fc I- O

BULLETIN OF

•& $\,

THE BRITISH MUSEUM
(NATURAL HISTORY)

ZOOLOGY

Vol. 31
1977

BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1977

DATES OF PUBLICATION OF THE PARTS

No. i . . . . -24 February 1977

No. 2 ..... 24 March 1977

No. 3 . . . . . .28 April 1977

No. 4 . . . . . .28 April 1977

No. 5 30 June 1977

No. 6 . . . . .30 June 1977

ISSN 0007-1498

Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU

CONTENTS

ZOOLOGY VOLUME 31

PAGE

No. I. Freeliving marine nematodes from southern Africa. By J. W.

COLES (PI. i) i

No. 2. The Accacoeliidae (Digenea) of fishes from the north-east Atlantic.

By R. A. BRAY and D. I. GIBSON 51

No. 3. The planktonic copepods of the northeastern Atlantic Ocean : some
taxonomic observations on the Oncaeidae (Cyclopoida) . By G. A.
BOXSHALL .......... 101

No. 4. Notes of deep-water Atlantic Crinoidea. By A. M. CLARK . . 157

No. 5. A revision of the lizard genus Scincus (Reptilia : Scincidae). By

E. N. ARNOLD and A. E. LEVITON (Pis 1-3) .... 187

No. 6. Miscellanea

The morphology of a common soil flagellate, Heteromita globosa
Stein (Mastigophorea : Protozoa). By C. M. MACDONALD, J. F.
DARBYSHIRE and C. G. OGDEN (Pis 1-6) ..... 255

Notes on the morphology and nomenclature of three members of the
Euplotidae (Protozoa : Ciliatea). By C. R. CURDS . . . 267

Notes on the Alpheids in White's List of the specimens of Crustacea
in the collections of the British Museum (1847). By A. H. BANNER
and D. M. BANNER ......... 279

Establishment of a new genus in the family Phoxocephalidae
(Crustacea : Amphipoda) and a description of a new species from
North Island, New Zealand. By A. A. FINCHAM . . . 285

A new species of the synaptid genus Rynkatorpa Rowe & Pawson,
1967 (Holothurioidea : Apodida) from the Great Barrier Reef. By

F. W. E. ROWE 293

A review of the family Centropomidae (Pisces, Perciformes) : an
appendix. By P. H. GREENWOOD ...... 297

A revision of the synonymy of three species of Leiolopismid skinks

from New Zealand. By J. ROBB 303

FREELIVING MARINE NEMATODES
FROM SOUTHERN AFRICA

J. W. COLES

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 31 No. i

LONDON : 1977

FREELIVING MARINE NEMATODES FROMW1^*^
SOUTHERN AFRICA

BY

JOHN W. COLES

Pp. 1-49 ; i Plate ; 14 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 31 No. i

LONDON: 1977

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Scientific
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. 31, No. i, of the Zoology series.
The abbreviated titles of periodicals cited follow those
of the World List of Scientific Periodicals.

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

ISSN 0007-1498

Trustees of the British Museum (Natural History), 1977

BRITISH MUSEUM (NATURAL HISTORY)

Issued 24 February, 1977 Price £4.10

FREELIVING MARINE NEMATODES FROM
SOUTHERN AFRICA

By JOHN W. COLES

CONTENTS

Page

SYNOPSIS ........... 4

INTRODUCTION .......... 4

SAMPLE DATA ........... 5

SPECIES STUDIED .......... 8

DISTRIBUTION OF SPECIES ........ 8

DESCRIPTIVE SECTION ......... 9

Family Leptosomatidae

Platycomopsis Ditlevsen, 1926 - Systematic notes .... 9

Platycomopsis dayi sp. nov. ........ 10

Platycomopsis barba (Inglis, 1964) comb. nov. .... 13

Macronchus shealsi Inglis, 1964 ....... 13

Deontostoma jae (Inglis, 1964) Hope, 1974 . . . . . 15

Deontostoma hopei sp. nov. . . . . . . . . 16

Thoracostoma campbelli Ditlevsen, 1921 . . . . . 18

Thoracostoma zeae Inglis, 1964 ....... 20

Thoracostoma ancorarium Vitiello, 1975 . . . . • 21

Thoracostoma sp. ......... 22

Family Phanodermatidae

Dayellus dayi Inglis, 1964 ........ 23

Crenopharynx eina Inglis, 1964 ....... 23

Crenopharynx afra Inglis, 1964 ....... 24

Family Enoplidae

Enoploides mandibularis sp. nov. ...... 25

Mesacanthoides magna sp. nov. ....... 28

Rhabdodemania nancyae Inglis, 1964 ...... 30

Family Oncholaimidae

Oncholaimus problematicus sp. nov. ..... 3°

Oncholaimus sp. ......... 33

Filoncholaimus capensis sp. nov. ....... 34

Pontonema yaenae Inglis, 1964 ....... 36

Pontonema spp. .......... 36

Family Enchelidiidae

Symplocostomella trichostoma sp. nov. ...... 37

Eurystomina sp. ......... 41

Polygastrophora sp. ......... 42

Family Cyatholaimidae

Gen. et sp. innom. ......... 43

Family Desmodoridae

Spirinia sp. .......... 43

Family Chromadoridae

Parapinnanema sp. ......... 43

4 J. W. COLES

Page
HISTORICAL ACCOUNT OF FREELIVING MARINE NEMATODES FROM

SOUTHERN AFRICA . '. . . . . ' . . . . 43

LIST OF SPECIES REPORTED FROM SOUTHERN AFRICAN WATERS . . 44

ACKNOWLEDGEMENTS ......... 48

REFERENCES ........... 48

SYNOPSIS

Thirty-two species of freeliving nematodes, belonging to nineteen genera and eight families,
have been found in eighty-two samples collected from off the coasts of southern Africa by the
University of Cape Town Ecological Surveys. Seven species (all belonging to the order Eno-
plida) are described as new : Platycomopsis dayi, Deontostoma hopei, Enoploides mandibularis,
Mesacanthoides magna, Oncholaimus problematicus, Filoncholaimus capensis and Symplocosto-
mella trichostoma. The genus Parabarbonema Inglis, 1964, is considered synonymous with
Platycomopsis Ditlevsen, 1926, and P. barba Inglis is therefore transferred to the former genus.
Thoracostoma campbelli Ditlevsen, 1921, originally described from Campbell Island, South
Pacific, is redescribed from Luderitz Bay, South West Africa. Species belonging to the genera
Thoracostoma (one), Oncholaimus (one), Pontonema (two), Eury stamina (one) and Polygastrophora
(one), not named specifically, are briefly described. Additional notes and measurements are
given to ten species previously described from off Cape Province : Platycomopsis barba (Inglis,
1964) comb, nov., Macronchus shealsi Inglis, 1964 ; Deontostoma jae (Inglis, 1964) Hope, 1974 '•
Thoracostoma zeae Inglis, 1964 ; Thoracostoma ancorarium Vitiello, 1975 ; Dayellus dayi Inglis,
1964 ; Crenopharynx eina Inglis, 1964 ; C. afra Inglis, 1964 ; Rhabdodemania nancyae Inglis,
1964 and Pontonema yaenae Inglis, 1964.

An historical account is given of freeliving marine nematodes reported from southern Africa,
followed by a list of species.

INTRODUCTION

THIS report is a continuation of work on freeliving marine nematodes collected from
South African waters by the University of Cape Town Ecological Surveys, under the
direction of Professor J. H. Day. Earlier samples from these surveys were studied
and reported on by Inglis (1963, 1964) and Vitiello (1975). I was entrusted with the
examination of the remaining samples and the results of this are given below. As
found by Inglis the species present are mainly members of the order Enoplida,
amongst which are found most of the larger freeliving nematodes. This may reflect
the inadequacies of the collecting and sorting techniques used rather than the fauna
present. The families of Enoplida represented in the following account are the
Leptosomatidae, the Phanodermatidae, the Enoplidae, the Oncholaimidae and the
Enchelidiidae. The non-enoplid species found appear to belong to the families
Cyatholaimidae, Desmodoridae and Chromadoridae. Of the latter families there
are not many specimens available and, as their condition is very poor, it has not been
possible to determine them specifically with any degree of certainty.

With the exception of the specimens collected from Luderitz Bay on the coast of
South West Africa at 26°S, I5°E (LU 19), the samples were collected from the west,
south-west and east coasts of the Republic of South Africa, between 24°S, 34°E and
36°S, 23°E. All these localities (with the exception of the Portuguese East Dredge
sample) are relatively close together and according to the work of ecologists and

NEMATODES FROM SOUTHERN AFRICA 5

others the water temperature of this region is quite cool and not affected by the
warm waters of the Mozambique/Algulhas Currents (Ekmann, 1953).

Thoracostoma campbelli Ditlevsen, 1921, reported here from Luderitz Bay, is the
only species in the present samples which has been found in localities outside South
African waters. It appears to be widely distributed in the southern hemisphere
(originally recorded from off Campbell Island) and is a subantarctic form which also
occurs further north in regions where cold-water currents maintain a fairly low
temperature. Wieser (1953) found it on the coast of Chile, which is even further
away from the type-locality than the present record. It therefore seems probable
that the other species described from South African waters also occur in the more
southerly regions.

Before the collection of freeliving nematodes was undertaken by the University
of Cape Town Ecological Surveys, very little appears to have been done on this
fauna from the South African coasts and a historical account will be given later in
this report.

Representatives of the species mentioned herein will be deposited in the collections
of the British Museum (Natural History).

LU 19 26°S, I5°E. io.vii.46.

SAMPLE DATA
Luderitz Bay

SB 214
SB 290
SB 299

SB 300
SB 304

SB 332
SB 334

SB 336
SB 339

SB 340
SB 348
SB 350

FAL 412
FAL 422
FAL 430

Saldanha Bay Dredgings

33°6'S, I7°56.6'E. In green sand and shell-gravel at a depth of 26 metres, i.vi.6o.
33°oi'S, i7°57.9'E. In fine khaki-coloured sand at a depth of 9 metres, 29.^.63.
33°O2.5'S, I7°57'E. In khaki-coloured sand and shell-gravel at a depth of 26 metres,

3o.iv.63 (grab sample).

33°oi.o'S, i7°58.9'E. In shell-gravel at a depth of n metres, 3o.iv.63.
33°O2.5'S, I7°58.9'E. In coarse shell-gravel at a depth of 20 metres, 3o.iv.63 (grab

sample) .

33°oi.2'S, I7°58.7'E. On a rock substratum at a depth of 13 metres, 25.^.64.
33°oi.2'S, i7°59.7'E. On a rock substratum at a depth of 13 metres, 25.^.64 (grab

sample) .

33°oo.3'S, I7°57'E. In fine sand and shell-gravel at a depth of 6 metres, 26.iv.64.
33°03.4'S, i8°o2.o'E. In fine green sand and shell-gravel at a depth of 7-6 metres,

2i.iv.64.
33°03.4'S, i8°o2.o'E.

(grab sample).
33°O2.9'S, i8°oi.6'E.

2g.iv.64.
33°O2.9'S, i8°oi.6'E.

In fine sand and shell-gravel at a depth of 7 metres, 27.iv.64
In shell-gravel and coarse sand at a depth of 13 metres,
In sand and shell-gravel at a depth of 13 metres, 2g.iv.64.

False Bay dredgings
In fine khaki-coloured sand at a depth of 31 metres, i6.v.6i

34°8.8'S, i8°33.5'E.

(grab sample).
34°i2.5'S, i8°37'E. In khaki-coloured sand and shell-gravel at a depth of 48 metres,

i5-v.6i (grab sample).
34°i5.2'S, i8°33.2'E. In shell-gravel at a depth of 41 metres, i6.v.6i.

J. W. COLES

FAL 436
FAL 449
FAL 613
FAL 617
FAL 622
FAL 630
FAL 633

FAL 639
FAL 643

FAL 648
FAL 65 1
FAL 654

FAL 657
FAL 660

FAL 661
FAL 666
FAL 674

FAL 676
FAL 679

FAL 684
FAL 688

FAL 689
FAL 696
FAL 700

FAL 704
FAL 706

FAL 709
FAL 713

FAL 721
FAL 725
FAL 726
FAL 729

FAL 731
FAL 738

FAL 748

34°07'S, i8°35'E.
34°o8'S, i8°28'E.
34°o8'S, i8°49'E.
34°i5'S, i8°55'E.

34°i5.2'S, i8°33.2'E. In shell-gravel at a depth of 41 metres, i6.v.6i (grab sample).
34°i3.3'S, i8°3i.3'E. In sand at a depth of 39 metres, 22.v.6i (grab sample).
34°i5'S, i8°35'E. In coarse sand and shell-gravel at a depth of 50 metres, 24.11.64.
In shell-gravel at a depth of 22 metres, 20.11.64.
In white sand at a depth of 26 metres, 21.11.64.
On a rock substratum at a depth of 7-9 metres, 21.11.64.
In coarse khaki-coloured sand and shell-gravel at a depth of

48 metres, 2.11.64 (grab sample).

34°io'S, i8°29'E. In fine green sand at a depth of 35 metres, 20.11.64 (graD sample).
34°o6'S, i8°33'E. In coarse white sand and shell-gravel at a depth of 15 metres,

20.11.64 (grab sample).

34°i3'S, i8°39'E. In green mud at a depth of 53 metres, 20.11.64 (grab sample).
34°o6'S, i8°33'E. In sand and shell-gravel at a depth of 15 metres, 20.11.64.
34°2i'S, i8°32'E. In khaki-coloured sand and shell-gravel at a depth of 75 metres,

20.11.64.
34°2i'S, i8°32'E. In khaki-coloured sand and shell-gravel at a depth of 75 metres,

24.11.64 (grab sample).
34°2i'S, i8°37'E. In fine khaki-coloured sand and shell-gravel at a depth of

97 metres, 24.11.64.

34°07'S, i8°36'E. In sand and shell-gravel at a depth of 26 metres, 20.11.64.
34°07'S, i8°36'E. In sand and shell-gravel at a depth of 26 metres, 20.11.64.
34°i7'S, i8°36'E. In khaki-coloured sand and shell-gravel at a depth of 66 metres,

24.11.64 (grab sample).

