and
lateral scales are outlined with larger and darker melanophores, more concentrated
in a vertical bar on the pocket of each scale to form a regular pattern. These
markings are heaviest on the lateral line scales each of which bears a narrow vertical

Range

Mean

Allometry

27-5-33-3
27-6-32-5
8-5-10-2

29-6
29-8
9-4

Negative
Negative

9'8-H -2
6-2-8-7

10-5

7-6

—

3-8-5-9

6-2-8-9
19-6-23-7

4'5

7-4
21-7

—

25-4-30-2

27-7

—

17-4-21-4

19-7

—

14-2-15-9

15-3

—

BARB US OF THE VOLTA REGION 107

stripe in the shape of a cupid's bow. A black mid-lateral band, one third of a scale
row in depth, runs from the side of the snout, across the operculum and along the
body to the tip of the caudal peduncle. There are scattered melanophores on the
pockets of the first and sometimes the second scale row below the lateral line but the
ventral surface is generally pigment-free. The anterior margin of the dorsal fin
appears greyish-black owing to a moderately heavy peppering of melanophores on
the membrane between the second and third simple rays, along the anterior margin
of the distal half of the third simple ray and to a lesser extent on the second simple
ray and on the membrane between the third simple and first branched rays. Pig-
mentation on the remainder of the fin is restricted to scattered and inconspicuous
melanophores chiefly on the distal parts.

DIAGNOSIS AND AFFINITIES : Dr. Greenwood recently compared the syntypes of
B. ablabes in the Leyden Museum with material from the Kotchwah River, south
Ghana (Brit. Mus. (N.H.) reg. no. 99 . 12 . 22 . 46-55) which has been used to supplement
the above description. He found (pers. comm.) that the two collections were
identical in barbel length and in details of pigmentation not lost in preservation.
There was no sign in the type specimens of particularly heavy pigment in the region
of the second simple dorsal ray (a heavily pigmented second simple ray is characteris-
tic of the closely related B. macrops).

Barbus ablabes resembles B. macrops in morphometric details and in pigmentation.
The most obvious difference is in the distribution of melanophores on the dorsal fin.
In B. ablabes moderately heavy pigment is distributed along the entire length of the
second and third simple rays and the anterior margin thus appears uniformly grey ;
in B. macrops the second simple ray and the tips of the first to third branched rays
are densely pigmented so that the dorsal fin has a broad black tip and a black streak
down the lower half of the anterior margin. The part of the third simple ray lying
between these two areas of dense pigment is almost clear of melanophores. The two
species also differ in the form of the pocket pigment on the lateral line scales ; in
B. ablabes the markings are relatively slender in the shape of a cupid's bow, whereas
in B. macrops the markings are broader and triangular. Barbus ablabes is readily
separated from West African populations of B. macrops by the relatively longer
posterior barbels (6-2-8-9 C-T- 2'5 to 5-1% S.L. in Ghanaian material). The relative
length of the posterior barbel in B. macrops, however, varies geographically and a
population from the Tibesti shows a slight overlap with B. ablabes (5-1-6-6 in B.
macrops c.f. 6-2-8-9% S.L.).

The relative diameter of the eye which Boulenger (1911) originally used to separate
the two species is no longer of diagnostic value. This character also varies geo-
graphically in B. macrops, and in the Tibesti population the eyes are relatively
smaller than in B. ablabes.

The geographical distributions of B. ablabes and B. macrops do not appear to
overlap. Barbus ablabes is recorded only from the forest rivers of south-west Ghana
and Liberia whereas B. macrops is widespread throughout the savannah rivers of
West Africa and the Chad basin.

Barbus ablabes is easily distinguished from B. spurrelli by the dark mid-lateral band

io8 A. J. & J. HOPSON

and the denseness of the pocket pigment on the lateral line scales. In B. spurrelli
the mid-lateral band is very faint and the pigmentation of lateral line scales is only
slightly heavier than that of more dorsal scales.

Barbus ablabes is compared with B. parablabes in the description of that species.

DISTRIBUTION : in Ghana, common and widespread in forest streams and rivers in
the southwest. Probably absent from the Volta Basin. Elsewhere, Dabo Crom,
Guinea (type locality) ; Liberia (no locality given, Boulenger, 1911). The record of
B. ablabes from the St. Paul basin, Liberia (Schultz, 1942) is doubtful ; Dr. Weitzman
of the Smithsonian Institution reports that the specimens appear to have no Beira-
barbus type cephalic pit-lines (pers. comm.}. Barbus ablabes probably belongs to a
faunal group restricted to the forest rivers of western Ghana, Cote d'lvoire and
Liberia. The group may also include Alestes longipinnis (Giinther), Petersius
occidentalis (Giinther), Nannocharax seyboldi Schultz, Barbus trispilus (Bleeker) and
Eutropius mentalis Blgr. among other species.

Barbus (Beirabarbus) spurrelli Boulenger, 1913
(Text-fig. 2)

Barbus spurrelli Boulenger, 1913, Proc. zool. Soc. Lond. : 51, pi. 3, fig. i ; Idem, 1916, Cat. Afr.

Fish. 4 : 264, fig. 162.
Barbus ablabes var. spurrelli Pellegrin, 1923, Les poissons des eaux douces de I'Afrique occidentale :

135, Paris.
Barbus spurrelli : Daget, 1952, Mem. Inst. France. Afr. Noire, no. 19 : 316, fig. 7.

LECTOTYPE : 67-8 mm. S.L.,B.M. (N.H.) reg. no. 1911.11.27.1, from near Dunkwa
(Prah Basin) south Ghana. We consider this fish to be the specimen illustrated in
Boulenger, 1916, fig. 162.

DESCRIPTION based on 12 fish, 25-46 mm. S.L. from the River Azubone (Volta
Basin), near Mpraeso, south Ghana. Body compressed. S.L. 3-25 to 3-5 times the
maximum depth and 2-9 to 3-5 times the length of the head. Predorsal profile
convex with a slight nuchal hump. Snout bluntly pointed, 0-85 to i-i times the
diameter of the eye. Mouth moderate, subterminal. Anterior barbel 0-3 to 0-7
times the diameter of the eye, barely overlapping the base of the posterior barbel.
Posterior barbel 0-6 to 0-95 times the diameter of the eye, the tip reaching the vertical
to the posterior margin of the pupil. Eyes inclined slightly upwards, the diameter
3-25 to 3-85 times in the length of the head and 1-2 to 1-5 times in the interorbital
width. Pit-lines of the Beirabarbus type present on the side of the snout, the cheek,
the operculum and on the dorsal surface of the head, between the eyes. Pectoral
fin 0-6 to 0-75 times as long as the head, sometimes (f.3) overlapping the pelvic origin.
Last simple dorsal ray smooth, flexible, 0-75 to 0-85 times the length of the head.
Apex of dorsal fin slightly rounded, distal margin concave. Dorsal fin rays III 7
(f.i) or III 8 (f.n). Anal fin rays III 5. The pelvic origin lies between the verticals
of the first and second branched dorsal rays. Caudal peduncle 1-2 to 1-55 times as
long as deep. The lateral line dips to a maximum of the depth of slightly over half
a scale row below the horizontal myoseptum in the anterior third of the body ; the

BARBUS OF THE VOLTA REGION

109

two converge above the posterior anal fin rays. Lateral line scales 24 to 26 (modal
number 25). Three and a half to 4! scales between the lateral line and the dorsal
origin, 3^ scales between the lateral line and the mid-ventral line immediately in
front of the pelvic origin and 2 to 2| scales between the lateral line and the pelvic
origin. Twelve scales round the caudal peduncle.

10mm.

S.L. ; measure-

FIG. 2. Barbus spurrelli.

Summary of morphometric data based on 12 fish, 25 to 46 mm.
ments are expressed as percentages of S.L.

Range Mean Allometry

Maximum depth . . . 28-3-30-5 29-3

Length head . . . 28-5-33-0 31-5 Negative

Diameter eye . . • 7 ' 4-9 '6 8-9 Negative

Interorbital width . . 10-8-13-0 11-7

Length snout . . .7-8-8-8 8-2

Length anterior barbel . . 3 • 2-5 -8 4-7

Length posterior barbel . . 5 • 7-7 -9 7-2

Length pectoral fin . . 18-6-24-6 22-2

Length dorsal fin . . . 22-6-28-1 25-8

Length caudal peduncle . . 18-5-23-3 20-8

Depth caudal peduncle . . 14-3-15-1 14-6

COLORATION : in formalin fixed specimens the dorsal surface is relatively heavily
pigmented with fine melanophores absent only from a narrow zone near the margin of
each scale. Dorsal and lateral scales are outlined with larger and darker melano-
phores ; this pigment is more concentrated in a vertical bar on the pocket of each
scale to form a regular pattern. Lateral line scales are marked with similar but
slightly heavier pigment ; each vertical bar is bisected horizontally by the lateral
line pore. Diffuse melanophores form an indistinct band, approximately one third
of a scale row in depth, running just below the horizontal myoseptum in the anterior
half of the body ; the two converge over the anal fin and run together onto the caudal

no A. J . & J. HOI'S ON

peduncle where the band ends in front of a small and inconspicuous terminal spot,
overlapping the base of the caudal fin. A narrow black band runs obliquely down-
wards from the lateral line origin to the base of the pectoral fin. The ventral surface
is generally pigment-free, with the exception of a few scattered melanophores on the
pockets of the first and sometimes the second row of scales below the lateral line.
The snout is marked with a dark lateral spot. The anterior margin of the dorsal fin
is moderately peppered with melanophores which are slightly heavier on the free
edge of the last simple ray. Melanophores are thinly distributed over the distal
half of the branched dorsal rays.

DIAGNOSIS AND AFFINITIES : these data are in close agreement (allowing for
allometry) with the type description. Barbus spurrelli is readily distinguishable from
other Beirabarbus occurring in West Africa either by the absence of a dark mid-
lateral band or by the lack of a dark spot on or near the tip of the dorsal fin. Detailed
comparisons with B. ablabes and B. pamblabes are given in the descriptions of those
species.

DISTRIBUTION : Ghana. The type locality of B. spurrelli is Dunkwa (Prah basin)
in the forest region of south Ghana. Our material was collected from the R. Azubone
near Mpraeso, a forest stream on the Volta side of the Prah-Volta watershed. Else-
where, Sierra Leone (Pellegrin, 1923) ; Mount Nimba, French Guinea (Daget, 1952).
The record of B. spurrelli from the St. Paul basin, Liberia (Schultz, 1942) is doubtful.
Dr. Weitzman of the Smithsonian Institution has kindly re-examined the material
and reports that no Beirabarbus cephalic pit-lines are visible (pers. comm.).

Barbus (Beirabarbus} macrops Boulenger, 1911
(Text-fig. 3)

Barbus macrops Boulenger, 1911, Ann. Mag. nat. Hist. (8) 7 : 374 ; Idem, 1916, Cat. Afr. Fish.

4 : 265, fig. 163.
Barbus deserti : Pellegrin, 1921 (Ennedi and Tibesti specimens only), Mem. Soc. Sci. nat. Maroc,

(1)2:143.
Barbus macrops : Pellegrin, 1923, Les poissons des eaux douces de VAfrique occidentale : 136,

Paris.

Barbus gambiensis Svensson, 1933, Kungl. Sven. vet. Handl. (12) 3 : 67, pi. 4, fig. i.
Mannichthys lucileae Schultz, 1942, Proc. U.S. Nat. Mus. 92 : 321, pi. 35, fig. 3.
Barbus deserti : Daget, 1954, Mem. Inst. France. Afr. Noire, no. 36 : 192, fig. 62.
Barbus ablabes : Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 132, fig. 13.

LECTOTYPE : a fish of 45-0 mm. S.L. (Brit. Mus. (N.H.) reg. no. 1912.4.1.110),
from above the rapids, Crobal River. Portuguese Guinea. We consider this to be the
specimen illustrated in Boulenger, 1916, fig. 163.

DESCRIPTION based on 24 fish, 31 to 56 mm. S.L. from a tributary of the White
Volta, 5 m. N. of Bawku, northern Ghana. Body moderately compressed. S.L.
3-3 to 3-9 times the maximum depth and 3-3 to 37 times the length of the head.
Predorsal profile convex with a slight nuchal hump. Snout bluntly pointed, 0-7 to
0-85 times the diameter of the eye. Mouth moderate, sub-terminal. Barbels
moderate, the anterior o-i to 0-35 times the diameter of the eye, the tip barely over-

B A R B US OF THE VOLT A REGION

lapping the base of the posterior barbel. Posterior barbel 0-25 to 0-6 times the eye
diameter, the tip reaching the vertical to the centre of the eye in a few individuals.
Eyes inclined slightly upwards, 2-85 to 3-1 times in the length of the head and 1-05
to 1-15 times in the interorbital width. Pit-lines of the Beirabarbus type fully
developed on the side of the snout, the cheek, the operculum and on the dorsal
surface of the head between the eyes. Pectoral fin 0-65 to 0-8 times the length of the
head, the tip always falling short of the pelvic origin. Last simple dorsal ray, smooth,
flexible, 0-9 to 1-05 times the length of the head. Apex of dorsal fin slightly rounded,
distal margin concave. Dorsal fin rays III 7 (f.i) or III 8 (f.23). Anal fin rays
III 5. The origin of the dorsal fin lies between the verticals to the last simple and
first branched dorsal rays. Caudal peduncle 1-3 to 1-6 times as long as deep. The
lateral line dips to a maximum depth of slightly over half a scale row below the
horizontal myoseptum in the anterior third of the body ; the two converge in the
vertical to the last anal fin rays. Lateral line scales 24 to 28 (modal number 25).
Three and a half scales between the lateral line and the dorsal origin, 3-J- scales between
the mid- ventral line immediately in front of the pelvic origin and 2-S- scales between
the lateral line and the pelvic origin. Twelve scales round the caudal peduncle.

FIG. 3. Barbiis macrops.

Summary of morphometric data based on 24 fish, 31 to 56 mm.
ments are expressed as percentages of S.L.

Maximum depth .
Length head
Diameter eye
Interorbital width
Length snout
Length anterior barbel
Length posterior barbel
Length pectoral fin
Length dorsal fin .
Length caudal peduncle
Depth caudal peduncle

S.L. ; measure-

Range

Mean

Allometry

25-5-30-2
26-4-29-7
8-3-10-2

27-1
27-6
9-1

Negative
Negative

9-2-12 -2

6-3-7-3

10-2

6-8

—

1-2-3-1

2-3

— •

2-5-5-1
17-8-21-8
25-2-28-8

l8 • I-2I -I

3'5
19-7
26-6
19-9

—

13-3-14-8

14-1

—

ii2 A. J. & J. HOPSON

COLORATION : in living specimens, greenish-brown above, silvery on the sides,
whitish below. A dark mid-lateral band extends from the snout to the tip of the
caudal peduncle. Dorsal and caudal fins bright orange-red, the dorsal with a con-
spicuous black tip. The other fins are frequently tinted orange-yellow. Iris, bright
orange-red above. In formalin fixed specimens the dorsal surface is densely pig-
mented with fine melanophores absent only from a narrow zone near the margin of
each scale. Dorsal and lateral scales are clearly outlined with larger and darker
melanophores which are more concentrated in a vertical line on the pocket of each
scale to form a regular pattern. These markings are heaviest on the lateral line
where each scale bears a relatively broad and triangular group of melanophores,
bisected horizontally by the lateral line pore. A conspicuous black band, one-third
of a scale row in depth, runs mid-laterally from the side of the snout, across the
operculum and along the body to the end of the caudal peduncle. In some speci-
mens an oblique dark stripe is noticeable between the origin of the lateral line and the
base of the pectoral fin. The pattern of scale pocket pigment, described above for
dorsolateral scales, is repeated more faintly on the first and sometimes the second
scale row below the lateral line. The ventral surface is generally pigment-free. The
apex of the dorsal fin bears a dense black spot extending over the distal third of the
first four branched rays but heaviest on the tips of the first two. Dense pigment is
also present in the vicinity of the second simple ray, particularly on the membrane
between it and the third ray. The third simple ray is less heavily pigmented and
closely packed melanophores are present only in the basal third along the anterior
margin and near the tip ; the central section of the third simple ray is almost pigment-
free.

GEOGRAPHICAL VARIATION : during the present investigations, B. macrops were
examined from most of the savannah region of West Africa, the Chad Basin, the
Hoggar in the western Sahara and the Tibesti in the eastern Sahara. Details of
pigmentation, particularly on the dorsal fin, proved to be constant throughout this
wide area. Marked geographical variation was however noted in certain morpho-
metric characters, especially eye diameter but also the relative length of the barbels
as demonstrated below :

B. macrops : geographical variations in eye diameter (N.B. — negative allometry).

No. Range S.L. Eye diameter

Locality examined (mm.) expressed as % S.L.

Tibesti 24 43-55 6-2-7-6

Niger (N. Nigeria) . 3 48-60 7-8-8-6

Sierra Leone . 2 61 8-0-8-4

Volta (N. Ghana) . . 24 31-56 8-3-10-2

Volta (S. Ghana) . . 17 21-52 8-2-10-6

Hoggar ... 4 26-39 8-6-10-0

Chad Basin (Nigeria) . 12 27-48 9-0-10-2

Port. Guinea . . 16 25-60 8-8-11-6

B. macrops : geographical variations in posterior barbel length.

BARBUS OF THE VOLTA REGION 113

No. Range S.L. Length post, barbel
Locality examined (mm.) expressed as % S.L.

Chad Basin (Nigeria) . 12 27-48 2-4-3-3

Volta (N. Ghana) . . 24 3!~56 2-5-5-1

Volta (S. Ghana) . . 17 21-52 2-7-6-0

Hoggar . 4 26-39 3'O-5'i

Port. Guinea . . 16 25-60 3'°-5'3

Sierra Leone . . 2 61 4 . 5-5 • i

Niger (N. Nigeria) . 3 48-60 4-8-5-4

Tibesti 24 43-55 5-1-6-6

Diagnosis and Affinities : there has been considerable confusion between B. macrops
and B. deserti Pellegrin. This undoubtedly originated from the inclusion of B.
macrops (localities Tibesti and Ennedi) in material used for a redescription of B.
deserti (Pellegrin, 1921).

We have examined type material of B. deserti from the Tassili des Azdjers, central
Sahara, both the syntypes in the Paris Museum (reg. no. 09.457-458) and a syntype
in the British Museum (N.H.) (reg. no. 1909.12.9.7) on which Boulenger (1911)
based his diagnosis. They all differ from B. macrops in the following details :

(a) In B. deserti there is no well-defined mid-lateral band running from the snout
across the operculum to the tip of the caudal peduncle. Barbus deserti is marked only
in the posterior half of the body with relatively faint dark pigment on the horizontal
myoseptum.

(b) The lateral line scales are relatively narrower and deeper in B. deserti (cf. B.
parablabes) ,

(c) The lateral line scale pockets of B. deserti are marked with narrow vertical
stripes (cf. B. parablabes), never with broader triangular spots as in B. macrops.

(d) The second simple dorsal fin ray (always dense black in B. macrops} is only
lightly pigmented.

(e) Further details of pigmentation of the dorsal fin in B. deserti are obscured by
damage to the tip in all the type material. Pellegrin (1909) noted that the dorsal fin
was tipped with black whereas Boulenger (op. cit.) records that a spot was present
near the tip. It seems likely that Boulenger's description is the more accurate of the
two. Three Barbus from the oasis of El Barkat, Fianferrari, Fezzan (within 500
miles of the type locality in the same region of the N. Central Sahara) in the British
Museum (N.H.) (reg. no. 1937.7.8.4-6), similar to the types in every respect, bear a
spot on the distal half of the last simple and first three branched dorsal rays, well
clear of the distal margin. This should prove a trenchant character in distinguishing
B. deserti from B. macrops where the black pigment extends to the tip of the anterior
dorsal rays.

Comparisons of B. macrops with B. ablabes and B. parablabes are included in the
descriptions of those species.

DISTRIBUTION AND HABITAT : in Ghana, restricted to the Volta basin where it is
the most common and widespread species of Barbus. Barbus macrops occurs in
rivers, well vegetated lakes and dams, over all substrates coarse and fine. Elsewhere,

H4 A.J.&J.HOPSON

distributed over most of the savannah area of West Africa and the Chad basin,
extending into the Sahara in the Hoggar to the west and the Tibesti to the east :
Portuguese Guinea (Boulenger, 1911, type locality) ; Gambia River (Svensson, 1933) ;
L. Kwarko, Sierra Leone (Brit. Mus. (N.H.) reg. no. 1958.9.18.54-55) ; St. Paul and
Bolor Rivers, Liberia (Schultz, 1942) ; Middle Niger (Daget, 1954) ; Arak Gorge,
Hoggar (Brit. Mus. (N.H.) reg. no. 1932.5.6.5-11); R. Katagum (Chad Basin)
N. Nigeria (Brit. Mus. (N.H.) reg. no. 1928.7.3.56-62) ; Sherda, Tibesti (Brit. Mus.
(N.H.) reg. no. 1960.6.7.111-160).

LIFE HISTORY : in the White Volta, B. macrops spawns during the rains. Gonads
start to mature during April and the majority of fish are ripe by mid- June when the
first post-larvae appear in the river. Spawning is over by the end of July and the
size-composition of samples taken in August suggests that most fish die at the age of
one year. Juvenile fish have grown to a modal length of c. 35 mm. by November.
Females, which ripen at a minimum S.L. of c. 50 mm, grow slightly larger than
males. Maximum size, a female of 98 mm. S.L.

Barbus (Beirabarbus) parablabes Blanc & Daget, 1957

(Text-fig. 4)
Barbus parablabes Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 134, fig. 13.

HOLOTYPE : a fish 62 mm. S.L. from between Tanguieta and Natitingou, Volta
basin, N. Dahomey, Paris Mus. reg. no. 56-108.

DESCRIPTION based on 24 fish, 39-52 mm. S.L. from the waterworks reservoir at
Tamale (Volta basin), northern Ghana. Body moderately compressed. S.L. 3-3 to
3-9 times the maximum depth and 3-2 to 3-6 times the length of the head. Predorsal
profile convex with a slight nuchal hump in the larger individuals (c. 45 mm. S.L. and
over). Snout bluntly pointed, 0-6 to 0-8 times the diameter of the eye. Mouth
moderate, subterminal. Barbels moderate, the anterior 0-25 to 0-4 times the dia-
meter of the eye, its tip reaching the base of the posterior barbel. Posterior barbel
0-45 to 0-6 times the diameter of the eye, the tip extending to between the verticals
to the anterior margin and the centre of the pupil. Eyes inclined slightly upwards,
the diameter 2-95 to 3-35 times in the length of the head and i-o to 1-15 times in the
interorbital width. Well developed Beirabarbus type pit-lines on the side of the
snout, the cheek, the operculum and on the dorsal surface of the head between the
eyes. Pectoral fin rather long, 0-7 to 0-85 times the length of the head, the tip
frequently overlapping the pelvic origin, particularly in smaller males. Last simple
dorsal ray smooth, flexible, 0-9 to i-o times the length of the head. Apex of dorsal
fin slightly rounded, distal margin concave. Dorsal fin rays III 7 (f.i) or III 8 (f.23).
Anal fin rays III 5. The pelvic origin lies between the verticals to the last simple and
first branched dorsal fin rays. Caudal peduncle slender, 1-4 to 1-8 times as long as
deep. The lateral line dips to a maximum of slightly more than the depth of half a
scale row below the horizontal myoseptum in the anterior third of the body ; the
two converge in the vertical to the anal fin, usually over the anterior rays. Lateral

BARB US OF THE VOLTA REGION 115

line scales 23 to 27 (modal number 25). Three and a half scales between the lateral
line and the dorsal origin, 3^ scales between the lateral line and the mid-ventral line
immediately in front of the pelvic origin and 2-2 1 scales between the lateral line and
the pelvic origin. Nine (f.i), 10 (f.i8) or n (f.5) scales round the caudal peduncle ;
the odd row of scales, when present always lies along the mid-ventral line (N.B. —
Daget records only 12 scales round the caudal peduncle in the type description).

10mm.

FIG. 4. Barbus parablabes.

Summary of morphometric data based on 24 fish, 39 to 52 mm. S.L. ; measure m nts
are expressed as percentages of S.L.

Range Mean Allometry

Maximum depth . . . 25-8-29-6 27-5

Length head . . . 27-3-30-7 28-9

Diameter eye . . . 8 • 9-9 -7 9-3

Interorbital width . . 8-9-10-4 9-8

Length snout . . . 6 • 4-7 -6 6-9

Length anterior barbel . . 2 • 5-3 -6 2-9

Length posterior barbel . . 4-1-5-7 4-9

Length pectoral fin . . 21-2-24-7 23-0

Length dorsal fin . . . 26-8-31-2 28-0

Length caudal peduncle . . 20-0-24-0 22-0

Depth caudal peduncle . . 13-1-14-4 13-7

COLORATION : in living specimens, dorsal surface greyish green, flanks silvery,
whitish below. A greyish mid-lateral band is punctuated with short dark vertical
streaks on the lateral line. Caudal and dorsal fins clear rose-pink, the dorsal less
brightly coloured than the caudal ; all other fins colourless. Iris reddish above.
In formalin-fixed specimens, dorsal surface with a moderately heavy scattering of
fine melanophores, free from a narrow zone near the margin of each scale ; the clear
zone becomes increasingly wide on the flanks. Dorsal and lateral scales are outlined

ZOOL. 13, 4 5

n6 A. J . & J. HOPSON

with larger and darker melanophores which are more concentrated in a vertical band
on the pocket of each scale to form, a regular pattern. Pocket pigment is particularly
heavy on the lateral line where each scale is marked with a conspicuous crescent-
shaped vertical stripe traversed by the lateral line pore. A well-marked band of
diffuse melanophores, one third of a scale row in depth, runs mid-laterally from the
side of the snout, across the operculum, to the tip of the caudal peduncle. The
ventral surface is generally pigment free with the exception of scattered melano-
phores on the pockets of the first and sometimes the second row of scales below the
lateral line. The dorsal fin is relatively lightly peppered with melanophores chiefly on
the second simple ray and on the distal halves of the third simple and first three
branched rays. This pigment is sometimes more concentrated towards the apex of
the fin but never enough to form a definite spot. Fine lines of black pigment are
frequently present on the anterior and posterior edges of the fin rays particularly in
the basal half of the fin ; this may produce a streaky appearance.

DIAGNOSIS AND AFFINITIES : Barbus pamblabes is readily distinguished from B.
macrops (with which it usually occurs in North Ghana) by the absence of a heavy
black spot at the tip of the dorsal fin, the relative lightness of the lateral band and
the slender crescentic markings on the pockets of the lateral line scales (broader and
triangular in B. macrops) . The lateral line scales themselves are narrower and deeper
in B. parablabes. The caudal peduncle is more slender in B. parablabes, and in
northern Ghana the lower number of scales round the caudal peduncle (9-11 cf. 12 in
B. macrops} is diagnostic. However, as noted above, 12 scales round the caudal
peduncle were recorded in the type material of B. parablabes from north Dahomey
(Blanc & Daget, 1957). The pectoral fin is relatively longer in B. parablabes, fre-
quently overlapping the pelvic origin in the smaller males (never in B. macrops.)
In living specimens the pale pink of the caudal fin is strikingly different to the bright
orange red of B. macrops.

Most of the above characters also serve to distinguish B. parablabes from B. ablabes.
Both species are, however, without a black tip to the dorsal fin and the stripes on the
lateral line scale pockets are only slightly broader in B. ablabes than in B. parablabes.
The shorter posterior barbel of B. parablabes (4-9-57 cf. 6-2-8-9% in B. ablabes) is
diagnostic.

Compared with B. spurrelli, B. parablabes has a shorter snout (6-4-7-6 cf. 7-8-8-8%
S.L.), shorter posterior barbels (4-1-5-7 cf. 5-7-7-9% S.L.) and a narrower caudal
peduncle (13-1-14-4 cf . 14-3-15-1% S.L.) . The dark mid-lateral band is much heavier
in B. parablabes and the vertical lines of pigment on the lateral line scales are narrower
and more emphasized ; in B. spurrelli there is relatively little contrast between the
pigmentation of the lateral line and more dorsal scales.

DISTRIBUTION AND HABITAT : in Ghana, widespread and common throughout the
northern region in rivers, streams and dams. Occurs over all substrates though
rarely in thickly vegetated habitats. No records from the south or from outside
the Volta basin. Elsewhere, recorded from a tributary of the River Oti (Volta basin)
in north Dahomey (type locality).

B A R BUS OF THE VOLTA REGION

117

NATURAL HISTORY : Barbus parablabes spawns during the rains. Gonads start to
mature during April and by mid- June the majority of fish are ripe. Juveniles appear
in the rivers during July and by mid- August the larger mature fish have disappeared,
suggesting that death occurs after breeding, at the age of approximately one year.
Females grow larger than males. Maximum size : a female of 53 mm. S.L.

Barbus hypsolepsis Daget, 1959

(Text-fig. 5)
Barbus hypsolepis Daget, 1959, Bull. Inst. France. Afr. Noire, 21 (A), 2 : 670, fig. i.

DESCRIPTION based on 21 specimens 14-27 mm. S.L. from the Black Volta, 3 miles
west of Lawra, north Ghana. Body moderately compressed. S.L. 3-65 to 4-15 times
the maximum depth and 3-05 to 3-90 times the length of the head. Pre-dorsal
profile smooth, convex. Snout somewhat pointed, 0-4 to 0-7 times as long as the
diameter of the eye. Mouth moderate, subterminal and protrusible obliquely
downwards. Weakly -developed pit-lines clearly visible on the cheeks. Anterior
barbel 0-03 to 0-25 times the diameter of the eye, the tip overlapping the base of the
posterior barbel only in the larger specimens. Posterior barbel 0-05 to 0*45 times
the diameter of the eye, the tip extending in larger specimens beyond the vertical to
the anterior margin of the pupil. Eyes large, inclined slightly upwards, 2-35 to
2-95 times in the length of the head and 0-85 to 1-05 times in the interorbital width.
Pectoral fin 0-55 to 0-75 times the length of the head, the tip falling short of the pelvic
origin. Last simple dorsal ray smooth, flexible, 0-9 to 1-05 times as long as the head.
Apex of dorsal fin pointed, distal margin concave. Dorsal fin-rays III 8. Anal
fin-rays III 5. Origin of pelvic fin below the vertical to the last simple dorsal ray.
Caudal peduncle slender, 1-55 to 1-9 times as long as deep. The lateral line scales
are unusually deep, particularly below the origin of the dorsal fin. The lateral line
dips to a maximum of the depth of a third of a scale row below the horizontal

10mm.

FIG. 5. Barbus hypsolepis.

"8 A. J. & J. HOPSON

myoseptum in the anterior third of the body ; the two converge in the vertical to the
base of the anal fin. Lateral line scales 22 to 24 (modal number 23). Two and a half
scales between the lateral line and the dorsal origin, z\ scales between the lateral line
and the mid- ventral line immediately in front of the pelvic origin and \\ scales
between the lateral line and the pelvic origin. Eight scales round the caudal
peduncle.

Summary of morphometric data based on 21 fish, 14 to 27 mm. S.L. ; measurements
are expressed as percentages of S.L.

Range Mean Allometry

Maximum depth . . . 23 -5-27 -8 25-5

Length head . . . 25-6-32-4 28-8 Negative

Diameter eye . . . 9-7-12-8 10-7

Interorbital width . . 9-7-11-0 10-4

Length snout . . . 4 • 7-7 • i 5-8

Length anterior barbel . . 0-1-2-6 1-2 Positive

Length posterior barbel . . o • 6-4 -6 2-4 Positive

Length pectoral fin . . 16-8-22-5 19-0

Length dorsal fin . . . 25-8-30-4 27-3

Length caudal peduncle . . 19-6-23-9 21-4

Depth caudal peduncle . . 1 1-8-14-0 12-5

COLORATION : in living specimens, body nacreous silver with a pinkish flush on the
dorsal surface. Scales clearly outlined with dark pigment. Dorsal and caudal fins
pale pinkish-orange, anal and paired fins tinged with the same colour. Iris with an
orange-red arc above. Daget (1959) noted that in B. hypsolepis from the Middle
Niger, females are less brightly coloured than males and have a greyish dorsal fin.
In formalin-fixed specimens scales of the lateral line and above are clearly outlined
with dark pigment which is emphasized in a slender, vertical arc of melanophores on
each scale pocket. A few scattered melanophores form an indistinct band on the
horizontal myoseptum, more noticeable in the posterior half of the body. There is
frequently a dark bar of pigment immediately behind the operculum, running
obliquely downwards from the lateral line to the base of the pectoral fin. Daget
(op. cit.} has observed that in B. hypsolepis this bar is present only in males.

DIAGNOSIS : these data agree closely (allowing for allometry) with the original
description of material from the Middle Niger (Daget, op. cit.}. Daget includes
B. hypsolepis in the subgenus Clypeobarbus which differs from other Barbus in having
exceptionally deep lateral line scales, particularly in the region below the origin of
the dorsal fin. He observes that when more material becomes available it may be
possible to consider B. hypsolepis as a subspecies of B. pleuropholis , a species occurring
in the Congo basin.

DISTRIBUTION : in Ghana local and uncommon, known only from the Black Volta
near Lawra and from the White Volta near Bolgatanga. Elsewhere, the Middle
Niger (Daget, op. cit.).

BARBUS OF THE VOLTA REGION ng

Barbus nigeriensis Boulenger, 1902
(Text-fig. 6)

Barbus nigeriensis Boulenger, 1902, Proc. zool. Soc. London, 2 : 327, pi. 28, fig. 3. ; Idem, 1910

(type only), Cat. Afr. Fish, 2 : 154, fig. 130.
[non Barbus nigeriensis : Trewavas, 1947, in Irvine, Fisheries and Fish of the Gold Coast, London ;

non Barbus nigeriensis : Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 135,

fig. 14.]

DESCRIPTION based on 20 fish, 23 to 50 mm. S.L., from the White Volta and its
tributaries near Bawku, northern Ghana. Body moderately compressed. S.L. 3-2
to 4-1 times the maximum depth and 3-2 to 4-1 times the length of the head. Pre-
dorsal profile convex with a slight nuchal hump. Snout rather pointed, 0-65 to 0-9
times in the diameter of the eye. Mouth relatively large, subterminal. Barbels
long, the anterior 0-65 to 1-2 times the diameter of the eye, and the posterior 0-8 to
i -4 times the diameter of the eye. In larger specimens the tip of the anterior barbel
reaches the vertical to the centre of the eye and the tip of the posterior barbel extends
beyond the posterior margin. Eyes inclined slightly upwards, relatively large, 2-5
to 3-2 times in the length of the head and 0-95 to 1-2 times in the interorbital width.
Weakly-developed pit-lines are visible on the cheek and operculum. Pectoral fin
0-55 to 0-75 times the length of the head, the tip not reaching the pelvic origin. Last
simple dorsal ray smooth, flexible, 0-8 to i-o times as long as the head. Apex of
dorsal fin somewhat rounded, distal margin slightly concave. Dorsal fin rays III 8
(f.ig) or III 9 (f.i). Anal fin rays III 5. The origin of the pelvic fin lies between
the verticals to the first and third branched dorsal fin rays. Caudal peduncle 1-35
to 1-55 times as long as deep. The lateral line dips to a maximum of approximately
one and a quarter scale rows below the horizontal myoseptum in the anterior third
of the body ; the two converge in the vertical to the base of the anal fin. Lateral
line scales 27 to 30 (modal number 29) . Four and a half scales between the lateral
lines and the dorsal origin, 3! to 4^ scales between the lateral line and the mid-
ventral line immediately in front of the pelvic origin and 2| scales between the lateral
line and the pelvic origin ; 12 scales round the caudal peduncle.

COLORATION : in living specimens greenish-brown above, silvery on the flanks and
whitish below. A dark narrow mid-lateral band is present. Dorsal and caudal fins
clear pale pink. The eye is marked with a red crescent over the pupil. In formalin-
fixed specimens scales of the dorsal surface with dense melanophores, free from a
narrow marginal band and less dense at the centre of each scale. This pattern, which
is darkest on the scales at the base of the dorsal fin, fades out rather abruptly on the
flanks. Dense melanophores lying over the horizontal myoseptum form a narrow
dark band running the entire length of the body. The band is generally less than a
scale row in depth, becoming slightly wider on the caudal peduncle and frequently
more dense at the tip to form a vague spot. The band may be interrupted below the
anterior rays of the dorsal fin but never enough to form distinct spots or streaks. The
scale pockets of the lateral line are each marked with a compact group of melano-
phores bisected horizontally by the lateral line pore. Similar markings may appear

120

A. J. & J. HOPSON

faintly on the scale row below the lateral line, but apart from an indistinct group of
melanophores at the base of the anal fin the ventral surface is generally pigment-free.

DIAGNOSIS AND AFFINITIES : the original description of B. nigeriensis (op. cit.}
was based on a poorly preserved holotype (Brit. Mus. (N.H.) reg. no. 1902.10.25.2)
lacking any obvious pigment. We believe that Boulenger (1911) later mistook
other superficially similar specimens for B. nigeriensis, resulting in his observations
that the markings are a " broad black lateral band which may be broken up into a
series of spots or streaks ". Such a broken band is not characteristic of B. nigeriensis
and the description has tended to confuse subsequent workers. Barbus nigeriensis
material in the British Museum (N.H.) thus included examples of 5. werneri Boulenger,
B. lawrae and B. sublineatus. The holotype is now in poor condition. Pigment is
restricted laterally to microscopic melanophores in a longitudinal line on the caudal
peduncle, grouped closer together at the posterior end to suggest a spot. Allowing
for features lost by deterioration, the holotype corresponds closely with our own
material from northern Ghana. Three fish from the Ogun River, Western Nigeria
(Brit. Mus. (N.H.) reg. no. 1956.9.6.57-59) must also be regarded as authentic
B. nigeriensis ; the pigmentation is identical with that of the specimens from Ghana.
Data fiom the three collections are tabulated below. Measurements are expressed as
percentages of the standard length. Means are given in parentheses.

Number of specimens .
Standard length (mm.)
Maximum depth
Length head*
Diameter eye*
Length snout
Interorbital width
Anterior barbel .
Posterior barbel .
Length pectoral fin
Length dorsal fin
Length caudal peduncle
Depth caudal peduncle
Number lateral line scales
Scales lateral line to dorsal

origin .
Scales lateral line to vent.

mid-line .

Holotype

Agberi River,

Ogun River,

White Volta,

Lower Niger

S. Nigeria

N. Ghana

45-50

24-50

25-2

32-2-35-0 (33-2)

24-5-29-7 (26-8)

23-7

22-7-23-7 (23-4)

24-5-3I-3 (28-7)

8-5

7-7-8-0 (7-8)

7-6-11-8 (9-9)

5-7

5-6-6-0 (5-8)

6-4-8-3 (7-5)

8- if

9-3-9-8 (9-5)

9-3-11-8 (10-4)

6'4t

7-9-8-6 (8-2)

7-4-10-0 (9-1)

7-8f

io-6-ii -o (10-8)

8-5-12-0 (10-8)

17-2

17-5-19-4 (18-6)

17-3-21-4 (19-6)

—

22-6-22-8 (22-7)

23-0-27-9 (25-0)

20-1

17-9-20-0 (19-0)

17-8-20-5 (19-3)

12-8

13-0-13-1 (13-0)

12-6-14-7 (13-7)

c. 27

28-29

27-30

4i
3i

4t
4i

4i
4i

* Negative allometry ; f somewhat shrivelled.

Barbus nigeriensis appears to have no close relatives. It resembles B. Morotaenia
Boulenger in pigmentation and barbel length but in that species the lateral line dips
less markedly below the longitudinal myoseptum and the mouth is smaller and set
more obliquely, with the lips folding characteristically round the base of the posterior

BARBUS OF THE VOLTA REGION

121

FIG. 6. Barbus nigeriensis.

barbel in the form of a socket. The combination of relatively large eyes, large mouth,
long barbels and the narrow and continuous lateral band widening on the caudal
peduncle should prevent the confusion of B. nigeriensis with other species.

HABITAT AND DISTRIBUTION : in Ghana, an uncommon species found in small
numbers in the White Volta and its tributaries in north-eastern Ghana. All examples
were from running water. Elsewhere, the Agberi River, a tributary of the Lower
Niger (type locality) and the Ogun River, both in south Nigeria.

Barbus lawrae Hopson, 1965.

(Text-fig. 7)
Barbus lawrae Hopson, 1965, Rev. Zool. Bot. Afr. 71 : 245.

HOLOTYPE : a female of 42 mm. S.L. from Lissa Dam on the Kamba River (a
tributary of the Black Volta), 15 miles north-east of Lawra, north-west Ghana,
25.x.6i, British Museum (N.H.) reg. no. 1964.9.8.1.

DESCRIPTION based on the holotype and 23 of the paratypes, 32-47 mm. S.L., from
Lissa Dam. Body compressed, particularly near the dorsal surface. S.L. 2-95 to
3-65 times the maximum depth and 3-4 to 3-75 times the length of the head. Pre-
dorsal profile convex with a slight nuchal hump. Snout rather pointed, 0-55 to 0-75
times the diameter of the eye. Mouth moderate, subterminal. Anterior barbel
o-i to 0-4 times the diameter of the eye, the tip barely overlapping the base of the
posterior barbel. Posterior barbel 0-3 to 0-8 times the diameter of the eye, the tip
reaching the vertical to the posterior margin of the pupil in the largest individuals.

Eyes inclined slightly upwards, 2-75 to 3-1 times in the length of the head and i-o
to i-i times in the interorbital width. Weakly-developed pit-lines visible on the
cheeks. Pectoral fin 0-7 to 0-8 times as long as the head, the tip falling short of the
pelvic origin. Last simple dorsal ray smooth, flexible, rather long, i-o to 1-2 times
the length of the head. Apex of the dorsal fin slightly rounded, distalmargin concave.
Dorsal fin rays III 8. Anal fin rays III 5. The origin of the pelvic fin lies between

122

A. J. & J. HOPSON

the verticals to the first and second branched dorsal rays. Caudal peduncle 1-3 to
1-6 times as long as deep. The lateral line dips to a maximum of the depth of one
and a third scale rows below the horizontal myoseptum in the anterior third of the
body ; the two converge in the centre of the caudal peduncle. Scales radially
striated. Lateral line scales 28 to 31 (modal number 30). Four and a half to 5^
scales between the lateral line and the dorsal origin, 4^ scales between the lateral
line and the mid-ventral line immediately in front of the pelvic origin and 2\ scales
between the lateral line and the pelvic origin. Twelve scales round the caudal
peduncle.

10mm.

FIG. 7. Barbus lawrae.

Summary of morphometric data based on 24 fish, 32 to 47 mm. S.L. ; measurements
are expressed as percentages of S.L.

COLORATION : in living specimens brownish above, silvery on the flanks and
whitish below. A dark mid-lateral band, often broken up into spots or streaks is
present. No conspicuous colour was noted on the fins. In formalin-fixed specimens

Range

Mean

Allometry

27-2-33-3
26-6-29-3
9-1-10-0

30-3
27-6

9-7

Negative
Negative

9-4-10-9
5-7-7-4

IO-I

6-5

—

1-2-3-9

2-2

—

3-3-7-8

5'5

—

19-5-22-4
28-4-31-6

20-8
29-5

—

19-3-24-0
I3-9-I5-8

21 -O
14-6

— .

BARBUS OF THE VOLTA REGION 123

each scale of the dorsal surface is marked with a broad band of fine melanophores,
free from a narrow marginal zone and from the centre of each scale. These markings
fade away quickly towards the flanks and a broad zone above the mid-lateral band
is more or less pigment-free. The mid-lateral band consists of heavy pigment,
approximately one third of a scale row in depth, originating at the level of the sixth
or seventh lateral line scale and running along the horizontal myoseptum to the tip
of the caudal peduncle. The band is frequently broken up into a series of rounded
spots or streaks, connected by more lightly scattered pigment. Spots, when present,
are irregular in arrangement and number ; only the first, at the level of the sixth or
seventh lateral line scale, and the last, on the tip of the caudal peduncle, are fixed in
position. A few scattered melanophores lie on the horizontal myoseptum anterior to
the mid-lateral band. Each pocket of the anterior lateral line scales may bear a
small group of melanophores, divided horizontally into two by the lateral line pore.
There are several indistinct spots of black pigment at the base of the anal fin.
Relatively dense pigment is present on the posterior margin of the second simple
dorsal ray and on the distal half of the third simple ray. Scattered melanophores
are present on the distal parts of all branched dorsal rays.

DIAGNOSIS AND AFFINITIES : material identified as B. nigeriensis from the R.
Katagum (Chad basin), Northern Nigeria in the British Museum (N.H.) reg. no.
1952.4.28.136-144, is now referred to B. lawrae. The Nigerian specimens differ
from the types in having slightly fewer lateral line scales (28 to 31, mode 29 cf. 29 to
31, mode 30) but are in other respects identical.

Barbus lawrae most closely resembles B. werneri, sewsw Daget, 1954 (from the Middle
Niger) but is more compressed laterally, has more lateral line scales (28 to 31, mode
29 and 30 cf. 25 to 29, mode 26 and 27) and much shorter barbels ; in B. lawrae the
anterior barbel barely extends beyond the base of the posterior barbel, whereas in
B. werneri, sensu Daget, the tip reaches the anterior margin of the pupil. Furthermore,
a heavier and more pronounced spot marks the tip of the caudal peduncle in B.
werneri, sensu Daget.

During the present studies, specimens of B. werneri, sensu Daget, 1954 from the
Middle Niger (Paris Museum reg. no. 51-246) were compared with a syntype of B.
werneri Boulenger from Fashoda on the White Nile (Brit. Mus. (N.H.) reg. no.
1907.12.2.1333). They were found to be a distinct species. Barbus werneri
BouJenger is easily distinguishable from 5. werneri, sensu Daget, and from B. lawrae
by the small terminal mouth, set very obliquely with the angle in front of the vertical
to the nostril. In B. werneri, sensu Daget, and in B. lawrae the mouth is subterminal
and set more horizontally with the angle overlapping the vertical to the nostril.
Barbus werneri Boulenger is unusual in the form of the band of scattered melano-
phores on the line of the mid-lateral row of spots ; the band veers upwards on the
caudal peduncle to end slightly above the terminal spot. In B. werneri, sensu Daget,
and B. lawrae the band merges horizontally with the terminal spot.

Barbus lawrae closely resembles B. lepidus in form and pigment pattern. The
mouth of B. lepidus is however more terminal, the barbels relatively longer and the
body less compressed laterally. In B. lepidus the lateral line dips further below the

124 A. J. & J. HOPSON

horizontal myoseptum and the two converge near the tip of the caudal peduncle (cf .
the centre of the caudal peduncle in B. lawrae}. The lateral spots of B. kpidus,
although variable in number, are always rounded and discrete ; spots when present
in B. lawrae are irregular and fragmented. The terminal spot in B. lawrae is never
as pronounced as in B. lepidus.

Morphometrically, B. lawrae is also similar to B. perince with which it occurs in
rivers of the Chad basin in Northern Nigeria (pers. record). Barbus perince is easily
distinguished from B. lawrae by the three rounded, black, mid-lateral spots, one in
the anterior third of the body, one below the last dorsal ray and one on the end of the
caudal peduncle. Barbus perince also has longer barbels than B. lawrae (the anterior
extending to the margin of the eye in B. perince) and the lateral line and horizontal
myoseptum converge near the tip of the caudal peduncle (cf. the centre in B. lawrae}.

DISTRIBUTION : in Ghana, probably rare and local ; known only from the type
locality in the Volta basin near Lawra. Elsewhere, rivers of the Chad basin in
Northern Nigeria (pers. records).

Barbus subinensis Hopson, 1965

(Text-fig. 8)
Barbus subinensis Hopson, 1965, Rev. Zool. Bot. Afr. 71: 249.

HOLOTYPE : a female of 33 mm. S.L. collected from the River Subin (Prah basin),
Juaso, south Ghana, g.xi.Gi, British Museum (N.H.) reg. no. 1964.9.8.256.

DESCRIPTION based on the holotype and 19 paratypes, 25-34 mm- S.L. from the
River Subin, Juaso. Body somewhat compressed. S.L. 3-1 to 3-9 times the
maximum depth and 3-4 to 4-1 times the length of the head. Predorsal profile
smooth, convex, becoming abruptly steep on the snout. Snout 0-6 to 0-75 times the
diameter of the eye. Mouth moderate, subterminal. Barbels small, the anterior
o-i to 0-3 times and the posterior 0-2 to 0-4 times the diameter of the eye. The tip
of the anterior barbel falls short of the base of the posterior barbel which extends
posteriorly slightly beyond the vertical to the anterior margin of the pupil. Eyes
lateral 2-65 to 3-05 times in the length of the head and 1-03 to 1-16 times in the
interorbital width. Weakly-developed pit-lines are visible on the cheeks and oper-
culum. Pectoral fin 0-7 to 0-85 times the length of the head, the tip overlapping the
pelvic origin in three smaller individuals. Last simple dorsal ray smooth flexible,
0-95 to i-i times the length of the head. Apex of dorsal fin somewhat pointed, distal
margin slightly concave. Dorsal fin rays III 8 ; anal fin rays III 5. The pelvic
origin lies on the vertical to the last simple dorsal ray. Caudal peduncle slender, 1-45
to 1-9 times as long as deep. The lateral line dips to a maximum depth of one and a
quarter scale rows below the horizontal myoseptum in the anterior third of the body ;
the two converge in the anterior half of the caudal peduncle. Scales radially
striated, 27 to 30 (modal number 28) in the lateral line. Five and a half scales
between the lateral l;ne and the dorsal origin, 4 to 4^ scales between the lateral line
and the mid-ventral line immediately in front of the pelvic origin and 2\ between

BARBUS OF THE VOLTA REGION

125

I Omm.

FIG. 8. Barbus subinensis.

the lateral line and the pelvic origin. Nine (f.i), 10 (f.5), n (f.i2) or 12 (f.2) scales
round the caudal peduncle.

Summary of morphometric data based on 20 fish, 25 to 34 mm. S.L. ; measurements
are expressed as percentages of S.L.

COLORATION : in formalin-fixed specimens a band of melanophores close to the
margin of each scale forms a regular pattern over the dorsal surface of the body,
fading abruptly on the flanks. A narrow continuous mid-lateral band of black pig-
ment is always present. The band, which originates on the operculum runs below the
myoseptum in the anterior half of the body ; the band and the myoseptum converge
between the verticals to the last dorsal ray and the anal origin and run together onto
the caudal peduncle. The ventral surface is pigment-free with the exception of a
group of melanophores at the base of the anal fin which continues backwards as a
narrow stripe along the ventral surface of the caudal peduncle. Snout with a con-
spicuous dark band running from eye to eye round the tip (cf. B. punctitaeniatus).
Scattered melanophores noticeable on the anterior rays of the dorsal fin.

Range

Mean

Allometry

25-6-32-2

28-9

—

24 -0-29 -2

27-1

Negative

8-5-10-4

9-4

— •

Q-6-II -2

10-3

—

5-9-7-4

6-4

—

1-2-3.0

2-1

—

2-0-3-9

3-o

—

19-4-24-6

21-3

—

25-0-30-0

27-0

—

21 -O-25 -O

22-5

• —

12 -4-15-2

13-9

—

126

A. J . & J. HOPSON

DISTRIBUTION : known only from the River Subin (Prah basin), near Juaso, south
Ghana.

REMARKS : Barbus subinensis appears to have no close relatives. The lateral
band and the markings on the snout recall B. boboi Schultz but the types of that
species bear a large black spot on the caudal peduncle, have considerably longer
barbels (anterior barbels 11-2-12-8 cf. 1-2-3-0% S.L. ; posterior barbels 11-8-13-0 cf.
2-0-3-9% S.L.) and only 4! scales above the lateral line compared with 5^ in B.
subinensis.

Barbus trispilus (Bleeker, 1863)
(Text-fig. 9)

Puntius (Barbodes) trispilos Bleeker, 1863, Nat. Verh. Wet. Haarlam, 23 : 113, pi. 23, fig. 3.

Barbus trispilus : Boulenger, 1910, Cat. Afr. Fish. 2 : 163, fig. 141.

Barbus trispilus : Daget, 1952, Mem. Inst. France. Afr. Noire, no. 19 : 314, fig. 4.

DESCRIPTION based on 24 fish, 27-65 mm. S.L. from the River Weiwei (Prah Basin),
Kumasi, south Ghana. Body slightly compressed. S.L. 3-0 to 3-75 times the
maximum depth and 3-3 to 3-85 times the length of the head. Predorsal profile
convex with a slight nuchal hump in larger individuals (over c. 40 mm. S.L.). Snout
rounded, 0-65 to 0-9 times the diameter of the eye. Mouth moderate, subterminal.
Barbels relatively long, the anterior 1-05 to 1-55 times the diameter of the eye, the
tip usually extending beyond the vertical to the posterior margin of the eye . Posterior
barbel 1-5 to 1-8 times the diameter of the eye, the tip extending beyond the posterior
margin of the preoperculum. Eyes inclined slightly upwards, the diameter 2-9 to 3-6
times in the length of the head and 1-15 to 1-65 times in the interorbital width.
Weakly-developed pit-lines present on the cheek and operculum. Pectoral fin 0-6 to
0-85 times the length of the head, the tip falling short of the pelvic origin Last
simple dorsal ray smooth, flexible, 0-75 to 0-95 times the length of the head. Apex
of dorsal fin slightly rounded, distal margin concave. Dorsal fin rays III 8. Anal

I 10mm. i

FIG. 9. Barbus trispilus.

BARBUS OF THE VOLTA REGION

127

fin rays III 5. The pelvic origin lies between the verticals to the last simple and
first branched dorsal rays. Caudal peduncle 1-2 to 1-5 times as long as deep. The
lateral line dips to a maximum of the depth of slightly over a scale row below the
horizontal myoseptum in the anterior third of the body ; the two converge in the
first third of the caudal peduncle. Lateral line scales 26 to 29 (modal number 28).
Four and a half scales between the lateral line and the dorsal origin, 4^ to 5| scales
between the lateral line and the mid-ventral line, immediately in front of the pelvic
origin and 2| scales between the lateral line and the pelvic origin. Twelve scales
round the caudal peduncle.

Summary of morphometric data based on 24 fish, 27 to 65 mm. S.L. ; measurements
are expressed as percentages of S.L.

Maximum depth .
Length head
Diameter eye
Interorbital width
Length snout
Length anterior barbel .
Length posterior barbel
Length pectoral fin
Length dorsal fin .
Length caudal peduncle .
Depth caudal peduncle .

COLORATION : in formalin-fixed specimens, dorsal surface with a dense peppering
of fine melanophores, more scattered towards the centre and free from a narrow zone
near the margin of each scale. The lateral line scale pockets are each marked with a
small, dark, triangular patch bisected horizontally by the lateral line pore. Similar
but lighter markings are present on most scale rows above the lateral line and on the
anterior and posterior scales of the row below. Three round, black, mid-lateral spots
are always present, the first at the level of the sixth or seventh lateral line scale, the
second immediately behind the vertical to the last dorsal ray and the third on the tip
of the caudal peduncle. The second spot is slightly larger than the first and last
spots which are approximately the depth of a scale row in diameter. Slight elonga-
tion of the second and third spots was noted in a few specimens. Scattered melano-
phores form a faint band over the horizontal myoseptum between the spots. The
base of the anal fin is marked with a few indistinct melanophores.

DIAGNOSIS : these data are in close agreement with previous descriptions of the
species. Barbus trispilus is superficially similar to B. perince (from the Nile and
Chad basins) which is also marked with three round mid-lateral spots. Barbus
trispilus has, however, fewer lateral line scales (26-29 cf. 28-32 in B. perince), a
lateral line which dips less markedly below the horizontal myoseptum (i cf. i| scale
rows) and relatively longer barbels (the anterior reaching the posterior margin of the
eye cf. the anterior margin in B. perince). The lateral spots are relatively larger in
B. trispilus.

Range

Mean

Allometry

26-2-32 -9

29-3

• —

25-8-30-0

27-7

Negative

7-4-10-0

8-8

Negative

IO-2-I2 -O

10-8

—

5-6-7-4

6-6

—

9-8-12-7

ii • i

—

11-7-15-4

13-8

— •

18-5-23-4

20-5

—

20-4-26-4

24-1

• —

17-4-22-6

20 -o

— •

13-6-15-6

14-8

—

128

A. J . & J . HUPSON

DISTRIBUTION : in Ghana, widespread and common in forest streams and rivers
(notably the Prah basin) in the south-west. Probably absent from the Volta.
Elsewhere Dabo Crom, Guinea (type locality) ; Sierra Leone (Norman, 1932) ;
Bolor River, Liberia (Schultz , 1942) ; Mt. Nimba, Rep. Guinea (Daget, 1952).

Barbus sublineatus Daget, 1954
(Text-fig. 10)

Barbus sublineatus Daget, 1954, Mem. Inst. France. Afr. Noire, no. 36 : 203, fig. 69.
Barbus sublineatus : Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 137, figs. 15
and 1 6.

DESCRIPTION based on 24 fish 28-40 mm. S.L. from a tributary of the White
Volta, 5 miles north of Bawku, north Ghana. Body slightly compressed. S.L.
3-8 to 4-4 times the maximum depth and 3-4 to 3-85 times the length of the head.
Predorsal profile convex with a slight nuchal hump. Snout rather pointed 0-6 to
0-8 times as long as the diameter of the eye. Mouth moderate, sub-terminal.
Barbels relatively long, the anterior 0-5 to 0-95 and the posterior 0-75 to 1-12 times the
diameter of the eye. The tip of the anterior barbel normally extends beyond the
centre and the posterior barbel beyond the posterior margin of the eye. Eyes
inclined slightly upwards, relatively large, 2-65 to 3-1 times in the length of the head
and 1-03 to 1-17 times in the interorbital width. Weakly-developed pit-lines are
present on the cheek and operculum. Pectoral fin 0-65 to 0-75 times the length of
the head, the tip falling short of the pelvic origin. Last simple dorsal ray smooth,
flexible, 0-75 to 0-95 times as long as the head. Apex of dorsal fin slightly rounded,
distal margin concave. Dorsal fin rays III 8 ; anal fin rays III 5. The origin of the
pelvic fin lies between the verticals to the first and second branched dorsal rays.
Caudal peduncle 1-35 to 1-7 times as long as deep. The lateral line dips to a maximum

lOmm.

FIG. 10. Barbus sublineatus.

BARB US OF THE VOLTA REGION 129

of approximately the depth of one scale row below the horizontal myoseptum in the
anterior third of the body ; the two converge in the vertical to the posterior anal rays
or in the anterior third of the caudal peduncle. Lateral line scales 27 to 31 (modal
number 29) . Four and a half scales between the lateral line and the dorsal origin, 4!
scales between the lateral line and the mid-ventral line immediately in front of the
pelvic origin and 2^ scales between the lateral line and the pelvic origin. Twelve
scales round the caudal peduncle.

Summary of morphometric data based on 24 fish, 28 to 40 mm. S.L. ; measurements
are expressed as percentages of S.L.

Range Mean Allometry

Maximum depth . . . 22-5-26-3 24-5

Length head . . . 25 -8-29 -2 27-4

Diameter eye . . . 8-7-10-3 9-5 Negative

Interorbital width . . 9 -7-11 -5 10-5

Length snout . . . 5 • 8-7 • o 6-5

Length anterior barbel . 5 • 3-9 • i 7-3

Length posterior barbel . 7 • 8-1 1-2 9-2

Length pectoral fin . . 18-7-20-3 19-5

Length dorsal fin . . . 22-5-25-3 23-9

Length caudal peduncle . 19-0-22-4 20-7

Depth caudal peduncle . . 12-0-14-3 13-8

COLORATION : in living specimens, yellowish fawn above, flanks silvery, belly
whitish, the body marked laterally with a row of four or more dark spots. Rays of
caudal fin with pale pink chromatophores, yellowish nearer the base ; dorsal and
pectoral rays with pale yellowish chromatophores ; pelvic and anal fins clear. The
iris is marked dorsally with an orange-red crescent. In formalin-fixed specimens
dorsal surface finely peppered with melanophores free from the margin and thinning
out towards the centre of each scale; the pattern fades out on more lateral rows. A
basic series of four black rounded spots lies over the horizontal myoseptum, the first
at the level of the seventh lateral line scale, the second below the last dorsal ray, the
third starting above the last anal ray and the fourth on the tip of the caudal peduncle.
The last three are usually slightly elongated. Additional spots up to four in number
may occur, usually between the second and the fourth spot. Occasionally two or
more of the posterior spots may coalesce to form a band. Each lateral line scale is
marked on the pocket with a small compact group of melanophores. Similar pigment
though less heavy is present on scales above the lateral line. The ventral surface is
generally pigment-tree with the exception of a faint dark spot at the base of the anal
fin continuing backwards as a slender line along the ventral surface of the caudal
peduncle.

DIAGNOSIS AND AFFINITIES : Blanc & Daget (1959) note variation in the pigmenta-
tion of B. sublineatus. Type specimens from the Middle Niger are marked with a
dark continuous mid-lateral band. The band is represented in more recent material
from the Volta and from the Haute Comoe (Blanc & Daget, 1957) by a series of spots
comparable with the markings of the present material from Ghana. In other

130 A. J. & J . HOPSON

respects our data agree closely (with allowances for allometry) with the type descrip-
tion. Barbus sublineatus is closely related to B. lineomaculatus Blgr. which occurs
in East and South Africa. Barbus lineomaculatus is similarly marked with a series of
lateral spots but never as few as four. In this species most of the first spot lies above
the horizontal myoseptum whereas in B. sublineatus it is situated chiefly below.
Barbus sublineatus also has relatively larger eyes, a narrower interorbital space and
the dorsal fin is inserted more posteriorly.

HABITAT AND DISTRIBUTION : in Ghana, widespread in the north, B. sublineatus
occurs sparingly in most of the rivers and larger streams over a substrate of gravel or
sand. It is apparently absent from lagoons, pools and dams. Elsewhere, recorded
from the Haute Comoe (Blanc & Daget, 1957) and the Middle Niger (Daget, 1954).

LIFE HISTORY : the occurrence of ripe females in June indicates that, as in other
Barbus, spawning takes place during the rainy season. Females grow larger than
males. Maximum size— a female of 74 mm. S.L.

Barbus macinensis Daget, 1954
(Text-fig, n)

Barbus macinensis Daget, 1954, Mem. Inst, France. Afr. Noire, no 36 : 200, fig. 67.
Barbus macinensis : Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 137.

DESCRIPTION based on 25 fish 26-32 mm. S.L. from the River Nahau, 5 miles north
of Bawku, northern Ghana. Body slightly compressed. S.L. 3-75 to 4-6 times the
maximum depth and 3-55 to 4-05 times the length of the head. Predorsal profile
well humped; snout rounded, 0-65 to 0-8 times the diameter of the eye. Mouth
moderate, subterminal. Anterior barbel 0-65 to 0-8 times the diameter of the eye,
its tip reaching the vertical to the anterior half of the pupil. Posterior barbel 0-8 to
i-2 times the diameter of the eye, the tip usually extending beyond the vertical to
the posterior margin. Eyes inclined slightly upwards, diameter 2-85 to 3-55 times
in the length of the head and 1-2 to 1-35 times in the interorbital width. Head with
weakly-developed pit-lines on the cheek and operculum. Pectoral fin 0-65 to 0-8
times the length of the head, its tip falling short of the pelvic origin. Last simple
dorsal ray smooth, flexible, 0-8 to i-o times as long as the head. Apex of dorsal fin
rounded, distal margin barely concave. Dorsal fin rays III 8 ; anal fin rays III 5.
The origin of the pelvic fin lies between the vertical to the ist and 2nd simple rays.
Caudal peduncle rather stout, 1-15 to 1-5 times as long as deep. The lateral line
dips to a maximum of slightly less than a scale row's depth below the horizontal
myoseptum in the anterior third of the body. The two converge in the anterior half
of the caudal peduncle. Lateral line scales 24 to 27 (modal numbers 25 and 26).
Three and a half scales between the lateral line and the dorsal origin, 4^ scales bet-
ween the lateral line and the mid-ventral line immediately in front of the pelvic
origin and 2\ scales between the lateral line and the pelvic origin. Ten (f.i), n (f.i5)
or 12 (f.g) scales round the caudal peduncle.

BARBUS OF THE VOLTA REGION

10mm.

Range

Mean

Allometry

24-7-27-8

26-2

— •

21 -4-26-6

24-6

—

7-8-9-2

8-4

Negative

10-3-11-8

10-9

Negative

5-7-6-7

6-2

—

3-5-5-9

4-9

—

6-8-10-3

8-4

—

17-9-22-9

19-9

—

22-4-26-3

24-3

—

18-2-21-8

2O -2

—

14-4-15-9

I5-I

—

FIG. ii. Barbus macinensis.

Summary of morphometric data based on 25 fish, 26 to 32 mm. S.L. ; measurements
are expressed as percentages of S.L.

COLORATION : in living specimens, dark brownish-yellow above, silvery on the
sides, pearly white below. Body marked dorsolaterally with three dark longitudinal
stripes. Three lateral spots are present and the lateral line bears a series of small
black dots. Basal two-thirds of the caudal fin, the dorsal, pectoral and pelvic fins
pale straw-yellow. Distal third of the caudal and the anal fin clear. The iris is
marked dorsal] y with an orange-red cresent. In formalin-fixed specimens body
marked dorsolaterally on each side with three dark bands running along the over-
lapping portion of adjacent scale rows and formed by a concentration of melanophores
in the upper and lower third of each scale. The two upper bands extend along the
entire length of their respective scale rows but the lowest, lying between the first and
second rows of scales above the lateral line, ends in front of the vertical to the dorsal
fin. The lowest band is frequently ill-defined and sometimes absent. One or two
round dark spots are often present on the dorsal mid-line between the head and dorsal
fin. Each scale of the lateral line, including the two overlapping the base of the
caudal fin, is marked with a small but heavy concentration of melanophores, usually
divided horizontally into two by the later? Uine pore. Three conspicuous black spots

ZOOL. 13. 4 6

132 A. J. & J. HOPSON

are constantly present on the horizontal myoseptum, the first at the level of the 7th
lateral line scale, the second below the last dorsal fin ray and the third at the tip of
the caudal peduncle. The second spot is somewhat elongated. Faint melanophores
occasionally form a noticeable band between the second and the last spots. Scales
of the row below the lateral line are often outlined with fine pigment and there is
always a faint subepidermal group of melanophores at the base of the anal fin. The
remainder of the ventral surface is pigment-free.

DIAGNOSIS AND AFFINITIES : stumpy and thick-set in appearance, B. macinensis
should be easily distinguishable from other voltaic species by the characteristic
pattern of pigmentation. Our data are in close accordance with the original descrip-
tion (Daget, 1954) based on specimens from the Middle Niger. Barbus macinensis
has close affinities with the morphometricaUy similar B. niokoloensis Daget, 1959
from the Gambia River, which is also marked longitudinally with dark dorsolateral
bands. That species, however, lacks the three lateral spots of B. macinensis and the
lateral line scales bear a dark stripe, extending forward onto the operculum.

Barbus macinensis is superficially similar to B. neefi Greenwood (1962) from the
Upper Zambesi which is also marked with dorsolateral stripes and lateral spots.
Barbus neefi, however, has more than three spots and in addition, horizontal stripes
below the lateral line. Compared with B. neefi, B. macinensis has fewer scales
between the lateral line and the dorsal origin (3^ cf. 4^-5^), shorter barbels (anterior
3'5-5'9 cf- 6-0-10-3% S.L. ; posterior 6-8-10-3 cf. 10-0-14-8% S.L. in B. neefi} and
a stouter caudal peduncle.

A comparison is made between B. macinensis and B. voltae in the description of
that species.

HABITAT AND DISTRIBUTION : in Ghana, B. macinensis is present in suitable localities
throughout the northern regions. The distribution suggests a preference for coarse
substrates and the species is particularly abundant in the sandy pools of dry season
river beds. Dams are sometimes colonized if there is a sandy or gravelly bottom
but the species is generally absent over muddy substrates and from areas of thick
vegetation. Elsewhere, recorded from the Haute Comoe (Blanc & Daget, 1957) and
from the Middle Niger (type locality). Blanc & Daget (op. cit.} note this species in
the Volta basin, Haute Volta.

LIFE HISTORY : Barbus macinensis breeds during the rainy season. Ripe and
ripening females were observed between May and August and the first juvenile stages
appeared in the rivers during July. Growth seems to be more rapid in females
than in males. All fish ripen at the end of their first year and few if any survive to
spawn a second time. Maximum size, a female of 5-9 cm. S.L.

Barbus voltae Hopson, 1965
(Text-fig. 12)

Barbus voltae Hopson, 1965, Rev. Zool. Bot. Afr. 71: 251

Barbus nigeriensis : Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 135, fig. 14.

HOLOTYPE : a female of 31 mm. S.L. from the River Nahau (tributary of the

BARB US OF THE VOLTA REGION

133

White Volta), 5 miles north of Bawku, northern Ghana, 27.xi.6i, British Museum
(N.H.) reg. no. 1964.9.8.73.

DESCRIPTION based on the holotype and 23 paratypes, 27 to 33 mm. S.L. from the
River Nahau, near Bawku. Unfortunately we have been unable to examine the
material identified by Blanc and Daget (op. cit.} as B. nigeriensis. Body somewhat
compressed. S.L. 3-3 to 3-7 times the maximum depth and 3-25 to 3-7 times the
length of the head. Predorsal profile smooth, convex. Snout bluntly pointed,
0-55 to 0-7 times the diameter of the eye. Mouth small, subterminal, protrusible and
opening obliquely downwards. Anterior barbel o-i to 0-25 times the diameter of the
eye, the tip falling short of the base of the posterior barbel. Posterior barbel 0-15
to 0-4 times the diameter of the eye, the tip extending to between the verticals to the
anterior margin of the eye and the anterior margin of the pupil. Eyes lateral, 3-35
to 3-7 times in the length of the head and i-o to 1-25 times in the interorbital width.
Weakly-developed pit-lines present on the cheek and operculum. Pectoral fin 0-55 to
0-8 times the length of the head, the tip falling short of the pelvic origin. Last
simple dorsal ray, smooth, flexible, 0-9 to 1-05 times as long as the head. Apex of
dorsal fin slightly rounded, distal margin concave. Dorsal fin rays III 7 (f.i) or
III 8 (f.23). Anal fin rays III 5. The origin of the pelvic fin lies in the vertical to
the first branched dorsal ray. Caudal peduncle 1-35 to 1-55 times as long as deep.
The lateral line dips to a maximum of slightly less than the depth of a scale row
below the horizontal myoseptum in the anterior third of the body ; the two converge
in the anterior third of the caudal peduncle. Scales radially striated, 25 to 29 (modal
number 27) in the lateral line. Three and a half scales between the lateral line and
the dorsal origin, 3! scales between the lateral line and the mid-ventral line immedi-
ately in front of the pelvic origin, and 2 to 2\ scales between the lateral line and the
pelvic origin. Eleven (f.i) or 12 (f.23) scales round the caudal peduncle.

lOmm.

FIG. 12. Barbus voltae.

ZOOL. 13, 4

134 A. J.&J.HOPSON

Summary of morphometric data based on 24 fish, 27 to 33 mm. S.L. ; measurements
are expressed as percentages of S.L.

Range Mean Allomctry

Maximum depth . . . 26 -8-29 -6 27-8

Length head . . . 26-7-30-6 28-2 Negative

Diameter eye . . . 9-4-11-1 10-0 Negative

Interorbital width . 9 • 6-1 2-9 11-2

Length snout . . . 5-8-7-3 6-4

Length anterior barbel . . i • 0-2 -6 i • 8

Length posterior barbel . . i • 6-3 -7 2-6

Length pectoral fin . . 16-5-22-5 19-8

Length dorsal fin . . . 25-3-31-1 27-7

Length caudal peduncle. . 20-0-25-0 21-5

Depth caudal peduncle . . 13-6-15-5 14-5

COLORATION : in living specimens brownish above, brilliant silver on the flanks,
whitish below. The body is marked mid-laterally with three small black spots ; a
fourth spot is present at the base of the anal fin. Caudal fin sandy-fawn, dorsal fin
pinkish-fawn, base of pectoral fin rays pale yellow. The remaining fins are colour-
less. Iris, red above. In formalin-fixed specimens the dorsal surface is peppered
with fine melanophores, free from the centre and margin of each scale. The pigment
is heaviest on the scales at the base of the dorsal fin and fades away on the second
row of scales above the lateral line. A moderately heavy band of melanophores,
approximately half a scale row in depth, runs along the horizontal myoseptum from
the operculum to end in a small, round, black spot on the tip of the caudal peduncle.
Two other mid-lateral spots are always present. They are small, usually slightly
elongated and lie within the lateral band, the first at the level of the sixth or seventh
lateral line scale and the second below the last dorsal fin ray. Although the band
varies in intensity and is sometimes broken into a series of irregular streaks, the three
spots are always discernible. Each lateral line scale is marked on the pocket with a
small but conspicuous group of melanophores, bisected horizontally by the lateral
line pore. Similar but lighter markings often appear on the first and second row of
scales above the lateral line. A conspicuous black spot is always present on the body
at the base of the anal fin.

DIAGNOSIS AND AFFINITIES : Barbus voltae is the species described by Blanc &
Daget (1957) as B. nigeriensis. Barbus voltae differs from B. nigeriensis Boulenger
in a number of important details : fewer scales between the lateral line and the dorsal
origin (3^- cf. 4! in B. nigeriensis), considerably shorter barbels (anterior 1-0-2-6 cf.
6-4-10-0% S.L. ; posterior 1-6-3-7 cf- 7'8-i2-o% S.L. in B. nigeriensis), smaller eyes
and a much smaller mouth. Furthermore, B. nigeriensis Blgr. has no trace of lateral
spots which are always present in B. voltae.

Barbus voltae is superficially similar to B. macinensis with which it frequently
occurs in the Volta basin. It is, however, easily distinguished by the smooth pre-
dorsal profile (humped in B. macinensis), shorter barbels (anterior i- 0-2-6 cf. 3-5-
5-9% S.L. ; posterior 1-6-3-7 c^- 6-8-10-3% S.L.), a smaller mouth, larger eyes

BARB US OF THE VOLT A REGION

135

(9-4-11-1 cf. 7-8-9-2% S.L. in B. macinensis} and a more obvious spot at the base of
the anal fin. Barbus voltae also lacks the dorsolateral stripes of B. macinensis.

HABITAT AND DISTRIBUTION : in Ghana, common in rivers, streams and dams
throughout the northern regions. Infrequent in well-vegetated situations. Barbus
voltae is more plentiful over muddy than over sandy bottoms, indicating a preference
for finer substrates. Elsewhere, recorded only from the Black Volta and its tribu-
taries in Haute Volta (Blanc & Daget, 1957).

Barbus atakorensis Blanc & Daget, 1957

(Text-fig. 13)
Barbus atakorensis Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 140, fig. 17.

DESCRIPTION based on 21 fish 27-36 mm. S.L., from Dahwenia Dam, 6 miles east
of Tema, south Ghana. Body moderately compressed. S.L. 3-3 to 3-9 times the
maximum depth and 3-5 to 4-0 times the length of the head. Predorsal profile
convex with a slight nuchal hump. Snout somewhat pointed, 0-7 to 0-95 times as
long as the eye diameter. Mouth moderate, subterminal. Anterior barbel 0-25 to
0-5 times the diameter of the eye, the tip extending beyond the base of the posterior
barbel. Posterior barbel 0-6 to 0-95 times the diameter of the eye, the tip extending
to between the verticals to the posterior margin of the pupil and the posterior margin
of the eye. Eyes lateral, the diameter 3-2 to 3-6 times in the length of the head and
1-2 to 1-5 times in the interorbital width. Weakly-developed pit-lines visible on the
cheeks and operculum. Pectoral fin 0-6 to 0-8 times as long as the head, the tip
falling short of the pelvic origin. Last simple dorsal ray smooth, flexible, 0-8 to
0-95 times the length of the head. Apex of dorsal fin rounded, distal margin slightly
concave. Dorsal fin rays III 7. Anal fin rays III 5. The origin of the pelvic fin

10mm.

FIG. 13. Barbus atakorensis.