34°n'S, i8°35'E. In fine khaki-coloured sand at a depth of 42 metres, 25.11.64.
34°i5'S, i8°4i'E. In fine khaki-coloured sand at a depth of 40 metres, 25.11.64

(grab sample).
34°io'S, i8°48'E.

sample) .
34°i9'S, i8°3i'E.

(grab sample).
34°u'S, i8°4i'E.
34°io'S, i8°43'E.
34°n'S, i8°39'E.

(grab sample).
34°io'S, i8°44'E.

(grab sample).
34°2i'S, i8°37'E.

sample) .

34°05'S, i8°38'E.
34°05'S, i8°38'E.

In white shelly sand at a depth of 29 metres, 21.11.64 (grab
In fine khaki-coloured sand at a depth of 61 metres, 24.11.64

In fine khaki-coloured sand at a depth of 53 metres, 25.11.64.

On a rock substratum at a depth of 36 metres, 29.11.64.

In coarse sand and shell-gravel at a depth of 44 metres, 25.11.64.

In coarse sand and shell-gravel at a depth of 39 metres, 25.11.64
In khaki-coloured sand at a depth of 87 metres, 24.11.64 (grab

In coarse sand and clay at a depth of 17 metres, 26.11.64.
In coarse green sand and shell-gravel at a depth of 17 metres,
26.11.64 (grab sample).
34°O7'S, i8°45'E. In shell-gravel on a rock substratum at a depth of 20-26 metres,

22.11.64.

34°i2'S, i8°45'E.
(grab sample).

In coarse sand and shell-gravel at a depth of 26 metres, 26.11.64

In fine khaki-coloured sand at a depth of 48 metres, 25.11.64

34°07'S, i8°o7'E.

(grab sample).
34°i5'S, i8°43'E.

(grab sample).
34°o8'S, i8°45'E.
340o6'S, i8°40'E.

26.11.64.
34°i4.3'S, i8°29.7'E. In

15.11.65 (grab sample).

In fine khaki-coloured sand at a depth of 54 metres, 25.11.64

In coarse sand at a depth of 29-26 metres, 26.11.64.
In coarse sand and shell-gravel at a depth of 29-26 metres,

coarse khaki-coloured sand at a depth of 40 metres,

NEMATODES FROM SOUTHERN AFRICA 7

FAL 758 34°i4.5'S, i8°38.8'E. In khaki-coloured sand at a depth of 53 metres, 17.11.65

(grab sample).
FAL 772 34°i7.o'S, i8°29.2'E. In coarse shelly sand at a depth of 27 metres, 15.11.65 (grab

sample).
FAL 798 34°2i.6'S, i8°38.9'E. In green mud and shelly sand at a depth of 87 metres,

22.11.65 (grab sample).
FAL 806 34°io.7'S, i8°26.3'E. In black mud and hydrogen sulphide at a depth of 18 metres,

17.11.65 (grab sample).
FAL 832 34°i4.3'S, i8°29.7'E. In coarse khaki-coloured sand at a depth of 40 metres,

I5.ii.65-

West Coast dredgings

WCD 41 33°6.5'S, I7°56'E. On a rock substratum at a depth of 18-33 metres, 2.v.6o.
WCD 71 33°6.5'S, i7°49'E. On a rock substratum at a depth of 91 metres, 22.ix.6o.
WCD 74 34°25'S, i7°36'E. In khaki-coloured sand at a depth of 1240 metres, 8.x. 60 (grab

sample) .
WCD 79 34°i7'S, i7°53'E. In green sand and mud at a depth of 320 metres, 8.x. 60 (grab

sample) .

WCD 86 33°6.4'S, I7°44.9'E. In green and black mud at a depth of 146 metres, 3.vii.6i.
WCD 87 33°6.2'S, I7°49'E. In dark green mud on a rocky substratum at a depth of 88 metres

3.vii.6i (grab sample).
WCD 90 32°5.o'S, i8°i6.7'E. In coarse white sand and shell-gravel at a depth of 39 metres,

2.vii.6i.
WCD 93 32°5.o'S, i8°i6.7'E. In coarse white sand and shell-gravel at a depth of 39 metres,

2.vii.6i (grab sample).
WCD 101 32°5.5'S, i8°i7.3'E. In coarse white sand and shell-gravel at a depth of 27 metres,

2.vii.6i.
WCD 108 32°o8'S, I7°39'E. In fine dark green mud at a depth of 172 metres, 2.vii.6i (grab

sample).
WCD 119 33°6.4'S, i7°44.9'E. In green and black mud at a depth of 146 metres, 3.vii.6i

(grab sample).
WCD 196 33°03.7'S, I7°45.2'E. In coarse sand and shell-gravel at a depth of 33 metres,

25.^.64.
WCD 206 33°o8.6'S, i7°57.3'E. On a shell substratum at a depth of 50 metres, 29.iv.64

(grab sample).
WCD 210 33°o8.4'S, i7°55.i'E. In shelly sand at a depth of 79 metres, 29.^.64.

South Coast dredgings

SCD 115 34°54.4'S, 22°i2.2'E. In coarse sand and shell-gravel at a depth of 107 metres,

26.xi.59.

SCD 134 34°29'S, 2i°49.5'E. In grey-green mud at a depth of 73 metres, 26.xi.59.
SCD 136 34°35'S, 2i°56'E. In coarse and fine shell-gravel at a depth of 78 metres, 28.viii.6o.

(grab sample).

SCD 197 34°07.5'S, 23°3i.7'E. In fine sand at a depth of 79 metres, 29.xi.6o.
SCD 233 36°28.5'S, 2i°n'E. In khaki-coloured sand at a depth of 183 metres, 4.xii.6o

(grab sample).

SCD 251 34°48'S, 23°39'E. On a rock substratum at a depth of 148 metres, 3O.xi.6o.
SCD 275 34°5i'S, 23°4i'E. In khaki-coloured sand at a depth of 183 metres, 2O.xi.6o.
SCD 298 33°09'S, 28°02'E. At a depth of 84 metres, 6.ii.62 (grab sample). Substratum type

unknown.

SCD 303 33°39'S, 27°i5'E. In hard sand at a depth of 88 metres, 6.11.62 (grab sample).
SCD 309 33°59'S, 25°5i'E. In coarse shell-gravel and mud at a depth of 50 metres, 9.11.62

(grab sample).

8 J. W. COLES

Portuguese East Dredge

PED 17 24°53'S, 34°56'E. In fine grey sand and rock at a depth of 55 metres, I9.viii.64.
PED 19 25°07'S, 34°34'E. In dark sandy mud at a depth of 112 metres, I9.viii.64.

SPECIES STUDIED

Thirty-two species, belonging to nineteen genera and eight families, have been
recognized or considered distinct. Of these, seven are described as new, one species
(Thoracostoma campbelli) is redescribed in detail, brief descriptions are given of six
species which could not be named specifically, and additional notes and measure-
ments are given of species originally described from South African waters by Inglis
and Vitiello.

DISTRIBUTION OF SPECIES

SPECIES

LEPTOSOMATIDAE
Platycomopsis dayi sp. nov.

Platycomopsis barba
Macronchus shealsi

Deontostoma jae

Deontostoma hopei sp. nov.
Thoracostoma campbelli
Thoracostoma zeae

Thoracostoma ancorarium
Thoracostoma sp. innom.
Thoracostoma sp. innom.

PHANODERMATIDAE
Dayellus dayi

Crenopharynx eina
Crenopharynx afra

ENOPLIDAE

Enoploides mandibularis
sp. nov.

LOCALITY

Saldanha Bay
False Bay

West Coast dredging
False Bay

West Coast dredging
False Bay

West Coast dredging

South Coast dredging

Saldanha Bay

False Bay

West Coast dredging

Saldanha Bay

Luderitz Bay, S.W. Africa

False Bay

South Coast dredging

False Bay

False Bay

Portuguese East dredge

False Bay

West Coast dredging

False Bay

South Coast dredging

False Bay

West Coast dredging
South Coast dredging

Saldanha Bay

False Bay

South Coast dredging

SAMPLE NOS

SB 348.

FAL6i3, 663, 666, 684, 704,

726, 748, 772, 832.

WCD 210.

FAL 617.

WCD 90, 93, 101, 196.

FAL 436, 622, 633, 704, 726,

772, 802, 806, 832.

WCD 196, 206, 210.

SCD 303.

SB 290, 332, 334, 350.

FAL 358, 651.

WCD 41, 74, 101, 196.

SB 334-

LUig.

FAL 617, 630, 661, 679.

SCD 298.

FAL 758.

FAL 832.

PED 19.

FAL 725, 798.

WCD 86.

FAL 613, 633, 689, 700.

SCD 309.

FAL 633, 654, 674, 679, 688,

713, 832.

WCD 86, 119.

SCD 134.

SB 214, 299, 300.
FAL 660, 706.
SCD 233.

NEMATODES FROM SOUTHERN AFRICA
DISTRIBUTION OF SPECIES (cont.)

SPECIES LOCALITY

Mesacanthion sp. innom. False Bay

Mesacanthoides magna sp. nov. West Coast dredging
Rhabdodemania nancyae False Bay

Rhabdodemania sp. innom. False Bay
Oxyonchus sp. innom. False Bay

ONCHOLAIMIDAE

Oncholaimus problematicus
sp. nov.

Oncholaimus sp. innom.
Filoncholaimus capensis

sp. nov.
Pontonema yaenae

Pontonema sp. innom.
Pontonema sp. innom.

ENCHELIDIIDAE

Symplocostomella trichostoma

sp. nov.

Eurystomina sp. innom.
Polygastrophora sp. innom.
Gen. et sp. innom.

CYATHOLAIMIDAE
Gen. et sp. innom.

DESMODIRIDAE

Spirinia sp. innom.

CHROMADORIDAE

Parapinnanema sp. innom.

False Bay

West Coast dredging
South Coast dredging
Saldanha Bay
West Coast dredging

False Bay

West Coast dredging

False Bay

West Coast dredging

West Coast dredging

False Bay

South Coast dredging

Portuguese East dredge

South Coast dredging

False Bay

SAMPLE NOS

FAL 449.
WCD 206, 210.
FAL 748.
FAL 666.
FAL 430.

FAL 613, 633, 648, 654, 674,

679, 689, 748, 772, 832.

WCD 86, 206.

SCD 134, 197, 233, 275.

SB 336.

WCD 74, 86, 119.

FAL 748, 832.
WCD 210.
FAL 613.
WCD 210

WCD 86.

FAL 412, 422.
SCD 251.
FED 17.

SCD 136.

FAL 684.

False Bay

West Coast dredging

DESCRIPTIVE SECTION

FAL 725, 748.
WCD 196, 206.

Note on measurements. Only absolute measurements of specimens are given,
with a range when more than two specimens of a species are present. The head-
diameter is given at the level of the cephalic setae. Measurements of the actual
lengths of the spicules (in the males) are given and not the chord.

Family LEPTOSOMATIDAE DeConinck & Stekhoven, 1935
Genus PLATYCOMOPSIS Ditlevsen, 1926

Dactylonema Filipjev, 1927 ; Parabarbonema Inglis, 1964.

Ditlevsen (1926) erected the genus Platycomopsis for a species collected during the
Danish Ingolf Expedition in the Arctic Ocean. He compared it with the genus
Platycoma Cobb, 1894, which has two flat seta-like structures around each amphid.

io J. W. COLES

Ditlevsen says : 'Cobb named the genus Platycoma because two flat hairs of unequal
length grow from the inner margin of the anterior border of each lateral organ'.
In the species described by Ditlevsen (Platycomopsis cobbii) there are four rounded
setae situated close together just behind each amphid, and these are about as long
as the cephalic setae. He also notes that in Platycoma a narrow buccal cavity is
present, whereas the specimens he examined lack a buccal cavity, as in Leptosomatum.
In other respects, it is similar to Platycoma. In the present species from off the coast
of South Africa there is no apparent buccal cavity, although there is some degree of
cuticularization of the walls of the oral cavity.

Inglis (1964) erected the genus Parabarbonema for a species (P. barba) from other
samples collected by the Ecological Surveys. This species has also been found in the
present samples. Inglis compared his genus with Barbonema Filipjev, 1927, but
made no reference to its similarity with Platycomopsis. Barbonema is characterized
by the very long cephalic setae, by an elongate tail, by the absence of a pre-cloacal
supplement in the male and by the position of its amphids. The details of the
structure of the spicules in Barbonema are uncertain, because Filipjev considered
that the spicules in the species he described were incompletely developed. Para-
barbonema is similar to Platycomopsis in nearly all respects, more particularly in the
form of the cephalic sense organs, in the position of the amphids and in the arrange-
ment of the group of setae behind each amphid. Moreover the spicules are of
similar form. Another important resemblance is the presence of a papillate pre-
cloacal supplementary organ.

Parabarbonema differs from Platycomopsis in minor details, which could really only
be considered specific differences. The differences that might be considered dis-
tinctive at a generic level are the presence of a cephalic capsule, and the more
strongly cuticularized walls of the oral cavity, forming a prominent dorsal tooth.
These differences are, however, only a matter of degree. The outline of the cephalic
capsule varies somewhat in its clarity, but this may depend on the age of the speci-
mens, as well as on their condition. Consequently, it seems that there is no justi-
fication for the retention of the name Parabarbonema.

Mawson (1956) reviewed the species of Platycomopsis, described two new species
from the Antarctic and included a key to all the species known at that time.

The genus Platycomopsis Ditlevsen, 1926, may be re-defined as follows : Lepto-
somatidae with unlobed cephalic capsule variably developed. Labial sense organs
papillate. Buccal cavity absent or very narrow, with its walls sometimes cuticu-
larized and modified as onchia. Amphids opening on the posterior edge of the
cephalic capsule, when this is seen. Sometimes two to four fairly long setae just
behind the amphids. Rows of cephalic setae on the dorso-lateral and ventro-
lateral surfaces, as well as on the lateral sides. A papillate supplementary organ
anterior to the cloaca in the male. Spicules fairly short and stout.