136 A. J. & J. HOPSON

lies between the vertical to the first and second branched dorsal rays. Caudal
peduncle relatively slender, 1-4 to 1-8 times as long as deep. The lateral line dips
to a maximum of the depth of one and a half rows below the horizontal myoseptum
in the anterior third of the body ; the two converge in the anterior third of the
caudal peduncle. Lateral line scales 27 to 32 (modal number 28). Four and a half
to 5 1 scales between the lateral line and the dorsal origin, 4! to 5| scales between the
lateral line and the mid-ventral line immediately in front of the pelvic origin, and 2!
to 3 scales between the lateral line and the pelvic origin ; 12 scales round the caudal
peduncle.

Summary of morphomdric data based on 21 fish, 27 to 36 mm. S.L. ; measurements
are expressed as percentages of S.L.

Range Mean Allometry

Maximum depth . . . 26 -5-30 -5 28-2

Length head . . . 24-9-27-5 26-0 Negative

Diameter eye . . . 6 • 9-8 -5 7-5 Negative

Interorbital width . . 9-7-10-9 10-3

Length snout . . . 5 • 4-6 -7 6-2

Length anterior barbel . . 2-1-3-3 2'7

Length posterior barbel . 5 • 0-6 -8 5-7

Length pectoral fin . . 17-8-20-4 19-3

Length dorsal fin . . . 20-9-24-5 23-3

Length caudal peduncle . 20-9-24-0 21-8

Depth caudal peduncle . . 13-2-15-0 14-1

COLORATION : in living specimens body greyish green, dark above, paler on the
sides and whitish on the belly. Fins colourless. The colour pattern is best seen in
formalin-fixed specimens where the dorsal surface is peppered with fine melanophores,
less heavy in the centre of each scale and absent from the free margin. The pig-
mentation becomes lighter on the flanks and fades out on the second row of scales
above the lateral line. Pockets of each lateral line scale are marked with a small
group of dense melanophores split horizontally into two by the lateral line pore.
Similar markings are repeated on the pockets of the first and sometimes the second
row of scales above the lateral line. The horizontal myoseptum, appearing as a fine
black line, is punctuated in most specimens with three black spots, the first at the
level of the 7th lateral line scale, the second on the vertical to the last dorsal ray and
the third at the tip of the caudal peduncle, partly overlapping the base of the caudal
fin. The third spot, present in all specimens, is elongated and nearly twice as long
as deep. The other two spots are variable in size and shape and one or both may be
missing. Occasionally a fourth spot may occur immediately behind the vertical to
the last anal ray. Scattered melanophores, usually heavier in the posterior half of
the body, lie in a diffuse band over the horizontal myoseptum. Underparts unpig-
mented with the exception of a faint black spot at the base of the anal fin and a thin
dark line along the ventral surface of the caudal peduncle.

DIAGNOSIS AND AFFINITIES : these data agree closely with the original description
(Blanc & Daget, 1957) of material from the Volta basin in north Dahomey. We

BARBUS OF THE VOLTA REGION 137

found 5| as well as 4! scale rows below the lateral line and a higher maximum
number of lateral line scales (32 cf. 29) although the modal number, 28, was the
same. The two collections differ principally in coloration. Whereas our specimens
normally have three lateral spots, the type material is characterized by a dark mid-
Jateral band, continuous from the operculum to the caudal peduncle. We noted,
however, that in the Paris Museum paratypes the band is constricted on the caudal
peduncle to form a rectangular terminal spot identical with the spot of our own
specimens. Although B. atakorensis is superficially similar to numerous species of
Barbus each with three lateral spots, we consider that it has no near relatives. The
presence of only seven branched dorsal rays, the relatively high number of scales and
the elongated spot on the slender caudal peduncle should prevent confusion with
other species.

DISTRIBUTION : in Ghana, at present known only from dams on the Accra plains
and from the River Azubone (Volta basin) in forest country near Mpraeso. The dams
are on small coastal rivers draining directly into the sea. No specimens of B.
atakorensis were collected in the northern regions of the country. Elsewhere, the
upper reaches of the River Oti (Volta basin), in north Dahomey (type locality), the
Ogun River, south-west Nigeria (personal observation), and the Kaduna River
(Niger Basin), Northern Nigeria (B.M. (N.H.) reg. no. 1936.11.24.27-29).

Barbus punctitaeniatus Daget, 1954
(Text-fig. 14)

Barbus punctitaeniatus Daget, 1954, Mem. Inst. France. Afr. Noire, no. 36 : 202, fig. 68.
Barbus punctitaeniatus : Blanc & Daget, 1957, Mem. Inst. France. Afr. Noire, no. 50 : 137.

DESCRIPTION based on 24 specimens 21-37 mm. S.L., 8 from tributaries of the
White Volta near Bawku and 16 from the Black Volta near Lawra, northern Ghana.
Body moderately compressed. S.L. 3-55 to 4-3 times the maximum depth and 3-55
to 4-1 times the length of the head. Predorsal profile convex with slight nuchal
hump, more pronounced in the largest individuals. Snout bluntly pointed, 0-55
to 0-9 times the diameter of the eye. Mouth moderate, subterminal. Barbels
relatively long, the anterior 0-4 to 0*85 times and the posterior 0-8 to 1-2 times the
diameter of the eye. Tip of the anterior barbel extending to about the vertical to
the centre of the eye and the posterior barbel extending beyond the posterior margin.
Eyes lateral, 2-75 to 3-6 times in the length of the head and i-o to 1-35 times in the
interorbital width. Cheeks and operculum with weakly-developed pit-lines.
Pectoral fin 0-65 to 0-85 times the length of the head, the tip falling short of the pelvic
origin. Last simple dorsal ray smooth, flexible, 0-8 to i-o times as long as the head.
Apex of dorsal fin slightly rounded, distal margin concave. Dorsal fin rays III 7
(f.i) or III 8 (1.23). Anal fin rays III 5. The pelvic origin lies on the vertical to
the first branched dorsal ray. Caudal peduncle 1-35 to 1-85 times as long as deep.
The lateral line incomplete in all but the three largest specimens (28-37 mm S.L.)
with 7 to 12 anterior scales perforated ; total scales in the row 23 to 27 (modal
number 25); this row dips to a maximum of slightly more than the depth of

133

A. J . cS: J . 1IOPSOX

half a scale row below the horizontal myoseptum in the anterior third of the body ;
the two converge at the anterior end of the caudal peduncle. Three and a half scales
between the lateral line and the dorsal origin, 3^ scales between the lateral line and
the mid-ventral line immediately in front of the pelvic origin and 2-2\ scales between
the lateral line and the pelvic origin. Nine (f.22) or 10 (f.2) scales round the caudal
peduncle.

FIG. 14. Barlus punctitaeniatus.

Summary of morphometric data based on 24 fish, 21-37 mm. S.L. ; measurements
are expressed as percentages of S.L.

COLORATION : in living specimens dorsal surface light fawn, silvery on the sides,
whitish below. A narrow, dark mid-lateral band runs from the tip of the snout to the
tip of the caudal peduncle. Yellow chromatophores are scattered over the dorsal
surface, on the operculum and most noticeably in a narrow zone above the lateral
band. Pectoral fins frequently tinged with yellow ; all other fins colourless. The
iris is marked dorsally with a yellow crescent. In formalin-fixed specimens pigment
on the dorsal surface consists of a relatively slender band of dense melanophores
near the margin of each scale fading abruptly on the flanks. A narrow black stripe

Range

Mean

Allometry

23-

1-28

•I

•5

—

24-

2-28

•o

•3

Negative

I-IO

•o

•7

Negative

3-i i

•o

•8

—

0-6-

•o

—

2-6-

•6

— •

O-II

•2

•8

—

18-

2-24

•2

•7

—

22'

0-25

•9

•4

• —

18-

8-23

•6

•3

—

12-

1-15

•o

•4

• — •

BARB US OF THE VOLT A REGION 139

originating on the tip of the snout (i.e. continuous round the tip from eye to eye) runs
obliquely upwards across the operculum and thence along the body. The stripe lies
slightly below the horizontal myoseptum in the anterior half of the body but the two
converge on the caudal peduncle. The stripe is intensified by small round spots of
dense pigment on the pockets of all lateral line scales. Melanophores form a faint
stripe at the base of the anal fin continuing backwards as a narrow line along the
ventral surface of the caudal peduncle.

DIAGNOSIS : our data are in close agreement with the type description (Daget,
1954) based on specimens from the Middle Niger. We examined six of the syntypes
in the Paris Museum and found that as in most of the material from Ghana, the lateral
line is incomplete, a point which Daget did not observe.

Barbus punctitaeniatus is readily distinguished from most West African species by
the black streak on the snout running without interruption round the tip. Barbus
boboi and B. subinensis are similarly marked but both have higher scale counts and a
pronounced dip in the lateral line ; in B. punctitaeniatus the dip is unusually shallow.
The differences between this species and B. bawkuensis are discussed in the description
of that species below.

DISTRIBUTION : in Ghana, widespread and reasonably common in the northern
regions, usually in rivers but occasionally in dams. Elsewhere, the Middle Niger
(Daget, 1954), the Volta basin in Haute Volta (Blanc & Daget, 1957) and rivers of
the Chad basin in Northern Nigeria (personal records) .

LIFE HISTORY : the spawning season is probably restricted to the rains. Ripe
females were noted in July and juveniles appeared in the rivers during September.
Females grow larger than males. Maximum size 37 mm. S.L.

Barbus bawkuensis Hopson, 1965

(Text-fig. 15)
Barbus bawkuensis Hopson, 1965, Rev. Zool. Bot. Afr. 71: 254

HOLOTYPE : a female of 26 mm. S.L. from the White Volta, Bazua Bridge, 10
miles west of Bawku, north Ghana, 28 . vi . 61, British Museum (N.H.) reg. no. 1964 .9.8.

278.

DESCRIPTION based on the holotype and 15 paratypes 20-29 mm- S.L. from the
White Volta, Bazua and on 17 specimens 21-26 mm. S.L. from the River Morago,
Nakpanduri, 45 miles south of Bawku. Body moderately compressed. S.L. 3-1
to 4-1 times the maximum depth and 3-55 to 4-2 times the length of the head. Pre-
dorsal profile convex with a slight nuchal hump. Snout rounded 0-65 to 0-95 times
the length of the head. Mouth moderate, slightly protrusible and subterminal.
Anterior barbel 0-25 to 0-7 times and the posterior barbel 0-6 to i-i times the diameter
of the eye. The tip of the anterior barbel usually extends to the vertical to the
anterior margin, and the tip of the posterior barbel beyond the centre of the eye. Eye
moderate 2-85 to 3-8 times in the length of the head and 1-05 to 1-4 times in the inter-

140

A . J . & J . H O P S O N

orbital width. Weakly-developed pit-lines visible on the cheeks and operculum.
Pectoral fin 0-6 to 0-8 times as long as the head, the tip falling well short of the pelvic
origin. The last simple dorsal ray smooth, flexible, 0-9 to i-i times the length of the
head. Apex of dorsal fin rather pointed, distal margin concave. Dorsal fin rays
III 8 ; anal fin rays III 5. The pelvic origin lies between the verticals to the first
and second branched dorsal rays. Caudal peduncle slender, 1-45 to 2-0 times as long
as deep. Scales radially striated, 24 to 28 in the lateral line row (modal number 26).
Lateral line incomplete, perforated scales usually consecutive, 4 to 9 in number at
the anterior end of the row. In a few individuals, however, up to 21 perforated scales
were noted ; the additional scales were not arranged consecutively but were scattered
irregularly amongst blank scales. The lateral line row dips to a maximum depth of
half a scale below the myoseptum in the anterior third of the body. There are 3^
scales between the lateral line and the dorsal origin, 3! scales between the lateral line
and the ventral midline immediately in front of the pelvic origin and 2 to 2\ scales
between the lateral line and the pelvic origin. Eight (f.i), 9 (f.24) or 10 (f.6) scales
round the caudal peduncle.

10mm.

FIG. 15. Barbus bawkuensis.

Summary of morphometric data based on 33 fish, 20 to 29 mm. S.L. ; measurements
are expressed as percentages of S.L.

Range

Mean

Allometry

24-4-31-1

27-7

—

23-8-28-5

25'9

—

7-1-9-0

7'7

—

8-9-11-5

9.9

—

5-7-6-9

6-3

—

1-9-4-6

3-o

—

4 • 4-8 • o

6-1

—

15-5-21-2

18-3

—

22-6-28-8

25-4

—

19-5-22-8

21-4

—

12-0-14-0

12-8

—

BARB US OF THE VOLTA REGION 141

COLORATION : in formalin-fixed specimens dorsal surface with a light scattering of
melanophores near the margin of each scale. A narrow, diffuse band of melano-
phores runs mid-laterally from the operculum to the end of the caudal peduncle. An
indistinct black spot lies at the base of the anal fin. The body is otherwise pigment-
free. The snout is marked laterally with a dark streak which does not extend
forwards onto the tip. Dark pigment is noticeable along the anterior margin of the
dorsal fin.

DIAGNOSIS AND AFFINITIES : morphometricalry, B. bawkuensis is very similar to
B. punctitaeniatus differing only in the shorter barbels (anterior 1-9-4-6 cf. 3-2-6-4%
S.L. ; posterior 4-4-8-8 cf. 6-0-11-2% S.L.) and in the slightly higher modal number
(26 cf. 25) of lateral line scales. Barbus bawkuensis, however, lacks the black streak
on the tip of the snout and the dark spots on the lateral line scales characteristic of
B. punctitaeniatus. Other minor differences in the newly described species are heavier
pigmentation on the last simple dorsal ray and a darker spot at the base of the anal
fin. Barbus bawkuensis and B. punctitaeniatus are remarkable for their high variability
in the number of perforated lateral line scales. This unusual feature has been noted
previously in B. anoplus (Groenewald, 1958) where, as in B. bawkuensis, the lateral
line is frequently interrupted, and also in B. pobeguini from Ksar Torchane, Maure-
tania (Daget, 1954) where some specimens had complete, otners rudimentary lateral
lines. Barbus bawkuensis and B. punctitaeniatus are unique among West African
species in the combination of a incomplete lateral line and two pairs of relatively
long barbels.

DISTRIBUTION : at present known only from the White Volta and its tributaries
near Bawku, northern Ghana.

Barbus pobeguini Pellegrin, 1911
(Text-fig. 1 6)

Barbus pobeguini Pellegrin, 1911, Bull. Soc. zool. France, 36 : 187.

Barbus pobeguini : Daget, 1954, Mem. Inst. France. Afr. Noire, no. 36 : 205, fig. 71.

DESCRIPTION based on 24 fish 29-36 mm. S.L., from Dedoro Tankara Dam, 8 miles
north-east of Navrongo (Volta basin), north Ghana. Body cylindrical, only slightly
compressed. S.L. 3-7 to 4-3 times the maximum depth and 3-5 to 3-8 times the length
of the head. Predorsal profile smooth, convex. Snout rounded, 0-85 to i-o times
the diameter of the eye. Mouth moderate, subterminal. The single pair of short
barbels o-i to 0-3 times the diameter of the eye. Eyes relatively small, inclined
slightly upwards, the diameter 3-45 to 4-0 times in the length of the head and 1-35 to
1-7 times in the interorbital width. Weakly-developed pit-lines present on the
cheeks. Pectoral fin 0-65 to 0-8 times the length of the head, the tip falling well short
of the pelvic origin. Last simple dorsal ray smooth, flexible, 0-8 to 0-9 times the
length of the head. Apex of dorsal fin rounded, distal margin slightly convex.
Dorsal fin rays III 7 ; anal fin rays III 5. The pelvic origin lies within the verticals
to the first or second dorsal rays. Caudal peduncle 1-45 to 1-75 times as long as deep.

14.! A. J . & J. HOPSON

The lateral line dips to a maximum depth of one scale row below the horizontal
myoseptum in the anterior third of the body ; the two converge in the first third of
the caudal peduncle. Twenty-seven to 32 lateral line scales (modal number 29).
Four and a half to 5| scales between the lateral line and the dorsal origin, 5^ scales
between the lateral line and the mid-ventral line immediately in front of the pelvic
origin and 2\ to 3 scales between the lateral line and the pelvic origin. Twelve (f.2i)
or 13 (f-3) scales round the caudal peduncle.

FIG. 1 6. Barbus pobeguini.

Summary of morphometric data based on 24 fish, 29 to 36 mm. S.L. ; measurements
are expressed as percentages of S.L.

Maximum depth .
Length head
Diameter eye
Interorbital width
Length snout
Length barbel
Length pectoral fin
Length dorsal fin .
Length caudal peduncle .
Depth caudal peduncle .

COLORATION : in living specimens silvery with a characteristic steel-blue iridescence.
A dark lateral band ends in a smah1 but pronounced spot on the caudal peduncle.
There is an oblique black stripe on the distal half of the anterior rays of the dorsal
fin. In formalin-fixed specimens dorsal surface well peppered with melanophores
absent from a narrow zone close to the margin and thinning out towards the centre of
each scale. Dark pigment fades out on the flanks, and scales below the lateral line
are clear. Pockets of anterior lateral line scales each with a small black streak

Range

Mean

Allometry

23-5-26-5
26-2-28-0
6-8-8-7

24-2

27-1

7-2

—

10-0-11-5
6-3-7-2

O • 7-2 • 2
I7-6-22-O

10-5

6-8

i-4
19-6

Positive

Negative

2I-5-25-5

23-4

—

21 '8-24-1

22-7

—

12 -9-15-2

13-9

—

BARBUS OF THE VOLTA REGION 143

divided horizontally into two by the lateral line pore. The upper halves of the streaks
persist on more posterior scales. A narrow black lateral band slightly below the
myoseptum extends backwards onto the caudal peduncle, stopping just short of a
round black spot on the tip. The spot overlaps the base of the caudal rays. Dorsal
fin with a narrow black stripe, parallel with the distal margin of the fin, on the upper
halves of the last simple and first three branched rays.

DIAGNOSIS : these data are in close agreement with descriptions of B. pobeguini
from the Middle Niger (Pellegrin, 1911 and Daget, 1954). Barbus pobeguini appears
to have no close relatives in West Africa. The presence of a single pair of barbels, 7
branched dorsal rays and the distinctive colour pattern should prevent confusion
with other species in the area. Pellegrin (op. cit.) has pointed out the similarity of
this species with B. afer Peters from South Africa.

HABITAT AND DISTRIBUTION : in Ghana, rivers, streams and pools throughout the
northern regions. The commonest Barbus in flood water ; often found in temporary
streams after storms and one of the first colonizers of newly built dams. Never
abundant in the larger rivers. Elsewhere, basins of the Niger (Daget, 1954) and Comoe
(Blanc & Daget, 1957) and in Mauretania (Daget, 1954).

LIFE HISTORY : Barbus pobeguini apparently breeds during the rains. Ripening
fish were observed in June and juvenile stages appeared in August. Growth is more
rapid in females than in males. Maximum size, a female 50 mm. S.L.

Barbus stigmatopygus Boulenger, 1903
(Text-fig. 17)

Barbus stigmatopygus Boulenger, 1903, Ann. Mag. nat. Hist. (7) 12 : 533 ; Idem, 1911, Cat. Afr.

Fish. 2 : 185, fig. 163.
Barbus stigmatopygus : Daget, 1954, Mem. Inst. France. Afr. Noire, no. 36 : 210, fig. 75.

DESCRIPTION based on 24 fish, 16-19 mm- S.L. from the Black Volta, 3 miles west
of Lawra, northern Ghana. Body compressed. S.L. 3-2 to 3-8 times the maximum
depth and 3-45 to 3-95 times the length of the head. Predorsal profile convex.
Snout short, bluntly pointed, 0-55 to 0-75 times in the diameter of the eye. Mouth
terminal, small. Eyes lateral, relatively large, 2-45 to 2-8 times in the length of the
head and 0-95 to 1-15 times in the interorbital width. A few weakly-developed pit-
lines present on the cheek and operculum. Barbels absent. Pectoral fin 0-6 to 0-75
times as long as the head, its tip falling short of the pelvic origin. Last simple ray of
dorsal fin smooth, flexible, 0-8 to i-o times the length of the head. Apex of dorsal fin
rounded, distal margin concave. Dorsal fin rays III 8. Anal fin rays III 5. Origin
of pelvic fin between the verticals to the last simple and first branched dorsal rays.
Caudal peduncle 1-25 to 1-5 times as long as deep. Lateral line incomplete, only the
first 4 to 7 scales in the row perforated. There are 23 to 25 (modal number 24)
scales in the lateral line row, 4! scales between the lateral line and the dorsal origin,
2|- scales between the lateral line and the mid-ventral line immediately in front of the
pelvic origin and ii scales between the lateral line and the pelvic origin. Nine (f.i)
or 10 (f.23) scales round the caudal peduncle.

144

A. J. & J. HO PS ON

10mm.

Range
26-1-31 -2
25-2-28-9
9-5-11-2
10-3-11-9

5-8-7-4
16-8-20-0
22-1-27-6

17-6-21 -2
I3-2-I5-0

Mean
28-5

27-3
10-3
u-i
6-7
18-6
24-9
19-9
14-2

Allometry

FIG. 17. Barbus stigmatopygus.

Summary of morphometric data based on 24 fish, 16 to 19 mm. S.L. ; measurements
are expressed as percentages of S.L.

Maximum depth .
Length head
Diameter eye
Interorbital width
Length snout
Length pectoral fin
Length dorsal fin .
Length caudal peduncle .
Depth caudal peduncle .

COLORATION : in living specimens dorsal surface finely patterned with yellow and
black chromatophores, scales of the lateral line and below, silvery. A round black
terminal spot on the caudal peduncle and a second spot at the base of the anal fin are
always present. Two additional spots frequently occur on the horizontal myoseptum
in the anterior half of the body. The fins are colourless. There is a yellow arc over
the pupil of the eye. Specimens from clear water are frequently transparent with the
vertebral column and viscera visible through the body wall. In formalin-fixed
specimens the scales of the dorsal surface are outlined with a fine lacework of melano-
phores which fade away on the flanks. A small round black spot at the tip of the
caudal peduncle is always present, as is a compact group of melanophores at the base
of the anal fin, which continues backwards as a thin black line along the ventral
surface of the caudal peduncle. Two further round black spots, smaller than the
one on the caudal peduncle, usually occur on the horizontal myoseptum, the first at
the level of the fifth lateral line scale, the second below the last ray of the dorsal fin.
One or both of these may be absent. Pockets of anterior lateral line scales are
frequently marked with a black dot above and below the canal.

BARBUS OF THE VOLTA REGION 145

DIAGNOSIS : these data correspond closely with previous descriptions of B.
stigmatopygus. Despite careful examination of the material described here we can
find no trace of the single pair of much reduced barbels which Dr. Trewavas recently
observed in specimens from the White Nile and Lake No (pers. com.). Barbus
stigmatopygus is easily distinguished by its markings from other small Barbus with
similarly reduced lateral lines and barbels.

HABITAT AND DISTRIBUTION : in Ghana, judging from our records, B. stigmatopygus
is a relatively rare and local species. With the exception of two fish from the River
Nasia near Tamale all records are from backwaters and tributaries of the Black Volta
in the Lawra district where it is not uncommon. The habitat is always deep,
relatively still water, free from vegetation. Elsewhere, widespread in the Sudanese
belt. Recorded from the White Nile (type locality), the Middle Niger (Daget, 1954),
Haute Volta (Blanc & Daget, 1957) and the Geba River, Portuguese Guinea (Boulen-
ger, 1911).

Barbus leonensis Boulenger, 1915

(Text-fig. 18)

Barbus leonensis Boulenger, 1915, Ann. Mag. nat. Hist. (8) 15 : 203 ; Idem, 1916, Cat. Afr. Fish.

4. : 273, fig. 167.
Barbus leonensis : Daget, 1954, Mem. Inst. France. Afr. Noire, no. 36 : 209, fig. 74.

DESCRIPTION based on 24 fish 27 to 33 mm. S.L. from the water-works reservoir,
Tamale, north Ghana (Volta basin). Body somewhat compressed. S.L. 3-1 to 3-7
times the maximum depth and 3-2 to 3-7 times the length of the head. Predorsal
profile smooth, convex. Snout pointed, 0-7 to 0-95 times the diameter of the eye.
Mouth small, slightly subterminal. Eyes lateral, 2-9 to 3-35 times in the length of

I Omm.

FIG. 1 8. Barbus leonensis.

146 A. J . & J . HOPSON

the head and 1-15 to 1-4 times in the interorbital width. Barbels absent. A few
weakly-developed pit-lines on the cheeks and on the operculum. Pectoral fin 0-55
to 07 times the length of the head, the tip falling short of the pelvic origin. The
last simple dorsal ray is smooth, flexible, 0-75 to 0-95 times as long as the head.
Apex of the dorsal fin slightly rounded, the distal margin straight. Dorsal fin rays
III 7 (f.2) or III 8 (f.22). Anal fin rays III 5 (f.2i) or 6 (f.3). The origin of the pelvic
fin lies between the verticals to the first and second dorsal rays. Caudal peduncle
1-2 to 1-5 times as long as deep. The lateral line is always incomplete ; 6 to 10
anterior scales are perforated. There are 22 to 26 scales (modal number 24) in the
lateral line scale row, 4! to 5| scales between the lateral line and the. dorsal origin,
2.\ scales between the lateral line row and the mid-ventral line immediately in front
of the pelvic origin, and i| scales between the lateral line and the pelvic origin.
Eight (f.i), 9 (f.i8) or 10 (f.5) scales round the caudal peduncle.

Summary of morphometric data based on 24 fish, 27 to 33 mm. S.L. ; measurements
are expressed as percentages of S.L.

Range Mean Allometry

Maximum depth . . . 27-0-31-8 29-3

Length head . . . 27-5-30-3 28-6

Diameter eye . . . 8 • 2-9 -6 8-9

Interorbital width . . 10-4-12-8 11-5

Length snout . . . 6 • 6-8 -2 7-3

Length pectoral fin . .16- 6-2 1 • o 1 8 • 6

Length dorsal fin . . . 23-0-26-0 23-8

Length caudal peduncle . . 18-2-21-0 19-6

Depth caudal peduncle . . 13-1-15-4 14-3

COLORATION : living specimens pale lemon yellow in colour with two rounded black
spots, one on the caudal peduncle the other on the anterior rays of the dorsal fin.
Yellow pigment brightest in an arc over the pupil of the eye and in the vicinity of
the two black spots. Living specimens frequently transparent, the vertebral column
and viscera clearly visible through the body wall. In formalin-fixed specimens dorsal
surface with an even scattering of melanophores thinning out and disappearing on
the flanks. Scales of the lateral line row and above lightly outlined with melano-
phores which are thicker in a vertical line on the pocket of each scale, thus forming a
regular pattern over the dorsal surface of the fish. Traces of the pattern may
persist on the row below the lateral line but the ventral surface is generally pigment-
free. The black spot on the caudal peduncle is equal in diameter to the depth of one
scale row and slightly overlaps the base of the caudal fin. The side of the snout is
marked with a narrow, horizontal, band of melanophores touching the anterior
margin of the eye. The spot on the dorsal fin lies between the last simple and the
third branched ray at a point midway between the base and the apex of the fin.

DIAGNOSIS : these data correspond closely with the description of the types from
Sierra Leone and also with data obtained by Daget (1954) from fish in the Middle
Niger. We, however, record 9 as the modal number of scales round the caudal
peduncle compared with 8 in the previous descriptions. Barbus leonensis is easily

BARB US OF THE VOLT A REGION 147

distinguished from B. anenia and B. stigmatopygus (both of which have incomplete
lateral lines and no barbels) by the characteristic pattern of pigmentation.

HABITAT AND DISTRIBUTION : in Ghana, widespread in well-vegetated pools,
oxbow lakes and dams throughout northern Ghana. Rare in rivers and streams.
Elsewhere, recorded from much of the savannah zone of West Africa — Sierra Leone
(type locality), Gambia River (Johnels, 1954), Niger basin (Daget, 1954), Comoe
(Blanc & Daget, 1957), Haute Volta (Blanc & Daget, op. cit.} and Chad (Daget, op.
cit.}.

LIFE HISTORY : spawning is confined to the rainy season, June to September. All
fish mature at the age of 8 to 10 months. Few individuals, if any, survive their
first spawning season and the maximum life span is probably 15 months. Growth
is slightly more rapid in females than in males. Maximum size, 33 mm. standard
length.

KEY TO THE VOLTAIC SPECIES OF Barbus

1 Well-developed pit-lines present ; individual pits bead-like in appearance, forming

parallel ridges on the side of the snout, the cheek, the operculum and on the
dorsal surface of the head between the eyes . . . subgenus Beirabarbus 2
Cephalic pits relatively few in number, sometimes orientated into lines, but with the
individual pits well-separated from one another and usually visible only on the
cheek and on the operculum .......... 3

2 Apex and second simple ray of dorsal fin dense black ; heavy melanophores form a

conspicuous mid-lateral band running from the side of the snout to the tip of the
caudal peduncle ; pockets of lateral line scales with relatively broad, triangular
spots ; 12 scales round the caudal peduncle ; the tip of the pectoral fin never
overlaps the pelvic origin ; snout 6-3-7-3% S.L. ; caudal fin in living specimens
orange-red B. macrops

No dense pigment patches on the dorsal fin ; mid-lateral band relatively light ;
pockets of lateral line scales with narrow and crescentic vertical stripes ; usually
10 or ii scales round the caudal peduncle ; the tip of the pectoral fin often over-
laps the pelvic origin ; snout 6-4-7-6% S.L. ; caudal fin pale pink in living
specimens B. parablabes

No dense black pigment patches on the dorsal fin ; mid-lateral band faint ; pockets
of lateral line scales with relatively narrow vertical bars of pigment ; 12 scales
round the caudal peduncle ; the tip of the pectoral fin sometimes overlaps the
pelvic origin ; snout 7-8-8-8% S.L. B. spurrelli

3 Lateral line scales markedly deep, over 2-7 times as deep as long ; dorsolateral scales

outlined with melanophores which are concentrated in a vertical arc on the pocket
of each scale ; mid-lateral band, if present, very faint ; an oblique black stripe
often lies between the origin of the lateral line and the base of the pectoral fin :
subgenus Clypeobarbiis ......... B. hypsolepis

Lateral line scales not markedly deep, under 2-4 times as deep as long ; pigment
pattern consisting of lateral stripes or spots ; no black stripe between the origin of
the lateral line and the base of the pectoral fin ....... 4

4 Barbels present ........ .5

Barbels absent ............ 13

5 Usually 8 branched dorsal rays ........ 6

Usually 7 branched dorsal rays . . . . . . . . . 12

i.fS A. J . & J. HO PS ON

6 Body marked with three or more mid-lateral spots ; one in the anterior third of the

body, one below the last dorsal ray and one on the caudal peduncle are nearly
always present ; the spots are sometimes partly obscured by a dark mid-lateral
band .....'......... 7

Body marked with a black mid-lateral band, sometimes broken into irregular spots or

streaks ............. 9

7 Body marked with three mid-lateral spots . . . . ... . . 8

Body marked mid-laterally with four or more large rounded black spots . B. sublineatus

8 Dorsolateral surface with two or three dark longitudinal stripes ; barbels relatively

long, the anterior clearly overlapping the base of the posterior ; mouth not
protrusible obliquely downwards ; spot at the base of the anal fin relatively
faint ............ B. macinensis

Dorsolateral surface without longitudinal stripes ; barbels short, the anterior barely
overlapping the base of the posterior ; mouth small, protrusible obliquely down-
wards ; black spot at the base of the anal fin conspicuous . . . B. voltae
g Lateral line complete ; 4! or 5^ scales between the lateral line and the dorsal origin 10

Lateral line usually incomplete ; 3 1 scales between the lateral line and the dorsal origin 1 1

10 Mid-lateral band more or less continuous from the tip of the operculum to the end of

the caudal peduncle ; mouth large, barbels long, the posterior extending to or
beyond the posterior margin of the eye ...... JB. nigeriensis

Mid-lateral band, originating between the operculum and the dorsal fin, frequently
broken into irregular spots or streaks ; mouth moderate in size ; barbels moderate,
the posterior not extending past the vertical to the posterior margin of the pupil

B. lawrae

1 1 Black pigment on the snout continuous round the tip from eye to eye ; conspicuous

black spots on the scale pockets of the lateral line B. punctitaeniatus

Black pigment on the snout confined to the sides ; scale pocket pigment of the

lateral line inconspicuous ........ B. bawkuensis

12 Two pairs of barbels ; mid-lateral pigmentation variable but three spots often

present ; no spot on the dorsal fin . . . . . . B. atakorensis

One pair of barbels ; a narrow continuous mid-lateral band ends in a spot on the tip
of the caudal peduncle ; dorsal fin marked with a black streak across the distal
half of the anterior rays ......... B. pobeguini

1 3 Dorsal fin marked with a round black spot ; body with a single spot on the tip of the

caudal peduncle .......... B. leonensis

No spot on the dorsal fin ; Body marked with two to four black spots, one at the

base of the anal fin, the others on the horizontal myoseptum . B. stigmatopygus

ACKNOWLEDGEMENTS

We wish to thank the following people in connection with this work : Dr. P. H.
Greenwood for his kind encouragement and advice, for many stimulating discussions,
for his invaluable criticism of the manuscript and for providing us with facilities to
work on material in the British Museum (N.H.) ; Dr. E. Trewavas for valuable help
and for allowing us to use her unpublished data on B. stigmatopygus ; Dr. M. Blanc
who kindly arranged for us to examine material in the Paris Museum ; Dr. S. H.
Weitzman of the Smithsonian Institution who re-examined the type of Mannichthys
lucileae on our behalf ; Mr. G. G. T. Harrison, formerly Chief Fisheries Officer, Ghana,
who suggested taxonomic work on Ghanaian fish ; Mr. D. Ofori-Adu, Fisheries
Assistant, Fisheries Division, Ghana, who helped to collect much of the data ; Mr.

BARBUS OF THE VOLTA REGION 149

Kposugbe Dutanyah, Fisherman, Fisheries Division, Ghana, who caught most of the
specimens ; Mr. David Barry of the Kwame Nkrumah University, Kumasi for help
with the collection of the forest species.

REFERENCES

BLANC, M. & DAGET, J. 1957. Les eaux et les poissons de Haute-Volta. Mem. Inst. France.

Afr. Noire, no. 50 : 99-169.
BOULENGER, G. A. ign. Catalogue of the freshwater fishes of Africa in the British Museum

(Natural History), 2, London.
1916. Catalogue of the freshwater fishes of Africa in the British Museum (Natural History),

4, London.
DAGET, J. 1952. La reserve naturelle integrate du Mt. Nimba. Mem. Inst. France. Afr.

Noire, no. 19 : 311-334.

1954- Les poissons du Niger sup6rieur. Mem. Inst. France. Afr. Noire, no. 36 : 391 pp.

1959. Les poissons du Niger superieur (ire note complementaire) . Bull. Inst. France.

Afr. Noire, 21, s6r. A, no. 2 : 664-688.
GREENWOOD, P. H. 1962. A revision of certain Barbus (Pisces, Cyprinidae) from east, central

and south Africa. Bull. Brit. Mus. (Nat. Hist.), Zool. 8, no. 4 : 151-208.
1962. A new species of Barbus (Pisces, Cyprinidae) from the Upper Zambezi River,

Rhodesia. Rev . Zool. Bot. Afr. 65 : 211-216.
1963. Notes on Barbus radiatus Peters 1853 and the subgenus Beirabarbus in east, south

and central Africa. Rev. Zool. Bot. Afr. 67 : 20-28.
HERRE, A. W. 1936. A new cyprinid genus and species and a new characin from Portuguese

East Africa. Proc. Biol. Soc. Washington, 49 : 99-102.
HOLDEN, M. J. 1963. The populations of fish in dry season pools of the Sokoto River. Col.

Off. Fish. Pub., London, no. 19 : 65 pp.
HOPSON, A. J. 1965. Four new species of Barbus (Pisces Cyprinidae) from Ghana. Rev. Zool.

Bot. Afr. 71 : 245-257.

IRVINE, F. R. 1947. The fishes and fisheries of the Gold Coast, London : 352 pp.
JOHNELS, A. G. 1954. Notes on fishes from the Gambia River. Arkiv. f. zool. Stockholm, (2)

6 : 327-411.
NORMAN, J. R. 1932. A collection of fishes from Sierra Leone. Ann. Mag. nat. Hist. (10)

10 : 180-185.
PELLEGRIN, J. 1909. Description d'un Barbus nouveau du Sahara. Bull. Mus. Nat. Hist.

nat. 5 : 18-19.
1911. Description d'un poisson nouveau de Guin6e Fran9aise appartenant au genre

Barbus. Bull. Soc. Zool. France, 36 : 187.
1921. Les poissons des eaux douces de 1'Afrique du Nord franyaise. Mem. Soc. Sci. Nat.

Maroc, 1 (2) : 216 pp.
1923. Les poissons des eaux douces de 1'Afrique occidentale, Paris : 373 pp.

SCHULTZ, L. P. 1942. The freshwater fishes of Liberia. Proc. U.S. Nat. Mus. 92 : 301-348.

SVENSSON, G. S. O. 1933. Freshwater fishes from the Gambia River (British West Africa).

Results of the Swedish expedition, 1931. Kungl. Sven. Vet. Akad. Handl. 12 (3) : 102 pp.

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NOTES ON THE CUPULADRIIDAE
(POLYZOA, ANASCA)

PATRICIA L. COOK

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 5

LONDON: 1965

NOTES ON THE CUPULADRIIDAE
(POLYZOA, ANASCA)

PATRICIA L. COOK

British Museum (Natural History)

Pp. 151-187 ; Plates 1-3 ; 6 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. 5

LONDON: 1965

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 13, No. 5 of the Zoological series.
The abbreviated titles of periodicals cited follow those of
the World List of Scientific Periodicals.

Trustees of the British Musi urn (Natural History) 1965

Issued October, 1965 Price Seventeen Shillings

NOTES ON THE CUPULADRIIDAE
(POLYZOA, ANASCA)

By PATRICIA L. COOK

Page

1 INTRODUCTION ............ 153

2 CUPULADRIIDAE LAGAAIJ .......... 154

3 STRUCTURE AND DEVELOPMENT OF THE ZOARIUM . . . . . -155

a. Early astogeny .......... 155

b. Growth. . . . . . . . . . . .156

c. Regeneration and deformation. . . . . . . . 157

4. CHARACTERS OF SYSTEMATIC VALUE . . . . . . . .159

a. The central area .......... 159

b. The cryptocyst . . . . . . . . . .160

c. The opercula ........... 160

d. The vibracula .......... 160

e. The basal surface . . . . . . . . . .161

5 The EFFECTS OF ZOARIAL FORM ON CHARACTERS . . . . . . 1 62

6 COMPARISON WITH THE STRUCTURE OF OTHER FREE-LIVING ZOARIA . . .164

7 GROUPING OF SPECIES .......... 167

A. Cupuladria canariensis group ........ 167

B. C. guineensis group . . . . . . . . .167

C. C. owenii group .......... 167

D. Discoporella umbellata group . . . . . . . .167

8 Cupuladria pyriformis (Busk) . . 3 . . . . . .168

9 C. indica n. sp. . . . . . . . . . . . . 169

10 C. guineensis (Busk) . . . . . . . . . . .170

11 The Discoporella umbellata-complcx . . . . . . . . 175

12 D. umbellata (Defrance) ... ...... 177

13 D. umbellata subsp. depressa (Conrad) . ...... 180

14 CUPULADRIIDAE, GENERAL COMMENTS . ...... 182

15 Setosellina capriensis (Waters) . . ...... 182

1 6 ACKNOWLEDGMENTS . . . . . . . . . . .184

17 SUMMARY ............. 184

18 REFERENCES ............ 184

i INTRODUCTION

RECENT work on the rich west African fauna of species belonging to the Cupulad-
riidae, has led to an examination of other species and to a general review of the
characters of the family. The following notes are an attempt to define the limits of
variability of some of these characters, and to delimit groups of species which appear
to be related.

Specimens in the Zoological Department of the British Museum are referred to by
registered number thus, 1899. 7. i. . . . ; in the Palaeontological Department thus,
D 6474. Measurements of zooecial length have been made from the distal rim of one
zooecium to that of the next succeeding radial zooecium, so including the vibraculum.

ZOOL. 13, 5. 8

154

P. L. COOK

Measurements are given thus :

Length of zooecium Lz

Width of zooecium Iz

Length of vibracular opesia Lvo

Length of vibracular seta Ls

Length of operculum Lo

Width of operculum lo

Length of zooecial opesia Lop

Width of zooecial opesia lop

DEFINITION OF TERMS USED. The majority of the terms used may be found denned
by Lagaaij (1952 : 12) and Bassler (1953 : 7).

Horizontal cryptocyst lamina. A porous lamina, formed from fused cryptocyst
denticles, extending from the descending cryptocyst across the opesia, parallel to,
and below, the frontal membrane.

Vestibular arch. The distal, cresentic, raised portion of the aperture.

Vicarious vibraculum. A large vibracular individual, taking the place of a
zooecium, and itself having a distal vibraculum.

2 CUPULADRIIDAE Lagaaij
Cupuladriidae Lagaaij, 1952 : 31.

Lagaaij denned the family to include only the genus Cupuladria. A series of
species is now known to exist (Cook (in press)) which links the membraniporan forms
such as C. canariensis (Busk) with the microporan forms like Discoporella umbellata
(Defrance). Both genera have therefore been included in the Cupuladriidae, the
diagnosis of which has been accordingly slightly modified.

DIAGNOSIS. Lunulitiform Anasca with vibracula alternating with zooecia in the
same radial series. Cryptocyst variously developed. Ovicells absent. Ancestrula
surrounded by seven zooecia and a distal vibraculum.

Lagaaij (1953 : 13) defined the term " lunulitiform " to include conical and discoid
zoaria such as those of Cupuladria, Discoporella, Lunulites and Selenaria. For
reasons given below, the term is here restricted to the first three of these genera, which
all bud radially, although it is not assumed that the Cupuladriidae and Lunulites are
closely related. The term " selenariiform " used by Harmer (1926 and 1957) for
both Anascan and Ascophoran conical colonies, is here restricted to zoaria with spiral
budding, e.g. Selenaria, Setosellina and Heliodoma. The term " conical " is used
here for all the remaining forms described by Harmer, some of which may be attached
to the substrate by rootlets, e.g. Conescharellina.

It must be stressed that, whereas the combination of characters found in the
Cupuladriidae is exclusive, and the limits defining the species are generally well
marked, the division of the family into two genera, Cupuladria and Discoporella, is
somewhat arbitrary, depending upon only one character, namely, the extent of the
development of the cryptocyst. Until work can be done on the life-histories and
breeding of the Cupuladriidae, the importance of this character in relation to, for
example, the size of the eggs, the larvae, and the other zooecial structures, cannot be
assessed, and Discoporella is therefore retained here.

THE CUPULADRIIDAE
3 STRUCTURE AND DEVELOPMENT OF THE ZOARIUM

155

Although many free-living colonies are similar in appearance, they differ con-
siderably in structure and development. In the Cupuladriidae both the early
astogeny and subsequent growth of the zoarium are distinct from that found in other
lunulitiform, selenariiform or conical colonies.

a. Early astogeny. The ancestrular area is here denned as comprising the ancestrula
and the primary circle of zooecia budded from it. The single ancestrula buds a distal
vibraculum, a pair of proximal-lateral zooecia and then one proximal zooecium (see
Marcus & Marcus, 1962 : 287). A pair of lateral zooecia follows, and a pair of distal-
lateral zooecia is the last to be formed (see Text-fig. iB). Lagaaij (ig63a : 182,
text-fig. 10) found that secondary proximal buds were developed before this last pair,
but colonies seem to be variable in development at this stage. Evidence has been
gained from very young colonies of Cupuladria sp. (Gulf of Mexico, 1959. 8. 20. 3,
Lagaaij Coll.) and D. umbellata (Madeira, 1963. 2. 28. 9, and S. Africa, 1949. n. 10.
668, Burrows Coll.). Waters (1926 : 426) described a double ancestrula in C.
canariensis, but none have been seen in any of the many specimens examined, and I
agree with Lagaaij (ig63a : 184) that it is possible that Waters regarded the proximal
bud as the second member of a paired ancestrula.

In one group (C. canariensis, group A, see p. 167), the entire ancestrular area may
consist of vicarious vibracula (see Smitt, 1873, pi. 2, fig. 70 ; Hastings, 1930 : 714 ;
Marcus & Marcus, 1962 : 289 ; Lagaaij, 19632. : 183 ; and Cook (in press)). Although
vicarious vibracula in the ancestrular area may sometimes be the result of regeneration
of a normal zooecium, most of those found in C. biporosa (Canu & Bassler) and
C. monotrema (Busk) are budded primarily as vibracular individuals. The " special
area " described by Canu & Bassler (1923 : 81, pi. 2, fig. 18) is not an ancestrular area,
but the growing edge of a regenerating, broken fragment (see below) .

The pattern of budding outside the ancestrular area appears to be variable.
Another variable character is found in C. Pyriformis (see 168) and a few colonies of
C. canariensis, where the vibracular seta of the ancestrula, when closed, is directed
towards the right side (viewed frontally) (see Text-fig. iC). In all specimens of

BCD

FIG. i. Early astogeny. a, ancestrula ; vibracula stippled. A. Setosellina. Spiral budding.
B. Cupuladriidae. Radial budding. C. Cupuladvia pyriformis (Busk), ist pair of
zooecia aborted. D. Lunulites. Radial budding: .

156 P. L. COOK

other species the seta is directed toward the left side, the seta of the zooecium budded
from the proximal end of the ancestrula having the same orientation (see Text-fig.
iB). The seta of the right-proximal-lateral bud is directed to the right, that of the
left-proximal-lateral bud to the left (see also Marcus & Marcus, 1962 : 297).

b. Growth. Each zooecium and vibraculum develops radially as a unit which
arises as a common bud, the vibracular chamber being divided off from the cavity of
the zooecium in the later stages of development. The relationship of the vibraculum
with the zooecium distal to it is comparable with that of the " recumbent " ovicell of
Crepidacantha (see Brown, 1954: 244, footnote). Here the ovicell is fully formed
before the development of the next distal zooecium, which grows round it from below.
The vibracular chamber in the Cupuladriidae is not terminal and does not reach the
basal surface of the zoarium. Thus, although apparently inserted between the zooecia
at the frontal surface, the vibracula are not strictly interzooecial, nor are they
adventitious or dependent.

The common bud is rarely present, even in specimens preserved in alcohol, as in
its peripheral position it is particularly susceptible to damage when the colony is dead,
and it is no longer protected by the extended peripheral vibracular setae (see Marcus
& Marcus, 1962, and Cook, 1963). The bud arises between peripheral zooecia. The
membranous ectocyst is closely applied frontally and basally to the neighbouring
zooecia, but over the bud it is convex and free from the underlying structures. A
line of demarcation occurs basally where the ectocyst is no longer in contact with the
calcareous basal walls (see Text-fig. 2A). On the basal side the bud originates from
the distal end of the proximal zooecium, below the vibracular chamber. On the
frontal side the upper surface of the bud appears at the distal wall of the vibracular
chamber. The membranous frontal part of the bud becomes the frontal membrane
of the zooecium, the basal part the endocyst lining the zooecial cavity (see Marcus &
Marcus, 1962, pi. 2, fig. 5). This can be seen in well-preserved specimens of C. indica
(1899. 5- I- 267) .' C. multispinata (1949. n. 10. 617) ; Discoporella umbellata
(MM I 39A) ; and D. u. depressa, (Terra Nova Coll., Stn. 42). The relationships of
the young common bud have also been observed in living colonies of D. umbellata
from Madeira.

As the bud enlarges and grows up between the two adjacent lateral zooecia, the
basal wall is covered by a calcified lamina, which develops from the point of origin of
the bud (see Text-fig. 2 A, B). The polypide is first differentiated at the proximal end
of the bud before it is completely covered basally by the lamina. As the lamina
advances distally, the cryptocyst develops beneath the frontal membrane growing
inward from the lateral walls. The lamina curves upward and inward, and the vibra-
cular chamber develops as an inner compartment from the distal end of the bud. The
walls of the chamber are thus double-layered laterally, with connecting bars of
calcification. Large pores are present in the outer layer (= " chambers " of Marcus
& Marcus, 1962 : 295). These calcify further, become smaller, and eventually com-
municate with the next distal-lateral zooecia to be developed. The vibracular
chamber is connected with its proximal zooecium through a large pore in the dividing

THE CUPULADRIIDAE 157

wall. An uncalcified area, which is left in the lamina as it curves up sharply distally
to the vibracular chamber, calcines, but leaves a small pore which eventually com-
municates with the succeeding radial zooecium (see Text-fig. 2). Specimens clearly
showing these structures are C. multispinata (S. Africa, 1949. n. 10. 94 B, Burrows
Coll.) and D-. u. depressa (Rio de Janeiro, Terra Nova Coll., Stn. 42).

Transverse sections of zoaria frequently show the stem-like extension of the zooe-
cium, below the vibracular chamber of the previous zooecium. Each zooecium thus
has a large distal pore communicating with the vibracular chamber, and 2-3 lateral
pores communicating with the adjacent distal-lateral zooecia. No pores enter the
vibracular chamber laterally, but it communicates with the next distal zooecium in a
radial row. Specimens showing the interzooecial pores clearly are : C. indica
(1899. 5. i. 267 pt.) and C. canariensis (section, 1929. 4. 26. 85 pt.) ; see also Marcus
& Marcus, 1962, pi. 2, fig. I.

Large colonies of D. umbellata (e.g. coast of Ghana, 506, Achimota Coll), and
C. canariensis (Senegal, MM II 3) show rings of growth (see Harmer, 1926 : 267).
Apparently these zoaria have grown evenly for a period and then ceased to grow.
New budding has been resumed simultaneously all round the periphery, so that a
series of concentric ridges may be seen on the basal side.

FIG. 2. Growth of the membranous common bud (c.b.) and calcareous basal lamina, viewed
from the basal side. A. Transparent, membranous common bud developing between
two peripheral zooecia. Note the uncalcified area in the distal wall of the vibracular
chamber. B. Calcareous curved basal lamina developing over the growing common bud.
Note the increased calcification of the previously developed proximal-lateral basal
laminae, and the development of the radial median groove.

c. Regeneration and deformation. A large number of colonies are regenerated from
broken fragments, and the budding by which a symmetrical colony is re-formed
follows a characteristic pattern. Stach (i936a : 63-65) and Dartevelle (1933 : 70-72)
have discussed the fragmentation of colonies, and the ecological significance of this
form of regeneration. Dartevelle (1935 : 559-561) also described the form of budding
in C. canariensis. He mentioned regeneration as occurring from " calcified zooecia "
(see below), and examination of many colonies shows that these closed zooecia are,
in fact, the primary buds regenerated from a broken edge of a fragment. The
original fragment can usually be seen on the basal side of such colonies, as secondary

158 P. L. COOK

calcification at the centre does not seem to occur. The broken zooecia which were
distal in position in a fragment regenerate zooecia distally, those which were proximal,
regenerate zooecia proximally, in which the zooecial orientation is reversed. Zooecia
growing from the lateral walls of old zooecia, are at right angles to the previous
direction of growth of the fragment. Each broken zooecium produces one bud
laterally from the mid-line of its basal wall, and another bud arises from beneath the
vibracular chamber. These zooecia are wide and often slightly distorted (see p. 175).
Subsequent budding of radial series, including intercalary rows, is exactly the same as
in zoaria developing from an ancestrula. The usual radial alternation in the asym-
metry of the vibracular opesia is established in the primary and secondary series of
buds (see Text-fig. 3 A). The colony of C. elongata Sakakura mentioned as distorted
by Marcus & Marcus (1962 : 288) is, in fact, such a regenerating fragment.

Marcus & Marcus (1962 : 301) have described a second form of regenerative or
" zoarial budding ", in which small, fan-shaped zoaria are produced from a single
zooecium at the periphery of the parent colony. This form of budding has been seen
so far only in the D. umbellata-complex (e.g. Madeira, 1912. 12. 21. 1004, Norman Coll.)
and in C. multispinata (Madeira, 1963. I. 2. 3 and 4, Norman Coll.). Marcus &
Marcus described the results of the fracture of the initial zooecium (" pseudo-
ancestrula "), which frees the young zoarium from the parent colony. It then
continues its development, eventually assuming a circular shape by the budding of
intercalary series. Colonies arising in this manner have no central substrate present,
e.g. a sand-grain or foraminiferan. They may also be recognized by the reversed
orientation of the regenerated zooecium developed from the broken pseudoancestrula,
and by their early fan-shaped budding.

The pseudoancestrula does not calcify fully on the basal side, and when still
attached to the zoarium has been seen to fold frontally when its vibracular seta closes
(see Cook, 1963 : 411). When fracture occurs, the basal lamina is detached with the
new zoarial bud, but the proximal part of the cryptocyst remains with the parent
colony (see Marcus & Marcus, 1962 : 302). The distal part of the pseudoancestrula
then regenerates a zooecium orientated in the opposite direction. In the Madeiran
material the fracture has not always occurred, and buds have continued to develop
attached to the parent colony. Deformed colonies have resulted, some of which
have been described as C. lowei by Busk (1854 : 99 — D. umbellata) and " C. deformis "
by Waters (1921 : 413 = C. multispinata).

Other deformed zoaria, especially of these two species, originate from regenerating
fragments, and here, deformation is the result of failure of development of one or
more radial rows of zooecia. The cause is unknown, but in some cases it appears to
be correlated with the growth of associated Sponges, belonging to the Plocamiidae,
on the basal side of the Polyzoan colony. The Sponge appears to have inhibited
growth at the origin of the chitinous zooecial bud, just below the penultimate series of
vibracular chambers. The radial rows of zooecia adjacent to the affected zooecium
or zooecia, produce intercalary series until the radial slit in the colony is closed when
the two sides touch. Frequently, these zooecia are unable to form interzooecial
connections, and a slit or furrow continues radially through the colony, which, as the

THE CUPULADRIIDAE

159

associated Sponge develops, becomes lined with spicules. No example of a spiral slit
has been seen in deformed colonies of the Cupuladriidae, and the budding pattern
should be compared with that of one deformed specimen of Selenaria maculata (Bass's
Straits, 1899. 7. i. 1497), where the slit is distinctly spiral (compare Text-figs. 36,
D). The basal surface of this zoarium also shows traces of a Sponge colony, and
although it is possible that the Sponge is a secondary and unrelated growth, the
correlation with this type of deformation is interesting.

A B

FIG. 3. Regeneration and deformation. Vibracula stippled. A. Cupuladria. Regenera-
tion from lateral walls of a radial series. B. Selenaria. Spiral regeneration. C.
Discoporella. Deformation (*), point where distal radial bud has not developed. D.
Selenaria. Deformation (*), point where lateral distal bud has not developed.

4 CHARACTERS OF SYSTEMATIC VALUE

a. The central area is here defined to include the ancestrular area and the centre of
the colony where the budding of intercalary radial rows is most frequent and closed
zooecia are found. In all groups of species but one (group A, see p. 167), the opesiae
of the zooecia of the central area become occluded by secondary calcareous growth ;
these zooecia do not then contain polypides (see Harmer, 1926 : 267). Closed zooecia
are often present even in fairly young colonies, and are also found in areas of rapid
budding at the edges of regenerating fragments (see above).

In the membraniporan group B (see below) the lamina is formed as a thin sheet
originating just below the level of the slight shelf formed by the descending
cryptocyst. It develops in a proximal direction from the distal end of the opesia,
later also growing out from the lateral walls. Although the lamina thickens it rarely
approaches close to the frontal membrane. Irregular pores are left, especially at the
lateral edges. In the denticulate forms (Group C, see p. 167), the lamina is formed by
extension and fusion of the existing cryptocystal denticles and first develops proxi-
mally, spreading progressively to the sub-opercular region. It is closer to the frontal
membrane in origin than the lamina in Group B, and when thickened by further
deposition may almost obscure the mural rims of the central zooecia. In Discoporella
where the opesia is restricted to a small sub-opercular area, the central cryptocystal
pores calcify first, the opesia then becoming occluded by denticulate growth. The
opesiules frequently remain uncalcified or only partially closed. The lamina in these

160 P. L. COOK

last two groups is rarely complete, irregular pores usually being present, but in
contrast to the first group, no pores are present in the subopercular area.

The existence and type of central area closure may be evidence of the group to
which a species belongs, and the association of closed zooecia with ancestrular and
regenerative budding in groups B, C and D is comparable to the occurrence of
vicarious vibracula in group A.

b. The cryptocyst is present in all species and, in well-preserved material, is a
useful specific character, especially when its form can be correlated with other
characters, such as those of the basal surface. Unfortunately, the cryptocyst is
among the first structures to be damaged or worn, although there are usually one or
two zooecia present in which it can be seen, even in fossil specimens.

The type of cryptocyst forms a series in the Cupuladriidae from the membrani-
poran C. canariensis to the microporan D. umbellata. In the C. canariensis and
C. guineensis groups (see below), it is simple, though variously extensive. In the
C. owenii group it is denticulate, the form and extent of the denticles being specific.
In Discoporella it is so extensive that it has been regarded as a generic character.
In development, however, the cryptocyst of D. umbellata shows great similarity to
that of the C. owenii group, as it is formed from the fusion of irregular denticles (see
Marcus & Marcus, 1962, pi. i, fig. 4). In Discoporella the final development of the
cryptocyst results in a pair of distal opesiules or opesiular indentations, through which
pass a pair of muscles. Other opesiular pores are present, but the opesia is always
limited proximally by a complete bar, formed by part of the cryptocyst.

c. The opercula of the majority of species of Cupuladriidae are similar to one another.
They are membraniporan with a strong, curved distal sclerite. Their mean dimen-
sions or proportions may give additional information toward the definition of a
species, but generally their range of variation is large. Furthermore, opercula are
absent not only in fossil specimens, but frequently from dead, Recent material which
is a little worn.

Groups C and D (see below) each include species in which the operculum has
a partial or complete proximal sclerite. In C. doma (Group C), the proximal sclerite
is variable, and may be incomplete. Its presence may be useful in distinguishing
small fragments from those of C. owenii, which may have similar zooecial characters,
but the basal surface is here more certain as a determinant of the species. Discoporella
(group D), has hitherto been distinguished by the possession of a proximal sclerite in
the operculum, but there is evidence (Cook (in press)) that even in this group the
opercula are not all of the same type and include at least one species in which they
are without a proximal sclerite.

d. The vibracula are also similar in character in all species of the family. The
opesia is auriform, with distinct condyles, which are alternately asymmetrical to
right or left in each member of a radial row of zooecia. The vibracular setae are long
and usually follow a sigmoid curve, they increase in length toward the periphery of
the colony. The setae have an asymmetrical basal sclerite, to which are attached
powerful muscles which fill the vibracular chamber. Although there are differences
between the setae of various species, especially in the size and shape of the basal

THE CUPULADRIIDAE

161

sclerite, the variation within the colony is large and they appear to be of little use in
specific determination. The basal sclerites of group A seem generally to be less
tuberculate and robust than those of the other groups. The detailed morphology of
the vibraculum was discussed by Marcus & Marcus (1962).

The seta is able to move in two directions and in one plane. Owing to its asym-
metrical articulation and curvature, these movements cover nearly half of the frontal
surface of the zooecia proximal and distal to any vibraculum. Live colonies examined
by me have not been seen to use their setae like tweezers, as described by Marcus &
Marcus (1962 : 299). The distribution of the setae, and their range of movement, is
such that they rarely come into contact (see Text-fig. 4). Those near the centre of
the colony may touch each other when closed, owing to the convergence of radial
rows, but should any detritus be held between two setae in this position, they
automatically lose all contact as soon as an opening movement begins. Particles on
the frontal surface are dislodged by the movement of individual setae. They roll
down the sloping surface of the colony, passing between the peripheral, supporting
setae. The zoarium is thus kept clear of deposits settling upon it (see Cook, 1963 :
410).

FIG. 4. Distribution of vibracular setae in Discoporella wnbellata. Membrane of vibra-
cular opesiae and opercula stippled. Setae pointing distally are in the " open " position ;
those pointing proximally are in the " closed " position.

e. The basal surface. Lagaaij (1952 : 34, 1953 : 13) stressed the importance of the
character of the basal surface in specific determination of fossil specimens, where the
frontal surface may have suffered wear. In those groups (A and B below) in which
rectangular or hexagonal sectors are present, the thickening consists of layers of

ib2 1'. L. COOK

calcification, pierced by regular series of kenozooecial chambers, or by small pores.
The shape and arrangement of the sectors, and the number of pores or chambers may
be specific, within defined limits. For example, the basal sectors of C. canariensis
s. s. consist of several layers of long irregular units, interspersed with occasional short
sectors ; whereas those of C. monotrema are single layered and invariably short and
square. The sectors of C. guineensis (see Hastings, 1930 : 714, and p. 170) may
vary, especially in large colonies, from square to hexagonal. In the first case the
characters of the basal surface are correlated with other distinguishing features, in the
second, the variability of the character is found within a single colony, and is not
correlated with any other character, and is therefore not specific. In the remaining
groups the basal surface consists of a series of alternating radial grooves and ridges,
the characters of which may also be specific, although they are variable, affected by
wear, and similar forms occur in both Cupuladria and Discoporella.

The ridges and grooves are the result of the form of growth of the zooecia. Each
curved basal lamina is grooved in the mid-line (see Text-fig. 2). As secondary cal-
cification increases the groove may be filled in, and in some cases may be marked by a
protuberant " thread " of heavier calcification. The tuberculation of the basal
surface may increase with calcification, but may also become changed by wear. In
some species, however, the form of tuberculation may be correlated with other charac-
ters, and be specific. For example, in C. owenii, the tubercles are consistently small
and regular, and in C. multispinata, they are large, coarse, and coalescent (Cook (in
press)).

5 THE EFFECTS OF ZOARIAL FORM ON CHARACTERS

The extreme variation in zoarial and zooecial characters of specimens attributed to
D. umbellata has been discussed by several authors, and Silen (1942 : 16) noted
variation in the shape of zoarium of his specimens and grouped his material into
localities where one type of zoarium was dominant. The type of zoarium appears to
have little effect on the zooecial characters except where the form of the colony is the
same as that found normally in C. doma (d'Orbigny) or that found in zoaria described
as C. peyroti by Duvergier (1924 : 19).

Doma-type zoaria occur in several species of Cupuladriidae (Cook (in press)) and
perhaps correspond to the Trochopora-habit of Lunulites (see Lagaaij, 1953 : 17).
Specimens belonging to the D. umbellata-complex. (see p. 175), frequently produce
doma-type colonies, and before discussion of specific differences, the effects of form of
growth must be considered.

The doma-type zoarium is small and usually high (diameter range 3-7 mm.) The
zooecia are small, and the basal surface solid or nearly filled in by secondary calcifica-
tion. In the majority of zoaria the peripheral zooecia are closed, and there may be
from 1-3 rows of enlarged peripheral vibracula.

In all lunulitiform zoaria the angle of the axis of the polypide cavity approaches a
right angle with the zoarial axis, and the planar intercept of the frontal of each
zooecium is thus longer in flatter colonies than in the doma-type zoarium, where there
is therefore a reduction of the zooecial dimensions usually found in the species, and

THE CUPULADRIIDAE 163

consequent changes in the development of the cryptocyst and the proportions of
various other characters. The basal surface is also frequently non-typical, owing to
the large amount of secondary calcification.

Canu & Bassler (1930 : 12) stated that specimens from abyssal depths of the
Pacific reported in 1929 (p. 142), were stunted. Photographs of these specimens
(1963. 3. 14. 14) show that they are not, however, either particularly small, or of the
doma-type. Canu & Bassler also mentioned that colonies of " var. conica " had been
found in the Helvetian of Touraine, but these colonies were not specifically mentioned
by Canu (1909 : 448) in his description of the zoaria from this locality. Marcus &
Marcus (1962 : 304) described colonies from the coast of Brazil as D. umbellata var.
conica Canu & Bassler. These are certainly the same form as those listed below as
doma-type zoaria (see p. 181).

The number of specimens examined of doma-type colonies is not at present sufficient
to determine whether they should be considered a subspecies of D. umbellata. It
appears more likely that they are a variant developed in response to specific, but
unknown, ecological conditions. The specimens examined fall roughly into 3 groups,
each corresponding to one of the 3 major forms in the D. umbellata-complex. (see
p. 176). The ecological factors involved in the occurrence of doma-type colonies are
probably not simple. There is, for example, no correlation with depth in the material
examined, although C. doma itself extends in distribution to greater depths than other
species of the Cupuladriidae. One doma-type specimen from Cape Frio, off Angola, is
the sole record from the south west coast of Africa, where the temperatures are lower
than, for example, east of the Cape of Good Hope. At Mowe point, just south of Cape
Frio, in 400 m., a temperature of 9° C. has been recorded (see Hart & Currie, 1960 :
173, text-fig. 29). The conditions producing doma-type growth may perhaps be
related to temperature.

In the D. umbellata-complex. the doma-type colonies are very small and the number
of opesiules is reduced. The size of opesiae and opercula are not, however, reduced
in the same proportion. The peripheral zooecia are closed and the peripheral
vibracula enlarged, as in C. doma. Generally, the effect is of a suppression of the
characters normally separating the populations in the complex. In nearly all cases,
however, the specimens retain traces of features consistent with those found in the
larger, more normally developed colonies (see pp. 180, 182).

Peyroti-type growth is similar to doma-type growth, and may also be caused by
ecological conditions at present unknown. Some colonies combine features of both
forms of growth (see p. 180), but whereas doma-type zoaria occur in all 3 populations
of the complex, Recent peyroti-type colonies are the characteristic form of one geo-
graphical area, namely the south-eastern coast of Africa. Duvergier (1924 : 19)
described, as Cupularia peyroti, colonies similar in character to Discoporella umbellata,
from the Miocene of France, in which the basal surface (described as " face superieur ",
following Canu's theoretical orientation of lunulitiform and selenariiform colonies)
was covered by kenozooecia and vibracula. Recent specimens from south and east
Africa (see p. 177) show stages in the development of the same form of growth, which
was briefly described by O'Donoghue (1924 : 39) and discussed by Hastings (1930 :

164 P. L. COOK

719), who doubted whether her south African specimens could be included in D.
umbellata. The large number of colonies now available show that although the
zoarial form is markedly different, the zooecial characters have much in common with
D. umbellata s.s.

The modifications to a specific character produced by peyroti-type growth are
principally in the appearance of the basal surface. Zoaria are usually large, unlike
do ma-type colonies, and have a smooth, slightly pitted base, which may be concave or
solid. There is no sign of the tuberculate, grooved basal surface usually found in
D. umbellata. In most zoaria the base is covered by a layer of kenozooecia and
vibracula proliferated from the peripheral zooecia and vibracula (see PI. 3, fig. 6).
The vibracular opesiae are as large as those of the peripheral vibracular zooecia but
their setae are not as long. The kenozooecia have no polypides and resemble the
closed zooecia of the frontal central area except that there are pores in the distal part
of the lamina which in normal closed zooecia is the solid, sub-opercular region (see
p. 160).

The peripheral frontal zooecia are also closed as in doma-type zoaria, but the peri-
pheral vibracula are not greatly enlarged nor do several rows of them occur.

The basal vibracula presumably act in a similar manner to those at the periphery,
and assist in stabilizing the zoarium and maintaining it just above the surface of the
substrate (see Cook, 1963).

The conditions determining peyroti-type. growth do not seem to be related to
temperature, as similar ranges occur east of the Cape of Good Hope to those off west
Africa (i.e. warm temperate to tropical, see Stephenson, 1947). No Cupuladriidae
have been reported from the colder waters immediately to the west of the Cape of
Good Hope.

6 COMPARISON WITH THE STRUCTURE
OF OTHER FREE-LIVING ZOARIA

Silen (1942 : 1-22) examined many free-living zoaria and considered that their
growth was spiral i.e. that the zooecia of each apparently radial row were budded
from the nearest proximal-lateral, not proximal, zooecium. Examination of many
lunulitiform, selenariiform and conical colonies has confirmed Silen's observations for
Setosellina, Setosella (principally an encrusting form) and Heliodoma, and Stach's
observations (i936a) for Selenaria, but has also shown that the method of growth in
the Cupuladriidae and the genus Lunulites is entirely different. The evidence is
summarized below.

a. In the spirally growing forms the ancestrula buds off both a distal and a proximal
(or proximal-lateral) vibraculum, each with an adjacent zooecium. A good example
was figured in Vibracellina caribbea1 by Osburn (1947 : n, pi. I, fig. 2). Two spirals
of zooecia and vibracula develop from these foci, so that although the ancestrula is
eventually surrounded by zooecia, unlike that of Lunulites and the Cupuladriidae, it
has not directly budded all of them (contrast Text-fig. lA with Text-figs. iB, C and

1 Lagaaij (ig63b : 172) also describes and figures the ancestrula region of this species, which he refers
to Setosellina goesi (Silen) .

THE CUPULADRIIDAE 165

D). The spiral budding is well illustrated in 5 zoaria of Selenaria squamosa (N.
Zealand, 1947. 8. 16. i), where the spiral is anti-clockwise. All colonies of other
species seen are developed in a clockwise direction. In Lunulites the budding
pattern is radial, not spiral, but differs from that of the Cupuladriidae in that the
ancestrula buds 6 zooecia, each of which then gives rise to a zooecium and a vibra-
culum. The ancestrula does not therefore directly bud a vibraculum (see Text-fig.
iD).

b. Silen based his conclusions as to the spiral nature of the zoaria of the Cupulad-
riidae on the assumption that morphologically the vibraculum was the distal bud
formed by each zooecium, and that a vibraculum could not bud off a zooecium distal
to itself. The zooecium and vibraculum of the Cupuladriidae originate as a unit,
and although there is evidence that the vibraculum is a modified zooecium (see Marcus
& Marcus, 1962 : 298), it is produced within the confines of the common bud, and is
not itself the product of distal budding. In spirally developed colonies (e.g. Setosel-
lina], the vibracular chamber can be seen to reach the basal side of the zoarium, and
the distal-lateral origin of the next zooecial bud can be seen.

c. In spirally developed colonies, the parent zooecium of an intercalary series forms
the second bud consistently on the side opposite to the direction of spiral growth.
In the Cupuladriidae the intercalary zooecia are produced from either distal-lateral
wall of the parent zooecium, in a random manner. The vicarious vibracula in the
C. canariensis group (see below) may be intercalary or may themselves produce
intercalary zooecia or further vicarious vibracula. Their appearance is also entirely
random, and has no spiral pattern. In Lunulites the radial rows of zooecia alternate
with those of the vibracula. There is no direct lateral relationship between the
budding of the types of individual, which are developed radially, not spirally.
Successive vibracula enlarge in a series and finally one buds a distal zooecium. New
intercalary rows of vibracula are produced from bifurcations of series distal to a zooe-
cium, not a vibraculum. In the central area the first vibracular rows are all budded
from the proximal zooecium in the same lateral relationship to the zooecial buds (see
Text-fig. iD). In subsequent bifurcations the vibracular bud arises to the right or
left of the zooecial bud in a random manner.

d. The growing edge of Selenaria bimorphocella (off Adelaide, 1928. 9. 13. 81),
shows no basal curved laminae, but a series of groups of young zooecia alternating
with developing vibracula, similar in appearance to the growing edge of an encrusting
colony. Similarly, the growing edges of colonies of Setosellina have no basal curved
laminae. The peripheral zooecia of Lunulites arise in a similar manner to those in
the Cupuladriidae and basal laminae have been seen. The resulting basal grooves
mark the junction of the lateral walls and do not have the alternating pattern of the
Cupuladriidae (see below). In Recent Lunulites the cavities of the vibracula reach
the basal surface, in the fossil specimens described by Lagaaij (1953), they do not.

e. As in the Cupuladriidae, the regenerative budding in Lunulites is at right angles
to the lateral walls of broken fragments. This is in contrast to colonies of Selenaria
regenerated from fragments (New Zealand, Discovery Coll., Stn. 936), which show
spiral budding from the lateral walls of old zooecia. Each new bud is consistently

1 66

P. L. COOK

distal-lateral, and no zooecium produces new zooecia at right angles to its long axis
(see Text-fig. 36). The budding of deformed colonies of Selenaria is also spiral (see

P-

FIG. 5. Relationships of basal grooves. Broken lines indicate the position of the frontals
of the zooecia on the upper surface. A. Cupuladriidae. Grooves running radially in
the mid-line of the zooecia, branching at the beginning of an intercalary series. B.
Selenaria. Grooves running between the zooecia, branching around the first zooecium of
an intercalary series.

f. The basal radial threads and radial sector boundaries branch frequently. The
origin of a branch may be traced and found to correspond exactly with the occurrence
of the first zooecium of an intercalary row on the frontal surface. The budding of an
intercalary row must be of the same nature as that of all other zooecia, and thus the
radial nature of the threads and sector boundaries may also be presumed to reflect
the radial development of the zoarium (see Text-fig. 5 A) . The formation of the basal
surface in the Cupuladriidae is exclusive, but apparently similar forms are present in
other families. An unnamed species of Lunulites described and figured by Lagaaij
(1953 : 18, pi. 2, figs, 6a, b), has a basal surface superficially like that of C. guineensis,
and specimens of Lunulites radiata show alternating tuberculate ridges and grooves
similar to those of C. owenii. Selenaria bimorphocella (S. Australia, 1928. 9. 13. 80,
and off Adelaide, 1928. 9. 13. 81, Siboga Coll.), shows radial undulating grooves with
large pores, which are occluded at the centre of the zoarium, except beneath the
vicarious vibracula, where they remain open. In section, these pores do not show the
serial kenozooecial structure found in Group A of the Cupuladriidae (see below).
The grooves exactly outline the position of the zooecia above them, and, at the peri-
phery, show indications of distal and proximal links, which correspond in position to
the distal and proximal walls of the zooecia. Thus the grooves are entirely different
in nature to, and do not have the alternating relationships of, the grooves, threads, or
radial sector boundaries of the Cupuladriidae, and are only secondarily radial in
appearance (see Text-fig. 56).

THE CUPULADRIIDAE 167

7 GROUPING OF SPECIES

The groups defined below do not include all the species which have been described,
but formulate the correlation of various characters found to be common to some
species. C. pyriformis and C. indica (see below) combine some of the characters
of more than one of these groups.

The groups are arranged in order of increasing extent of cryptocystal development.

A. C. canariensis group. Cryptocyst simple, without denticles, central zooecia
without closures by calcareous laminae. Vestibular arch absent. Basal surface
consisting of rectangular sectors, with successive series of porous kenozooecial cham-
bers. Vicarious vibracula sometimes present. (Examples : C. canariensis (Busk),
C. monotrema (Busk), C. biporosa Canu & Bassler).

B. C. guineensis group. Cryptocyst simple, without denticles, central zooecia
closed by calcareous lamina developing proximally from below level of cryptocyst.
Vestibular arch absent. Basal surface of hexagonal or quadrilateral sectors, with
very small pores. Vicarious vibracula not present. (Examples : C. guineensis
(Busk), C. elongata Sakaura2.)

C. C. owenii group. Cryptocyst with denticles, central zooecia closed by lamina
developing distally from fusion of denticles. Vestibular arch present. Basal surface
with alternating ridges and grooves, tuberculate. (Examples : C. owenii (Busk),

C. haidingeri (Reuss), C. multispinata (Canu & Bassler), C. doma (d'Orbigny), ? C.
misrai (Tewari et al.s.)

D. D. iimbellata group. Horizontal cryptocyst lamina complete proximally,
with pores. Opesia with opesiules or opesiular indentations. Opesiaeof central area
closed by a complete lamina. Vestibular arch present. Basal surface with alter-
nating grooves and ridges, or short grooves and lozenges, tuberculate. (Examples :

D. reussiana (Manzoni), D. umbellata (Defrance), D. umbellata subsp. depressa (Con-
rad)).

There are two species which cannot be included in the groups defined above.
Although superficially alike, they are probably not closely related. C. Pyriformis
has a similar cryptocyst to that found in groups A and B, but differs from both in the
character of its basal surface, and in the possession of a distinct vestibular arch.
C. indica is close to C. guineensis, but differs completely in the character of its basal
surface ; its geographical distribution is also distinct.

Cupularia capriensis Waters differs from the Cupuladriidae in important respects
and is here referred to Setosellina (see p. 182).

2 Cupuladria elongata Sakakura (1935 : 6, pi. i, figs. 1-3) was described with extremely elongated basa
sectors, with small pores, from the Pleistocene of the Bozo peninsula, Japan. Recent specimens from
Japanese waters are all attributable to C. guineensis.

3 Discoporella misrai Tewari ct al. (1958 : 234, pi. 46, figs. 1-4, pi. 47 fig. 2), a Miocene fossil, was
described to include specimens from Kutch, India, and those previously reported from the Miocene of
Travancore as Cupularia spp. (Jacob & Sastri 1953 : 348, pi. 14, figs. 5a, b, pi. 15, figs, ya, b, pi. 16,
figs. 10, n). The specimens appear to have a denticulate cryptocyst and tuberculate basal surface.

168 P. L. COOK

8 Cupuladria pyriformis (Busk)
(PI. i, figs. 3, 4, Text-fig. iC)

Cupularia pyriformis Busk, 1854 : 100, pi. 124, St. Vincent (Windward Is.).
Cupuladria canariensis (Busk) : Sildn (part), 1942 : 14, pi. 4, figs. 15, 16, text-fig. 9, Anguilla,
Leeward Is., 40-150 m. (not text-fig. 8).

LECTOTYPE, chosen here, specimen on slide B.M. 1840. 10. 23. 80 (figured by Busk,
1854, pl- I24> basal surface), St. Vincent.

LECTOPARATYPE, 1840. 10. 23. 81, on the same slide.

OTHER MATERIAL. Naturhistoriska Riksmuseet, Stockholm, FBD 533, Anguilla,
Leeward Is., 45-150 m., Goes Coll., i specimen.

Zoarium small (3-5-4-5 mm. diameter). Cryptocyst without spines or denticles.
Vestibular arch well developed. Opercula without proximal sclerite. One pair of
circum-ancestrular zooecia aborted. Central zooecia closed by irregularly per-
forated lamina. Vibracula raised, with salient condyles. Basal surface with deep
grooves and raised ridges, with very small tubercles.

DIMENSIONS. Lz 0-35-0-40 mm. ; Iz 0-20-0-30 mm. ; Lop 0-12-0-17 mm. ; lop
0-18-0-27 mm. ; Lvo 0-09-0-12 mm.

C. pyriformis has been reported twice with certainty, and only 3 specimens are at
present known to exist in Museum collections.

The zooecia are narrow, tapering distally, with a vestibular arch distinctly raised
above the zoarial surface. The vibracula are also prominent, with narrow opesiae
and slightly hooked condyles. The central zooecia are closed by a thin, minutely
granular lamina, originating just below the salient, shelf-like part of the cryptocyst.
The lamina is perforated by 4-7 irregular pores, which in several cases are arranged
parallel to the edge of the cryptocyst, but which may also be scattered.

In all 3 specimens the circle of zooecia budded by the ancestrula is unlike that of
other Cupuladriidae in that pair No. i is nearly aborted. Each minute zooecium has
a small distal vibraculum (see Text-fig. iC). Silen (1942, text-fig. 9) showed only one
abortive zooecium and did not clearly indicate its relationship to the ancestrula, but
the figure shows the salient character of the vibracula.

The basal surface is deeply grooved, with sharp alternating ridges. The surface is
finely tuberculate but not porous. Busk described C. pyriformis with basal pores,
but the faint markings are in fact very small tubercles ; his figure shows the other
basal characters well, and they are the same in the specimen from Anguilla ; un-
fortunately Silen's photograph does not show the ridges which are in fact present
(see PI. i, fig. 4).

The position of the locality given by Busk for his material, " St. Vincent " (" St.
Vincents " on the slide label), is open to several interpretations. Although much of
the material from "St. Vincent " described in the 1854 Catalogue was from St.
Vincent, Cape Verde Islands, the specimens registered as 1840. 10. 22 ... and
23 ... consist almost entirely of slides labelled " St. Vincents, W.I.". There is a
St. Vincent Island off the west coast of Florida but the most likely locality is St.
Vincent Island in the Windward Islands. This supposition is strengthened by the

THE CUPULADRIIDAE 169

occurrence of the one other specimen from the neighbouring Leeward Islands. Silen
only doubtfully included this colony in C. canariensis , and examination of the speci-
men shows certainly that it belongs to C. pyriformis.

Studer (1889) listed specimens identified by Kirchenpauer from Madeira as
Cupularia pyriformis Busk, on p. 7 and from near Cape Verde as Cupularia pyriformis
Gray (sic), on p. 13. It is possible that Studer's specimens belonged either to C.
canariensis or to Cupuladria sp. (Cook (in press)), which, superficially, have similar
zooecial characters. Prof. E. Voigt (Geologische Staatsinstitut, Hamburg), has
informed me that Kirchenpauer's specimens were almost certainly destroyed during
the 1939-1945 war.

Lagaaij (i963a : 182, pi. 25, figs. 2a, b) mentioned a species of Cupuladria, probably
related to C. Pyriformis, from the Miocene of East Venezuela.

The character of the basal surface and the presence of a distinct vestibular arch
indicate an affinity between C. pyriformis and the C. owenii group of species ; but the
absence of denticles on the cryptocyst and the form of the central closures show a
similarity with the C. guineensis group.

9 Cupuladria indica sp. n.

(PL 2, figs. I, 2)

Cupularia umbellata Defrance : Hincks, 1887 : 125, Mergui Archipelago.

Cupuladria guineensis (Busk) : Hastings (part), 1930 : 714, 719 (specimens from Mergui, Burma).
Silen (part), 1942 : 8, pi. 4, figs. 13, 14 (specimens from locality i, Gulf of Aden).

HOLOTYPE. Mergui, Burma, B.M. 1899. 5. i. 23A, Hincks Coll.
PARATYPES, as above, 1899. 5. i. 23B-H, 1899. 5. i. 267, Hincks Coll.

OTHER MATERIAL. Mergui, 1963. 3. 18. 3, Anderson Coll.

" John Murray " Coll. (all specimens from a sandy or muddy substrate) : Stn. 80,
S.E. of Muscat, 22° 13' 30" N., 59° 49' 42" E., 3O.xi.33, 16-22 m., ZigiA. Stn. 77,
as above, 22° 13' 30" N., 59° 52' E., 421 m., ZI9&A. Stn. 178, N. of Somali Horn,
12° oo' 36" N., 50° 40' 06" E., 2.V.34, 91 m., Z20iA (with Anoteropora latiroitris
Silen). Stn. MB I d, Hanish Is., Red Sea, 13° 39' 30" N., 42° 43' E., I7.ix.33, 29 m.,
Zi46A and Z4iE. Stn. Mb I b, as above, Zi3gA. Stn. MB II c, Khorya Morya
Is., 17° 33' 30" N., 56° 01' 30" E., 28.x. 33, 26 m., Zi6iA.

Naturhistoriska Riksmuseet, Stockholm, FED 822, Gulf of Aden, 30 m. (with
Setosellina sp.), photograph registered B.M. 1963. 3. 14. i.

Zoarium frequently flattened, discoid, average diameter 7 mm. Cryptocyst simple,
well developed laterally, descending gently, edge serrate but not denticulate. Oper-
culum without proximal sclerite. Central zooecia closed by irregular denticles form-
ing a lamina at the level of the cryptocyst. Basal surface with radial ridges and
large, regular tubercles, no rectangular or hexagonal sectors or pores present.

DIMENSIONS. Lz 0-40-0-50 mm., av. 0-45 mm. ; Iz 0-30-0-40 mm., av. 0-35 mm. ;
Lop 0-25-0-30 mm., av. 0-28 mm. ; lop 0-16-0-21 mm., av. 0-19 mm. ; Lvo 0-12-
0-15 mm. ; Ls 0-50-0-70 mm.

ZOOL. 13, 5 . 9

170 P. L. COOK

Hincks described his specimens as C. umbellata and his material in the British
Museum was originally labelled as this species. Hincks noted that the majority of
the zooecia showed " no trace of a lamina ", but that the central zooecia were closed
by one. The peripheral pores in the cryptocyst of the closed central zooecia in C.
indica do somewhat resemble the opesiules of D. umbellata, as does the basal surface
with its tuberculate ridges and grooves, of which Hincks gave a very full description.

The specimens show little variation ; just over half of the zoaria are extremely
flat, the remainder slightly more conical. The central closures differ from those of
C. guineensis in that they are formed from denticles or laminae which appear to
originate from the salient shelf-like edge of the descending cryptocyst, not just below
it (see p. 159). This is particularly noticeable at the proximal edge of the opesia,
which is the last part to be occluded. However, the later thickening results in a very
similar appearance in both species. Young colonies, and those formed from regenera-
ted fragments, have no closed zooecia. Generally, the cryptocyst is narrower than
that of C. guineensis.

The basal surface shows little variation in the size of the tubercles, and the radial
grooves are marked by a slight thread in all specimens, which distinguish C. indica
from those specimens of C. guineensis which have heavy basal tuberculation.

The basal structure of C. indica is similar to that of the C. owenii group. There
are no sectors, and no pores in the basal lamina, which is not thickened as in C.
guineensis. In other respects, C. indica closely resembles C. guineensis, and its
zooecial dimensions fall within the range of varation of C. guineensis ; they are,
however, consistently at the smaller end of that range.

The two species also appear to be allopatric, their geographical distributions being
mutually exclusive. The occurrence of the conical Anoteropora latirostris and the
selenariiform Setosellina sp. from the same stations as the lunulitiform C. indica
illustrates the correlation between various types of free-living zoaria with a sandy
or muddy substrate, as discussed by Harmer (1957 : 649).

10 Cupuladria guineensis (Busk)
(PI. i, figs, i, 2, PI. 2, figs. 3, 4, Text-fig. 6)

Cupularia guineensis Busk, 1854 : 98, pi. 114, figs. 1-5, New Guinea. Busk, 1884 : 206, pi. 14,
fig. 6, N.E. Australia, 8 fath. Marcus, 1921 : 8, 2° 30' S., 107° 10' E., 15 Fd.

Cupularia stellata Busk, 1854 : 99, pi. 118, figs. 1-4, Philippines. Marcus 1921 : 8.

Cupularia canariensis Busk, Waters, 1885 : 308, Aldinga, Tertiary, S. Australia. 1887 : 201,

Princess Charlotte Bay, N.E. Australia, 12 fath. 1921 (part) : 410. 1926 (part) : 426, pi. 18,
fig. 10, Princess Charlotte Bay.

Cupuladria guineensis (Busk) Harmer, 1926 : 266, text-fig. 9, pi. 16, figs. 5-7, S. of Mindanau,
0-36 m. Canu & Bassler, 1929 : 74. Hastings, 1930 : 714 (not specimens from Mergui,
= C. indica). 1932 : 412, Great Barrier Reef. Silen, 1938 : 366, pi. 17, fig. 119. 1941 : 61,
East China Sea, 90 m. (cf. locality 7 in 1942). 1942 (part) : 8, text-figs, 5-6, pi. 3, figs. 10-12,
localities 2-7 (not locality i, pi. 4, figs. 13-14 =C. indica). Androsova, 1963 : 23, 39, pi. i,
fig. 3, S. China Sea.

Cupuladria intermedia Canu & Bassler, 1929 : 74. Hastings, 1932 : 412.

Cupuladria brevipora Canu & Bassler, 1929 : 74. Hastings, 1932 : 412.

THE CUPULADRIIDAE 171

Cupuladria transversata Canu & Bassler, 1929 : 74, 75, pi. 3, figs. 3, 4. Stn. D 5579, Borneo 175

fath., sand.
Cupuladria dentifera Canu & Bassler, 1929 : 74, 75, pi. 3, figs. 5-8, Stn. D 5230, Limasaua Is.

118 fath., sand.
Cupuladria tuberosa Canu & Bassler, 1929 : 76, pi. 4, figs. 1-4, Stns. D 5134, Sulu Archipelago,

25 fath., sand, and D 5144, Jolo, 19 fath., sand.
Cupuladria grandis Canu & Bassler, 1929 : 77, pi. 4, figs. 10-13, Stn. D 5161, Sulu Archipelago,

1 6 fath., sand.
Cupuladria granulosa Canu & Bassler, 1929 : 74, 78, pi. 4, figs. 5-9, Stn. D 5358, Jolo Sea, 39 fath.,

mud. Kataoka, 1961 : 224, pi. 30, figs. 2, 5, Pleistocene, Japan.
Cupuladria hexagonalis Canu & Bassler, 1929 : 74 (as hexagona) 78, pi. 5, figs. 1-4, Stns. D 5141,

Jolo, 29 fath., sand, and D 5147, Sulu Archipelago, 21 fath., sand.
Cupuladria microdenticulata Kataoka, 1961 : 225, pi. 29, figs, i, 2, Pleistocene, Japan.

HOLOTYPE, B.M. 1842. ii. 30. 14, New Guinea.

OTHER MATERIAL. Where the predominant type of basal sector present is hexa-
gonal (see Text-fig. 6B), the specimens listed below are marked "A", where it is
nearly square (see Text-fig. 61)), they are marked " B " ; where pores have been
seen, and where tubercles are present, the letters " P " and " T " are added.

British Museum. New Guinea 1842. u. 30. 14 (APT). 1859. 2. 25. 11 (BP).
S. of Mindanau, 15 fath. 1928. 3. 6. 74 (APT). Torres Straits 1862. 6. 5. 14 (BP),
10 fath., sand. 1882. 2. 23. 37-40 (BP). ?N. Australia 1855. 12. 27. 38 (APT),
1872. 6. 21. 21 (B). Holothuria Bank, N.W. Australia, 34 fath., 1892. i. 28. 125
(BPT). 1963. 2. 2. i (BP), 1891. 12. 16. 1-4 (BP). Cossack Is., N.W. Australia,
8 fath., 1891. 12. 16. 31-37 (BP), Baudin Is., 15 fath., 1891. 12. 16. 21 (BP), 22
(BP), 23 +24 (B + AP), 25-30 (BP + BPT), 1892. i. 28. 128. 8-15 fath., (B +
APT). Cape York N.E. Australia, 10° 30' S., 142° 18' E., 8 fath., sand, Chall.
Stn. 186, 1887. 12. 9. 819 (BP). Low Is. off Port Douglas, Gt. Barrier Reef, 1932 .
4. 20. 112 (BPT). Bass's Straits, S. Australia, 1899. 7. i. 1269. Busk Coll.
(BPT) Australia 1929. 5. 10. i (AP), 1899. 7. i. 1252 Busk Coll. (labelled C. stellata).
Philippines, 1854. I]C- I5- I47 (tyPe °f C. stellata), 1899. 7. i. 1231, Busk Coll.
(AT). Philippines, Stn. D. 5161, 1931. 12. 30. 9 (AP) and Stn. D 5156, off Tinakta
Is., 18 fath., 1931. 12. 30. 10 (AP) (topotypes of C. grandis), Canu & Bassler Coll.
Sulu Archipelago, " Albatross " Stn., 1)5134, 1963. 3. 14. 2 (AP) ; Suluade Is., Stn.
D 5147, 1963. 3. 14. 3 (APT) ; and Philippines, 26 fath., Stn. D 5181, 1963. 3. 14. 4
(AT), Bassler Coll. Photographs of type specimens from the Bassler Collection,
United States National Museum, all X 20. C. transversata, Holotype, 1963. 3. 14. 5
and paratypes, 1963. 3. 14. 6 (AP). C. dentifera, Cotypes, 1963. 3. 14. 7 (APT).
C. grandis, Syntypes, 1963. 3. 14. 8 (AP). C. granulosa, Syntypes, 1963. 3. 14. 9
(APT). C. hexagonalis, Syntypes, 1963. 3. 14. 10 (APT). C. tuberosa, Syntypes,
1963. 3. 14. ii (AP).

Manchester, Museum, Waters Coll. Princess Charlotte Bay, 12 and 13 fath.,
Australia, (BP).

Naturhistoriska Riksmuseet, Stockholm. Forty-five miles W.S.W. off Cap
Jaubert, N.W. Australia, 100 m., FBS 157 (BPT), FBS 109 (BPT). S. China Sea,
between Nagasaki and Hongkong 75 m., mud, FBS 904 (BPT). Malayan Archi-
pelago, Java Sea, 2° 30' S., 107° 10' E., 27 m., FBS 639 (AT).

ZOOL. 13, 5. 9§

172 P. L. COOK

United States National Museum. The Museum was unable to lend material named
by Canu & Bassler, but made available photographs of the type specimens (see
above) and also unnamed specimens from various " Albatross " Pacific Stations,
2 of which were the same as those from which some of Canu and Bassler's species
were obtained. D 5134 Sum Archipelago, 6° 44' N., 121° 48' E., (APT)
(Locality for C. tuberosa). D 5147 Suluade Is., Sulu Archipelago, 21 fath., (APT)
(Locality for C. hexagonalis). D 5145, Jolo, 23 fath. (APT). D 5159, Philippines
(AP). D 5181, Philippines 26 fath. (APT). D 5314, China Sea nr. Hong Kong,
122 fath. (AT).

Zoarium frequently very large (maximum observed diameter 24 mm.). Zooecia
with broad tuberculate cryptocyst, opesiae often almost rectangular. Operculum
without a basal sclerite. Central area of zooecia closed by growth of a lamina from
beneath the shelf-like salient part of the cryptocyst. Basal surface smooth or
tuberculate, divided into quadrilateral or hexagonal sectors, variable. Pores present
in peripheral sectors, becoming sunk in pits, occluded or hidden by tubercles.

DIMENSIONS. Lz 0-40-0-75 mm., av. 0-60 mm. ; Iz 0-30-0-50 mm., av. 0-40 mm. ;
Lop 0-30-0-40 mm., av. 0-35 mm. ; lop 0-15-0-25 mm., av. 0-20 mm. ; Lvo 0-12-
0-20 mm. ; Ls 0-5-1-5 mm.

C. guineensis was denned by Hastings (1930 : 714-5), who distinguished it from the
C. canariensis-complex of species (Cook (in press)) by the nature of the basal surface
and the extent of the cryptocyst. Of the 8 species introduced by Canu & Bassler
(1929 : 74-9), two, C. intermedia and C. brevipora, were placed in the synonymy of
C. guineensis by Hastings (1932 : 412), who also agreed with Marcus (1921 : 8) in
regarding C. stellata Busk as the young form of C. guineensis.

Before considering the remaining species introduced by Canu & Bassler, the charac-
ters of the material described by Busk in 1854 and 1884 must be re-examined. Two
groups are evident in the large number of specimens available (see list above) ;
these differ principally in the pattern of the basal sector boundaries, and fall very
roughly, but not exclusively, into two areas of geographical distribution.

The first, more northerly form (A), is that originally described by Busk. The
basal sectors are hexagonal, pierced by very small pores, which are frequently
obscured by tubercles. Busk described the basal surface as " divided into hexagonal
areas, each corresponding to a cell ", and his fig. 3 showed irregular, tuberculate,
hexagonal sectors ; he did not mention the pores, which are, however, visible at the
periphery of the holotype. The sectors, as described above (p. 162) do not, in fact,
each correspond to a zooecium, but do have a definite relationship with the zooecia
frontal to them (see Text-fig. 6) . Busk's specimen from the Challenger Collection is
of a second type (B), the basal sectors being quadrilateral, and the surface smooth.
Pores are visible in the peripheral zooecia, which become immersed in pits and finally
occluded toward the centre of the colony. The basal surface of his specimen was
described by Busk " with shallow radial sulci, ridges quite flat, divided into moveable
quadrangular areas, each of which has four or five shallow pits ". Figure 6b showed
these, and (p. 207) Busk mentioned that they might " represent occluded pores ".

THE CUPULADRIIDAE

173

Busk's description of " moveable " areas may have referred to a dry, loose, basal ecto-
cyst, which, like the calcareous surface, shows the sector boundaries, although his
specimen from the Challenger Collection has no ectocyst present. However, it may
perhaps have been a reference to the variation in shape of the basal sectors of other
specimens Busk had examined.

Both types of basal sector may be modified by tuberculation which obscures the
sector boundaries and pores. Zoaria of the B-type tend to be less tuberculated, and
are also usually larger than the A-type. In several cases they show central areas
with irregular and hexagonal sectors (e.g. 1891. 12. 16. 23 + 24 and 1892. i. 28.
128).

The variation in zooecial size and proportion, the extent of the cryptocyst, and the
shape of the zooecial and vibracular opesiae appear to include all the forms listed in
the synonymy above. Canu & Bassler introduced C. intermedia for C. guineensis
Busk, 1884, which they distinguished from the holotype by the possession of porous
basal sectors. As noted above, pores are present in both A and B forms, and this is
not a specific distinction. C. brevipora was introduced for C. guineensis Harmer, but
the Siboga specimens are not distinct (see Hastings, 1932 : 412).

The remaining 5 species described by Canu & Bassler all have hexagonal basal
sectors and are only distinguished from each other by the form of the opesiae and the
degree of basalt uberculation. Silen (1942 : 8-9) remarked on the variations within
the limits of the species and on the similarity of character given in Canu & Bassler's
descriptions. One species was introduced on the evidence of one fragment of a
zoarium (see Canu & Bassler 1929 : 75), and examination of photographs of the type
specimens, and of unsorted Philippine material, shows that all 5 species fall within a
continuous range of variability.

C. transversata, which was described from a single fragment, showed " a great
variety of aspect of the inner face ", and had a smooth basal surface with some
peripheral tubercles, and pores. The figure shows a somewhat worn, moderately
tuberculate colony, and photographs of the holotype (1963. 3. 14. 5, which comprises
only 32 zooecia), show that it has characters extremely like those of the photo-
graphs of C. hexagonalis.

C. dentifera was described as differing from C. stellata in its tuberose basal surface.
Busk's fig. 3 shows C. stellata without basal tubercles, but one of the specimens
numbered 1854. I][- I5- X47» is mounted on glass, and it may be seen that fine
tubercles are present. Specimens from the type-locality of C. tuberosa show similar
ranges of variation in size of colony, zooecia, opesiae and tuberculation of basal sur-
face, as those from the type-locality of C. hexagonalis. Canu & Bassler's figure of the
frontal surface of C. hexagonalis is of a worn specimen, and while some of the zoaria
from Stns. D 5147 and D 5314 resemble it, others are very similar to the figures of
C. transversata. Photographs of the cotype specimens of C. dentifera show that Canu
& Bassler's figure (pi. 3, fig. 7) has been retouched, and that the irregular shape of the
opesiae is due to the development of the cryptocyst laminae closing the zooecia.

C. granulosa, in which the sector boundaries were ' scarcely visible ", is an extremely
tuberculated form. The specimen FBS 639 shows this type of development, but the

FIG. 6. Basal sectors in Cupuladria guineensis. Ovals in broken lines indicate the position
of the frontals of the zooecia on the upper surface. A. " C. dentifera ". Sector boun-
daries little developed. B. C. guineensis, " A " form. Hexagonal sector boundaries
divided by radial boundaries. C. " C. transversata ". Wide hexagonal sectors where
the radial sector boundaries are intermittently obscured. D. C. guineensis, " B " form.
Quadrilateral sectors.

sector boundaries may be seen at the periphery of the colony, and by lighting the
surface obliquely. Heavily tuberculated variants of the granulosa-type are super-
ficially similar to C. indica, but may be distinguished by the absence of any radial
threads on the basal side (see above).

The features distinguishing C. grandis were its large zoarium and smooth basal
surface. The largest zoaria of C. guineensis examined are all shallow, with square
basal sectors ; the topo types of C. grandis are large, strongly domed, and have hexa-
gonal sectors. Domed, B-type zoaria, of similar size (e.g. FBS 157, 18 mm.) and
flatter, smaller, A-type colonies (e.g. 1928. 3. 6. 74) indicate that zoarial size and
shape are not correlated with type of basal sector, although there is a tendency to
reduction of tuberculation in large zoaria of both types. Canu & Bassler figured, but
did not comment further on, two distinct forms of vibracular seta in C. grandis ;
"recurved" (text-fig. I3A) and "rectilinear" (text-fig. 136). Two kinds of seta
have been found in C. canariensis s.s., the differences in which are reflected in the
calcareous parts (Cook (in press)). No sign of any such dimorphism has been seen
in any of the many colonies with setae, or in the opesiae of specimens of the topotypes
of C. grandis and the other specimens attributed to C. guineensis. The seta in Canu
& Bassler's text-fig. I3C somewhat resembles that of a regenerating seta figured by
Marcus & Marcus (1962, pi. 2, fig. 7).

The variation in type and degree of calcification of the curved laminae may result
in differing appearances of the basal surface. Where the hollows between the
laminae are not filled in by secondary calcification, and where the pores are obscured,
the surface is like that described in C. dentifera (see Text-fig. 6A). Where the radial
sector boundaries are intermittently obscured, large, wide, hexagonal sectors are
produced, as in C. transversata (see Text-fig. 6C). Differential development of
tuberculation and pitted pores, together with the effects of wear, produce many other
intermediate types of basal surface, but the characters of the zooecia are similar in all
zoaria.

THE CUPULADRIIDAE 175

The specimens described as C. granulosa and C. microdenticulata by Kataoka
(1961 : 224, 5) appear to be referable to C. guineensis. The basal surface of both
species had sectors, which were porous in C. microdenticulata, and the zooecia had the
well-developed cryptocyst of C. guineensis.

The zoarium of C. guineensis attains a larger size than that of any other species in
the Cupuladriidae. The largest colonies (Torres Straits 1862. 6. 5. 14 and S. China
Sea FBS 904) are 24 mm. in diameter, and many of the other zoaria, especially those
with square basal sectors, exceed 15 mm. in diameter.

The size of zooecia varies considerably, but the Lz does not seem to exceed 0-75
mm. and the Iz 0-50 mm. The range of Lz given by Canu & Bassler for C. transversata
includes the dimensions given for all the other species. Measurements taken from
their figures indicate that the Lz for C. dentifera, C. grandis and C. gmnulosa may be as
low as 0-50 mm., which is the figure given in the text for the Lz of C. transversata,
C. tuberosa and C. hexagonalis. The opesiae of these last 3 species are rectangular in
contrast to the opesiae of the first 3 species, which are often oval, but A-type zoaria
having zooecia with both forms of opesiae are present from Stn. D 5181, and B-type
zoaria with the same degree of opesial variation from the Holothuria Bank 1963. 2.
2.1, and Low Island, 1932. 4. 20. 112. Generally the more peripheral zooecia of the
larger zoaria of both types tend to have more oval opesiae and more steeply descending
cryptocysts. Some zooecia have distinctly serrated cryptocysts.

As noted above, the vibracular setae appear to be of one kind, and are similar to
those found in all other species of the family. Proportionately they are shorter than
those of Discoporella umbellata, the larger zoaria of which approach those of C.
guineensis in size.

Two forms of C. guineensis therefore exist, but the characters of each are not
exclusively correlated and cannot be regarded as specific.

ii THE DISCOPORELLA U M B ELL AT A -COMPLEX

The characters defining species in the Cupuladriidae, where the common zoarial
form determines similarity in the development of all structures, must of necessity be
judged using fine criteria. To define a species adequately there must be a recog-
nizable degree of correlation between each character considered, morphologically,
geographically, and if possible, palaeontologically. In each of these respects it is
possible to divide records of D. umbellata into two distinct groups. The problem is
whether these groups are of specific or subspecific rank, and the principal criterion
applied below is that of character-consistency throughout the time range of the group
considered.

Canu & Bassler (1923, text-fig. 13) noted the variability of their material assigned
to D. umbellata, and figured wide zooecia in Miocene material from Santo Domingo
(pi. 2, fig. 18) . The part of the zoarium figured showed the growing edge of a regener-
ating fragment, where the zooecia of all species are frequently wide and slightly
distorted (see pp. 155, 158). Photographs of this material (1963. 3. 14. 12) show that
subsequent zooecia are narrower. Canu & Bassler's figures (including fig. 17)
emphasized the different appearance of Recent specimens from Oran (p. 76, text-

176 P. L. COOK

fig. I3A) and the increase in size of the zooecia compared with those of Miocene
material from France (text-figs. 136-6).

C. robertsoniae was introduced by Canu & Bassler for specimens named C. canarien-
sis by Robertson (1908 : 314). Her figures showed narrow zooecia with 5-7 opesiules
and were certainly referable to the D. umbellata-complex (see Hastings, 1930 : 719).
In renaming Robertson's specimens Canu & Bassler were apparently aware of basic
differences between " typical " D. umbellata and their Pleistocene material from the
Pacific coast, which was included in C. robertsoniae, as they gave the narrow zooecia
and large, proximally curved opesia as diagnostic characters. Comparison of Canu
& Bassler's figures of their Pacific Pleistocene material (1923, pi. 34, figs. 5-7) with those
they gave of West Indian Miocene specimens (1923, pi. 2, figs. 15-19, as C. umbellata)
shows striking similarities and indicates a strong affinity between the two groups of
specimens.

Osburn (1950 : 113) concluded that the wide range of variation found in American
specimens justified their inclusion in one species with European D. umbellata, but
apparently he did not compare his specimens with European material.

Generally, fossil material from the Miocene of Europe and America shows con-
sistently correlated differences in character, which have become even more divergent
in Recent specimens from the two areas (see PI. 3, figs. 1-4). Applying the criteria
used in defining the other species described above, the American fossils cannot be
included in typical D. umbellata, and, as the Recent specimens differ still further, all
American records have been separated here as D. umbellata subsp. depressa (Conrad).

The constant characters of the European material are the wide zooecia, the presence
of a pair of opesial denticles, the proportionately large size of the vibracular opesiae,
and the numerous small opesiules. There has been an increase, in size in Recent
specimens, where the proximal edge of the vibracular opesia is denticulate, and the
smaller pores in the centre of the horizontal cryptocyst persist.

The constant characters of the American material are the narrow zooecia, the non-
denticulate opesiae, with curved proximal edge, the relatively small size of the vibra-
cular opesia, which is never denticulate, and the smaller number of large opesiules.
Some fossil specimens have central cryptocystal pores, but these are rarely persistent
in Recent material. In contrast to D. umbellata, the zooecia of fossil specimens of
D. u. depressa are, on average, larger than those of Recent colonies.

It might be argued that the narrower zooecia of D. u. depressa themselves directly
result in the smaller number of opesiules and the lack of central cryptocystal pores,
but, as the two species exhibit similar ranges in size and shape of zoarium, there must
be some inherent difference producing the narrower zooecia of D. u. depressa. The
nature of the zooecial and vibracular opesiae are, however, not affected by conditions
of growth or width of zooecia, and these are the most important features which show
correlated character-consistency in the two forms.

When the peyroti-type zoaria listed below are included, records of D. umbellata
show three separate mean types of population. These forms may overlap at
the ends of their ranges of variation, but the Recent forms of each group are geo-
graphically distinct. By the application of a discriminant function technique to

THE CUPULADRIIDAE 177

further morphometric studies, it is intended to investigate the nature and degree of
relationship of the members of this complex.

12 Discoporella umbellata (Defrance)
(PL i, fig. 7 ; PL 3, figs, i, 3, 5, 6 ; Text-fig. 4)

Lunulites umbellata Defrance, 1823 : 361, pi. 47, figs. I, la, ib. " Lunulite en-parasol " de

Blainville, 1834 : pi. 72, fig. la, ib.
Discoporella umbellata (Defrance) d'Orbigny, 1853 : 473, pi. 717, figs. 1-5. Lagaaij, 1953 : 16,

pi. i, fig. 3, Miocene, Netherlands. Buge (part), 1957 : J77> pi- IO- ng- 4- Miocene, S.W. France.
Discoporella berardana d'Orbigny, 1853 : 474, Recent, Algeria, 14 m.
Cupularia lowei Busk, 1854 : 99, pi. 116, figs. 1-6, Madeira. C. lowei Gray (sic) Norman,

1909 : 290, pi. 37, figs. 7-12, Madeira 50-70 fath. Waters, 1921: 412, pi. 30, figs. 1-6, 26-29,

Madeira.
Cupularia umbellata (Defrance) : Manzoni, 1869 : 26, pi. 2, fig. 16, Pliocene, Italy. 1875 : 39,

pi. 5, fig. 67, Miocene Italy. Calvet 1907 : 393. Canaries, 80 m., and Cape Verde Islands,

1900 m. Canu, 1909 : 448, pi. 16, figs. 16, 17, Burdigalien, Helvetien, Miocene, S.W. France.

Waters, 1921 : 414, Pliocene, Italy. Canu & Bassler, 1923 : text-figs. I3A, Oran, Recent ;

B. C. D. E France Miocene, ?I Oran, Recent (as C. multispinata) .
Peyroti-type zoaria : Cupularia peyroti Duvergier, 1921 : 124. 1924 : 19, pi. i, figs. 6— 10,

Helvetien, Miocene, France.

Cupularia umbellata Defrance : O'Donoghue, 1924 : 39, Cape St. Blaize, S. Africa.
Discoporella umbellata (Defrance) part Hastings, 1930 : 719, specimens from S. Africa.

MATERIAL. British Museum. Bay of Funchal, 1892. 8. 6. i (type material of

C. lowei}. Madeira, 1911. 10. i. 648, 1004 and 1963. i. 16. 10 and n, Norman Coll.
Madeira, 1932. 8. 23. i. Canaries, 1899. 7. i. 1250, Busk Coll. Near Cape Frio,
Angola, 12° E., 18° 30' S., 1899. 7. i. 1241, Busk Coll. (doma-type zoarium).

Palaeontology Dept. Coll. D. 25005, 25003, Burdigalian, L. Miocene, Bordeaux.

D. 39047-96, Miocene, Reinbeck, Germany. D. 38947-9, Miocene, Beeringen.
D. 39884-91 (pt), Albena, Bordighera, Pliocene, Italy.

Voigt Coll. Reinbeck, L. Miocene. Boring, Bockstedt, Middle Miocene (i doma-
type zoarium present).
Peyroti-type zoaria.

MATERIAL. British Museum. Cape St. Blaize, S. Africa, 1923. 7. 26. 9, O'Donog-
hue Coll. S. Africa, 1963. i. 16. 9, O'Donoghue Coll. S. Africa, 34° 29' S., 21° 02'
E., 26. ii. 59, 73 m., sand, SCD ii4C, 1963. 1. 16. 6 ; 34° 29' S., 22° 18' E., 28. 8. 60,
io6m., sand, SCD 1470, 1963. i. 16. 7; False Bay, 31. i. 59, 50 m., FAL 34iD (B),
1963. i. 16. 8, Univ. Cape Town Ecological Survey Coll. Cape of Good Hope, 1842.
12. 2. 20, Belcher Coll. S. Africa, Cape Infanta N.E. by N|N, 13-5 miles, 43 fath.,
1949. ii. 10. 618 ; Stalwart Point N.N.W. 9 miles, 53 fath., 1949. n. 10. 652A ;
Nanquas Peak N. 46° E., 19 miles, 63 fath., 1949. ii. 10. 674A, and N.E. |N., 15
miles, 59 fath., 1949. ii. 10. 675 (including 2 doma-type zoaria), N.W. £N. 15 miles
49 fath., 1949. ii. 10. 668 ; Cape Infanta N.E. by Nig miles, 46 fath., 1949. ii. 10.
622A, 623A, N.E. by N.|N. 14-5 miles, 45 fath., 1949. ii. 10. 620 ; Cape St. Blaize
N. 7-5 miles, 37 fath., 1949. ii. 10, 624A, 62$A, 626A, 62^A and 94A, N.E. JN. n

178 P. L. COOK

miles, 40 fath., 1949. n. 10. 629 ; Tugela River mouth, N.N.W. 11-75 miles, 37 fath.,
1949. ii. 10. 644 (including 2 doma-type zoaria) ; Umtwalumi River mouth N. by W.
7 miles, 50 fath., 1949. 11. 10. 651 ; False Island N. 8-5 miles, 53 fath., 1949. 11. 10.
66gA. ; Glendower Beacon N.|W. 6 miles, 39 fath., 1949. n. 10. 660, Burrows Coll.
John Murray Coll. (all from sandy substrates). Stn. 103, 5° 39' 30" S., 39° n' 30"
E.,N. of Zanzibar, ii. 1.34. 101 m., Z. 198 C. Stn. 104., 5° 37' 54" S., 39° n' 36" E.,
ii. i. 34, 207 m. Z. I99A.

Zoarium usually large (maximum observed diameter 22 mm.). Zooecia often as
wide as long. Opesia small, with straight proximal border with a pair of minute
denticles. Operculum with a straight proximal sclerite. Descending cryptocyst
wide, tuberculate, horizontal cryptocyst lamina with some small irregular central
pores and 8-14 small opesiules, frequently slit-like. Vibracular opesia very large
(X 1-75 of Lop), finely denticulate proximally in Recent specimens. Basal surface
with short grooves, pits and ridges, tuberculate, or smooth, with a layer of keno-
zooecia and vibracula encroaching from the frontal surface.

DIMENSIONS. Lz 0-50-0-70 mm. ; Iz 0-40-0-63 mm. ; Lop 0-10-0-15 mm. ; lop
0-12-0-20 mm. ; Lo o-io-o-n mm. ; lo 0-12-0-15 mm. ; Lvo 0-18-0-25 mm. ;
Ls 0-6-1-45 mm.

The records of European Miocene and Pliocene specimens agree in describing and
figuring D. umbellata with wide zooecia and a large number of opesiules. Although
generally the zooecial measurements of fossils are smaller than those of Recent
specimens, the proportions and general characters of the species seem constant, and
are certainly distinct from those of D. u. depressa.

The majority of zooecia are as wide or occasionally wider than long, and the
number of opesiules is most frequently between 8 and 10. Defrance figured 8-9,
d'Orbigny 10, de Blainville 8-9, and Manzoni (1869), 10-11, although the number
given in 1875 was 7. In Recent specimens the number is often as many as 14, and
the opesiules are slit-like. D'Orbigny described, as D. berardana, Recent specimens
from Algeria with denticulate cryptocysts, transverse, semi-lunar orifices, and very
large vibracular opesiae. D. umbellata was figured by Canu & Bassler (1923, text-
fig. I3A) from Oran, and it is possible that d'Orbigny's record referred to worn speci-
mens of this species. Canu & Bassler's photograph of C. multispinata (1923, text-
fig. 13!.) showed opesiules and a basal sclerite in the operculum, and is perhaps also
of D. umbellata .

The specimens described as C. lowei were distinguished from Z). umbellata by the
deformation of the zoarium (see p. 158, and Marcus & Marcus, 1962 : 294).

The paired denticles on the proximal edge of the opesia are present in all well-
preserved fossil and Recent material. Waters (1921 : 415) noted their occurrence,
and Lagaaij (1953 : 16) included them in his diagnosis of the species.

The proximal edge of the vibracular opesia of Recent specimens is finely denticulate
(see pi. 3, fig. i), a character which has not been seen in fossil material. The vibra-
cular opesiae are extremely large, as noted by d'Orbigny (1853 : 474, in D. berardana},
their length being significantly greater than that of the zooecial opesiae,

THE CUPULADRIIDAE 179

The operculum is strongly flanged laterally, with a straight, robust, proximal
sclerite.

The specimen from Cape Frio has a doma-type zoarium. The basal surface is solid
and there are peripheral kenozooecia, but the zooecia are worn, and the character of
the opesiae cannot be clearly seen. However, the width of the zooecia (Iz 0-3-0-48
mm.) and the number of the opesiules (5-7), are nearer the higher end of the range of
variation than those of many of the corresponding doma-type specimens from the
west Atlantic (see below), and thus reflect the differences found in zooecia of normal
zoaria of the two forms. One fossil zoarium (Voigt Coll.) is also extremely worn, but
shows two rows of peripheral vibracula and a basal surface almost filled by secondary
calcification.

The development of the cryptocyst in the young peripheral zooecia shows the close
relationship of D. umbellata both to D. u. depressa and to the denticulate species of
Cupuladria (group C). Fine, irregular denticles grow first from the proximal and
lateral edges of the cryptocyst and then develop progressively in a distal direction. At
this stage the zooecia greatly resemble specimens of C. owenii and C. doma. Further
calcification produces a distal bar cutting off the opesia, and, as the more proximal
denticles fuse, leaving a series of denticulate, irregular opesiules, and a number of
pores in the centre of the horizontal cryptocyst lamina thus formed (see also Marcus
& Marcus, 1962, pi. i, fig. 4). The appearance of the zooecia at this stage is then
very similar to that of Miocene specimens of D. u. depressa from Oak Grove, Florida,
where this type of development was the final stage. Other specimens of D. u. depressa
from the Miocene of Cercado de Mao, Santo Domingo, show more complete calcifica-
tion, with rounded opesiules and no central pores. The calcification of eastern
Atlantic fossil and Recent specimens of D. umbellata is even greater, and the opesiules
are further reduced.

The distribution of D. umbellata s.s. is now confined to the northern and north
western coasts of Africa, and the neighbouring islands.

D. umbellata and C. haidingeri are frequently associated in samples from the
European Miocene. When worn they are extremely difficult to distinguish from
each other. The basal surface of D. umbellata is not as regularly grooved as that of
C. haidingeri, and generally the zooecia of D. umbellata are wider. Even when the
central part of the cryptocyst lamina is broken, the vestibular arch surrounding the
distal part of the opesia is smaller and more incurved in D. umbellata than in C.
haidingeri. At the edge of the central area of closed zooecia the most recent closured
frequently show the straight bar, proximal to the opesia, which is absent in C.
haidingeri.

Recent specimens from south and east Africa correspond almost exactly with
C. peyroti as described by Duvergier. The dimensions he gave fall within the range
of those now obtained from a large number of specimens, and the photograph shows
kenozooecia exactly like those of the Recent material. In one respect, Duvergier's
account may be confusing, as he followed Canu is describing the frontal, convex
surface of the colony as basal, and the concave or planar kenozooecial surface as
frontal, Q'Donoghue mentioned the invasion of the basal surface by kenozooecia

180 P. L. COOK

and vibracula, and Hastings, who examined some of O'Donoghue's material, expressed
doubt as to its inclusion in D. umbellata s.s. The Miocene specimen of D. umbellata
figured by Buge shows a hollow basal surface covered by kenozooecia and vibracula.
The specimens from south Africa show the complete range of development, those
from east Africa, the young stages only. Young zoaria are flat, with a solid, smooth
base, and older colonies are more domed, with the basal surface either concave (e.g.
1963. i. 16. 6-9, 1949. n. 10. 625A) or completely filled with secondary calcification
(e.g. 1949. ii. 10. 66gA and 652A). In neither state is the surface ridged or tuber-
culate. The budding of the basal kenozooecia appears to be random and irregular,
and the calcified cryptocyst is developed like that of the zooecia, from the fusion of
irregular denticles.

The zooecial opesiae are often slightly curved proximally, and the denticles are often
reduced or absent, but the opesiae are never sinuate as in D. u. depressa. The horizon-
tal cryptocyst lamina differs from that of typical D. umbellata in the large number of
frontal pores, which are often indistinguishable from the marginal opesiules. The
operculum is like that of D. umbellata, and is flanged laterally, with a strong, straight
proximal sclerite.

A few colonies combine some characters of both doma- and peyroti-type growth.
They are small (diameter 2 mm.) and high, with solid, smooth bases. The peripheral
zooecia are closed, but only one row of peripheral vibracula is present, and there is no
invasion of the basal surface by kenozooecia. The zooecia are small, approximately
half the length of zooecia in a normal colony, but the opesiae are not reduced in
proportion, being about f of the length of normal opesiae. The zooecia have an average
of 8 opesiules and a number of irregular frontal pores in the cryptocyst. The
proximal edge of some vibracular opesiae are finely denticulate. The doma-type
colonies thus have many of the characters of the normal zoaria from the south east
African coast. Their occurrence does not seem in this case to be correlated with low
temperature, as all the records are confined to warm waters, extending from the
warm-temperate zone, eastward to the subtropical zone, as described and discussed
by Stephenson (1947 : 215).

As noted above (p. 163), no specimens of typical D. umbellata have been found
from the south and east coasts of Africa, nor have peyroti-type zoaria been recorded
from the west African coast. Specimens of Cupuladria owenii (Gray) and C. multi-
spinata (Canu & Bassler), both of which also occur from these two areas, show no
examples of peyroti-type growth.

13 Discoporella umbellata subsp. depressa (Conrad)
(PI. 3, figs. 2, 4)

Lunulites depressa Conrad, 1841 : 348, Miocene, N. Carolina.

Discoporella denticulata (Conrad) Gabb & Horn, 1862 : 142, pi. 20, fig. 25, Miocene, New Jersey to
S. Carolina.

THE CUPULADRIIDAE 181

Cupularia umbellata (Defrance) Smitt, 1873 : 14, pi. 3, figs. 75-80, Cape Fear River, N. Carolina,
7 fath. C. umbellata (Manz.) (sic) : Verrill, 1878 : 305. Canu & Bassler, 1918 : 118, pi. 53,
figs. 2-4, 1919 : 85, pi. i, figs. 5-7, pi. 2, figs. 17-21. 1923 : 68, pi. 2, figs. 15-19, text-fig.
joG, Lower Miocene, west Indies. 1928 : 64, pi. 7, figs. 1-3, Gulf of Mexico. 1929 : 142, pi. 15,
figs. 5-11, text-figs. 35A-K (as C. lowei), between California and Hawaii, 2723 fath. 1930 : n,
Galapagos and Hawaii. Sil6n, 1942 (part) : 15, text-figs. 10-12, localities 1-17.

Cupularia punctata Canu, 1904 : 10, pi. 2, fig. 23, Miocene, Patagonia.

Cupularia canariensis Busk : Robertson, 1908 : 314, pi. 24, figs. 90, 91, California.

Cupularia lowei Busk : Osburn, 1914 : 194, Florida, 12 and 22 fath.

Cupularia robertsoniae Canu & Bassler, 1923 : 82, pi. 34, figs. 5-7, Pleistocene, California.

Discoporella umbellata (Defrance) Hastings, 1930 : 718, pi. n, fig. 54, Gorgona, Galapagos,
Balboa, 15-30 fath. Mcguirt, 1941 : 65, pi. i, figs. 4, 7, 9-11, Miocene, Louisiana. Osburn,
1950 : 113, pi. n, figs. 7-10, California to Ecuador, 5-100 fath. Soule, 1959 : 34, Gulf of
California, 5-46 fath. Soule & Duff, 1957 : 99> Pleistocene, California. Marcus & Marcus,
1962 : 290, pi. i, fig. 4, pis. 2-5, Brazilian coast, 4-150 m.

MATERIAL. British Museum. Gorgona, 1929. 4. 26. 102, 103, 106 ; Balboa,
1929. 4. 26. 104 ; Galapagos, 1929. 4. 26. 105. " St. George " Coll. Galapagos,
1933. 12. 10. 15, Canu & Bassler Coll. Abrolhos Bank, off Brazil, 1899. 7. i. 1260,
1261 and 1265 (doma-type zoarium). Berbice, British Guiana, 25 fath. (21 zoaria, 4
of doma-type). ' Terra Nova " Coll. Stn. 42, off Rio de Janeiro.

Palaeontological Dept., D 33911, D 33912, D 33916, D 33910, Miocene, San
Domingo. D 41118, Bowden marl, Miocene, Jamaica. D 34392-409, Miocene,
Duplin, S. Carolina.

Cheetham Coll. Miocene, Oak Grove, Okaloosa Co., Florida.

Philadelphia Academy Coll., Wilmington, N. Carolina.

Zoarium large (maximum diameter observed 18 mm.) . Zooecia generally narrower
than in D. umbellata, lateral cryptocyst descending more steeply. Horizontal crypto-
cyst lamina not usually porous at centre (in Recent specimens) . Opesia large, curved
proximally or slightly sinuate, denticles absent. Opesiules large, 3-9 in number.
Operculum with delicate basal sclerite curved distally. Vibracular opesiae little
larger than zooecial opesiae, not denticulate proximally. Basal surface tuberculate
with grooves, or short furrows and pits.

DIMENSIONS. Lz 0-45-0-7 mm. ; Iz 0-22-0-35 mm. ; Lop 0-11-0-13 mm- ) 1°P
0-12-0-14 mm. ; Lo 0-07-0-08 mm. ; lo o-io-o-n mm., Lvo 0-15-0-18 mm. ; Ls
0-60-1-10 mm.

Conrad's description of L. depressa mentioned that " many of the larger pores "
(i.e. zooecia) were " filled with a minutely porous plate or diaphragm, solid in the
centre ". Specimens examined from the Miocene of Wilmington, N. Carolina, have
narrow zooecia, with 7 large opesiules and an opesia with a curved proximal edge. A
few central pores may be seen from the underside of the cryptocyst, but most of these
are occluded by secondary calcification. Natural Well, the type locality, and Wil-
mington, are near the Cape Fear River, from which Smitt described the first Recent
specimens from Pourtales's Collection. Canu's figure of C. punctata showed the
frontal surface of a worn specimen ; it had the characters of D. u. depressa, with 3-7
opesiules, and a rounded opesia.

The magnification of some of the figures of D, u. depressa (as D, umbellata} given in

i8a P. L. COOK

two of the works of Canu & Bassler is confusing. The same series of photographs
were used throughout ; but those of 1919 (pi. 2, fig. 17 (nat. size), figs. 18 (X25),
19, 20 and 21 (xig)) are smaller than those of 1923 (pi. 2, figs. 15, 16, 17, 18, 19)
which are described as having the same magnification.

Osburn (1950) confirmed that Verrill's record (1878) of D. umbellata from Fort
Macon, N. Carolina, was correct, as he had examined material from the same locality.

The zooecia of D. u. depressa are generally narrower than those of D. umbellata.
The figures of American material all agree in showing narrow zooecia with large,
rounded opesiae and opesiules, and Soule & Duff described the zooecia as " elongate ".
The opesia is always curved proximally, and may be distinctly sinuate ; denticles are
absent. The bars between the opesiules are narrow. The proximal sclerite of the
operculum is delicate and slightly curved in a distal direction. The vibracular
opesiae are never denticulate proximally, as in D. umbellata , and proportionately, they
are smaller ; the setae do not appear to attain the length of those of D. umbellata.

Most of the 7 doma-type colonies examined have large, rounded opesiae and narrow
zooecia (Iz 0-20-0-35 mm.), with 2-7 opesiules, and are thus similar to more normal
specimens of D. depressa. They differ from the doma-type colony of D. umbellata
principally in their narrower zooecia and larger opesiae.

14 CUPULADRIIDAE, GENERAL COMMENTS

The number of species of Cupuladriidae from the Pacific Ocean is thus two,
C. guineensis and D. u. depressa, which last is confined to the eastern part. Two
species occur in the Indian Ocean ; C. indica in the north and west, and D. umbellata
(peyroti form) in the south-west. Ten forms may be distinguished in the Atlantic
Ocean (Cook (in press)), only the subspecies D. u. depressa occurring in both the
Atlantic and Pacific.

The family is sharply defined, its principal characters being the distinctive asto-
geny, radial development and type of vibracula. Species in which any character
shows a major discrepancy should be assignable to other families. This is demon-
strated by examination of the characters of Cupularia capriensis Waters.

HINCKSINIDAE Canu & Bassler 1927
Hincksinidae Canu & Bassler, 1927 : 3, Bassler, 1953 : G 159, Lagaaij, ig63b : 171.

SETOSELLINA Calvet, 1906 : 157

15 Setosellina capriensis (Waters)
(PL i, figs. 5, 6)

not Setosellina roulei Calvet, 1907 : 395, pi. 26, figs. 5, 6, Cape Verde Is., 1900 m., Capo Blanco,

2330 m.
" Cupularia minima (Busk MS) " Waters, 1921 : 419, as Heliodoma implicata Calvet, specimens

from the Aegean Sea. (not H. implicata, see below).
Setosellina roulei Calvet : Waters, 1925 : 350, pi. 21, fig. 4, Capri, 50 fath. ; Gautier, 1962 : 69,

Tunisia, 150 m., Rhone Delta, 120 m.

THE CUPULADRIIDAE 183

Cupularia capriensis Waters, 1926 : 432, pi. 18, figs. 8, g, Capri.
Cupuladria capriensis (Waters) Marcus & Marcus, 1962 : 288.

LECTOPARATYPES, chosen here, Manchester Museum, Faraglione, Capri, 150 fath.,
Waters Coll.

OTHER MATERIAL. British Museum. Aegean Sea, 130 fath., 1899. 7. i. 1253 and
5122 ; 100 fath., 1899. 7. I. 1251, collected Spratt. Ras el-Amouch, 1899. 7. i.
4701 ; Mediterranean, 30 fath., 1899. 7. i. 1491, collected " Porcupine ". Busk
Coll.

Zoarium small, selenariiform, free, growing on a sand-grain. Zooecia with oval
opesia. Each zooecium with a distal interzooecial vibraculum, reaching the basal
wall of the zoarium. Opesia of ancestrular and central area closed by a calcined
lamina with a single central pore. Ancestrula with one distal and one proximal-
lateral vibraculum, and surrounded by 6 zooecia, budded spirally.

DIMENSIONS. Lz 0-45-0-60 mm., av. 0-54 mm. ; Iz 0-30-0-34 mm., av. 0-32 mm. ;
Lop 0-32-0-35 mm., av. 0-33 mm. ; lop 0-17-0-20 mm., av. 0-19 mm.

Waters (1921) described, as Heliodoma implicate Calvet (1906 : 157, 1907 : 396,
pi. 26, figs. 7, 10), some specimens from the Aegean Sea in the Busk Collection
(1899. 7. i. 1253). He particularly noted that the form of the zoarial spiral differed
from that figured by Calvet. The zooecia of the ancestrular area of H. implicata are
not occluded by a lamina, which is however, present in the specimens on slide 1899.
7. i. 1253. These zoaria are identical with those from Capri described by Waters
in 1926 as Cupularia capriensis, which has almost complete opesial closures in the
ancestrular area, a small central pore only remaining open.

Waters does not at any time appear to have considered the Aegean specimens
either as distinct from H. implicata or as identical with his new species, C. capriensis.
His quotation of Busk's MS name for them in 1921 cannot therefore be regarded as a
specific designation.

C. capriensis Waters is certainly referable to Setosellina Calvet. Calvet (1907 :
395) described Setosellina roulei with a salient calcareous lamina and figured (pi. 26,
fig. 6) zooecia which were partially closed, with a small central pore. The early
astogeny of specimens of 5. capriensis was described (as 5. roulei} by Waters (1925 :
350, pi. 21, fig. 4, Capri, 50 fath.). The budding is similar to that of Otionella and
Selenaria figured by Stach (i936b : 64, text-figs. 1-7). The same type of spiral
budding is found in Setosellina constricta (Borneo Bank, 1928. 3. 6. 75 and N. New
Guinea, 1928. 3. 6. 76) described by Harmer (1926 : 264), and 5. goesi (Silen) des-
cribed by Lagaaij (i963b : 172). The basal surface of 5. capriensis shows that the
vibracular chambers are inserted between, and extend to the same depth as the
zooecia. They are thus truly interzooecial, and completely unlike those found in the
Cupuladriidae.

Gautier (1962 : 69) commented on the probable identity of C. capriensis Waters
with Setosellina roulei Calvet. The dimensions of the abyssal specimens figured by
Calvet are significantly smaller thanthoseof 5. capriensis given above. They average :
Lz 0-36 mm. ; Iz 0-25 mm. ; Lop 0-23 mm. ; lop 0-13 mm. Measurements taken from

184 P. L. COOK

Waters's figure of 5. roulei from the Mediterranean (1925, pi. 21, fig. 4), combined
with those given by Gautier, give a range of Lz 0-45-0-50 mm. ; Iz 0-32-0-35 mm. ;
Lop 0-25-0-28 mm. ; lop 0-16-0-18 mm. These dimensions are far closer to those of
S. capriensis given above than to those of 5. roulei as originally described. The
geographical and bathymetrical distributions of the two forms are also distinct ;
S. roulei occurring off the Cape Verde Islands at 1900 m., and off Capo Blanco at
2330 m. The larger S. capriensis has been recorded from the Mediterranean and
Aegean only, at a greatest depth of 150 m. Until further material from both areas
can be examined, all the Mediterranean records are here considered to be distinct and
referable to 5. capriensis.

S. capriensis differs from the Cupuladriidae in its early astogeny, and related to
this, in the spiral development of the zoarium. The relationship of the vibracular
chamber to the zooecium, and the type of opesial closure of the central zooecia, are
also fundamentally different from those found in the Cupuladriidae.

16 ACKNOWLEDGMENTS

My thanks are due to Dr. A. Anderson (Naturhistoriska Riksmuseet, Stockholm),
Dr. R. S. Boardman (Smithsonian Institution, Washington) and Dr. D. E. Owen
(Manchester University Museum), for their generous assistance in lending many of
the specimens examined. The photographs were taken by Messrs J. V. Brown and
P. Green at the British Museum (Natural History), and at the Smithsonian Institution
(see PI. i for details). I am also indebted to Dr. A. Cheethan (Louisiana State
University), Dr. R. Lagaaij (Shell Exploratie en Produktie Laboratorium, The
Netherlands), and Herr Prof. Dr. E. Voigt (Geologisches Staatsinstitut, Hamburg), for
much helpful discussion and correspondence. Lastly, I wish to thank Dr. A. B.
Hastings and Dr. J. P. Harding (British Museum, Natural History) for their valuable
help and encouragement throughout the course of this work.

17 SUMMARY

The principal characters and mode of development of the Cupuladriidae are
described and discussed, the family is defined, and groups of species with similar
characters are listed. The effects of zoarial form of zooecial characters, and the
occurrence and possible causes of abnormal zoaria similar to those found in C. doma,
and those described as C. peyroti, are discussed. Descriptions are given of C.
pyriformis, C. indica and C. guineensis. The complex of forms previously assigned
to D. umbellata is examined in the light of character-consistency of fossil and Recent
records. Cupularia capriensis Waters is described and referred to Setosellina.

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i86 P. L. COOK

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THE CUPULADRIIDAE 187

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PLATE i
Cupuladria, Setosellina and Discoporella

FIG. i. Cupuladria guineensis (Busk). Philippines, United States National
Museum. Syntype of " C. granulosa ", 1963. 3. 14. 9. Photograph by
courtesy of the United States National Museum. x 14.5.

FIG. 2. C. guineensis. Philippines, United States National Museum. Cotype of
" C. dentifera " , 1963. 3. 14. 7. Photograph by courtesy of the United States
National Museum. xi6.

FIG. 3. C. pyriformis (Busk). Anguilla, Naturhistoriska Riksmuseet, FBD 533,
frontal surface. x 10.

FIG. 4. C. pyriformis. The same specimen, basal surface. x 10.

FIG. 5. Setosellina capriensis (Waters). Aegean, 1899. 7. i. I253pt., Busk Coll.,
frontal surface. x 1 7 . 4.

FIG. 6. S. capriensis. The same specimen, basal surface. XI7-4.

FIG. 7. Discoporella umbellata (Defrance). Peyroti-type zoarium, S. Africa,
1949. ii. 10. 94, Burrows Coll., basal surface, showing kenozooecia and vibracula
growing over the basal surface. X4-3.

Bull. 13. M. (N.H.) Zool. 13, 5

PLATE 1

PLATE a
Cupuladria indica and C. guineensis

FIG. i. Cupuladria indican. sp. Burma, 1899. 5. i. 267 pt., Hincks Coll., paratypc,
frontal surface. x 1 3 . 4.

FIG. 2. C. indica. The same specimen, basal surface. xio.5.

FIG. 3. C. guineensis (Busk). Sulu Archipelago, 1963. 3. 14. 2, U.S.N.M. Coll.
(from type-locality of " C. tuberosa "}, frontal surface. xg.

FIG. 4. C. guineensis. The same specimen, basal surface. xy.

Bull. B.M. (N.H.) Zoo/. 13, 5

PLATE 2

PLATE 3
The Discoporella umbellata-complex

FIG. i. Discoporella umbellata (Defrance). Ghana, Achimota Coll., II, jA.

Zooecia and vibracula. x 40.
FIG. 2. D. umbellata subsp. depressa (Conrad). Rio de Janeiro, Discovery Coll.,

SygA. Zooecia and vibracula. X55-
FIG. 3. D. umbellata. S.W. France, L. Miocene, 025003. Zooecia and vibracula.

X3i.
FIG. 4. D. umbellata subsp. depressa. Florida, Miocene. Zooecia and vibracula.

Xi8.5.
FIG. 5. D. umbellata. Peyroti-typezoa,num.S. Africa, 1949. n. 10. 94 pt., Burrows

Coll. Zooecia and vibracula. X43-

FIG. 6. D. uwibellata. Peyroti-type zoarium. The same specimen, basal surface,
showing kenozooecia and vibracula. X3y.

Bull. B.M. (N.H.) Zool. 13, 5

PLATE 3

PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING

POLYZOA FROM WEST AFRICA

THE CUPULADRIIDAE
(CHEILOSTOMATA, ANASCA)

PATRICIA L. COOK

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 6

LONDON: 1965

POLYZOA FROM WEST AFRICA

THE CUPULADRIIDAE
(CHEILOSTOMATA, ANASCA)

PATRICIA L. COOK

British Museum (Natural History)

Pp. 189-227 ; Plates 1-3 ; 4 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. 6

LONDON: 1965

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 13, No. 6 of the Zoological series.
The abbreviated titles of periodicals cited follow those of
the World List of Scientific Periodicals.

Trustees of the British Museum (Natural History) 1965

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued October, 1965 Price Eighteen Shillings

POLYZOA FROM WEST AFRICA

THE CUPULADRIIDAE
(CHEILOSTOMATA, ANASCA)

By PATRICIA L. COOK

CONTENTS

Page

INTRODUCTION AND METHODS .....

. lyi

CUPULADRIIDAE LAGAAIJ .....

. 192

GENERAL NOTES ON LUNULITIFORM COLONIES

• 193

KEY TO SPECIES .......

196

Cupuladria Canu & Bassler .....

. 197

C. canariensis (Busk) ......

. 197

C. biporosa Canu & Bassler .....

. 203

C. monotrema (Busk) ......

209

C. multispinata (Canu & Bassler) ....

210

C. owenii (Gray) .......

. 213

C. owenii subsp. disciformis n. subsp.

. 215

C. doma (d'Orbigny) ......

. 216

Discoporella d'Orbigny ......

219

D. reussiana (Manzoni) ......

219

D. ocellata n. sp. .......

220

D. umbellata (Def ranee) ......

221

ACKNOWLEDGMENTS ......

• 223

SUMMARY ........

. 223

REFERENCES. .......

224

i INTRODUCTION

THE Collections from which the specimens originate have been described by Cook
(1964 : 44) ; they comprise the " Calypso " Collections (Collection I from Senegal to
the Bay of Biafra, Collection II from the Cape Verde Islands) ; the Marche-Marchad
Collections (from Senegal) ; and the Achimota Collections (from Ghana). The
Collections are rich in lunulitiform colonies belonging to the family Cupuladriidae ;
8 species occur round the west African coast, and it is estimated that approximately
2,000 colonies have been available for examination.

The lunulitiform and selenariiform types of zoarium are found among several
unrelated genera of Polyzoa (see Harmer, 1926 and 1957, and Cook, 1965) ;
they are both associated with sandy or muddy substrates, upon which other Polyzoa
are usually unable to grow directly. Harmer (1957 : 649, 726, 885, 891 and 1009)
discussed the correlation between substrate and selenariiform habit. A large number
of the specimens in these Collections are known to be from a sandy or muddy bottom ;
for example, those from the Achimota Collection are nearly all from stations close to,
or included in, the " silty sand community " of Buchanan (1958 : 16, 26) and
Bassindale (1961 : 492). Stations where the bottom is known to be of this type are
marked with an asterisk in the lists of material examined. The lunulitiform zoarium

ZOOL. 13, 6 10

iga P. L. COOK

has been discussed by Waters (1921), Harmer (1926 : 261 and 1957) and Lagaaij
(1952 : 31, 43 ; 1953 ; and 1963). Briefly it may be described as free, discoidal or
subconical, with zooecia arranged in radial rows. In the Cupuladriidae each zooecium
is associated with a distal vibraculum, which has a long seta.

METHODS. Colonies were treated with eau de javelle and fragments mounted dry.
Others were decalcified in dilute acid, sometimes only partially, and stained to show
the relationship of muscles to the calcareous parts, etc. Dry specimens were treated
with trisodium phosphate solution to restore shape to shrunken chitinous parts
before preparations were made.

The length of the zooecia were measured from the distal edge of the aperture to the
distal edge of the next succeeding radial zooecial aperture, thus including the
vibraculum.

The dimensions quoted give the range of variation of 50 measurements (where
possible). Generally, measurements of the central area or of very young colonies
have not been included. It is hoped to make a survey of measurements useful for
specific determination, with a morphometric analysis of at least one population
complex, in the near future.

The measurements made are :

Length of zooecium Lz Length of vibracula seta Ls

Width of zooecium Iz Length of vicarious seta Lvs

Length of ancestrula Length of operculum Lo

(including vibraculum) La Width of operculum lo

Length of vibracular opesia Lvo Length of opesia Lop

Length of vicarious vibraculum Lv Width of opesia lop

Definition of terms and symbols. The registration numbers of specimens in the
British Museum (Zoology Department) are given thus : 1899. 7. i . . . ; (Palaeon-
tological Department) thus : D 6764 . . .

Horizontal cryptocyst lamina. A porous lamina, formed from fused cryptocystal
denticles, extending from the descending cryptocyst across the opesia, parallel to, and
below, the frontal membrane.

Vestibular arch. The distal, raised portion of the aperture. Usually accompanied
in the Cupuladriidae by a pair of distal cryptocystal denticles.

Vicarious vibraculum. A large vibracular individual, taking the place of a zooe-
cium, and itself having a distal vibraculum.

In the lists of material examined, the presence of vicarious vibracula is indicated
thus : (V) ; of Acrothoracid Cirripedes thus : (C), see p. 194 ; and of a sandy or
muddy substrate thus : *.

2 CUPULADRIIDAE Lagaaij
Cupuladriidae Lagaaij, 1952 : 31. Cook, 1965: 154.

The series of species described below links the membraniporan forms assigned to
Cupuladria with the microporan forms represented by Discoporella, and both genera
have therefore been placed in the family Cupuladriidae.

POLYZOA FROM WEST AFRICA 193

Diagnosis. Lunulitiform Anasca with vibracula alternating with zooecia in the
same radial series. Ancestrula surrounded by seven zooecia and a distal vibraculum.
Cryptocyst variously developed. Ovicells absent.

The systematic position of the two genera here included in the Cupuladriidae has
been discussed by Canu & Bassler (1923 : 75), Harmer (1926 : 266), Hastings (1930 :
714, 717), Lagaaij (1952 : 32, and 1953), Marcus & Marcus (1962), and Cook (1965 :
154). Bassler (1953) placed Cupuladria in the Membraniporidae (p. G 156) and
Discoporella in the Calpensiidae (p. G 171), but the small distinction between the
membraniporan and microporan forms had already been indicated by Canu & Bassler
(1923 : 75) who remarked " The union of spines is not a generic character. In fact
it may be accidental (Cupuladria denticulata] , partial (Cupuladria reussiana) , almost
complete (Cupuladria, umbellata] ". D. ocellata n. sp. links the irregular opesiular
indentations and fused cryptocystal denticles typical of D. reussiana with the
microporan D. umbellata (see p. 221).

3 GENERAL NOTES ON LUNULITIFORM COLONIES

a. Budding. The method of budding and development of lunulitiform colonies has
been discussed by Silen (1942 : 7-13), Lagaaij (1963) and Cook (1965 : 155), and the
regeneration of broken colonies by Dartevelle (1935 : 559-561). A large number of
the colonies in these collections are regenerated from broken fragments ; approxi-
mately 80% of D. umbellata in the " Calypso " Collection are of this type, as are many
specimens of the other species. It is interesting that the drawings of C. owenii by
Gray (1828, pi. 3, figs. i5a) and of D. umbellata by d'Orbigny (1853, pi. 717, figs. 3, 4)
are of regenerated colonies.

b. Sexual reproduction. Ovicells are absent ; large eggs (" about 0-5 mm. long "),
were seen in C. canariensis and C. doma (as C.johnsoni) by Waters (1921 : 404 and 414),
and smaller eggs, which may not have been fully developed, by Hastings (1930 : 726)
in D. umbellata depressa (as D. umbellata}. These eggs (Gorgona, 1929. 4. 26. 103
pt.), have an average diameter of 0-05 mm. (9% of Lz) ; those found in zooecia of
C. multispinata (Stn. 299, Cape Verde Is., Discovery Coll.), have an average diameter
of 0-30 mm. (45% of Lz). Eggs in C. owenii disciformis (" Calypso " Coll., C47A)
average 0-20 mm. (60% of Lz), and one egg in C. biporosa (" Calypso " Coll., C49G)
has a diameter of 0-20 mm. (50% of Lz). The date of breeding shows great variation
and is probably dependent upon several unknown factors. Marcus & Marcus (1962 :
297) found no evidence of germ cells in material from the Brazilian coast collected in
December, January, March, April, June and August. The " Calypso " specimen of

C. biporosa was breeding on 26th May and that of C. owenii disciformis on 26th
July, 1956. The Madeiran C. doma. described by Waters was received from Norman,
who collected in March and May (see Norman, 1909 : 275) . The fertile C. multispinata
from the Cape Verde Islands was collected on 4th September, 1927, and Hastings's

D. umbellata depressa from Gorgona, was breeding in July, 1924. Specimens of
D, umbellata collected in March, 1963, from Funchal, Madeira, were not breeding,

HJ4 1'. L. COOK

but one very young, recently settled, colony was found. This suggests that perhaps
the breeding season here was in early spring.

Generally, the present evidence is that the eggs of the membraniporan and denti-
culate species are larger than those of the microporan species. This may be con-
nected with the development of the cryptocyst and the size of the opesia, but until
observations are made upon living, breeding colonies, the importance of these differen-
ces cannot be assessed. The relatively large size of some of the eggs may indicate
that the larvae have a yolk and therefore a short free-living existence ; but Lagaaij
(1963 : 178) considered that the larval life may persist for some time. Settlement is
upon a sand grain, small stone, or Foraminiferan shell, and the ancestrula buds one
proximal, a pair of proximal-lateral zooecia, and a distal vibraculum. A pair of
lateral zooecia follows, and the ancestrular area is later completed by a pair of distal-
lateral zooecia (see Lagaaij, 1963, text-fig. 10 ; and Cook, 1965, text-fig. iB).

c. Mode of life. The mode of life of lunulitiform colonies was virtually unknown,
but living specimens have now been observed (Marcus & Marcus, 1962 ; and Cook,
1963). Their orientation is normally with the zooecial face upward. All the
evidence at present available indicates that colonies belonging to the Cupuladriidae,
at least, are not capable of free-swimming movement through the water, but are
maintained by their peripheral vibracular setae in a position just above the surface of
the substrate. The zooecia of the central area do not appear to have either a " hydro-
static " or " radicular " function (cf. Canu & Bassler, 1920 : 75, 1923 : 238, and 1929 :
144). The single instance of 9 colonies being taken at the surface was recorded by
Silen (1942 : 13, in the Atlantic, 1881, 27° 16' N., 23° 21' W.), who commented on the
lack of any similar occurrence. These specimens have been re-examined (see p. 207).
All have vibracular setae and intact frontal membranes ; the zooecia have polypides.
They were therefore presumably alive when collected, but I agree with Silen, who
later remarked (1947 : 10), " the free, swimming life of C. canariensis is not at present
to be accepted as proved fact ".

d. Associated Cirripede. West African Cupuladriidae are often hosts to individuals
of an Acrothoracid Cirripede, the presence of which is indicated on the basal side of
the colony by slightly thicker calcification, which in C. canariensis obscures the basal
kenozooecial pores. On the frontal side a slit, surrounded by a calcified border,
marks the opening through which the cirri of the Acrothoracid protrude for feeding
(see PI. 3, fig. 3). Members of this group of Cirripedes have been reported from
localities including Cadiz and South Africa (see Utinomi, 1950 : 5), inhabiting bar-
nacle plates and corals, and producing similar slit openings in these hosts. Most of
the slits are found near the central area of the colony. The majority of Acrothoracid
specimens have been found in large colonies of C. canariensis and D. iimbellata, but
they also occur in C. multispinata, C. doma and D. reussiana. Other conical zoaria
with slits evidently made by similar Acrothoracids are those of Selenaria maculata
Busk (Bass's Straits, 1854. u. 15. 52, incinerated specimen) and Stylopoma duboisii
(Audouin) (Holothuria Bank, 1892. i. 28. 43, see Cook, I965a, in press).

The majority of other Polyzoa of the silty sand community encrust the large
Foraminiferan, Jullienella foetida, which has an argillaceous test. Neither these nor

POLYZOA FROM WEST AFRICA 195

the associated large, erect branching Polyzoan colonies of Metrarabdotos unguiculatum
Canu & Bassler and Cleidochasma oranense (Waters), show any evidence of being
inhabited by the Acrothoracid Cirripede.

Without observation of living specimens it is difficult to establish whether the
association described above is one of symbiosis, commensalism or parasitism, but it
should be noted that many of the zooecia surrounding the slit and thus directly
overlying the cavity containing the Acrothoracid appear to be unaffected, and have
well-developed polypides. The successful functioning of the Cirripede suggests that
the Polyzoan zoarium has a degree of stability surprising in an unattached organism.
Inhabited specimens are marked in the lists of material examined thus, (C) .

e. Epifauna. The colony may provide a substrate for the settlement of larvae of
species of Polyzoa which would otherwise not be able to establish themselves success-
fully in a sandy or muddy habitat. Osburn (1914 : 190) described Beania cu-pularien-
sis and (1950 : 176) Membraniporella pulchra, growing on the basal side of colonies of
Cupuladria, and Soule (1959 : 22) reported Chaperiella condylata on C. canariensis and
D. umbellata. Smittipora levinseni (Canu & Bassler) has been found on C. canariensis
from the " Calypso " Coll. (C 726), and a specimen of Onychocdla angulosa (Reuss)
budded from a central ancestrula completely covers the frontal surface of a colony of
C. biporosa from the Canaries (1962. 10. 8. 7), the peripheral zooecia of which can be
seen beneath the growing edge of the Onychocella.

The colonies are frequently the substrate for other groups, notably small barnacles,
tube-worms, hydroids and sponges. The barnacles and worms are found on the basal
side of zoaria with a well domed cavity, where there is presumably sufficient room for
their cirri and branchiae to be protruded. Hydroid stolons are found on the frontal
surface, running between the zooecia, and sponges on the basal side. Apparently
these last may cause deformation of the zoaria, by the suppression of growth of one or
more radial rows of zooecia. C. multispinata (see p. 210) was referred to by Waters
(1921 : 413) under Busk's MS. name, Cupularia deformis, and specimens labelled "C.
deformis ' ' by Busk nearly all show fragments of sponges covering the area where the
zooecia have failed to develop. Deformed colonies of D. umbellata were described as
C. lowei by Waters, and his type-material and other specimens in the British Museum
Coll. have sponge colonies on the basal side. Zoaria of D. umbellata (" Calypso "
Coll. C56G and Marche-Marchad Coll. I 390) are also covered basally with a sponge
belonging to the Plocamiidae. Here the sponge appears to prevent the connection
of the intercalary rows of zooecia produced on either side of the abortive row ; the
sponge eventually lines the radial slit thus produced. This may, of course, be a
secondary effect, but the correlation of the occurrence of deformity and sponges is
significant (see Cook, 1965 : 158).

The geographical, bathymetrical and palaeontological distribution of the Cupula-
driidae is very wide. Recent specimens of C. canariensis have been found from the
S.W. Mediterranean to the Gulf of Mexico ; " D. umbellata " has been reported from a
depth of 2,723 fathoms (Canu & Bassler, 1929 : 144) ; and the majority of the species
have also been recorded from the Tertiary, D. reussiana and D. umbellata having
originally been described as fossils.

196 P. L. COOK

4 KEY TO THE SPECIES OF CUPUL AD RI I D AE DESCRIBED BELOW

Note on the identification of worn specimens. Lunulitiform colonies are frequently
found to be worn and identification under these conditions is extremely difficult.
Lagaaij (1952 : 34 and 1953 : 13) stressed the importance of the character of the basal
surface, which is less susceptible to wear, and this is emphasized in the following key.
Characters not found in fossil or worn specimens are placed in parentheses. All other
features have been found to be present in a few zooecia at least of moderately worn
Recent and fossil specimens examined.