Platycomopsis dayi sp. nov.

(Fig. la-c)
TYPE-LOCALITY. Off West coast of Cape Province, South Africa.

NEMATODES FROM SOUTHERN AFRICA n

MATERIAL STUDIED. Of the nine specimens available (from different samples)
only three males are present. Of these, only one (WCD 206) is reasonably well
preserved. Most of the females are also in rather poor condition.

Saldanha Bay : i? SB 348.

False Bay : i^ FAL 613 ; 2?? FAL 726 ; 2?? FAL 684 ; i<$ FAL 704 ;

i? FAL 633 ; i larva FAL 666 ; i? FAL 748 ; i larva FAL 772 ;

i? FAL 832.
West Coast Dredgings : i<£ i? WCD 206 ; 3^, 4?$ WCD 210.

SYNTYPES. BM(NH) reg. nos 1975.701-709.
MEASUREMENTS (in mm).

c?c? 9?

Body-length 15-60-22-80 21-00-22-20

Body-breadth 0-18 - 0-21 0-25 - 0-26

Head-diameter 0-05 - 0-06 0-06 - 0-07

Length of oesophagus 1-67- 2-42 2-18- 2-20

Length of tail 0-21 - 0-24 0-21 - 0-26

Distance of nerve-ring from anterior end of body 0-53 - 0-73 0-67 - 0-72

Anal diameter 0-12 - 0-13 0-12 - 0-14

Length of spicules 0-12 - 0-14

Length of gubernaculum 0-12

Distance of vulva from anterior end of body 13-10-13-70

DESCRIPTION. The body is very long and slender and the cuticle is quite smooth
throughout its length. The head is truncate and without any marked constriction.
A distinct buccal-cavity is not apparent - only a very slight thickening, a little
wider than the oesophageal lumen, where the walls are slightly cuticularized. In
apical view three rod-like thickenings have been observed, one on each lobe of the
tri-radiate mouth opening. The cephalic capsule is not very well defined, and only
a very faint outline of its posterior extremity was seen in some specimens.

There is a crown of six papillae situated far anteriorly, surrounding the mouth.
Posteriorly to this crown there is a ring of ten setae, two of which are situated close
together on each of the dorso-lateral and ventro-lateral surfaces of the head and a
single seta on each lateral surface (Fig. la).

The amphids are, as is usual in this genus, c. 30/u,m from the anterior end of the
body. They are somewhat boat-shaped and measure c. 22/u.m in diameter. Just
behind each amphid there are three setae situated very close together. They are a
little shorter than the cephalic setae. In the hinder part of the cervical region there
are longitudinal rows of setae, two rows on the sub-dorsal and two rows on the
sub-ventral surfaces. The setae are c. 20 p,m long, but gradually become shorter
and less numerous towards the end of the oesophageal region. Setae also occur on
the lateral surfaces, but they are not arranged in such regular rows and are more
sporadic in their distribution. They also tend to be shorter. The width of the
oesophagus is fairly even throughout its length.

The nerve-ring occurs at about the anterior third of the body. An excretory pore
or ventral gland has not been made out.

12

J. W. COLES

FIG. i. Platycomopsis dayi sp. nov. (a) Anterior end of body (lateral view) ; (b) Spicules
and gubernaculum ; (c) Posterior end of male.

Male. The spicules are fairly short and only slightly curved. The proximal end
of each spicule is broad and truncate, while the distal end terminates in a point
although a rounded expansion occurs just in front of the tips (Fig. ib). A sheath-
like gubernaculum surrounds the spicules and commences quite near to their proxi-
mal end. There is a papillate supplementary organ anterior to the cloaca at a
distance of approximately the length of the tail. A row of setae occurs on each
ventro-lateral surface. Each row commences just anteriorly to the supplementary
organ and consists of ten setae, six anteriorly to the cloaca and four on the tail. The
tail is conical, with a short cylindrical tip (approximately a third of the tail-length)
bearing two small setae. Setae also occur sporadically on other parts of the tail
region.

NEMATODES FROM SOUTHERN AFRICA 13

Female. No gravid specimens were found. The vulva is situated a little behind
the middle of the body. The tail is similar in shape to that of the male, namely,
conical with a short cylindrical tip.

REMARKS. This species is characterized by having three setae situated close
together behind each amphid, combined with the size and form of the spicules. It
can be readily distinguished from P. barba, also present in the samples collected by
U.C.T.E.S., by the shape of the tail. Further, in P. dayi the tail terminates in a
short cylindrical portion, whilst in P. barba the tail is blunter with quite prominent
setae. This character applies to both sexes.

Platycomopsis barba (Inglis, 1964) comb. nov.

Parabarbonema barba Inglis, 1964

TYPE-LOCALITY. Off West coast of Cape Province, South Africa.

MATERIAL STUDIED.

False Bay : i? FAL 617.

West Coast Dredgings : i<? WCD 90 ; i£, 3$$ WCD 93 ; i? WCD 101 ;

5$? WCD 196.

MEASUREMENTS (in mm).

<?<? ??

Body-length 9-10-9-80 10-60-15-80

Body-breadth 0-09 0-09 - 0-14

Head-diameter 0-03 0-03 - 0-04

Length of oesophagus 1-20 - 1-25 1-50 - 1-90

Length of tail 0-17-0-18 0-18 - 0-30

Distance of nerve-ring from anterior end of body 0-41 - 0-42 0-45 - 0-56

Anal diameter 0-07-0-08 0-08- 0-12
Length of spicules 0-08
Length of gubernaculum

Distance of vulva from anterior end of body 6-80 - 10-30

REMARKS. These specimens agree very well with the description given by Inglis
(1964), except that there is much greater variation in the size of the females. This
species is readily distinguished from Platycomopsis dayi by the shape of the tail,
which is lacking a short cylindrical portion as in the case in the other species of this
genus.

GEOGRAPHICAL DISTRIBUTION. Coast of South Africa (Inglis, 1964 ; present
report.)

Genus MACRONCHUS Inglis, 1964
Macronchus shealsi Inglis, 1964

(Fig- 2)
TYPE-LOCALITY. Off West coast of Cape Province, South Africa.

J. W. COLES

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NEMATODES FROM SOUTHERN AFRICA

MATERIAL STUDIED.

False Bay : 6<$3, 5??, 2 larvae FAL 622 ; i<?, 8?$, 7 larvae FAL 633 ; 2?? FAL

726 ; 3?? FAL 704 ; i? FAL 436 ; 2?$ FAL 806 ; i<? FAL 832.
West Coast Dredgings : 2$$, 6$? WCD 196 ; 63$, 15?$, 2 larvae WCD 206 ;

6$? WCD 210.

South Coast Dredging : i<j> SCD 303.

REMARKS. The specimens in the present samples agree in most respects with the
description given by Inglis (1964), although there is a much greater variation in the
dimensions. In the sample from False Bay (FAL 622), the six males and five
females are of approximately similar size to the types, with the exception of one
female which measures 24-3 mm in length. The other females vary from 11-5 to
13-0 mm. In the West Coast dredging sample (WCD 206) the females vary from
22-5 to 26-6 mm. The males in this sample are also somewhat larger than those
from False Bay. For this reason measurements are given separately for samples
FAL 622 and WCD 206. Specimens in the remaining samples fall within these
limits of variation.

In the smaller males of the present samples, the anterior supplement contains six
to seven rods on each side but in the larger males there are up to ten rods on each
side (see Fig. 2). In the male type-specimens of this species there are nine rods on
each side of the anterior supplement.

Fig. 2. Macronchus shealsi Inglis, 1964. Vari-
ation in anterior supplement of male.

GEOGRAPHICAL DISTRIBUTION. Coast of South Africa (Inglis, 1964 ; present
report).

Genus DEONTOSTOMA Filipjev, 1916
Deontostoma jae (Inglis, 1964) Hope, 1974

Thoracostoma jae Inglis, 1964

TYPE-LOCALITY. Off West coast of Cape Province, South Africa.
MATERIAL STUDIED.

Saldanha Bay : i? SB 290 ; i<$, 4$$, i larva SB 332 ; i<$, 2$?, 2 larvae SB 334 ;

i? SB 350.

False Bay : 23$, 2?? FAL 738 ; i? FAL 651.

West Coast Dredgings : 5<J(J, 2??, i larva WCD 196 ; i larva WCD 101 ; i?

WCD 74 ; i(J WCD 41.

16 J. W. COLES

MEASUREMENTS (in mm).

Larvae

Body-length 8-50-11-00 8-60-11-30 6-30-7-10

Body-breadth 0-15- 0-20 0-15- 0-20 0-15-0-17

Head-diameter 0-04 0-04 0-04

Length of oesophagus 1-40— 1-60 1-41— 1-81 1-31-1-64

Length of tail o-io - 0-14 o-io 0-08-0-10

Distance of nerve-ring from anterior end of body 0-47 - 0-50 0-58 - 0-61 0-44 - 0-49

Distance of eye-spots from anterior end of body o-ii - 0-13 o-ii - 0-18 0-11-0-12

Anal diameter o-n - 0-15 o-n- 0-12 o-ii

Length of spicules 0-16 - 0-20

Length of gubernaculum o-ii- 0-15

Distance of vulva from anterior end of body 6-90 - 7-20

REMARKS. The specimens in the present sample agree well with the description
given by Inglis (1964). Some of these are, however, a little longer. The pre-
cloacal supplement in the males is situated at about the same position (0-09 mm
anteriorly from the cloaca) as in the type-specimens. There are up to six pairs of
ventro-lateral papillae present anteriorly to the pre-cloacal supplement. Several
gravid females are present and the eggs vary in size from 0-17x0-06 to
0-38 x 0-17 mm.

Hope (1974) transferred Thoracostoma jae to the genus Deontostoma because of the
lack of a 'trophis' (an anteriorly directed tooth-like process, usually hollow, arising
from the internal surface of the ventral side of the cephalic capsule).

GEOGRAPHICAL DISTRIBUTION. Coast of South Africa (Inglis, 1964 ; present
report).

Deontostoma hopei sp. nov.
(Fig. 3a-c, PI. i)

TYPE-LOCALITY. Saldanha Bay, Cape Province, South Africa.

MATERIAL STUDIED.

Saldanha Bay : i<$, 3$$, 2 larvae SB 334.

MEASUREMENTS (in mm).

<J ?? Larvae

Body-length 22-50 19-60-26-10 13-50-15-70

Body-breadth 0-13 0-12- 0-14 o-io - 0-13

Head-diameter 0-04 0-03 — 0-04 0-03

Length of oesophagus 1-76 1-60- 1-76 1-50

Length of tail o-io 0-08 - 0-09 0-07

Distance of nerve-ring from anterior end of body 0-48 0-42-0-45 0-40

Anal diameter 0-09 0-09— o-io 0-08

Length of spicules 0-14

Length of gubernaculum 0-07

Distance of vulva from anterior end of body 13-10 - 17-20 —

NEMATODES FROM SOUTHERN AFRICA

DESCRIPTION. Readily separated from Deontostoma jae (present in the same
sample) by being very long and slender. The cuticle is smooth and setae on the
body are scarce. The form and arrangement of the cephalic sense organs are typical
of the family, although the more anterior crown of six papillae is very indistinct.
The ring of ten setae lying further posteriorly are only about 5 /nm long. Of these
setae there are two on each dorso-lateral and ventro-lateral surface and a single
seta on each lateral surface.

The cephalic capsule, although clearly denned, is more feebly developed than in
other species of the family so far known to occur in South African waters. There are
six posterior lobes to the capsule and these are uneven in outline. The fenestrae are
wide between the lobes on the lateral sides, where the amphids are situated (Fig. 3a) .
No loculi were seen on the posterior lobes. A cephalic ring surrounds the anterior
end of the capsule and is very noticeable. There are six uneven rows of short
setae situated posteriorly to the cephalic capsule, but they extend only for a short
distance along the body. The walls of the buccal-cavity are only slightly cuti-
cularized and no definite teeth have been made out.

An excretory pore has not been observed, but the nerve-ring is quite distinct and
encircles the oesophagus at its anterior third. The oesophagus is typical of the
family for it is without a bulbous posterior swelling.

FIG. 3. Deontostoma hopei sp. nov. (a) Lateral view of head ; (b) Posterior end of male

(c) Spicule and gubernaculum.

i8 J. W. COLES

Male. The extremities of the testes have been difficult to make out, but they
appear to be paired, as is usual in this family. The two spicules are equal and quite
distinctive in shape. They are stout and non-alate. They bend sharply towards
the ventral side just behind the middle of each spicule (Fig. 3c). A gubernaculum
encloses the distal part of the spicules with processes pointing anteriorly. A mid-
ventral papillate supplement occurs 0-07 mm in front of the cloaca, and anteriorly
to this lie two rows of ventro-lateral papillae (about nine to ten in each row), some
with setae projecting from them. They vary in size and the more anterior ones are
not very clearly denned. They extend to about 0-33 mm anteriorly from the
cloaca. There are also setae surrounding the cloaca. The tail is short and bluntly
rounded with a short seta.

Female. The ovaries are paired, opposed and reflexed. One very elongate egg
has been observed in the posterior region of the uterus of one specimen and measures
0-9 x 0-12 mm. The vulva is situated posteriorly to the mid-body region and the
tail is short and rounded.

REMARKS. D. hopei may be distinguished from others of the genus principally
by the size and form of the spicules.

This species is named after Dr W. Duane Hope, who has reviewed the genus
Deontostoma and established its validity (Hope, 1967, 1974).

GEOGRAPHICAL DISTRIBUTION. Off the coast of South Africa (present report).

Genus THORACOSTOMA Marion, 1870
Thoracostoma campbelli Ditlevsen, 1921

(Fig. 4a-d)

TYPE-LOCALITY. Campbell Island, South Pacific.
MATERIAL STUDIED. Luderitz Bay : 2&J, 3??, i larva, LU 19.

MEASUREMENTS (in mm).