1 Basal surface divided by radial and tangential boundaries into sectors ... 2
Basal surface not divided into sectors ........ 4

2 Basal sectors nearly always (95%) small, square, with 1-6 pores, vicarious vibracula
frequently present (Operculum longer than wide) . . . . . . ' . 3

- Basal sectors irregular, majority (75%) long with 6-20 pores. Vicarious vibracula

infrequent, zooecial vibracula of two kinds. (Operculum wider than long)

C. canariensis (p. 197)

3 Basal sectors with 1-3 pores per sector, one layer only of basal kenozooecia, vicarious

vibracula present throughout colony. (Operculum within a thickened area of the
frontal membrane) ....... C. monotrema (p. 209)

- Basal sectors with 1-6 pores per sector, basal surface frequently filled in by many

layers of kenozooecia, vicarious vibracula near central area. (Operculum longer
than wide) .......... C. biporosa (p. 203)

4 Colonies small (3-7 mm. diameter), steep-sided, solid basally, or with a small con-

cavity lined with spinous tubercles. Peripheral zooecia closed, several rows of en-
larged peripheral vibracula present. (Operculum with incomplete proximal
sclerite) .......... C. doma1 (p. 216)

- Colonies flatter, concave basally. Cryptocyst with denticles and spinules, or with

horizontal cryptocyst lamina formed of fused denticles ..... 5

5 Cryptocyst denticles fused to form horizontal cryptocyst lamina, with pores

(Discoporelta) 6
Cryptocystal denticles not fused to form horizontal cryptocyst lamina ... 8

6 Opesia small with a pair of closed opesiules ....... 7

- Opesia with a pair of opesiular indentations, trifoliate. Zoarium high. Basal

surface with large tubercles ....... D. reussiana (p. 219)

7 Opesia sinuate proximally, cryptocyst with a few large pores. Basal surface

tuberculate with radial threads. (Operculum without a proximal sclerite)

D. ocellata (p. 220)

8 Opesia straight proximally with a pair of small denticles, basal surface tuberculate

with pits and grooves. (Operculum with a proximal sclerite) D. umbellata (p. 221)

9 Zooecia small Lz 0-37-0-47 mm., sides of vestibular arch convergent. Cryptocyst

with large distal denticles. Basal surface with very small, regular tubercules, or
smooth and glassy. (Operculum longer than wide) C. owenii (p. 213)

- Sides of vestibular arch straighter, distal denticles not well developed. (Operculum

not longer than wide) ........... 9

10 Zooecia large, Lz 0-55-0-76 mm., denticles spinulose. Basal surface with large,
irregular tubercles and salient threads. (Operculum as long as wide)

C. multispinata (p. 210)

1 Small, doma-type zoaria may occur in C. biporosa, C. owenii and D. umbellata (see Cook, 1965 : 162).
C. biporosa may be distinguished here by the basal surface (fork i), C. owenii by the absence of closed
peripheral zooecia and of a proximal sclerite in the operculum (fork 4), and D. umbellata by the presence
of a complete horizontal cryptocyst lamina (fork 4).

POLYZOA FROM WEST AFRICA 197

Zoarium flat, zooecia smaller, Lz 0-37-0-53 mm., denticles small, simple, few. Basal
surface with small regular tubercles. (Operculum wider than long)

C. owenii subsp. disciformis (p. 215)

5 CUPULADRIA Canu & Bassler

Cuptiladria Canu & Bassler, 1919 : 77, 1920 : 103. Hastings, 1930 : 718. Lagaaij, 1952 : 32.
Cheetham & Sandberg, 1964 : 1020.

TYPE SPECIES. Cupularia canariensis Busk, Madeira, Recent.

Zoarium lunulitiform. Zooecia with cryptocyst variously developed, frequently
with denticles, which do not normally fuse. Vibraculum distal to each zooecium.
Vicarious vibracula and basal sectors present in some species.

Vicarious vibracula have been found to occur only in the C. canariensis group of
species (group A, Cook, 1965 : 167). They are marked in the lists of material
examined thus : (V).

6 Cupuladria canariensis (Busk)

(Pl. i, figs. lA, B, PL 3, fig. 4, Text-figs. la-f)

Cupularia canariensis Busk (part), i85ga : 66 (not pi. 23, figs. 7, 8 — C. biporosa see p. 203),

Madeira, Canaries.
Cupularia canariensis Busk : Manzoni, 1869 : 26, pi. 2, fig. 17, Pliocene, Italy. 1877 : 24, pi. 17,

figs. 5a, b, c, Miocene, Austria and Hungary.
IMembranipora canariensis (Busk) Smitt (part), 1888 : 79, text-fig. 326, 120 fath. and over,

Florida.
ICupularia canariensis Busk : Angelis (part), 1899 : xxxiii, pi. B, figs. 7, 8, Pliocene, Spain (not

fig. 6 = ?C. biporosa).
Cupularia canariensis Busk : Neviani 1891 : 130, Post Pliocene, Italy. 1895 : 101, Miocene,

Italy.

ICupularia canariensis Busk : Calvet (part), 1907 : 393, Canaries and Cap Blanc, 80-259 m.
Cupularia guineensis Busk : Norman (part), 1909 : 289, pi. 37, figs. 3 and 6, Madeira (not figs.

4, 5 = C. biporosa).

?CupTtlaria guineensis Busk : Osburn (part), 1914 : 195, Tortugas Islands, Florida, 10 fath.
^.Cupularia canariensis Busk : Faura & Canu (part), 1916 : 133, Miocene, Spain (not pi. 3, fig. 8).
Cupularia canariensis Busk : Canu, 1917 : 137, pi. 3, figs. 4-6, Burdigalien, L. Miocene, France.
Cupularia canariensis Busk : Waters (part), 1921 : 410 pi. 30, figs, n, 12 Liberia, (not pi. 29,

fig. 15 = C. biporosa).
Cupuladria canariensis (Busk) : Canu & Bassler (part), 1928 : 16, pi. i, figs. 7-9, Recent, Gulf of

Mexico (not text-fig. 2 = C. biporosa).

Cupuladria canariensis (Busk) : Dartevelle, 1935 : 560, pi. 19, figs, i, 2, Ras el-Amouch, Algeria.
Cupuladria canariensis (Busk) : Sil6n (part), 1942 : 13, Pliocene, Italy ; Recent, Azores, West

Indies (not text-fig. 8 = C. biporosa ; not text-fig. 9, pi. 4, figs, 15, 16 = C. pyriformis).
Ciipuladria canariensis (Busk) : Lagaaij, 1952 : 33, pi. 2, figs. la, b, Pliocene, Netherlands.

T953 : T5. pl- I. fig- T» Miocene, Pliocene, Netherlands.
Cupuladria canariensis (Busk) : Buge, 1957 : I39> pl- 9> fig- 5- Miocene, S.W. France (not pl. 10,

fig- 3)-

Cupuladria canariensis (Busk) : Gautier, 1962 : 53, Algeria, 100-300 m.
Cupuladria canariensis (Busk) : Lagaaij, (part), 1963, pl. 25, figs, i, 3-5, pl. 26, figs. 2, ?3.
Cupuladria canariensis (Busk) Annoscia, 1963 : 225, pl. 9, fig. i, pl. 10, fig. i, pl. n, figs, ra, ib,

pl. 12, figs. la, ib, Quaternary, Italy.
Cupuladria canariensis (Busk) Cheetham & Sandberg, 1964 : 1021, text-figs, n, 13, Quaternary,

Louisiana,

ig8 P. L. COOK

MATERIAL. LECTOTYPE (chosen here), the top left-hand specimen on the slide
B.M. 1899. 7. i. 4697 (A), Busk Coll., M' Andrew, Canaries. Lectoparatypes, the top
right-hand specimen on the same slide and 1962. i. 24. 3, Busk Coll., M'Andrew,
Canaries.

"Calypso " Coll. I. Stn. 7*, 9° 40' N., 13° 53' 5" W., 17^.56, 18 m., C4F. Stn.
17*, 5° N., 5° 28' 30" W., 2i.v.56, 27 m., C$6] (C). Stn. 29*, 4° 3' N., 6° 12' E.,
26.V.56, 32 m., C49B (V). Stn. 45*, o° 25' N., 9° o' E,, 8.vi.56, 73 m., C48B (V).

Coll. II. Stn. 24, 15° 16' 34" N., 23° 47' 44" W., i8.xi.59, 55-6° m-> C86A.
Stn. 26, He Sao Tiago, 15° 16' 30" N., 23° 47' 31" W., i8.xi.59, 50-65 m., C65G.
Stn. 73, C72B (V).

Marche-Marchard Coll. I. Konakrey, Guinee Ise., iF, 2E. Flor de la bouteille,
Guinee Ise., 21.1.53, 8 m., 36 (V). Baie de Goree, 1954, 46-48 m., 6B. Sud de
Goree, 27.x. 53, 38-42 m., 76 (CV) ; and 24.11.53, 40-41 m., nH (CV). S.W.
Madeleines, i5.ix.53, 48 m., 2oB ; 45-46 m., 26G ; and 21.1.54, 46-48 m., 46E
(V). Sud de presque File du Cap Vert, 18.11.54, 95 m-> 29^ (CV), 33C (V). Baie de
Seminoles, Goree, 8.xii.53, 38 m., 3gC (V).

Coll. II. Baie de Goree, g.ix.55, 190-220 m., 3A (V) ; and 210-220 m., 33A (V).
S.W. large du Cap Vert, 9.ix.55, 100 m., uA (V), I4E (V). Banque de Fagaque
(Joal), J-5-V.53, 5 m., I5A. S.W. Madeleines, 9.1.54, 47-5 m., 3iD ; and i5.ix.53,
48 m., 4oA (V). Devant le Cap Manuel, Oct., 1952, 35 m., 37A (CV).

Coll. III. Sud de presque File du Cap Vert, i8.ii.54, 4^-5° m-> IC (V). Sud
de Goree, I3.xi.53, 33-35 m., 96 ; and 34-37 m., 176. Either as above, 34-37 m.,
or S.W. Madeleines, I5.ix.53, 48 m., i6E. Dragage i " Gerard Freca " i8.ii.54,
23F, dragage 4, 97 m., 28G ; dragage, 5, 156. 27.xi.53, 33-34 m., 29C. No infor-
mation 24G.

Achimota Coll. I, Stn. 69, Dredge haul No. 5, 22.1.51, 22 m., 90. III. C.

British Museum Coll. Madeira, 1912. 12. 21. 1002 (V), 1911. 10. i. 647 (V) and
1962. i. 28. 2 (V), Norman Coll. Bay of Funchal, Madeira, 1962. i. 28. i (V),
Norman Coll ; 50 m., 1963. 2. 28. 10 Cook Coll., mud*. Tangier Bay, 35 fath.,
1899. 7. i. 1225-1229, Busk Coll. Ras el-Amouch, Algeria, 45 fath. and upward,
" Porcupine ", 1899. 7. i. 84, 41 fath. 1226 ; 45 fath., 1222, 1264 and 1228 ; 4693 and
4694, Busk Coll. Mediterranean, 45 fath., 1899. 7. i. 1223 and 1230, Busk Coll.
Cape Rosa, Algeria, 95 fath., 1899. 7. i. 1227, Busk Coll. Cape Sagraas, Portugal,
1899. 7. i. 1221, Busk Coll. West Indies, " Blake ", 1879,1911. 10. I. 1722, Norman
Coll. Barbados, 73 fath., 9.iii.79, and 23° 13' N., 89° 10' W., 84 fath., " Blake "
1911. 10. i. 644, Norman Coll. St. James Coast, Barbados, 1962. i. 26. 2, Saunders
Coll. Gulf of Mexico, 28° 58' N., 89° 9' W., mud lump SP 5, 1961. n. 2. 4gA,
Cheetham Coll.

" Discovery " Coll., Stn. 279, off Cape Lopez, io.vii.27, 58-67 m., mud and fine
sand*, (CV).

Waters Coll. Manchester Museum. Four slides from Oran, i slide from " Post
Pliocene ", Pisa.

Naturhistoriska Riksmuseet, Stockholm Coll. (numbers in Silen, 1942 : 13-14 in
parentheses). St. Agata, Piemonte, Italy, Pliocene, FED 737 (No. 2). Azores, off

POLYZOA FROM WEST AFRICA 199

Punta Delgada, FBS, 964 (No. 6). West Indies, 52 miles off Florida, 40 m., FBD
548, i specimen only (No. 16).

Specimens from Dr. R. Lagaaij. Beeringen, Netherlands, Miocene. Karsy,
Poland, Miocene. Antwerp, Belgium, Pliocene. Gulf of Mexico " Atlantis " Stn.
161, 22 fath., Recent.

Specimens from Prof. Voigt. Rostej Banat, M. Miocene. Hamburg, Miocene.
Pinneberg, nr. Hamburg. U. Miocene.

Zoarium frequently large (maximum diameter measured, 23 mm.). Cryptocyst
narrow, descending steeply, finely tuberculate, not thickened in central zooecia.
Operculum slightly wider than long with no proximal sclerite. Zooecial vibracula of
two kinds, one with a short hooked seta. Vicarious vibracula sometimes present
surrounding ancestrula, and in the central area, with a short, stout hooked seta.
Basal surface divided into irregular rectangular sectors, the majority long, with from
4-20 pores (usually 6-12). Tentacles 17-19 (see p. 203).

DIMENSIONS. Lz 0-50-0-80 mm., Iz 0-30-0-47 mm., La 0-45-0-60 mm., Lvo
0-12-0-19 mm., Lv 0-32-0-45 mm., Ls 0-70-1-50 mm., Lvs 0-17-0-30 mm., Lo
0-10-0-13 mm., lo 0-12-0-15 mm., Lop 0-35-0-47 mm., lop 0-21-0-32 mm.

Lagaaij (1952 : 33) first chose Busk's slide (B.M. 1899. 7. i. 4697) as lectotype of
C. canariensis, but did not indicate upon which of the 6 specimens present his
description was based. The two specimens chosen above as lectotype and lecto-
paratype are definitely C. canariensis as described by Lagaaij, and as generally
understood by European authors ; although only part of Busk's description and
neither of his magnified figures (iSsga, pi. 23, figs. 7, 8) refer to this species. I have
here followed the Recommendation 74A of the International Code of Zoological
Nomenclature (1961 : 79), with regard to the " agreement with previous restriction "
of the species.

Smitt2 (1888 : 79, text-fig. 326) recorded Membranipora canariensis from 120
fathoms and over, from Florida. The figured colony is 18 mm. in diameter, and as
C. biporosa does not appear to attain this size, is perhaps C. canariensis. Specimens
sent to Norman by Smitt, from the " Blake " Collection from the West Indies, consist
of a mixture of C. canariensis and C. biporosa, and presumably Smitt's record refers
to both species, as perhaps does Osburn's from the Tortugas Islands, where some of the
specimens were very large (0-75 in.). Calvet (1907 : 393) included " C. guineensis
Kirkpatrick "3 (sic) in his synonymy ; it may be presumed that part of his material,
at least, was true C. canariensis.

Waters (1921 : 399) received material from Norman, whose figures (1909, pi. 37,
figs. 3 and 6) are of C. canariensis. Norman's specimens from Madeira in the British
Museum consist of a mixture of C. canariensis and C. biporosa.

The material described by Dartevelle is C. canariensis (Ras el-Amouch, 1899. 7.
i. 84, Busk Coll.), and shows the method of budding of zooecia from a broken frag-
ment particularly well.

2 For Smitt's authorship see Agassiz, 1888 : xxi.

3 Kirkpatrick (1890 : 612) only listed C. guineensis from the Torres Straits, his record is certainly not
C, canariensis (see Hastings, 1930 : 714).

200 P. L. COOK

Much of the material listed by Silen (1942 : 13-14) has been re-examined, namely
specimens from localities 2, 6, 7, 8, 9, 10, 15, 16 and 18. Of these, specimens 2 and 6,
and one of the zoaria from 16, are C. canariensis. The remaining specimens belong to
three other species, C. biporosa and C. owenii (see pp. 203 and 213 below), and C.
Pyriformis (Busk). Silen expressed doubt as to the identity of specimen 18, from
Anguilla, West Indies ; it is certainly referable to C. pyriformis (see Cook 1965 : 168).

The figures given by Canu & Bassler (1928) of specimens from the Gulf of Mexico
are all of C. canariensis with the exception of text-fig. 2, of an operculum, which shows
the elongated appearance typical of C. biporosa. Presumably their Recent material
consisted of both species.

The figures of C. canariensis given by Manzoni (1869, Pliocene, Italy), Manzoni
(1877, Miocene, Austria and Hungary) and Canu (1917, Lower Miocene, France) all
show steeply descending cryptocysts and long basal sectors. Neviani (1895)
recorded large colonies (2 cm. diameter) from the Italian Miocene, and (1891) had
mentioned the elongated basal sectors with numerous pores of his post-Pliocene
specimens. Both the specimens from the European Pliocene and Miocene lent by
Dr. Lagaaij, and his description and figures (1952 and 1953) have the same consistent
characters and thus contrast significantly with the figures of American Miocene
specimens given by Canu & Bassler (see p. 206), all of which are attributable to

C. biporosa. Buge (1957 : 177) stated that the figure of the basal side (as " face
superieure non zoeciale ") of D. umbellata given by D'Orbigny (1853, pi. 717, fig. 4)
was in reality of C. canariensis, and showed " secteurs radiaux avec pores ".
D'Orbigny's figure shows radial grooves typical of D. umbellata, and irregular
tubercles, with no sign of tangential sector boundaries or biserial pores. It is,
moreover, in outline, a mirror image of figure 3, which shows the frontal surface of

D. umbellata, and the two figures were obviously drawn from the same colony with
great attention to detail, as the budding pattern of the regenerated fragment from
which the zoarium originates (see p. 193), may be seen to correspond on the frontal
side, with the shape of the fragment visible on the basal side. D'Orbigny's explana-
tion of pi. 717 (p. 473), however, gives the Plate number as " 747 " (which contains
drawings of Cyclostomes only), and the descriptions of figures 2 and 3 have been
interchanged. Buge found that d'Orbigny's material of D. umbellata contained one
zoarium of C. canariensis. The frequent association of this pair of species has been
noted, (see p. 209), and may have occurred in Buge's material from the Redonian, as
his figure (pi. 10, fig. 3) of the frontal surface of his " C. canariensis " shows a well-
developed vestibular arch and wide zooecia, and is perhaps a worn fragment of
D. umbellata or C. haidingeri ; it has not the character of C. canariensis, but the basal
surface of a specimen figured on pi. 9, fig. 5, has irregular sectors with 4-8 pores, and
is similar to other Miocene specimens of C. canariensis.

The figure given by Faura & Canu (pi. 3, fig. 8) shows evidence of a worn denticulate
cryptocyst in some zooecia and is not of a specimen of C. canariensis. Their text-
figures 4a, b, are reproductions of those of Pliocene specimens given by Angelis
(1899), and do not represent their own material. Canu & Lecointre (1925, pi. 3)
figured C, canariensis from the Burdigalian of the Gironde, but the photographs have

POLYZUA FROM WEST AFRICA 201

been retouched (in fig. n sector boundaries have been drawn in through basal pores),
and little of the character of the specimens can be deduced.

The measurements given by Gautier (1962 : 53) show that the zooecia of his speci-
mens had wide opesiae. The zoaria were large (12-5 mm. in diameter), and were thus
very probably of C. canariensis.

Lagaaij's extremely full and detailed study of C. canariensis (1963), includes a
bibliography which combines references to both C. canariensis and to C. biporosa, as
limited here.

Zoaria from west Africa are generally larger than those from the Gulf of Mexico,
and of a flattened cone shape. In all the specimens examined the sides of the opesia
are curved and the lateral cryptocyst descends steeply, and is so little salient that in
cleaned specimens the pores in the lateral walls can be seen from a frontal view.

The ancestrula is frequently slightly larger than the zooecia of the primary circle
budded from it (see Canu & Bassler, 1928, pi. i, fig. 7), and the zooecia of the ances-
trular area never show any thickening of the cryptocyst as in C. biporosa and C,
monotrema. The zooecial vibracula of Recent specimens of C. canariensis are of two
distinct kinds. The majority are similar to those found in all other species of the
family ; they have auriform opesiae, rounded proximally, and long vibracula setae.
A proportion of between I : 10-1 : 30 individuals are small, have short, hooked setae,
and more symmetrical opesiae, with prominent narrow rostra pointing proximally
(see pi. 3, fig. 4). This kind of zooecial vibraculum has been found to occur in the
lectotype, the majority of west African colonies, and in a few zoaria from the Gulf of
Mexico. The occurrence thus differs from that of the vicarious vibracula with hooked
setae (see below) . No other species of Cupuladriidae has been seen in which two such
types of vibracula occur. Canu & Bassler (1929 : text-fig. 136) figured a second
type of seta in C. grandis, but did not describe it. Their figure is somewhat similar
to that of a regenerating seta in D. umbellata (subsp. depressa), given by Marcus &
Marcus (1962, pi. 2, fig. 7). The opesiae of regenerated vibracula are not of the
narrow, symmetrical type described above.

Vicarious vibracula are commonest in the central area of zoaria in which they occur,
but may also be present among zooecia budded from a regenerating edge. They
completely replace a zooecium and each has an ordinary small, distal vibraculum,
like the zooecia. The opesia of the vicarious vibraculum of C. canariensis is quite
unlike that of C. biporosa, which is auriform (see below). In C. canariensis a long,
acute, raised, beaked rostrum protrudes proximally beyond the short gymnocyst.
It supports a wide, short, strongly hooked seta, which is slung between the asym-
metrical condyles. The basal sclerite differs slightly from that of the normal vibra-
cular seta, but the individual is still basically a vibraculum (see Hastings, 1963 : 180).

The basal sectors of C. canariensis are unlike those of C. biporosa, being irregular in
length and rarely square in outline. The majority are elongated rectangles, with
6-12 pores, and occasionally more (see Text-fig. 21). Interspersed with these sectors
are short wide ones with 3-6 pores, these occur at the origin of an intercalary row of
zooecia. In basal view colonies of C. canariensis do not show the regular concentric
series of sectors which is so striking in C. biporosa. At the growing edge, the minute

202 P. L. COOK

pores in the basal wall can be seen in the developing zooecia to be without any sur-
rounding kenozooecial chamber. Further toward the centre of the colony, a cresent
of calcification with the concavity pointing distally grows round the pore ; when the
arms of the cresent meet, this forms the first kenozooecial chamber. In young
colonies the tangential divisions between the radial sector boundaries are not developed
until at least 4 radial series of zooecia have been budded. Thus even young colonies

FIG. i. Cupuladria canariensis and C. biporosa. a-f, C. canariensis. a. Vicarious
vibraculum and normal distal vibraculum, treated with eau de javelle. Marche-Marchad
Coll., I, 39 C. b. Hooked seta of vicarious vibraculum. c. Seta of normal vibraculum,
central part omitted, d. Operculum. b-e, " Calypso " Coll., I, C 56 J. f. Basal surface,
treated with eau de javelle, showing elongated sector with porous kenozooecial chambers.
Marche-Marchad Coll., I, 39 C. g-h, C. biporosa. g. Vicarious vibraculum and normal
distal vibraculum, treated with eau de javelle. 1962. i. 26. i. h. Scimitar-shaped seta
of vicarious vibraculum, in lateral view. i. Operculum. j. Basal surface, treated with
eau de javelle, showing small, square sectors with porous kenozooecial chambers. Marche-
Marchad Coll., Ill, 24 G.

POLYZUA FROM WEST AFRICA 203

may be distinguished from those of C. biporosa where the tangential sector boun-
daries are seen as soon as the zooecia grow out beyond the original substrate.

Waters (1921 : 411) gave the number of tentacles as 14, but this is the number
found in sections of C. biporosa (1929. 4. 26. 85 pt.). Waters's material included
both C. canariensis and C. biporosa, and the number of tentacles in C. canariensis
sections (MM II, 3A) is 17-19. C. doma (see Waters, 1921 : 411, and Cook, 1963 :
409) and the D. timbellata-complex (see Marcus & Marcus, 1962 : 295, and Cook,
1963 : 409) have 13-16 tentacles, and the same number has been seen in sections of
C. owenii disciformis (" Calypso " Coll. C47A).

Thus C. canariensis differs from C. biporosa chiefly in the character of the basal
surface, the cryptocyst, the number of tentacles, in the possession of two kinds of
zooecial vibracula, and the form of the vicarious vibracula. The distribution of the
two species also appears to have been completely distinct in the Miocene (see below) .
C. canariensis does not occur south of Barbados in Recent collections, all Brazilian
records being of C. biporosa or C. monotrema.

7 Cupuladria biporosa Canu & Bassler
(PL i, figs. 2A, B, 3A, B, 4A, B, 5, 6A, B, Text-figs, ig-j)

Cupularia canariensis Busk (part) i85ga : 66, pi. 23, figs. 7, 8, Madeira and Canaries.

Cupularia canariensis Busk : Busk, 1859 : 87, pi. 13, figs. 2 a-e, Coralline Crag, Pliocene, Britain.

Membranipora canariensis (Busk) Smitt, 1873 : 10, pi. 2, figs. 69-71, 10-14 fath. Florida. (Ppart),

1888 : 79, 1 20 fath. and over, Florida.

ICiipularia canariensis Busk : Angelis (part), 1899 : 33, pi. B, fig. 6 only, Pliocene, Spain.
Cupularia canariensis Busk : Canu, 1908 : 275, pi. 5, figs. 8, 9, 10, Pampean, Pliocene, Argentina.
Cupularia giiineensis Busk : Norman (part), 1909 : 289, pi. 37, figs. 4, 5, Madeira (not figs. 3, 6

= C. canariensis).

ICupularia guineensis Busk : (part) Osburn, 1914 : 195, Tortugas Islands, Florida, 10 fath.
Cupularia canariensis Busk : Canu & Bassler, 1918 : 119, pi. 53, figs. 5-7, Miocene, Costa Rica

and Jamaica.
Cupuladria canariensis (Busk) Canu & Bassler, 1919 : 78, pi. i, figs. 8-10, Lower Miocene, Costa

Rica, 1920 : 103, text-fig. 240. 1923 : 28, pi. i, figs. 7-9, Lower Miocene, Florida, Jamaica,

Santo Domingo, Costa Rica ; Miocene and Pliocene, Florida.
Cupularia canariensis Busk : Waters (part), 1921 : 410, pi. 29, fig. 5, Madeira.
Cupuladria biporosa Canu & Bassler, 1923 : 29, pi. 47, figs. 1-2, Miocene, Santo Domingo.
Cupuladria canariensis (Busk) : Canu & Bassler, 1928 : 16, text-fig. 2, Pliocene Panama ; and

Recent, Gulf of Mexico.
Cupuladria canariensis (Busk) : Hastings, 1930 : 714, pi. 8, figs. 38, 40, Gorgona, Colombia,

15-30 fath.
Cupuladria canariensis Busk (sic) McGuirt, 1941 : 46, pi. i, figs. 1-3, 5-6, 8, Miocene & Pliocene,

Louisiana.

Cupuladria canariensis (Busk) : Silen (part), 1942 : 13, text-fig. 8, West Indies.
Cupuladria canariensis (Busk) : Osburn, 1950 : 33, pi. 3, figs. 2, 3, Lower California to the

Galapagos Islands, shallow water to 40 fath.

Cupuladria canariensis (Busk) : Soule, 1959 : 8, California, 7-40 fath.

Cupuladna canariensis (Busk) : Galopim de Carvalho, 1961 : 97, pi. i, figs. 1-3, Pliocene, Portugal.
Cupuladria canariensis Marcus & Marcus, 1962 : 285, pi. i, figs. 1-3, off Sao Paulo, 150 m., near

Cabo Frio, 3 m., off mouths of the R. Amazon, 70 1^-71-5 m., dead.
Cupuladria canariensis (Busk) : Lagaaij, part, 1963, pi. 26, figs. 4, 5.
Cupuladria sp. Cheetham & Sandberg, 1964 : 1021, Quaternary, Louisiana.

204 !'• 1- COOK

MATERIAL. Holotype U.S.N.M. 68425. (Photographs, B.M.N.H. 1963. i. 2. i, 2)
Miocene, Bowden Marl, Santo Domingo.

" Calypso " Coll. I. Stn. 7*, 9° 40' N., 13° 53' 5" W., 17^.56, 18 m., C4l(V).
Stn. 29*, 4° 3' N., 6° 12' E., 26^.56, 32 m., C49G (V).

Coll. II. Stn. 26, 15° 16' 30" N., 23° 47' 31" W., i8.ii.59, 50-65 m., C65R.
Stn. 75, 16° 04' 20" N., 22° 58' 10" W., 45 m., CiogA.

Marche-Marchad Coll. I. Flor de la bouteille, Guinee Ise, 21.1.53, 8 m., 3E (V).
Sud de Goree, 27.x. 53, 38-42 m., jD. Sud de presque Tile de Cap Vert, 95 m.,
33Q. S.W. Madeleines, 21.1.54, 46-48 m., 46G (V).

Coll. II. S.W. Madeleines, 9.1.54, 77-5 m., 3iR.

Coll. III. Dragage 5, "Gerard Freca ", iS.ii. 54, 15! ; Dragage 4, 97-98 m.,
28K (V). 33-34 m., 27.11.53, 29! (V). No information, 24R (V).

British Museum Coll. Canaries, M'Andrew, 1899. 7. i. 46976, Busk Coll.
Canaries, M'Andrew, 1899. 7. i. 4702, 1962. i. 24. 4 and 5, Busk Coll. Madeira,
1879. 5- 28. 6. Bay of Funchal, Madeira, 1962. i. 24. 6, 1963. i. 16. 73, Madeira,
1962. i. 24. 7, and 9 (V), Norman Coll ; Funchal, 50 m., 1963. 2. 28. 6, Cook Coll.,
mud.* Ras el-Amouch, Algiers " Porcupine ", 4 fath. and upwards, 1962. i. 24. i.
Tangier Bay " Porcupine ", 1911. 10. i. 646, Norman Coll. Barbados, 73 fath.,
9.iii.79, and 23° 13' N., 89° 10' W., 84 fath., " Blake ", 1962. i. 24. 8, Norman Coll.
(V) West Indies, 1962. i. 24. 2, Norman Coll (V). Gulf of Mexico, 91° W., 28° 6'
25" N., 37 fath., 1959. 8. 20. 3, Lagaaij Coll. (V). 89° 9' W., 28° 58' N., 1961.11.2.
496, Cheetham Coll. Gorgona, Colombia, 1929. 4. 26. 85. 86, 263 (V), 87 (V).
St. James coast, Barbados, 1962. i. 26. i,SaundersColl.(V). Miocene, Santo Domingo,
1899. 7. i. 1259, Busk Coll. (V).

B.M. Palaeontological Dept. Coll. Coralline Crag, Pliocene, Britain, D 39968,
39969, D 6767, 6762, 6764, B 1624. Astigiana, Pliocene, Italy, 1848. 6. 12. 3-7.

U.S. National Museum Coll. Holotype, see above (V). 80747, Miocene, Bowden
Marl, Jamaica (V).

Dr. A. Cheetham Coll. Forest Hill Clay, Alabama, L. Miocene. Oak Grove,
Yellow River, Okaloosa County, Florida, L. Miocene. Cercado de Mao, Santo
Domgino, L. Miocene (V). Tamboo Trace, Shell Bed, nr. Talparo, Trinidad, L.
Miocene (V). Manzanilla, Manzanilla coast, Trinidad, Miocene (V).

Dr. R. Lagaaij Coll. Miocene, Limestone Creek, Missouri. Miocene, Cubagua,
Venezuela. Miocene, Bowden Marl (V), Jamaica. Pleistocene, Gulf of Mexico
" Neptune " i. Gulf of Mexico, " Atlantis " Stn. 163, 20 fath. (V).

Naturhistoriska Riksmuseet, Stockholm Coll. (Nos. in Silen, 1942 : 13-14 in
parentheses). Atlantic, 27° 16' N., 23° 21' W. at the surface, FBS 321 (No. 7).
(One of the 9 specimens is C. owenii qv) West Indies, S.W. off Tortugas 40 m.,
FED 239 (No 8) (V). As above, East Key, 20 m., FED 258 (No. 9) (V). As above,
nr. Rebecca shoal, 15 m., FED 281 (No. 10) (V). Yucatan, 35 miles, N.N.E. off C.
Catocha, 40 m., FED 696 (No. 15) (V). 52 miles off Florida, 40 m., FED 548 (No.
16) (V) (One colony of C. canariensis present).

C. biporosa is not present in the Achimota Collection.

Zoarium generally smaller than that of C. canariensis, not exceeding 16 mm. in

POLYZOA FROM WEST AFRICA 205

diameter, zooecia also smaller. Operculum frequently longer than wide with no basal
sclerite. Cryptocyst descending gently, sides of opesia straight. Vicarious vibracula
frequent, usually present in ancestrular area, occasionally throughout zoarium ; seta
straight, elongated, broad, slightly hooked, scimitar shaped. Basal surface divided
into concentric series of small, nearly square sectors with 1-6 pores, usually 4.
Basal concavity flattened and filled by many layers of kenozooecial chambers.
Tentacles 14-15 (see above).

DIMENSIONS. Lz 0-40-0-65 mm., Iz 0-25-0-35 mm., La 0-40-0-47 mm., Lvo 0-07-
0-13 mm., Lv 0-45-0-55 mm., Ls 0-40-1-40 mm., Lvs 0-25-0-30 mm., Lo o-io-o-n
mm., lo 0-09-0-10 mm., Lop 0-25-0-32 mm., lop 0-15-0-19 mm.

Dr. R. S. Boardman (in litt. 13.11.1962) has found that the only specimen referred
to C. biporosa in the U.S. National Museum Collection, is the holotype-fragment of
which part was figured by Canu & Bassler (1923, pi. 47, figs, i, 2). Enlarged photo-
graphs of the entire fragment show that one vicarious vibraculum is present, and that
the opesiae of the zooecia in the published figure (fig. i) have been enlarged by re-
touching of the photograph. This was presumably done to erase the image of
detritus lodged in the opesiae, but has unfortunately given the impression that the
cryptocyst descends more steeply than it in fact does.

Specimens of complete young zoaria labelled " C. canariensis ", from the same
horizon in Jamaica as the holotype (U.S.N.M. 80747) are strikingly similar in all
characters to young Recent colonies from the West Indies (see pi. i, figs. 4, 6).

Busk's description (i85ga : 66) of the occurrence of 2-4 pores in the sectors on the
basal side of his specimens, and his figures (pi. 23, figs. 7, 8) showing the small square
basal sectors (fig. 7), and well-developed cryptocyst (fig. 8), are certainly of C.
biporosa. The major part of the material from the Canaries collected by M' Andrew
in the Busk Collection (see Busk, i85ga : 67) is also of C. biporosa, but specimens of
C. canariensis are mixed with it, and it is probably these which Busk described as
0-5 in. in width, as none of his specimens of C. biporosa reach this size. The speci-
mens on slide 1899. 7. i. 4697 A -f B, of C. canariensis and C. biporosa, are mounted
on glass, and the distinctive characters of both surfaces of the colonies of the two
species can be seen clearly.

Busk's figures of specimens from the British Crag (1859 : 87, pi. 13, figs. 2a-e),
show typical C. biporosa characters, namely the straight sides of the opesiae and the
gently descending cryptocyst. The figure of the basal side shows some irregular
sectors, but specimen D 6764 has square sectors, with 4 pores, and many layers of
kenozooecial chambers, as well as the frontal character of C. biporosa. Busk's
specimens are particularly interesting as the. only other records of C. biporosa from
the Pliocene of Europe are from Spain, Portugal and Italy. Pliocene specimens
from the Netherlands are all referable to C. canariensis.

Smitt (1873 : 10) described the " inner lamina " (i.e. cryptocyst) of his specimens
as being similar to that of Farcimia cereus (p. 3, pi. i, figs. 55, 56), which has a well-
developed cryptocyst. His figure 69 shows the slightly elongated operculum,
figure 70 the cryptocyst, and the vicarious vibracula in the ancestrular area (see
Hastings, 1930 : 714). Figure 71 shows the basal sectors, some of which have 4

ZOOL. 13, 6 n

206 P. L. COOK

pores, although others are shown bearing more. The specimens listed by Silen
(1942, Nos. 8, 9 and 10) have been examined ; they include those described and
figured by Smitt. No. 8 (FED 239) is the original of figure 71 and the actual sectors
have been recognized. The basal sectors are square and more regular than in
Smitt's drawing and the long irregular sectors do not occur, but are composed of
several smaller sectors. Figure 70 is also drawn from this specimen and confirms
that the irregular opesiae shown in the figure are those of vicarious vibracula. In
fact, not only the ancestrular, but all the 7 individuals surrounding it are of this
type. This development is not unusual in C. biporosa and also occurs in colonies from
FED 696, FED 548, Dr. Lagaaij's specimen from the Gulf of Mexico, and 1962. i. 26.
i. from Barbados. Specimen FED 281 is the original of Smitt's figure 69 ; it is a
large colony, 10 mm. in diameter.

The figures given by Angelis (1899, pi. B, figs. 7, 8) of the basal surface of his
specimens from the Pliocene of Catalonia, show long irregular sectors, and are
attributable to C. canariensis. Figure 6 (in which the zooecia are shown upside-
down) has the appearance of C. biporosa, with a wide cryptocyst and straight-sided
opesia. The figure is very similar in appearance to that of Busk's Crag specimens.
Angelis's descriptions however, are taken from that of Manzoni (1877) and refers to
C. canariensis. No fossil Spanish material has been seen, but specimens figured by
Galopim de Carvalho (1961, pi. i, figs. 1-3, Pliocene, Portugal) also show the straight-
sided opesia and small basal sectors of C. biporosa. Pliocene material from Italy also
consists of both C. canariensis and C. biporosa, (see pp. 198 and 204), and the cor-
relation of the characters present is respectively consistent with the diagnoses of
the two species.

The specimens figured from the Pliocene of Argentina by Canu (1908, pi. 5, figs.
8, 9, 10) were extremely worn, but figure 8 perhaps shows 2 vicarious vibracula (in the
bottom right-hand corner).

Both Norman (1909, pi. 37, figs. 4, 5), and Waters (1921, pi. 29, fig. 5), who received
his Madeiran material from Norman, show 2-4 pores on the basal sectors of their
specimens. Examination of the plentiful Madeiran colonies in the Norman Collection
shows that they, too, consist of a mixture of both C. biporosa and C. canariensis.
Norman and Waters both stated that they considered C. canariensis andC. guineensis
(Busk) to be synonymous. In view of the mixed nature of their material it should be
noted that the salient lateral cryptocyst and small square basal sectors of C. biporosa
are very similar to some specimens of C. guineensis, which was, however, distin-
guished and defined by Hastings (1930 : 714).

The material from Costa Rica figured by Canu & Bassler (1918, 1919) as C.
canariensis, belongs to C. biporosa. The same set of photographs was reproduced
in both these papers and was again used to illustrate American fossil " C. canariensis "
in 1920 and 1923. The photographs show i) small colonies, natural size, with flat-
tened bases ; 2) zooecia with well-developed lateral cryptocysts, and with one
vicarious vibraculum at the lower right-hand edge ; 3) the basal sectors, each with
2-4 pores. The introduction of C. biporosa was illustrated by material from Santo

POLYZOA FROM WEST AFRICA 207

Domingo (1923). The fossils from the Gulf of Mexico region (1928), were illustrated
by photographs of Recent C. canariensis from the same area.

The specimens described by Hastings (1930 : 714) from Gorgona, consist of colonies,
the majority of which have regenerated from broken fragments. Where an ances-
trula is present, the vicarious vibracula near it have smaller, more auriform opesiae
than those occurring elsewhere in the colony.

McGuirt (1941 : 46) mentioned the difficulty in distinguishing his Miocene material
from Louisiana, attributed to C. canariensis, from C. biporosa. The specimens
figured show vicarious vibracula (pi. i, figs, i, 3) and the small, square basal sectors
(figs. 2, 5), and are certainly of C. biporosa. The zooecia in figure 8 have unusually
narrow cryptocysts, and may have been worn.

Of the colonies collected at the surface of the Atlantic off the west coast of Africa
(see Silen, 1942 : 13, specimen FBS 321 above, and p. 194), 8 colonies belong to
C. biporosa. The ninth is a specimen of C. owenii (see p. 213).

Osburn's figure (1950, pi. 3, figs. 2, 3) shows small, square sectors with 1-3 pores
(fig. 3), the zooecia in figure 2 have the well-developed lateral cryptocyst of C. biporosa,
and Soule (1959 : 8) mentioned vicarious vibracula in his Californian material.

Marcus & Marcus (1962) gave a short description of the central vicarious vibracula,
and, on pi. i, fig. 2, illustrated the basal surface of one of their colonies with small
segments arranged concentrically. Their material was collected from the northern
and southern coasts of Brazil, no specimens being found in the intermediate area.
The shallow depths at which some specimens occurred (3 m. near Cabo Frio) were at
" sheltered localities ", where there must have been little turbulence.

Lagaaij's figures (1963, pi. 26, figs. 4, 5) of material from the Gulf of Mexico are of
C. biporosa, as is the unnamed species mentioned by Cheetham and Sandberg (1964).

The zoarium rarely attains a diameter of more than 10 mm. ; fragments of large
colonies estimated at 16 mm. diameter were found in fossil material from the U.S. Nat.
Mus. Collection, 80747, and a colony of n mm. diameter, covered with 3 small
colonies of Onychocella angulosa (Reuss) occurs in material from Madeira, 1962.
i. 24. 7.

Unlike C. canariensis, the sides of the opesiae are almost straight, and in the great
majority of specimens the cryptocyst descends gently forming a salient lateral shelf
and obscuring the pores in the lateral walls, which are not seen in frontal view as in
C. canariensis. In some fossil specimens the cryptocyst descends more steeply, but
this seems to be confined to zooecia of fragments of large zoaria, and does not occur
in the smaller, whole colonies. The cryptocyst of the zooecia of the central area are
sometimes slightly thickened by secondary calcification, but a lamina is never formed
as is found in C. guineensis (see Cook, 1965 : 172) ; a similar thickening is found in
C. monotrema.

The ordinary vibracula are of one kind only and vicarious vibracula are nearly
always present, especially near the centre of the colony ; their opesiae are auriform,
and there is a distinct, prominent gymnocyst. The setae of Recent specimens have
the same type of basal sclerite and are similar to those of the ordinary vibracula (see
also Hastings, 1930 : 714) ; they have no sigmoid curve, however, and in lateral view

2o8 P. L. COOK

they are broad, scimitar-shaped, and hooked at the tip (see Text-fig, ih). They thus
differ from the short, hooked setae of the vicarious vibracula of C. canariensis. Both
Marcus & Marcus (1962 : 288, pi. i, fig. 2) and Lagaaij (1963 : 183) described the
vicarious vibracula of the ancestrular area as regenerated. Although the original
zooecia may regenerate as vibracula in some cases, many specimens (notably those
from Gorgona and the Gulf of Mexico) , show that the vicarious vibracula are budded
primarily as part of the normal development of a colony. Vicarious vibracula are
often associated with areas of regenerating growth (see Cook, 1965 : 160), and here
also, they are some of the first individuals to be budded, not the results of regeneration
of the older zooecia.

The development of the basal porous sectors in C. biporosa differs from that of
C. canariensis in that the chambers appear to develop more quickly, and large pores
are rarely seen at the growing edges of young colonies. The tangential sector
boundaries are always present on the basal side of the young peripheral zooecia,
unlike C. canariensis (see above).

The layers of kenozooecial chambers are more numerous than C. canariensis, even
in young colonies. They frequently conceal the original substrate, and in many
colonies are so thick that the basal surface is flattened. In section these zoaria show
successive rows of chambers up to 14 deep. A slide of sectioned colonies of C. biporosa,
where the species is mounted with C. guineensis for comparison (1879. 5- 2^- 6,
Madeira + 1929. 5. 10. i, Australia), was figured by Hastings (1930, pi. 8, fig. 38, as
C. canariensis}. The identity of the species in no way affects her description of the
chambers as a distinction from C. guineensis, as those of C. canariensis are exactly the
same in appearance, although they rarely reach a comparable number of layers, even
in large colonies of this species.

One of the specimens from the Miocene of Santo Domingo (U.S.N.M. Coll.) shows a
form of growth similar to normal colonies of C. doma (see Cook, 1965 : 162). The
zoarium is small and high, the basal side is half filled by secondary calcification,
and there are 3 rows of peripheral vibracula, the last of which encroaches on the
basal side (see PI. i, fig. 5).

The geographical distribution of specimens of C. canariensis and C. biporosa, with
and without vicarious vibracula, falls into distinct groups. The first, northerly area,
consists of records from the S.W. Mediterranean, Spain, Portugal, and the Canaries ;
no vicarious vibracula have been found in colonies of either species from these
localities. Material of both species from Madeira, the Cape Verde Islands and the
coast of Senegal has colonies both with and without vicarious vibracula ; whereas all
the specimens from the southerly localities in the Gulf of Guinea and the Bay of
Biafra have vicarious vibracula. Recent and fossil specimens of C. biporosa from the
Gulf of Mexico are rarely without vicarious vibracula, whereas none have been found
in the Recent colonies of C. canariensis from this area.

C. biporosa differs from C. canariensis consistently in the character of the basal
surface, of the cryptocyst, in the number of tentacles, in the absence of a second kind
of ordinary vibraculum, and in the form of the vicarious vibraculum. The fossil
record shows whereas there are apparently no certain descriptions of C. biporosa

POLYZOA FROM WEST AFRICA 209

occurring before the Pliocene in south-western Europe, the only American records
of C. canariensis, from the Gulf of Mexico and Florida, are Recent. The Miocene
faunas of the eastern and western Atlantic were distinct.

C. canariensis and C. biporosa have similar geographical distributions, live in
similar depths, under the same ecological conditions, and are thus frequently associ-
ated in Recent collections. There is a similar correlation between their occurrence
and that of the D. umbellata-complex (see Canu & Bassler, 1918 : 119), and as many
as 5 lunulitiform species occur together at Konakrey and Cape Verde, and Stn. 45,
" Calypso " Coll. I, off Cape Lopez, in the Bay of Biafra.

The confusion between C. canariensis and C. biporosa, and the occurrence of
vicarious vibracula in both species, make it advisable to compare them with the only
other species with similar characters, namely C. monotrema (Busk).

8 Cupuladria monotrema (Busk)

Cupularia monotrema Busk, 1884 ; 207, pi. 14, figs. 5, 5a, 5b, off Bahia, 10-20 fath.
Cupulana canariensis Busk : Waters, 1888 : 36, pi. 3, fig. 2, Bahia, 10-80 fath.
Cupuladria monotrema (Busk) Hastings, 1930 : 715.

MATERIAL. LECTOTYPE (chosen here), B.M., 1887. 12. 9. 820. British Museum
Coll. Off Bahia, 10-20 fath., 1887. 12. 9. 820, 82oA, Challenger Coll. (V). As above,
1944. i. 8. 346, Busk Coll. (V). As above, 10-80 fath., 1889. 12. 12. 9 (V) Suppl.
Challenger Coll. As above, 1890, 3. 22. n (V). Bahia and Pernambuco, 11° 45' S.,
37° 15' W., 16-28 fath., 1890. i. 30. 33-44 pt., and 13° 50' S., 38° 46' W., 32 fath.,
1890. i. 30. 33 pt. (VV).

British Antarctic Expedition Coll. Stn. 42, off Rio de Janeiro, 40 fath., 878A.

C. monotrema does not occur in any of the west African Collections.

Zoarium fairly large, maximum diameter 10 mm. Zooecia with cryptocyst des-
cending quite steeply, lateral pores just visible in frontal view. Operculum opening
within a strongly chitinized area of the frontal membrane, which is surrounded by a
thickened band. Operculum frequently heavily chitinized, but proximal sclerite
absent. Basal surface with only one layer of kenozooecia. Sectors small and square,
with 1-3 pores. Vicarious vibracula numerous, nearly as large as the zooecia, with
auriform opesia and long proximal gymnocyst. Vicarious vibracula often inter-
calary, reaching the penultimate peripheral circle of zooecia. Setae scimitar-shaped,
similar to those of C. biporosa.

DIMENSIONS. Lz 0-40-0-50 mm., Iz 0-26-0-37 mm., La 0-40-0-45 mm., Lvo
0-09-0-12 mm., Lv 0-35-0-45 mm., Ls 0-50-1-00 mm., Lvs 0-30-0-40 mm., Lo o-io-
o-n mm., lo 0-12-0-13 mm., Lop 0-25-0-30 mm., lop 0-15-0-20 mm.

The opesiae of the zooecia of the central area are slightly reduced by secondary
thickening as in C. biporosa. The vicarious vibracula are similar to those of C.
biporosa, but are far more numerous. The opercular area is unlike that of other
species, although the operculum itself is like that of C. biporosa in shape. Frequently
its dark brown colour ends abruptly at the junction with the frontal membrane, giving
the appearance of a curved proximal sclerite. In many colonies, a band of thickening

2io P. L. COOK

surrounds the distal part of the operculum and then extends proximally and across
the frontal membrane. This whole area, with the operculum opening within it, is
more heavily chitinized than the remaining frontal membrane surrounding it. It is
particularly noticeable in dry specimens (such as those seen by Busk) , that the area
enclosed by the thick chitinous band shrinks differentially in relation to the rest of
the frontal membrane, producing the " hippocrepian " shape described by Busk,
who was thus misled into thinking that this part of the whole frontal area was the
operculum. In his explanation of terms (p. xvi), Busk did not give a defini-
tion of " hippocrepian ", but his "coarctate " form of orifice is similar to the lepra-
lioid shape of the frontal area in C. monotrema. The chitinized band is not present in
all specimens, nor in all zooecia of the zoaria in which it may occur.

As stated by Hastings (1930 : 715), C. monotrema is clearly distinct from C. canarien-
sis. It is very similar to C. biporosa, and may indeed be synonymous, but until more
material is available, it is distinguished by the opercular area, the more numerous
vicarious vibracula, and the single layer of basal kenozooecial chambers.

Marcus & Marcus (1962 : 286) found no specimens of C. monotrema in collections
ranging from Sao Paulo north to the mouths of the Amazon. C. biporosa was found
only at the extreme North and South of the collecting area, not in the Bahia area.

9 Cupuladria multispinata (Canu & Bassler)
(PI. 2, figs. 2A, B, Text-figs. 2d)

Cupularia owenii (Gray) Busk, 1884 : 207, St. Vincent, Cape Verde Islands, n fath., mud (not

C. owenii (Gray) see p. 213).
Cupularia johnsoni Busk ; Norman part, 1909 : 290, pi. 38, figs. 5, 6, Madeira (not figs. 1-4

= C. doma, see p. 216).
Cupularia denticulata Waters, 1921 : 413, listing specimens labelled " C. deformis " by Busk, in

the British Museum, from Portugal and the Mediterranean, 50 m. to 45 fath.
Cupularia multispinata Canu & Bassler, 1923 : 78, text-fig. I3H, Oran, 105 m. (not text-fig. 13 I

= D. umbellata, see p. 221). 1928 : 22.
HCupularia haidingeri (Reuss) : Canu & Bassler, 1923 : 77, pi. i, figs. 13-17, Miocene, Europe

(not C. haidingeri (Reuss) see Lagaaij, 1952 : 35).
Cupularia denticulata var. multispinata Waters, 1926 : 427, pi. 18, figs. 2, 4, 7, Oran (C. denticulata

in the explanation of the plate) .
Cupuladria owenii (Gray) : Sil6n, 1942 : 15, Madeira.

MATERIAL. "Calypso" Coll. I. Stn. 7*, 9° 40' N., 13° 53' 5" W., 17^.56,
18 m. C4E.

Coll. II. Stn. 26, 15° 16' 30" N., 23° 47' 31" W., i8.xi.59, 50-65 m., C65H.

Marche-Marchad Coll. I. Konakrey, Guinee Ise., 2E. S.W. Madeleines, 15 .ix.53,
2oA ; g.i-54, 45-46 m., 26E. Sud de presque 1'ile du Cap Vert, i8.ii.54, 95 m., 33 A.
Baie de Seminoles, Goree, 8.xii.53, 38 m., 3gA (C) (+ Hippoporina lacrimosa Cook)

Coll. II. S. Baie de Goree, i8.ii.54, 95 m., 3oC. Baie de Seminoles, Goree,
8.xii.53, 38 m., 36A.

Coll. III. Sud de Goree, 13.^.53, 33-35 m., 9C ; 34-37 m., I7C. Either S.W.
Madeleines, I5.ix.53, 48., or Sud de Goree, 13.11.53, 34-37 m., i6G. Dragage i
"Gerard Freca ", i8.ii.54, 23G ; Dr. 4, 97-98 m., 28!; Dr. 5, 27C. 33-34 m.,

POLYZOA FROM WEST AFRICA 211

27 . ii . 53, 2gE. Mission Guinee Pte. 50, Parages des Fles de Los, Konakrey, 21 . i . 53,
19 m., 300 (C). No information, 24!.

British Museum Coll. St. Vincent, Cape Verde Islands*, n fath., mud, 1887.
12. 9. 821, Challenger Coll. Port of Oratava, Canaries, 1899. 7. i. 1224, Busk Coll.
Bay of Funchal, Madeira, 1963. i. 16. 4, Busk Coll ; 50 m., 1963. 2. 28. n. Cook
Coll., mud*. Madeira, 1963. i. 16. 2 and 3, Norman Coll. Mediterranean, 1899. 7. i.
1236, and on 1223 with C. canariensis , " Porcupine ", Busk Coll. Spain, 1899. 7- I-
1239, Busk Coll. Cape Rosa, Algeria, 1899. 7. i. 1233, Busk Coll. Ras el-Amouch,
Algeria, 45 fath., 1963. i. 16. i. Tangier Bay, 1890. 4. 15. 23. Cape of Good Hope,
1842. 12. 2. 19, Belcher Coll. South Africa, False Bay, 50 m., fine sand*, 1963. i.
16. 5, Univ. Cape Town Coll. S. Africa, False Island N. 8-5 miles, 53 fath., 1949. n.
10. 6696 ; Bakhoven Rock W., \ N. f mile, 22 fath., 1949. n. 10. 617 ; Cape St.
Blaize, 7-5 miles N. 37 fath., 1949. n. 10. 946, 6256, 6266, 6276 ; Cape Infanta
N.E. by N. 19 miles, 46 fath., 1949. n. 10. 6226, 6236 ; N.E. by N. J N., 45 fath.,
1949. ii. 10. 6216 ; Stalwart Point, N.N.W. 9 miles, 53 fath., 1949. 11. 10. 6526 ;
Nanquas Peak N. 46° E. 9 miles, 63 fath., 1949. n. 10. 6746. Burrows Coll.

Discovery Coll. Stn. 299, I4.X.27, 7-11 m., Tarrafal, S. Antonio, Cape Verde
Islands, 86 A.

Naturhistoriska Riksmuseet, Stockholm, Madeira, FED 36.

C. multispinata is not represented in the Achimota Collection.

Zoarium frequently large, (maximum diameter measure, 16 mm.), zooecia with a
large range in size. Sides of vestibular arch straight, salient distal denticles absent.
Operculum without a proximal sclerite. Descending lateral cryptocyst tuberculate,
with 4-8 irregular denticles, themselves ending in fine spinules ; proximal spinules
infrequent. Central zooecia closed. Basal surface with large, irregular, coalescent
tubercles and grooves frequently marked by a salient calcareous thread.

DIMENSIONS. Lz 0-55-0-76 mm., Iz 0-32-0-47 mm., La 0-45-0-52 mm., Lvo 0-15-
0-21 mm., Ls 0-60-1-90 mm., Lo 0-12-0-14 mm., lo 0-15-0-19 mm.

Busk's specimen from the Cape Verde Islands (Challenger Collection) differs
slightly from the other west African specimens of C. multispinata. The colony is
7-5 mm. in diameter, but the peripheral zooecia are smaller than in other colonies of
comparable size. The sides of the vestibular arch are straight, however, with no
evidence of a distinct pair of distal denticles, as in C. owenii. The basal surface is
worn, and almost smooth, but two areas show large irregular tubercles and faint
threads marking the radial grooves. The specimen certainly belongs to C. multi-
spinata.

Norman (1909 : 290) remarked that specimens of Cupularia oweni (sic) were not
found by him from Madeira. The Madeiran specimens which he referred to C.
johnsoni are, however, a mixture of C. multispinata and C. doma. He included C.
reussiana Manzoni and C. doma Smitt (sic) in his synonymy of C. johnsoni (= C. doma
(d'Orbigny)). Part of the material which he referred to C. lowei (= D. umbellata,
see p. 221), is also C. multispinata, as is the " variety " of C. johnsoni he described in
which " the radiating ridges are greatly developed, and bear two rows of tubercles

212

P. L. COOK

regularly placed and of much larger size than those of the type ". These specimens
were figured on pi. 38. figs. 5, 6, and show the typical character of C. multispinata.

Waters (1921 : 413), in discussing Cupularia lowei (see p. 221), listed several British
Museum specimens of another species, which had similarly deformed zoaria. He
proposed to place these " under denticulate ". Waters's publication of the manuscript
name on the labels of Busk's specimens in the British Museum was not accompanied
by a description, definition nor an indication (see International Code of Zoological
Nomenclature, 1961, Articles 12 and 16). The specimens he listed are all identical
with C. multispinata, as was suggested by Canu & Bassler (1923 : 78). Waters later
(1926 : 427) confirmed this, and although in the explanation of pi. 18, on p. 432, he
still referred to the species as Cupularia denticulata, he added the note " This is the
C. multispinata, Canu & Bassler ". On p. 427 he referred to it as " Cupularia denti-
culata var. multispinata", C. denticulata is a synonym of C. owenii (see below).
The distribution of C. multispinata given by Canu & Bassler (1928 : 23) was derived
from Waters's list of specimens given in 1921.

FIG. 2. Development of the cryptocyst in Cupuladria and Discoporella. Drawings semi-
diagrammatic, a-e, Cupuladria. a, b. C. owenii disciformis . a. Form with vestibular
arch with convergent sides, and several cryptocystal denticles, b. Form with vestibular
arch with divergent sides, and very reduced cryptocystal denticles, c. C. owenii. d.

C. multispinata e. C. doma. f-h. Discoporella f. D. reussiana. g. D. ocellata. h.

D. umbellata.

POLYZOA FROM WEST AFRICA 213

Nearly all the deformed colonies originate from regenerating fragments, and the
irregularities are due to one or more radial rows failing to develop (see p. 195).
Canu & Bassler's figure purporting to be of the frontal surface of C. multispinata with
chitinous parts intact (1923, text-fig. 13 I) shows opercula with a strong basal sclerite
and a horizontal cryptocyst with opesiules in one of the zooecia ; it is almost certainly
referable to D. umbellata (see p. 221).

Canu & Bassler (1923 : 77) described, as C. haidingeri, fossil specimens from Europe,
in which the zooecia were large, like those of C. multispinata. C. haidingeri as
defined by Lagaaij (1952 : 35) has small zooecia like C. owenii. Canu & Bassler gave
a Lz of 0-60 mm., and measurements taken from their pi. i, fig. 15 give an average
length of 0-58 mm. It should be noted here that whereas the measurements given by
Canu & Bassler for their C. haidingeri apparently include the vibraculum, those given
for their C. multispinata are found (when compared with text-figure I3H), not to
include it.

The specimens from Madeira referred by Silen to C. owenii have been re-examined,
they belong to C. multispinata.

In west African collections where both C. multispinata and C. owenii are found, the
basal surface of unworn colonies is so different that the two species may be dis-
tinguished by it even in young specimens.

The distribution of Recent specimens of C. multispinata is restricted to two areas :
the northerly, from the south west Mediterranean to the cost of Senegal, and the
southerly, off the the South African coast. Although there are differences between
the two populations, the South African colonies being larger and more domed in shape,
the characters of the zooecia are similar. C. multispinata has been recorded from
depths of 7-105 m.

10 Cupuladria owenii (Gray)
(PL 2, figs. 3A, B, Text-fig. 2c)

Lunulites owenii Gray, 1828 : 8, pi. 3 figs. 15, i5a, I5b Coast of Africa.
Lunulites denticulata Conrad, 1841 : 348, Miocene, N. Carolina.

Lunulites denticulata Conrad : Lonsdale, 1845 : 503, text-figs, a, b, Miocene, N. Carolina.
Cupularia owenii (Gray) Busk, 1854 : 99, pi. 115, figs. 1-5, Coast of Africa.
Not Cupularia denticulata (Conrad) Busk, 1859 : 85 (= C. haidingeri (Reuss), see Lagaaij, 1952).
Cupularia denticulata (Conrad) Canu & Bassler, 1923 : 79, pi. 15, figs. 6-10, Miocene, N. Carolina
and Florida, Pliocene, S. Carolina and Florida.

Lectotype, see Hastings (in press).

MATERIAL. " Calypso " Coll. I, Stn. 7*, 9° 40' N., 13° 53' W., 17^.56, 18 m.,
C4H (C). Stn. 45*, o° 25' N., 9° o' E., 8.vi.56, 73 m., C55G.

Marche-Marchad Coll. I. Konakrey, Guinee Ise., 2E (C). Flor de la bouteille,
Guinee Ise, 21.1.53, 8 m., 3A. S.W. Madeleines, 21.1.54, 46-48 m., 46C.

III. Au sud de presque 1'ile du Cap Vert, iS.ii. 54, 46-50 m., lE. Mission Guinee
Pte 50, Parages des Fles de Los, Konakrey, ,'1.1.53, *9 m-> 3<>A (C).

British Museum Coll. West coast of Africa, 1899. 7. i. 4879, received from Gray,

214 !'• L- COOK

Busk Coll. Canaries, M'Andrew, 1899. 7. i. 1249, Busk Coll., and 1892. 6. 17. 1-4.
Port of Oratava, Canaries, 1899. 7. i. 1224, Busk Coll. South Africa, 1963. i. 16.
12, O'Donoghue Coll. S. Africa, Cape Infanta N.E. by N. 19 miles, 46 fath., 1949.
ii. 10. 622C, 623C, N.E. by N. \ N., 13 miles, 43 fath., 1949. n. 10. 6i8B, N.E. by
N. | N., 14-5 miles, 45 fath., 1949. n. 10. 62iA ; Great Fish River lighthouse N.E.,
9 miles, 51 fath., 1949. n. 10. 653A, Burrows Coll. Palaeontological Department,
2 miles S.W. Magnolia, S. Carolina, Duplin marl, Miocene, D 34410-18 and D 34419-
30.

Naturhistoriska Riksmuseet, Stockholm. Atlantic, 27° 16' N., 23° 21' W., at the
surface, FBS 312 part, (No. 7) (see p. 194).

Philadelphia Academy, Conrad Coll. Wilmington, N. Carolina.

U.S. Nat. Mus., Bassler Coll. Pliocene (Waccamaw marl), Waccamaw River, S.
Carolina, No. 80710, Plesiotypes C. denticulata, fig'd. Canu & Bassler, 1923. Caloo-
sahatchee marl, Pliocene, Florida.

C. owenii is not represented in the Achimota Coll.

Zoarium small (average diameter 5 mm.), zooecia small. Sides of vestibular arch
curved, convergent, with a pair of well developed distal cryptocystal denticles.
Operculum longer than wide, without a proximal sclerite. Descending cryptocyst
finely tuberculate, with 4-6 denticles varying from wide, irregular denticles to narrow
spinules. Central zooecia closed. Basal surface grooved, finely and regularly tuber-
culate or smooth and glassy.

DIMENSIONS. Lz 0-37-0-47 mm., Iz 0-20-0-37 mm., La 0-32-0-35 mm., Lvo 0-08-
0-13 mm., Ls 0-60-1-40 mm., Lo 0-07-0-09 mm., lo 0-09-0-10 mm.

The slide, 1899. 7. i. 4879, which is labelled " West coast of Africa ", was received
by Busk from Gray. It consists of two fragments, one of which was figured by Busk
(1854, pl- I3:5> ng- 3), see Hastings (in press). The frontal of Gray's specimen is worn,
but shows the small zooecia and convergent vestibular arch. The basal surface is
smooth, glassy and grooved, a condition found in many specimens from the Marche-
Marchad Collection. Specimens from the Achimota Collection have a finely tuber-
culate basal surface, but also show that the smooth type is not an effect of wear.
Some colonies have the more central parts smooth and the later growth tuberculate ;
whilst in others the position of these types of surface is reversed.

The identity of L. denticulata Conrad with L. owenii Gray was suggested by
Lonsdale (1845 : 503), who also described the closed, central zooecia. Specimens of
C. denticulata from Conrad's Collection, from Wilmington, N. Carolina, and from
Bassler's Collection, from S. Carolina and Florida, agree in all respects with the type of
C. owenii and thus extend its range in time and space to the Miocene and Pliocene of
North America. The fossil material shows a similar range in variation of characters
of both frontal and basal surface to that of Recent zoaria from west Africa. The
specimens from the Caloosahatchee marl are small, the largest zoarium having a
diameter of 4 mm., the smallest of 2 mm. ; and those from the Waccamaw marl are
generally larger and flatter. A few zoaria from Caloosahatchee have a growth form
approaching that of C, doma ; the basal concavity being nearly filled by secondary

POLYZOA FROM WEST AFRICA 215

calcification. However, the surface is still finely tuberculate, and there are no
peripheral closed zooecia or enlarged vibracula (see below).

Generally, the small size of the zooecia distinguishes colonies of C. owenii from those
of C. multispinata, even to the naked eye. In young colonies, the size of some of the
zooecia may be similar, but the zoaria may then be separated by the basal surface
(see above). In well preserved colonies of C. owenii, the strongly incurved vestibular
arch, with the distinct distal denticles, is completely unlike that of C. multispinata,
and resembles that of C. doma, from which C. owenii is distinguished by the absence
of any trace of proximal sclerite on the operculum, the basal surface, and by the
absence of peripheral kenozooecia or enlarged vibracula. C. owenii is very similar
to the European fossil C. haidingeri (Reuss) (see Lagaaij, 1952 : 35, 1953), but differs
in its finely tuberculate or smooth basal surface. A specimen of D. owenii (Marche-
Marchad Coll. Ill, 3oA) has a regenerated zoarium where many of the first zooecia
budded from the edge of the fragment have become closed in the same manner as the
ancestrular zooecia of other colonies.

The S. African zoaria are slightly larger and more domed than those from west
Africa, and the distal pair of denticles is less well developed. The small zooecia and
the glassy basal surface distinguish them from S. African colonies of C. multispinata.

The Recent distribution of C. owenii appears to be restricted to the African coast
where it is known from the Canaries to the Bay of Biafra, and off S. Africa. It has
not been found from depths of more than 95 m., and for this reason is unlikely to have
been the species listed by Calvet (1907 : 393) from 1900 m., Cape Verde Islands, as
" Cupularia umbellata ... a la forme de denticulata de Conrad " (see p. 222).

ii Cupuladria owenii subsp. disciformis n. subsp.
(PI. 2, figs. lA, B, Text-figs. 2a, b)

HOLOTYPE. Museum d'Histoire naturelle, Paris, C43A, pt.

PARATYPES. " Calypso " Coll. I. Stn. Pi, entre Pta da Mina et I. S. Ana,
2i.vi.56, 10-12 m., C44A. Stn. ?7*, entre Pta da Mina et P. Novo Destino, 26. vi.
56, 6 m., C47A. Stn. P8, I. S. Ana, 27.vi.56, 2-4 m., C43A.

C. owenii disciformis is not present in the Marche-Marchad nor the Achimota
Collections.

Zoarium flattened, disc-like, average diameter 7 mm. Zooecia small, narrow,
Cryptocyst descending steeply, with 2-4 small, simple denticles. Sides of vestibular
arch not strongly convergent, distal denticles weakly developed. Central zooecia
closed. Opercula wider than long. Basal surface with grooves and fine tubercles.
Tentacles 13-16 (see p. 203).

DIMENSIONS. Lz 0-37-0-53 mm., Iz 0-20-0-35 mm., La 0-40 mm., Lvo 0-09-0-10
mm., Ls 0-60-0-90 mm., Lo 0-05-0-06 mm., lo 0-11-0-12 mm.

At first sight, this form is so unlike C. owenii that it might be thought to be a distinct
species. Detailed examination of the colonies shows, however, that some zooecia
have a vestibular arch with convergent sides, and more numerous cryptocystal

216 P. L. COOK

denticles, than others. These zooecia approach those of C. owenii in appearance.
The vestibular arch varies in character in C. owenii (see above), and it is possible that
a series of forms might be found linking the two extremes of variation.

This population was collected from one small area in the Bay of Biafra, from very
shallow water. The zoaria all have polypides, and chitinous parts intact. They
were associated with many specimens of Caulibugula sp., which were not, however,
attached to the lunulitiform zoaria.

C. owenii disciformis is separated here because each of the large number of colonies
examined (over 100), shows a high level of consistency of correlation of the characters
distinguishing them from typical C. owenii. These characters are : the flattened
shape ; the slightly larger zooecia and coarser basal tubercles ; the shallow opercula ;
and the reduction in cryptocystal denticles. In some zoaria, this reduction is such
that the zooecia resemble those of C. pyriformis (Busk).

Studer (1889 : 7, 13) listed C. pyriformis from west Africa, but it seems unlikely
that it occurs anywhere in the eastern Atlantic (see Cook, 1965, 169). Although
Studer's record may have belonged to C. canariensis or C. biporosa, it is also possible
that the specimens were of C. o. disciformis, which, like C. pyriformis, has a ridged,
finely tuberculate basal surface.

Five unnamed fossil specimens from France have similar characters to those of
C. o. disciformis. Two zoaria are from the Burdigalian of Leognan (Lower Miocene,
S. of Bordeaux, Gironde, D 25002 and 25004) ; they are very flat, regenerated frag-
ments. The remaining three zoaria are from the Pliocene of Biot (near Antibes,
D 49301-3) ; they are slightly more domed, and originate from ancestrula. The
zooecia of all the specimens are slightly larger than those of C. o. disciformis (Lz
0-40-0-57 mm.), but like them, nearly all have a vestibular arch with divergent sides,
and rarely more than 3 small lateral cryptocyst denticles. The basal surface of all
the colonies is finely tuberculate. No reference has been found in the literature to
this form, and further material would have to be examined before its relationship
with C. o. disciformis could be established.

12 Cupuladria doma (d'Orbigny)
(PI. i, fig. 7, Text-figs. 2e, 3)

Discoflustfella doma d'Orbigny, 1853 : 561, Recent, Algeria.

Cupularia johnsoni Busk, i85ga : 67, pi. 23, figs. 1-5, Madeira. Norman (part), 1909: 290, pi. 38,
figs. 1-4, Madeira (not figs. 5-6 = C. multispinata, see p. 210). Canu, 1917 : 139. Waters,
1921 : 413, pi. 29. fig. 17, pi. 30, figs. 23, 30, 31, Oran, Madeira.

Cupularia doma (d'Orb.) Smitt, 1873 : 15, pi. 3, figs. 81-84, Florida, 29 fath. Canu & Bassler,
T923 : 77, pi. i, fig. 15, pi. 15, figs. 1-5, Miocene, N. Carolina. 1928 : 64, pi. 6, figs. 2-5,
Florida, 56 fath. ig28a : 23, pi. 2, figs. 1-4, Cap Blanc, 20-30 m.

^Cupularia reussiana Manzoni : Neviani (part), 1895 : IO2» Recent, Algeria.

Discoporella doma (d'Orbigny) McGuirt, 1941 : 64, pi. 2, figs, i, 4, Miocene, Louisiana. Maturo,
1957 •' 41. fig3- 48, 49, N. Carolina. Cheetham & Sandberg, 1964 : 1022, text-fig. 15, Quater-
nary, Louisiana.

Cupuladria doma (d'Orbigny) Gautier, 1962 : 54, Algeria, 75-80 m.

POLYZOA FROM WEST AFRICA 217

MATERIAL. " Calypso " Coll. I, Stn. i*, 21° 05' N., 17° 14' W., io.v.56, 43-45 m.,
CiH.

Marche-Marchad Coll. I, Konakrey, Guinee Ise, 2D. Flor de la bouteille, Guinee
Ise, 21.1.53, 8 m., 3D. Sud de Goree, 27.x. 53, 38-42 m., 7A;24.xi.53, 40 m.,
8A ; 39-5 m., gF ; 40-41 m., nG. Bale de Seminoles, 8.xii.53, 38 m., 396. S.W.
Madeleines, i5.ix.53, 48 m., 22A ; 9.1.54, 45-49 m., 26F ; 21.1.54, 46-48 m., 46!.).
Sud de presque Tile du Cap Vert, i8.il. 54, 95 m., 336.

Coll. II. Alignment Goree, Cap Manuel, hauteur de Madeleines, I9-X.56, 35-42
m., gA. S.W. Madeleines, 9.1.54, 47-5111., 3iC. Bale de Seminoles, 8.xii.53, 38 m.,
36B.

Coll. III. Sud de presque Tile du Cap Vert, iS.ii. 54, 46-50 m., iF. Sud de Goree,
I3.xi.53, 33-35 m., gD. Either as above or S.W. Madeleine, I5.ix.53, 48 m., i6H
and 170. Dragage i "Gerard Freca ", iS.ii. 54, 23!!; Dr. 4, 97-98 m., 28F ;
Dr. 5, 150, 27E. 27.xi.53, 33-34 m., 2gF. No information, 24J.

British Museum Coll. Madeira, 1879. 5- 2&- 6, and 1911. 10. i. 649, 650, 1912. 12.
21. 1003, Norman Coll ; Bay of Funchal, 50 m., 1963. 2. 28. 5, Cook Coll., mud*.
Canaries, 1899. 7. i. 1240, Busk Coll. Mediterranean, 1899. 7. i. 1243, 152 fath. ;
1242, 5-51 fath. ; 1245, 151 fath., Busk Coll. Tangier, 1899. 7. i. 1244, 164, Busk
Coll. Ras el-Amouch, Algeria, 1899. 7. i. 1247 '> I245> 45 fath., Busk
Coll. Oran, 1903. 3. 6. 1-4, and 1911. 10. i. 645, Norman Coll. Gulf of Mexico,
28° 45' N., 85° 2' W., 30 fath., Stn. 2405, " Albatross ", 1932. 3. 7. 102, Canu &
Bassler Coll. S.S.W. John's Pass, Florida, 34 fath. 1959. i. 6. 5, Lagaaij Coll.

C. doma is not represented in the Achimota Collection.

Zoarium small (average diameter 4-5 mm.), steep-sided, sometimes solid basally.
Zooecia small, vestibular arch strongly curved, with well developed distal crypto-
cystal denticles. Operculum with a complete or partial proximal sclerite. Crypto-
cyst descending gently, with wide denticles which occasionally fuse proximally.
Basal surface frequently solid, grooved or smooth, or with a small central cavity
with elongated spiny tubercles. Central zooecia and those at the periphery closed
in fully grown colonies. Peripheral vibracula enlarged, occasionally more than one
series developed. Tentacles 13-16 (see p. 203).

DIMENSIONS. Lz 0-34-0-42 mm., Iz 0-30-0-37 mm., La 0-34-0-37 mm., Lvo 0-09-
0-19 (peripheral) mm., Ls 0-50-1-70 mm. (peripheral), Lop 0-07-0-10 mm., lop
0-10-0-12 mm.

D'Orbigny 's description of Discoflustrdla mentions the " Gros pore special "
(= vibraculum) in front of each zooecium, and his specimens of D. doma were examined
and their identity confirmed by Smitt (see below). D'Orbigny also described and
figured Discoflustrellaria doma (p. 509, pi. 722, figs. 6-10), a Cretaceous fossil, probably
referable to Lunulites (see Waters, 1921 : 408).

Unfortunately, Canu and Bassler (1923 : 77) confused the two references and
quoted " Discoflustrellaria doma d'Orbigny, p. 561 " in their synonymy, a combination
followed by Gautier (1962 : 54), and by Annoscia (1963 : 227, as Disco frustrettaria
(sic)).

218

P. L. COOK

Smitt compared his specimens from Florida with Algerian material from Busk ;
he also examined specimens in d'Orbigny's Collection (see Canu & Bassler, 1923 : 78),
as did Canu (1917 : 139), who, nevertheless, still placed C. doma in the synonymy of
C. johnsoni. There has been some confusion of C. doma with Discoporella reussiana
(see below). Specimens labelled " C. reussiana ", some from Levinsen (1903. 3. 6.
1-4), others in the Norman Collection (1911. 10. I. 645), are referable to C. doma.