2cJcJ ?? Larva

Body-length 16-40, 18-20 15-40-18-20 13-20

Body-breadth 0-27, 0-27 0-22 — 0-23 0-21

Head-diameter 0-06, 0-05 0-05

Length of oesophagus 2-51 2-61 2-55 - 2-62 2-20

Length of tail 0-12, 0-12 0-12 o-io

Distance of nerve-ring from anterior end of body 0-71, 0-71 0-65 - 0-67 0-60

Anal diameter 0-18, 0-17 0-15 - 0-16 0-15

Length of spicules 0-25, 0-27

Length of gubernaculum 0-14, 0-19

Distance of vulva from anterior end of body 10-00 - n-io

DESCRIPTION. This species is a comparatively large member of the genus Thora-
costoma, and specimens in the present sample measure up to 18-2 mm in length.
The cuticle is fairly smooth, although very fine longitudinal striations appear to be
present. Transverse striations have not been made out. As noted by Ditlevsen

NEMATODES FROM SOUTHERN AFRICA 19

(1921), there is a well-marked constriction of the head in the region of the cephalic
setae.

The arrangement of the cephalic sense organs is typical of the genus, and there is a
crown of six papillae situated far anteriorly and surrounding the mouth. Further
back there is a ring of ten short cephalic setae, two of which are close together on
each of the dorso-lateral and ventro-lateral surfaces, and a single seta on each lateral
surface.

The cephalic capsule extends posteriorly to c. 40-45 /am from the anterior end of
the body. It is divided posteriorly into six lobes, separated by furrows which widen
to form six fenestrae. The two lateral fenestrae in which the amphids are situated
are markedly rounded or oval (Fig. 4a). The four fenestrae on the latero-dorsal
and latero-ventral sides are slightly less pronounced and more elongate horizontally.
On each of the lobes lying between the furrows there are several openings or loculi
that vary in shape and number. Sometimes there are three or four loculi, some of
which may not be very wide, and sometimes only two transversely elongate openings.
As Ditlevsen noted in his specimens there does not appear to be any regular arrange-
ment of the shape and number of the loculi. The anterior region of the cephalic
capsule is divided into six lobes, but they can only be seen clearly in an apical view.
A fairly well-defined cephalic ring is seen to surround the mid-region of the capsule.

Posteriorly to the cephalic capsule and surrounding the head there are several
rows of small granules which form a band c. 10 /zm wide. There are several setae
in the cervical region, although they are not arranged in clearly-defined rows.

There are two prominent cordiform cuticular structures on each lateral side of the
tri-radiate mouth. The walls of the buccal cavity are lined with a thickened
cuticle.

An excretory pore has not been made out. The nerve-ring is quite pronounced
and surrounds the oesophagus fairly far forward. There is no bulbous swelling at the
posterior end of the oesophagus.

Male. The testes appear to be paired, but the extremity of one of them could
not be clearly seen in the two specimens available. The two spicules are com-
paratively short and stout and compare well with Ditlevsen's illustration. The
gubernaculum which surrounds the distal ends of the spicules also compares quite
well with Ditlevsen's description. A mid-ventral supplementary organ lies opposite
the mid-region of the spicules, a feature typical of the genus Thoracostoma. There
are two rows of about twenty to thirty setae situated anteriorly to the cloaca and
extending just posteriorly to the latter in the ventro-lateral lines. Anteriorly to
the supplement there are two rows of papillae, six pairs in one specimen and seven
in the other. There are several small setae on the rounded posterior end of the tail.

Female. The ovaries are paired, opposed and reflexed. Two eggs have been
observed in the uterus of one specimen. They measure 0-53 x 0-19 mm and 0-53
x 0-15 mm. The vulva is situated about two-thirds of the total length of the body
from the anterior end. The tail is fairly short and the tip bluntly rounded, similar
to that in the male.

REMARKS. This species, originally found off Campbell Island, 52°3o'S, i69°io'E
(Ditlevsen, 1921), is widely distributed in the southern hemisphere. The present

20

J. W. COLES

FIG. 4. Thomcostoma campbelli Ditlevsen, 1921. (a) Lateral view of head ; (b) Apical view
of head ; (c) Ventral view of head ; (d) Spicule and gubernaculum (a and b to same scale) .

specimens from Luderitz on the coast of South West Africa agree in nearly all
respects with Ditlevsen's description. They, however, lack pigment spots in the
cervical region. Ditlevsen's specimens possessed cyathiform pigment spots, a
feature quite common in the genus Thoracostoma. Ditlevsen also noted that as there
was no lens to be seen it may be supposed that it had been diluted by the preservative
fluid, or had disappeared in some other way, which often happens to preserved
specimens of freeliving nematodes. In the specimens from Luderitz it seems that
the pigment spots, as well as the lens, have completely dissolved out.

GEOGRAPHICAL DISTRIBUTION. Campbell Island, South Pacific (Ditlevsen, 1921),
Kerguelen Islands (Stekhoven & Mawson, 1955 ; Mawson, 1958 ; Platonova, 1958,
1968) ; Macquarie Island (Stekhoven & Mawson, 1955 ; Mawson, 1958), Heard
Island (Mawson, 1958), off the coast of Chile (Wieser, 1953), off the coast of South
West Africa (present report).

Thoracostoma zeae Inglis, 1964
TYPE-LOCALITY. Off West coast of Cape Province, South Africa.

NEMATODES FROM SOUTHERN AFRICA 21

MATERIAL STUDIED.
False Bay : i<£ 3??, i larva FAL 617 ; i<? FAL 630 ; 2$? FAL 661 ; i$ FAL

679.
South Coast Dredging : i<j> SCD 298.

MEASUREMENTS (in mm).

<?<? ?? Larva

(? 4th stage)

Body-length 17-80-18-90 14-20-21-20 I4'3°

Body-breadth 0-24 - 0-25 0-22 - 0-23 0-23

Head-diameter 0-06 - 0-07 0-06 - 0-07 0-05

Length of oesophagus 2-40 - 2-88 2-40 - 3-06 2-20

Length of tail 0-18 0-12 0-12

Distance of nerve-ring from anterior end of body 0-65 - 0-66 0-70 - 0-78

Distance of eye-spots from anterior end of body 0-15 - 0-18 0-15 - 0-18 0-16

Anal diameter 0-18 - 0-19 0-14- 0-16 0-12

Length of spicules 0-31 — 0-32

Distance of vulva from anterior end of body 7-00 - 12-7

REMARKS. The specimens in the present samples, although in rather poor con-
dition, agree reasonably well with the description given by Inglis (1964) which was
based on only one male and three larval specimens. The specimens studied here,
however, are greater in length. The pre-cloacal supplement in the males occurs at
about the same position as in the type specimen (c. 0-13 mm anteriorly to the cloaca)
and the more anterior pair of papillae occurs at c. 0-62 mm anteriorly to the cloaca.

Females are recorded for the first time. They are typical of the genus with a very
short rounded tail. A gravid specimen is present in sample FAL 617 and has two
eggs, one of which measures 0-55 x 0-16 mm and the other 0-54 x 0-18 mm.

GEOGRAPHICAL DISTRIBUTION. Off the coast of South Africa (Inglis, 1964 ;
present report).

Thoracostoma ancorarium Vitiello, 1975

TYPE-LOCALITY. Lamberts Bay, Cape Province, South Africa.

MATERIAL STUDIED.
False Bay: i<? FAL 758.

MEASUREMENTS (in mm).

Body-length 20-00

Body-breadth 0-32

Head-diameter 0-07

Length of oesophagus 2-87

Length of tail 0-13

Distance of nerve ring from anterior end of body 0-77

Distance of eye-spots from anterior end of body 0-19

Cloacal diameter 0-15

Length of spicules 0-31

Length of gubernaculum 0-17

Distance of supplement anterior to cloaca 0-16

22

J. W. COLES

REMARKS. The only available specimen, a male, agrees very closely with the
description given by Vitiello (1975), except that pigmented eye-spots are present
on the lateral surfaces at a distance of 190 /*m from the anterior end of the body.
As noted in the case of T. campbelli, the presence or absence of eye-spots appears to
be variable and therefore unreliable as a specific character.

There are six pairs of pre-cloacal papillae compared with five in Vitiello's speci-
mens. The most anterior pair are, however, small, but distinct.

GEOGRAPHICAL DISTRIBUTION. Off Cape Province, South Africa ; Lamberts Bay
(Vitiello, 1975) ; False Bay (present report).

Thoracostoma sp.

(Fig. 5)

MATERIAL STUDIED.

South Coast Dredging : i<j> (immature) SCD 115.

MEASUREMENTS (in mm).

Body-length
Body-breadth
Head-diameter
Length of oesophagus

Distance of nerve-ring from anterior end of body
Length of tail
Anal diameter
Length of cephalic setae

Distance of posterior edge of cephalic capsule from
anterior end

30-70
0-18
0-05
2-31
0-55
o-io
0-13
o-oi
0-04

REMARKS. Although this specimen is very long it does not appear to be mature.
The cephalic capsule, in having six posterior lobes, is typical of the genus. The two
lateral fenestrae are wide and contain very prominent amphids (see Fig. 5). The

FIG. 5. Thoracostoma sp. Lateral view of head.

NEMATODES FROM SOUTHERN AFRICA 23

cephalic sense organs are typical. Cervical setae are present, scattered rather
unevenly at the level of the anterior end of the oesophagus. Eye-spots have not
been observed. The shape of the tail is typical of the genus, very short and rounded.

Until more specimens become available a more precise identification cannot be
attempted.

GEOGRAPHICAL DISTRIBUTION. Off the coast of South Africa (present report).

Family PHANODERMATIDAE Stekhoven, 1935

Genus DAYELLUS Inglis, 1964

Dayellus dayi Inglis, 1964

TYPE-LOCALITY. False Bay, Cape Province, South Africa.

MATERIAL STUDIED.

False Bay : i? FAL 725 ; i<$ FAL 798.
West Coast Dredging : i<? WCD 86.

MEASUREMENTS (in mm).

<? ?

(WCD 86)

Body-length 9-70 9-20

Body-breadth 0-15 0-15

Head-diameter 0-015 0-015

Length of oesophagus 1-50 1-40

Length of tail 0-42 0-35

Distance of nerve-ring from anterior end of body 0-40 0-34

Anal diameter 0-09 0-08

Distance of vulva from anterior end of body 5-10

Length of cephalic setae 0-02 0-02

REMARKS. These specimens agree well with the description given by Inglis (1964).

GEOGRAPHICAL DISTRIBUTION. Off the coast of South Africa (Inglis, 1964 ;
present report).

Genus CRENOPHARYNX Filipjev, 1934

Crenopharynx eina Inglis, 1964

(Fig. 6)

TYPE-LOCALITY. Off South coast of Cape Province, South Africa.

MATERIAL STUDIED.

False Bay : i<J, i? FAL 613 ; i(J, i$, i larva FAL 633 ; i? FAL 689 ; I& i?

FAL 700.
South Coast Dredging : i$ SCD 309.

24 J. W. COLES

MEASUREMENTS (in mm). icj

Body-length 7 '5°

Body-breadth 0-14

Head-diameter 0-018

Length of oesophagus 1-81

Length of tail 0-37

Distance of nerve-ring from anterior end of body 0-54

Distance of excretory pore from anterior end of body o-n

Anal diameter 0-07

Length of spicules 0-38

Length of gubernaculum 0-07
Distance of vulva from anterior end of body

I?

i larva

8-00

6-20

0-16

0-14

0-019

0-017

1-47
0-28

1-14
0-28

0-58

O-II

0-08

o-io

0-07

5-80

REMARKS. These specimens very closely resemble the description by Inglis
(1964). The spicules in one male specimen are protruded and, as in C. afra, the
distal ends do not appear to be so needle-like and the tips are slightly hooked (see
Fig. 6). The sharp-pointed barb directed towards the distal end of the spicule is
clearly seen. The cephalic setae are a little shorter in the present specimens, being
c. 10 /u,m in length.

GEOGRAPHICAL DISTRIBUTION. Off the coast of South Africa (Inglis, 1964 ;
present report).

FIG. 6. Crenopharynx eina Inglis, 1964. Distal part of spicules and gubernaculum.
FIG. 7. Crenopharynx afra Inglis, 1964. Distal ends of spicules.

Crenopharynx afra Inglis, 1964

(Fig- 7)
TYPE-LOCALITY. Off South coast of Cape Province, South Africa.

NEMATODES FROM SOUTHERN AFRICA

MATERIAL STUDIED.

False Bay : i<$, 2?? 3 larvae FAL 633 ;

2??, i larva FAL 679 ; i<£

FAL 832.

West Coast Dredging : 4^, 3?? WCD 86
South Coast Dredging : i$ SCD 134.

MEASUREMENTS (in mm).

Body-length

Body-breadth

Head-diameter

Length of oesophagus

Length of tail

Distance of nerve-ring from anterior end of body

Distance of excretory pore from anterior end of

body

Anal diameter
Length of spicules
Length of gubernaculum
Distance of vulva from anterior end of body

4^?, 4$$ FAL 654 ; 2??, FAL 674 ;
, 2?$ FAL 689 ; i? FAL 713 ; i$

; 3^, 8??, 2 larvae WCD 119.

33 ?? Larvae

7

•50

-8-40

7

•60-

10-00

6

50

-6

80

0

•22

- 0-26

o

•22 -

0'

25

o

18

- o

•22

0

•O2

- 0-025

O-O2 -

0-03

O-O2

- o

O2 1

i

•68

- 1-90

I

•67-

I

85

I-

50

- I

•52

0

•40

-0-49

O

•38-

0

•47

O

40

- o

•42

0

'44

-o-53

o-49- 0-55

o

•48

-0

•50

0

•13

-0-15

o

•12 -

o

15

o

12

- o

•16

O-II -0-12
0-40 - 0-49
0-07 - 0-o8

O-IO- O-II

4-00- 5-64

O-IO

REMARKS. This species occurs in nine samples, and all the specimens compare
well with the description by Inglis (1964), who had only one male and one female
at his disposal. In one of the males the spicules protrude from the cloaca and they
do not appear to narrow like a needle at the distal ends, as they do when completely
enclosed in the body (see Fig. 7). The slight swelling of the spicules at the point
where they narrow, as noted by Inglis, was seen in some specimens, but with difficulty.
The cephalic setae vary from 0-09 to 0-012 mm.

Several gravid female specimens are present. Up to five eggs occurred in one
specimen and they varied in size from 0-20 x 0-08 to 0-28 x o-n mm.

Family ENOPLIDAE Baird, 1853

Genus ENOPLOIDES Saveljev, 1912

Enoploides mandibularis sp. nov.

(Fig. 8a-c)

TYPE-LOCALITY. Saldanha Bay, Cape Province, South Africa.

MATERIAL STUDIED.

Saldanha Bay : i$ SB 214 ; 5&?, 3$$ SB 299 ; 4?$ SB 300.
False Bay : i? FAL 706 ; i<$, i? FAL 660.
South Coast Dredging : i^ SCD 233.

SYNTYPES. BM(NH) reg. nos 1975.809-820.

26 J. W. COLES

MEASUREMENTS (in mm).

33 9?

Body-length 4-80-6-40 3-80-5-70

Body-breadth 0-20-0-24 0-19-0-23

Head-diameter 0-05 - 0-06 0-05 - 0-06

Length of oesophagus i-oo - i-io 0-72 - i-oi

Length of tail 0-32 - 0-40 0-21 - 0-36

Anal diameter 0-09 - o-io 0-06 - 0-08

Length of spicules 0-40 - 0-46

Distance of pre-cloacal supplement anteriorly from cloaca 0-32 - 0-36

Distance of vulva from anterior end of body 1-80 - 2-50

DESCRIPTION. The body is of even width throughout most of its length and is of
average size for this genus. The head has high lips, whilst the tail is long and
tapering. The cuticle is transversely striated and the striations are evident on the
head and lip regions - a characteristic feature of the genus.

Surrounding the tri-radiate mouth, just posteriorly to the base of the lips, there is
a crown of six setae, each c. 10 /mi long. Further posteriorly on the head there are
two setae situated close together on each dorso-lateral and each ventro-lateral
surface. One of each pair is c. 30 /tm long and the other c. 20 ^m long. A single
seta c. 30 /x,m long is also situated on each lateral surface at the same level. In the
males, sub-cephalic setae (c. 20 pm long) are also present in eight bundles of two to
four setae at the base of the cephalic capsule. As the number of setae is not regular,
it seems possible that some have become lost during fixation or by some other cause.
Sub-cephalic setae also occur in the females, but they are rather irregular in their
distribution. Additional setae are also sometimes present on the cuticle at the level
of the anterior end of the oesophagus in both sexes.

The cephalic capsule is well defined and ornamented with granular markings. Its
posterior extremity lies at c. 60-70 pm from the anterior end of the body. The
three mandibles, each of which is c. 30 /am in length, are of the 'solid' type and
expand laterally at the anterior end into two laterally-directed arms. There is also
a broad central expansion on the side facing the buccal cavity (see Fig. 8b). A
slender tooth (about half the length of the mandibles) arises from the inner surface
of the base of each mandible along the onchial cavity. The teeth appear to be of
equal length. The cephalic ring is well defined with processes extending anteriorly
on either side of each mandible (see Fig. 8a). Amphids were not observed.

The oesophagus is of the form usually found in the genus Enoploides. The nerve-
ring is very difficult to make out and has been seen only in a few specimens where
it encircles the oesophagus at its anterior third (at c. 0-45 mm from the anterior end
of the body). An excretory pore was not observed.

Male. The spicules are long, of simple form and about one and a third as long as
the length of the tail. Their distal ends are pointed with a slight swelling on the
ventral side shortly before the extremity. A very short gubernaculum is weakly
developed (Fig. 8c). A tubular supplement is situated mid-ventrally (c. o-i mm in
length) c. 0-35 mm anteriorly from the cloaca.

NEMATODES FROM SOUTHERN AFRICA

27

FIG. 8. Enoploides mandibularis sp. nov. (a) Anterior end oi body (dorso-lateral view) ;
(b) Apical view oi head ; (c) Posterior end ot male (a and b to same scale).

Female. The ovaries are paired, opposed and reflexed. Several gravid specimens
are present in the samples, each specimen with up to four eggs. They measure
c. 0-15 x 0-12 mm. The vulva is situated a little anteriorly to the middle of the body.

REMARKS. This species is characterized by the form of the mandibles, combined
with the relatively long spicules when compared with the length of the tail and the
weakly-developed gubernaculum.

GEOGRAPHICAL DISTRIBUTION. Off Cape Province, South Africa (present report).

28 J. W. COLES

Genus MESACANTHOIDES Wieser, 1953

Mesacanthoides magna sp. nov.

(Fig. ga-b)

TYPE-LOCALITY. Off West coast of Cape Province, South Africa.
MATERIAL STUDIED.
West Coast Dredgings : 3^, 9?$ WCD 206 ; i? WCD 210.

SYNTYPES. BM(NH) reg. nos 1975.821-830.
MEASUREMENTS (in mm).

Body-length

Body-breadth

Head-diameter

Length of oesophagus

Length of tail

Anal diameter

Length of spicules

Length of gubernaculum

Distance of vulva from anterior end of body

I2-2O - 15-50

O-2O - O-22

0-12 - 0-I3

2-I5 - 2-50

0-175- 0-185

0-085 — 0-090

0-65 - 0-81

0-08 - o-io

11-50- 17-20

0-17-
0-05 -

2-OO -
0-I9-
0-10-

0-25
0-08
2-50
0-32
0-14

6-80 - 9-90

DESCRIPTION. This species is very large compared with all those previously
described and assigned to this genus. The body is of even width throughout most of
its length. The head is truncate. The tail is conical, terminating in a short cylin-
drical portion in both sexes, not normally bent dorsally as shown in Fig. gb. The
cuticle is finely striated transversely.

The three lips are high and prominent. Surrounding the mouth there is a crown
of six, short, stout setae c. 10-15 P™ m length. The arrangement of the cephalic
setae surrounding the head more posteriorly is typical of the family, inasmuch as
two lie close together on each dorso-lateral and ventro-lateral surface and a single
seta on each lateral surface - ten setae in all. One of each pair of setae on the
dorso-lateral and ventro-lateral sides is shorter than the remainder, being c. 40 ju,m
long. The longer setae, including those on the lateral surfaces vary from 50 to
100 /Am in length. Sub-cephalic setae have been seen only in the males. They are
c. 40 /urn. long and occur on or near the posterior edge of the cephalic capsule. In
one case three setae were seen close together, whilst of the remainder only two were
near each other. It has not been possible to be certain about the arrangement of
these setae around the body, as some appear to be missing.

The cephalic capsule is well defined and ornamented with fine granules. Larger
pigmented granules also occur in some specimens. The posterior extremity of the
capsule lies at 0-13 mm from the anterior end of the body. The three mandibles
are large (c. 45-50 /AHI in length). They are of the 'solid' type and sculptured, as in
other species of this genus. As noted by Wieser (1959) in M. sinuosus, there is an
anterior medium segment set off from the remainder of the mandible. There are
also present small apophyses on the external sides of the tips. Arising from the
inside base of each mandible, and curving inwards, is a very prominent tooth.

NEMATODES FROM SOUTHERN AFRICA

29

a b

FIG. 9. Mesacanthoides magna sp. nov. (a) Anterior end of body ; (b) Posterior end of male.

The three teeth are of about equal size and about half the length of the mandibles.
The cephalic ring surrounds the head at about the level of the mid-region of the
mandibles, but it appears to form an indentation, indicated posteriorly between the
mandibles, to join a small curved, cup-like cuticular structure at about the level of
the cephalic setae. On the other hand, the cup-like structure might be a mere fold
at the base of the indentation of the cephalic ring. Surrounding each of the cup-like
structures there is a transversely elongate cuticular thickening. Lying anteriorly
to the points where the ring curves posteriorly a crescentic space in the wall of the
buccal-cavity is to be seen between each mandible. Arising from the anterior
margins of these spaces are finger-like extensions. The amphids are situated near
the bases of the lateral cephalic setae. They are pyriform and rather difficult to
make out.

The oesophagus is of even thickness throughout its length. Its outer walls have
a wavy outline. The nerve-ring was very difficult to make out and was seen only
in one specimen. In this specimen (a female) it occurs at 0-47 mm from the anterior
end of the body. A ventral gland or excretory pore has not been made out.

Male. The spicules are long and narrow and of equal length. They are not very
prominent and poorly cuticularized. The gubernaculum consists of a pair of curved
elongate cuticular structures. In the middle region of each gubernaculum the
ventral surface gives rise to a sharply-pointed triangular projection, and the distal
end of each gubernaculum is somewhat curved ventrally (Fig. gb). Several setae
are scattered on the tail including the posterior cylindrical portion. A vestigial
short tubular supplement lies in the median line at c. 0-12 mm anteriorly to the
cloaca, but it was seen in only one specimen.

30 J. W. COLES

Female. The ovaries are paired, opposed and reflexed. The vulva occurs a little
anteriorly to the middle of the body.

REMARKS. This species is distinguished from others known to belong to the genus
Mesacanthoides by its very large size (about twice the length of those previously
described), the length of the spicules, the form of the gubernaculum and the structure
of the head.

GEOGRAPHICAL DISTRIBUTION. Off Cape Province, South Africa (present report).

Rhabdodemania nancyae Inglis, 1964

TYPE-LOCALITY. Off West coast of Cape Province, South Africa.

MATERIAL STUDIED.

False Bay : i<$ FAL 748.

MEASUREMENTS (in mm).

Body-length 5-00

Body-breadth 0-08

Head-diameter 0-02

Length of oesophagus 0-57

Length of tail 0-16

Distance of nerve-ring from anterior end of body 0-27
Distance of excretory pore from anterior end of body 0-22

Cloacal diameter 0-05

Length of spicules 0-07

Length of gubernaculum 0-04

REMARKS. The male specimen present in this sample is a little larger than the
specimens originally described by Inglis (1964), but there is no doubt that it belongs
to the same species. The buccal-cavity is typically conical, 20 /mi deep. The
longer cephalic setae are c. 15 /mi. The shape of the spicules and gubernaculum and
the form of the tail agree very well with the original description.

GEOGRAPHICAL DISTRIBUTION. Off the coast of South Africa (Inglis, 1964 ;
present report.)

Family ONCHOLAIMIDAE Baylis & Daubney, 1926

Genus ONCHOLAIMUS Dujardin, 1845

Oncholaimus problematicus sp. nov.

(Fig. loa-d)

TYPE-LOCALITY. False Bay, Cape Province, South Africa.

MATERIAL STUDIED.

False Bay : 2?$ FAL 613 ; 2?$ FAL 633 ; i? FAL 648 ; itf i? FAL 654 ;
i$ FAL 674 ; ic? 3$$ FAL 679 ; i<? FAL 689 ; 2$? FAL 748 ;
i? FAL 772 ; i? FAL 832.

NEMATODES FROM SOUTHERN AFRICA 31

H

West Coast Dredgings : 3?$ WCD 86 ; 5?? WCD 206.

South Coast Dredgings : 2&? SCD 134 ; i<£ 22?? SCD 233 ; 2?$ SCD 197 ;

i(J SCD 275.

MEASUREMENTS (in mm). 33 ??

Body-length 6-00 - 8-90 6-40 - 9-90

Body-breadth 0-18 -0-31 0-23-0-32

Head-diameter 0-05 - 0-06 0-05 - 0-06

Length of oesophagus 0-80 - 0-91 0-77 - 0-95

Distance of nerve-ring from anterior end of body 0-42 - 0-45 0-42 - 0-50

Distance of excretory pore from anterior end of body o-n -0-15 0-13-0-15

Length of tail 0-20 - 0-30 0-24 - 0-32

Anal diameter 0-063 - °'°8 0-08 - o-io

Length of spicules 0-09 -0-12

Distance of vulva from anterior end of body 4-60 - 6-90

DESCRIPTION. This species appears to belong to the 'Paralangruensis-group' of
the genus, as described by Wieser (1953), because the tail is more or less straight and
slender and of three or more anal diameters in length, of which more than the distal
half is cylindrical. The cuticle is smooth and setae occur on the body principally
on the tail as well as in the cervical region, where they are not very numerous or
conspicuous. There is a crown of papillae around the mouth and a ring of short
setae c. 6-8 /*m in length further posteriorly. The arrangement of these cephalic
setae is typical of the family, i.e. two on each dorso-lateral and ventro-lateral
surface and a single seta on each lateral surface.

The buccal cavity is large, measuring up to 0-07 mm in length and 0-04 mm in
width. There are three teeth present in the cavity, one dorsally and two sub-
ventrally. The two sub-ventral teeth are slightly unequal in size, as is usual in this
genus, the shorter one about as long as the dorsal tooth. In one of the samples
(FAL 679) a female specimen had all three teeth more or less of equal size. In all
other respects, however, it compares well with the other specimens (Fig. loa-b).
The amphids are somewhat boat-shaped and quite readily seen. They are 10-15 I"-111
in diameter and occur at about the level of the middle region of the buccal-cavity.

The oesophagus is typical of the family in being without any distinct posterior
swelling. The excretory pore lies at a distance of twice the length of the buccal-
cavity from the anterior extremity of the body, whilst the nerve-ring surrounds the
oesophagus at a distance of approximately half its length.

Male. The spicules are paired, simple and slightly curved ventrally. The
proximal ends are a little expanded and knob-like while the distal ends are simply
pointed. A gubernaculum was not detected. There are two longitudinal rows of
setae (about 15-18) one on each side of the cloacal region. A little more than half
the length of the tail is cylindrical and rather narrow, and as stated above setae are
scattered over this region.