The west African material shows a range in development of the proximal opercular
sclerite, which is, for example, complete in Marche-Marchad Coll. I 396, but only
partial, and absent centrally in Coll. II gA.

The basal surface of C. doma may consist of a concavity, with long spinous tubercles,
or be flat, and smooth or tuberculate. Although the solid state is apparently an effect
of age, it is not correlated with growth in a simple manner, as very small colonies may
have flattened bases, whereas in larger zoaria they may be concave. There seems,
however, to be a correlation between the occurrence of peripheral kenozooecia and a
flattened, solid basal surface, and presumably a colony at this stage does not grow
further, as no zoarium has been found with new zooecia budding from closed periphe-
ral kenozooecia. In these fully grown colonies, there may be several series of peripheral
vibracula ; they are enlarged, and their setae may be much longer than those of the
rest of the colony. The intervening kenozooecia are still present, but have become
progressively smaller, until the zoarium has the appearance of possessing two or
three rows of vibracula only at the periphery. A section through a zoarium shows
that the cavity of each zooecium is present, but that the angle of its longitudinal axis
in relation to the axis of the colony is such, that its projection on to the plane of the
zoarial surface is very small. Thus, when the opesia is closed by calcification, the
kenozooecial frontal is hardly noticeable, and the enlarged vibracular chambers
appear to be adjacent to each other (see Text-fig. 3).

FIG. 3. Cupuladvia doma, closed peripheral zooecia, drawings semi-diagrammatic, a.
Section through a zoarium, x 20. b. Frontal view of enlarged peripheral vibracula and
closed peripheral zooecia, X4O.

POLYZOA FROM WEST AFRICA 219

There may be some relationship between the presence of closed peripheral keno-
zooecia and the form of the zoarium. Although the closed zooecia are typical of
C. doma they may also occur in specimens of other species which develop apparently
aberrant small, high zoaria with a solid base (see Cook, 1965 : 162, and p. 208).
Duvergier (1924 : 20) mentioned that some specimens of C. porosa Busk (= C.
haidingeri (Reuss), see Lagaaij, 1952 : 36) had two rows of peripheral zooecia.

C. doma is very similar in many features to C. owenii, which occasionally develops
small high colonies (see p. 215), but it may be distinguished by its slightly smaller
zooecia, its basal surface, and the presence of a proximal sclerite on the operculum.

The African distribution of C. doma is confined to the more northerly coasts from
Algeria to Senegal, and includes the Canaries and Madeira, but not the Cape Verde
Islands. It extends to greater depths than C. owenii, and has been recorded from
8-369 m.

13 DISCOPORELLA d'Orbigny

Discoporella d'Orbigny, 1852 : 472. Hastings 1930 : 718. Cheetham & Sandberg, 1964 : 1022.

TYPE-SPECIES Lunulites umbellata Defrance, 1823, Miocene, France.

Zoarium lunulitiform. Zooecia with a well-developed vestibular arch. Horizontal
cryptocyst lamina complete, formed of fused denticles. Opesiular indentations or
closed opesiules present. Vibraculum distal to each zooecium. Vicarious vibracula
absent. Basal surface grooved and tuberculate, not divided into sectors. Central
zooecia closed.

14 Discoporella reussiana (Manzoni)
(PI. 3, fig. i, Text-fig. 2f)

Cupularia reussiana Manzoni, 1869 : 27, pi. 2, figs. 19, ig1 (as 18, iS1 in text), Tertiary, Italy.

Waters, 1878 : 16, Pliocene, Italy. Canu & Bassler, 1923 : 78, pi. i, figs. 19-22, Pliocene,

Italy.
Cupuladria reussiana (Manzoni) Annoscia, 1963 : 226, pi. 9, fig. 2, pi. 10, fig. 2, pi. 13, fig. i,

pi. 14, figs, i a, ib, Quaternary, Italy.

MATERIAL. " Calypso " Coll. Stn. 45*, o° 25' N., 9° o' E., 8.vi.56, 73 m., C48U
(C), with Cleidochasma porcdlanum (Busk), C55L.

D. reussiana is not represented in the Marche-Marchad nor Achimota Collections.
Zoarium domed, sometimes higher than wide (average diameter 6 mm.). Zooecia

wide, proximal part of vestibular arch strongly curved and flanged. Cryptocyst
descending gently at first, coarsely tuberculate. Horizontal cryptocyst formed by
fusion of wide, irregularly shaped denticles, with 2-4 pores. Distal pair of denticles
forming a bar, with a wide tooth protruding distally into the opesia, forming two
lateral opesiular indentations. Basal surface grooved with large tubercles, which
obscure the slight threads marking the grooves in peripheral zooecia.

DIMENSIONS. Lz 0-48-0-60 mm., Iz 0-30-0-42 mm., La 0-36 mm., Lvo 0-14-0-17
mm.

220 I'. L. COOK

Norman (1909 : 290) included C. reussiana Manzoni in his synonymy of C. johnsoni,
and Waters (1878 : 16) placed C. doma Smitt (sic) in the synonymy of C. reussiana.
Neviani (1895 : 102) included C. doma in the distribution of C. reussiana, giving the
localities of Recent specimens as Algeria and Florida.

There is no doubt that the " Calypso " specimens are referable to Manzoni's
species, which has hitherto been described only from European Tertiary and Quater-
nary deposits. Manzoni's fig. 19 shows the large cryptocystal pores and the toothed
distal bar which is not present in the majority of the zooecia figured by Canu & Bassler.

D. reussiana resembles C. doma and C. owenii in the strongly flanged vestibular
arch, but the horizontal cryptocyst lamina is similar to that of D. ocellata, (see
below) and forms a link in the series of species of which the end term is D. umbellata.

The specimens of C. canariensis, C. owenii, D. reussiana and D. ocellata from
" Calypso " Stn. 45, in the Bay of Biafra, are all dark grey in colour, quite unlike
those of the other west African localities, which are light brown. Those of D. reussiana
have no chitinous parts and some are worn, but, as polypides are present in the zooecia
of the majority of zoaria of the other 3 species, it is improbable that the specimens of
D. reussiana are displaced fossils, and therefore it must be presumed that the species
has existed continuously from Tertiary times.

15 Discoporella ocellata n.sp.

(PI. 3, fig. 2, Text-fig. 2g)

MATERIAL. HOLOTYPE, Museum d'Histoire naturelle, Paris, C48C pt.

" Calypso " Coll. I, Stn. 45*, o° 25' N., 9° o' E., 8.vi.s6, 73 m., C48C, C55K
Lagaaij Coll., Nigeria D. 220.

D. ocellata is not represented in the Marche-Marchad nor Achimota Collections.

Zoarium fairly large (average diameter 8 mm.). Zooecia with lateral cryptocyst
descending moderately at first, then salient, forming a narrow shelf. Horizontal
cryptocyst lamina formed by fusion of wide, irregular denticles originating on the
frontal side of the shelf. Vestibular arch with two curved flanges uniting centrally
with each other below the orifice, and with the fused distal lateral denticles, forming a
sinuate opesia with two closed lateral opesiules. Operculum without a basal sclerite.
Basal surface grooved, grooves marked by a thread, not obscured by the irregular
tubercles.

DIMENSIONS. Lz 0-55-0-63 mm., Iz 0-36-0-42 mm., Lvo 0-10-0-14 mm., Ls 0-70-
2-00 mm., Lo 0-10-0-12 mm., lo 0-12-0-15 mm.

The zoaria are more flattened than those of D. reussiana. In worn specimens of
D. ocellata the proximal bar between the opesia and opesiules may be broken, pro-
ducing a trifoliate opesia very similar to that of D. reussiana. The slope of the
cryptocyst and position of the origin of the cryptocystal denticles are then the only
features distinguishing the species on the frontal surface. In D. ocellata the crypto-
cyst descends moderately at first, then becoming salient, forming a shelf, which may
be seen distally and below the horizontal denticles which originate above the angle
produced by the change of slope (see Text-fig. 2g). In D. reussiana the descent of the

POLYZOA FROM WEST AFRICA 221

cryptocyst is at first gentle and then, after the origin of the horizontal denticles, steep,
so that no shelf is visible below them (see Text-fig. 21).

There is a superficial resemblance between D. ocellata and the photographs of
Cupularia bioculata Canu (1904 : 10, pi. 2, figs. 21, 22, Miocene, Patagonia). C.
bioculata is shown with a sinuate opesia and a pair of opesiules. None of the zooecia
has a distal vibraculum, and the occasional large individuals may be broken vicarious
vibracula. The absence of small vibracula excludes it from the Cupuladriidae, and it
is perhaps referable to Selenaria.

D. ocellata is related to both D. reussiana and D. umbellata and forms an important
link between those species. It differs from D. reussiana in its closed opesiules and
basal surface, and from D. umbellata in its sinuate opesia, basal surface, and oper-
culum, which has no proximal sclerite.

16 Discoporella umbellata (Defrance)
(PI. 3, fig. 3, Text-fig. 2h)

Lunulites umbellata Defrance, 1823 : 361, pi. 47, figs. la, ib, Miocene, France.

Cupularia lowei Busk, 1854 : 99, pi. 116, figs. 1-6.

Discoporella umbellata (Defrance) : Lagaaij, 1953 : 16, pi. i, fig. 3, Miocene, Netherlands. Cook,

1965: i?7. Pi- i. fig- 7. Pi- 3, figs- i, 3, 5, 6, text-fig. 4.

MATERIAL. " Calypso " Coll. I, Stn. 7*, 9° 40' N., 13° 53' 5" W., 17^.56, 18 m.,
C4G. Stn. 17, 5° N., 5° 28' 30" W., 21 .v.56, 27 m., C$6G (C). Stn. 18, 5° 2' 5" N.,
5° 24' 4" W., 21 .v.56, 20-25 m., C$V. Stn. 19, 5° 2' 30" N., 5° 24' 40" W., 21. v.56,
21-27 m., C57l. Stn. 29*, 4° 3' N., 6° 12' E., 26. v.56, 32 m., C4gA. Stn. 45*,
o° 25' N., 9° o' E., 8.vi.56, 73 m., C48A (with Labioporella dipla Marcus and Cleido-
chasma porcellanum (Busk)), C55H. Stn. 63, o° 20' N., 6° 47' E., I7.vi.56, 54-40 m.,
C28A. Stn. 77, o° 25' 40" N., 6° 40' 10" E., 2i.vi.56, 50 m., C45A. Stn. 90*,
i° 36' 55" N., 7° 22' E., 26.vi.56, 30 m., C37B. Stn. Pi4*, Dans 1'axe de la Baie,
Principe, 29.vi.56, 15 m., C5gA.

Coll. II. Stn. 73, C72A. Stn. 88, 16° 15' N., 22° 56' 5" W., 26.xi.59, 54 m.,
C8oA.

Marche-Marchad Coll. I. Konakrey, Guinee Ise, iG, 2F. Flor de la bouteille,
Guinee Ise, 21.1.53, 8 m., 3C. Sud de Goree, 27.x. 53, 38-42 m., yC. S.W.
Madeleines, i5.ix.53, 48 m., 2oC ; and 21.1.54, 46-48 m., 46F (C). Presque Tile du
Cap Vert, i8.ii.54, 95 m., 33? (C). Baie de Seminoles, 8.xii.53, 38 m., 390 (C).

Coll. II, as above, 36C. Alignment Goree-Cap Manuel, lantern de Madeleines,
I9-X.56, 35-42 m., 96 (C), 20-25 miles au large de Saloum, 8.iii-55, 35-37 m., 26A
(C).

Coll. III. Au sud de presque Tile du Cap Vert, i8.ii.54, 46-50 m., iD. Either
S.W. Madeleines, i5.ix.53, 48 m., or Sud de Goree, 34-37 m., i3.xi.53, i6F.
Dragage 4, " Gerard Freca ", 97-98 m., 28H ; Dr. 5, i8.ii.54, J5C, 270. 33-34 m.,
27.xi.53, 2gD. Museum Guinee, parages de Flos de Los, Konakrey, 21.1.53, 9 m.,
306. No information, 24!!.

Achimota Coll. I, Stn. 47*, Dredge Haul No. i, 4.1.51, 44 m., I4d. Stn. 48*,

ZOOL. 13, 6 12

222 P. L. COOK

Dredge Haul No. 2, as above, 22C. Stn. in*, Agassiz Trawl No. 2, 4.iv.5i, 43 m.,
49?. Stn. 132,* as above, 2.v.5i, 44 m., 42E and 506 (C). Stn. 133*, Agassiz
Trawl No. 3, as above, 51 m., 45!!. Coll. II, Stn. 133*, see above, 7A. Stn. in*,
see above, I2A.

Zoarium frequently very large (maximum diameter measured, 30 mm.). Zooecia
wide, opesia small, nearly straight proximally, with a pair of small denticles. Oper-
culum with a stout proximal sclerite. Lateral cryptocyst descending gently, tuber-
culate, horizontal cryptocyst a complete lamina, with 8-14 small peripheral opesiules
and small scattered central pores, which are frequently occluded. Vibracular opesia
large, proximal edge denticulate. Basal surface with short grooves, irregular pits
and tubercles. Tentacles 13-16 (see p. 203).

DIMENSIONS. Lz 0-60-0-70 mm., Iz 0-45-0-65 mm., La 0-40-0-43 mm., Lvo
0-20-0-25 mm., Ls 0-90-1-65 mm., Lo 0-10-0-12 mm., lo 0-12-0-15 mm., Lop 0-13-
0-15 mm., lop 0-15-0-20 mm.

C. lowei Busk differs from D. umbellata only in its deformed zoaria (see p. 195 and
cf. C. multispinata, p. 212). The majority of the specimens from the Marche-
Marchad Collections II and III are deformed, most of those from Collection I, the
" Calypso " Collections, and the Achimota Collection, are not.

The tubules which extend from the basal wall of the cystid cavity to the basal
surface of the zoarium have been described by Marcus & Marcus (1962 : 295, pi. 2,
fig. 5). In decalcified specimens these appear as minute strands, and were first
described by Waters (1921 : 412, pi. 30, figs. 3, 5, 6). Their presence in D. umbellata
(subsp. depressa) was noted by Hastings (1930 : 719), and they are clearly seen in
specimens from Rio de Janeiro (British Antarctic Expedition Coll., Stn. 42). The
pore in the basal surface described by Waters is present in specimens of D. umbellata
from Senegal (Marche-Marchad Coll. I 39D). It is present in all peripheral zooecia
and remains open in the majority of zooecia of young colonies. In older colonies it
becomes obscured by the increasing development of tuberculate secondary cal-
cification. A short groove may be seen in peripheral zooecia running proximally and
distally from the pore. As stated by Waters (who described them as muscles), the
tubules appear to be inserted into the basal wall along or near the groove. Decalcified
specimens from Rio de Janeiro show a small granular mass which corresponds in
position to the basal pore.

Calvet (1907 : 393) recorded D. umbellata from Madeira (80 m.) and a specimen
from the Cape Verde Islands (1900 m.) which was described as "a la forme de
denticulata de Conrad ". In view of the depth given, it seems unlikely to be C. owenii
(see p. 213), which does not extend to great depths, and it may have been a colony of
D. umbellata, in which the cryptocyst was worn away or damaged.

Specimens of D. umbellata from west Africa are consistent in character, and al-
though larger, agree with the European fossil material. The wide zooecia, the nearly
straight proximal edge of the opesia, with its two denticles, and the denticulate edge
of the proximal vibracula opesia, are all features not found in the fossil and Recent
specimens from the western Atlantic and eastern Pacific which have been separated

POLYZOA FROM WEST AFRICA 223

as D. umbellata subsp. depressa (Conrad) (see Cook, 1965 : 180). The development of
the horizontal cryptocyst lamina in young peripheral zooecia, however, shows the
close relationship of D. umbellata to both the subspecies depressa and to the denticu-
late species of Cupuladria.

The distribution of D. umbellata in these collections extends from Senegal to the
Bay of Biafra, and includes the Cape Verde Islands. Recent specimens have also
been recorded from Madeira and the Canaries. The recorded bathymetrical range is
from 8-130 m.

Dr. R. Lagaaij (in litt] 20.X.64 has drawn my attention to the two distinct species
figured as Lunulites rhomboidalis by Miinster (in Goldfuss, 1829 : 105, pi. 37, figs. 7a-c),
from the German Tertiary deposits. Figure 7a greatly resembles a young colony of
Discoporella umbellata, although no opesiules, and very few vibracula, are figured.
Figures 7b and c are the frontal and basal views respectively of a regenerated fragment
of a species of Cupuladria. Dr. Lagaaij remarks " Of the two localities mentioned
on p. 106, that of Kassel is now considered the type locality of the Chattian (Upper
Oligocene) " (see Drooger, 1964 : 372), " this would not fit with the presence of
D. umbellata which first appears in the Aquitanian (Lower Miocene) ". In this
connection, it should be noted that neither Philippi (1844 : 3) nor Reuss (1865 : 685),
who both listed L. rhomboidalis, had found specimens in their material.

17 ACKNOWLEDGMENTS

I should like to thank Dr. A. Andersson (Naturhistoriska Riksmuseet, Stockholm),
Dr. R. Boardman (Smithsonian Institution, Washington), Prof. Dr. E. Voigt (Geo-
logisches Staatsinstitut, Hamburg), and Dr. H. G. Richards (Academy of Natural
Sciences, Philadelphia), for the loan and presentation of specimens and photographs.
My thanks are also due to Dr. A. Cheetham (Louisiana State University) and Dr. R.
Lagaaij (Shell Exploratie en Produktie Laboratorium, the Netherlands), both for
stimulating discussion and criticism, and for the loan of specimens. The photo-
graphs were taken at the British Museum by Mr. J. V. Brown and Mr. P. Green.
Finally, I am deeply indebted to Dr. A. B. Hastings and Dr. J. P. Harding (British
Museum, Natural History) for their continual interest and encouragement.

18 SUMMARY

Nine species belonging to the family Cupuladriidae are described, eight of which
occur in west African waters.

Cupuladria canariensis is defined and distinguished from C. biporosa ; C. mono-
trema is also described, and the occurrence of vicarious vibracula in the three species is
discussed.

Three species with denticulate cryptocysts, C. multispinata, C. owenii and C. doma,
are defined and distinguished, and a new subspecies, C. owenii disciformis, characteris-
tic of very shallow waters in the Bay of Biafra, is described.

The genus Discoporella is defined, and three species are described, in which the
development of the horizontal cryptocyst lamina is increasingly complex. D.
reussiana, hitherto recorded as a fossil only, shows similarities with both the denti-

224

I'. L. COOK

culate species of Cupuladria and with D. umbellata. D. ocellata, a new species, links
the zooecial characters of D. reussiana and D. nmbellata.

FIG. 4. Distribution of Recent Cupuladriidade. i. Cupuladvia pyriformis . 2. C. canarien-
sis. 3. C. biporosa. 4. C. monotrema. 5. C. guineensis, A = A form, B — B form.
6. C. indica. 7. C. multispinata. 8. C. owenii. 8A. C. owenii disciformis. g. C. doma.
10. Discoporella reussiana. n. D. ocellata. 12. D. umbellata. I2A. D. umbellata,
peyroti-type zoaria. 126. D. umbellata depressa. (For notes on species i, 5, 6, I2A and B,
see Cook, 1965).

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226 P. L. COOK

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POLYZOA FROM WEST AFRICA 227

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Hist., (9) 18 : 424-433.

PLATE i
Cupuladria, zoaria treated with eau de javelle

FIG. i. C. canariensis (Busk) Barbados, 1962. i. 26. 2. A. Frontal view. B. Basal view.

x 7.4.
FIG. 2. C. biporosa Canu & Bassler. Barbados, 1962. i. 26. i. A. Frontal view, showing

zooecia with wide lateral cryptocysts, and large vicarious vibracula. B. Basal view.

x 7.4.
FIG. 3. C. biporosa. Yucatan, Naturhistoriska Riksmuseet, No. FBD 696. A. Frontal view.

B. Basal view, showing basal kenozooecia covering the original substrate, cf . C. canarien-
sis, fig. iB. x 7.4.
FIG. 4. C. biporosa. West Indies, Naturhistoriska Riksmuseet, No. FBD 548, a Recent

specimen, cf. figs. 6A, B, of a Miocene colony. A. Frontal view. B. Basal view.

x 7.4.
FIG. 5. C. biporosa. Jamaica, Miocene, Bowden marl, United States National Museum, No.

80747 pt. Colony with aberrant doma-type growth, basal view, showing rows of

peripheral vibracula. x 7.4.
FIG. 6. C. biporosa. Jamaica, Miocene, Bowden marl, United States National Museum, No.

80747 pt-» cf- n£s- 4-A, B. A. Frontal view. B. Basal view. X 8.1.
FIG. 7. C. doma (d'Orbigny). Senegal, Marche-Marchad Coll., I, 33 B. Lateral view, showing

closed peripheral zooecia. x 12.

Bull. 13. M. (N.H.) Zwl. 13, 6

PLATE i

PLATE 2
Cupuladria, zoaria treated with eau do javelle

FIG. i. C. owenii disciformis n. subsp. Bay of Biafra, "Calypso" Coll., C^jA.. A. Frontal
view, showing closed central zooecia, and reduced cryptocystal denticles of other
zooecia. B. Basal view, showing fine tubercles. x 9.

FIG. 2. C. multispinata (Canu & Bassler). Senegal, Marche-Marchad Coll., I, 33 A. A.
Frontal view, showing numerous cryptocystal denticles. B. Basal view, showing
large tubercles and radial " threads ". x 7.5.

FIG. 3. C. owenii (Gray). Senegal, " Calypso " Coll., I, C4.H. A. Frontal view, showing closed
central zooecia, and well-developed vestibular arches of other zooecia. B. Basal view,
showing both smooth and finely tuberculate forms of basal calcification. x 10.

N.B. Figures 2 and 3 are not reproduced at the same magnification, so that the marked difference

in size between the zooecia of C. multispinata and C, owenii is not immediately apparent.

Hull. n.M. (N.H.) Zoo/. 13, 6

PLATE 2

PLATE 3
Cupuladria and Discoporella, zooecia and vibracula treated with eau de javelle.

FIG. i. D. reussiana (Manzoni). Bay of Biafra, " Calypso " Coll., I, C48D. Zooecia showing

toothed cryptocystal bar and opesiular indentations. x 49.
FIG. 2. D. ocellata n. sp. Bay of Biafra, " Calypso " Coll., C48C. Zooecia showing sinuate

opesia and paired distal opesiules. x 53.
FIG. 3. D. umbellata, (Defrance). Ghana, Achimota Coll., II, yA. Showing the slit indicating

the former presence of an Acrothoracid Cirripede. x 32.
FIG. 4. C. canariensis (Busk). Senegal, Marche-Marchad Coll., II, 3 A. Showing position of

small, hooked vibracula thus : ***. x 24.

Bull. B.M. (N.H.) ZooL 13, 6

PLATE 3

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DESCRIPTIONS OF SOME

STRONGYLES (NEMATODA) FROM

MAMMALS IN EAST NEPAL: WITH

RECORDS OF OTHER PARASITIC

NEMATODES

WILLIAM G. INGLIS & COLIN G. OGDEN

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 7

LONDON: 1965

4 HOY \(

DESCRIPTIONS OF SOME STRONGYLES

(NEMATODA) FROM MAMMALS IN EAST

NEPAL: WITH RECORDS OF OTHER

PARASITIC NEMATODES

WILLIAM G. INGLIS

AND

COLIN G. OGDEN

Pp. 229-245 ; 50 Text-figures

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. 7

LONDON: 1965

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
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Parts will appear at irregular intervals as they become
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within one calendar year.

In 1965 a separate supplementary series of longer
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This paper is Vol. 13, No. 7 of the Zoological series.
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Issued October, 1965 Price Eight Shillings

DESCRIPTIONS OF SOME STRONGYLES

(NEMATODA) FROM MAMMALS IN EAST

NEPAL: WITH RECORDS OF OTHER

PARASITIC NEMATODES

By WILLIAM G. INGLIS & COLIN G. OGDEN

INTRODUCTION

ALTHOUGH the main zoological interest of the British Museum (Natural History)
East Nepal Expedition : 1961-62 was in soil inhabiting invertebrates (Sheals &
Inglis, 1965) the opportunity was taken, when possible, to collect helminth parasites.
Among the nematodes collected are several new or rare species of Strongylina, which
are described below, together with a list of the other species which were collected.
Two new, non-strongyle species have been described elsewhere (Inglis £ Ogden, 1965,
1965*).

List of All Species Collected

All host identifications were carried out by the staff of the British Museum (Natural
History).

Felis bengalensis horsfieldi Gray, 1842. <$. Hatia (27° 44' N., 87° 21' E.), Arun
River, East Nepal. (22.xii.i96i ; 7,000 feet alt.).

Toxocam mystax (Zeder, 1800) ex small intestine.
Mastophorus muris (Gmelin, 1790) ex stomach.
Molineus (?) patens (Dujardin, 1845) ex stomach.
Molineus springsmithi sp. nov. ex small intestine.
Arthrostoma felineum Cameron, 1927 ex small intestine.
Arthrostoma tunkanati sp. nov. ex small intestine.

Callosciurus Pygerythrus lokroides Hodgson, 1836. <$. Popti La (27° 47' N., 87° 21'
E.), Arun River, East Nepal. (22.xii.i96i ; 9,600 feet alt.)

Mastophorus sp. (°-$ only) ex small intestine.

Rictularia dhanra Inglis & Ogden, 1965 ex intestine.

Brevistriata (?) skrjabini (Schulz & Lubimov, 1932) ex small intestine.

Longistriata gold sp. nov. ex small intestine.

CaUosciurus macclellandi macclellandi Horsfield, 1839. ?• Hatia (27° 44' N.,
87° 21' E.), Arun River, East Nepal. (i7.xii.i96i ; 7,500 feet alt.)

Citellina himalensis Inglis & Ogden, 1965 ex small intestine.
Brevistriata (?) skrjabini (Schulz & Lubimov, 1932) ex small intestine.
Longistriata (?) gola ($9, only) ex small intestine.

ZOOL. 13, 7 I3

232 W. G. INGLIS & C. G. OGDEN

Suncus murinus caerulescens Shaw, 1800. $. Sanghu (27° 21' N., 87° 33' E.),
Maewa Khola, East Nepal. (25. xi. 1961 ; 6,500 feet alt.)

Capillaria minuta Chen, 1937 ex stomach.
Rattus rattus sikkimensis Hinton, 1919. juv. <$. Sanghu (as above). (17. x. 1961).

Nippostrongylus bmsiliensis (Travassos, 1914) ex small intestine.
Rattus r. sikkimensis <$. At same locality (6.xi.i96i).

Nippostrongylus brasiliensis ex small intestine.
Rattus r. sikkimensis <$. At same locality.

Heterakis spumosa (Schneider, 1866) ex caecum.
Rattus r. sikkimensis $. At same locality (21. x. 1961).

Nippostrongylus brasiliensis ex small intestine.

Mastophorus muris ex stomach.

Heterakis spumosa ex intestine and caecum.

Rattus r. sikkimensis unsexed, at same locality (6.1.1962).
Heterakis spumosa ex caecum.

In addition two Suncus m. caerulescens and six Rattus r. sikkimensis, all caught
at Sanghu between 7.xi.i96i and 5.^.1962, were searched without finding any
parasites.

Descriptive Section
Molineus (?) patens (Dujardin, 1845)

MATERIAL STUDIED. 7 <$, 3 $ ex stomach Felis bengalensis horsfieldi. Hatia,
Arun River, East Nepal. B.M. (N.H.), Reg. Nos. 1965. 1-5.

MEASUREMENTS (in mm.). Males: Body length: 2-56; 2-64; 2'68 ; 278;
2-87 ; 2-91 ; 3-34. Body breadth : 0-053 ; 0-053 ; 0-055 ', 0-050 ; 0-051 ; 0-055 ;
0-054. Diameter of head : 0-022 ; 0-020 ; 0-017 ; 0-022 ; 0-022 ; 0-019 ; 0-023.
Diameter of cephalic vesicle : 0-038 ; 0-030 ; 0-030 ; 0-034 > 0<035 '> O'OS1 > 0-036.
Length of cephalic vesicle : 0-060; 0-053 ; 0-054 > 0-047 ; 0-051 ; 0-056 ; 0-054.
Distance of cervical groove from anterior end of body : 0-153; 0-177 > 0tI59 > °'I5& ',
0-188 ; 0-183 ; 0-177. Oesophagus length : 0-348 ; 0-347 > °'352 > 0-366 ; 0-358 ;
0-298 ; 0-398. Length of spicules : 0-099 J 0-107 > 0-089 ; o-ioo ; 0-090 ; 0-103 ;
0-105. Length of gubernaculum : 0-051 ; 0-058 ; 0-047 J 0-060 ; 0-046 ; 0-059 I
0-064.

Females : Body length : 3-03 ; 3-55 ; 3-62. Body breadth : 0-063 '> 0-050 ;
0-072. Diameter of head : 0-023 '> 0-020 ; 0-022. Diameter of cephalic vesicle :
0-040 ; 0-030 ; 0-043. Length of cephalic vesicle : 0-056 ; 0-044 '> 0-060. Distance
of cervical groove from anterior end of body : 0-160 ; 0-154 i 0-201. Oesophagus
length : 0-378 ; 0-316 ; 0-420. Length of tail : 0-064 ', 0-056 ; 0-068. Distance of
vulva from posterior end of body : 0-588 ; 0-568 ; 0-622. Size of eggs : 0-054 x
0-027 to 0-075 x 0-031.

STRONGYLES FROM XEPALESE MAMMALS

2.33

FIGS. 1-13. Mo/wews (?) ^afews. Fig. i. Dorsal view of head showing the cephalic
vesicle. Fig. 2. Ventral view of male bursa. Fig. 3. Detail of dorsal ray. Fig. 4.
Spicule from the left. Fig. 5. Spicule from the right. Fig. 6. Dorso-lateral view of
spicule. Fig. 7. Lateral view of gubernaculum. Fig. 8. Terminal region of female
tail (lateral). Fig. 9. Terminal view of female tail (ventral). Fig. 10. Lateral view
of vulvar region. Fig. n. Lateral view of bursa from right-hand side (teratological) .
Fig. 12. Ventral view of bursa (teratological) . Fig. 13. Lateral view of bursa from left-
hand side (teratological).

234 W. G. IXGLIS & C. G. OGDEN

The head bears the typical cephalic vesicle (Text-fig, i).

The cuticle is marked by ten to twelve longitudinal ridges and the cervical groove is
complete round the body although it is rather faint on the dorsal surface. No
cervical papillae have been seen.

Male. The bursa is typical with the ventro-lateral and latero-ventral rays arising
from a common base and the postero- and medio-laterals arising from a common base.
The antero-lateral (or externo-lateral) ray does not reach the edge of the bursa. The
externo-dorsal ray is given off some distance along the dorsal ray, and the latter ray
bifurcates terminally twice with the internal small ray further divided just before
the edge of the bursa (Text-figs. 2, 3 and 13).

The gubernaculum is fairly simple with a slight kink in its lateral outline (Text-
figs. 7, ii and 13) while the complex spicules are relatively slim and terminate
posteriorly in three needle-like points, of which the median, and the longest, splits the
spicule for about half its length (Text-figs. 4, 5 and 6). A small genital cone is
present which carries two small papilla-like structures on its posterior edge (Text-
figs. 2 and 12).

One teratological male is present in which there is an additional ray on the right-
hand side of the bursa (Text-figs. 11 and 12) which arises between the typical externo-
lateral and the pair of postero- and medio-laterals, and reaches the edge of the
bursa (Text-fig, n). The rays on the left-hand side are typical (Text-fig. 13).

Female. The vulva, uterus and eggs are typical of the genus (Text-fig. 10) while
the tail carries a pair of ventro-lateral swellings just anterior to the fine posterior
terminal process (Text-figs. 8 and 9).

DISCUSSION. The specimens match the description of Molineus patens (Dujardin,
1845) Petrov, 1928 although they are smaller than any previously reported. Ad-
ditional points of difference appear to be the lack of any small hooks or cuticular
bumps on the inner surface of the bursa and the slightly elaborate genital cone.
Comparison of the specimens described above with the descriptions of M. patens
given by Petrov (1928), Zunker (1929), Leiper (1936), Travassos (1937) and Skrjabin
et al. (1954) shows that the body is only half, the spicules two-thirds and the guber-
naculum half the lengths of the corresponding dimensions for the smallest specimens
previously recorded. Leiper (1936) observed that there appeared to be a tendency
towards a host : parasite size relationship for specimens of M. patens recovered from
the stoat and weasel in England, but Table 2 in Petrov (1928) shows that there are
only slight variations between the measurements of specimens from five different
hosts examined in the U.S.S.R.

As the relative proportions of the specimens we described above are in general
agreement with those already reported and the structure of the spicules appears to
be the same, we treat our specimens as M. patens, although with some reservations.

Molineus springsmithi sp. nov.

MATERIAL STUDIED. 6 £, 6 $ ex small intestine of Felis bengalensis horsfieldi.
Hatia, Arun River, East Nepal. B.M. (N.H.), Reg. Nos. 1965. 6-25.

STRONGYLES FROM NEPALESE MAMMALS

235

FIGS. 14-26. Molinens springsmithi sp. nov. Fig. 14. Lateral view of male bursa.
Fig. 15. Lateral view of spicule and gubernaculum. Fig. 16. Detail of whole spicule.
Fig. 17. Detail of posterior end of spicule. Fig. 18. Lateral view of female tail. Fig.
19. Ventral view of male bursa. Fig. 20. Lateral view of vulvar region of female.
Figs. 21-26. Ventral views of dorsal rays showing variation of major bifurcation.

236 W. G. IXGLIS & C. G. OGDEN

MEASUREMENTS (in mm.). Males. Body length : 2-27 ; 2-52 ; 2-63 ; 2-68 ;
2-69 ; 3-01. Body breadth : 0-054 J 0-067 '> 0-044 '> 0052 ; 0-046 ; 0-072. Dia-
meter of head : 0-021; 0-022; 0-022; 0-022; 0-028; 0-025. Diameter of cephalic
vesicle : 0-029 > 0-040 ', 0-030 ; 0-028 ; 0-036 ; 0-033. Length of cephalic vesicle :
0-047 » 0-042 ; 0-044 '> 0'°45 ', O'OS1 ', 0-044. Oesophagus length : 0-347 » °'353 J
0-307 ; 0-381 ; 0-340 ; 0-397. Distance of cervical groove from anterior end of
body : 0-125 ', 0-147 ; 0-141 ; 0-144 ; 0-139 ; 0-157. Length of spicules : 0-070 ;
0-065 i 0-068 ; 0-068 ; 0-062 ; 0-065. Length of gubernaculum : 0-024 •' 0-024 '>
0-025 i 0-021 ; 0-024 J 0-021.

Females. Body length : 3-15 ; 3-52 ; 3-57 ; 3-79 ; 3-94 ; 3-95. Body breadth :
0-052 ; 0-057 '> 0-052 ; 0-054 > 0-062 ; 0-057. Diameter of head : 0-023 I 0-025 '>
0-024 » 0-024 J 0-025 I 0-024. Diameter of cephalic vesicle : 0-032 ; 0-031 ; 0-032 ;
0-028 ; 0-033 ', 0-030. Length of cephalic vesicle : 0-041 ; 0-057 '> 0-047 ; 0-045 ;
0-046; 0-051. Oesophagus length : 0-298; 0-343; 0-374; 0-360; 0-422; 0-372.
Distance of cervical groove from anterior end of body : 0-120 ; 0-155 i 0>I73 >
0-154 > 0-172 ; 0-172. Distance of vulva from posterior end of body : 0-592 ;
0-565 ; 0-522 ; 0-500 ; 0-532 ; 0-543. Length of tail : 0-068 ; 0-059 >' 0-048 ;
0-051 ; 0-072 ; 0-058. Size of eggs : 0-055 * 0-026 to 0-071 x 0-031.

The cuticle is marked by eleven to fifteen longitudinal ridges and the cervical groove
is distinct completely round the body. No cervical papillae have been seen.

Male. The bursa is typical with the ventro-lateral and latero-ventral and the
postero- and medio-lateral rays arising in pairs from common bases. The first
bifurcation of the dorsal ray is rather deep while the terminal bifurcations, the inner
one double, are somewhat variable (Text-figs. 21-26). The gubernaculum is small
and slightly hooked anteriorly (Text-figs. 14-15) and the spicules are simple with
three terminal processes. The major division of the processes extends only for about
one third the length of the spicules (Text-figs. 15-17). The bursa is not divided into
lobes and there is a small pre-cloacal supplementary membrane which is supported by
an internal Y-shaped thickening (Text-fig. 19).

Female. The reproductive system is typical (Text-fig. 20) and the tail ends in a
small spike (Text -fig. 18).

DISCUSSION. This species is very distinct in the extreme depth of the first bi-
furcation of the dorsal ray, the characteristic shape of the gubernaculum and the
relative simplicity of the spicules.

Brevistrata ? skrjabini (Schulz & Lubimov, 1932)

MATERIAL STUDIED. 2 £, 5 $ ex small intestine of Callosciurus pygerythrus
lokroides. Popti La, Arun River, East Nepal. B.M. (N.H.) 1965. 26-28.

2 <£, 7 $ ex small intestine Callosciurus macclellandi macclellandi. Hatia, Arun
River, East Nepal. B.M. (N.H.) 1965. 29-31.

STRONGYLES FROM NEPALESE MAMMALS 237

MEASUREMENTS (in mm.). In two groups separated by full stop. First group
from Popti La sample. Males. Body length : 3-31 ; 373. 4-35; 5-32. Body
breadth : 0-067 '< 0-063. 0-116 ; 0-116. Diameter of head : 0-020 ; 0-020.
0-028; 0-026. Diameter of cephalic vesicle : 0-030; 0-030. 0-040; 0-036. Length
of cephalic vesicle : 0-042 ; 0-045. 0-046 ; 0-057. Oesophagus length : 0-347 ;
0-327. 0-314 ; 0-388. Length of spicules : 0-450 ; 0-442. 0-570 ; 0-548. Length
of gubernaculum : 0-040 ; 0-044. °'°53 '> 0-060.

Females. Body length : 3-43 ; 3-98 ; 4-13 ; 4-39 ; 4-59. 4-63 ; 5-32 ; 5-33 ;
5'36 ; 5-40 ; 5-45 ; 5-59. Body breadth : 0-065 > 0-065 ; 0-063 '> 0-065 ; 0-072.
0-089 i °'I23 '> 0-109 ; 0-135 ; 0-098 ; 0-098 ; 0-104. Diameter of head : 0-022 ;
0-022 ; 0-023 '> 0-023 i 0-025. 0-025 I 0-029 J 0-028 ; 0-025 '> 0-030 ; 0-030 ; 0-028.
Diameter of cephalic vesicle : 0-026 ; 0-028 ; 0-023 .' 0-029 '> 0'027- 0-033 ', 0-038 ;
0-040 ; 0-032 ; 0-030 ; 0-030 ; 0-042. Length of cephalic vesicle : 0-042 ; 0-048 ;
0-042 ; 0-044 ; 0-044. 0>043 » 0-050 ; 0-048 ; 0-047 '> 0-050 ; 0-048 ; 0-051. Oeso-
phagus length : 0-300 ; 0-353 ', 0-335 ; 0-338 ; 0-322. 0-382 ; 0-436 ; 0-400 ; 0-410 ;
0-388 ; 0-378 ; 0-364. Length of tail : 0-049 '> 0'°37 ', 0-042 ; 0-042 ; 0-044.
0-049 i 0-049 '> °'°54 i 0-040 ; 0-057 i O'OS1 '> 0-056. Distance of vulva from
posterior end of body : 0-085; 0-094; 0-096; 0-093; o-no. 0-098; 0-115; 0-107;
0-079 '> °'112 ; 0-107 ; 0-117. Size of eggs : 0-055 x 0-033 to 0-076 x 0-040.

The worms are coiled into spirals with two or three turns. The head carries a small
cephalic vesicle and the mouth is simple without any obvious cavity between it and
the beginning of the oesophagus. The cuticle bears the typical broken longitudinal
ridges which alternate down the body.

Male. The bursa has a small distinct dorsal lobe (Text-fig. 34). The ventral rays
of the bursa arise together, as do the three lateral rays (Text-figs. 27 and 28). All
these rays reach the edge of the bursa. The lateral rays differ in size with the stout
externo- and medio-laterals lying close together, while the postero-lateral is much
slimmer and is directed posteriorly away from the other two. The thin externo-
dorsal rays arise some distance from the origin of the dorsal ray (Text-fig. 28).
The dorsal ray bifurcates twice towards its posterior end with the inner branch of the
final bifurcation bearing a further relatively small inner branch (Text-fig. 30) .

The spicules are long and filiform with bifurcate posterior ends (Text-figs. 31 and
32). The main terminal branch of the spicules ends in a hook-liked projection while
the other branch is blunt. The gubernaculum is small and roughly square in
outline when viewed from the ventral surface (Text-fig. 29).

Female. The single uterus and ovojector open through the vulva which lies very
near the anus (Text-fig. 33) . The tail is relatively short and conical with the phasmids
near the tip.

DISCUSSION. The three species currently referred to the genus Brevistriata:
B. skrjabini (Schulz & Lubimov, 1932), B. sinensis Li, 1941 and B. callosciuri
Supperer & Kutzer, 1963 ; all occur in Asian squirrels. Supperer & Kutzer (1963)
tabulate what they consider to be the diagnostic characters for these three species.

W. G. INGLIS & C. G. OGDEN

FIGS. 27-34. Brevistriata skrjabini. Fig. 27. Lateral view of male bursa. Fig. 28.
Ventral view of male bursa. Fig. 29. Ventral view of gubernaculum. Fig. 30. Dorsal
ray of bursa. Fig. 31. Ventral view of distal end of spicule from right. Fig. 32.
Ventral view of distal end of spicule from left. Fig. 33. Lateral view of female
tail showing vulva. Fig. 34. Detail of small dorsal lobe of bursa.

STRONGYLES FROM NEPALESE MAMMALS 239

B. sinensis is characterized by extremely long spicules with non-bifurcate posterior
ends and by the externo-dorsal rays arising about half-way along the length of the
dorsal ray. B. callosciuri is unique in the extreme division of the caudal bursa into
two distinct lobes, the spicules are bifurcate distally forming two, unequal pointed
branches and the gubernaculum is complex.

The specimens described above are, however, in good agreement with the original
description of B. skrjabini with the exception of the bifurcation of the spicules.
Schulz & Lubimov (1932) state that the spicules are bifurcate " near the proximal
end ", but these structures are not figured and it would appear that " proximal " is
an error for " distal ". Travassos (1937) and Supperer & Kutzer (1963) state that
the spicules are bifurcate distally although they apparently derive their data from
the original description. The specimens described above agree with the original
description in sufficient detail to warrant their reference, at least provisionally, to
B. skrjabini.

Longistriata gola sp. nov.

MATERIAL STUDIED. 4 ^, 7 $ ex small intestine Callosciurns macclellandi mac-
clellandi. Hatia, Arun River, East Nepal. B.M. (N.H.) Reg. Nos. 1965. 32-35.

MEASUREMENTS (in mm.). Males. Body length : 4-71 ; 4-97 ; 5-12 ; 5-51.
Body breadth : 0-149 '> 0-128 ; 0-129 '> 0<I37- Diameter of head : 0-027 > 0-028 ;
0-028; 0-028. Diameter of cephalic vesicle : 0-042; 0-049; °'°55 '> 0-048. Length
of cephalic vesicle : 0-051 ; 0-062 ; 0-055 '> o-°57- Oesophagus length : 0-351 ;
0-349 '> 0*367 ; 0-380. Length of spicules : 0-664 > 0-692 ; 0-646 ; 0-661. Length
of gubernaculum : 0-028 ; 0-034 » 0-030 ; 0-036.

Females. Body length : 6-52 ; 6-73 ; 7-04 ; 7-21 ; 7-23 ; 7-32 ; 7-48. Body
breadth : 0-120 ; 0-123 i o*1^1 ', O-I39 ', 0-129 » 0-116 ; 0-148. Diameter of head :
0-032 ; 0-031 ; 0-030 ; 0-030 ; 0-031 ; 0-028 ; 0-039. Diameter of cephalic vesicle :
0-051 ; 0-050 ; 0-046 ; 0-044 » 0'045 '> 0-049 i °'°57- Length of cephalic vesicle :
0-058 ; 0-054 I 0'055 ', 0'°54 ', 0-056 ; 0-060 ; 0-058. Oesophagus length : 0-400 ;
0-425 ; 0-383 ; 0-373 ; 0-426 ; 0-443 ; 0-338. Length of tail : 0-048 ; 0-058 ;
0-050; 0-036; 0-052; 0-054; °'°57- Distance of vulva from posterior end of body :
o-i2i ; 0-135 ; 0-171 ; 0-157 ; 0-168 ; 0-118 ; 0-108. Size of eggs : 0-072 X 0-044
to 0-054 X 0-032.

The body is coiled four or five times in a tight spiral. The head has the characteris-
tic cephalic vesicle. The cuticle bears typical longitudinal ridges, twenty just
posterior to the cephalic vesicle increasing to twenty-six at the mid-body region and
diminishing in number posteriorly.

Male. The bursa is symmetrical and bilobed (Text-fig. 39). The ventral rays
arise together from a common base as do the three lateral rays (Text-figs. 38 and 39),
and all reach the edge of the bursa. The externo-dorsal rays arise close to the root of
the dorsal ray but do not reach the edge of the bursa. The dorsal ray bifurcates
at the extreme distal end to give two short branches which again bifurcate (Text-fig.
39). The genital cone is prominent and bears a pair of processes (Text-fig. 40)

240

\V. G. IXGLIS & C. G. OGDKX

FIGS. 35-41. Longistriata gola sp. nov. Fig. 35. Anterior end of female. Fig. 36.
Spicules showing sheath. Fig. 37. Ventral view of gubernaculum. Fig. 38. Lateral
view of male bursa. Fig. 39. Ventral view of male bursa. Fig. 40. Lateral view of
genital cone showing process. Fig. 41. Lateral view of female tail showing vulvar and
cuticular modification.

STRONGYLES FROM NEPALESE MAMMALS 241

slightly ventral to the genital opening. The spicules are equal, long, thin, being
finely pointed distally and enclosed in a sheath for most of their length (Text-fig. 36) .
The gubernaculum is small with the outer edges folded inwards to form a groove
(Text-fig. 37).

Female. The tail is typical of the genus (Text-fig. 41), with the vulva situated just
anterior to the anus. The extreme tip of the tail bears two small phasmids. The
cuticle at the posterior end appears to be very loose and in some specimens completely
surrounds the posterior end of the body, Mawson (1961) figures this in some detail.

DISCUSSION. Keys to the species of the genus Longistriata have been published by
Dickmans (1935), Skrjabin, Shikhobalova & Schultz (1954) and Mawson (1961)
in which the main criteria for specific separation are cuticular specialization (i.e.
presence or absence of lateral alae) and the structure of the male caudal apparatus
(symmetry of bursa and shape of the dorsal ray). Using these criteria more than
half of the fifty-odd described species have no lateral alae and possess a symmetrical
bursa as in L. gola. However, of these only five are similar to L. gola in the shape of
the dorsal ray of the bursa, thus : L. bathyergi, L. beta, L. leporis, L. schidzi and
L. seurati.

L. bathyergi Ortlepp, 1939 is similar to L. gola in overall size and the length of the
spicules, but differs in having pre-bursal papillae, an indefinite gubernaculum, in the
thickness of the dorsal rays and in the thickening at the tips of the spicules. L. beta
(Travassos, 1918) differs in being only half the size of L. gola, in having smaller
spicules and in the shape of the gubernaculum. L. leporis Schulz, 1931, and L.
schulzi Schachnasarova, 1949 also have different spicules, those of the former being
longer and distally bifurcate, whilst those of the latter are almost three times as long
as those of L. gola. L. seurati Travassos & Darriba, 1929 differs in having only
fourteen longitudinal ridges on the cuticle, in the spicules being united distally and in
having an asymmetrical gubernaculum.

Arthrostoma tunkanati sp. nov.

MATERIAL STUDIED. 4 $, 7 $ ex small intestine. Felis bengalensis horsfieldi. Hatia,
Arun River, East Nepal. B.M. (N.H.) Reg. Nos. 1965. 36-40.

MEASUREMENTS (in mm.). Males. Body length : 2-37 ; 3-23 ; 3-62 ; 3-74.
Body breadth : 0-182 ; 0-145 ; 0-142 ; 0-163. Length of buccal cavity : 0-065 ;
0-074 > 0-064 > 0-080. Diameter of buccal cavity : 0-040 ; 0-061 ; 0-056 ; 0-053.
Distance of excretory pore from anterior end of body : 0-327 ; 0-325 ; 0-293 ; 0-328.
Oesophagus length : 0-438 ; 0-471 ; 0-483 ; 0-491. Breadth of oesophagus (maxi-
mum) : 0-094 ; 0-068 ; 0-070 ; 0-095. Length of spicules : 0-270 ; 0-410 ; 0-383 ;
0-376. Length of gubernaculum : 0-054 '> 0>045 '> 0-040 ; 0-048.

Females. Body length : 2-46 ; 2-56 ; 2-90 ; 2-94 ; 3-85 ; 4-27 ; 4-28. Body
breadth : 0-216 ; 0-205 ; 0-215 ; 0-211 ; 0-176 ; 0-192 ; 0-151. Length of buccal
cavity : 0-063 ; 0-063 ; 0-070 ; 0-066 ; 0-086 ; 0-088 ; 0-074. Diameter of buccal
cavity: 0-046; 0-045; 0-051; 0-045; 0-059; 0-056 ; 0-056. Distance of excretory
pore from anterior end of body : 0-306 ; 0-276 ; 0-288 ; 0-308 ; - - ; 0-360 ; — -.

\V. G. 1XGL1S & C. G. OGDEN

FIGS. 42-48. Arthrostoma tunkanati sp. nov. Fig. 42. Lateral view of head, additional
plate marked X. Fig. 43. Dorsal ray. Fig. 44. Lateral view of male bursa. Fig. 45.
Ventral view of male bursa. Fig. 46. Lateral view of vulvar region of female showing
papilla and cuticular flap. Fig. 47. Ventral view of gubernaculum. Fig. 48. Lateral
view of gubernaculum and tips of spicules. Fig. 49-50. Arthrostoma feline um. Fig.
49. Lateral view of gubernaculum and tips of spicules. Fig. 50. Ventral view of
gubernaculum.

STRON GYLES FROM NEPALESE MAMMALS 243

Oesophagus length : 0-420 ; 0-503 ; 0-457 ; 0-477 > °'595 '> 0-567 ', 0'55°. Breadth
of oesophagus (maximum) : 0-094 ; 0-087 > 0-097 ; 0-113 .' 0-102 ; 0-166 ; 0-091.
Length of tail : 0-080 ; 0-086 ; 0-081 ; 0-098 ; 0-088 ; 0-083 ', 0-087. Distance of
vulva from posterior end of body : 0-83 ; 0-81 ; 0-94 ; i-oi ; 1-06 ; 1-15 ; 1-14.
Size of eggs : 0-059 x 0-026 to 0-083 X 0-043.

Short narrow worms with the anterior end of the body bent dorsally. The
lining of the buccal cavity is in the form of plates arranged in the same way as
Arthrostoma felineum Cameron, 1927 but differing in the presence of an additional
lateral plate (marked X in Text-fig. 42). That is, there are ten plates of which one
forms a cone round the posterior of the buccal cavity, one triangular plate is ventral
in position, and the remaining plates are paired, one large L-shaped pair ventro-
lateral, one ovoid pair dorso-lateral and two pairs wholly lateral.

The mouth opening is ovoid without elaboration. The oesophagus is the typical
club-shape.

Male \ The bursa is very similar to that of A . felineum with prominent prebursal
papillae, a distinct small dorsal lobe, small ventral rays which arise from a common
base, large lateral rays which all arise from the same base. The externo-dorsal ray
is markedly different from that of A . felineum in being narrow over its whole length
and in arising some distance from the base of the dorsal ray. The dorsal ray also
differs slightly in the terminal bifurcations, all of which reach the edge of the bursa
(Text-figs. 43, 44 and 45). An unmodified genital cone is present.

The spicules are simple and needle-like. The gubernaculum is short, swelling
slightly posteriorly where there are small ventro-lateral processes (Text-figs. 47 and 48).
This is in marked contrast to the relatively larger gubernaculum of A . felineum in
which the posterior end is sharply pointed with two lateral pointed processes (Text-
figs. 49 and 50).

Female : The vulva is identical with that of A. felineum with a flap developed from
the dorsal lip which is supported by dense cuticle (Text-fig. 46), and with a single
papilla-like structure posterior and slightly lateral in position.

DISCUSSION. There are at present only two species referable to the genus Arthro-
stoma: A. felineum Cameron, 1927 and A. cheni Kou, 1958. These species are very
similar but are distinguishable by the size of the spicules and the form of the guber-
naculum in the males. Further the female of A. cheni has a wart-like projection just
anterior to the vulva in addition to the usual single lateral papilla.

The species described above is similar to both those already referred to the genus
but is very distinct in the additional plate in the head, in the size of the spicules and
in the form of the gubernaculum.

Arthrostoma felineum Cameron, 1927

MATERIAL STUDIED. 2 ^ ex small intestine Felis bengalensis horsfieldi. Hatia,
Arun River, East Nepal. B.M. (N.H.) Reg. Nos. 1965. 41-42. 3 $, 3 $. Cameron's
type specimens. B.M. (N.H.) 1928.9.27.1-6.

^44 W. G. 1XGLIS & C. G. OGDKX

MEASUREMENTS (in mm.). Males. Body length : 4-23 ; 6-01. Body breadth :
0-166 ; 0-182. Length of buccal cavity : 0-068 ; 0-073. Diameter of buccal
cavity : 0-044 I 0-048. Distance of excretory pore from anterior end of body : — ;
0-382. Oesophagus length : 0-619 '> °'620. Breadth of oesophagus (maximum) :
0-108 ; 0-120. Length of spicules : 1-07 ; 1-08. Length of gubernaculum : 0-083 ',
0-080.

Corrected measurements of Cameron's type material :

Males : Oesophagus length : 0-359 '> °'36o ; 0-379. Breadth of oesophagus
(maximum) : 0-085 ; 0-094 ; 0-073.

Females : Oesophagus length : 0-420 ; 0-449 > °'456. Breadth of oesophagus
(maximum) : o-ioo ; 0-097 ; 0-089.

The two male specimens from Nepal are in good agreement with those described by
Cameron (1927), with the exception of the length of the oesophagus and the description
of the spicules. As the differences in length are more than can reasonably be attributed
to differences in methods of fixation, in view of the otherwise close similarity between
the Nepalese material and Cameron's description, the types of Arthrostoma felineum
were re-examined (B.M. (N.H.) Reg. Nos. 1928.9.27.1-6).

The oesophagus in both sexes is much longer than originally stated by Cameron
(see above) and the spicules do not form a single fine point (Text-fig. 49), but are
quite separate posteriorly. The gubernaculum is distinct (Text-fig. 50).

A. felineum has now been reported from Sumatra (Cameron, 1927), Palestine
(Witenberg, 1934) and Nepal.

REFERENCES

CAMERON, T. W. M. 1927. On Arthrostoma felineum, gen. et sp. nov., a Parasite of Cats.

/. Helminth. 5 : 143-148.
DICKMANS, G. 1935. New nematodes of the genus Longistriata in rodents. /. Wash. Acad.

Sci. 25 : 72-81.
INGLIS, WILLIAM G. & OGDEN, COLIN G. 1965. Observations on the nematode genus Citellina :

with the description of a new species, Citellina himalensis. J. Helminth. 39 : 11-18.
- 1965(3. Miscellanea nematodologica. V. Rictularia dhanra sp. nov. from a squirrel in

Nepal. Zool. Anz. 174, 227-236.
Kou, C. C. 1958. Studies on parasitic nematodes of mammals from Canton. I. Some new

species from Paradoxurus minor exitus Schwarj, Paguma larvata larvata (Hamilton Smith)

and Manis pentadactyla aurita Hodgson. Acta zool. Sinica. 10 : 60-71.
LEIPER, J. W. G. 1936. The occurrence of Molineus patens (Dujardin, 1845) in English Stoats

and Weasels. /. Helminth. 14 : 119-126.
Li, S. Y. 1941. On two new species of nematodes from China. Peking Nat. Hist. Bull. 15 :

195-199.
MAWSON, PATRICIA M. 1961. Trichostrongyles from rodents in Queensland with comments on

the genus Longistriata (Nematoda : Heligmosomatidae) . Aust. J. Zool. 9 : 791-826.
ORTLEPP, R. J. 1939. South African Helminths, Part VI. Some helminths, chiefly from

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animals of the U.S.S.R.] Trud. Cos. Inst. Eksp. Vet. 5 : 1-15. (In Russian.)
SCHULZ, R. E. & LUBIMOV, M. P. 1932. Longistriata skrjabini n. sp. (Nematoda, Tricho-

strongylidae) from the Ussuri Squirrel. Parasit. 24 : 50-53.

STRONGYLES FROM NEPALESE MAMMALS 245

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zoo. 13, 7 14

PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING

STUDIES ON THE BRITISH

DERMANYSSIDAE
(ACARI : MESOSTIGMATA)

PART I EXTERNAL MORPHOLOGY

G. OWEN EVANS

AND

W. M. TILL

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 8

LONDON : 1965

STUDIES ON THE BRITISH DERMANYSSIDAE

(ACARI : MESOSTIGMATA)
PART I EXTERNAL MORPHOLOGY

G. OWEN EVANS

AND

W. M. TILL

British Museum (Natural History)

Pp. 247-294 ; 21 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. 8

LONDON: 1965

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 13, No. 8 of the Zoological series.
The abbreviated titles of periodicals cited follow those of
the World List of Scientific Periodicals.

Trustees of the British Museum (Natural History) 196-5

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued December, 1965 Price £i is.

STUDIES ON THE BRITISH DERMANYSSIDAE
(ACARI : MESOSTIGMATA)*

PART I EXTERNAL MORPHOLOGY

By G. OWEN EVANS & W. M. TILL

CONTENTS

Page

SYNOPSIS ............ 249

INTRODUCTION ........... 249

EXTERNAL MORPHOLOGY ......... 251

Gnathosoma ........... 251

Idiosoma ........... 262

Legs ............ 279

DISCUSSION ............ 292

REFERENCES ........... 293

ABBREVIATIONS USED IN TEXT-FIGURES ....... 293

SYNOPSIS

The external morphology of the British representatives of the acarine family Dermanyssidae
is reviewed with particular reference to the modifications in morphology associated with the
adoption of a parasitic mode of life.

INTRODUCTION

THE family Dermanyssidae (including Laelapidae, Haemogamasidae and Macronys-
sidae) comprises free-living species inhabiting soil and humus as well as forms dis-
playing various degrees of association with vertebrate and invertebrate animals.
Associations with other animals range from predatory species living in the nests of
insects, birds and mammals to facultative and obligatory ectoparasites of the nesting
animal. At present the classification of the Dermanyssidae is largely based on the
study of the parasitic forms whose economic importance as vectors or potential
vectors of disease has overshadowed the taxonomic importance of their free-living
relatives. Recent key works on the parasitic Dermanyssidae have been produced by
Bregetova (1956), Strandtmann & Wharton (1958) and by Zumpt & Till (1961).

The British representatives of this family number about 75 species and these
exhibit a wide range of structural and biological adaptations to the variety of
ecological niches they have successfully colonized. This first contribution to a
taxonomic revision of the British Dermanyssidae deals with the external morphology
of the group.

* This study was supported, in part, by Research Grant No. £4656 from the National Institutes of
Health, United States Public Health Service.

ZOOL. 13, 8 15

250

G. OWEN EVANS & W. M. TILL

LARVA

, PROTO-, DEUTONYMPH + FEMALE

PROTO- DEUTONYMPH * FEMALE

MALE

LARVA PROTO- DEUTONYMPH + FEMALE

MALE

LARVA . . PROTONYMPH , DEUTONYMPH

FEMALE

MALE

FIG. i. Chelicerae of the immature and adult stages of : A. Holostaspis vitzthumi (Womersley) ;
B. Laelaps echidnina Berlese ; c. Dermanyssus gallinae (Degeer) ; D. Ornithonyssus bacoti (Hirst).

THE BRITISH DERM AN YSSID AE (ACARI) 251

EXTERNAL MORPHOLOGY

Gnathosoma

Chelicerae: The chelicerae, paired pre-oral trophic appendages, of the Der-
manyssidae are three segmented (Text-fig, i). The short proximal segment to which
the cheliceral retractor muscles are attached is articulated to the longer second seg-
ment by a condylar process located on its internal (paraxial) face. Distally the
second segment forms the fixed digit (/. d.) of the chelicera and the third segment or
movable digit (m. d.} is articulated ventrally to the fixed digit by arthrodial mem-
brane and two condyles developed on the limb of the second segment which engage
acetabula on the movable digit (Text-fig. 2E). The movable digit is operated by a
pair of opposed muscles originating within the proximal segment and connected to
the digit by a dorsal levator tendon and a ventral depressor tendon. The arthrodial
membrane at the base of the movable digit is usually produced into setiform processes.

In the free-living and the majority of the nest-inhabiting members of the family, the
cheliceral shaft is of approximately equal diameter throughout its length and the
ratio of the length of the first to the second segment is in the region of I : i'S-3'5.
The external (antiaxial) face of the chelicera bears a dorsal seta (d.s.),a. lateral fissure
(lat. l.f.) and a simple pilus dentilis (p. d.}1, the latter being situated on the distal half of
the fixed digit (Text-fig 2A-B). A lyriform fissure (d. l.f.) associated with the dorsal
seta occurs on the dorsal surface of the second segment at the origin of the fixed digit.
In some forms this fissure extends to the lateral face of the segment. The fixed digit
is provided with a variable number of teeth, but the movable digit in the nymphae
and females is normally bidentate. In the male the movable digit, usually uni-
dentate, carries on its external face a grooved spermadactyl (sp.). This process
shows considerable diversity in form but its distal portion is invariably free (Text-fig.
2F-G). At the larval stage the chelicerae are well-developed and have a full com-
plement of setae and fissures. The digits are weaker and less strongly dentate than
in subsequent developmental stages since this stage is non-feeding (Text-fig. IA).
The chelicerae of the nymphae and female are essentially similar in form, the only
differences being in their relative sizes and occasionally in the number of teeth on the
fixed digit.

This basic type of chelicera, encountered in the polyphagous free-living forms, has
become variously adapted for specialized feeding in the facultative and obligatory
parasitic species of the family. Modifications are evident in the form of the cheliceral
shafts, the digits and associated structures, and in the ontogenetic development of
the chelicerae.

The form of the shaft (first and second segments) in the larva, nymphae and female
of the facultative parasites, for example Androlaelaps , Laelaps and Haemogamasus , is
fundamentally the same as in the free-living forms although in the males there is a
tendency for a shortening of the second segment (Text-fig. IB). In obligatory
parasites the relative lengths of the first and second segments of the chelicera in the

1 Van der Hammen (1964) has suggested replacing the term pilus dentilis by " cheliseta " since the
former is " too long ". To change such a well-established and universally accepted term for such a
trivial reason is unwarranted!

252

G. OWEN EVANS & W. M. TILL

non-feeding larva are similar to those in the same developmental stage of the free-
living forms and facultative parasites but the distal half of the second segment
generally shows some attenuation. The feeding nymphae and female, with few

m.d.

acef.

FIG. 2. Cheliceral digits of certain Dermanyssidae. A. Eulaelaps stabularis (Koch) female;
B. Androlaelaps fahrenholzi (Berlese) female ; c. Haemogamasus hirsutus Berlese female ;
D. Ornithonyssus bacoti (Hirst) female ; E. condylar articulation of the movable digit
in a free-living Gamasine mite ; F. Hypoaspis (H.) krameri (Canestrini) male ; G. Laelaps
hilaris Koch male.

THE BRITISH DERM AN YSSID AE (ACARI) 253

exceptions, show a distinct lengthening of the second segment in relation to the first.
Both nymphae and female of Dermanyssus have the second segment enormously
elongated so that the chelicerae resemble stylets (Text-fig, ic). A less spectacular
elongation of this segment is also evident in the feeding stages of the Macronyssinae
(Text-fig. ID) in which the mesial surface of the segment is distinctly flattened. An
interesting phenomenon in this group is the marked difference between the chelicerae
of the feeding protonymph and the non-feeding deutonymph, the chelicerae of the
latter reverting to the form in the non-feeding larva. Two main types of male
chelicerae are present, one (Dermanyssus} in which the second segment is not more
than twice the length of the first and the other (Ornithonyssus) in which the second
segment is about three to four times the length of the first. This difference in the
length of the second segment appears to be related to the length of the spermadactyl
(see below) .

The modifications of the digits and their associated structures are diverse and often
complex. With few exceptions, the digits of the larva, nymphae and female of the
facultative parasites retain their chelate-dentate form, the major specialization in this
group being apparent in the degree of development and form of the setae, fissures and
arthrodial processes. The short slender pilus dentilis of the free-living forms is often
enlarged and inflated as in post-larval stages of certain species of Androlaelaps (Text-
fig. 2B), Laelaps and Haemogamasus. The dorsal seta is usually relatively short and
simple, but may be lacking in certain Androlaelaps. In Haemogamasus, on the
other hand, this seta is greatly enlarged and may be simple or spatulate. It tends to
migrate ventrally onto the external face of the chelicera and in the male of H. horridus
(Berlese) it has moved anteriorly onto the fixed digit. The arthrodial processes may
be simple and arranged in the form of a " coronet " as in the free-living species, but in
certain members of the Androlaelaps-Laelaps complex there is a tendency for a
reduction in the number of the processes accompanied by their elongation, for
example in Hyperlaelaps amphibia Zachvatkin. Lyriform fissures are present in the
normal position. Exceptions to the chelate-dentate form of the chelicerae occur in
the nymphae and females of some species of Androlaelaps and Haemogamasus. In
the ethiopian species A ndrolaelaps cryptomia (Radford), for example, the fixed digit
is reduced to a weakly sclerotized, terminally unhooked process bearing a long pilus
dentilis, whilst the movable digit is bipartite, comprising a strongly sclerotized,
edentate, hooked digit with a less strongly sclerotized, dentate, digitiform process
arising externally from its basal portion in much the same way as a spermadactyl on
a male chelicera. Both digits in Haemogamasus hirsutus Berlese, on the other hand,
are somewhat elongated and edentate, and the dorsal seta is lacking (Text-fig. 20).

Two main types of chelicerae are present in the males of the facultative parasites.
The least specialized form occurs in Eulaelaps and Haemogamasus in which the
movable digit, usually unidentate, retains its hooked form and the grooved sperma-
dactyl, free distally, rarely extends beyond the tip of the digit by more than one-third
the length of the digit. Arthrodial processes are reduced or lacking on the chelicerae
of this sex. Considerably greater specialization is apparent in the second type which
occurs in males of the Androlaelaps-Laelaps complex. These are characterized by

254 G OWEN EVANS & W. M. TILL

the reduction of the fixed digit to a relatively slender, distally tapering process
bearing the enlarged pilus dentilis and by the size of the grooved spermadactyl which
forms the greater part of the movable article of the chelicera (Text-fig. 2G). The
typical form of the movable digit is lost although, in most species, its distal portion is
recognizable as a slender pointed process protruding dorsally from the grooved
spermadactyl. As in the Haemogamasus-type the arthrodial processes often show
marked reduction in number and, rarely, the dorsal seta is lost.

As one would expect, the obligatory parasites feeding on the tissue fluids of their
hosts show the highest degree of specialization of the chelicerae. The digits in the
larva are small, subtriangular, edentate and functionless as trophic appendages. They
lack setae and, apparently, fissures (Text-figs, ic and D). With the exception of the
Dermanyssinae, in which the digits are minute but dentate (Text-fig, ic), the digits
of the feeding nymphs and females are very weakly dentate or edentate and lose the
hooked form typical of the polyphagous species. In the Macronyssinae, the fixed
digit is generally slender and often terminates in a rounded hyaline boss whereas the
movable article has an elliptical trough formed by cuticular outgrowths of the digit
along its dorsal surface, the normal dentate surface of an unmodified chelicera (Text-
fig. 2D). Both the digits are dorsoventrally flattened and grooved along their
" cutting " faces in Hirstionyssus. We have been unable to see a pilus dentilis on
the fixed digit although, in some species, a small depression occurs on the digit at the
normal site of this sensory seta. The dorsal seta is reduced to a microseta or is
lacking. There are no distinct arthrodial processes. The forms of the chelicerae in
the males are comparable to the two types described for the facultative parasites. All
macronyssines and Hirstionyssus have a relatively short spermadactyl showing
varying degrees of fusion with the digit. In the dermanyssines, however, the fixed
digit is strongly reduced and it is difficult to distinguish the movable digit from the
proximal body of the elongated spermadactyl (Text-fig, ic). Unlike the facultative
parasites, the chelicerae of the males of the obligatory parasites lack a pilus dentilis
and the dorsal seta, as in the female, is present as a microseta or is lost. The shorten-
ing of the second cheliceral segment in the forms with an extremely long spermadactyl
(see above) has the effect of strengthening the shaft to counteract the increased weight
of the movable digit and its appendage.

In those obligatory parasites (Dermanyssinae and Hirstionyssus) in which the two
nymphal stages and the female are active and feeding the form of the digits is
determined at the protonymphal stage and is retained by the deutonymph and female.
In the macronyssines in which the deutonymph is relatively inactive and non-feeding,
the specialized functional digits of the protonymph are not developed in the deuto-
nymph whose chelicerae revert to the larval form. At the succeeding feeding female
stage, however, the chelicerae resemble those of the protonymph.

Tectum capituli: The tectum2, which forms the roof of the gnathosoma and

2 At present the term tectum i s also used to describe a shelf-like structure occurring below the vertex
and forming the roof of the camerostome in some Uropodina. It is not a gnathosomal structure and
appears to provide a supporting (and protective) shelf for the first pair of legs when the animal is at rest.
As such, and to avoid confusion, it seems appropriate to re-name this structure pedotectum-I in the
Uropodina.

THE BRITISH DE RM AN YSSI D AE (ACARI)

255

overlies the chelicerae (Text-figs, 3, 45 and D), shows considerable variety in its degree
of development and in the form of its anterior margin. In the free-living species and
facultative parasites, with the exception of the Eulaelaps-Haemogamasus group, the
anterior margin is smooth (Melittiphis), basically trispinate (Pseudolaelaps, Text-fig.
35) or multidenticulate (Hypoaspis, Text-fig. 3A), and does not extend anteriorly to
completely cover the hypostome and its associated structures (Text-fig. 4B) . Members
of the Eulaelaps-Haemogamasus group, however, have the free margin of the tectum

FIG. 3. Tectum capituli of : A. Hypoaspis (Gaeolaelaps) praesternalis Willmann ; B.
Pseudolaelaps doderoi (Berlese) ; c. Haemogamasus ambulans (Thorell) ; D. Dermanyssus
gallinae (Degeer).

strongly fimbriated and considerably elongated so that it entirely covers the hypo-
stome from above (Text-fig. 30, 40). This anterior extension of the tectum is also
characteristic of the obligatory parasites (Dermanyssinae, Macronyssinae) and takes
the form of a triangular lobe, smooth or denticulate at its tip, whose down-turned
lateral margins overlap the lateral walls of a trough formed by the modified internal
and external malae of the hypostome (Text-figs. 30 and 50).

Basis capituli and hypostome: The basis capituli is formed mainly by the
enlarged coxae of the pedipalps which are separated mid-ventrally by a shallow
capitular groove whose floor is considered to represent the sternite of the pedipalpal
segment (the deutosternum) . A longitudinal series of antrose deutosternal denticles
(d. d.) occurs on the floor of the capitular groove and the number and form of the
denticles show little intraspecific variation (Text-figs. 4-5). Each member of the
longitudinal series of denticles may be simple (one denticle) or comprise two to eight
denticles arranged in a transverse row (Text-figs. 4-5) . In free-living forms the larva
appears to have a full longitudinal complement of denticles, but the number of denticles
in each transverse row may vary in the succeeding developmental stages. Larvae of
obligatory parasites, however, have the deutosternal denticles markedly reduced or
absent and it is at the protonymphal stage that the full complement is determined.
The number of denticles comprising the longitudinal series appears to be characteris-
tic for certain groups of genera of the Dermanyssidae. Free-living forms and

ZOOL. 13, 8 is§

256

G. OWEN EVANS

W. M. TILL

members of the Androlaelaps-Laelaps groups have six, rarely seven, transverse rows
of denticles /longitudinal series (Text-fig. 4A) with the number of denticles in the

D C

FIG. 4. A-B. Gnathosoma of female of Cosmolaelaps claviger (Berlese), A. ventral and B.
lateral view. C-D. Gnathosoma of female of Haemogamasus hirsutus Berlese, c. ventral
and D. lateral view.

THE BRITISH DERM AN YSSID AE (ACARI) 2.57

transverse rows ranging from two to eight, the lower numbers being present in the
facultative parasites of the Androlaelaps-Laelaps group. On the other hand, mem-
bers of the facultatively parasitic Eulaelaps-Haemogamasus group always have more
than seven transverse rows of denticles /longitudinal series, the usual number ranging
between nine and fourteen (Text-fig. 40). In the obligatory parasites the floor of the
capitular groove contains about nine or ten small triangular denticles in a single
longitudinal series ; rarely, two denticles form a transverse row (Text-fig. 5). A
similar longitudinal series of denticles occurs in Melittiphis and Myonyssus whose
feeding habits are uncertain, but in these genera the corniculi (external malae) and
the chelicerae do not show the marked modifications of the obligatory parasites
(Dermanyssinae and Macronyssinae) .

The basis capituli bears ventrally a pair of setae, the capitular setae, which first
appear at the protonymphal stage (Evans, 1957).

The hypostome, divided mid-ventrally by the anterior extension of the capitular
groove, bears three pairs of setae (hyp. i, 2, 3) in the nymphae and adults. Setae
hyp. 2 and 3 form a more or less transverse row of four setae (Text-figs. 4-5). At the
larval stage only two pairs of setae are present on the hypostome (hyp. I and 2) ;
the third pair (hyp. 3) first appears at the protonymphal stage. In the free-living
forms and in the majority of the facultative parasites the hypostome bears antero-
laterally a pair of large, horn-like structures (hypertrophied setae) termed corniculi
(or external malae}. The floor of the capitular groove in the region of the hypostome
is considered by some acarologists to represent the sternite of the cheliceral segment
and is referred to as the protester num. It is divided into two parts by a longitudinal
suture and anteriorly each part is produced into variously shaped process (Text-fig.
4) termed internal malae (or hypostomal processes) .

The main external features of the hypostome in the obligatory parasites are its
marked elongation (Text-figs. 5B, c and D) and the modification of the internal malae
and corniculi to form a pre-oral trough. The corniculi, analagous structures to the
rutetta of the Cryptostigmata, already show certain deviations from their normal
horn-like form in the facultative parasite Haemogamasus hirsutus Berlese, where they
become modified into flat lobe-like structures (Text-figs. 40 and D) . Further speciali-
zation of the corniculi results in the complex flange-like structures forming the lateral
walls of the hypostomal trough in the obligatory parasites. The ventral floor of this
trough in the protonymph and female is formed by the fusion of the internal malae
along the longitudinal suture on the floor of the capitular groove (Text-fig. 5B).
Owing to the complex nature of the internal malae and corniculi in these specialized
parasitic forms, it is difficult to determine their homology with those in the free-
living forms. At the non-feeding deutonymphal stage of Ornithonyssus bacoti (Hirst)
fusion of the internal malae does not occur and although the internal malae and the
corniculi are less complex than in the feeding stages it is still difficult to elucidate their
homology (Text-fig. 5A) . Two interpretations are possible, namely, one in which the
short, pointed, internal process is homologous with the internal malae of the free-
living forms and the two outer processes with the corniculus, and the second in which
only the external article is homologous with the corniculus, the internal pair being

G. OWEN EVANS & W. M. TILL

.int. maL

corn.
int mal.

_s.s.

FIG. 5. Gnathosoma of Ornithonyssus bacoti (Hirst) : A. female deutonymph, ventral
and lateral views ; B. female, ventral view ; c. female, lateral view ; D. male, ventral
view.

THE BRITISH D ERM AN YSSID AE (ACARI) 259

the internal malae. In the male of 0. bacoti the "internal malae " are in the form of
two blades which overlap anteriorly and thus differ quite markedly from the form in
the protonymph and female (Text-fig. 50).

Labrutn: The labrum (epipharynx of some authors), an extension of the dorsal-
wall of the pharynx, appears as a long tapering structure between the corniculi and
dorsal to the internal malae (Text-fig. 4) . In the free-living forms and the facultative
parasites the labrum projects beyond the tip of the hypostome but it is usually hidden
by the elongated hypostome in the obligatory parasites. Its margin may be fim-
briated and in the Androlaelaps-Laelaps group it has a distinct median longitudinal
groove.

Salivary styli: The paired salivary styli lie laterally, immediately ventral to the
chelicerae, and carry the ducts of the salivary glands located in the idiosoma. They
are large and particularly well-sclerotized in the obligatory parasites (Text-figs. SB
and D).

In the forms having horn-like corniculi the salivary stylus lies in a lateral or
dorsal channel along the length of the corniculus. Thus, the corniculi function as
protective guides for the styli.

Pedipalps: The pedipalps have six free segments (trochanter, femur, genu,
tibia, tarsus and apotele) of which the terminal segment, the apotele, is represented
by a tined seta-like process at the inner basal angle of the tarsus (Text-fig. 6A) . Weak
condylar processes are present externally on the trochanter and dorsally on the
distal margins of the femur and genu. A conspicuous lyriform fissure occurs proxi-
mally on the dorsal surface of the genu. These appendages have a well-defined
chaetotaxy.

The normal ontogenetic sequence of development of the setae of the pedipalp has
been defined by Evans (1964) and is characteristic of the free-living forms and of the
majority of the facultative parasites of the Dermanyssidae. At the larval instar the
trochanter lacks setae ; the femur bears four (al, dv dz and pi], the genu five (al1,
dl-ds and pi), the tibia 12 and the tarsus n setae. One seta (v^ is added to the
trochanter, and three setae to the tarsus at the protonymphal stage. A single seta is
further added to the trochanter (vz), femur (d3) and genu (alz), and two setae to the
tibia at the deutonymphal stage. The deutonymphal chaetotactic pattern is
retained by the adults. Thus the segmental formulae for the pedipalp (excluding the
apotele which is present in all post-embryonic developmental stages) are : L.
(0-4-5-12-11) ; p. (I_4_5_I2_I5) ; D. Ad. (2-5-6-14-15).

Deviations from the normal setal complement occur in both facultative and
obligatory parasites as well as in certain myrmecophilous forms and are the result,
in many cases, of the retention of the larval complement of setae on a segment in
succeeding developmental stages. For example, in the adults of Laelaspulus the
tibia retains the larval number of setae throughout ontogeny, with the result that the
adult formula is (2-5-6-12-15), whilst Hyperlaelaps amphibia Zachvatkin retains
the larval number on the femur and genu (Text-fig. 6s) . In the latter the tibia is also

2&0

G. OWEN EVANS & W. M. TILL

E F G H

FIG. 6. A. Right pedipalp of Hypoaspis (Gaeolaelaps) aculeifer (Canestrini), dorsal view ;
B. right pedipalp of Hyperlaelaps amphibia Zachvatkin, dorsal view ; c. palptrochanter
of Ornithonyssus bacoti (Hirst) female, external (antiaxial) view ; D. palptrochanter of
Eulaelaps stabularis (Koch) female, ventral view ; E-F. right palptibia and tarsus of
Ornithonyssus bacoti (Hirst), larva (E), protonymph (F), deutonymph (G) and adult (H),
in dorsal view.

THE BRITISH DERM AN YSSID AE (ACARI) 261

unideficient in the deutonymph and adult (2-4-5-13-15). Amongst the macronys-
sines the normal pedipalpal chaetotaxy has been observed only in Macronyssus.
Members of the other genera of this group show a deficiency in their tibial chaetotaxy.
For example, Sauronyssus, Steatonyssus , Ophionyssus and the majority of the species
of Ornithonyssus have only 13 setae on this segment. Some species of Pellonyssus,
Ornithonyssus (0. sylviarum) and Hirstionyssus retain the larval complement of 12 in
the nymphae and adults.

The greatest diversity in palpal chaetotaxy occurs in Dermanyssus. Only D.
intermedius Evans & Till appears to have the normal number of setae. Deutonymphs
and adults of the other species, except D. alaudae (Schrank), have the formula
(2-4-6-12-15), indicating that the larval number has been retained on the femur and
tibia. In D. alaudae the chaetotaxy of the four proximal segments is (2-4-5-7/8) ;
thus, the femur and genu show no increase in setation from the larval condition whereas
the tibia displays larval specialization, that is, the segment never bears the complete
larval number.

The ontogenetic development of the pedipalpal chaetotaxy in the macronyssines
follows the normal pattern on the trochanter, femur and genu only, that is L. (0-4-5),
P. (1-4-5) and D. Ad. (2-5-6) . At the larval stage of 0. bacoti, for example, ten setae
are present on the tibia and nine setae on the tarsus (Text-fig. 6E). There is an
increase to the normal complement of 12 and 15 respectively on these segments in
the protonymph whilst in the deutonymph, the protonymphal complement of 12
setae is retained on the tibia and the tarsus shows a regression to about 12 setae. In
the adult 13 (unideficient) and 15 (normal) setae are present on the tibia and tarsus
respectively. An interesting feature of the chaetotaxy of these distal segments is
the reduction in the size and in the number on the tarsus of the hollow eupathidia-
like setae in the non-feeding larva and deutonymph (Text-figs. 6E and G).

There is considerable variety in the form of the palpal setae (simple, pilose, leaf-
like, spinose etc.) which provides stable taxonomic criteria. Hollow setae super-
ficially resembling eupathidia occur on the tibia and tarsus, and are probably
chemo-receptors. A pair is located dorsally on the distal margin of the tibia and a
group of nine or ten on the distal half of the tarsus (Text-fig. 6A and H) .

A median, longitudinal, keel-like process occurs ventrally on the trochanter of
certain macronyssines (Text-fig. 6c) . It appears to occur in females only of Steato-
nyssus s. str., some Pellonyssus and Ornithonyssus, but in both sexes and the pro-
tonymphs of Macronyssus. Its function is not known.

Both sexes of Eulaelaps have a unique organ of probable sensory function on the
ventral surface of the trochanter (Text-fig. 60). It was first observed by Fonseca

(1935).

The apotele may be two or three-pronged. A three-pronged apotele occurs in the
genera Ololaelaps, Pseudoparasitus, Gymnolaelaps and in some species (H. hirsutus
and H. pontiger) of the genus Haemogamasus. The prongs may be slender with the
proximal member small and relatively inconspicuous as in Ololaelaps and Pseudo-
parasitus or the three prongs may be large and spatulate as in some Gymnolaelaps.
All the other species we have examined have a two-pronged apotele.

262

G. OWEN EVANS & W. M. TILL

\ 4| rf

lc -3 Ms

THE BRITISH DERM AN YSSID AE (ACARI)

Idiosoma

263

Dorsal sclerotization : The ontogenetic development of the sclerotization of the
dorsum of the idiosoma in the free-living forms and in the facultative ectoparasites
follows a definite and constant sequence. In the few larvae we have examined the
dorsal sclerotization is relatively indistinct, the podonotal shield being the only one
discernible (Text-fig. SA). At the protonymphal stage a large anterior podonotal
shield and a smaller posterior pygidial shield are always present, and on the striated
cuticle between these two scutal elements lie a variable number of mesonotal scutellae
(Text-fig. 70). These scutal elements coalesce at the deutonymphal stage ; pre-
sumably the pygidial shield and the mesonotal scutellae first unite and the resulting
opisthonotal shield fuses with the podonotal shield except at its lateral margins to
form a single dorsal shield with a pair of lateral incisions (Text-fig. 76). Further
fusion occurs at the adult stage resulting in the disappearance of the lateral incisions
(Text-fig. 70).

In order to allow for the expansion of the idiosoma which is necessary to accom-
modate the large quantities of tissue fluids taken during feeding, the obligatory
parasites show a decrease in idiosomal sclerotization. This is first evident at the
protonymphal stage by the reduction in the size of the pygidial shield (Text-fig, yd) .
No distinct scutal elements are present in the larva (Text-fig. SB). In the macro-
nyssines the podonotal shield and the mesonotal scutellae of the protonymph resemble

pod. sh.

A B

FIG. 8. Dorsal view of the larva of : A. Holostaspis vitzthumi (Womersley) ; B. Ornithonys-

sus bursa (Berlese).
ZOOL. 13, 8 i5§§

264 G. OWEN EVANS & W. M. TILL

the condition in the same instar of the free-living forms, but the pygidial shield is
markedly reduced in size as is evident from its chaetotaxy (Text-figs. IOA and c).
The non-feeding deutonymphal stages (male and female) of this group are very weakly
sclerotized and it is difficult to differentiate the sclerotized area from the sur-
rounding cuticle. It is probable that the extent of the weakly sclerotized areas is
similar to that in their respective adult stages. All males, with the exception of
Steatonysella, which has two dorsal shields, have an entire dorsal shield formed as in
the free-living species by the fusion of the podonotal, mesonotal and pygidial shields.
This is also the condition in the female of Macronyssus, Sauronyssus and the
majority of Ornithonyssus (Text-fig, jh}. The females of Steatonyssus and
Pellonyssus, on the other hand, retain a distinct podonotal shield and have a smaller
opisthonotal shield consisting of the pygidial and mesonotal elements (Text-fig. 7g).
Ornithonyssus aridus Furman & Radovsky is exceptional amongst its congeners in
retaining more or less the same form of dorsal sclerotization as the protonymph
(Text-fig. 7/). The most specialized form occurs in females of Ophionyssus where
distinct podonotal, mesonotal and pygidial elements are present and the pygidial
shield shows a decrease in size in comparison with the protonymph (Text-fig. 70).

Protonymphs of the Dermanyssinae show a further reduction of the pygidial
shield (Text-figs, ji and /). In the genus Liponyssoides the extremely small pygidial
shield is either retained by the deutonymph and female as a separate scute (Text-
figs. 7; and&) or fused with the remainder of the dorsal shield whilst in Dermanyssus
the pygidial shield is lacking. The protonymphs of both genera have a strong posterior
prolongation of the podonotal shield which extends between the mesonotal scutellae
and serves as a more posterior attachment site for the retractor muscles of the greatly
elongated cheliceral shafts. At the deutonymphal and adult stages of Dermanyssus
an entire dorsal shield is usually present and this is probably formed by the posterior
extension of the podonotal shield and its fusion with the mesonotal scutellae (Text-
figs, jm and n). Even within this genus there is a considerable variety in the dorsal
sclerotization of the deutonymph and female, depending on the degree of development
of the posterior prolongation of the podonotal shield. For example, in D. scutatus
Krantz the posterior extension of the podonotal shield is very weak and bears only
the first pair of dorsal opisthonotal setae, and the mesonotal scutellae are retained.
The males of the Dermanyssinae have an entire dorsal shield.

Dorsal chaetotaxy: The dorsum of the idiosoma has a well defined chaetotaxy
whose ontogenetic development is relatively stable. Deviations from the normal
setal complement (the holotrichous condition) are relatively common in the specialized
facultative and obligatory parasites and may be the result of hypo- or hypertrichy.
The system of nomenclature adopted for the dorsal chaetotaxy in this work is that
proposed by Lindquist & Evans (1965) and is a modification of the system used by
Hirschmann (1957). In the adult holotrichous condition the dorsum is considered to
bear 44 pairs of setae arranged in four longitudinal rows of n setae, a dorsal (/, /),
a median (z, Z), a lateral (s, S) and a marginal series (r, R), on each side of the median
longitudinal axis of the idiosoma, or alternatively in n transverse rows of eight setae.

THE BRITISH DERM AN YSSID AE (ACARI)

265

The setae of the podonotal region are denoted by letters in lower case and those of the
opisthonotum by capitals (Text-fig, gc).

Van der Hammen (1964) has been extremely critical of the " artificial " (we should

'«ir6

«;R2

FIG. 9. Semi-diagrammatic representation of the dorsal chaetotaxy in the protonymph
(A), deutonymph (B) and adult, dorsal (c) and lateral (D) view, of a free-living or nest-
inhabiting dermanyssid (based on Laelaps}.

266 G. OWEN EVANS & W. M. TILL

prefer to call it " practical ") system of setal nomenclature used in the Mesostigmata
on the basis that the division of the idiosoma into two regions is probably artificial
and useless for notation, since there is no evidence that the posterior margin of the
podonotal shield and the anterior margin of the opisthonotal shield represent borders
of tagmata. According to this author only a metameric arrangement of setae, that
is in transverse rows, has phylogenetic value.

Theoretically, a metameric arrangement of setae would be ideal, but how is one to
determine metamerism in a group which shows no evidence of idiosomal segmentation
during post-embryonic development or for which there is, at present, no embryological
evidence upon which to base metamerism? Certainly, the porotaxy of the dorsum
shows such variability both in number and form of the " pores " as to be of doubtful
value or significance, and we can see no basis for van der Hammen's statement that
the " pores " show a metameric arrangement because their position corresponds to
the borders of segments. In fact, any attempt to define metameric segmentation in
the Mesostigmata on present evidence would be entirely speculative, and a system of
setal terminology based upon it could be just as " artificial " as the system in current
use. Such terms as " metameric arrangement " and " phylogenetic value "used at
present in reference to a system of setal nomenclature in the Mesostigmata can only
be classed as " red herrings " whose sole value is to add a pseudo-academic flavour to
the text.

The main criterion for a system of setal nomenclature, in the absence of evidence
of metamerism or of a primitive setal pattern, should be practicability. By this we
mean that reference, for taxonomic purposes, can be made to individual setae and a
comparison is possible between the setal patterns of related forms on the basis of
" positional homology ". The modified Hirschmann system of setal terminology
proposed by Lindquist & Evans (1965) enables one to do this except in cases of
extreme hypo- or hypertrichy of the dorsal chaetotaxy. This system retains the
division of the chaetotaxy into a podonotal and an opisthonotal series since it is of
practical value. The podonotal shield is a characteristic feature of the larval and
protonymphal stages of the Mesostigmata and is retained as such (except for an
increase in sclerotization laterally) in the deutonymphal and adult stages of many
species. It can be readily defined by its chaetotaxy. No claim is made that the
podonotal and opisthonotal shields define tagmata although it would indeed be
surprising if the widespread occurrence of a division of the dorsal sclerotization of the
idiosoma in the Mesostigmata at the podonotal-opisthonotal junction was without
some functional significance.

Larvae of the Dermanyssidae are weakly sclerotized although it is usually possible
in the free-living forms to define a podonotal shield which bears nine pairs of setae
(ji, J3-J6, Z2, Z4, z$ and 54). A tenth pair of podonotal setae (s6) occurs on the
striated cuticle lateral to ;6 (Text-fig. SA). The chaetotaxy of the opisthonotum is
considerably less stable ; for example, four pairs of setae are present in this region in
some Holostaspis and seven pairs in some Hypoaspis s. lat. We have not attempted
to name these setae. Few larvae of the obligatory parasites have been examined.
In Ornithonyssus bacoti, for example, the podonotal region shows a deficiency of two

THE BRITISH D E RM AN YSS I D AE (ACARI)

pod. sh,

267

laf. in.

mes. sc.

pyq. sh.

t'r'V • <

FIG. 10. Dorsum of : A. Macronyssus flavus (Kolenati) protonymph ; B. Dermanyssus
gallinae (Degeer) protonymph ; c. Ornithonyssus bacoti (Hirst) protonymph ; D. Haemo-
gamasus nidi Michael, deutonymph.

268 G. OWEN EVANS & W. M. TILL

pairs of setae (^'3 and s6) and the opisthonotum bears only three pairs of long, slender
setae. Setae j$, however, are present in some larvae of the Macronyssinae but the
presence of only six opisthonotal setae appears to be characteristic for the group.

At the protonymphal stage of the free-living forms and the majority of the facul-
tative parasites, the number of podonotal setae is increased to 16 pairs of which eleven
pairs (J2 and 55 being additional to the nine pairs occurring on the shield in the larva)
are situated on the podonotal shield and five pairs (r2, r3,r$, s6 and r6) on the cuticle
lateral to the shield (Text-fig. QA) . Typically, the pygidial shield bears eight pairs of
setae (/3-/5, Z^-Z^, 54 and 55) and the unsclerotized cuticle of the opisthonotum
six pairs (Ji, J2, Zi, Z2, 52 and 53) making a total of 14 pairs of opisthonotal setae
and a dorsal complement of 30 pairs. In the facultative parasite Eulaelaps stabularis
(Koch) the pygidial shield is reduced in size and bears only six pairs of setae (/3-/5
and Z3-Z5), setae 54 and 55 lying on striated cuticle. The total number of setae on
the opisthonotum is sixteen pairs, an increase of two pairs over the normal. This
may be the beginning of the hypertrichy of the dorsum which is so marked in the
deutonymph and adult of this species. There is no hypertrichy of the podonotum at
this instar. On the other hand, in the protonymphs of Haemogamasus nidi Michael
there is marked hypertrichy of the lateral and marginal series in both the podonotal
and opisthonotal regions.

The podonotal chaetotaxy of the protonymphs of the obligatory parasites is
essentially the same as in the free-living forms (Text-fig. IOA) . However, there is a
tendency for hypotrichy of the _;' series by the suppression of j^, for example in some
species of Dermanyssus, Ornithonyssus and Pellonyssus, or of j$ and;4 as in Dermanys-
sus alaudae (Schrank) . Varying degrees of hypo- and hypertrichy are evident on the
opisthonotum. In Dermanyssus gallinae (Degeer) the normal complement of / and Z
series of setae is present but the 5 series is deficient, comprising only two pairs of
setae (Text-fig. IOB). 0. bacoti (Text-fig, ice), on the other hand, shows a deficiency
in the number of / setae (/4 and /5 being suppressed) whilst Macronyssus flavus
(Kolenati) has the full complement of / and Z setae, and marked hypertrichy in the
region of the 5 and R series. The widespread occurrence of hypo- and hypertrichy of
the opisthonotum in the dermanyssines and macronyssines makes it difficult to use
the relative positions of setae as a basis for comparing the chaetotaxy of this region
with that of free-living forms, and we have restricted the naming of setae to those
regions unaffected by these phenomena.

The full complement of dorsal setae is attained at the deutonymphal stage except
possibly in some forms showing extreme hypertrichy (Text-figs. QB-D). In the
majority of the free-living forms and facultative parasites the dorsal, median and
lateral series situated on the dorsal shield form six longitudinal rows of eleven setae
(or eleven transverse rows of six setae). Setae added to these series at the deutonym-
phal stage are : zi, 23, z6, 51-53 and Si (Text-fig. 93). The marginal series on the
podonotum is fairly stable and comprises setae r2-r6 (74 being added in the deutonym-
phal instar and r6 remaining on the lateral integument) although in certain nest
inhabiting forms (Pneumolaelaps) and facultative parasites (Laelaps) hypertrichy of
this series is not uncommon. This series (R) and the submarginals (UR) are con-

THE BRITISH DERM AN YSS ID AE (ACARI) 269

siderably more variable on the opisthonotum and are commonly affected by hypo-
or hypertrichy. Hypoaspis praesternalis Willmann, for example, has only one pair
of R setae whilst in Pneumolaelaps marginalis Willmann both series are markedly
hypertrichous and each comprises over 25 setae! Differences also occur in the
number of opisthonotal marginal series (R) between the sexes of a given species and
this is first evident at the deutonymphal stage, the male deutonymph and adult
male having fewer setae in this series than the female deutonymph and adult female.
Two further interesting features of the dorsal chaetotaxy are exhibited by the laela-
pines. The first is the appearance of a supernumerary row of setae between the / and
Z series of the opisthonotum. This series, which is absent in Stratiolaelaps , some
Androlaelaps and Hyperlaelaps , comprisises a maximum of three setae /row, referred
to as posterior accessories (pxi-3) by Costa (1961). The maximum complement of px
setae occurs in some Androlaelaps, but the usual number present is two pairs, p%2
and px$ (Text-fig. 90). Only one pair of the series (px2)is present in some Hyper-
laelaps. The second feature is the tendency for hypertrichy to occur in the region of
the / series in forms associated with other animals either as nest-inhabitants or
facultative parasites. These unpaired setae show considerable intraspecific varia-
bility in number and in length. General hypertrichy of the podonotum and opis-
thonotum occurs at the deutonymphal stage of the Eulaelaps-Haemogamasus group
and completely obscures the primary chaetotaxy (Text-fig. IOD).

Even within the free-living and facultative forms some hypotrichy of the dorsal
and median series of setae is evident. For example, in Ololaelaps the deutonymphal
setae zi and 23 are suppressed and in Ondatralaelaps3 j$ and 23 are lacking on
the podonotum and /3, /4 and Z3 on the opisthonotum. Rarely, in some Androlae-
laps, a supernumerary seta occurs on the podonotal region of the dorsal shield lateral
to $4 and has been referred to as an anterior accessory seta (ax) by Costa.

The dorsal chaetotaxy of the deutonymphs of the obligatory parasites is charac-
terized by the tendency for the retardation in the development of the chaetotaxy of
the podonotum and the occurrence of hypertrichy in the lateral and marginal series.
Further, in the non-feeding deutonymphs of the macronyssines two distinct types of
dorsal chaetotaxy are present, corresponding to the form in the adult male and
female (Text-figs. IIA and c). Reference has already been made to the hypotrichy of
the dorsal series at the larval and protonymphal stages of both dermanyssines and
macronyssines, and those setae of the series which are suppressed at the protonymphal
stage do not appear in subsequent developmental stages. However, the main feature
of retardation in the development of the podonotal chaetotaxy in the deutonymphal
stage is most apparent in the suppression of certain setae which normally first appear
at this instar, for example, the para verticals (zi), 23, S2 and, particularly in the male,
z6 (Text-figs, ii A and c). The chief difference between the chaetotaxy of the male and
female non-feeding deutonymphs of the macronyssines is seen in the greater hyper-
trichy of the idiosoma in the region of the lateral and marginal series of setae in the
female.

In all the British members of the Dermanyssidae, the adults appear to retain the

3 Ondatralaelaps gen. nov. (type : Laelaps multispinosa Banks).

270

G. OWEN EVANS & W. M. TILL

FIG. ii. Ornithonyssus bacoti (Hirst). Dorsum of A. male deutonymph ; B. male ; c. female

deutonymph ; D. female.

THE BRITISH DERM AN YSSID AE (ACARI)

271

dorsal idiosomal chaetotaxy of the corresponding male or female deutonymph.
Males of the obligatory parasites show an increase in the area of dorsal sclerotization
compared with the female with the result that certain of the r series and of the z
series occurring on unsclerotized cuticle in the female are incorporated on the dorsal
shield in the male (Text-figs. IIB and D).

An exception to the basic dorsal chaetotaxy of the Dermanyssidae occurs in the
adults (the only known stages) of Pseudolaelaps doderoi (Berlese) whose systematic
position is problematical. The dorsum is markedly hypotrichous, the podonotum
bearing only 15 pairs of setae and the opisthonotum nine pairs.

Some intraspecific variability in the primary chaetotaxy of the opisthonotum is
apparent in many obligatory parasites and in some facultative parasites and takes the
form of the suppression of one seta of a pair or an asymmetrical arrangement of the
setae of a pair. This is particularly evident in members of the genera Dermanyssus
and Ondatralaelaps.

FIG. 12. Ornithonyssus bacoti (Hirst) . Tritosternum of larva (A), pro tony mph (B), deutonymph

272 G. OWEN EVANS & W. M . TILL

The dorsal setae in the majority of the Dermanyssidae are slender and smooth or
weakly pilose. Exceptions occur in the genera Stratiolaelaps and Cosmolaelaps where
they tend to become leaf-like or spatulate, and in facultative parasites of the Laelaps-
group in which they may be modified as short robust spines. The vertical setae (_;'i)
sometimes differ in form from the other setae in the region of the vertex.

Dorsal porotaxy: The basic adult and deutonymphal complement of " pores "
on the dorsum of the idiosoma appears to be 22 pairs (n podonotal and n opis-
thonotal) and their distribution in Laelaps is shown in Text-fig. 90. Considerable
diversity occurs in the pattern and number of " pores " throughout the family and
both hyper- and hypoporosity are not uncommon, particularly in the obligatory
parasites.

Tritosternum: The tritosternum4 lies between coxae I on the venter of the
idiosoma and is present as a biramous structure in all British Dermanyssidae (Text-
figs. 12, 15, 16) . Its function is unknown. In life, the laciniae lie along the capitular
groove and the fine processes (pilae) along their margins possibly engage the denticles
of the deutosternum. The basal portion of the tritosternum is longer than wide and
the laciniae in the free-living and facultatively parasitic forms are invariably pilose.
Many obligatory parasites (Dermanyssinae and Macronyssinae) have a transparent,
marginally denticulate border to the tritosternum (Text-fig. I2D). The non-feeding
larva and deutonymph of the Macronyssinae have the laciniae of the tritosternum
markedly reduced (Text-figs. I2A-D).

Sclerotization and chaetotaxy of the venter: At the larval stage in the free-
living forms and in most of the facultative parasites two sclerotized areas are present
ventrally ; a sternal shield, of varying degrees of sclerotization, inter-coxally and a
subtriangular anal shield surrounding the anus which is provided with a pair of valves
(Text-fig. I3A) . The sternal shield carries three pairs of setae (st. 1-3) and the anal
shield three setae, comprising a pair of paranals (pan.) and an unpaired postanal
(pon.). There are no euanal setae, that is, setae situated on the anal valves. Four
pairs of setae occur on the unsclerotized cuticle of the opisthogaster and, according
to the system proposed by Lindquist & Evans (1965), these consist of three pairs of
internal ventrals (Jvi, 2 and 5) and one pair of medio-lateral ventrals (Zvz) arranged
as in Text-fig. I3A. Stigmata, peritremes and podal sclerites are not present in the
larva. In the obligatory parasites there is a marked reduction in sclerotization to
the extent that only rarely can one define distinct sternal and anal shields. However,
the normal three pairs of sternal setae and the three setae associated with the anal

4 Van der Hammen (1964) has rejected the term tritosternum (for furca) on the ground that it is " no
sternum at all, because a sternum is a distinct plate ". However, he considers (without giving any
evidence) that the small sclerite occurring in the normal position of a tritosternum in some of the highly
specialized bat parasites of the family Spinturnicidae represents a " real sternal plate belonging to segment
III "! The reason for this must be, by inference, that the structure is a " plate " although, even in this
group, the tritosternum is not always a simple plate [cf. Meristaspis jordani (Radford)]. Perhaps the
tritosternum in some Trachytidae (Uropodina) will also satisfy his " criterion " of a sternum? At
present, the origin of the tritosternum is unknown but it seems unnecessary to change such a well-
established term, certainly on the " evidence " presented by van der Hammen.

THE BRITISH DERM AN YSSI D AE (ACARI)

273

FIG. 13. Semi-diagrammatic representation of the venter of the larva (A), pro tony mph (B),
deutonymph (c) and female (D) of Hypoaspis (Gaeolaelaps) aculeifer (Canestrini) .

274

G. OWEN EVANS & W. M. TILL

region are present, but there is a retardation in the development of the opisthogastric
setae, only one pair (Jvi) being developed in some Dermanyssinae and two pairs
(Jvi and 2) in the Macronyssinae (Text-fig. I4A).

The sternal and anal shields in the protonymph are generally well-defined and carry
the same complement of setae as in the larva (Text-fig. 133). A pair of genital setae
(sometimes present as incipient setae in the larva) and two pairs of lyriform fissures
(p. i and^>. 2) first appear at this instar. In the free-living forms the larval number
of setae on the opisthogastric cuticle is retained, but in the obligatory parasites the
larval deficiency in opisthogastric setae is rectified and the protonymph now acquires
its full complement with the appearance of Zv2 and Jv$ (Text-fig. 146). Hyper-
trichy of the opisthogaster may occur at this stage in some Macronyssinae, for
example, Macronyssus flavus (Kolenati). Stigmata with short, anteriorly directed
peritremes never extending beyond the anterior margin of coxae III are present
lateral to the third coxal interspace. No distinct podal shields occur at the proto-
nymphal stage.

Increased sclerotization and setation of the venter are evident at the deutonymphal
stage. The sternal shield, bearing the three pairs of setae and two pairs of fissures
present in the protonymph and a fourth pair of setae, the metasternals (st. 4) with
their associated pores, which first appear at this instar, occupies almost the entire

FIG. 14. Ornithonyssus bacoti (Hirst). Venter of larva (A) and protonymph (B

THE BRITISH DERM AN YSSI D AE (ACARI) 275

length of the intercoxal region. Its tapered, posterior region extends between the
genital setae (Text-fig. 130) . Hypotrichy of the sternal region has been observed in
some obligatory parasites, for example, certain species of Dermanyssus and Pellonys-
sus, and this is characterized by the absence of the metasternal setae which fail to
develop at their normal stage in ontogeny. Certain species of Haemogamasus, on
the other hand, show a distinct hypertrichy of the sternal chaetotaxy, the secondary
setae usually being shorter than the primary. The anal shield more or less retains
the form present in the larva and protonymph.

Considerable variation occurs in the chaetotaxy of the opisthogaster in the deuto-
nymph. Normally, the holotrichous condition for this region is seven pairs of setae
(Zvi, Jv3 and Jvq. added to the protonymphal complement) and this occurs in certain
of the free-living forms, for example, some species of Hypoaspis s. lat., Ololaelaps,
Pseudoparasitus and Stratiolaelaps. More commonly, however, the region shows
some degree of hypertrichy, particularly in the Eulaelaps-Haemogamasus group and
in the obligatory parasites. As in the case of the marginal setae of the dorsum, the
male deutonymph often has fewer opisthogastric setae than the female deutonymph .
Further, there may be considerable intraspecific variability in the number of opistho-
gastric setae in deutonymphs of the same " sexual phase ". The stigmata are
situated in the same position relative to coxae IH and IV as in the protonymph, but
in all free-living forms and the majority of the facultative parasites the peritreme is
considerably longer and usually extends up to or beyond coxa I. Exceptions to this
occur in certain of the obligatory parasites (Dermanyssus, Steatonyssus, Pellonyssus,
Ophionyssus) where the protonymphal form of the peritreme may be retained at the
deutonymphal stage. In the free-living forms a weakly developed peritrematal
shield is also present and its post-stigmal portion bears a conspicuous pore (Text-
fig. 130). Podal shields are absent except for a weakly sclerotized strip along the
posterior border of the coxal cavity of leg IV. Weakly sclerotized metapodalia may
be present. An anal shield with the normal three setae is characteristic of the family
although in densely hypertrichous forms (Haemogamasus} it may carry a number of
secondary unpaired setae.

With the exception of the hypertrichous forms showing intraspecific variability of
the opisthogastric setae, the adult setal complement of the venter is determined at the
deutonymphal stage (male or female) . The degree of sclerotization and the size and
outline of the shields at the adult stage, however, show great diversity, greater in fact
than within any other family of the Mesostigmata. In the female of the free-living
forms and the facultative parasites the sternal shield normally bears three pairs of-
setae (st. 1-3) and two pairs of lyriform fissures (p. I and^>. 2). It is fused laterally
with the endopodal shields of coxae II. More rarely it bears, in addition, the meta-
sternal setae (st. 4) and their associated pores (p. 3) as in Ololaelaps (Text-fig. 150).
The posterior margin of the sternal shield may extend as far as the level of the pos-
terior margin of coxa III, as in G. aculeifer. Its anterior margin is often indistinct
owing to the sclerotization of the cuticle immediately anterior to the shield. This
pre-sternal area of sclerotization may be uniformly sclerotized and reticulated (Text-
fig. 153) or regionally more heavily sclerotized to form pre-endopodal sclerites (Text-

276

G. OWEN EVANS

W. M. TILL

ant.sp.

ant. sp.

FIG. 15. Venter of female of : A. Laelaps agilis Koch ; B. Myonyssus gigas (Oudemans)
c. Ophionyssus natricis (Gervais) ; D. Ololaelaps placentula (Berlese).

THE BRITISH DE RM AN YSSID AE (ACARI) 277

fig. 150) . We have not observed true jugularia in the British members of this family.
The surface of the sternal shield is variously ornamented, the ornamentation being in
the form of a reticulate pattern of lines, or punctate lines, or scattered punctures.
The metasternal setae and pores are normally situated posterior to the sternal shield
and lie on striated cuticle or on small metasternal shields which may be free or fused
with the endopodal shields of coxae III. In the obligatory parasites the sternal
shield is less well developed and rarely fused with the endpodal shields of coxae II.
It may bear only two pairs of setae (st. I and 2) and pores (p. I and 2) or the normal
three pairs of setae and two pairs of pores. In outline it is generally sub-rectangular
but in some Pellonyssus it is reduced to a narrow, transverse, band-like sclerite.
Subcircular " porose " areas occur between sternal setae i and 2 in some species of
Macronyssus. The metasternal setae in this group are usually situated on striated
cuticle. They are absent in the adults whose deutonymphs lack these setae.

The female genital orifice (with its apodemes) appears as a transverse slit at the
level of the fourth coxae. Associated with this orifice, and extending posterior to it,
is a characteristic genital shield bearing the genital setae. Anteriorly the shield has
a hyaline flap of varying degrees of development. In its least specialized form the
genital shield is more or less flask-shaped and bears only the genital setae (Text-fig.
130) but, with the exception of the obligatory parasites, there is a general tendency
for an increase in the size of the shield and the incorporation with it of a number of
opisthogastric setae. Such a shield is termed genito-ventral (Text-figs. I5A-B).
Marked hypertrichy of the genital shield occurs in Eulaelaps, Haemogamasus and
Myonyssus. The genital shield shows some reduction in size in the obligatory
parasites and tapers in its posterior half. In some macronyssines the reduction in
size and the posterior tapering of the shield is very marked ; the resulting shield is
slender and does not carry the genital setae (Text-fig. 150). The anterior hyaline
flap is usually extensive in these obligatory parasites and its tapering anterior region
may extend beyond the anterior margin of the sternal shield.

All the British dermanyssids have an anal shield bearing the three setae normally
associated with the anus. It is typically sub-triangular or pear-shaped. An
exception occurs in the genus Myonyssus (Text-fig. 155) in which it is considerably
broader than long. The region posterior and lateral to the postanal seta is invariably
provided with aciculae. Secondary unpaired setae occur on the anal shield of some
members of the genus Haemogamasus.

The ventral sclerotization of the female of Ololaelaps is exceptional in that there
is a fusion of the genito-ventral, metapodal and anal elements to form a compound
shield occupying almost the entire area of the opisthogaster (Text -fig. 150).

The endo- and exopodal shields form a more or less complete framework enclosing
coxal cavities II-IV in many of the free-living forms and facultative parasites. In
Pseudoparasitus, Ololaelaps, Laelaspulus and some species of Holostaspis and Gymno-
laelaps, the podal elements behind coxae IV become greatly enlarged (Text-fig. 150).
Varying degrees of reduction of the exopodal shields occur in the genera Hypoaspis,
Haemogamasus and Laelaps and in the obligatory parasites only fragments of the
podal shields remain, the most conspicuous being the one behind coxa IV.

278

G. OWEN EVANS & W. M. TILL

sf-q. sh

v -a. sh.

.an. sh.

D C

FIG. 16. Venter of male of : A. Hypoaspis (Gaeolaelaps) aculeifer (Canestrini) ;
B. Hyperlaelaps microti (Ewing) ; c. Macronyssus sp. ; D. Ophionyssus natricis
(Gcrvais).

THE BRITISH DERM AN YSSI D AE (ACARI) 279

Metapodal shields are variable in form and number. In most free-living forms
they are small, oval, elliptical or elongate bodies, but in Eulaelaps they become large
subtriangular structures. Obligatory parasites usually have only one pair which is
weakly sclerotized and difficult to discern. The metapodals may fuse with the
genito-ventral shield as in Ololaelaps (Text-fig. 150).

Stigmata and peritremes are of basically the same form as in the deutonymph.
The peritrematal shield is fused with the dorsal shield anteriorly, except in some
obligatory parasites. Posteriorly it may be free or fused with the podal shield in the
region of coxa IV (Text-figs. i5A-c).

Typically, the males have a holoventral shield formed by the fusion of the sternal,
metasternal, endopodal, genital, ventral and anal shields. It carries the four pairs of
sternal setae, the genital setae, the three setae associated with the anus and a variable
number of opisthogastric setae (Text-fig. i6A). In the free-living forms and the
facultative parasites it is generally expanded posterior to coxae IV, but in some
macronyssines it shows little or no expansion in this region. Deviations from the
typical holoventral shield are apparent in some facultative and obligatory parasites.
Hyperlaelaps, for example, has a compound sternito-genito-ventral shield and a
separate anal shield (Text-fig. i6B). Further reduction of the ventral sclerotization
occurs in Ophionyssus natricis (Degeer), the sclerotized areas comprising a sternito-
genital and a separate anal shield (Text-fig. 160). Certain species of Macronyssus
show another variant in having a sternito-genital and a separate ventro-anal shield
(Text-fig. i6c). The genital orifice, closed by a single valve, is always situated near
the anterior margin of the " sternal shield " and is flanked by the first pair of sternal
setae.

Legs

Segmental chaetotaxy: Without exception the legs are seven-segmented, the
terminal segment being represented by the ambulacrum. The segments, from coxa
to tarsus, have a well-defined chaetotaxy and the system of nomenclature and
formulae for the segmental chaetotaxy follows that proposed by Evans (1963). This
system, based on the observed ontogenetic development of the leg chaetotaxy in over
100 species of Gamasina and which has subsequently been found to be applicable to
the Uropodina and Antennophorina, has been criticized by van der Hammen (1964)
who, with little success, attempted to apply Grand] cans' nomenclatural system for
the leg chaetotaxy of the Oribatei and allied Actinochaeta to an adult macrochelid
mite. His criticisms are : (a) that the existence of a basic whorl of setae on a seg-
ment was ignored ; (b) that no attention was paid to the presence of the unpaired
dorsal seta, d, and (c) that the segmental formulae are of little or no comparative
value. Are these criticisms of the Evans system valid in the light of the observed
segmental chaetotactic patterns on the legs of the Mesostigmata? In our opinion
they are not, for the following reasons :

Grandjean (1940) considers the basic element of the segmental chaetotaxy to be a
ring or whorl of setae consisting of a maximum of seven (oribatologists refer to this
condition as " primitive "), namely, an unpaired dorsal, two pairs of laterals and one

280

G. OWEN EVANS & W. M. TILL

pair of ventrals. The existence of a whorl of setae on the segment was not ignored
by Evans and is implicit in his segmental formula

av j

which van der Hammen overlooked and instead based his criticism on a simpler
formula

(, dorsals , A
al — r- — $1]
ventrals /

which was adopted only in those cases where the extreme non-cylindrical form of the
segments (the femora and trochanters) made it impossible to distinguish the anterior
and posterior members of the dorsal and ventral series. As regards the presence of
an unpaired dorsal seta, there is no evidence from observed segmental patterns that
such a seta exists on any leg segment (excluding perhaps tarsus I which has not been
studied in detail) of the larva or any segments, excluding the tarsi, in subsequent
developmental stages of the Gamasina and Uropodina. Van der Hammen's state-
ment that Evans " did not pay attention to the definition of the unpaired dorsal
seta d. . . " is contrary to the facts and one wonders whether the author has read

FIG. 17.

Diagrammatic representation of the setal positions on a leg segment of a
dermanyssid mite.

THE BRITISH DE RM AN YSSID AE (ACARI) 281

beyond the " introduction " in Evans' 1963 paper! In this paper attention is drawn
to the appearance on tarsi II-IV of an unpaired dorsal and ventral seta at the
protonymphal and deutonymphal stages respectively, and these are referred to as
medio-dorsal (md) and medio- ventral (mv). If the observed position of these
unpaired setae reflects their primary position on these segments, then the maximum
complement of a segmental whorl in the Mesostigmata would be eight setae as shown
diagrammatically in Text-fig. 17. Owing to a regression in the dorsal and ventral
setae, the maximum complement of a whorl is apparently not present on other seg-
ments of the legs.

Van der Hammen's attempt to define the unpaired dorsal seta on the leg segments
is characterized by such statements as " impossible to conclude whether the three
plumose setae with a dorsal position are really d", " possibly dorsal plumose setae "
and " probably dorsal setae ". This uncertainty is inevitable and arises from an
attempt to make the chaetotactic patterns fit a pre-conceived system. It is sur-
prising that van der Hammen has used " probably dorsal setae " only in reference to
certain setae on the dorsal surface of femur II, one of the segments showing an
extreme non-cylindrical form which affects the primary chaetotactic pattern of the
segment. In those forms which have a less specialized femur II, that is, the segment
is more or less cylindrical, the longitudinal arrangement of the setae on its dorsal
surface resembles that of genu II for which van der Hammen finds it impossible to
define unpaired dorsal setae! The " migration " of setae from their normal (primary)
position relative to other setae on a segment is common and is not only due to the
asymmetrical development of the segment. More often the " migration " occurs as
the result of the absence of one member of a " pair " of setae on the segment. The
remaining seta often develops in an intermediate position along an imaginary line
which would have connected the bases of the " pair ". For example, four setae
(avlt avz, pvlt pv2) whose bases form the corners of a rectangle often occur on the
ventral surface of tibia I in the Gamasina, but in some groups pvz is lacking and the
remaining member of the pair (av2) may migrate to occupy a median position on the
segment at the level of the line joining the bases of avz and PVZ in the forms with four
ventral setae. Such deviations from the normal distribution of setae on a segment
add to the difficulty of applying any system of setal nomenclature based on" positional
homology ".

We reject van der Hammen's criticism of the segmental formulae which have
proved to be of considerable practical value in referring to the segmental chaetotaxy.
This author's alternative formula, which he considers " more logical " (?), depends on
the recognition of unpaired dorsal setae and is completely impracticable.

It seems appropriate, in the present context, to refer to another fundamental
difference between the system of nomenclature proposed by Evans and that used for
the Actinochaeta. Although in both systems the positions of setae are defined in
relation to certain faces of the segment, the lateral faces in the Grandjean system are
defined according to the natural position of the limb relative to the longitudinal axis
of the body, the anteriorly directed first two pairs of legs and the posteriorly directed
legs III and IV having paraxial and antiaxial faces. In the system used in the Meso-

282 G. OWEN EVANS & W. M. TILL

stigmata, on the other hand, the definition of the lateral faces (anterior and posterior)
is not based on the natural, but on the artificial position of the limb, that is, when it is
extended laterally at right angles to the longitudinal axis of the idiosoma. Adopting
Grand] can's terminology for the lateral faces of the legs in the Mesostigmata would
necessitate referring to the anterior and posterior faces of legs I and II as paraxial
and antiaxial respectively, since these legs are directed anteriorly, retaining anterior
and posterior for legs III which are directed laterally, and referring to the anterior
and posterior faces of legs IV as antiaxial and paraxial respectively, since these limbs
are directed posteriorly. In comparison with the existing system of setal terminology
in the Mesostigmata, the advantages gained, if any, by basing the system on two (or
three) different terminologies for the lateral faces of the limbs, would be at the ex-
pense of simplicity and practicability.

The segmental chaetotaxy of the legs in the larval dermanyssids we have examined
is constant and normal for the Gamasina, as defined by Evans (1963). At the
protonymphal stage one seta is added to the ventral surface of femur II and one
medio-dorsal seta to each of tarsi II-IV, giving the following chaetotaxy : —

I II III IV

2 i

I — \ — o I — \ — u

i — f — o i — § — o

coxa

trochanter

. i — § — i

i-i-i

femur .

. 2—1—2

i-f-i

genu .

• i-!>!-i

x-§.fr-

tibia

.I-f,f-I

I-M-

tarsus5.

—

o 313

o — ^r> r>> ^r~

i— i, f— i i— \, f— i

3— f , £, §—3 3— |, £, §—3 3— f , i, §—3

Genu IV in the protonymphae of Ophionyssus and Sauronyssus, is exceptional
in having six setae (i — §, § — i).

The deutonymphal and adult6 segmental complements in the free-living forms, for
example, Ololaelaps, Pseudoparasitus, some Hypoaspis s. lat. and Cosmolaelaps is
extremely constant. The segmental chaetotaxy may be summarized as follows : —

III

coxa

trochanter

. I — ij — I

i-*-x

I— I— 0

I— 1—0

femur .

25 />
—5 2

2-f-I

I-f-I

i-f-i

genu .

<» . 3 3 o

2 3. 1 2

2-f,f-I

2-f , §-I

tibia .

233 /o
2"' T — Z

? 2 2 ^
2 TJ T 2

2-i,f-I

/7 1 3 o

a ]• T T

tarsus5.

3313 o
"£> T» V 0

3 — f » T» 1 3

3313 /-
~"^r- T ^"~ -"

Certain nest-inhabiting forms and facultative and obligatory parasites show con-
siderable diversity in the segmental chaetotaxy of the legs at the deutonymphal and
adult stages and these are discussed below. In the semi-diagrammatic representa-
tions of the segmental chaetotaxy in Text-fig. 18 the patterns marked (a) are con-
sidered to be the normal types for the family.

5 The chaetotaxy of tarsus I has not been studied in detail.

• The deutonymphal complement of setae is invariably retained by the adult.

THE BRITISH DERM AN YSSID AE (ACARI)

283

III

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FIG. 1 8. Semi-diagrammatic representation of the chaetotaxy of selected segments of the legs

of the Dermanyssidae.

284 G. OWEN EVANS & W. M. TILL

Coxae I-IV: In the larva coxae I-III each carry two setae and this condition
is retained in succeeding developmental stages. Coxa IV bears a single seta in post-
larval instars. The coxal setae are normally simple but in many of the facultative
parasites (Laelaps-group) certain of them, particularly on coxa I, become stout spines
or spurs as the result of hypertrophy. Processes of non-setous origin may also
occur on certain of the coxae. For example, in many facultative or obligatory
parasites a stout, spine-like structure develops on the distal margin of the anterior
face of coxa II (Text -figs. 15 A and B) and in Hirstionyssus stout spur- or ridge-like
processes are present on coxae II and III and sometimes IV. The distal margin of
the coxa is produced into a dorsal and a ventral condyle.

Trochanters I-IV : Trochanters I-III at the larval and protonymphal stages and
trochanter IV in the protonymph each have four setae. In the deutonymphs and
adults trochanter I normally carries six setae and trochanters II-IV five setae. An
exception occurs in Dermanyssm alaudae (Schrank) which has only five setae (i — § — i)
on trochanter I.

Femora I-IV (Text-fig. 18) : Three variants of the normal complement of setae
(2 — f , §-2) of femur I have been observed. This segment (fig. b) in Laelaspulus
flexuosus (Michael) shows an addition of one antero-ventral seta to the normal
number, giving the formula (2 — f , f — 2). Many obligatory parasites [Dermanyssus
chelidonis Oudemans, D. hirundinis (Hermann) and D. quintus Vitzthum, Sauronys-
sus saurarum (Oudemans) and Ophionyssus natricis (Deeger)] show a decrease in the
number of ventral setae to three (fig. c) by the suppression of one postero-ventral
seta (2 — f , f — 2). D. alaudae is exceptional in retaining the larval complement of
setae (2 — f, f — 2) on this segment throughout ontogeny (fig. d). No setae occur
proximal to the circumsegmental fissure.

Exceptions to the normal pattern (2 — f , f — i) of femur II have been observed in
Haemogamasus hirsutosimilis Willmann (2 — f , § — i) which has an additional postero-
ventral seta (fig. b), in 0. natricis, D. quintus and 5. saurarum which lack al2 (i — f ,
| — i), fig. c, and in D. alaudae which differs from the latter group in having only two
setae in the antero-dorsal series (i — f , f — i), fig. d. Seta al2> when present, is situa-
ted proximal to the circumsegmental fissure.

The normal complement of setae on femur III is six (i — ^ — i), of which the
antero-ventral is situated proximal to the circumsegmental fissure. Although this
number of setae also occurs on femur III in Pellonyssus reedi (Zumpt and Patterson)
the ventral seta is situated distad of the fissure (fig. c) . Two other types of chaetotaxy
have been observed ; in H. hirsutosimilis two ventral setae are developed (i — f — i)
fig. b, whilst in D. alaudae, chelidonis and hirundinis, and in 5. saurarum, the larval
complement (i — f — o) is retained throughout ontogeny (fig. d).

The number of setae on femur IV is remarkably stable and the only exception we
have found to the normal pattern (i — f — i) is in D. alaudae which has only five setae
(i-f-o) fig. b.

Genua I-IV (Text-fig. 18) : The apparent great diversity in the segmental chaeto-
taxy of the genua in the Dermanyssidae is somewhat misleading since the majority
of the " variants ", except on genu IV, occur in a single genus, Dermanyssus.

THE BRITISH DE RM AN YSSID AE (ACARI) 285

Exceptions to the normal chaetotaxy (2 — f , f — 2) of gemi I are found, with the
exception of Pseudolaelaps doderoi Berlese, only amongst the obligatory parasites.
In S. saurarum, 0. natricis and D. chelidonis, the setation is deficient in one postero-
dorsal member (2 — f , f— 2, fig. b) whilst in P. doderoi and in some populations of
0. sylviamm an antero-ventral seta is lacking (2 — f, f — 2, fig. c). The remaining
three types of chaetotaxy have been observed in D. hirundinis (2 — f, f — 2, fig. d),
D. alaudae (2 — f, f — -2, fig. e) and Z). quintus (2 — f, f — i, fig./).

The normal pattern (2— f , f — 2) of genu II occurs in all the British dermanyssids
except P. doderoi and some species of Dermanyssus . In P. doderoi the antero-ventral
seta is lacking (2 — §, f — 2) fig. 6) whilst in D. chelidonis the antero-dorsal series is
unideficient (2 — f, f — 2) fig. c.). Further specializations are seen in D. quintus
(2 — f, -f — i, fig. d) and in D. alaudae which retains the larval complement of setae
(i — §, g — i, fig. g) throughout ontogeny. The chaetotaxy of this segment shows
considerable intraspecific variation in D. hirundinis (2 — f , f — i, i — f , f— i, fig. e
and i— f, \— i, fig./).

Five different chaetotactic patterns have been observed on genu III, excluding the
normal (2 — f, f- — i). Thus, members of the genera Ophionyssus and Pellonyssus
and Ornithonyss^ls bacoti (Hirst) and 5. saurarum have ten setae(2 — f , f— 2, fig. b)
on this segment ; D. chelidonis and 0. sylviarum have nine setae, but differ from the
normal form in lacking PVI and having two postero-laterals (2 — f, § — 2, fig. c) ;
Hyperlaelaps amphibia Zachvatkin and H. microti (Ewing), and P. doderoi bear 8
setae (2 — f, § — i, fig. d), D. hirundinis has seven setae (i — f, § — i, fig. e) and
finally, D. alaudae retains the larval complement (i — §, § — -i, fig./).

Genu IV shows the greatest diversity in chaetotaxy of all leg segments in the
British Dermanyssidae. In addition to the normal pattern (2 — f , § — i), six other
types of chaetotaxy have been observed, namely : —

fig. b, (2 — f, f — 2) in Pellonyssus, Laelaps agilis Koch, 0. bacoti and 0.

natricis.

fig. c, (2 — f , $• — 2) in Laelaps echidnina, hilaris, muris ; Hyperlaelaps
amphibia and microti ; Eulaelaps stabularis and nova ;
Androlaelaps casalis and fahrenholzi ; Hypoaspis (G.)
sardoa ; Ondatralaelaps multispinosa and Laelaspulus
flexuosus.

fig. d, (2 — f , f • — i) Pneumolaelaps , Hypoaspis (G.) lubrica ; Melittiphis
alvearius ; Macronyssus ; Steatonyssus periblepharus and
Ornithonyssus sylviarum.
fig. e, (2 — f, f- — 2) in Sauronyssus saurarum
fig. /, (2 — f , § — 2) in Dermanyssus chelidonis
fig. g, (i — f , § — i) in Dermanyssus alaudae, hirundinis and quintus.

Tibiae I-IV (fig. 18) : With the exception of P. doderoi, deviations from the normal
setal pattern of tibia I (2 — f , f — 2) have been noted only in the obligatory parasites.
Three of the " variants " occur in the genus Dermanyssus, namely, (2— f , f — 2, fig. d)
in D. chelidonis and hirundinis; (2 — f , f — 2, fig. e} in D. alaudae and (2 — f , f — i) fig.

286 G. OWEN EVANS & W. M. TILL

/) in D. quintus. The remaining two types (2 — f , f — 2, fig. b) and (2 — f , f — 2, fig. c)
have been found in Pellonyssus, Sauronyssus saurarum and Ophionyssus natricis, and
in P. doderoi and some forms of 0. sylviarum, respectively.

The only exceptions to the normal segmental chaetotaxy of tibia II (2 — f , f — 2)
occur in Dermanyssus alaudae, chelidonis, hirundinis and quintus which retain the
larval complement of setae throughout ontogeny (i — \, f — i, fig. b).

In addition to the normal form (2 — y, f — i), the following three types of chaetotaxy
are found on tibia III :

fig. b, (2 — f , f — 2) in the ethiopian species Steatonyssus tibialis Till and

Evans.

fig. c, (2 — y, f — 2) in Pellonyssus , Steatonyssus (except tibialis} , Or nithonyssus
bacoti and sylviarum ; S. saurarum, 0. natricis,
Melittiphis alvearius, Hypoaspis (G.) sardoa and
Laelaspulus flexuosus.

fig. d, (i — -}, f — i) in Hyperlaelaps amphibia and Dermanyssus alaudae,
chelidonis, hirundinis and quintus. This is the larval
pattern.

Only in the obligatory parasites have we observed variants of the normal chaeto-
taxy of tibia IV (2 — \, f — 2). S. tibialis has eleven setae on this segment (2 — f , f — 2,
fig. b), S. saurarum nine setae (2 — y, f — 2, fig. c) and Dermanyssus alaudae, chelidonis,
hirundinis and quintus have the protonymphal complement of seven setae (i — T, T—

/-/F : The chaetotaxy and the form of the distal sensory region of tarsus I
have not been studied in detail in the present work. The ontogenetic development
of the chaetotaxy of tarsi II-IV is extremely constant, the basic adult complement of
eighteen setae (3 — f , T, f — 3) being present in the majority of the species. In the
obligatory parasites there is a tendency for the reduction in the size of setae ad1
and pd± which may be represented by microsetae or be lacking.

Ambulacra (Text-figs. IQA-D) : Little attention has been paid previously to the
structure of the pretarsus and the ambulacrum in the Mesostigmata. The following
account of the structure and mode of functioning of the ambulacral apparatus is
based on that of Haemogamasus hirsutus Berlese.

The striated, sclerotized components of the pretarsus form a more or less rigid
sheath. In the distal half of the pretarsus the sclerotized elements of the floor are
thickened medially to form two slender, internal, longitudinal struts, the median
pretarsal apodemes (ptr. apod.). Anterior to the apodemes lies a well-defined,
dorsally directed U-shaped basilar sclerite (bas. scl.}. It is pivoted by its base to
the anterior processes of the pretarsal apodemes. A tendon (basilar tendon,
Text-fig. 190) is attached to the basal region of the basilar sclerite but its origin
is difficult to trace in whole mounts of the tarsus and pretarsus. It appears to
run ventrally through the tarsus and to be connected to a muscle originating in
the tibia. A second unpaired sclerotized structure, the pulvitractor sclerite (pulv.
scl.} is situated medially within the pulvillar mass immediately anterior and

THE BRITISH DERM AN YSSID AE (ACARI)

287

dorsal to the basilar sclerite. In H. hirsutus it forms a dorsally directed hook-like
process with a bifid base. Its basal limbs are movably articulated with the dorsally
directed arms of the basilar sclerite. The paired lateral claws, each with a deep
proximo-internal acetabulum, appear also to be articulated with the limbs of the
pulvitractor and/or basilar sclerite. Lying dorsal to the apodemes and sclerites is a
strong median tendon (pulvitractor tendon) connected distally with the pulvillus and
by way of the pulvillus to the dorsal hook of the pulvitractor sclerite. It runs the
entire length of the pretarsus and is connected to a strong retractor muscle originating
within the basi-tarsus. Near the junction of the pretarsus and tarsus the tendon
passes between the limbs of a dorsally directed Y-shaped apodeme (pulvitractor
guide). The lateral elements of the pretarsal sheath are produced distally into
denticulate lobes which Krantz (1962) has referred to in the Macrochelidae as the
*' laterodistal pretarsal elements ". In view of their apparent function we propose
renaming these lobes the pretarsal opercula (ptr. op.). The true form of the pulvillus

bas.tend,

\ A

tar.
bas.tend.

FIG. 19. Ambulacrum and pretarsus of Haemogamasus hirsutus Berlese. A. Dorsal view
from a compressed preparation ; B. partially retracted ambulacrum in lateral view ;
c. fully retracted ambulacrum in lateral view ; D. Basilar sclerite with tendon.

288 G. OWEN EVANS & W. M. TILL

is difficult to ascertain since it is invariably distorted during preparation for study.
From compressed mounts it appears to comprise ventral, dorsal and lateral elements.
It is provided with weakly sclerotized " veins " (Text-fig. IQA).

The function of the tarsal muscle and its tendon is to withdraw the ambulacrum
into the pretarsal sheath and this is made possible by the nature of the articulations
between the apodemes of the pretarsus and the basilar sclerite, and between the limbs
of the basilar sclerite and the pulvitractor sclerite. As the muscle contracts the
backwardly directed pull of the pulvitractor tendon on the pulvillus and on the
pulvitractor sclerite causes the pulvillus and the claw-complex (basilar and pulvi-
tractor sclerites and claws) to be retracted into the pretarsal sheath (Text-fig. IQB).
At maximum retraction the basilar sclerite and the pulvitractor sclerite lie ovei the
rigid median pretarsal apodemes with the hook of the pulvitractor sclerite directed
ventrally and the claws directed anteriorly (Text-fig. 190). Also at this stage the
anterior margins of the pretarsal opercula meet and together form a protective
" lid " to the pretarsal sheath.

The mode of extension of the ambulacral apparatus will remain somewhat prob-
lematical until the definite origin of the tendon of the basilar sclerite has been eluci-
dated. If, as it appears, it is the tendon of a tibial protractor muscle, then contrac-
tion of this muscle would have the effect of pushing forward the basilar sclerite on its
pivot when tension through the pulvitractor tendon is released. Thus, the ambulac-
rum would be operated by a pair of antagonistic muscles, the pulvillar retractor and
the basilar protractor.

The ambulacral apparatus has this basic form throughout the Dermanyssidae
although there is considerable diversity in the shape of the sclerotized structures,
particularly the basilar and pulvitractor sclerites and the pretarsal opercula, and in
the degree of the development of the claws. Normally in the dermanyssines paired
claws are present on all legs and those of legs II-IV are usually stronger and larger
than those of leg I which is chiefly sensory and not ambulatory in function. Claws
are absent or strongly reduced on leg I of Stratiolaelaps miles Berlese and Eulaelaps
nova Vitzthum and on legs I-IV of Ondatralaelaps multispinosa (Banks) and
Laelaps muris Koch.

Form of the leg setae: The leg setae are generally simple or pilose in form but
throughout the family there is a tendency for certain setae, particularly the ventral
setae of the femora, genua and tibiae of leg II and the distal setae of tarsi II-IV to
enlarge and form stout spine- or spur-like structures (Text-fig. 200). Hypertrophy
of setae on leg II is not restricted to the male as is usual in the Gamasina. Seta av1
on femur II in both sexes of certain Androlaelaps may be affected and, to a lesser
extent, seta av^ on the genu and tibia of this leg. In Hypoaspis (Gaeolaelaps] nidi-
corva, on the other hand, seta avz of femur II hypertrophies (Text-fig. 20A). On the
tarsi, setae allf pl^ av± and pv± usually show some enlargement in comparison with
other tarsal setae and in Eulaelaps nova setae adz, pv2 and mv on tarsus II are also
spur-like (Text-fig. 20B). Some members of Hirstionyssus have the tarsal setae av-^
and PV-L developed into stout hook-like structures.

THE BRITISH DERM AN YSSID AE (ACARI)

289

Certain coxal setae are often enlarged in the Laelaps-group and in many of the
facultative parasites seta adl and to a lesser extent, seta pdt on femur I and II are
considerably stronger and longer than other femoral setae. The relative increase in

FIG. 20. A. Femur II, anterior view, of Hypoaspis (Gaeolaelaps] nidicorva sp. nov. female ;
B. dorsal view of tarsus II of Eulaelaps nova Vitzthum, female ; c. anterior view of
leg II (femur to tarsus) of Hypoaspis (H. )krameri (Canestrini) male ; D. dorsal view of
leg IV (femur to tarsus) of Hypoaspis (H.) krameri (Canestrini).

2QO

G. OWEN EVANS & W. M. TILL

length of specific leg setae is also apparent in Hypoaspis (H.) krameri, for example, on
leg IV seta ad± on the femur and setae ad2, pdz and pd3 on the tarsus are long and whip-
like (Text-fig. 20D).

Cuticular processes of non-setous origin: Reference has already been made
to the occurrence of setiform or spiniform cuticular processes on the chelicerae at the
junction of the second segment and the arthroidal membrane at the base of the
movable digit. Such processes are by no means restricted to the trophic appendages
but commonly occur at the distal margins of certain pedipalpal and leg segments.
Generally, they are better developed on the basal segments of the appendages,
particularly the coxae. The processes show considerable diversity in form and num-
ber. In their least specialized form (Text-fig. 21 A), they appear as a more-or-less

a.m.

,a.m.

FIG. 21. A. Coxa I of Haemogamasus hirsutus Berlese, dorsal view, showing processes on
distal margin of segment ; B. processes free from distal margin of segment ; c. processes
fused with distal margin of segment ; D-F. processes on distal margin of anterior (paraxial)
face of coxa II in D. Haemogamasus hirsutus Berlese ; E. Laelaps agilis Koch ; F. Mac-
ronyssus flavus (Kolenati).

THE BRITISH DE RM AN YSSID AE (ACARI) 291

complete circumsegmental fringe of spines on the distal margin of the segment. The
spines appear to have their origin on the arthrodial membrane in close proximity to
the distal margin of the segment (Text-fig. 2IB), but by the sclerotization of the
narrow strip of membrane between the segment and the processes, they become
incorporated with the segment (Text-fig. 2ic) . Reduction in the number of processes
is common to the extent that only one, two or three spines of the circumsegmental
fringe remain and these are usually located on the anterior face of the limb. The
remaining process (or processes) is better developed than the individual members of a
circumsegmental fringe. It is possible to trace the regression in the number of
marginal processes on the anterior face of coxa II in the Dermanyssidae. In free-
living species, for example members of the genera Ololaelaps and Pseudoparasitus,
the fringe is composed of spine-like processes of approximately equal size, but in
certain of the facultative parasites a medial process of the fringe hypertrophies as in
Haemogamasus hirsutus (Text-fig. 2ID). A similar hypertrophy of a medial process
occurs in Laelaps agilis but this is also accompanied by a reduction in the number and
size of the remaining processes (Text-fig. 2IE). Finally, in certain obligatory para-
sites (Macronyssus and Steatonyssus) only the hypertrophied medial process remains
and this forms the stout anterior coxal spine (Text-figs. I5A and B, 2ir).

Van der Hammen (1964) has referred to the cuticular processes on the distal margin
of the external face of the basis capituli and on the antero-distal margin of coxa I
as the latero-coxal setae and has unhesitatingly homologized them with the latero-
coxal spines e an ei in the Actinochaeta. These are not setae and each represents a
cuticular process originating from the junction of the segment and the arthrodial
membrane as described above. In many forms of Mesostigmata a fringe of spines
occurs on the distal margins of the lateral face of the basis capituli to which the
pedipalp trochanter is articulated and throughout the group there is a tendency for a
reduction in the number of spines. To designate such a single process, which is rarely
present in the group, a latero-coxal seta, seems unwarranted. There is no available
evidence upon which to base the homology of the cuticular processes in the Meso-
stigmata with the latero-coxal spines of the Actinochaeta and we deprecate the
application of the symbols e and ei to any of the cuticular processes associated with
the pedipalpal " coxae " or the coxae of leg I, respectively.

Rows of spine-like cuticular processes also occur on the unsclerotized pre-sternal
cuticle and regionally on the leg segments and basis capituli of many facultative
parasites such as Haemogamasus. Spur-like structures, not hypertrophied setae, are
present on certain of the coxae of Hirstionyssus. The functions of these various
cuticular processes are not known.

Lyriform fissures : Fissures occur on the femora and tarsi of legs I-IV. Two
lyriform fissures in the basal third of each femur are connected and form a complete
or interrupted circumsegmental fissure which secondarily divides the segment into
proximal and distal sections (Text-figs. 20A and c). The fissures on the basal third
of the tarsi similarly divide these segments into two regions, but the circumsegmental
fissure on tarsus I is usually less well developed than on the other tarsi. For example,
in many obligatory parasites the fissure is interrupted dorsally and the isolated

292 G. OWEN EVANS & W. M. TILL

proximodorsal fissure assumes the characteristic form of a lyriform fissure. A dorsal
lyriform fissure, distinct from those forming the circumsegmental fissures, is also
present on tarsi II-IV (Text -fig. 2OB).

DISCUSSION

Members of the family Dermanyssidae show considerable diversity in morphology
and biology largely as the result of the specializations exhibited by the parasitic forms
which have become adapted to life in the great variety of niches afforded by their
hosts. Within the family the free-living species, on the one hand, and the obligatory
parasites, on the other, appear to form the extremes of a morphological continuum,
the connecting link between them being provided by the morphological characteristics
of nest inhabitants and facultative parasites. Species of these extreme groups have
different morphological properties : the free-living forms show little diversity in
morphology in comparison with the obligatory parasites which display marked
morphological plasticity. This morphological plasticity is evident in the grades
of specialization found amongst congeners, for example the degree of specialization
of the gnathosoma in Haemogamasus , and in the higher incidence of intraspecific
variability in morphological " characters ".

The most spectacular morphological adaptations are associated with changes in
feeding habits, and may be seen in the specialization of the trophic appendages and
associated gnathosomal structures and in the reduction of idiosomal sclerotization in
the monophagous obligatory parasites. Taxonomic characters based on these
adaptive morphological structures have long played a dominant role in the supra-
generic classification of the family. Less spectacular, but nevertheless of consider-
able taxonomic importance, are the differences in chaetotaxy between the free-
living and parasitic forms. Deviations from the normal adult chaetotaxy, particu-
larly of the segments of the pedipalps and legs, are due to a retardation in the normal
ontogenetic development of the chaetotaxy. The retention of the larval complement
of setae on a segment throughout ontogeny (localized neoteny) is not uncommon,
although in most cases deficiency in adult chaetotaxy is due to the subnormal number
of setae added to the protonymphal complement at the deutonymphal stage. Retarda-
tion in normal ontogenetic development also affects the dorsal idiosomal sclerotization
in the obligatory parasites which tend to retain, to a greater or lesser degree, the form
of the protonymphal scutal elements in succeeding developmental stages. In
addition to the structural adaptations, many of the facultative and obligatory para-
sites show some degree of specialization of the life cycle. There is a general tendency
for ovoviviparity and, more rarely, viviparity to occur and in the Macronyssinae the
normally active, feeding, deutonymphal stage of the family is replaced by an inactive,
non-feeding instar which shows considerable degeneration of the feeding and sensory
organs.

The morphological and biological diversity displayed by the Dermanyssidae
provides the taxonomist with numerous readily definable characters which are pre-
eminently suitable for the practical differentiation of the taxa, that is, for a classifica-
tion emphasizing differences between species. This is essentially the basis of the
present classification of the family where the exaggerated importance given to one or

THE BRITISH DERM AN YSSID AE (ACARI)

293

two morphological characters has resulted in a plethora of supraspecific taxa. Such
a classification, which can be readily expressed in key form, serves a very useful
purpose as a tool for identification. A more natural classification of the family
requires emphasis to be made on resemblances rather than differences between its
members, but in the elaboration of such a classification the considerable diversity in
the morphology of facultative and obligatory parasites, resulting from variations in
the degree of specialization of various parts of the body, not only between groups of
species but also amongst closely related (congeneric) species, will inevitably add to the
difficulty of defining distinct supraspecific categories.

REFERENCES

BREGETOVA, N. G. 1956. Gamasid Mites (Gamasoidea) . Acad. Sci. U.S.S.R. Moskva.
COSTA, M. 1961. Mites associated with rodents in Israel. Bull. Brit. Mus. (nat. Hist] Zool.

8 : 1-70.
EVANS, G. O. 1957. An introduction to the British Mesostigmata (Acarina) with keys to

families and genera. /. Linn. Soc. (Zool.) 43 : 203-259.
- 1963. Observations on the chaetotaxy of the legs in the free-living Gamasina (Acari :

Mesostigmata). Bull. Brit. Mus. (nat. Hist.) Zool. 10 : 277-303.
— 1964. Some observations on the chaetotaxy of the pedipalps in the Mesostigmata (Acari).

Ann. Mag. nat. Hist. (13) 6 : 513-527.
FONSECA, F. DA. 1935. Notas de Acareologia XII. Eulaelaps vitzthumi, sp.n. (Acarina,

Laelaptidae) . Mem. Inst. Butantan S. Paulo 9 : 33-38.
GRANDJEAN, F. 1940. Les poils et les organes sensitifs portes par les pattes et le palpe chez les

Oribates. Deuxieme partie. Bull. Soc. Zool. France 45 : 32—44.
HAMMEN, L. VAN DER. 1964. The morphology of Glyptholaspis confusa (Foa, 1900) (Acarida,

Gamasina). Zool. Verh. No. 71 : 1-56.
HIRSCHMANN, W. 1957- Gangsystematik der Parasitiformes. Teil i. Rumpfbehaarung und

Ruckenfldchen. Schriftenreihe fur vergleichende Milbenkunde. 20 pp.
KRANTZ, G. W. 1962. A review of the genera of the family Macrochelidae Vitzthum 1930

(Acarina : Mesostigmata). Acarologia 4 : 143-173.
LINDQUIST, E. E. & EVANS, G. O. 1965. Taxonomic concepts in the Ascidae with a modified

setal nomenclature for the idiosoma of the Gamasina (Acari : Meso-stigmata). Mem. ent.

Soc. Canada (in press).
STRANDTMANN, R. W. & WHARTON, G. W. 1958. A manual of mesostigmatid mites parasitic on

vertebrates. Inst. Acarology, Maryland, U.S.A. Contr. No. 4.
ZUMPT, F. & TILL, W. M. in ZUMPT et al. 1961. The arthropod parasites of vertebrates in

Africa south of the Sahara (Ethiopian Region). 1 (Chelicerata) . Publ. 5. Afr. Inst. med.

Res. 9 (i) : 18-86.

ABBREVIATIONS USED IN TEXT-FIGURES

a.m. arthrodial membrane

ac. aciculae

acet. acetabulum

adi-g antero-dorsal setae

alj.o antero-lateral setae

an. sh. anal shield

ant. sp. anterior spine

ar. pr. arthrodial processes

av,_, antero-ventral setae

bas. scl.
bas. tend.

c. gr.
c. s.
ch.
cl.
corn.

basilar sclerite
basilar tendon

capitular groove
capitular seta
chelicera
claw
corniculi

294

G. OWEN EVANS & W. M. TILL

dj-3 dorsal setae

d.d. deutosternal denticles

d.l.f. dorsal lyriform fissure

d.s. dorsal seta of chelicera

end. sh. endo podal shield

exp. sh. expodal shield

f.d. fixed digit

fern. femur

g.b. gnathosomal base

g-v. sh. genito-ventral shield

g-v-a. sh. genito-ventro-anal shield

gen. genital seta

gen. orf. genital orifice

gen. sh. genital shield

hoi. sh. holoventral shield

hyp. 1-3 hypostomal setae

int. mal. internal malae

ji-6 dorsal series of podonotal setae

Ji-5 dorsal series of opisthonotal

setae

Jvi-5 internal ventral setae

1. f. lyriform fissure

lat. in. lateral incision

lat. 1. f. lateral lyriform fissure

Ib. labrum

m. d. movable digit

md. medio-dorsal seta

mes. sc. mesonotal scutellae

met. sh. metasternal shield

mp. sh. metapodal shield

p. 1-3 pores

p.d. pilus dentilis

pan. paranal seta

pdi-s postero-dorsal setae

pd. podal shield

per. peritreme

per. sh. peritrematal shield

pli_2 postero-lateral setae

pod. sh. podonotal shield

pon. postanal seta

pr. process on palp trochanter

ptr. pretarsus

ptr. apod. pretarsal apodeme

ptr. op. pretarsal operculum

pulv. pulvillus

pulv. gd. pulvitractor guide

pulv. scl. pulvitractor sclerite

pulv. tend, pulvitractor tendon

pvt_2 postero- ventral setae

px2_3 posterior accessory setae

pyg. sh. pygidial shield

T2-6 marginal series of podonotal

setae

Ri-y marginal series of opisthonotal
setae

si-6

lateral series of podonotal

setae

Si-5

lateral series of opisthonotal

setae

s.s.

salivary stylus

sg-

stigma

sp.

spermadactyl

st. 1-3

sternal setae

st. 4

metasternal seta

st-g. sh.

sternito-genital shield

st-g-v. sh.

sternito-genito-ventral shield

st. met. sh.

sterno-metasternal shield

st. sh.

sternal shield

tar.

tarsus

tect.

tectum capituli

ten. dep.

depressor tendon

ten. lev.

levator tendon

troch.

trochanter

trt.

tritosternum

UR submarginal series of opis-

thonotal setae

Vj-2 ventral setae

v-a. sh. ventro-anal shield

x supernumerary seta

zi-6 median series of podonotal

setae
Zi-5 median series of opisthonotal

setae
Zvi-2 medio-lateral ventral setae

PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING

A REVIEW OF THE MURIDAE
(ORDER RODENTIA) OF UGANDA

'/?AL

M. J. DELANY and B. R. NEAL

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 13 No. 9

LONDON: 1966

A REVIEW OF THE MURIDAE
(ORDER RODENTIA) OF UGANDA

M. J. DELANY* and B. R. NEAL

Department of Zoology, University of Southampton
Present Address : Department of Zoology, Makerere College, Uganda

Pp. 295-355 ; 20 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 13 No. 9

LONDON: 1966

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Vol. 13, No. 9 of the Zoological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

Trustees of the British Museum (Natural History) 1966

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 2 March, 1966 Price £i js.

A REVIEW OF THE MURIDAE
(ORDER RODENTIA) OF UGANDA

By M. J. DELANY & B. R. NEAL

CONTENTS

INTRODUCTION .......... 297

KEY TO THE GENERA OF UGANDA MURIDS ..... 300

SYSTEMATIC ACCOUNT ......... 302

GAZETTEER ........... 348

REFERENCES .......... 352

INDEX OF GENERA, SPECIES AND COMMON NAMES .... 353

INTRODUCTION

ALTHOUGH Uganda has an exceptionally rich mammal fauna, there has up to the
present time been very little work published on the systematics and biology of these
animals. Apparently, the only check list of Uganda mammals was that given by
Oldfield Thomas in 1902 and published in Sir Harry Johnston's The Uganda Pro-
tectorate. The present work restricts itself to a review of the existing information
on the murid rodents of Uganda. Thomas gave sixteen species occuning in the same
general area (Uganda's boundaries have been changed since 1902) whereas the
present list numbers forty-four. Even so, it is uncertain that this list is complete
particularly in view of the fact that as recently as 1961 a new genus (Delanymys) was
discovered in the south-west of the country and in 1963 a previously unrecorded
genus (Zelotomys) was collected in Ankole. The main sources of information have
been the collections in the British Museum (Natural History), the Coryndon Museum,
Nairobi, published and unpublished literature and the collections made by Delany
in 1961 and 1963. Of the unpublished, Hopkins manuscript entitled The Known
Wild Rodents of Uganda has been particularly useful.

Descriptions of genera and species are given with keys to their identification. As
we found it possible to identify almost every species without reference to cranial
morphology or internal anatomy it has been decided to base the descriptions entirely
on external characters. Such features as grooving of the incisors are included as
their examination does not necessitate dissecting the animal in any way. Only in the
identification of the species of Otomys has it been necessary to refer to other than
external characters. All the measurements given in the text have been obtained
from animals collected in Uganda. Animals that from their external measurements
are obviously very young have been omitted.