Female. There is a single reflexed ovary and the vulva occurs posteriorly to the
middle of the body. Several gravid specimens are present. Up to five eggs have
been seen in the uterus. They are all approximately of the same size, the largest

J. W. COLES

FIG. 10. Oncholaimus problematicus sp. nov. (a-b) Lateral view of head in two specimens
showing variation in position of teeth (from same sample) ; (c) Head of a specimen from
a different sample ; (d) Posterior end of male (a, b and c to same scale).

of which measured 0-15 x 0-12 mm. A Demanian system is present. Although all
the details could not be made out, a uvette was clearly seen. It is evenly rounded
and similar to that found in 0. brachycercus and illustrated by Rachor (1969). The
efferent pores of the system on the lateral sides occur at c. 0-35-0-40 mm anteriorly
to the anus, where the body is a little swollen.

REMARKS. This species differs from others belonging to the 'Paralangruensis-
group' of the genus Oncholaimus in the greater length of the spicules, combined with

NEMATODES FROM SOUTHERN AFRICA 33

the total length of the worm, in the position of the excretory pore and in the size and
arrangement of the sub-ventral teeth.

As noted above, the difference in length of the sub- ventral teeth is not very great
and in one specimen they are more or less equal. The genus Metaparoncholaimus
De Coninck & Stekhoven, 1933, is characterized by the two sub-ventral teeth being
equal, by having a single ovary and by the presence of a Demanian system. The
present species has, nevertheless, been compared also with those assigned to Meta-
paroncholaimus. It appears that the differences between the species belonging to the
'Paralangruensis-group' of Oncholaimus and to Metaparoncholaimus are not very
great, and a close study might possibly show them to be synonymous.

GEOGRAPHICAL DISTRIBUTION. Off Cape Province, South Africa (present report).

Oncholaimus sp.

MATERIAL STUDIED.

Saldanha Bay : 2?? SB 336.

MEASUREMENTS (in mm). 2^

Body-length 3-00, 3-30

Body-breadth 0-07, 0-08

Head-diameter 0-027, °'°3

Length of oesophagus 0-42, 0-46

Distance of nerve-ring from anterior end of body 0-19, 0-22

Distance of excretory pore from anterior end of body 0-07, 0-08

Length of tail 0-12, 0-13

Anal diameter 0-03, 0-04

Distance of vulva from anterior end of body 2-17, 2-40

DESCRIPTION. This species is very similar to Oncholaimus problematicus , except
that it is much smaller (less than half the average length of the other species).
Males have not been found and the two females are mature. One of them is gravid,
the larger specimen having three eggs in the single uterus. These eggs measure
c. o-io x 0-05 mm. A Demanian system is present and the exit pores are situated
at approximately the tail length distance anteriorly from the anus. Amphids are
present at the level of the posterior part of the buccal-cavity. They are boat-
shaped, as in 0. problematicus, and are c. 8 /urn in width. The cephalic setae are
about a fifth of the corresponding head-diameter in length (6 /xm). There are
several longitudinal rows of short setae in the cervical region.

Because no males have been found and the females are different in length from
the species described above, this unnamed species must be considered a species
inquirendum, until males are studied. The three teeth in the buccal-cavity are about
the same size, their apices occurring at about the same level, as in some specimens
of 0. problematicus. As noted above this is, however, a characteristic of the genus
Metaparoncholaimus.

GEOGRAPHICAL DISTRIBUTION. Saldanha Bay, Cape Province, South Africa
(present report).

34 J- W. COLES

Genus FILONCHOLAIMUS Filipjev, 1927

Filoncholaimus capensis sp. nov.

(Fig. na-b)

TYPE-LOCALITY. Off West coast of Cape Province, South Africa.
MATERIAL STUDIED.

West Coast Dredgings : i? WCD 74 ; 2&? 5?? WCD 86 ; 2&? 2?$ WCD 119.
SYNTYPES. BM(NH) reg. nos 1975.855-859.

MEASUREMENTS (in mm).

33 ??

Body-length 6-10 - 7-80 8-20 - 10-50

Body-breadth 0-11—0-14 0-14 — 0-20

Head-diameter 0-04 0-04 - 0-05

Length of oesophagus 0-90-1-15 i-oo - 1-27

Distance of nerve-ring from anterior end of body 0-33 - 0-42 0-42 - 0-50

Distance of excretory pore from anterior end of body 0-14-0-15 0-14 - 0-18

Length of tail 0-40-0-53 1-13 - 0-90

Anal diameter 0.19 0-075 - 0-09

Length of spicules 0-19 - 0-20

Length of gubernaculum 0-06 - 0-07

Distance of vulva from anterior end of body 3-90 - 5-50

DESCRIPTION. The body is slender, being of even width throughout most of its
length. The tail is long and filiform in both sexes, more especially in the female.
The cuticle is quite smooth. The setae on the body occur principally in the cervical
region and on the tail of the male, but they are never very numerous nor prominent.

The arrangement of the cephalic sense organs is typical of the family, i.e. a crown
of six papillae near the mouth and a ring of ten setae, each c. io/xm in length, situated
further posteriorly on the head. Two of these setae lie close together on each side
of the dorso-lateral and each of the ventro-lateral surfaces, and a single seta is
situated on each lateral surface.

The buccal-cavity is large, measuring from 0-055 x °'°33 to 0-08 x 0-03 mm. The
dorsal and subventral teeth are all approximately of similar length, and they extend
anteriorly to near the level of the cephalic setae. The amphids are situated at
about a level with the anterior ends of the teeth. They are oval and measure
c. 15 /u,m in diameter (Fig. na).

The ventral gland is not very prominent, but it has been seen in most of the
specimens. The position of the excretory pore is more or less constant, situated at a
distance of c. 0-14 mm from the anterior end of the body in nearly all the specimens
examined. The oesophagus very gradually widens towards its posterior end, but is
without a distinct bulb. The nerve-ring surrounds the oesophagus just anteriorly
to the middle of its length.

Male. The spicules are long and somewhat curved. The distal ends are claw-like,
while the proximal extremities are slightly expanded and blunt. The gubernaculum

NEMATODES FROM SOUTHERN AFRICA

35

FIG. n. Filoncholaimus capensis sp. nov. (a) Lateral view of head ; (b) Posterior end

of male.

consists of two simple pieces which are similar in shape to the spicules (Fig. lib).
There are two precloacal papillate supplements situated in the median line
resembling those inF. ditlevseni Kreis, 1932. One lies at a distance of c. 0-08 mm in
front of the cloaca and the second at c. 0-13 mm further on. The more anterior
supplement is larger and measures c. 0-05 mm in length. There are two pairs of
setae situated on papillae quite close to the cloaca, and there are small setae scat-
tered over the tail. Posteriorly to the cloaca the tail narrows suddenly and
throughout most of its length it is filiform.

Female. The ovaries are paired, opposed and reflexed. The vulva is situated
approximately in the middle of the body. The tail is very long and filiform, but it
seems that in some specimens the ends have broken off. As in other members of
this genus, a Demanian system is not present.

REMARKS. This species is distinguished from others assigned to the genus
Filoncholaimus in having all three teeth in the buccal-cavity of equal length. More-
over, only in one other species, F. prolatus Hopper, 1967, does the male possess a
gubernaculum, but it is of a different shape and has a dorsal piece as well as a pair of
lateral pieces.

GEOGRAPHICAL DISTRIBUTION. Off West coast of Cape Province, South Africa
(present report).

36 J. W. COLES

Genus PONTONEMA Leidy, 1855
Pontonema yaenae Inglis, 1964

TYPE LOCALITY. Off West coast of Cape Province.
MATERIAL STUDIED.

False Bay : 2?$, I larva FAL 748 ; i? FAL 832.

West Coast Dredging : 2$$ WCD 210.

MEASUREMENTS (in mm). $$ Larva

Body-length 10-00-11-30 7-20

Body-breadth 0-22 — 0-25 0-20

Head-diameter 0-07 - 0-08 0-06

Length of oesophagus 1-23- 1-70 i-io

Length of tail 0-22 - 0-27 0-20

Distance of nerve-ring from anterior end of body 0-50 0-40

Distance of excretory pore from anterior end of body 0-13 - 0-15

Anal diameter 0-09 — o-io 0-07

Distance of vulva from anterior end of body 4-10 - 5-40 —

Length of buccal-cavity o-n - 0-14 o-n

Width of buccal-cavity 0-05 - 0-06 0-04

Length of cephalic setae o-oi o-oi

REMARKS. Only female specimens and one larva are present in the samples.
They are larger than those described by Inglis (1964) and appear to belong to P.
yaenae because of the following features : the form of the buccal-cavity, including
the size, shape and position of the teeth, amphids and cephalic setae. They also
agree with this species in the position of the excretory pore, length and shape of the
tail.

GEOGRAPHICAL DISTRIBUTION. Off West coast of Cape Province (Inglis, 1964) ;
False Bay (present report).

Pontonema sp.
MATERIAL STUDIED.

False Bay : i? FAL 613.

MEASUREMENTS (in mm).

Body-length 5-90

Body-breadth 0-18

Head-diameter 0-07

Length of oesophagus 1-15

Distance of nerve-ring from anterior end of body 0-40
Distance of excretory pore from anterior end of body 0-14

Length of tail 0-20

Anal diameter o-io

Distance of vulva from anterior end of body Not seen

Length of cephalic setae o-oi

Length of buccal-cavity 0-12

Width of buccal-cavity 0-055

NEMATODES FROM SOUTHERN AFRICA 37

REMARKS. This specimen is typical of the genus, but as no males are present it
cannot, at the moment, be identified more precisely. The six labial papillae are
slightly setose. The amphids are about n /mi in diameter and situated at about
the level of the anterior end of the dorsal tooth.

GEOGRAPHICAL DISTRIBUTION. False Bay, Cape Province, South Africa (present
report).

Pontonema sp.

MATERIAL STUDIED.

West Coast Dredging : i$ WCD 206.

MEASUREMENTS (in mm).

Body-length 1 1 -oo

Body-breadth 0-20

Head-diameter 0-09

Length of oesophagus 1-61

Distance of nerve-ring from anterior end of body 0-58
Distance of excretory pore from anterior end of body 0-18

Length of tail 0-28

Anal diameter 0-12

Distance of vulva from anterior end of body 5-40

Length of cephalic setae o-oi

Length of buccal-cavity 0-13

Width of buccal-cavity 0-07

REMARKS. This specimen is coiled and in very poor condition. Its structure
is, however, typical of the genus Pontonema but as no males are present it cannot
be identified specifically. The cephalic sense organs appear to be typical. The
amphids are quite prominent, about 20 (xm in diameter, and are situated at about
the level of the anterior end of the dorsal tooth. Three eggs are present in the
uterus. They measure c. 0-30 x 0-12 mm. There are eight rows of cervical setae
measuring up to 10 (xm in length.

GEOGRAPHICAL DISTRIBUTION. Off West coast of Cape Province, South Africa
(present report).

Family ENCHELIDIIDAE Filipjev, 1918

Genus SYMPLOCOSTOMELLA Micoletzky, 1930

Symplocostomella trichostoma sp. nov.

(Fig. i2a-c)

TYPE-LOCALITY. Off West coast of Cape Province, South Africa.

MATERIAL STUDIED.

West Coast Dredging : i<$ 2?? WCD 86.

SYNTYPES. BM(NH) reg. nos 1975.860-862.

38 J. W. COLES

MEASUREMENTS (in mm).

c? 29?

Body-length 9-01 8-00, 9-90

Body-breadth 0-15 0-29, 0-30

Head-diameter 0-04 0-04, 0-05

Length of oesophagus 1-50 1-41, 1-65

Distance of nerve-ring from anterior end of body 0-50 0-40, 0-39

Distance of excretory pore from anterior end of body 0-09 0-08, 0-09

Length of tail 0-28 0-25, 0-35

Anal diameter o-n 0-12, 0-12

Length of spicules 0-225 —

Length of gubernaculum 0-06 —

Distance of vulva from anterior end of body 4-30, 5-20

DESCRIPTION. Female. The body is of even width throughout most of its
length. The head narrows a little, while the tail is conical and terminates with a
short cylindrical portion, about a third of the total length of the tail. The cuticle
appears to be smooth, although in one of the specimens exceedingly fine transverse
striations are present in parts of the oesophageal and mid-body regions.

The mouth is surrounded by a cuticular ring, as in other members of the genus
Symplocostomella and related genera. Around this ring lies a crown of fine seta-like
elements, reminiscent of the leaf-crown found at the entrance to the mouth in
certain members of the parasitic nematode family Strongylidae. A little further
back on the head a crown of six papillae is situated, and lying at 10 /zm posteriorly
from the anterior end of the body there is a ring of ten setae, each 10-12 /mi long.
Two of these setae are disposed close together on each of the dorso-lateral and the
ventro-lateral surfaces, and a single seta on each lateral surface. There are several
uneven longitudinal rows of short setae on the body in the cervical region. In one
of the females there are two prominent papillae situated close together on the ventral
side, at 0-4 mm from the anterior end of the body. Comparable, but much smaller
and less conspicuous, structures have also been seen in the other female specimen at
approximately the same position.

The buccal-cavity is large, measuring 60 /xm long and 35 /mi at its greatest width.
It is sub-divided at the anterior end by two rings, which are rather difficult to make
out. At about the level of the cephalic setae where the buccal-cavity widens a
little, there is a ring of somewhat pyriform cuticular rodlets. Three teeth are
present in the cavity, two sub-ventral and one dorsal. One of the sub-ventral teeth
is long and spear-like, and arises from the base of the buccal-cavity. The other
sub- ventral tooth is smaller and projects from the buccal-wall about halfway along
its length. The dorsal tooth is wide with a serrated anterior margin. The apices of
all three teeth lie at approximately the same level. Numerous small denticles
occur on the wall of the buccal-cavity close behind the ring of cuticular rodlets. The
amphids are quite clearly seen and are shaped somewhat like an inverted cup. Just
posteriorly to the amphids there are small triangular pieces that appear to be con-
nected to the amphids by a stalk, which may, however, be an artifact.

The oesophagus very gradually widens towards its posterior end, but does not
form a muscular bulb. The ventral gland and duct are very prominent and the

NEMATODES FROM SOUTHERN AFRICA 39

excretory pore is situated at about one and a half times the length of the buccal
cavity from the anterior end of the body.