The systematics include the reference to the original description of the genus and
the name of the type species. The reference to the original description of each
species is given together with the type locality. If the latter is outside Uganda the
location follows the place name ; if it is in Uganda then the location appears in the

ZOOL. 13, 9 16

2g8 M. J. DELANY & B. R. NEAL

gazetteer. Synonyms based on material collected in Uganda are also included. We
have not attempted any revisions of the systematics although in several cases they
are clearly required. Subgeneric names have been avoided. Whether forms such as
Grammomys, Praomys, Myomys, Mastomys or Hylomyscus are genera or subgenera
seems a matter of constant debate. We have given them generic status in accordance
with fairly general usage, but we fully appreciate that future and more extensive
studies may show them to be of lower taxonomic rank. Furthermore, the paucity
of information on the variation shown by individual species has prompted us to omit
any reference to subspecies. Only very much more collecting from the whole country
can reveal the nature of the subspecies, clines and variability that may be present.

In the following account we have used Simpson's (1945) definition of the Muridae.
His classification of the Myomorph rodents recognizes three super families. They are,
the Muroidea which includes the rats, mice, voles, gerbils and lemmings ; the
Gliroidea containing the dormice, and the Dipodoidea containing the jumping " mice "
and the jerboas. None of the third group occur in Uganda whilst the Gliroidea are
represented by the genus Graphiurus Smuts. The dormice are easily recognized by
their small size and very bushy tail ; although normally grey in colour, brown and
buff forms also occur.

The Muroidea are split into four families ; the Muridae which are being considered
in the present account, the Spalacidae or mole rats of the Mediterranean region, the
Rhizomyidae which has only one species recorded from Uganda (Tachyoryctes ibeanus
Thomas) and the Cricetidae in which group are included the gerbils. Tachyoryctes,
one of the African mole rats, is easily recognized and not readily confused with the
murids. It is highly adapted for burrowing. The limbs are very short and the front
legs broad and slightly flattened for digging ; external ears are almost completely
absent and the eyes are very small. The fur is long, soft and dense and usually
brown to black in colour. The tail is very short.

The gerbils are the forms most likely to be confused with the murids and can only
be separated from them by a combination of several characters. The hind limbs are
long in proportion to the fore limbs (not so obvious in Tatera), the feet have long
claws, the fur is soft and dorsally sandy or buff in colour. The belly fur and the backs
of the hands and feet are pure white and a patch of short white hairs occurs behind
the ear. The tail is long and hairy and often tufted. The upper incisors are grooved
but on occasion the grooving is so shallow as to make it hardly perceptible. Two
genera (Tatera and Taterillus) have been recorded from Uganda.

Fetter (1964) has recently suggested that the genera Cricetomys and Saccostomus
should be placed in the Cricetidae.

In the sections on distribution the locality from which animals have been obtained
is given with the District except in Buganda where the reference is to the Kingdom.
The latitude and longitude of each locality appears in the gazetteer. In addition
the distributions have been mapped ; the location of Buganda and the Districts
outside this Kingdom are shown in Text-fig. I. The distribution records are un-
doubtedly inadequate as almost every species probably has a wider and more con-
tinuous distribution than is suggested from the maps. This can only be remedied by

THE MURIDAE OF UGANDA

299

considerably more collecting. Some distribution records are very vague and may
refer to no more than a District ; in which case the District is given in quotation
marks. Various parts of Uganda appear to have been more rigorously collected than
others with particularly large gaps occurring in Acholi, Busoga, east Toro and Ankole
and in the north along the length of the Sudan border.

There is not a great deal of information available on the biology and breeding.
Some data have been included from outside Uganda. However, there seemed little
reason for including times of breeding of animals outside the country as this pheno-

-A'N.

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31°

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i 100 miles i

35°E.

FIG. i. Map of Uganda showing the location of the Kingdom of Buganda and the Districts

outside the Kingdom (underlined) .

300 M. J. DELANY & B. R. NEAL

menon is probably influenced by local climatic conditions. As these vary con-
siderably from one part of Africa to another (as well as in Uganda itself) the inclusion
of times of breeding from elsewhere could give an erroneous picture of when breeding
took place in Uganda.

We are particularly indebted for the very considerable help Mr. R. W. Hayman
of the British Museum (Natural History) has given us with this work. We are
grateful to the Royal Geographical Society for assistance in tracing some of the
localities. The collecting trips made by M. J. D. in 1961 and 1963 were financed by
grants from the Royal Society, the Percy Sladen Memorial Fund and the University
of Southampton.

ABBREVIATIONS

The following abbreviations have been used in the text : —

cm. centimetres Mt. Mount

Co. County N. North

E. East Q.E.P. Queen Elizabeth National Park

g. grams S. South

h. & b. head-and-body length strm. stream

h. f. hind foot length W. West

meas. measured wt. weight

M.N.P. Murchison Falls National Park $ male

m. metre (s) $ female

mm. millimetres

Months have been abbreviated to their first three letters.

The number of records of the number of foetuses have been indicated in brackets ;
thus (2 X i) signifies that two animals each had one foetus.

KEY TO THE GENERA OF UGANDA MURIDS
Large size (h. & b. 300-450 mm.). Long tail, distal portion white, proximal portion

dark . CRICETOMYS (p. 338)

Smaller with tail not showing this pattern of coloration ..... 2

Dorsal fur either spiny or very bristly ........ 3

Dorsal fur neither spiny nor very bristly ........ 6

Fur modified into distinct spines along dorsal surface . . ACOMYS (p. 334)

Fur bristly but not spiny .......... 4

Dorsal fur chestnut, bristly and stiff; tail about 1-5 times h. & b. DEOMYS (p. 343)

Dorsal fur stiff and brush-like, reddish-brown; tail approximately equal to or less

than h. & b 5

Backs of hands and feet brown or black; belly red-brown or buff

LOPHUROMYS (p. 331)
Backs of hands and feet white, belly white .... URANOMYS (p. 336)

Either upper and lower incisors grooved or just the upper ones .... 7

Neither upper nor lower incisors grooved . . . . . . . . n

Both upper and lower incisors deeply grooved; fur long and soft; tail appreciably

shorter than h. & b OTOMYS (p. 345)

Upper incisors at least faintly grooved; lower incisors not grooved ... 8

THE MURIDAE OF UGANDA 301

8 Large forms (h. & b. over 100 mm.) ......... 9

Small forms (h. & b. under 100 mm.) ........ 10

9 Fur coarse and harsh; belly hairs dirty yellow-grey . . . PELOMYS (p. 310)
Fur thick and soft ; belly hairs white at the tips and sharply demarcated from those

at the flanks . . . MYLOMYS (p. 305)

10 Tail very short, less than 40 mm STEATOMYS (p. 342)

Tail appreciably longer, over 60 mm. .... DENDROMUS (p. 339)

1 1 With red tip to snout in sharp contrast to rest of body . . OENOMYS (p. 305)
Colour of snout neither bright red nor in sharp contrast with rest of body . . 12

12 Fur with metallic iridescent lustre ......... 13

Fur without metallic iridescent lustre . . . . . . . . 14

13 Fur rough and shaggy looking ....... DASYMYS (p. 307)

Fur closely applied to the body AETHOMYS (p. 317)

14 With at least one dorsal stripe .......... 15

Without stripes ............ 17

15 A single black mid-dorsal stripe ...... HYBOMYS (p. 316)

Several stripes present ........... 16

16 Four black stripes along back; mid-dorsal line pale . RHABDOMYS (p. 315)
Numerous pale stripes along back (may be composed of lines of spots) ; mid-dorsal

line dark . . . . • . . . . LEMNISCOMYS (p. 313)

17 Tail considerably longer than h. & b. . . . . . . . . 18

Tail approximately equal to h. & b. or shorter ....... 23

1 8 Tip of tail with small but distinct pencil of hairs . . . . . . 19

Tip of tail without pencil of hairs ......... 20

19 Hind foot relatively broad THAMNOMYS (p. 302)

Hind foot not broadened GRAMMOMYS (p. 304)

20 Very small (h. & b. less than 60 mm.) ; tail relatively long (about 100 mm.)

DELANYMYS (p. 343)
H. & b. over 60 mm.; tail relatively not so long . . . . . . 21

21 Medium size (h. & b. 105 to 138 mm.) ..... PRAOMYS (p. 321)
Smaller size (h. & b. 71 to 107 mm.) ......... 22

22 Hairs of belly with white tips and grey bases . . . HYLOMYSCUS (p. 322)
Hairs of belly pure white ....... MYOMYS (p. 324)

23 Medium size (h. & b. 127 to 160 mm.); tail £ to £ h. & b. SA CCO S TOM US (p. 337)
Small or medium size ; if the latter tail is not short ...... 24

24 Hairs coarse, strongly annulated black on buff, producing a " pepper and salt " effect ;

tail shorter than h. & b. but never less than f h. & b. length; ears hairy

ARVICANTHIS (p. 309)
Without " pepper and salt " effect ......... 25

25 Medium size (h. & b. 133 to 171 mm.) ; tail about h. & b. length; hind feet very large

(35 to 40 mm.) MALACOMYS (p. 327)

Variable size but with appreciably smaller hind feet . . . . . .26

26 Upper incisors pro-odont (projecting forwards) . . . ZELOTOMYS (p. 327)
Upper incisors not pro-odont .......... 27

27 Small size (h. & b. less than 93 mm.) ...... MUS (p. 329)

Larger size (h. & b. over 90 mm.) ......... 28

28 Texture of fur very soft ; flanks brown, belly grey with clear demarcation between the

two MASTOMYS (p. 325)

Texture of fur coarse, dorsally brown to grey with a gradual transition in colour
from back to belly with no sharp line of demarcation between flanks and belly

RATTUS (p. 319)

302 M. J. DELANY & B. R. NEAL

SYSTEMATIC ACCOUNT

Family MURIDAE Gray
1821. Muridae Gray, London Med. Reposit. 15 : 303.

Subfamily MURINAE Murray
1866. Murinae Murray, The Geographic Distribution of Mammals : 359.

Genus THAMNOMYS Thomas. Thicket Rats

1907. Thamnomys Thomas, Ann. Mag. nat. Hist., (7) 19 : 121. Genotype Thamnomys venustus
Thomas.

The genera Thamnomys and Grammomys are very similar. The tail is very long in
both and with a pencil of hairs at its tip. The same is true of the gerbils but they
can be separated from these two genera on other characters (p. 298). The hind foot
of Thamnomys is broader than in Grammomys. Examination of the Uganda material
shows differences in the colour of the belly fur in the two genera. In Grammomys
the belly is pure white whilst in Thamnomys it is whitish washed with buff, or greyish
with white tips. This character has not been found to hold good for all the specimens
examined from other parts of Africa. We have recognized T. kempi as conspecific
with T. venustus.

Hairs of belly white to bases, washed with buff ..... T. rutilans

Hairs of belly grey with white tips ....... T. venustus

Thamnomys rutilans (Peters)

1876. Mus rutilans Peters, Monatsb. K. preuss. Akad. Wiss., Berlin 1876 : 478. Lambarene,
Gaboon [o° 45' S. 10° 15' E.].

DESCRIPTION. Only specimen examined from Uganda has a rich suffusion of buff
to the white hairs of the belly.

DISTRIBUTION. Zika Forest, Buganda. Text-fig. 2.

MEASUREMENTS. One ^ h. & b. 145 mm. ; tail 172 mm. ; h. f. 25 mm. ; ear 17
mm. ; weight 54 g.

BREEDING. No information available.
HABITAT. Typically a forest species.
BIOLOGY. No information available.

Thamnomys venustus Thomas

1907. Thamnomys venustus Thomas, Ann. Mag. nat. Hist., (7) 19 : 122. Mubuku Valley, Toro.
1911. Thamnomys kempi Dollman, Ann. Mag. nat. Hist., (8) 8 : 658. Buhamba, near Lake
Kivu, Congo [i° 32' S. 29° 19' E.].

DESCRIPTION. Tips of belly hairs white, bases grey. Fur thicker and softer texture
than T. rutilans.

THE MURIDAE OF UGANDA

303

DISTRIBUTION. Echuya Forest, Impenetrable Forest, Kigezi ; Mihunga Swamp,
Mubuku Valley, Toro. Text-fig. 2.

MEASUREMENTS. One <$ h. & b. 125 mm. ; tail 181 mm. ; h. f. 25 mm. ; ear 18
mm. Three $ h. & b. 151 mm., 131 mm., 141 mm. ; tail 183 mm., 162 mm., 184 mm. ;
h. f. 26 mm., 26 mm., 28 mm. ; ear 19 mm., 20 mm., 19 mm. ; weights 66 g., 51 g.,
56 g.

BREEDING. No information available.

o Thamnomys rutilans
• Thamnomys venustus
^ Grammomys dolichurus

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SUDAN

Edward,

MT.
MUHAVURA

FIG. 2. Distribution of Thamnomys rutilans, T. venustus and Grammomys dolichurus.

304 M. J. DELANY & B. R. NEAL

HABITAT. High altitude (including moist montane) forest.
BIOLOGY. Arboreal. Nocturnal (Allen & Loveridge, 1942).

Genus GRAMMOMYS Thomas. Tree Rats

1915. Grammomys Thomas, Ann. Mag. nat. Hist., (8) 16 : 150. Genotype, by original desig-
nation, Mus dolichurus Smuts.

Separated, on external characters, from Thamnomys on the narrower hind foot.
Long tail with a pencil of hairs at its tip. In the Uganda specimens examined the
belly hairs are pure white without a buff suffusion. The dorsal fur is brown to grey-
brown with, in some specimens, a buff line where flank and belly fur meet. Allen
(1939) includes Grammomys as a subgenus of Thamnomys.

Grammomys dolichurus (Smuts)

1832. Mus dolichurus Smuts, Enumerat. Mamm. Capens. : 38. Type locality given as " near
Cape Town ", South Africa but this is of uncertain accuracy (see Ellerman, Morrison-Scott &
Hayman, 1953)-

1907. Thamnomys dryas Thomas, Ann. Mag. nat. Hist., (7) 19 : 123. Mubuku Valley, Toro.

1907. Thamnomys macmillani Wroughton, Ann. Mag. nat. Hist., (7) 20 : 504. Wouida, north of
Lake Rudolf, Ethiopia.

1908. Thamnomys surdaster Thomas & Wroughton, Proc. zool. Soc. Lond. 1908 : 550. Zomba,
Malawi [15° 22' S. 35° 22' E.].

1910. Thamnomys discolor Thomas, Ann. Mag. nat. Hist., (8) 5 : 283. Kakamega Forest,
Kisumu, Kenya [o° 19' N. 34° 51' E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Maramagambo Forest, Ankole ; Kalule, Kampala, Lialo, Nkya-
nuna, Buganda ; Salalira, " south Bugisu ", Bugisu ; " Bukedi " ; Kotido, Lotome,
Moroto, Karamoja ; Echuya Forest, Kumba, Nyalusanje, Kigezi ; Serere, Teso ;
Ilumia, Mihunga, Mubuku Valley, Mweya, Wasa River, Toro ; Rhino Camp, West
Nile. Text-fig. 2.

MEASUREMENTS. Three $ h. & b. 120 mm., 98 mm., 126 mm. ; tail 165 mm.,
134 mm., 183 mm. ; h. f. 23 mm., 22 mm., 27 mm. ; ear 15 mm., 14 mm., 17 mm. ;
weights ?, 24 g., 51 g. $ h. & b. 113-9 mm- (8 meas., range 99 to 130 mm.) ; tail
173-7 mm. (8 meas., range 150 to 205 mm.) ; h. f. 23-6 mm. (8 meas., range 22 to 25
mm.) ; ear 16-2 mm. (8 meas., range 15 to 20 mm.) ; weight 41-2 g. (5 weighed, range
33 to 53 g.).

BREEDING. Foetuses (1x3), Mweya ; (2x3) Moroto, Nov. In Zambia (3 X
4) (Ansell, 1960). Watson (1950) reports usually 3 young in a litter.

HABITAT. Scrub, bush and various types of forest.

BIOLOGY. Arboreal ; nocturnal Build nests in bushes and trees. Outer part of
nest of coarse grass with a lining of finely chewed grass. Brown house snake (Boaedon
lineatus) reported to prey on it (Allen & Loveridge, 1942). Vegetarian, including
seeds.

THE MURIDAE OF UGANDA 305

Genus OENOMYS Thomas. Rusty-nosed Rats

1904. Oenomys Thomas, Ann. Mag. nat. Hist., (7) 13 : 416. Genotype, by original designation,
Mus hypoxanthus Pucheran.

Medium sized rats easily recognized by their rusty-red nose. Upper surface brown
or grey often with tinge of olive, this colour produced by the hairs having long buff or
rufous tips and dark grey bases. Rump suffused with rusty-red colour. Backs of
hands and feet pale brown. Flanks paler than back sharply demarcated from white
of underside. Belly hairs white to roots. Tail grey-brown above, much paler below.

Oenomys hypoxanthus (Pucheran)

1855. Mus hypoxanthus Pucheran, Rev. Mag. Zool. 2 : 206. Gaboon.

1910. Oenomys bacchante Thomas and Wroughton, Trans, zool. Soc. Lond. 19 : 509. Mubuku
Valley, Toro.

DESCRIPTION. As for genus.

DISTRIBUTION. Kampala, Buganda ; Bubungi, Buyobo, Sipi, Bugisu ; Fadjao,
Bunyoro ; Impenetrable Forest, Kigezi ; Benet, Sebei ; Bundibugyo, Bundimali,
Ilumia, Kilembe, Kyatwa, Mihunga, Mubuku Valley, Sara, Toro. Text-fig. 3.

MEASUREMENTS. H. & b. <J 142-4 mm. (n meas., range 105 to 167 mm.), $
143-2 mm. (13 meas., range 131 to 159 mm.) ; tail $ 172-6 mm. (n meas., range 150
to 205 mm.), $ 170-5 mm. (13 meas., range 135 to 187 mm.) ; h. f. $ 29-9 mm. (n
meas., range 24 to 33 mm.), $ 29-7 mm. (13 meas., range 28-5 to 31 mm.) ; ear <$
19-2 mm. (n meas., range 18 to 21 mm.), $ 18-8 mm. (13 meas., range 17 to 21 mm.) ;
weight cJ 70 g. 90 g., $ 57 g., 76 g. (only four weighed).

BREEDING. Foetuses (5 x 3, 4 X 4) Kampala (Hopkins MS.) ; (1x3) in Congo
(Misonne, 1963). Litter of three blind nestlings in Congo (Allen & Loveridge, 1942).

HABITAT. Tall grass bordering marshes and streams ; swamps amongst Cyperus
latifolius and fern.

BIOLOGY. Climbs easily and makes nests in grass.

Genus MYLOMYS Thomas

1906. Mylomys Thomas, Ann. Mag. nat. Hist., (7) 18 : 224. Genotype, by original designation,
Mylomys cuninghamei Thomas.

Medium sized rats with the dorsal surface a bright olive-gold, heavily lined with
black ; flanks with less black but sharply demarcated from the white of the under-
side. Tail generally a little shorter than the head-and-body length, black above,
yellow buff or whitish below. Grooved upper incisors.

Mylomys cuninghamei Thomas

1906. Mylomys cuninghamei Thomas, Ann. Mag. nat. Hist., (7) 18 : 225. East of Aberdare

Mountains, Kenya [approximately o°-i° S. 36° 45' E.].
1915. Mylomys lutescens Thomas, Ann. Mag. nat. Hist., (8) 16 : 149. Nyalasanje, Kigezi.

DESCRIPTION. As for genus.

306

M. J. DELANY & B. R. NEAL

DISTRIBUTION. Paraa, Acholi ; Lutoto, north of Maramagambo Forest, Ankole ;
Kampala, Mabira Forest, Buganda ; Walasi Bugisu ; Budama, Kidoko, Mulanda,
Bukedi ; Kanaba, Kiduha, Nyalusanje, Kigezi ; Serere, Teso ; Butiti, Crater Track,
Toro. Text-fig. 3.

MEASUREMENTS. H. & b. <$ 154-3 mm. (20 meas., range 122 to 183 mm.), £
153-1 mm. (21 meas., range 125 to 180 mm.) ; tail $ 141-4 mm. (20 meas., 104 to 180
mm.), $ 142-2 mm. (20 meas., range 119 to 156 mm.) ; h. f. $ 33-2 mm. (20 meas.,
range 29 to 36 mm.), $ 32-4 mm. (21 meas., range 30 to 35 mm.) ; ear J 18-5 mm.

• Oenomys hypoxanthus
v Mylomys cuninghamei

SUDAN

MT.
MUHAVURA

'yf\S \

RUANDA ^
L;

100 miles

TANGANYIKA

FIG. 3. Distribution of Oenomys hypoxanthus and Mylomys cuninghamei.

THE MURIDAE OF UGANDA 307

(17 meas., range 14 to 22 mm.), $ 18-8 mm. (20 meas., range 15 to 22 mm.) ; weight
<J 103-1 g. (18 weighed, range 50 to 165 g.), $ 99-4 g. (20 weighed, range 46 to 160 g.).

BREEDING. Foetuses (i X 3, wt. less than i g.), (i x 5, wt. 18 g.) Jul. Crater
Track, (i X 5, wt. i g.), (i X 4, wt. 4 g.) Aug. north of Maramagambo Forest,
(i x 3), (i X 4 wt. i g.) Aug. Lutoto.

HABITAT. Typically thick grassland, heath and scrub.
BIOLOGY. Diurnal ? Herbivorous, eating grass leaves and stems.

Genus DASYMYS Peters. Shaggy Swamp Rats

1875. Dasymys Peters, Mber. preuss. Akad. Wiss. Berl. 1875 : 12. Genotype, by monotypy,
Dasymys gueinzii Peters = Mus incomtus Sundevall.

Moderate sized rats with long, soft and untidy fur and sparsely haired tail. The
animal has a flattened appearance. Dorsal hairs inconspicuously annulated black
and buff giving the general effect of darkish grey tinged with brown or greyish brown.
Backs of the hands and feet dark coloured, almost naked. Underside slaty grey
with a greater or lesser amount of whitish admixture due to the pale tips of the hairs.
Eyes small. Ears hairy. Ellerman (1941) and Hopkins (MS.) believe there is only
one species of Dasymys in Uganda.

Dasymys incomtus (Sundevall)

1846. Mus incomtus Sundevall, 6/vers. VetenskAkad. Fork. Stockh. 3 : 120. " E. Caffraria

prope Port Natal " (= Durban, Natal) [29° 53' S. 31° oo' E.].
1875. Dasymys gueinzii Peters, Mber. preuss. Akad. Wiss. Berl. 1875 : 13. Interior of " Port

Natal " (= Durban, Natal).

1906. Dasymys medius Thomas, Ann. Mag. not. Hist., (7) 18 : 143. Mubuku Valley, Toro.
1906. Dasymys montanus Thomas, Ann. Mag. nat. Hist., (7) 18 : 143. Mubuku Valley, Toro.
1911. Dasymys orthos Heller, Smithson, misc. Coll. 56 : 13. Butiaba, Bunyoro.

DESCRIPTION. As for genus.

DISTRIBUTION. Asuya, Gulu, Acholi ; near Kagambah, Lutoto, Ankole ; Kaku-
miro, Kampala, Mengo, Buganda ; Budama, Bukedi ; Butiaba, Bunyoro ; Echuya
Swamp, Ingezi, Kiduha, Kumba, Nyalasanji, Kigezi ; Moyo, Madi ; Amuria, Serere,
Teso ; Mihunga, Mubuku Valley, Toro ; Rhino Camp, West Nile. Text-fig. 4.

MEASUREMENTS. H. & b. $ 154-9 mm- (X3 meas., range 146 to 166 mm.), $ 145-0
mm. (n meas., range 130 to 170 mm.) ; tail $ 132-5 mm. (13 meas., range in to 150
mm.), $ 127-4 mm. (n meas., range 105 to 150 mm.) ; h. f. <£ 29-9 mm. (13 meas.,
range 26 to 32-5 mm.), $ 28-8 mm. (n meas., range 27 to 31 mm.) ; ear $ 20-6 mm.
(13 meas., range 19 to 23 mm.), $ 20-4 mm. (n meas., range 17-5 to 25 mm.) ; one <j>
weighed 59 g.

308

M. J. DELANY & B. R. NEAL

BREEDING. Foetuses (i x 4, i X 5) in Zambia (Ansell, 1960) ; (3 x 2, 3 x 3)
Hopkins (MS.). Usually 2 to 4 in a litter in South West Africa (Shortridge, 1934).

HABITAT. Typically swamps, reed beds and river valleys. Occurs at various
elevations ; recorded from boggy moss covered ground between 12,500 and 14,000
feet on Ruwenzori. Misonne (1963) has found this species in mixed savanna in the
Congo.

BIOLOGY. Vegetarian. Nests made of grass on surface of the ground.

• Dasymys incomtus

MT. ;_,• V \

MUHAVURAJ?^ / N

Edward,

FIG. 4. Distribution of Dasymys incomtus.

THE MURIDAE OF UGANDA 309

Genus ARVICANTHIS Lesson. Unstriped Grass-mice

1842. Arvicanthis Lesson, Nouv. Tabl. Regne Anim. Mammif. : 147. Genotype, by monotypy
and original designation, Lemmus niloticus Geoffroy, 1803 = Hypudaeus variegatus Lich ten-
stein, 1823 = Arvicola niloticus Desmarest, 1822.

1843. Isomys Sundevall, K. svenska VetenskAkad. Handl. 1842 : 219.

Medium sized animals, with fur composed of rather harsh hairs which are annulated
with brown and buff in such a way as to produce a " pepper and salt " effect. The
genus might be confused with Mylomys but Arvicanthis has a grisly coloured under-
surface and upper incisors without grooves. The tail is usually distinctly shorter
than the head and body.

Arvicanthis niloticus (Desmarest)

1822. Arvicola niloticus Desmarest, Mammalogie 2 : 281. Egypt.

1842. Mus abyssinicus Riippell, Mus. Senckenbergianum 3 : 104. Entschetqab, Simen Province,
Ethiopia [13° 15' N. 38° 20' E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Kitgum, Acholi ; Congo Road, north of Maramagambo Forest,
Burumba, Kagambah, Mbarara, Ankole ; Buruli, Entebbe, Kakumiro, Kampala,
Kisingo, Mabira Forest, near Masaka, Nalweyo, Buganda ; Budongo Forest, Bulisa,
Butiaba, Hoima, Kibiro, Masindi, Bunyoro ; Amudat, Anamuget, Bokora, Kam-
churu, Kotido, Locihotome, Moroto, Moruita, Nabilatuk, Karamoja ; Nyakabande,
Nyalusanje, Kigezi ; Kibusi, Lango ; Ajeluk, Amuria, Serere, Teso ; Bugoye,
Bundibugyo, Crater Track, Hakitengya, Ilumia, Kamulikwezi, Kilembe, Kimara,
Makoga, Mpanga Forest, Mubuku Valley, Mweya, south-east Ruwenzori, Toro ;
Adropi, Arua, Offude, Rhino Camp, West Nile. Kibandama, Patong. Text-fig. 5.

MEASUREMENTS. H. & b. <£ 146-3 mm. (54 meas., range 120 to 184 mm.), $
141-6 mm. (59 meas., range 120 to 167 mm.) ; tail <£ no-8 mm. (54 meas., range 83
to 134 mm.), $ 109-7 mm- (58 meas.; range 86 to 129 mm.) ; h. f. ^ 28-5 mm. (55
meas., range 23 to 32 mm.), $ 27-9 mm. (58 meas., range 25 to 32-5 mm.) ; ear ^
16-7 mm. (52 meas., range 13 to 20 mm.), $ 16-3 mm. (57 meas., range 13 to 20 mm.) ;
weight <$ 80-0 g. (23 weighed, range 57 to 120 g.), $ 77-5 g. (33 weighed, range 50 to

BREEDING. Foetuses (i x 4) Aug. Congo Road, (i x 4) Sep. Kamulikwezi.
Watson (1950) reports a possible maximum breeding season towards the end of the
rains as they are very numerous during the early months of the dry season.

HABITAT. Typically a grassland species but also common in bush and cultivated
land. May be found in native huts and grain stores.

BIOLOGY. Nocturnal and diurnal. Herbivorous, eating leaves and stems,
especially of Ameranthus polygamus. The black-shouldered kite (Elanus coeruleus)
has been reported to prey on this species. It digs burrows in which it nests, often in
banks or rubbish heaps or at the foot of bushes, with tunnels leading through the
thick grass from them. They are made of fine grass and often placed four or five
together, forming a warren. Surface nests also occur, as well as burrows, but it is not

3io

M. J. DELANY & B. R. NEAL

known in what circumstances they are used. The underground nests are usually 8
inches to 2 feet deep. The surface nests are usually in a thick tussock of grass.

Genus PELOMYS Peters. Creek Rats

1852. Pelomys Peters, Mber. preuss. Akad. Wiss. Berl. 1852 : 275. As a subgenus of Mus

Linneaus ; genotype, by monotypy, Mus. (Pelomys) fallax Peters.
1910. Desmomys Thomas, Ann. Mag. not. Hist., (8) 5 : 284. Pelomys harringtoni Thomas,

valid as a subgenus.
1924. Komemys de Beaux, Ann. Mus. Stor. nat. Genova 51 : 207. Komemys isseli de Beaux,

valid as a subgenus.

Arvicanthis niloticus

Edward

TANGANYIKA
I

FIG. 5. Distribution of Arvicanthis niloticus.

THE MURIDAE OF UGANDA 311

The creek rats are medium sized rats. The colour of the dorsal surface rather
resembles Arvicanthis from which they can be distinguished by the grooved upper
incisors. They may or may not have a distinct dorsal stripe. The underside is
dirty yellow, buff or greyish and not sharply demarcated from the dark coloured
flanks. The relative length of the tail varies in the different species. P. isseli is
included in the subgenus Komemys and the other two Uganda species in the subgenus
Pelomys.

1 With very distinct black dorsal stripe ........ 2

Dorsal stripe absent .......... P. fallax

2 Tail about i -5 times length of head and body ...... P. isseli

Tail about as long as head and body ...... P. hopkinsi

Pelomys fallax (Peters)

1852. Mus (Pelomys} fallax Peters, Mber. preuss. Akad. Wiss. Berl. 1852 : 275. Caya District,
Zambezi River, Portuguese East Africa = Sena, Mozambique [17° 20' S. 35° 10' E.].

DESCRIPTION. Golden-yellow above, the hairs heavily annulated with black,
giving a " pepper and salt " effect as in Arvicanthis ; no dark dorsal stripe. Under-
side olive buff. Tail just shorter than head-and-body length.

DISTRIBUTION. Kagambah, Mbarara, Ankole ; Kiduha, Nyalasanje, Kigezi.
Text-fig. 6.

MEASUREMENTS. H. & b. <$ 142-3 mm. (6 meas., range 135 to 146 mm.), $ 147 mm.
(i meas.); tail <$ 133-7 mm. (6 meas., range 127 to 137 mm.), $ 135 mm. ; h. f. $
29-7 mm. (6 meas., range 29 to 30 mm.), $ 29-5 mm. ; ear <$ 17-8 mm. (6 meas.,
range 17 to 18 mm.), $ 18 mm. ; none weighed.

BREEDING. Foetuses, Ansell (1960) reports (i x 9, i X 7) ; juveniles and sub-
adults caught throughout the year suggests no fixed breeding season in Zambia.

HABITAT. Reported from swamps, reed beds, river banks and damp places.

BIOLOGY. Reported to be diurnal in South West Africa (Shortridge, 1934), but
Ansell (1960) suggests that it is mainly nocturnal in Zambia. The South West
African form of this species is reported to feed on reed shoots and other swamp
vegetation, whilst in Tanzania it is stated to be destructive to grain crops. The
species is a very able swimmer, and in Angola is reported to make deep burrows
although no signs of any nests or holes have been found in other localities.

Pelomys isseli (de Beaux)

1924. Komemys isseli de Beaux, Ann. Mus. Star. nat. Genova 51 : 207. Kome Island, Buganda.

DESCRIPTION. Upperside buff, sprinkled with black and with a very distinct black
mid-dorsal stripe. Underside dirty whitish or buff. Tail nearly 1-5 times the length
of the head and body.

ZOOL. 13, 9 17

312 M. J. DELANY & B. R. NEAL

DISTRIBUTION. Bugala Island, Rome Island, Buganda. Text-fig. 6.

MEASUREMENTS. Only two $ measured, h. & b. 100 and 106 mm. ; tail 143 and
148-5 mm. ; h. f. 29 and 29 mm. ; ear 16 and 18-5 mm. ; not weighed.

BREEDING. No information available.
HABITAT. No information available.
BIOLOGY. No information available.

• Pelomys fallax
v Pelomys isseli
o Pelomys hopkinsi ')

SUDAN

Edward,

FIG. 6. Distribution of Pelomys spp.

THE MURIDAE OF UGANDA 313

Pelomys hopkinsi Hayman

1955. Pelomys hopkinsi Hayman, Rev. Zool. Bot. afr. 52 : 323. Rwamachuchu, Kigezi.

DESCRIPTION. General appearance that of a small Arvicanthis with a distinct
black mid-dorsal stripe. Underside generally buffy. Tail about as long as head
and body.

DISTRIBUTION. Rwamachuchu, Kigezi. Text-fig. 6.

MEASUREMENTS. One <$ meas., tail 135 mm. ; h. f. 32 mm.

BREEDING. No information available.

HABITAT. Edge of papyrus swamp.

BIOLOGY. No information available.

Genus LEMNISCOMYS Trouessart. Striped Grass-mice

1881. Lemniscomys Trouessart, Bull. Soc. Sci. Angers 10 : 124. As a subgenus of Mus Lin-
naeus ; genotype, by subsequent designation (Thomas, 1916, Ann. Mag. nat. Hist., (8) 18 :
67), Mus barbarus Linnaeus.

The striped grass mice are characterized by the presence of a dark mid-dorsal line
and numerous white dorsal and dorso-lateral stripes which may or may not be broken
up into spots. The underside is pure white. The tail is hairy and as long as, or
longer than the head and body. The fifth finger is shortened.

White dorsal and dorso-lateral stripes continuous .... L. barbarus

White dorsal and dorso-lateral stripes broken into spots ... L. striatus

Lemniscomys barbarus (Linnaeus)

1766. Mus barbarus Linnaeus, Syst. Nat. i2th ed., pt. 2, add. not paged. Morocco.
DESCRIPTION. Slightly smaller than L. striatus ; continuous stripes along body.

DISTRIBUTION. Paraa, Acholi ; Nabilatuk, Karomoja ; Ajeluk, Malera, Serere,
Teso ; Rhino Camp, Wadelai, West Nile. Text-fig. 7.

MEASUREMENTS. H. & b. ^ 105-7 mm- (7 meas., range 98 to 118 mm.), one $
105 mm. ; tail <$ 107-3 mm. (7 meas., range 95 to 116 mm.), $ 118 mm. ; h. f. 23-3 mm.
(7 meas., range 22 to 25 mm.), $ 23 mm. ; ear <$ 12-9 mm. (7 meas., range 12 to 15 mm.),
$ 12 mm. ; weight <$ 30-0 g. (6 weighed, range 23 to 36 g.), $ 41 g.

BREEDING. Foetuses (i x 5) Oct., Nabilatuk.
HABITAT. Typically grass and scrub in dryer areas.
BIOLOGY. No information available.

Lemniscomys striatus (Linnaeus)

1758. Miis striatus Linnaeus, Syst. Nat. loth ed., pt. i : 62. " India " = Sierra Leone (Thomas

1911, Proc. zool. Soc. Lond. 1911 : 148).
1910. Arvicanthis macculus Thomas & Wroughton, Trans, zool. Soc. Lond. 19 : 515. Muhokya,

Toro.
1919. Lemniscomys macculus Hollister, Bull. U.S. nat. Mus. 99 : 138.

314

M. J. DELANY & B. R. NEAL

DESCRIPTION. Slightly larger of the two species. The white stripes broken into
spots which may be joined to each other.

DISTRIBUTION. Awack, Fort Patiko, Pamdero, Acholi ; Congo Road, Marama-
gambo Forest, north of Maramagambo Forest, between Rwempuno and Kaizi Rivers,
Kagambah, Ankole ; Entebbe, Kabanyolo, Kabula, Kajansi, Kampala, Kasai
Forest, Kisingo, Lunyo, Mabira Forest, Nabugabo, Buganda ; Lwakaka, Bugisu ;
Tororo, Bukedi ; Busingiro, Hoima, Masindi, Bunyoro ; Moroto, Namalu, Karamoja ;

• Lemniscomys barbarus
T Lemniscomys striatus

SUDAN

-I I

' v- Ml.

A'ELGON .£•
<<J

RUWENZORI

Edward,

MT. i i
MUHAVURAJ?- \
s'

RUANDA

FIG. 7. Distribution of Lemniscomys spp.

THE MURIDAE OF UGANDA 315

Bugoye, Bundibugyo, Crater Track, Ilumia, Mihunga, Mpanga Forest, Mubuku
Valley, Muhokya, Mweya, south east Ruwenzori, Tokwe, Toro ; Rhino Camp, West
Nile. Text-fig. 7.

MEASUREMENTS. H. & b. ^ 111-7 mm- (27 meas., range 93 to 131 mm.), $ 113-9
mm. (24 meas., range 91 to 142 mm.) ; tail <$ 119-4 mm. (25 meas., range 92 to 144
mm.), $ 124-2 mm. (18 meas., range 103 to 141 mm.) ; h. f. ^ 24-2 mm. (28 meas.,
range 20-5 to 27-5 mm.), 24-4 mm. $ (24 meas., range 20-5 to 27 mm.) ; ear ^ 15-3 mm.
(21 meas., range 14 to 17-5 mm.), $ 15-8 mm. (15 meas., range 14 to 17 mm.) ; weight
c? 36'7 g- (13 weighed, range 27 to 46 g.), $ 38-2 g. (12 weighed, range 18 to 68 g.).

BREEDING. Foetuses (i x 5) Aug. Maramagambo Forest ; Allen & Loveridge
(1942) report (i X 5) Dec. Kibale Forest, (i X 5) Dec. Bundibugyo. Juveniles — 4
nestlings Nov. Mabira Forest, 2 blind nestlings Jan. Mihunga, 2 young late Jan.
Bugoye. Misonne (1963) reports litters of 3 to 6 in Congo.

HABITAT. Inhabits grassland, savanna, dense scrub and cultivated land. It is
also apparently found in quite thick forest.

BIOLOGY. Nocturnal and diurnal. Omnivorous, observation of stomach contents
showed leaf and stem remains, seeds and insects to be numerous. " Natives state
that this species eats young shoots of grass, sweet potatoes, cassava and fallen
maize cobs. The nest is often placed at the base of a tussock of tall grass, and is
made from grass " (Hopkins MS.).

Genus RHABDOMYS Thomas. Four-striped Grass-mice

1916. Rhabdomys Thomas, Ann. Mag. nat. Hist., (8) 18 : 69. Genotype, by original designation,
Mus pumilio Sparrman.

Characterized by having four black stripes along length of body and a pale mid-
dorsal line. Yellow to grey-brown ground colour. Tail black above, light brown
below.

Rhabdomys pumilio (Sparrman)

1784. Mus pumilio Sparrman, K. svenska VetenskAkad. Handl. 1784 : 236. Sitzicamma Forest-
on Snake River, east of Cape of Good Hope, South Africa [33° 55' S. 23° 47' E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Mudangi, Bugisu. Text-fig. 8.

MEASUREMENTS. H. & b. $ 106 mm., $ 103 mm. ; tail <J 82 mm., $ 81 mm. ;
h. f. $ 22 mm., $ 20 mm. ; ear <$ 12 mm., $ n mm.

BREEDING. Foetuses (2 x 5, i X 3) in Kenya. From South West Africa 6 and
7 foetuses have been reported (Shortridge, 1934). Breed at 3 months.

HABITAT. In East Africa only recorded from high altitudes. In South West
Africa frequent in bushy and semi-dry vlei country, mainly in scrub, long grass and
forest edges (Shortridge, 1934).

3i6

M. J. DELANY & B. R. NEAL

BIOLOGY. Ground-dwelling but also, to a limited extent, arboreal. Makes bur-
rows. Diurnal. Mainly vegetarian but also recorded as eating snails, insects and
eggs and nestlings of birds.

Genus HYBOMYS Thomas. Back-striped Mice

1910. Hybomys Thomas, Ann. Mag. nat. Hist., (8) 5 : 85. Genotype, by original description,
Mus univittatus Peters.

1911. Typomys Thomas, Ann. Mag. nat. Hist., (8) 7 : 382. Genotype, by original description,
Mus trivirgatus Temminck. Synonym of Hybomys (Ingoldby, Ann. Mag. nat. Hist., (10)
3 : 522).

v Rhabdomys pumllio
• Hybomys univittatus

Edward,

MT.
MUHAVURA

TANGANYIKA

FIG. 8. Distribution of Rhabdomys pumilio and Hybomys univittatus.

THE MURIDAE OF UGANDA 317

Medium sized rats characterized by having a rather indistinct mid-dorsal black
line running from between the ears to the base of the tail. Upperside light brown,
usually with a strong reddish tinge which is particularly well marked on the rump.
Underside light buffy-grey contrasting sharply with the flanks. Scales of tail not at
all obscured by the short hairs. Tail slightly shorter than head-and-body length.
Hindfeet with long toes. Incisors not grooved.

Hybomys univittatus (Peters)

1876. Mus univittatus Peters, Monatsb. K. preuss. Akad. Wiss. Berl. 1876 : 479. Donghila,
Gaboon [o° 12' N. 9° 44' E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Kalinzu Forest, Maramagambo Forest, Ankole ; Malabigambo
Forest, Mpanga Forest, Buganda ; Impenetrable Forest, Kigezi ; Mpanga Forest,
Mubuku Valley, north Ruwenzori, Toro. Text-fig. 8.

MEASUREMENTS. H. & b. ^ 118-5 mm- (8 meas., range 103 to 128 mm.), $ 117-9
mm. (n meas., range 108 to 127 mm.) ; tail $ 105-5 mm- (8 meas., range 85 to 116
mm.), $ 103-9 mm- (I][ meas., range 86 to 115 mm.) ; h. f. <$ 27-2 mm. (8 meas.,
range 26 to 29 mm.), $ 26-4 mm. (n meas., range 24 to 28 mm.) ; ear $ 15-6 mm. (8
meas., range 15 to 17 mm.), $ 15-5 mm. (n meas., range 14 to 17-5 mm.) ; weight <$
50-7 g. (6 weighed, range 46 to 61 g.), $ 49-4 g. (7 weighed, range 38 to 56 g.).

BREEDING. Foetuses (2 X 3) (i lactating) Oct. Impenetrable Forest, (1x2)
lactating Jul. Mpanga Forest (Buganda). Two $ lactating Sep. Maramagambo
Forest. Misonne (1963) reports two embryos in Congo animals.

HABITAT. Apparently confined to forest being found in rain forest, transition
forest and mountain forest but not bamboo forest. It has not been found in gallery
forest. It is more abundant in secondary vegetation than in primary forest ; ap-
parently prefers the wetter parts of forest.

BIOLOGY. Reported by the Congo expedition to eat " the red fruit of a lily-like
plant of which chimpanzees are so fond " (Hatt, 1940). Two stomachs examined by
Delany (19646) contained only vegetable matter. Apparently a good swimmer.

Genus AETHOMYS Thomas. Bush Rats

1915. Aethomys Thomas, Ann. Mag. nat. Hist., (8) 16 : 477. As a subgenus of Epimys Troues-
sart = Rattus Fisher ; genotype, by original designation, Epimys hindei (Thomas).

Medium sized rats with tail usually a little shorter than head-and-body length.
Incisor teeth ungrooved. Fur soft, smooth and tidy, but not silky and with a
metallic lustre. Hairs of underside with pure white tips, with long slate grey bases,
mottled when hairs displaced. The backs of the hands and feet are white.

Tail sparsely haired, nearly as long as, or occasionally slightly longer than head and

body; dorsal pelage light brown in colour . . . . . . A. kaiseri

Tail more hairy, barely longer than body without head; dorsal pelage rich warm

brown ............ A. nyikae

3i8 M. J. DELANY & B. R. NEAL

Aethomys kaiseri (Noack)

1887. Epimys kaiseri Noack, Zool. Jb. Syst. 2 : 228. Qua Mpala (Marungu), southern Congo
[6°46'S. 29°32'E.].

DESCRIPTION. Tail sparsely haired ; upper side dark brownish grey or brown ;
underside mainly white, grey bases of hairs show if the fur is ruffled.

DISTRIBUTION. Chua, Fort Patigo, Paraa, Acholi ; Entebbe, Kabanyolo, Kabu-
lamuleri, Kakumiro, Kampala, Kikonda, Kisingo, Lialo, Nabugabo, Nkyanuna, Zika
Forest, Buganda ; Tororo, Bukedi ; Hoima, Bunyoro ; Moroto, Nabilatuk, Namalu,
Karamoja ; Ajeluk, Serere, Teso ; Kimara, Wanka R., Wassa R., Toro ; Nebbi,
Ngal, Offude, Pakwach, Rhino Camp, West Nile. Text-fig. 9.

MEASUREMENTS. H. & b. <$ 160-4 mm. (18 meas., range 140 to 184 mm.), $
149-5 mm. (10 meas., range 135 to 169 mm.) ; tail <$ 156-8 mm. (18 meas., range 140
to 186 mm.), $ 145-8 mm. (10 meas., range 121 to 180 mm.) ; h. f. ^ 29-2 mm. (17
meas., range 26 to 32 mm.), $ 29-0 mm. (10 meas., range 26 to 32 mm.) ; ear <£ 19-0
mm. (13 meas., range 17 to 21 mm.), $ 19-0 mm. (8 meas., range 16 to 23 mm.) ;
weight <$ 109-0 g. (10 weighed, range 62 to 150 g.), $ 82-4 g. (7 weighed, range 58 to
100 g.).

BREEDING. Foetuses (i x 3) Sep. Kabanyolo, (3 x 3) Kampala (Hopkins MS.).
Three large young observed attached to nipples of $ in Tanzania (Allen & Loveridge,
1942).

HABITAT. Open country, thick grassland. Southern & Hook (19630) report
finding this rat in the ground layer of seasonal forest, the swamp edge of lakeside
forest and in deserted shambas at the edge of forest.

BIOLOGY. Occasionally enters houses (Hopkins MS.).

Aethomys nyikae (Thomas)

1897. Mus nyikae Thomas, Proc. zool. Soc. Lond. 1897 : 431. Nyika Plateau, northern Nyasa-

land [approximately 10° 30' S. 33° 52' E.].
1907. Mus walambae Wroughton, Mem. Manchr. lit. phil. Soc. 51 : 21. Msofu River, Rhodesia

[close to 13° 30' S. 29° E.].

DESCRIPTION. Tail more hairy and the underside a darker grey than in A.
kaiseri. Dorsal pelage a rich warm brown. Tail shorter in proportion to head-and-
body length than A . kaiseri.

DISTRIBUTION. Kagambah, Mbarara, near Lake Nakivali, Ankole ; Koki Co.,
Buganda ; " Kigezi ". Text-fig. 9.

MEASUREMENTS. Two ^ h. & b. 159 mm., 162 mm. ; tail 124 mm., 150 mm. ;
h. f. 28 mm., 28-5 mm. ; ear 21 mm., 22 mm. ; none weighed. $ h. & b. 167-9 mm-
(7 meas., range 148 to 196 mm.) ; tail 138-4 mm. (7 meas., range, 131 to 151 mm.) ;
h. f. 30-3 mm. (7 meas., range 28 to 33 mm.) ; ear 22-3 mm. (7 meas., range 20 to 24
mm.) ; none weighed.

THE MURIDAE OF UGANDA

319

BREEDING. Foetuses, in Zambia, (i x 2), (i x 4), (i X 5) (Ansell, 1960).
HABITAT. In Zambia, usually in ant hills in woodland (Ansell, 1960).
BIOLOGY. No information available.

Genus RATTUS Fischer. House Rats

1803. Ruttus [sic] Fischer, Natmus. Naturg. Paris 2 : 128. Genotype, by subsequent designa-
tion (Hollister, 1916, Proc. biol. Soc. Wash. 29 : 126), Mus decumanus Pallas = Mus norvegicus
Berkenhout.

1881. Epimys Trouessart, Bull. Soc. Sci. Angers 10 : 117. As a subgenus of Mus Linnaeus ;
genotype Mus rattus Linnaeus.

• Aethomys kaiseri
v Aethomys nyikae

SUDAN

Edward,

MT.
MUHAVURA

TANGANYIKA
!

FIG. 9. Distribution of Aethomys spp.

320 M. J. DELANY & B. R. NEAL

Medium sized rats with incisor teeth ungrooved. Dorsal pelage not striped, rather
long and slightly harsh ; varying shades of grey and brown. Underside varies in
colour from creamy to dark slate, never pure white. No clear demarcation between
flanks and belly. Tail usually longer than head and body, and uniformly dark coloured
along length.

Rattus rattus (Linnaeus)

1758. Mus rattus Linnaeus, Syst. Nat., loth ed., i : 61. Uppsala, Sweden [59° 55' N. 18° 08'

E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Patiko, Acholi ; Gayoza, Kichwamba, Ankole ; Entebbe,
Kampala, Mabira Forest, Masaka, Buganda ; Biso, Budongo Forest, Butiaba,
Masindi Port, Bunyoro ; Iriri, Moroto, Namalu, Karamoja ; Nyakabande, Kigezi ;
Kapiri, Serere, Teso ; Bundibugyo, Fort Portal, Isungo, Katwe, Mihunga, Muhokya,
south east Ruwenzori, Toro ; Rhino Camp, West Nile. Probably occurs in all towns
throughout Uganda. Text-fig. 10.

MEASUREMENTS. H. & b. ^ 142-2 mm. (9 meas., range 109 to 193 mm.), 3 £
measured, 126 mm., 150 mm., 170 mm. ; tail <$ 172-6 mm. (9 meas., range 148 to 198
mm.), $ 124 mm., 189 mm., 192 mm. ; h. f. ^ 31-7 mm. (9 meas., range 31 to 33 mm.),
$ 29 mm., ?, 33 mm. ; ear $ 21-3 mm. (9 meas., range 19 to 24 mm.), $ 21 mm., ?,
22-5 mm. ; weight <$ 62-5 g. (5 weighed, range 47 to 92 g.), $ 58 g., 100 g., ?.

BREEDING. Breeding data from Kenya and Congo (Misonne, 1963) indicate
that Rattus breeds throughout the year, with a peak during the long rains in the
early part of the year and with a second smaller peak during the later rains.
Breeding at a minimum during the dry seasons. The average number of foetuses in
54 pregnant females collected in Kampala was 6 (Hopkins MS.). Watson (1950)
reports that the average number in a litter is 6 or 7 in Uganda. The young when born
are blind with the external ears sealed down. The eyes open on the fourteenth day,
weaning takes place about a month after birth. The gestation period is 21 days.

HABITAT. Essentially a dweller in huts and houses ; when found in open usually
in close proximity to buildings. Generally found in thatch of huts, but occasionally
may be found in the walls and floor.

BIOLOGY, Exclusively nocturnal. Omnivorous, food includes all kinds of grain,
groundnuts, cotton seed, meat, potatoes etc. Competes with Mastomys, the other
commensal rat, which it has now excluded from most towns. It is a vector of the flea
Xenopsilla and there is no doubt that it is also the principle vector of plague in
Uganda (Hopkins MS.). Allen & Loveridge (1942) report that predators include the
owl (Bubo africanus), brown house snake (Boaedon lineatus), gaboon viper (Bitis
gabonica), nose-horned viper (Bitis nasicornis) and the black-necked cobra (Naja
nigricollis) .

THE MURIDAE OF UGANDA

Genus PRAOMYS Thomas. Soft-furred Rats

321

1915. Praomys Thomas, Ann. Mag. nat. Hist., (8) 15 : 477. As a subgenus of Epimys Troues-
sart = Rattus Fischer ; genotype, by original designation, Epimys tullbergi (Thomas).

Similar to Mastomys with fur of silky texture ; tail appreciably longer and only
three pairs of mammae present. Tail almost naked. Fur brown to black dorsally,
greyish white ventrally.

Rattus rattus
Praomys morio

SUDAN

Edward,

MT
MUHAVURA

FIG. 10. Distribution of Rattus rattus and Praomys morio.

322 M. J. DELANY & B. R. NEAL

Praomys mono (Trouessart)

1881. Mus morio Trouessart, Bull. Soc. Sci. Angers 10 : 121. Cameroon Mountains [4° 13' N.

9° 10' E.].
1897. Mus jacksoni de Winton, Ann. Mag. nat. Hist., (6) 20 : 318. Entebbe.

DESCRIPTION. As for genus.

DISTRIBUTION. Maramagambo, Kalinzu Forests, Ankole ; Entebbe, Kabanyolo,
Kabulamuliro, Kampala, Kikandwa, Kisimbiri, Lunyo Forest, Malabigambo Forest,
Mpanga Forest, Nabugabo, Zika Forest, Buganda ; Budongo, Bugoma Forests,
Bunyoro ; Echuya, Impenetrable Forests, Kumba, Kigezi ; Benet, Sebei ; Bundi-
bugyo, Bwamba, Kimara, Makoga, Mihunga, Mongiro, Mpanga Forest, Mubuku
Valley, Wasa River, Toro. Text-fig. 10.

MEASUREMENTS. H. & b. ^ 119-9 mm- (5° meas., range 97 to 140 mm.), $ 115-3
mm. (45 meas., range 95 to 135 mm.) ; tail <$ 138-4 mm. (49 meas., range 121 to 160
mm.), $ 136-1 mm. (45 meas., range 94 to 160 mm.) ; h. f. ^ 25-0 mm. (43 meas., range
22 to 27 mm.), $ 24-2 mm. (41 meas., range 21 to 27 mm.) ; ear <$ 17-5 mm. (43 meas.,
range 15 to 20 mm.), $ 17-4 mm. (39 meas., range 15 to 19 mm.) ; weight <$ 44-1 g.
(40 weighed, range 21 to 57 g.), $ 37-0 g. (40 weighed, range 21 to 55 g.).

BREEDING. Foetuses (1x5 lactating) Oct. Echuya ; (1x3) Sep. Chambura ;
(i X 5), (i X 4 lactating) Sep. Maramagambo. Three lactating, Mpanga (Buganda),
May. Litters 2 to 6, normally 3 or 4 (Hopkins MS.). Foetuses (i X 5) in Zambia.

HABITAT. Typically medium and high altitude forest.

BIOLOGY. Nocturnal. Omnivorous ; plant and insect remains found in stomachs.

Genus HYLOMYSCUS Thomas. Climbing Wood-mice

1926. Hylomyscus Thomas, Ann. Mag. nat. Hist., (9) 17 : 178. Genotype, by original designa-
tion, Epimys aeta Thomas.

The climbing mice are small with the tail always longer than the head-and-body
length ; they are very like Praomys. They differ from the latter in their smaller
size and in the hind foot rarely attaining a length of 22 mm. whereas in Praomys it is
always at least 22 mm.

Underside whitish grey with buff, flanks grey to dull brown . . H. denniae

Underside white or silvery grey, flanks rich buff ..... H. Stella

Hylomyscus denniae (Thomas)
1906. Mus denniae Thomas, Ann. Mag. nat. Hist., (7) 18 : 144. Mubuku Valley, Toro.

DESCRIPTION. Dorsal surface grey to buff, the fur soft and rather woolly. Under-
side whitish grey touched with buff, well demarcated from the flanks. Tail longer
than head and body.

DISTRIBUTION. Mpanga Forest, Buganda ; Impenetrable Forest, Kigezi ; Mubuku
Valley, Toro. Kokanjiro. Text-fig, n.

THE MURIDAE OF UGANDA

323

MEASUREMENTS. H. & b. ^ 89-8 mm. (21 meas., range 71 to 103 mm.), $ 87-2 mm.
(14 meas., range 76 to 99 mm.) ; tail <J 125-6 mm. (21 meas., range 84 to 154 mm.),
$ 121-2 mm. (14 meas., range 97 to 145 mm.) ; h. f. <$ 20-1 mm. (21 meas., range 18 to
22 mm.), $ 19-7 mm. (14 meas., range 18 to 21-5 mm.) ; ear <$ 16-9 mm. (21 meas.,
range 13 to 21 mm.), $ 15-4 mm. (14 meas., range 13 to 20 mm.) ; weight <$ 17-3 g.
(12 weighed, range 8 to 24 g.), <j> 20-0 g. (n weighed, range 12 to 42 g.).

BREEDING. No data available. Large numbers of males with small testes sug-
gesting large juvenile population in October in Kigezi.

BIOLOGY. No information available.

Hylomyscus denniae

SUDAN
Hylomyscus stella

TANGANYIKA

(-1

100 miles i

FIG. ii. Distribution of Hylomyscus denniae, H. stella and Myomys fumatus.

324 M. J. DELANY & B. R. NEAL

Hylomyscus Stella (Thomas)

1911. Rattus stella Thomas, Ann. Mag. not. Hist., (8) 7 : 590. Between Mawambi and Avakubi,
Ituri, E. Congo [between i° oo' N. 28° 55' E. and i° 18' N. 27° 32' E.].

DESCRIPTION. Slightly larger in size than H. denniae ; upperside and flanks a
bright ochraceous colour ; underside a whitish or silvery grey colour well demarcated
from the flanks. Tail longer than head and body.

DISTRIBUTION. Kalinzu Forest, Maramagambo Forest, Ankole ; Mabira Forest,
Malabigambo Forest, Mpanga Forest, Zika Forest, Buganda ; Bwamba, Toro.
Text-fig, ii.

MEASUREMENTS. H. & b. ^ 98-3 mm. (6 meas., range 89 to 104 mm.), $ 90-3 mm.
(7 meas., range 85 to 101 mm.) ; tail $ 125-3 nun. (6 meas., range 121 to 133 mm.),
? 131-9 mm. (7 meas., range in to 140 mm.) ; h. f. ^ 17-8 mm. (5 meas., range 17 to
19 mm.), $ 18-0 mm. (7 meas., range 17 to 20 mm.) ; ear <$ 15-0 mm. (5 meas., range
14 to 16 mm)., $ 15-1 mm. (7 meas., range 14 to 16 mm.) ; weight ^ 18-6 g. (6 weighed,
range 16 to 23-5 g.), $ 17-4 g. (7 weighed, range 15 to 22 g.).

BREEDING. Hatt (1940) reports a female from the Congo containing three embryos.
HABITAT. Typically found in rain forest, especially around the bases of trees.
BIOLOGY. No information available.

Genus MYOMYS Thomas. African Meadow Rats

1915. Myomys Thomas, Ann. Mag. nat. Hist., (8) 16 : 447. As a subgenus of Epimys Troues-
sart = Rattus Fischer ; type, by original designation, Epimys colonus (Smith) = Mus colonus
Brants.

The meadow rats have a similar dorsal coloration to Mastomys but are smaller in
size ; the tail is longer than the head and body ; the underside is pure white.

Myomys fumatus (Peters)

1878. Mus fumatus Peters, Mber. preuss. Akad. Wiss. Berl. 1878 : 200. Ukamba, Kenya
[i°-2°S. 37°-38°E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Fort Patiko, Acholi ; Kotido, Nakiloro, Karamoja. Text-fig, n.

MEASUREMENTS. H. & b. <$ 98-0 mm. (5 meas., range 88 to 105 mm.), two $ 75
mm., 103 mm. ; tail <$ 126-4 mm. (5 meas., range no to 150 mm.), $97mm., 130 mm. ;
h. f. <$ 21-8 mm. (5 meas., range 21 to 24 mm.), $ 20 mm., 21 mm. ; ear ^ 16-2 mm.
(5 meas., range 14 to 18 mm.), £ 13 mm., 14 mm. ; weights ^ 30-4 g. (5 weighed, range
24 to 35 g.), $ 12 g., 31 g.

BREEDING. No information available.

HABITAT. Dry savanna.

BIOLOGY. No information available.

THE MURIDAE OF UGANDA 325

Genus MASTOMYS Thomas. Multimammate Rats

1915. Mastomys Thomas, Ann. Mag. nat. Hist., (8) 16 : 477. As a subgenus of Epimys Troues-
sart = Rattus Fischer ; type by original designation, Epimys coucha Smith — Mus marik-
quensis Smith, a race of Mus natalensis Smith.

The multimammate rats have no distinctive markings being very like Praomys in
general appearance. The colour of the dorsal pelage is very variable, usually a grey-
brown, but melanic specimens, entirely coal-black are common, especially in Kigezi.
The underside is also of very variable colour ranging from a silvery grey to a pale
grey touched with buff. The fur is distinctly soft and silky. Tail generally shorter
than head and body. Female with teats in a continuous row, numbering up to 12
pairs, not divided into pectoral and inguinal sets.

Mastomys natalensis Smith

1834. Mus natalensis Smith, 5. Afr. quart. J. 2 : 156. " About Port Natal " = Durban,
Natal [29° 53' S. 31° oo' E.]. The nomenclature of this species is very confusing ; Swyn-
nerton & Hayman (1950) argue cogently in favour of the use of the specific name coucha
Smith.

1836. Mus coucha Smith, Kept, Exp. C. Afr. : 43. 'The country between the Orange River and
the Tropic' of Capricorn.

1897. Mus ugandae De Winton, Ann. Mag. nat. Hist., (6) 20 : 377. Entebbe, Buganda.

1923. Rattus somereni Kershaw, Ann. Mag. nat. Hist., (9) 11 : 594. Kaborini, Bukedi.

DESCRIPTION. As for genus.

DISTRIBUTION. Fort Fatiko, Gulu, Kitgum, Pamdero, Paraa, Acholi ; Burumba,
Kichwamba, north of Maramagambo Forest, Ankole ; Chagwe, Entebbe, Kabanyolo,
Kabulamuliro, Kampala, Kikandwa, Kikonda, Kisimbiri, Kisingo, Lialo, Mabira
Forest, Mengo, Mubende, Buganda ; Kabaroni Camp, Mbale, Bugisu ; Busia, Tororo,
Bukedi ; Fadjao, Hoima, Masindi, Bunyoro ; Isegero, Kama Island, Busoga ;
Amudat, Iriri, Kamchuru, Kotido, Moroto, Nabilatuk, Nakiloro, Namalu, Karamoja ;
Kumba, Kigezi ; Kacheba, Kibusi, Ngai, Lango ; Ajeluk, Serere, Teso ; Bundi-
bugyo, Crater Track, Kamulikwezi, Kimara, Makoga, Mubuku Valley, Mweya, Wasa
River, Toro ; Login, Rhino Camp, Vurra, Yumbe, West Nile. Usaga. Text-fig. 12.

MEASUREMENTS. H. & b. <$ 122-8 mm. (57 meas., range 90 to 154 mm.), $ 125-1
mm. (53 meas., range 95 to 148 mm.) ; tail <$ 112-3 mm- (57 meas., range 88 to 150
mm.), $ 114-0 mm. (52 meas., range 95 to 135 mm.) ; h. f. ^ 23-7 mm. (59 meas.,
range 18 to 30 mm.), $ 23-3 mm. (49 meas., range 20 to 27 mm.) ; ear $ 16-6 mm.
(58 meas., range 14 to 24 mm.), $ 16-8 mm. (50 meas., range n to 21 mm.) ; weight
<J 48-0 g. (32 weighed, range 23 to 70 g.), $ 44-6 g. (34 weighed, range 20 to 60 g.).

BREEDING. Female with 2 or 3 embryos Jul., Murchison Falls National Park.
Two females lactating Jun. and Jul. from Mweya and Tororo respectively. Hopkins
(MS.) states that number of embryos varies between 3 and 12, but 16 has been
recorded. First litters are usually small, young females frequently containing 2 to 4
foetuses. In Tanzania, Chapman, Chapman & Robertson (1959) note that the popu-

326

M. J. DELANY & B. R. NEAL

lation drops to a minimum at the end of the dry season and that breeding is at its
maximum towards the end of the rains. Large catches from Uganda in the dry
season and not in breeding condition confirm this.

HABITAT. Practically in all types of habitat including buildings, swamps and
cultivation. It was formerly the prevalent hut-rat throughout Uganda until driven
out by the invading Rattus rattus.

BIOLOGY. Nocturnal. Omnivorous, plant remains were the commonest material
observed in the stomachs ; seeds and insects have been observed. The species is a

Mastomys natalensis

SUDAN

,,?f

Edward,

MT.
MUHAVURA

FIG. 12. Distribution of Mastomys natalensis.

THE MURIDAE OF UGANDA 327

good burrower and climber although less active than R. rattus. Nests in burrows.
This species is a vector of the flea Xenopsylla and in the past has been an efficient
vector of plague. The gaboon viper (Bitis gabonica) has been observed to prey on it
(Allen & Loveridge, 1942).

Genus MALACOMYS Milne-Edwards. Swamp Rats

1877. Malacomys Milne-Edwards, Bull. Soc. philom. Paris 12 : 10. Genotype, by original
designation, Malacomys longipes Milne-Edwards.

The swamp rats are characterized by the great elongation of the hind feet. The
metatarsals are loosely held together so as to splay out on soft ground. The tail is
longer than the head and body ; the underside of the body is greyish.

Malacomys longipes Milne-Edwards

1877. Malacomys longipes Milne-Edwards, Bull. Soc. philom. Paris 13 : 9. Gaboon River, West
Africa [approximately o° 10' N. 10° oo' E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Kalinzu Forest, Ankole ; Kabanyolo, Mpanga Forest, Zika
Forest, Buganda ; Mpanga Forest, Toro. Text-fig. 13.

MEASUREMENTS. H. & b. ^ 154-2 mm. (n meas., range 120 to 172 mm.), $
155-6 mm. (14 meas., range 127 to 183 mm.) ; tail $ 172-6 mm. (12 meas., range 158
to 190 mm.), $ 165-9 mm- (T4 meas., range 151 to 186 mm.) ; h. f. <$ 38-2 mm. (n
meas., range 35-5 to 40 mm.), $ 37-6 mm. (14 meas, range 35 to 40 mm.) ; ear <$ 24-2
mm. (n meas., range 21 to 29 mm.), $ 23-2 mm. (14 meas., range 19 to 28 mm.) ;
weight (J 94-4 g. (9 weighed, range 65 to 120 g.), <j> 93-3 g. (13 weighed, range 50 to 130
g-)-

BREEDING. Foetuses (i X 3) reported from Congo (Hopkins MS.) ; Ansell (1960)
reports (i x 3) from Zambia.

HABITAT. Wetter parts of forests.

BIOLOGY. Believed to be nocturnal. Omnivorous ; food includes vegetable
matter, insects, slugs and toads (Hopkins MS.). Makes a grass nest on the ground,
also a climber (Misonne, 1963).

Genus ZELOTOMYS Osgood. Broad-headed Mice

1910. Zelotomys Osgood, Publ. Field Mus. Nat. Hist., Zoo/., ser. 10, no. 2 : 7. Genotype Mus
hildegardeae Thomas.

Medium sized with distinctly pro-odont upper incisors. Dorsal pelage grey-brown,
individual hairs with grey bases and brown tips. Backs of hands and feet brown.
Tail covered in scales ; hairs very sparse ; appreciably shorter than length of head and
body.

ZOOL. 13, 9 !8

328 M. J. DELANY & B. R. NEA

Zelotomys hildegardeae (Thomas)

1902. Mus hildegardeae Thomas, Ann. Mag. nat. Hist., (7) 9 : 219. Machakos, Kenya [i° 31'
S. 37°i5'E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Between Rwempuno and Kaizi Rivers, Ankole ; Crater Track,
Toro. Text-fig. 13.

MEASUREMENTS. Two <$ h. & b. 124 mm., 120 mm., two $ 129 mm., 136 mm. ;
tail $89 mm., 86mm., $92 mm., 90 mm. ; h. f. ^22 mm., 25 mm. ,$21 mm., 22 mm. ;
ear ^ 15 mm., 13 mm., $ 15 mm., 13 mm. ; weights ^ 64 g., 57 g., $ 56 g., 64 g.

Malacomys longipes
Zelotomys hildegardeae
i

Edward,

MT.
MUHAVURA

FIG. 13. Distribution of Malacomys longipes and Zelotomys hildegardeae.

THE MURIDAE OF UGANDA 329

BREEDING. Foetuses (i x 5) Jul., Crater Track.
HABITAT. Imperata grassland and scrub.

BIOLOGY. Examination of stomachs of four animals indicates an insectivorous
diet.

Genus MUS Linnaeus

1758. Mus Linnaeus, Syst. Nat., loth ed. i : Genotype Mus musculus Linnaeus.

1837. Leggada Gray, Charles worth's Mag. nat. Hist., 1 : 586. Genotype Leggada booduga Gray.

The forms of this genus are all small mice (head-and-body length usually under
90 mm.), with the tail almost always shorter than the head and body. The fur is
fairly soft but also quite crisp, falling back stiffly into place after being ruffled. We
have included grata and tenellus with the species minutoides.

1 Underside grey or tinged with buff ......... 2

Underside pure white or nearly so; very small size .... M. minutoides

2 Underside grey ........... M. triton

Underside grey tinged with buff ........ M. bufo

Mus bufo (Thomas)
1906. Leggada bufo Thomas, Ann. Mag. nat. Hist., (7) 81 : 145. Mubuku Valley, Toro.

DESCRIPTION. Large dark species with underside well washed with ochraceous-
buff. Bases of belly hairs slate grey. General colour above dark coppery brown.
Tail shorter than head and body.

DISTRIBUTION. Echuya Swamp, Echuya Forest, Impenetrable Forest, Kumba,
Kigezi ; Mihunga, Mubuku Valley, Toro. Text-fig. 14.

MEASUREMENTS. H. & b. $ 72-3 mm. (n meas., range 68 to 79 mm.), $ 68-9 mm.
(9 meas., range 61 to 73 mm.) ; tail $ 62-1 mm. (n meas., range 53 to 69 mm.),
$ 63-2 mm. (9 meas., range 56 to 68 mm.) ; h. f. <$ 16-3 mm. (n meas., range 15 to 18
mm.), $ 15-6 mm. (9 meas., range 13-5 to 16-5 mm.) ; ear <£ 10-8 mm. (n meas.,
range 9 to 12 mm.), $ 11-9 mm. (9 meas., range 10 to 13 mm.) ; weight <$ 9-1 g. (8
weighed, range 7 to 12 g.), $ 9-3 g. (3 weighed, range 7 to 12 g.).

BREEDING. No data available.

HABITAT. Caught in bamboo, moist montane forest and at edge of sedge swamp.

BIOLOGY. No information available.

Mus minutoides Smith

1834. Mus minutoides Smith, 5. Afr. quart. J. 2 : 157. Cape Town [33° 56' S. 18° 28' E.].

1910. Leggada grata Thomas & Wroughton, Trans, zool. Soc. Lond. 19 : 507. Mubuku Valley,
Toro.

1911. Mus tenellus Heller, Smithson. misc. Coll. 56 : 6. Rhino Camp, West Nile.
1911. Mus bellus Heller, Smithson. misc. Coll. 56 : 8. Rhino Camp, West Nile.

1911. Mus musculoides Heller, Smithson. misc. Coll. 56 : 28. Kabulamuliro, Buganda.

DESCRIPTION. Small species with pure white underside.

330

M. J. DELANY & B. R. NEAL

DISTRIBUTION. Paraa, Acholi ; Congo Road, Kalinzu Forest, Lutoto, Marama-
gambo Forest, north of Maramagambo Forest, Ankole ; Kabanyolo, Kabulamuliro,
Kampala, Kikandwa, Kikonda, Kisimbiri, Kisingo, Lunyo, Mpanga Forest, Nabu-
gabo, Nkyamma, Buganda ; Butiaba, Hoima, Kajuia, Masindi, Bunyoro ; Lotome,
Moroto, Nabilatuk, Karamoja ; Echuya Swamp, Kigezi ; Crater Track, Mweya,
Mihunga, Mubuku Valley, Toro ; Rhino Camp, Wadelai, West Nile. Text-fig. 14.

MEASUREMENTS. H. & b. ^ 60-6 mm. (13 meas., range 45 to 76 mm.), $ 60-8 mm.
(n meas., range 49 to 77 mm.) ; tail $ 43-8 mm. (12 meas., range 35 to 54 mm.),
$ 45-5 mm. (n meas., range 35 to 63 mm.) ; h. f. ^ 13-1 mm. (10 meas., range 12 to

o Mus bufo

• Mus minutoides

v Mus trlton

SUDAN

FIG. 14. Distribution of Mus spp.

THE MURIDAE OF UGANDA 331

14-5 mm.), $ 13-1 mm. (8 meas., range 12 to 14 mm.) ; ear <$ 9-7 mm. (9 meas., range
8 to ii mm.), $ 8-7 mm. (8 meas., range 7 to 10 mm.) ; weight ^ 6-4 g. (7 weighed,
range 4 to 10 g.), $ 5-9 g. (9 weighed, range 2-5 to 10 g.).

BREEDING. Foetuses (i x 5) Jul. Mpanga, (i x 4) Aug. Maramagambo Forest,
(i x 4) Nov. Moroto. Allen & Loveridge (1942) report (i X 7) foetuses and groups
of 3, 6, 7 and 8 blind nestlings in Tanzania. Ansell (1960) reports (i x 3) well
developed foetuses and groups of 4 and 5 juveniles from Zambia.

HABITAT. Very varied ranging from dry sandy ground to forest.

BIOLOGY. Nocturnal. Makes shallow burrows with a bed of grass in the bottom.
Predators include the brown house snake (Boaedon lineatus), the burrowing viper
(Atractaspis bibroni), mamba (Dendraspis sp.) and viper (Atheris nischei). Omni-
vorous, the stomachs containing leaves, seeds and insect remains.

Mus triton (Thomas)

1909. Leggada triton Thomas Ann. Mag. not. Hist., (8) 4 : 548. Kirui, southern foothills of Mt.
Elgon, Kenya [approximately o° 45' N. 34° 40' E.].

DESCRIPTION. Differs from Mus bufo in the belly fur not being tinged with buff
and the tail being appreciably shorter.

DISTRIBUTION. Kichwamba, Lutoto, north of Maramagambo Forest, Ankole ;
Entebbe, Kikandwa, Kisimbiri, Masaka, Buganda ; Siroko Valley, Bugisu ; Masindi
Port, Bunyoro ; Echuya Swamp, Ingezi, Kigezi ; north Ruwenzori, Toro. Text-fig.
14-

MEASUREMENTS. H. & b. <$ 78-3 mm. (n meas., range 57 to 93 mm.), $ 73-8 mm.
(10 meas., range 66 to 85 mm.) ; tail ^ 48-5 mm. (10 meas., range 42 to 53 mm.),
$ 46-8 mm. (9 meas., range 44 to 48 mm.) ; h. f. ^ 15-4 mm. (11 meas., range 14 to 19
mm.), $ 14-9 mm. (9 meas., range 14 to 16 mm.) ; ear <$ 10-6 mm. (9 meas., range 9 to
12 mm.), $ 10-7 mm. (9 meas., range 9 to 11 mm.) ; weight <$ 11-4 g. (9 weighed, range
10 to 13 g.), $ 9-4 g. (9 weighed, range 7 to 13 g.).

BREEDING. No data available.

HABITAT. Found in grassland, heath and scrub principally in wetter areas.

BIOLOGY. Nocturnal. Makes nests of both fine and coarse rootlets and leaves of
grass. Loveridge (1953) reports that the brown house snake (Boaedon lineatus) is a
predator.

Genus LOPHUROMYS Peters. Harsh-furred Mice

1866. Lasiomys Peters, Mber. preuss. Akad. Wiss. Berl. 1866 : 409. Genotype = Mus sikapusi

Temminck. Not Lasiomys 1854, in Mammalia (Octodontidae) .
1874. Lophuromys Peters, Mber. preuss. Akad. Wiss. Berl 1874 : 234. Lasiomys afer Peters =

Mus sikapusi Temminck. To replace Lasiomys Peters, preoccupied.

The texture of the fur is very characteristic. It is smooth, sleek, stiff and brush-
like. This is particularly obvious when the fur is stroked against the lie of the hairs.