The ovaries are paired, opposed and reflexed. In one of the specimens two eggs
are present in each uterus, and in the other specimen there are three in one uterus
and two in the other. They are all approximately of the same size, and the largest
measures 0-35 x 0-20 mm. The vulva occurs just a little posteriorly to the middle
of the body.

Male. The only male specimen in the sample of this species is in rather poor
condition. Although it appears that males of the genus Symplocostomella have not
been described hitherto, the present specimen does resemble males of closely-related
genera. Sexual dimorphism is known to occur in the family Enchelidiidae, and,
having been found together, it seems reasonable to assume that this specimen is
conspecific with the females described above. The body is about the same length in
both sexes, the tail is very similar and the position and prominence of the ventral
gland and excretory pore are also similar. The body of the male is, however,
rather stout and very gradually tapers towards the anterior and posterior ends,
presenting a fusiform appearance.

The head is small and bulbous and separated from the rest of the body by a
constriction. There is no crown of fine seta-like elements as found in the female.
There appear to be six small papillae surrounding the simple mouth and further
posteriorly a ring of ten setae. Two pairs of setae lie close together on the dorso-
lateral surfaces and the setae on the lateral surfaces are a little longer than those on
the ventro-lateral surfaces. A buccal-cavity is absent and the head is very simple
with a few cuticular processes. The amphids are very prominent and different
from those in the female. They are cup-shaped, 12 /mi wide and 10 /mi deep (see
(Fig. i2a). Setae occur sporadically in the cervical region.

The internal structures are in very poor condition and may be somewhat mace-
rated. There appears to be a single testis. The spicules are narrow and slender
and slightly curved. They are of even width throughout their length. The proxi-
mal ends are only very slightly expanded, whilst the distal ends are bluntly pointed.
The gubernaculum is simple with a sharp bend in the middle (see Fig. i2c). Extend-
ing 0-98 mm anteriorly from the cloaca there is a row of 19 mid- ventral papillae,
each with a short seta projecting from it. The tail is very similar in shape to that
in the female with a short terminal cylindrical portion. There are short setae
scattered on the tail.

DISCUSSION. Micoletzky (1930) erected the genus Symplocostomella for a new
species, 5. javaensis, and characterized it by the absence of ocelli and lenses and by
the presence of a row of small rodlets at the border of the anterior and posterior
portions of the buccal-cavity. Vitiello (1970) reviewed the genus and included a
key to the species. Only five species were known at that time, and he divided them
into two main groups. In the first group the cephalic setae are greater than 50
per cent of the corresponding head-diameter in length and in the second group they
are 40 per cent or less. The present species falls in Vitiello's second group and
shows some resemblance to 5. mediterranea Schuurmans Stekhoven, 1950, in which
the excretory pore is about 1-6 buccal-cavity lengths from the anterior end of the

4o

J. W. COLES

body. S. trichostoma sp. nov. differs, however, from S. mediterranea in being about
twice as long, as well as in the presence of denticles in the buccal-cavity, in the shape
and size of the amphids and in the ring of seta-like elements around the mouth.

As mentioned above, males of this genus have not been found previously, and the
single male found in the present sample appears to belong to the same species of
females described above, although it cannot yet be proved conclusively. In general

FIG. 12. Symplocostomella trichostoma sp. nov. (a) Head of male ; (b) Head of female

(c) Tail of male.

NEMATODES FROM SOUTHERN AFRICA 41

form it is similar to other males belonging to the family Enchelidiidae, in which
sexual dimorphism is known to occur.

GEOGRAPHICAL DISTRIBUTION. Off Cape Province, South Africa (present report).

Genus EURYSTOMINA Filipjev, 1921
Eurystomina sp.

(Fig. 13)

MATERIAL STUDIED.
False Bay : i$ FAL 412 ; i? FAL 422.

MEASUREMENTS (in mm).

Body-length 6-30, 7-40

Body-breadth 0-08, 0-08

Head-diameter 0-05, 0-045

Length of oesophagus 1-30, 1-60

Distance of nerve-ring from anterior end of body 0-26, 0-26

Length of tail 0-08, 0-09

Anal diameter 0-07, 0-05

Distance of vulva from anterior end of body 3 -60, 4-50

Length of cephalic setae 0-012, o-ou

Length of buccal-cavity 0-04, 0-035

Greatest width of buccal-cavity 0-032, 0-031

Length of cylindrical portion of tail 0-036, 0-030

FIG. 13. Eurystomina sp. Lateral view of head.

DESCRIPTION. Both specimens are in poor condition. The body is of even width
throughout most of its length. The mouth opening is surrounded by a crown of six
papillae, and further posteriorly there is a ring of cephalic setae, most of which appear
to be missing so that their number and arrangement could not be ascertained. The
buccal-cavity is wide and spacious and is divided into two sectors. The anterior
sector is wide and surrounded at its posterior region by four transverse rows of
denticles - those on the right lateral side are large and gradually become smaller and
more numerous towards the left side. At the base of the posterior sector of the
6

42 J. W. COLES

cavity there are two dorso-lateral teeth, one a little longer than the other. Cervical
setae, c. 10 pm long, are arranged in eight longitudinal rows extending to about the
level of the nerve-ring. The internal structures are somewhat macerated, but the
ovaries appear to be paired and opposed. The tail terminates with a short cylindri-
cal portion.

Until males are found and in good condition, a more precise identification does
not seem possible.

GEOGRAPHICAL DISTRIBUTION. False Bay, Cape Province, South Africa (present
report).

Genus POLYGASTROPHORA De Man, 1922
Polygastrophora sp.

(Fig. 14)
MATERIAL STUDIED.

South Coast Dredging : i? SCD 251.

MEASUREMENTS (in mm).

Body-length

Body-breadth

Head-diameter

Length of oesophagus

Distance of nerve-ring from anterior end of body

Length of tail

Anal diameter

Distance of vulva from anterior end of body

6-50
0-08
0-02
i -20
0-52

0-21

0-05

3-70

FIG. 14. Polygastrophora sp. Lateral view of head.

REMARKS. The only female specimen available is in very poor condition. It
appears to be typical of the genus and has six oesophageal bulbs, the first of which
commences at 0-87 mm from the anterior end of the body. There are four teeth in
the buccal-cavity, two situated dorsally and two sub-ventrally. Cephalic setae
were not seen. Only one papilla was found situated laterally near the mouth. Until
more specimens are found no more information on this form can be given.

GEOGRAPHICAL DISTRIBUTION. Off South coast of Cape Province, South Africa
(present report).

NEMATODES FROM SOUTHERN AFRICA 43

Family CYATHOLAIMIDAE De Coninck & Stekhoven, 1935
Gen. et sp. innom.

South Coast Dredging sample : i? SCD 136.
In poor condition. Not considered further.

Family DESMODORIDAE Steiner, 1927
Spirinia sp.

False Bay : 2^, i$ FAL 684.

Although the specimens are in rather poor condition this species will be included
in a review of the genus to be undertaken at a later date.

Family CHROMADORIDAE Filipjev, 1917
Parapinnanema sp.

False Bay : i<J FAL 725 ; i? FAL 748. West Coast Dredging samples : i<j>
WCDi96; i?WCD2o6.

All the specimens which appear to belong to this genus are in very poor condition
and have not been determined more specifically.

HISTORICAL ACCOUNT OF FREELIVING MARINE NEMATODES
FROM SOUTHERN AFRICA

Our knowledge of freeliving marine nematodes from southern Africa began in
1908 when Otto von Linstow reported on a collection of nematodes from west and
central southern Africa. This report included several species of marine nematodes
from Luderitz Bay on the coast of South West Africa. Several years later, Steiner
(1918) published a report on marine nematodes from the west coast of Africa, and
included four species also taken from Luderitz Bay. No other records of freeliving
marine nematodes from southern Africa appear to have been published until 1961.
Professor J. Omer-Cooper, whilst on a visit to the British Museum (Natural History)
in 1960, brought to the attention of Dr W. G. Inglis a small collection of freeliving
nematodes from an interesting locality at the mouth of the Greater Kleinemonde
River in the Bathurst District of Cape Province. Although taken in brackish water
these forms were found to belong to genera which were otherwise known only from
wholly marine localities (Inglis, 1961). A year or so later Professor J. H. Day
presented to the British Museum (Natural History) collections of marine animals
made by the University of Cape Town Ecological Surveys and included quite a large
number of samples of freeliving marine nematodes. These were studied and
reported on in two papers by Inglis (1963, 1964). Later Dr W. D. Oliff from the
South African Institute for Water Research brought to the attention of Dr Inglis a
collection of nematodes from off the coast of Durban, and some of these worms were
also studied and reported upon by Inglis (1966).

6*

44 J- W. COLES

Reports have also been published on marine nematodes from the coast of
Madagascar by Gerlach (1953, 1958). It is very likely that the same species reported
by Gerlach also occur along the coast of Mozambique. Nothing appears to be known
of freeliving marine nematodes on the coast of Angola.

The following list is of the freeliving marine and brackish water forms reported
from the coasts of South West Africa and the Republic of South Africa. They cover
a range from as far north as Luderitz on the west coast to Durban on the east coast.

Specimens described or reported as far as genus only are not included.

LIST OF SPECIES REPORTED FROM SOUTHERN AFRICAN WATERS

Order ENOPLIDA Filipjev, 1929

Family IRONIDAE De Man, 1876

Pheronous ogdeni Inglis, 1966 Inglis, 1966 : 93. Beach-sand in surf-zone,

near Durban.

Thalassironus jungi Inglis, 1964 Inglis, 1964 : 337. West coast of Cape Pro-

vince.

Trissonchulus janetae Inglis, 1961 Inglis, 1961 : 294. Brackish water, mouth of

Greater Kleinemonde River.

Family TRIPYLOIDIDAE FiUpjev, 1918

Bathylaimus deconincki Inglis, 1966 Inglis, 1966 : 103. Beach-sand in surf-zone,

near Durban.

Family LEPTOSOMATIDAE DeConinck & Stekhoven, 1933

Anticoma chitwoodi Inglis, 1964 Inglis, 1964 : 324. West coast of Cape Pro-

vince.
Platycoma sudafrica Inglis, 1966 Inglis, 1966 : 83. Beach-sand in surf-zone,

near Durban.

Platycomopsis barba (Inglis, 1964) Inglis, 1964 : 326. West coast of Cape

(syn. Parabarbonema barba Inglis, 1964) Province.

Present report : False Bay.
Platycomopsis dayi sp. nov. Present report : False Bay. West coast of

Cape Province.
Macronchus shealsi Inglis, 1964 Inglis, 1964 : 328. West coast of Cape

Province.
Present report : False Bay. West coast of

Cape Province.
Deontostoma hopei sp. nov. Present report : Saldanha Bay, Cape

Province.

Deontostoma jae (Inglis, 1964) Hope, 1974 Inglis, 1964:332. West coast of Cape

(syn. Thoracostoma jae Inglis, 1964) Province.

Present report : Saldanha Bay. False Bay.

West coast of Cape Province.
Deontostoma parantarcticum Vitiello, 1975 Vitiello, 1975:341. Lamberts Bay, Cape

Province.
Thoracostoma ancorarium Vitiello, 1975 Vitiello, 1975 : 345. Lamberts Bay, Cape

Province.
Present report : False Bay, Cape Province.

NEMATODES FROM SOUTHERN AFRICA
Thoracostoma angustifissulatum Mawson, 1956
Thoracostoma campbelli Ditlevsen, 1921
Thoracostoma zeae Inglis, 1964

45

Inglis, 1964 : 331. West coast of Cape

Province.
Present report : Luderitz Bay, South West

Africa.
Inglis, 1964 : 334. West coast of Cape

Province.
Present report : False Bay. South coast of

Cape Province.

Family PHANODERMATIDAE Stekhoven, 1935

Dayellus dayi Inglis, 1964

Phanoderma unica Inglis, 1964
Crenopharynx afra Inglis, 1964

Crenopharynx eina Inglis, 1964

Inglis, 1964 : 304. False Bay, Cape Province.
Present report : False Bay. West coast of

Cape Province.
Inglis, 1964 : 309. West coast of Cape

Province.
Inglis, 1964 : 308. South coast of Cape

Province.
Present report : False Bay. West and South

coasts of Cape Province.
Inglis, 1964 : 306. South coast of Cape

Province.
Present report : False Bay. South coast of

Cape Province.

Family ENOPLIDAE Baird, 1853

Enoplus bisetosus v. Linstow, 1908
Enoplus harlockae Inglis, 1964
Enoplus macrolaimus v. Linstow, 1908
Enoplus michaelseni v. Linstow, 1896
Enoploides mandibularis sp. nov.

Mesacanthoides magna sp. nov.
Afracanthion nudus, Inglis, 1964

Enoplolaimus mus Inglis, 1964
Epacanthion oliffi Inglis, 1966
Mesacanthion cavei Inglis, 1964
Mesacanthion ceeus Inglis, 1964
Mesacanthion frica Inglis, 1966
Mesacanthion studiosa Inglis, 1964
Oxyonchus ditlevseni Inglis, 1964

v. Linstow, 1908 : 27. Luderitz Bay, South

West Africa.
Inglis, 1964 : 320. West coast of Cape

Province,
v. Linstow, 1908 : 27. Luderitz Bay, South

West Africa.
Inglis, 1964 : 321. West coast of Cape

Province.
Present report : Saldanha Bay, False Bay.

South coast of Cape Province.
Present report : West coast of Cape Province.
Inglis, 1964 : 316. West coast of Cape

Province.
Inglis, 1964 : 312. West coast of Cape

Province.
Inglis, 1966 : 88. Beach-sand in surf-zone,

near Durban.
Inglis, 1964 : 314. South coast of Cape

Province.
Inglis, 1964 : 314. South coast of Cape

Province.
Inglis, 1966 : 87. Sand in surf-zone, near

Durban.
Inglis, 1964 : 315. West coast of Cape

Province.
Inglis, 1964 : 311. West coast of Cape

Province.