332 M. J. DELANY & B. R. NEAL

Dorsally the fur is red to brown whilst the underside is rather paler in colour. Eller-
man, Morrison-Scott & Hayman (1953) list two species of short-tailed Lophuromys
both of which occur in Uganda. In addition, two species of long-tailed Lophuromys
have been described from mountainous regions of western Uganda but as they are
obviously very similar they are included here under L. woosnami which is the prior
name.

1 Tail short, usually less than 80 mm. ........ 2

Tail long, usually more than 100 mm. . . . . . . L. woosnami

2 Fur finely speckled ......... L. flavopunctatus

Fur not speckled .......... L. sikapusi

Lophuromys flavopunctatus Thomas

1888. Lophuromys flavo-punctatus Thomas, Proc. zool. Soc. Land. 1888 : 14. Shoa, Ethiopia
(probably obtained at Ankober [9° 32' N. 39° 43' E.], Thomas, 1903, Proc. zool. Soc. Lond.,
1902 : 314).

1892. Mus aquilus True, Proc. U.S. nat. Mus., 15 : 460. Kilimanjaro, Tanzania [3° oo' S.

37° 25' E.].

DESCRIPTION. Dorsal fur dark brown-red speckled with yellow or buff. Belly
buff washed with red, overall effect buffy-pink.

DISTRIBUTION. Burumba, Lutoto, Kalinzu and Maramagambo Forests, between
Rwempuno and Kaizi Rivers, Ankole ; Entebbe, Kabanyolo, Kabulamuliro, Kam-
pala, Kikonda, Kisingo, Lialo, Mabira Forest, Nabugabo, Nalweyo, Zika Forest,
Buganda ; Hoima, Bunyoro ; Echuya Forest, Echuya Swamp, Impenetrable Forest,
Nyakabande, Kigezi ; Bundibugyo, Mihunga, Mpanga Forest, Mubuku Valley, Toro ;
Rhino Camp, West Nile. Text-fig. 15.

MEASUREMENTS. H. &b. ^125-3 mm. (18 meas. .range 117 to 141 mm.),? 125-0 mm.
(20 meas., range 112 to 144 mm.) ; tail <$ 62-8 mm. (17 meas., range 46 to 69mm.),
$ 62-6 mm. (19 meas., range 55 to 94 mm.) ; h. f. $ 20-3 mm. (17 meas., range 19 to
21 mm.), $ 20-1 mm. (18 meas., range 18 to 21 mm.) ; ear <$ 15-2 mm. (15 meas., 13 to
17 mm.), $ 15-5 mm. (18 meas., range 14 to 18 mm.) ; weight <£ 52-0 g. (18 weighed,
range 40 to 62 g.), $ 46-1 g. (18 weighed, range 36 to 56 g.).

BREEDING. Foetuses (2 x 3) Jun. Entebbe ; (4x2) Sep. Echuya. Embryos
near full term weigh 4-5 g. Ansell (1960) reports 3 embryos from Zambia. Litter
size i to 4 (average 2-4) of 43 gravid females examined in Malawi (Hanney, 1964).
Allen & Loveridge (1942) report foetuses (i X 4) from Tanzania and (i X 3) from
Congo.

HABITAT. Moist situations in scrub and forest ; recorded at altitudes up to
12,000 ft.

BIOLOGY. Diurnal and nocturnal. Omnivorous, mainly arthropods ; slugs,
snails, seeds, worms, frogs or toads and birds also recorded. Nests at base of grass
tussocks (Hanney, 1964). Numerous scars on back of Mabira specimens. •

THE MURIDAE OF UGANDA

Lophuromys sikapusi Temminck

333

1853. Mus sikapusi Temminck, Esq. Zool. Cote de Guine : 160. Dabacrom, Ghana [7° 40' N.

2° 58' W.J.

191 1. Lophuromys pyrrhus Heller, Smithson, misc. Coll. 56 : 10. Rhino Camp, West Nile.

DESCRIPTION. Dorsal fur a rich red-brown without any form of speckling.
Underside usually a richer red than L. flavopunctatus . Tail short.

DISTRIBUTION. Kagambah, Kichwamba, between Rwempuno and Kaizi Rivers,
Rutanda, Ankole ; Entebbe, Kampala, Mpanga Forest, Buganda ; Bubungi,

• Lophuromys flavopunctatus
v Lophuromys sikapusi
o Lophuromys woosnami

SUDAN

Edward*

MT.

MUHAVURA
x

FIG. 15. Distribution of Lophuromys spp.

334 M. J. DELANY & B. R. NEAL

Lwakaka, Bugisu ; Butiaba, Bunyoro ; Jinja, Busoga ; Nyalasanje, Kigezi ; Moyo,
Madi ; Serere, Teso ; Bwamba, Crater Track, Kamulikwezi Swamp, Fort Portal,
Ruwenzori, Wasa River, Toro ; Rhino Camp, West Nile. Text-fig. 15.

MEASUREMENTS. H. & b. <$ 133-1 mm. (27 meas., range 100 to 153 mm.), $ 136-6
mm. (16 meas., range 105 to 159 mm.) ; tail <$ 71-6 mm. (26 meas., range 49 to 91 mm.),
$ 72-7 mm. (16 meas., range 56 to 82 mm.) ; h. f. <$ 23-0 mm. (27 meas., range 20 to
25 mm.), $ 22-8 mm. (16 meas., range 21 to 24 mm.) ; ear ^ 14-8 mm. (23 meas.,
range 12 to 17-5 mm.), $ 14-1 mm. (13 meas., range 12 to 16 mm.) ; weight $ 76-8 g.
(20 weighed, range 50 to 104 g.), $ 79-0 g. (14 weighed, range 43 to 100 g.).

BREEDING. Foetuses (i X 2) Aug. Kichwamba. Two $ lactating Jul. Q. E. P.
Usually 2 to a litter (Watson, 1950).

HABITAT. Heavily grassed bush country.

BIOLOGY. Diurnal and nocturnal. Examination of contents of 37 stomachs
suggests that ants form major food. Insectivorous.

Lophuromys woosnami Thomas

1906. Lophuromys woosnami Thomas, Ann. Mag. nat. Hist., (7) 18 : 146. Mubuku Valley,

Toro.
1911. Lophuromys prittiei Thomas, Ann. Mag. nat. Hist., (8) 8 : 377. Mufumbiro region,

Kigezi.

DESCRIPTION. Easily separated from the other two species by its relatively long
tail. The dorsal fur is not as richly coloured as in the other species tending to be
rather more grey-brown. The speckling is sparse or absent. Underside without
trace of pink or red and much more grey to brown.

DISTRIBUTION. Echuya Swamp, Impenetrable Forest, Muhavura, Kigezi ;
Mihunga, Mubuku Valley, north Ruwenzori, Toro. Text-fig. 15.

MEASUREMENTS. H. & b. ^ 118-0 mm. (5 meas., range 114 to 126 mm.), $ 107-2 mm.
(n meas., range 84 to 123 mm.) ; tail $ 113-0 mm. (5 meas., range no to 117 mm.),
$ 107-1 mm. (n meas., range 97 to 118 mm.) ; h. f. $ 23-6 mm. (5 meas., range 22 to
25 mm.), $ 23-1 mm. (n meas., range 22-5 to 26 mm.) ; ear $ 19-0 mm. (5 meas.,
range 19 to 23 mm.), $ 19-1 mm. (n meas., range 17 to 22 mm.) ; weight <£ 45 g. (i
weighed), $ 36-5 g. (7 weighed, range 23 to 48 g.).

BREEDING. Foetuses (i x 2) Sep. Echuya ; (2 x 2) Oct., Impenetrable.

HABITAT. Obtained in moist situations in scrub and forest at altitudes of 6,000 to
8,200 feet.

Genus ACOMYS Geoffroy. Spiny Mice

1838. Acomys Geoffroy, Ann. Sci. nat. Zool. 10 : 126. Genotype, by monotypy, Mus cahirinus
Desmarest.

The spiny mice are small mice characterized by having the hair of the dorsal surface
converted into coarse spines. They inhabit arid semi-desert country. Two species

THE MURIDAE OF UGANDA 335

have been collected in Karamoja from apparently similar types of habitat. Tail
hard and scaly. Incisors ungrooved.

Dorsal pelage uniform grey -brown not speckled . . . . . A. percivali

Dorsal pelage speckled light and dark brown . . . . . A. wilsoni

Acomys percivali Dollman

1911. Acomys percivali Dollman, Ann. Mag. nat. Hist., (8) 8 : 126. Chanler Falls, N. Guaso
Nyiro, Kenya [o° 47' N. 38° 03' E.].

DESCRIPTION. Dorsal pelage uniform grey-brown, tail relatively long, hind foot
large.

DISTRIBUTION. Kotido, Namalu, Karamoja. Text-fig. 16.

MEASUREMENTS. Two $ h. & b. 82 mm., 94 mm. ; tail 67 mm., 84 mm. ; h. f.
15 mm., 15 mm. ; ear n mm., ? ; weights 19 g., 5 g. Two $ h. & b. 93 mm., 74 mm. ;
tail 79 mm., 48 mm. ; h. f. 14 mm., 14 mm. ; ear 12 mm., n mm. ; weights 33 g.,
"g.

BREEDING. Foetuses (i x i) Nov. Kotido.

HABITAT. Dry savanna, semi-desert.

BIOLOGY. No information available.

Acomys wilsoni Thomas

1892. Acomys Wilsoni Thomas, Ann. Mag. nat. Hist., (6) 10 : 22. Mombasa, Kenya [4° 3' S.
39°40'E.].

DESCRIPTION. Dorsal pelage annulated light and dark brown ; annulations may
tend to disappear posteriorly. Tail short ; hind foot small.

DISTRIBUTION. Amudat, Kachere, Lorengikipi, Lotome, Manimani, Moroto
Forest, Nabilatuk, Karamoja. Text-fig. 16.

MEASUREMENTS. Two ^ h. & b. 87 mm., 83 mm. ; tail 43 mm., 45 mm. ; h. f.
12 mm., 13 mm. ; ear 10 mm., 10 mm. ; weights, 22 g., 21 g. Three $ 96 mm.,
86 mm., 84 mm. ; tail 48 mm., 47 mm., 41 mm. ; h. f. 13 mm., 14 mm., 13 mm. ;
ear 12 mm., 10 mm., 10 mm. ; weights 25 g., 27 g., 19 g.

BREEDING. Foetuses (i x 2) Oct. Nabilatuk, (i x i) Nov. Moroto, (i X 3)
Nov. Amudat.

HABITAT. Dry savanna semi-desert. Watson (1950) only found it in less arid
parts or alongside rivers.

BIOLOGY. No information available.

336

M. J. DELANY & B. R. NEAL

Genus URANOMYS Dollman

1909. Uranomys Dollman, Ann. Mag. not. Hist., (8) 4 : 155. Genotype Uranomys ruddi
Dollman.

Texture of fur brush-like ; hairs harsh and long measuring about 17 mm. Back
grey-brown paling to buff on sides and on upper surfaces of limbs. Nasal region and
head darker than back. Similar to Lophuromys but distinguished from it by the
pure white backs to the hands and feet (in Lophuromys they are at least tinged with
black or brown). In Uranomys the belly fur is white and in Lophuromys brown,
orange, red or grey. Upper incisors are orthodont in Lophuromys and slightly pro-
odont in Uranomys.

Acomys percivali
Acomys wilsoni

FIG. 16. Distribution of Acomys percivali, A. wilsoni and Uranomys ruddi.

THE MURIDAE OF UGANDA 337

Uranomys ruddi Dollman

1909. Uranomys ruddi Dollman, Ann. Mag. nat. Hist., (8) 4 : 552. Kirui, southern foothills of

Mt. Elgon, Kenya [approximately o° 45' N. 34° 40' E.].
1911. Uranomys ugandae Heller, Smithson, misc. Coll. 56 : 12. Kikonda, Buganda.

DESCRIPTION. As for genus.

DISTRIBUTION. Kikonda, Buganda ; Lwakaka, Bugisu ; Budama, Bukedi.
Text-fig. 16.

MEASUREMENTS. Two $ h. & b. 104 mm., 95 mm. ; tail 50 mm., 66 mm. ; h. f.
17 mm., 16 mm. ; ear 15 mm., 13 mm.

BREEDING. No information available.

HABITAT. No information available.

BIOLOGY. No information available.

Genus SACCOSTOMUS Peters. Pouched Mice

1846. Saccostomus Peters, Ber. Verh. preuss. Akad. Wiss. Berl. 1846 : 258. Genotype, by
monotypy, Saccostomus campestris Peters.

Medium sized with long, soft, silky fur and a relatively short tail. Cheek pouches
present. Grey dorsally with brown tinge in some specimens ; paler on flanks ; belly
hairs with white tips and slate bases. Ears small and hairy ; tail dark above, paler
below. Backs of hands and feet white. Ellerman, Morrison-Scott & Hayman (1953)
believe there is only one species in this genus.

Saccostomus campestris Peters

1846. Saccostomus campestris Peters, Ber. Verh. preuss. Akad. Wiss. Berl. 1846 : 258. Tete,

Portugese East Africa [16° 10' S. 33° 35' E.].
1936. Saccostomus cricetulus Allen & Lawrence, Bull. Mus. comp. Zool. Harv. 79 : 100. South

bank of Greek River, Sebei.

DESCRIPTION. As for genus.

DISTRIBUTION. Amudat, Lotome, Moroto, Nabilatuk, Karamoja ; Greek River,
Sebei. Text-fig. 17.

MEASUREMENTS. H. & b. $ 116-0 mm. (4 meas., range 94 to 130 mm.), $ 144-9
mm. (7 meas., 127 to 157 mm.) ; tail $ 45-3 mm. (4 meas., range 34 to 55 mm.), $
53-9 mm. (7 meas., range 50 to 58 mm.) ; h. f. <$ 21-0 mm. (4 meas., range 19 to 25
mm.), $ 21-9 mm. (7 meas., range 21 to 22 mm.) ; ear £ 18-3 mm. (4 meas., range 14
to 25 mm.!), 18-3 mm. (7 meas., range 16 to 20 mm.) ; weight $, 2 weighed, 24 g.,
34 g., $ 65-2 g. (6 weighed, range 41 to 84 g.).

BREEDING. Foetuses (i x 7) Nov. Amudat. Ansell (1960) reports (i x 7,
1x6) from Zambia and Shortridge (1934) (i X 8) from South West Africa.

HABITAT. Dry savanna. Attracted to cultivated areas.

BIOLOGY. Nocturnal, very slow moving. From contents of cheek pouches food
apparently largely of seeds, grain, fruits and also occasionally, insects. Lives in
burrows.

338

M. J. DELANY & B. R. NEAL

Genus CRICETOMYS Waterhouse. Giant Rats

1840. Cricetomys Waterhouse, Proc. zool. Soc. Lond. 1840 : 2. As a subgenus of Mus Linnaeus ;
genotype, by original designation, Cricetomys gambianus Waterhouse.

Very large rat with head-and-body length usually exceeding 300 mm. The tail is
longer than the head and body ; it has a dark proximal portion and a white distal
portion. Cheek pouches are present.

v Saccostomus campestris
• Cricetomys gambianus

SUDAN

Edward,

MT. i V \"

MUHAVURAA- v /' S
/ '.'-

L
\Kivu

RUANDA ,.^

<?
^.

FIG. 17. Distribution of Saccostomus campestris and Cricetomys gambianus.

THE MURIDAE OF UGANDA 339

Cricetomys gambianus Waterhouse

1840. Cricetomys gambianus Waterhouse, Proc. zool Soc. Lond. 1840 : 2. River Gambia, West
Africa [13° 30' N. 13° 30'-! 6° 40' W.].

DESCRIPTION. As for genus.

DISTRIBUTION. Aiago River, Fort Patigo, Acholi ; Kampala, Mabira Forest,
Malabigambo Forest, Buganda ; " Bunyoro " ; Jinja, Busoga ; Moroto, Namalu,
Karamoja ; Mt. Sabinio, Kigezi ; Mt. Elgon, Sebei ; Ngora Rest House, Ongino,
Serere, near Soroti, Teso ; Bundibugyo, Mihunga, Mongiro, Mubuku Valley, Toro.
Text-fig. 17.

MEASUREMENTS. Two <$ h. &b. 350 mm., 350 mm. ; tail 380 mm., 418 mm. ; h. f.
70 mm., 72 mm. ; ear 37 mm., 42 mm. Two $ h. & b. 335 mm., 330 mm. ; tail 370
mm., 390 mm. ; h. f. 66 mm., 66 mm. ; ear ? mm., 42 mm. ; weight 910 g., ?.

BREEDING. Foetuses (i X i) Mt. Sabinio (Hopkins MS.) ; (i X i) Congo
(Misonne, 1963) ; (1x4) Zambia (Ansell, 1960).

HABITAT. Ubiquitous, being found in rain forest, mountain forest, gallery forest
and very dry savanna.

BIOLOGY. Strictly nocturnal. Herbivorous, feeding mainly on seeds of trees.
They live in deep burrows in the ground and do serious damage to agriculture. It is
reported that they often climb trees and shrubs in search of fruit. Infected with an
ectoparasitic Dermapteran (Hemimerus talpoides Walker) peculiar to itself.

Subfamily DENDROMURINAE Allen

1939. Dendromurinae Allen, Bull. Mus. comp. Zool. Harv. 83 : 349.

Genus DENDROMUS Smith. African Tree Mice

1829. Dendromus Smith, Zool. J. 4 : 438. Genotype, by original designation, Dendromus typus
Smith = Mus mesomelas Brants.

1830. Dendromys Fischer, Synop. Anim. Add. : 658. Substitute for Dendromus Smith.

1916. Poemys Thomas, Ann. Mag. not. Hist., (8) 18 : 238. As a subgenus of Dendromus
Smith ; type, by original designation, Dendromus melanotis Smith.

Small mice. The fore feet have three well-developed digits only ; hind feet narrow
with very short hallux and fifth digit nearly as long as second. There is often a single
dark, dorsal stripe along the length of the body. The upper incisors are grooved.
Bohmann's (1942) revision of the genus has been adopted together with Ellerman,
Morrison-Scott & Hayman's (1953) use of the specific name mystacalis in place of
pumilio.

1 No dorsal stripe .......... D. mystacalis

Dark dorsal stripe running the length of the body ...... 2

2 Small size; dorsal stripe broken to form spot on the head; tail shorter than h. & b. ;

hind foot less than 18 mm. ........ D. melanotis

Larger size; dorsal stripe not broken to form a spot on the head; tail longer than

h. & b. ; hind foot more than 19 mm. ...... D. mesomelas

340 M. J. DELANY & B. R. NEAL

Dendromus melanotis Smith

1834. Dendromus melanotis Smith, S. Afr. quart. J. 2 : 158. Near " Port Natal " = Durban,

Natal [29° 53' S. 3i°oo'E.].
1911. Dendromus spectabilis Heller, Smithson. misc. Coll. 56 : 3. Rhino Camp, West Nile.

DESCRIPTION. Colour of upper parts light brown, gradually turning a grey-brown
on the sides to a pale grey on the under parts. A wide black median dorsal stripe
runs from the shoulders to the base of the tail, widest anteriorly and narrowing
gradually posteriorly. A median black spot occurs on the forehead between the ears
and eyes. At the anterior base of the ears are a few white hairs and a larger white
patch just below the ear. Hind foot less than 18 mm.

DISTRIBUTION. Hoima, Bunyoro ; Mweya, Toro ; Rhino Camp, West Nile.
Text-fig. 18.

MEASUREMENTS. Three $ h. & b. 67 mm., 73 mm., 61 mm. ; tail 65 mm., 68 mm.,
? ; h. f. 16 mm., 17 mm., 16 mm. ; ear 13 mm., 12 mm., n mm. ; weight 5 g., 7 g.,
6 g. Two $ h. & b. 91 mm., 56 mm. ; tail 67 mm., 64 mm. ; h. f. 16 mm., 16 mm. ;
ear 10 mm., ? ; weight 8 g., 7 g.

BREEDING. Foetuses (i x 3) Jul. Mweya. Ansell (1960) reports (4 x 3, 4 X 4,
i X 5, 4 X 6) from Zambia.

HABITAT. Found in short herbs where D. mystacalis is uncommon. Typically in
dry savanna.

BIOLOGY. Nocturnal. Herbivorous, although Ansell (1960) reports that they
also eat insects. Apparently nest in burrows in the ground in Zambia but in the Congo,
Misonne (1963) reports that they build nests in herbs at heights ranging from 10 cm.
to i m. above the ground. (Further information under D. mystacalis.}

Dendromus mesomelas (Brants)

1827. Mus mesomelas Brants, Het Geslacht der Miuzen : 122. " Near Zondags River " (Sundays
River, just east of Port Elizabeth, Eastern Cape Province) [approximately 33° 45' S. 25° 45'

E.].

DESCRIPTION. Colour of upper parts light brown, the sides not grey-brown as in
D. melanotis. Dorsal stripe distinctly narrower than in D. melanotis. No white
patch as base of ears. Tail longer than head and body. Hind foot longer than 19
mm.

DISTRIBUTION. Echuya Swamp, Kumba, Kigezi ; Mubuku Valley, Toro. Text-
fig. 18.

MEASUREMENTS. H. & b. ^ 80-8 mm. (4 meas., range 76 to 87 mm.), $ 81-3 mm.
(3 meas., range 78 to 87 mm.) ; tail $ 89-8 mm. (4 meas., range 86 to 93 mm.), $
97-0 mm. (3 meas., range 93 to 102 mm.) ; h. f. <$ 21-0 mm. (4 meas., range 20 to 22
mm.), $ 21-0 mm. (3 meas., all 21 mm.) ; ear ^ 12-5 mm. (4 meas., range n to 15 mm.),
$ 13-0 mm. (3 meas., range 12 to 14 mm.) ; weight <$ 13-7 g. (3 weighed, range n to
15 mm.), $ two weighed, 13 g., 15 g.

THE MURIDAE OF UGANDA 341

BREEDING. Allen & Loveridge (1942) record litters of 3 and 4 young.
HABITAT. Found in swamp and associated vegetation in Kigezi.

BIOLOGY. Ansell (1960) reports that they feed largely on grass seeds but are to
some extent insectivorous. Live in tall grass in which they are arboreal ; also semi-
terrestrial. Misonne (1963) claims that they are often caught around villages in the
Congo. Apparently less communal than D. mystacalis and D. melanotis.

v Dendromus melanotis
o Dendromus mesomelas
• Dendromus mystacalis

SUDAN

MI. IQ y x-

MUHAVURAAr \ ^Jf V

s * *

Edward,

FIG. 1 8. Distribution of Dendromus spp.

342 M. J. DELANY & B. R. NEAL

Dendromus mystacalis Heuglin

1863. Dendromus mystacalis Heuglin, NovaActaLeop. Carol. 30, art. 2. suppl. : 5. Ifag, east of

Lake Tana, Ethiopia [12° 15' N. 37° 45' E.].
1911. Dendromus lineatus Heller, Smithson, misc. Coll. 56 : 4. Rhino Camp, West Nile.

DESCRIPTION. No dorsal stripe running the length of the body. Tail about one
and a third times length of the head and body.

DISTRIBUTION. Buligi, Kampala, Kawenge, Buganda ; Budadin Camp, Bugisu ;
Hoima, Kajuia, Masindi, Bunyoro ; Kamchuru, Karamoja ; Sebei Camp, Sebei ;
Bubukwanga, Bugoye, Bumatta, Bummaddu, Bundibugyo, Bundimali, Fort Portal,
Humya, Kyabombo, Mihunga, Tokwe, Toro ; Rhino Camp, West Nile. Text-fig. 18.

MEASUREMENTS. H. & b. <$ 69-2 mm. (20 meas., range 59 to 68 mm.), $ 61-7 mm.
(6 meas., range 50 to 73 mm.) ; tail <? 90-4 mm. (20 meas., range 81 to 101 mm.),
? 86-0 mm. (6 meas., range 75 to 95 mm.) ; h. f. <£ 16-6 mm. (21 meas., range 14 to 19
mm.), $ 16-2 mm. (6 meas., range 16 to 17 mm.) ; ear <$ 12-8 mm. (21 meas., range 10
to 14 mm.), $ 12-3 mm. (6 meas., range 10 to 13 mm.) ; weight $ 8-7 g. (15 weighed,
range 7-5 to 10-5 g.), ? 7-6 g. (4 weighed, range 6-5 to 9 g.).

BREEDING. Normally 3 in a litter, occasionally 4 (Hopkins MS.). Foetuses
(i x 5) in Zambia (Ansell, 1960). Three naked, blind nestlings, Jan. Mihunga ; 4
juveniles Jan. Bugoye. Seven naked nestlings and 4 furred with eyes open in
Tanzania (Allen & Loveridge, 1942). Juveniles in Zambia : (5 x 4, i X 6) (Ansell,
1960).

HABITAT. Common in banana shambas where the nests are usually among the leaf-
bases (Hopkins MS.). Also in grass and amongst herbage.

BIOLOGY. Nocturnal and herbivorous. Quarrelsome and aggressive animals.
D. mystacalis and D. melanotis fight constantly when placed together ; in the course
of the fighting the subordinate animal puts itself on its back and defends itself ;
D. mesomelas is the less aggressive. Dendromus walks with the first and fifth digits
at right angles. Dendromus dominates Mus minutoides and M. triton when they are
together. They climb easily, the tail is prehensile and their light weight allows them
to scale the lighter herbage, where they may construct nests.

Genus STEATOMYS Peters. Fat Mice

1846. Steatomys Peters, Ber. Verh. preuss. Akad. Wiss. Berl. 1846 : 258. Genotype, by mono-
typy, Steatomys pratensis Peters.

The fat mice are small with relatively short tails. Their plump appearance is due
to a layer of fat beneath the skin. Upper incisors grooved. Fur of the back light
brown-fawn ; sharp line of demarcation between flanks and belly ; hairs of the latter
pure white. Hairs of back grey with brown tips. Backs of hands and feet white.
Tail brown above, white below, moderately haired.

THE MURIDAE OF UGANDA 343

Steatomys parvus Rhoads

1896. Steatomys parvus Rhoads, Proc. Acad. not. Sci. Philad. 1896 : 529. Reshiat, Lake
Rudolf, Ethiopia [approximately 4° 33' N. 36° oo' E.].

DESCRIPTION. As for genus.

DISTRIBUTION. Lotome, Nabilatuk, Napyananya, Karamoja. Text-fig. 19.

MEASUREMENTS. Two <$ h. & b. 59 mm., 60 mm. ; tail 37 mm., 35 mm. ; h. f.
13 mm., 15 mm. ; ear 10 mm., 13 mm. ; weight 5 g., ?.

BREEDING. No information available.
HABITAT. Dry savanna, semi-desert.
BIOLOGY. No information available.

Genus DEOMYS Thomas

1888. Deomys Thomas, Proc. zool. Soc. Lond. 1888 : 130. Genotype Deomys ferrugineus
Thomas.

Similar to Grammomys with rufous upperside, white underside and pencilled tail 1-5
times head-and-body length. Differences include the presence of stiff fur along the
back (soft in Grammomys), two faint grooves on the upper incisors (none in Gram-
momys), long and narrow snout of the skull (short and broad in Grammomys) and
elongate feet (Grammomys short).

Deomys ferrugineus Thomas

1888. Deomys ferrugineus Thomas, Proc. zool. Soc. Lond. 1888 : 130. Lower Congo.

DESCRIPTION. As for genus.

DISTRIBUTION. Bwamba Forest, Toro. Text-fig. 19.

MEASUREMENTS. One $ h. & b. 122 mm. ; tail 191 mm. ; h. f . 33 mm. ; ear 24 mm.

BREEDING. Foetuses (2 X 2) in Congo (Hatt, 1940).

HABITAT. In the Congo typical of primary forest ; does not penetrate Cynometra
or transitional forest ; rare (Misonne, 1963).

BIOLOGY. Eight stomachs examined of animals caught in the Congo contained
grasshoppers, ants, termites and flesh resembling that of a rat (Hatt, 1940).

Genus DELANYMYS Hayman. Delany's Swamp-mice

1962. Delanymys Hayman, Rev. Zool. Bot. afr. 65 : 129. Genotype, by original designation,
Delanymys brooksi Hayman.

A very small mouse with a relatively long tail. Front feet very small, hind feet
long and narrow. Tail thinly covered with short stiff hairs. Fur very dense and
soft with stout guard hairs on dorsal surface. Basal two-thirds of dorsal and ventral
hairs slate grey ; dorsally, tips of hairs russet or hazel ; ventrally, warm buff.
Black patch between each eye and nostril.

ZOOL. 13, 9 19

344 M. J. DELANY & B. R. NEAL

Delanymys brooksi Hayman

1962. Delanymys brooksi Hayman, Rev. Zool. Bot. afr. 65 : 132. Echuya Swamp, Kigezi.

DESCRIPTION. As for genus.

DISTRIBUTION. Echuya Swamp, Kigezi. Text-fig. 19.

MEASUREMENTS. One ^ h. & b. 57 mm. ; tail 100 mm. ; h. f . 17 mm. ; ear 10 mm. ;
weight 5 g.

BREEDING. No information available.

• Steatomys parvus

v Deomys ferrugineus

o Delanymys brooksi .. X

Edward,

MT.
MUHAVURA

FIG. 19. Distribution of Steatomys parvus, Deomys ferrugineus and Delanymys brooksi.

THE MURIDAE OF UGANDA 345

HABITAT. Sedge swamp in bamboo and montane forest. Occurs in similar
habitat in the Congo (Hayman, 1962).

BIOLOGY. No information available.

Subfamily OTOMYINAE Thomas
1897. Otomyinae Thomas, Proc. zool. Soc. Lond. 1892 : 1017.

Genus OTOMYS Cuvier. Swamp Rats

1823. Otomys Cuvier, Dents. Mamm. : 168. Genotype, by subsequent designation (Sclater,

1899, Ann. S. Afr. Mus. 1 : 195), Otomys irroratus (Brants).
1918. Anchotomys Thomas, Ann. Mag. nat. Hist., (9) 2 : 204. As a subgenus of Otomys Cuvier ;

type, by monotypy and original designation, Euryotis anchietae Bocage.

The genus Otomys is easy to identify as both upper and lower incisors are deeply
grooved. The fur is long, thick and soft. A well haired tail appreciably shorter
than head-and-body length. The ears are small and the face blunt. The colour is
very variable. Swamp rat is probably a misnomer as they can occur in dry situations
some distance from water. Many species of this genus have been described although
according to Bohmann (1952) there are probably only three occurring in Uganda.
In making specific identifications reference has to be made to the number of trans-
verse laminae on the third upper molars. The systematics of this genus are in need
of further investigation.

1 Lower incisors with two deep grooves ....... O. typus

Lower incisors with a deep outer groove and a shallow inner groove ... 2

2 Five or six lamellae on third upper molar ....... O. denti

Seven lamellae on third upper molar ...... O. irroratus

Otomys denti Thomas

1906. Otomys denti Thomas, Ann. Mag. nat. Hist., (7) 18 : 142. Ruwenzori East, Toro.

1915. Otomys kempi Dollman, Ann. Mag. nat. Hist. (8) 15 : 152. Burunga, Mt. Mikeno, Congo

[i° 28' S. 29° 25' E.].

DESCRIPTION. Dark coloured ; dorsal surface brown-black speckled with copper
buff. Backs of hands and feet blackish brown. Ventral surface slaty-black slightly
speckled with buff. Tail black above and below. Lower incisor with a shallow
inner groove, five or six transverse lamellae on third upper molar.

DISTRIBUTION. Mabira Forest, Buganda ; Echuya Swamp and Forest, Kigezi ;
Kibale Forest, Mubuku Valley, Toro. Text-fig. 20.

MEASUREMENTS. Two ^ h. & b. 170 mm., 150 mm. ; tail 96 mm., 95 mm. ; h. f.
26 mm., 27 mm. ; ear 23 mm., 21 mm. ; weight 125 g., ?. Three $ h. & b. 167 mm.,
167 mm., 157 mm. ; tail 94 mm., 94 mm., 89 mm. ; h. f. 26 mm., 27 mm., 27 mm. ;
ear 23 mm., 25 mm., 21 mm. ; weight 120 g., ?, ?.

BREEDING. Litters of two (Misonne, 1963).

346

M. J. DELANY & B. R. NEAL

HABITAT. Difficult to define in view of the limited number of records but ap-
parently occurs in mixed vegetation and forest at various elevations.

BIOLOGY. Preyed on by harrier (Circus macrourus), grass owl (Tyto capensis] and
leopard in Malawi (Loveridge, 1953).

Otomys irroratus (Brants)

1827. Euryotis irroratus Brants, Het Geslacht der Muizen : 94. Uitenhage, Cape of Good Hope

[33° 41' S. 25° 25' E.].
1915. Otomys tropicalis Dollman, Ann. Mag. nat. Hist., (8) 15 : 157. Mt. Kenya, Kenya

[o°o8'S. 37°i5'E.].
1915. Otomys rubeculus Dollman, Ann. Mag. nat. Hist., (8) 15 : 161. Kagambah, Ankole.

o Otomys denti
• Otomys irroratus
^ Otomys typus

SUDAN

Edward,

MT.
MUHAVURA

FIG. 20. Distribution of Otomys spp.

THE MURIDAE OF UGANDA 347

DESCRIPTION. Colour variable but not usually as dark as 0. denti. Distinguished
from the other species on tooth characters.

DISTRIBUTION. Gulu, Acholi ; Burumba, between Rwempuno and Kaizi Rivers,
Ankole ; Bugala Island, Kampala, Masaka, Mbanga Forest, Mubende, Buganda ;
Beelrungi, Bubungi, Likima, Mbale, Bugisu ; " Bunyoro " ; Echuya Swamp,
Kiduha, Kumba, Kigezi ; Fort Portal, Humya, Toro. Kasiba. Text-fig. 20.

MEASUREMENTS. H. & b. $ 168-1 mm. (18 meas., range 124 to 201 mm.), $
164-7 mm- (T4 rneas., range 139 to 183 mm.) ; tail <$ 93-0 mm. (18 meas., range 83 to
112 mm.), $ 84-4 mm. (14 meas., range 69 to 96 mm.) ; h. i. J 29-5 mm. (18 meas.,
range 27 to 34 mm.), 9 28-3 mm. (14 meas., range 26 to 30 mm.) ; ear <$ 22-0 mm.
(18 meas., range 20 to 25 mm.), ? 22-0 mm. (13 meas., range 20 to 25 mm.) ; weight
<$ 95 g., no g. (only 2 weighed), $ 101-3 g- (6 weighed, range 60 to 120 g.).

BREEDING. Foetuses (i x 2) Jul. Mbanga Forest ; (1x2) between Kaizi and
Rwempuno Rivers, Aug. ; (i x i) Oct. Echuya Swamp. One $ from Echuya
lactating in Sep. Two embryos obtained in August weighed 25 g. Apparently no
fixed breeding season in Zambia (Ansell, 1960) ; litter size 2 to 3. Hair and incisors
erupted at birth. Litter size 2 to 4 (never more) in South West Africa where the
young have been found in rough grass nests in dense reed-growth (Shortridge, 1934) .

HABITAT. Grassland, scrub where herbage is fairly dense. Has been obtained at
relatively large distances from water.

BIOLOGY. Herbivorous ; stomachs of nine animals contained leaves and stems.

Otomys typus (Heuglin)

1877. Oreomys typus Heuglin, Reise in Nordost. Afrika 2 : 77. Highlands of Simyen, Ethiopia

[i3°-i4° N. 38°-39° E.].

1891. Otomys jacksoni Thomas, Ann. Mag. nat. Hist., (6) 7 : 304. [Crater of] Mt. Elgon.
1906. Otomys dartmouthi Thomas, Ann. Mag. nat. Hist., (7) 18 : 141. Mubuku Valley, Toro.

DESCRIPTION. Similar to 0. irroratus but separated on tooth characters.

DISTRIBUTION. Mudangi, Bugisu; Arugot (Mt. Elgon), Sebei ; Mubuku Valley,
Toro. Text-fig. 20.

MEASUREMENTS. H. & b. <$ 142-3 mm. (4 meas., range 132 to 150 mm.), $ 140-0
mm. (4 meas., range 135 to 143 mm.) ; tail J 91-0 mm. (4 meas., range 82 to 101 mm.),
$ 80-5 mm. (4 meas., range 64 to 93 mm.) ; h. f. ^ 26-3 mm. (4 meas., range 25 to 27
mm.), $ 24-6 mm. (4 meas., range 21 to 26-5 mm. ); ear $ 24-3 mm. (4 meas., range
22'5 to 25 mm.), $ 23-7 mm. (3 meas., range 21 to 25 mm.). None weighed.

BREEDING. No information available.

HABITAT. Typically an animal of high altitudes occurring at over n,ooo feet on
Mt. Elgon and over 12,000 feet on Ruwenzori. Occurs in Senecio-Lobelia zone
(Misonne, 1963).

BIOLOGY. Apparently active in the early morning and evening. Does not dig
burrows but lives in long tunnels under moss and Sphagnum (Misonne, 1963).

348 M. J. DELANY & B. R. NEAL

GAZETTEER
As much information as was obtained is given for places not located.

Adropi, West Nile ....

Aigo River, Acholi

Ajeluk, Teso ....

Amudat, Karamoja

Amuria, Teso . . .

Anamuget, Karamoja

Arua, West Nile ....

Arugot, north-east Mt. Elgon, Sebei .

Asuya, Acholi ....

Awack, Acholi ....

2° 48'
2° 21'

i°3o'
i°58'

2° 01'

2° 25'

3° or'

2° 57'
2° 39'

N.
N.
N.
N.
N.
N.

3i°i5'E.
3i°55'E.
33° 50' E.

34° 57' E.
33° 38' E.
34° 30' E.
30° 55' E.

not located
N. 32° 36' E.

N. 33° 27' E.

Beelrungi, south Bugisu .

Benet, Sebei .

Biso, Bunyoro

Bokora, Karamoja

Bubukwanga, Toro

Bubungi, Bugisu .

Budadin Camp, Bugisu .

Budama, Bukedi .

Budongo Forest, Bunyoro

Bugala Island, Buganda

Bugiongolo .

Bugoma Forest, Bunyoro

Bugoye, Toro

Buligi, Buganda

Bulisa, Bunyoro

Bumatta, Bwamba Co., Toro

Bummaddu = Bumadu, Toro

Bundibugyo, Toro

Bundimali, Toro

Buruli Co., Buganda

Burumba, Ankole .

Busia, Bukedi

Busingiro, Bunyoro

Butiaba, Bunyoro .

Butiti, Toro .

Buyobo, Bugisu

Bwamba Co., Toro

Bwamba Forest, Toro

i° 20'
i°45'

2° 25'

o°45'

I°05'

o° 40'

i°45'
o° 24'

I" IS'

o° 17'
o° 27'

2° 07'

o° 43'
o° 42'

o°55'

I°25'
1° 00'

0° 28'

i°44'
i°49

o°39
i° 10

o°55
o° 8

not located
N. 34°

N. 31°

N. 34°

N. 30°

N. 34°

not located
N. 34°

N. 31°

S. 32°

not located
N. 31°

N. 30°

N. 31°

not located
N. 30°

N.
N.
N.
S.

N.
N.
N.
N.
N.
N.
N.

30'
3»'
32'
30'
34'

30'
34'
3°
3°

33' E.
25' E.
25' E.
05' E.
20' E.

03' E.
36' E.
10' E.

oo' E.
07' E.
53' E.
25' E.

05' E.
04' E.
03' E.
25' E.
50' E.
05' E.
28' E.
19' E.
32' E.
17' E.
03' E.
06' E.

Chagwe = Kyagwe Co., Buganda
Chua Co., Acholi .
Congo Road (Q.E.P.) Ankole .
Crater Track (Q.E.P.), Toro

o° 24' N.
3° 20' N.
o° 14' S.
o° 07' S.

32°45'E.
33° 05' E.
29° 59' E.
29° 54' E.

Echuya Forest, Kigezi
Echuya Swamp, Kigezi
Elgon, Mt., Sebei .
Entebbe, Buganda

14' S.
14' S.
o8'N.
03' N.

29° 46' E.
29° 46' E.

34° 33' E.

32° 28' E.

THE MURIDAE OF UGANDA

349

Fadjao, Bunyoro ....
Fort Patiko = Baker's Fatiko, Acholi
Fort Portal, Toro ....

Gayaza, Ankole ....
Greek River, south bank, Sebei
Gulu, Acholi ....

Hakitengya, Toro ....
Hoima, Bunyoro ....
Humya — Humiya, Toro

Ilumia, Toro ....

Impenetrable Forest, Kigezi .
Ingezi, Kigezi ....
Iriri, Teso .....
Isegero, Busoga ....
Isungo, Toro ....

Jinja, Busoga ....

Kabanyolo, Buganda

Kabaroni Camp, north Bugisu

Kabula, Buganda ....

Kabulamuliro, Buganda .

Kacheba, Lango ....

Kacheri, Karamoja

Kagambah, Ankole

Kaizi River (Q.E.P.), Ankole .

Kajansi, Buganda ....

Kajuia, Bunyoro ....

Kakumiro, Buganda

Kalinzu Forest, Ankole .

Kalule, Buganda ....

Kama Island, Busoga

Kamchuru, Karamoja

Kampala, Buganda

Kamulikwezi, Toro

Kamulikwezi Swamp, Toro

Kanaba Gap, Kigezi

Kapiri, Teso ....

Kasai Forest = Kasa Forest, Buganda

Kasiba .....

Katwe, Toro ....

Kawenge = Kawempe, Buganda

Kibale Forest = Mpanga Forest, Toro

Kibandama, ? Kigezi

Kibiro, Bunyoro ....

Kibusi — Kibuji, Lango

Kichwamba, Ankole

Kidoko, Bukedi ....

Kiduha, Kigezi ....

Kikandwa, Buganda

2° 15' N.

31° 40' E.

3° 02' N.

32° 21' E.

o° 40' N.

30° 18' E.

o° 45' S.

30° 47' E.

i°36'N.

34° 20' E.

2° 47' N.

32° i8'E.

o° 45' N.

30° 05' E.

i° 25' N.

3i°2i'E.

o° 46' N.

30° 02' E.

o° 53' N.

30° 03' E.

i° 05' S.

29° 49' E.

i° oo' S.

29° 50' E.

2° 06' N.

34° 12' E.

o° 50' N.

33° 35' E.

o° 30' N.

30° 21' E.

o° 27' N.

33° 12' E.

o° 27'

0° 22'

o° 42'

3° 10'
i° oo'

0°25'
0° 12'

o°48'

0° 22'

o°38'
o° 09'

2° 40'

o° 19'
o° 06'
o° 05'

I°I4'

1° 40'
o°i3'

o° 08'
o° 20'
o°33'

I" 53

o° 14

o°52
1° 15

o°37

N. 32° 36' E.

not located
S. 31° 10' E.
N. 32° 13' E.

not located
N. 33° 56' E.
S. 30° 15' E.
S. 29° 51' E.
N. 32° 32' E.

not located

31° 20' E.
30° 07' E.
32° 32' E.
33° 54' E.
33° 35' E.
32° 35' E.
30° 04' E.
30° 09' E.
29° 46' E.
33° 48' E.
32° 02' E.

N.
S.
N.
S.
N.
N.
N.
N.
S.
N.
N.

not located
S. 29° 52' E.
N. 32° 35' E.
N. 30° 24' E.

not located
N. 3i°i5'E.
32° 23' E.
30° 06' E.

34° °7' E.
29° 41' E.

N.
S.
N.
S.

32° 07' E.

350

M. J. DELANY & B. R. NEAL

Kikonda, Buganda . .

Kilembe, Toro . . . . . .

Kimara, Toro . ...

Kisimbiri, Buganda .....

Kisingo, Buganda ......

Kitgum, Acholi ......

Kokanjiro, Mt. Elgon .....

Koki Co., Buganda .....

Kome Island, Buganda .....

Kotido, Karamoja .....

Kumba, Kigezi ......

Kyabombo, Toro ......

Kyatwe, Toro ......

Lialo, Buganda ......

Likima, Bugisu ......

Locihotome, Karamoja .....

Lorengikipi strm., Karamoja ....

Lotome, Karamoja .....

Lungo, Bukedi ......

Lunyo, Bugando ......

Lutoto, Ankole ......

Lwakaka — Lwakhakha, Bugisu

Mabira Forest, Buganda ....

Makoga, Toro ......

Malabigambo Forest, Buganda

Malera, Teso ......

Manimani, Karamoja .....

Maramagambo Forest (Q.E.P.) Ankole
Maramagambo Forest, north of (Q.E.P.), Ankole
Masaka, Buganda .....

Masindi, Bunyoro ......

Masindi Port, Bunyoro .....

Mbale, Bugisu ......

Mbanga Forest = Mpanga Forest, Buganda
Mbarara, Ankole ......

Mengo District, Buganda ....

Mfumbiro — Mufumbiro = Bufumbiro region, Kigezi
Mihunga, Toro ......

M.N.P. = Murchison Falls National Park in south-west

Bunyoro
Mongiro, Toro ......

Moroto, Karamoja .....

Moroto Forest, Karamoja ....

Moruita, Karamoja .....

Moyo, Madi .......

Mpanga Forest — Mbanga Forest, Buganda
Mpanga Forest = Kibale Forest, Toro
Mubende, Buganda .....

Mubuku Valley = Ruwenzori East, Toro
Mudangi, Bugisu ......

1°

16'

31°

31'

0°

12'

30°

or'

0°

50'

30°

15'

0°

24'

32°

29'

0°

45'

3i°

57'

3°

17'

32°

53'

not located

0°

43'

3i°

20'

06'

32°

45'

01'

34°

06'

i°

08'

29°

54'

not located

0°

27'

30°

13'

0°

53'

3i°

57°

£.

not located

2°

20'

33°

5i'

24'

34°

31'

0°

25'

34°

oo'

03'

32°

28'

0°

20'

30°

06'

0°

48'

34°

22'

0°

30'

33°

00'

01'

30°

22'

57'

3i°

33'

1°

26'

34°

08'

2°

19'

34°

39'

0°

25'

29°

52'

0°

15'

30°

03'

0°

20'

3i°

44'

4i'

3i°

43'

x°

42'

32°

05'

!•

04'

34°

ii'

0°

ii'

32°

16'

0°

37'

30°

39'

0°

17'

32°

35'

!•

22'

29°

39'

0°

22'

30°

03'

st Acholi

and north west

<>•

50'

30°

10'

2°

33'

34°

39'

33'

34°

44'

i°

55'

34°

45'

39'

3i°

42'

ii'

32°

1 6'

0°

33'

30°

24'

0°

35'

3i°

23'

0°

22'

30°

01'

10'

34°

29'

THE MURIDAE OF UGANDA 351

Muhavura Mt., Kigezi ......

I°23'S.

29° 40' E.

Muhokya, Toro .......

o° 06' N.

30° 04' E.

Mulanda, Bukedi .......

o° 42' N.

34°oi'E.

Mweya, Toro .......

o° ii' S.

29° 54' E.

Nabilatuk, Karamoja ......

2° 03' N.

34° 35' E.

Nabugabo, Buganda ......

0° 22' S.

3i°53'E.

Nakiloro, Karamoja ......

2° 37' N.

34° 44' E.

Nakivali Lake, Ankole ......

o° 47' S.

30° 53' E.

Nalweyo, Buganda ......

i° 07' N.

31° i6'E.

Namalu, Karamoja ......

i° 49' N.

34° 38' E.

Napyananya, Karamoja .....

i°52'N.

34° 35' E.

Nebbi, West Nile

2° 30' N.

31° 06' E.

Ngai, Lango ........

2° 30' N.

32° 29' E.

Ngal, West Nile

2° 26' N.

3I°29'E.

Ngora Rest House, Teso .....

i°3o'N.

33° 45' E.

Nkyanuna = Kyanuna, Buganda ....

o° 33' N.

32° 14' E.

Nyakabande, Kigezi ......

i° 18' S.

29° 43' E.

Nyalusanje, Kigezi ......

1° GO'S.

29° 58' E.

Offude, West Nile

3° 13' N.

30° 58' E.

Ongino, Teso .......

i°33'N.

33° 59' E.

Packwack, West Nile ......

2° 27' N.

31° 29' E.

Pamdero (M.N.P.), Acholi

2° 22' N.

31° 40' E.

Paraa (M.N.P.), Acholi

2° 15' N.

3i°35'E.

Patiko = Baker's Fatiko, Acholi ....

3° 02' N.

37°2i'E.

Patong, River Naam ......

not located

Q.E.P. = Queen Elizabeth National Park in Kigezi, west Ankole

and south Toro.

Rhino Camp, West Nile .....

2° 58' N.

31° 24' E.

Rukiga Co., Kigezi ......

i° 05' S.

30° 02' E.

Rutanda (Q.E.P.) , Ankole

o° 15' S.

30° 04' E.

Ruwenzori East = Mubuku Valley of British Museum

Ruwenzori Expedition, 1906 ....

O° 22' N.

30° 01' E.

Ruwenzori North, Toro .... about

o° 40' N.

30° 10' E.

Ruwenzori South East, probably around Muhokya .

o° 06' N.

30° 04' E.

Rwamachuchu — Rwamucucu, Kigezi

i° 10' S.

30° 02' E.

Rwempuno River (Q.E.P.), Ankole ....

o° 23' S.

29° 53' E.

Sabinio Mt., Kigezi ......

i°23'S.

29° 36' E.

Salalira, Bugisu .......

i° 14' N.

34° i?' E.

Sara, Toro ........

o° 47' N.

30° 05' E.

Sebei Camp, Teso .......

i° 31' N.

33° 26' E.

Sipi, Bugisu ........

i° 20' N.

34° M' E.

Siroko Valley, Bugisu ......

1° 21' N.

34° H' E.

Soroti, Teso .......

i°44'N.

33° 36' E.

Tokwe, Toro .......

o° 48' N.

30° 02' E.

Tororo, Bukedi .......

o° 41' N.

34° 10' E.

Usaga ......... not located

352 M. J. DELANY & B. R. NEAL

Vurra, West Nile 2° 53' N. 30° 53' E.

Wadelai, West Nile ; . . . . . 2° 42' N. 31° 27' E.

Walasi, Bugisu . . . . . . . i° n' N. 34° 13' E.

Wanka River, Toro o° 51' N. 30° 16' E.

Wasa River, Toro . . . . . . . o° 47' N. 30° 15' E.

Yumbe, West Nile 3° 28' N. 31° 15' E.

Zika Forest, Buganda o° 10' N. 32° 28' E.

REFERENCES

ALLEN, G. M. 1939. Check list of African Mammals. Bull. Mus. comp. Zool. Harv. 83 : 1-763.
ALLEN, G. M. & LOVERIDGE, A. 1942. Scientific results of a fourth expedition to forested areas

in East and Central Africa. I. Mammals. Ibid. 89: 147-214.

ANSELL, W. F. H. 1960. Mammals of Northern Rhodesia. Lusaka: The Government Printer.
BERE, R. M. 1962. The Wild Mammals of Uganda. London : Longmans Green.
BOHMANN, L. 1942. Die Gattung Dendromus. Zool. Anz. 139 : 33-60.

1952. Die afrikanische Nagergattung Otomys Cuvier. Z. Saugetierk. 18 : 1-80.

BRAMBELL, F. W. R. & DAVIS, D. H. D. 1941. Reproduction in the multimammate mouse

(Mastomys erythroleucus] of Sierra Leone. Proc. zool. Soc. Lond. 11 IB : i-n.
CAMBRIDGE NABUGABO BIOLOGICAL SURVEY. 1962. Preliminary Report, pp. 22.
CHAPMAN, B. M., CHAPMAN, R. F. & ROBERTSON, I. A. D. 1959. The growth and breeding of

the multimammate rat, Mastomys natalensis (Smith) in Tanganyika Territory. Proc. zool.

Soc. Lond. 133 : 1-9.
DELANY, M. J. 1964^. A study of the ecology and breeding of small mammals in Uganda.

Ibid. 142 : 347-37°-
19646. An ecological study of the small mammals in the Queen Elizabeth Park, Uganda.

Rev. Zool. Hot. afr. 70 : 129-147.
ELLERMAN, J. R. 1941. The Families and Genera of Living Rodents Vol. II. London : British

Museum (Nat. Hist.).
ELLERMAN, J. R., MORRISON-SCOTT, T. C. S. & HAYMAN, R. W. 1953. Southern African

Mammals 1758-1951 : A Reclassification. London: British Museum (Nat. Hist.).
HANNEY, P. 1964. The harsh-furred rat in Nyasaland. /. Mammal 45 : 345-358.
HATT, R. T. 1940. Lagomorpha and Rodentia other than Sciuridae, Anomaluridae and

Idiuridae collected by the America Museum Congo Expedition. Bull. Amer. Mus. Nat. Hist.

76 : 457-604.
HAYMAN, R. W. 1962. The occurrence of Delanymys brooksi (Rodentia, Muridae) in the Congo.

Bull. Inst. Sci. nat. Belg. 38 : 1-4.
HOLLISTER, N. 1919. East African mammals in the U.S. National Museum II. Rodentia,

Lagomorpha and Tubulidentata. Bull. U.S. nat. Mus. 99 : 1-184.

HOPKINS, G. H. E. Undated. The known Wild Rodents of Uganda. Manuscript, 70 pp.
JOHNSON, Sir H. 1902. The Uganda Protectorate. 2 Vols. London : Hutchinson.
JOHNSON, H. L. & OLIFF, W. D. 1954. The oestrous cycle of female Mastomys natalensis

(Smith) as observed in the laboratory. Proc. zool. Soc. Lond. 124 : 605-613.
LANGDALE-BROWN, I. 1962. Vegetation : Atlas of Uganda. Entebbe : Department of Lands

and Surveys.
LOVERIDGE, A. 1953. Zoological results of a fifth expedition to East Africa. VII. Itinerary

and conclusions. Bull. Mus. comp. Zool. Harv. 110 : 3-80.
MISONNE, X. 1963. Les rongeurs du Ruwenzori et des regions voisines. Exploration du Pare

National Albert, Deuxieme Serie 14 : 1-164.

THE MURIDAE OF UGANDA 353

MOREAU, R. E., HOPKINS, G. H. E. & HAYMAN, R. W. 1964. Type localities of some African

mammals. Proc. zool. Soc. Lond. 115 : 387-447.
OLIFF, W. O. 1953. The mortality, fecundity and intrinsic rate of natural increase of the

multimammate mouse Mastomys natalensis in the laboratory. /. Anim. Ecol. 22 : 217-226.
FETTER, F. 1964. Affinites du genre Cricetomys. Une nouvelle sous-famille de Rongeurs

Cricetidae, les Cricetomyinae. C.r. hebd. Seanc. Acad. Sci,, Paris 258 : 6516-6518.
SCHOUTEDEN, H. 1948. Faune de Congo Beige et du Ruanda-Urundi I. Mammiferes. Ann.

Mus. Congo. Beige., Zool. 1 : 1-331.
SHORTRIDGE, G.C. 1934. The Mammals of South West Africa ; a biological account of the forms

occurring in that region. Vol. i. London : Heinemann.
SIMPSON, G. G. 1945. The principles of classification and a classification of mammals. Butt.

Amer. Mus. nat. Hist. 85 : 1-350.
SOUTHERN, H. N. & HOOK, O. 1963^. A note on small mammals in East African forests. /.

Mammal. 44 : 126-129.
- 1 9636. Notes on breeding of small mammals in Uganda and Kenya. Proc. zool. Soc.

Lond. 140 : 503-515.

SWYNNERTON, G. H. & HAYMAN, R. W. 1950. A checklist of the land mammals of the Tan-
ganyika Territory and the Zanzibar Protectorate. /. E. Afr. Ug. nat. Hist. Soc. 20 : 274-

392.
THOMAS, O. 1906. Descriptions of new mammals from Mount Ruwenzori. Ann. Mag. nat.

Hist., (7) 18 : 136-147.
1907. On further new mammals obtained by the Ruwenzori Expedition. Ibid. 19 : 118-

123.

THOMAS, O. & SCHWANN, H. 1904. On mammals collected during the Uganda Boundary Com-
mission by the late Mr. W. G. Doggett and presented to the British Museum by Col. Delme-

Radcliffe. Proc. zool. Soc. Lond. 1904 : 459-465.
THOMAS, O. & WROUGHTON, R. C. 1909-10. Ruwenzori Expedition reports, 17 : Mammalia.

Trans, zool. Soc. Lond. 19 : 481-554.
WATSON, J. M. 1950. The wild mammals of Teso and Karamoj a. IV. Uganda J. 14 : 53-84.

INDEX

Acomys, 334 bufo, Mus, 329

aeta, Epimys, 322 Bush Rats, 317
aeta, Hylomyscus, 322

Aethomys, 317 cahirinus, Mus, 334

afer, Lasiomys, 331 campestris, Saccostomus, 337

African Meadow Rats, 324 Climbing Wood-mice, 322

African Tree Mice, 339 colonus, Epimys, 324

anchietae, Euryotis, 345 colonus, Mus, 324

Anchotomys, 345 coucha, Epimys, 325

aquilus, Mus, 332 coucha, Mastomys, 325

Arvicanthis, 309 coucha, Mus, 325

Creek Rats, 310

bacchante, Oenomys, 305 Cricetomys, 338

Back-striped Mice, 316 cricetulus, Saccostomus, 337

barbarus, Lemniscomys, 313 cuninghamei, Mylomys, 305
barbarus, Mus, 313

bellus, Mus, 329 dartmouthi, Otomys, 347

booduga, Leggada, 329 Dasymys, 307

Broad-headed Mice, 327 decumanus, Mus, 319

brooksi, Delanymys, 344 Delanymys, 343

bufo, Leggada, 329 Delany's Swamp-mice, 343

354

Dendromus, 339

Dendromys, 339

denniae, Hylomyscus, 322

Deomys, 343

denti, Otomys, 345

Desmomys, 310

discolor, Thamnomys, 304

dolichurus, Grammomys, 304

dolichurus, Mus, 304

Dormice, 298,

dryas, Thamnomys, 304

M. J. DELANY & B. R. NEAL

jacksoni, Mus, 322
jacksoni, Otomys, 347

kaiseri, Aethomys, 318
kaiseri, Epimys, 318
kempi, Otomys, 345
kempi, Thamnomys, 302
Komemys, 310

Euryotis, 345

Epimys, 318, 319, 321, 322, 324, 325.

fallax, Mus, 311
Fat Mice, 342
ferrugineus, Deomys, 343
flavopunctatus, Lophuromys, 332
Four Striped Grass-mice, 315
fumatus, Mus, 324
fumatus, Myomys, 324

gambianus, Cricetomys, 339
Gerbils, 298
Giant Rats, 338
Grammomys, 304
Graphiurus, 298
grata, Leggada, 329
gueinzii, Dasymys, 307

harringtoni, Pelomys, 310
Harsh-furred Mice, 331
hildegardeae, Mus, 328
hildegardeae, Zelotomys, 328
hindei, Epimys, 317
hopkinsi, Pelomys, 313
House Rats, 319
Hybomys, 316
Hylomyscus, 322
hypoxanthus, Mus, 305
hypoxanthus, Oenomys, 305
Hypudaeus, 309

ibeanus, Tachyoryctes, 298
incomtus, Dasymys, 307
incomtus, Mus, 307
irroratus, Euryotis, 346
irroratus, Otomys, 346
Isomys, 309
isseli, Komemys, 310
isseli, Pelomys, 311

Lasiomys, 331
Leggada, 329
Lemmus, 309
Lemniscomys, 313
lineatus, Dendromus, 342
longipes, Malacomys, 327
Lophuromys, 331
lutescens, Mylomys, 305

macculus, Arvicanthis, 313
macculus, Lemniscomys, 313
macmillani, Thamnomys, 304
Malacomys, 327
marikquensis, Mus, 325
Mastomys, 325
medius, Dasymys, 307
melanotis, Dendromus, 340
mesomelas, Dendromus, 340
mesomelas, Mus, 340
minutoides, Mus, 329
montanus, Dasymys, 307
morio, Mus, 322
morio, Praomys, 322
Multimammate Rats, 325
Mws, 304, 305, 307, 309, 311, 313, 315. 317.
318, 319, 320, 322, 324. 325. 328, 329, 331.

332, 333. 340
musculoides, Mus, 329
musculus, Mus, 329
My/owrys, 305
Myomys, 324
mystacalis, Dendromus, 342

natalensis, Mastomys, 325
natalensis, Mus, 325
niloticus, Arvicanthis, 309
niloticus, Arvicola, 309
niloticus, Lemmus, 309
norvegicus, Mus, 319
nyikae, Aethomys, 318
nyikae, Mus, 318

Oewowys, 305
Oreomys, 347

INDEX

355

orthos, Dasymys, 307
Otomys, 345

parvus, Steatomys, 343
Pelomys, 310
percivali, Acomys, 335
Poemys, 339
Pouched Mice, 337
Praomys, 321
pratensis, Steatomys, 342
prittiei, Lophuromys, 334
pumilio, Mus, 315
pumilio, Rhabdomys, 315
pyrrhus, Lophuromys, 333

Rattus, 319, 324, 325
Rhabdomys, 315
rubeculus, Otomys, 346
ruddi, Uranomys, 337
Rusty-nosed Rats, 305
rutilans, Mus, 302
rutilans, Thamnomys, 302

Saccostomus, 337
Shaggy Swamp-rats, 307
sikapusi, Lophuromys, 333
sikapusi, Mus, 333
Soft-furred Rats, 321
somereni, Rattus, 325
spectabilis, Dendromus, 340
Spiny Mice, 334
Steatomys, 342
stella, Hylomyscus, 324
stella, Rattus, 324
striatus, Lemniscomys, 313
striatus, Mus, 313

Striped Grass-mice, 313
surdaster, Thamnomys, 304
Swamp Rats, 345

Tachyoryctes, 298
Tatera, 298
Taterillus, 298
tenellus, Mus, 329
Thamnomys, 302, 304
Thicket Rats, 302
Tree Rats, 304
triton, Leggada, 331
triton, Mus, 331
trivirgatus, Mus, 316
tropicalis, Otomys, 346
tullbergi, Epimys, 321
Typomys, 316
typus, Dendromus, 339
typus, Oreomys, 347
typus, Otomys, 347

ugandae, Mus, 325
ugandae, Uranomys, 337
univittatus, Hybomys, 317
univittatus, Mus, 317
Unstriped Grass-mice, 309
Uranomys, 336

variegatus, Hypudaeus, 309
venustus, Thamnomys, 302

walambae, Mus, 318
wilsoni, Acomys, 335
woosnami, Lophuromys, 334

Zelotomys, 327

INDEX TO VOLUME 13

The page numbers of the principal references and the new taxonomic names arc printed in bold type.

For index to Number 9 see page 353.

ablabes, Barbus

acanthina, Amphiacantha

acanthinobrachia, Ophiobyrsa

acanthinotata, Ophiacantha

Acaronychidae

aculeifer, Gaeolaelaps

aculeata, Ophiopolis

adiaphora, Ophiacantha

agilis, Laelaps

alaudae, Dermanyssus

albomaculata, Ophionereis

alvearius, Melittiphis

amphibia, Hyperlaelaps

Amphiuridae

Ampullariidae

anchista, Pectinura

ancistrotus, Amphioplus

Ancylidae

Androlaelaps . 251, 253,

anema, Barbus

angulata, Ophiothrix

angulosa, Onychocella

angustus, Segmentorbis .

Anisus

aphidinus, Parhypochthonius

arcystata, Amphiura

ardua, Rhysotritia

aridus, Ornithonyssus

aspilus, Barbus

asterictus, Amphioplus .

Asteroschema

atakorensis, Barbus 102,

aurantiacus, Barbus

1 02-105-z 08

. , . .66

40-4i

. 3-io
• 275
. 67
. 67
285, 291

261, 268, 284-286
70

285-286

253, 259, 285-286
. ' . 43-6o

90-91
64-65
51-54
. 90

256-259, 269, 288
104

. 69

. 195

. 82-83, 93

93
10

44-45

22-23

264

104

54-55

. 69

104, 135-137, 148
104

bacoti, Ornithonyssus

Barbus

bawkuensis, Barbus
Beirabarbus .
bengalensis horsfieldi, Felis
berlesei, Oribotritia
bidentata, Ophiacantha
bimorphocella, Selenaria
Biomphalaria
biporosa, Cupuladria

257-259, 261, 266, 268,
285-286
101-149

IO2, 105, 139-141, 148
103-104

193,

147

. 231
26

. 67

165-166

-95-l9(>,
203-209

Bithyniidae .
brachyactis, Ophiopholis .
brasiliensis, Nippostrongylus

callipterus, Barbus
camerunensis, Biomphalaria
camerunensis, Bulinus
Camissidae
canariensis, Cupuladria

154-

165

capriensis, Cupularia
caribbea, Vibracellina
casalis, Androlaelaps
cercops, Barbus
charischema, Ophioleuce
chelidonis, Dermanyssus
chiajei, Amphiura
chimalae, Himalacarus
cinereum, Ophioderma .
collarti, Barbus
condylata, Chaperiella
Conescharellina
congesta, Ophialcaea
congicus, Barbus .
conspicua, Bathypectinura
coretus, Anisus
Cosmochthoniidae
Cosmolaelaps .
costulatus, Gyraulus
craterodmeta, Amphiodia
Crepidacantha
cryptomia, Androlaelaps
Cupuladria . 154, 162

Cupuladriidae
cupulariensis, Beania
cylindrica, Epilohmannia

66
232

104

77-79, 93. 95
87-88
. 33

155, 157, 160, 162,
, 167, 193-197-203
167, 182-184
164
. 285
104

. 67
284-286
. 69
. 3-io
. 69
104

• 195

• 154
41-42

104
70

79-82, 93
10

272, 282
82, 93
58, 67
. 156

• 253

193, 195, 197-219
151-187, 189-227

• 195

danae, Ophiothela ..... 69
deformis, Cupuladria . . . 158, 195
denticulata, Cupuladria .... 193
Dermanyssidae .... 247-294
Dermanyssus 253, 261, 264, 268, 271, 275,

284-285
deserti, Barbus , 103-104

358

INDEX

dhanra, Rictularia . . . .231

digitula, Amphiura . . . 49-5 1

Discoporella 154, 159-160, 162, 193, 196,

219-223

doderoi, Pseudolaelaps . . 271, 285-286
doma, Cupuladria 160, 162-163, I93~I95. 216-

219
duboisii, Stylopoma .... 194

echidnina, Laelaps
elongata, Cupuladria
Eniochthoniidae
Epilohmanniidae .

285

158

Eulaelaps 253, 255, 257, 261, 269, 275-279

Eulohmanniidae ..... 26
euopla, Amphiura . . . 45-47

Euphthiracaridae .... 18-26

eusteira, Ophiothrix .... 60

fahrenholzi, Androlaelaps
felineum, Arthrostoma .
Ferrissia
fissa, Amphiodia
flavus, Macronyssus
flexuosus, Laelaspulus
foetida, Jullienella
forskali, Bulinus
fragilis, Ophiothrix
freethi, Potadoma

Gabbia

gallinae, Dermanyssus
gelida, Ophiurolepis
gigantica, Fasciola
gola, Longistriata
gorgonia, Ophiarachnella
guineensis, Cupuladria

Gymnolaelaps

. 285

231,243-244

. 90

58-6o

268, 274

284-286

. 194

87, 93

. 69

. 91-92, 95

91, 93

. 268
70

231, 239-241
66

160, 162, 166-167,
170-175
261, 277

Haemogamasus 251, 253, 255, 257, 269,

275-277, 291

haematobium, Schistosoma . . 93-94

Heliodoma ..... 154, 164
hilaris, Laelaps ..... 285
himalensis, Citellina . . . .231
Hincksinidae .... 182-184

Hirstionyssus . 254, 261, 284, 288, 291

hirsuta, Macrophiothrix ... 69

hirsutosimilis, Haemogamasus . . 284

hirsutus, Haemogamasus 253, 257, 261,

286-287, 291

hirundinis, Dermanyssus. . . 284-286

Holostaspis ..... 266, 277
horridus, Haemogamasus . . . 253

Hyperlaelaps .... 269, 279

Hypoaspis . . 255, 266, 275-277, 282

Hypochthoniidae ..... 10
hypsolepis, Barbus 102, 104, 117-1 18, 147

indica, Cupuladria

inepta, Amphiura .
inglisi, Euphthiracarus

intermedius, Dermanyssus
intorta, Ophiobyrsa
iridoides, Amphiura
iris, Amphiura

jae, Barbus .
japonicus, Amphioplus .
japonicus, Ophioplocus

kanisaensis, Segmentorbis
kessleri, Barbus
kinbergi, Ophiura .
koreae, Amphiura
koreana, Ophiothrix
krameri, Hypoaspis

156, 167, 169-170
48-4Q

18-22

261
39-40
. 43
43.44

104

55-58
66

. 83-85, 93

104

47-48

61-62

290

Laelaps 251, 253, 256-259, 268, 272, 277, 284,

288

Laelaspulus ..... 259, 277
laevigatus, Phthiracarus . . .15

lanatus, Cosmochthonius ... 10

lapponica, Camisia . . . .33

lawrae, Barbus . 102-104, 121-124, J48

leonensis, Barbus 102, 104-105, 145-147, 148
lepidevaspis, Amphiura .... 67
leptoctenia, Ophiura .... 68
levinseni, Smittipora . . . .195

libycus, Lanistes .... 90-9 1

Liponyssoides ..... 264
longicauda, Ophioderma ' . . .70
longifissus, Asteronyx .... 68
longispina, Ophiozonella ... 67

loveni, Asteronyx ..... 68
lowei, Cupuladria . . . . .158

lubrica, Hypoaspis .... 285

Lunulites . . . 154, 162, 164-166

Lymnaeidae .... 88-90

macclellandi macclellandi, Callosciurus . 231

macinensis, Barbus 102, 105, 130—132, 148

macraspis, Amphioplus .... 68

macrolepidota, Ophiactis

Macronyssus .

macrops, Barbus

maculata, Ophiura

maculata, Selenaria

maerkeli, Mesotritia .

Malaconothridae

Mannicthys

mansoni, Schistosoma

marenzelleri, Ophiothrix

marginalis, Pneumolaelaps

Mastophorus

Melittiphis

microti, Hyperlaelaps

miles, Stratiolaelaps .... 288

minuta, Capillaria . 232

42-43

261, 264, 277-279, 285, 291

102-104, 110-H4, J47

159, 194
24-26
. 33
. 103

94-95
66
269
- 231
• 255, 257
. 285

INDEX

359

minutissimus, Eniochthonius ... 10
mirabilis, Ophiopholis .... 66
mixta, Ophiomastix .... 66
monodactylus, Malaconthrus . . 33

monotrema, Cupuladria 155, 162, 196, 209-2 10
multispinata, Cupuladria 156-158, 162,

193-196, 210-213

multispinosa, Ondatralaelaps . . 285, 288
murinus caerulescens, Suncus . . 232

muris, Laelaps .... 285, 288
muris, Mastophorus . . . 231—232
Myonyssus ..... 257, 277
mystax, Toxocara . . . .231

nana, Nanhermannia
Nanhermanniidae .
natalensis, Lymnaea
natricis, Ophionyssus
nepalensis, Hoplophthiracarus
nidi, Haemogamasus
nidicorva, Hypoaspis
nigeriensis, Barbus
nodosa, Stegophiura
Nothridae
nova, Eulaelaps

. 33
. 33

. 88-90, 95

279, 284-286
15-i8
. 268

. 288

102-104, 119-121, 148

68
26-32

285, 288

ocellata, Discoporella 193, 196,220-221

Ololaelaps . 261, 269, 275-279, 282, 291

Ondatralaelaps .... 269-271

Onychocella . . . . . 195

Ophiacanthidae .... 41-42

Ophiactidae ..... 42-43

Ophiodermatidae .... 68-65

Ophiomyxidae .... 39-41

Ophionyssus . . 261, 264, 275, 282, 285

Ophiotrichidae .... 60-63

Ophiuridae ...... 65

oranense, Cleidochasma . . . .195

Ornithonyssus . . 253, 261, 264, 268

owenii, Cupuladria 160, 162, 166-167, 196,

213-215
owenii disciformis, Cupuladria 193, 197,

215-216

palustris, Barbus . . . . .103

palustris, Nothrus . . . .31

panchyendyta, Ophiothrix . . .62

parablabes, Barbus 102-103, 114-H7, 147

Paragonimus ..... 95

Parhypochthoniidac .... 10
patens, Molineus . . . 231-232-234
Pellonyssus 261, 264, 268, 275-277, 285-286

peltifer, Platynothrus . . . .33
pentagona, Ophiacantha ... 66

periblepharus, Steatonyssus . . . 285

perince, Barbus . . . . .103

permixta, Ophioconis . . . 68-64

pfeifferi, Biomphalaria . . . 79, 95

Phthiracaridae .... Il-i8

Planorbidae . 77-88

pleuropholis, Barbus . . . .104

peyroti, Cupuladria . . . 163-164

Pneumolaelaps .... 268, 285
pobeguini, Barbus . 102, 105, 141-143, 148

pontiger, Haemogamasus . . .261

praesternalis, Hypoaspis . . .269

projecta, Ophiozonella .... 67
pseudognathodon, Barbus . . .104
Pseudolaelaps ..... 255

Pseudoparasitus . 261, 275-277, 282, 291

pteropoma, Ophiactis .... 66
pulchra, Membraniporella . . .195
pumilus, Barbus ..... 104
punctitaeniatus, Barbus 102-104, 137-139, 148
pygerythrus lokroides, Callosciurus . . 231
pyriformis, Cupuladria . 155, 167-168-169

quadrispina, Ophiura .... 68
quintus, Dermanyssus . . . 284-286

radiata, Lunulites
radiatus, Barbus
rattus sikkimensis, Rattus
reedi, Pellonyssus .
reussiana, Cupuladria
rhachophora, Ophiacantha
ribagai, Eulohmannia
robertsi, Phthiracarus
rohlfsi, Bulinus
rufulus, Hypochthonius
rugosum, Ophiopsammium

sagaminum, Astrodendrum

sardoa, Hypoaspis .

sarsi, Ophiura

sarsi vadicola, Ophiura

saurarum, Sauronyssus

Sauronyssus

savigniyi, Ophictais

scolopendrina, Ophiocoma

scutatus, Dermanyssus .

Selenaria

Setosella

Setosellina . . 154,

simplex, Ophiactis

skrjabini, Brevistriata

sladeni, Stegophiura

sobrina, Amphipholis

springsmithi, Molineus

springsmythi, Nothrus

spumosa, Heterakis

spurrelli, Barbus . 102-

squamosa, Selenaria

stabilis, Ophiothrix

stabularis, Eulaelaps

Steatonysella .

Steatonyssus

stigmatopygus, Barbus 102,

Stratiolaelaps

striculus, Steganacarus .

261,

. 166

104

• 232

. 284

193-196, 219-22O

41
26

11-15

85-86, 93-94

. 63

. 65

285-286

284-286
264, 282
. 69
. 69
264

154, 164, 166
164

164-165, 182-184

. 69

231, 236-239

. 67

231, 234-236

26-3i

232

-104, 108-no, 147

. 165

. 62

268, 285

264

261, 264, 275, 291

105, 143-145, 148

269, 272, 275

INDEX

subinensis, Barbus
sublineatus, Barbus
svenssoni, Barbus .
sylviarum, Ornithonyssus

tectorum, Trhypochthonius
Thiariidae

tibialis, Steatonyssus
trachydisca, Arnphiura .
Trhypochthoniidae .
trispilomimus, Barbus
trispilus, Barbus
tropicus, Bulinus .
tunkanati, Arthrostoma

102, 105, 124—126
105, 128-130, 148
104
261, 285-286

. 33

91-Q2
.286
. 44
. 33

104

102, 104-105, 126-128
86-87, 93-94
231, 241-243

umbellata depressa, Cupuladria 156—157,

180-i82, 193

umbellata, Discoporella 154-156, 158, 160,

162-164, I07, i75-177-i8o, 193-196, 221-223
unguiculatum, Metrarabdotos . . .195

uniumbonata, Aspidophiura . . .65

urostigma, Barbus . . . .104

vivipara, Stegophiura
voltae, Barbus

werneri, Barbus

102-104, 132-135,

103

PRINTED IN GREAT BRITAIN
BY ADLARD & SON LIMITED
BARTHOLOMEW PRESS, DORKING

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