46 J. W. COLES

Rhabdodemania dura Inglis, 1966 Inglis, 1966 : 91. Polluted sediments in the

mouth of Durban Harbour.
Rhabdodemania nancyae Inglis, 1964 Inglis, 1964 : 322. West coast of Cape

Province. Present report : False Bay.
Trileptium ayum Inglis, 1964 Inglis, 1964 : 317. West and South coasts of

Cape Province.
Trileptium longisetosum Inglis, 1966 Inglis, 1966 : 85. Beach-sand in surf -zone,

near Durban.
Thoracostomopsis carolae Inglis, 1964 Inglis, 1964 : 319. West coast of Cape

Province.

Family ONCHOLAIMIDAE Baylis & Daubney, 1926

Oncholaimus problematicus sp. nov. Present report : False Bay. West and

South coasts of Cape Province.

Oncholaimus spiralis v. Linstow, 1908 v. Linstow, 1908 : 27. Luderitz Bay, South

West Africa.

Filoncholaimus capensis sp. nov. Present report : West coast of Cape Province.

Metoncholaimus murphyi Inglis, 1966 Inglis, 1966 : 95. Beach-sand in surf-zone,

near Durban.

Pontonema yaenae Inglis, 1964 Inglis, 1964 : 336. West coast of Cape Pro-

vince.

Present report : False Bay. West coast of
Cape Province.

Wiesoncholaimus mawsonae Inglis, 1966 Inglis, 1966 : 97. Polluted sediments in the

mouth of Durban Harbour.

Family ENCHELIDIIDAE Stekhoven, 1935

Symplocostomella trichostoma sp. nov. Present report : West coast of Cape Province.

Eurystomina sudensis Inglis, 1964 Inglis, 1964 : 335. West coast of Cape

Province.
Polygastrophora omercooperi Inglis, 1961 Inglis, 1961 : 309. Brackish water, Greater

Kleinemonde River, Cape Province.

Order CHROMADORIDA Filipjev, 1929
Family CYATHOLAIMIDAE Filipjev, 1918

Longicyatholaimus dayi Inglis, 1963 Inglis, 1963 : 342. West coast of Cape

Province.
Xyzzors fitzgeraldae Inglis, 1963 Inglis, 1963 : 344. West coast of Cape

Province.

Family CHONIOLAIMIDAE Stekhoven & Adam, 1931

Choniolaimus wieseri Inglis, 1963 Inglis, 1963 : 539. West coast of Cape

Province.

Family DESMODORIDAE Filipjev, 1922

Sigmophoranema brevispiculatum (Inglis, Inglis, 1963 : 537. West coast of Cape

1963) Hope & Murphy, 1972 (syn. Province.

Sigmophora brevispiculata Inglis, 1963)

Desmodora cuddlesae Inglis, 1963 Inglis, 1963 : 532. West coast of Cape

Province.

NEMATODES FROM SOUTHERN AFRICA

47

Desmodora michaelseni Steiner, 1918

Desmodora (Xenodesmodora) nini (Inglis,
1963) Wieser & Hopper, 1967 (syn.
Bla nini Inglis, 1963)

Steiner, 1918 : 17. Luderitz Bay, South West

Africa.
Inglis, 1963 : 534. West coast of Cape

Province.

Family MONOPOSTHIIDAE Filipjev, 1934

Nudora omercooperi Inglis, 1961 Inglis, 1965 : 176. In sand from a small

spring at Claytons Rocks about two kilo-
metres from the Kleinemonde River, Cape
Province.

Family COMESOMATIDAE Filipjev, 1918

Mesonchium janetae Inglis, 1963 Inglis, 1963 : 547. West coast of Cape

Province.

Inglis, 1961 : 301. Brackish water, mouth of
Greater Kleinemonde River, Cape Province.

Mesonchium nini Inglis, 1961

Family CHROMADORIDAE Filipjev, 1917

Hypodontolaimus angelae Inglis, 1961
Euchromadora africana v. Linstow, 1908
Euchromadora dubia Steiner, 1918
Euchromadora longicaudata Steiner, 1918
Euchromadora luderitzi Steiner, 1918

Parapinnanema shirleyae (Coles, 1965)
Warwick & Coles, 1975
(syn. Euchromadora shirleyae Coles, 1965
Austranema shirleyae (Coles, 1965)
Inglis, 1969)

Inglis, 1961 : 305. Brackish water, mouth of

Greater Kleinemonde River, Cape Province,
v. Linstow, 1908 : 28. Luderitz Bay, South

West Africa.
Steiner, 1918 : 10. Luderitz Bay, South West

Africa.
Steiner, 1918 : 15. Luderitz Bay, South West

Africa.
Steiner, 1918 : 12. Luderitz Bay, South West

Africa.
Coles, 1965 : 182. West coast of Cape Pro-

Order ARAEOLAIMIDA DeConinck & Stekhoven, 1933
Family LEPTOLAIMIDAE Oerley, 1880

Plectolaimus juliani Inglis, 1966 Inglis, 1966 : 100. Beach-sand in surf-zone,

near Durban.

Order MONHYSTERIDA Filipjev, 1929
Family LINHOMOEIDAE Filipjev, 1922

Linhomoeus timmi Inglis, 1963 Inglis, 1963 : 550. West coast of Cape

Province.

Family SPHAEROLAIMIDAE Filipjev, 1918

Sphaerolaimus anterides Inglis, 1961 Inglis, 1961 : 313. In brackish water, mouth

of Greater Kleinemonde River, Cape Pro-
vince.

48 J. W. COLES

OF UNCERTAIN TAXONOMIC POSITION

Rhaptothyreus typicus Hope & Murphy, 1969 Hope & Murphy, 1969 : 81. Off south west

coast of South Africa (abyssal).

ACKNOWLEDGEMENTS

I wish to thank Mr S. Prudhoe for his help and valuable criticism throughout this
study.

Thanks are also due to Dr R. M. Warwick of the Institute for Marine Environ-
mental Research, Plymouth, for advice on the families Leptosomatidae and Eno-
plidae ; to Mr C. G. Ogden, of the Electron Microscope Unit of this Museum, for
preparing a specimen of Deontostoma hopei sp. nov. for scanning electron microscopy,
and to my wife for help with the final drawings.

REFERENCES

COLES, J. W. 1965. A critical review of the marine nematode genus Euchromadora De Man,

1886. Bull. Br. Mus. nat. Hist. (Zool.) 12 : 157-194.
DITLEVSEN, H. 1921. Marine free-living nematodes from the Auckland and Campbell Islands.

(Papers from Dr. Th. Mortensen's Pacific Expedition 1914- 16. III.) Vidensk. Medd. dansk.

naturh. For en. 73 : 1-32.

1926. Free-living Nematodes. Dan. Ingolf Exped. 4 (6) : 1-42.

EKMANN, S. 1953. Zoogeography of the Sea. London.

GERLACH, S. A. 1953. Recherches sur la faune des eaux interstitielles de Madagascar. III.

Sur quelques nematodes libres des eaux souterraines littorales de Madagascar. Mem. Inst.

scient. Madagascar Ser. A. Biol. anim. 8 : 73-86.
1958. Recherches sur la faune interstitielle des sediments marins et d'eau douce de

Madagascar. X. Deuxieme contribution a la faune des nematodes des eaux interstielles

littorales de Madagascar. Mem. Inst. scient. Madagascar Ser. F. Oceanogr. 2 : 342-365.
HOPE, W. D. 1967. Free-living nematodes of the genera Pseudocella Filipjev, 1927, Thora-

costoma Marion, 1870 and Deontostoma Filipjev, 1916 (Nematoda Leptosomatidae) from the

west coast of North America. Trans. Amer. micr. Soc. 86 : 307-334.
1974- Deontostoma timmerchioi n. sp., a new marine nematode (Leptosomatidae) from

Antarctica, with a note on the structure and possible function of the ventro-median sup-
plement. Trans. Amer. micr. Soc. 93 : 314-324.
& MURPHY, D. G. 1969. Rhaptothyreus typicus n. g., n. sp., an abyssal marine nematode

of uncertain taxonomic position. Proc. biol. Soc. Wash. 82 : 81-92.
INGLIS, W. G. 1961. Free-living Nematodes from South Africa. Bull. Br. Mus. nat. Hist.

(Zool.) 7 : 291-319.
1963. New marine nematodes from off the coast of South Africa. Bull. Br. Mus. nat.

Hist. (Zool.) 10 : 529-552.
1964. The marine Enoplida (Nematoda) a comparative study of the head. Bull. Br.

Mus. nat. Hist. (Zool.) 11 : 265-376.
1965. Nudora omercooperi sp. nov. (Nematoda : free-living) from an unusual locality in

South Africa. Hydrobiologia 25 : 176-180.
1966. Marine nematodes from Durban, South Africa. Bull. Br. Mus. nat. Hist. (Zool.)

14 : 79-106.
LINSTOW, O. VON 1908. Helminthes. Nematoden und Acanthocephalen aus dem westlichen

und zentralen Sudafrika. Denkschr. med.-naturw. Ges. Jena 13 : 19-28.

NEMATODES FROM SOUTHERN AFRICA 49

MAWSON, P. M. 1956. Free-living nematodes. Section i. Enoploidea from Antarctic

Stations. Rep. B.A.N.Z. antarctic Res. Exp. 6B (3) : 37-74.
1958. Free-living nematodes. Section 3. Enoploidea from Subantarctic Stations.

Rep. B.A.N.Z. antarctic Res. Exp. 6B (14) : 307-358.
MICOLETZKY, H. 1930. Freilebende marine Nematoden von den Sunda-Inseln. I. Eno-

plidae. (Papers from Dr. Th. Mortensen's Pacific Expedition 1914-1916, 53, edited by

H. A. Kreis.) Vidensk Medd. dansk. naturh. Foren. 87 : 243-339.
PLATONOVA, T. A. 1958. Contribution to the Nematode fauna of the family Leptosomatidae

from Kerguelen Island. (In Russian.) Int. Byull. sov. antarkt. Eksped. 1955-1958.

3 : 59-61.
1968. Marine freeliving nematodes of the family Leptosomatidae from Kerguelen Island.

Resultaty biologitscheskich issledowanij sowjetskoj antarktitscheskoj expeditzii 1955-1958.

4. (In Russian.) Issledowanija fauny morej 6 (14) : 5-24.
RACHOR, E. 1969. Das de Mansche Organ der Oncholaimidae, eine genitointestinale Ver-

bindung bei Nematoden. C. Morph. Okol. Tiere 66 : 87-166.
STEINER, G. 1918. Neue und wenig bekannte Nematoden von der Westkuste Afrikas. Zoo/.

Am. 47 : 322-351.

STEKHOVEN, J. H. SCHUURMANS & MAWSON, P. M. 1955. On some free-living marine Nema-
todes from Kerguelen Island. /. Helminth. 29 : 87-104.
VITIELLO, P. 1970. Ne'matodes libres marins des vases profondes du Golfe du Lion. I. Eno-

plida. Tethys 2 : 139-210.
1975- Deontostoma parantarcticum n. sp. et Thoracostoma ancorarium n. sp. nouvelles

especes de Leptosomatidae (Nematoda) d'Afrique du Sud. Trans. Roy. Soc. South Africa

41 : 339-350.

WIESER, W. 1953. Reports of the Lund University Chile Expedition 1948-1949. 10. Free-
living marine nematodes. I. Enoploidea. Ada Univ. lund. N. F. 49 (6) : 1-155.
1959. Freeliving nematodes and other small invertebrates from Puget Sound beaches.

Univ. Wash. Publ. Biol. 19 : 1-179.

J W. COLES

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7 5BD

PLATE i
Head of Deontostoma hopei sp. nov. (Stereoscan electron micrographs.)

Bull. Br. Mus. nat. Hist. (Zool.) 31, i

PLATE i

1

0

a
3"

A LIST OF SUPPLEMENTS
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OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
(Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates.
1965. (Out of Print.)

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy
of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates
77 Text-figures. 1969. £4.50.

4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80.

5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72
Text-figures. 1973. £9.70.

6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa: the
Biology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures.
I974- £3-75- Hardback edition £6.

Printed in Great Britain by John Wright and Sow Ltd. at The Stonebridge Prett, Bristol B&4 jNU

THE ACCACOELIIDAE (DIGENEA) OF
FISHES FROM THE NORTH-EAST

ATLANTIC

R. A. BRAY

AND

D. I. GIBSON

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 31 No. 2

LONDON: 1977

I'BSARY

THE ACCACOELIIDAE (DIGENEA) OF FISHES
FROM THE NORTH-EAST ATLANTIC

BY

RODNEY ALAN BRAY

AND

DAVID IAN GIBSON

Pp. 51-99 ; 15 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)

ZOOLOGY Vol. 31 No. 2

LONDON : 1977

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Scientific
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. 31, No. 2, of the Zoology series.
The abbreviated titles of periodicals cited follow those
of the World List of Scientific Periodicals.

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

ISSN 007-1498

Trustees of the British Museum (Natural History), 1977

BRITISH MUSEUM (NATURAL HISTORY)

Issued 24 March, 1977 Price £4.20

THE ACCACOELIIDAE (DIGENEA) OF FISHES
FROM THE NORTH-EAST ATLANTIC

By RODNEY A. BRAY and DAVID I. GIBSON

CONTENTS

Page

SYNOPSIS .... 53

INTRODUCTION ..... 53

MATERIALS AND METHODS . . 54

SYSTEMATIC SECTION ... 54

Aspects of Biology . . 55

Family Accacoeliidae ...... 56

Subfamily Accacoeliinae . . 57

Accacoelium contortum ...... 58

Accacladium serpentulum ...... 62

Accacladocoelium nigroflavum ..... 65

Accacladocoelium alveolatum .... .68

Accacladocoelium macrocotyle ... 72

Accacladocoelium petasiporum .... 75

Rhynchopharynx paradox a ...... 78

Odhnerium calyptrocotyle ....... 81

Tetrochetus raynerii . . ... 87

Subfamily Paraccacladiinae . ..... 90

Paraccacladium jamiesoni . ...... 91

Paraccacladium sp. (immature) .... 95

HOST-PARASITE LIST ...... 96

ACKNOWLEDGEMENTS ......... 97

REFERENCES ........... 97

SYNOPSIS

The family Accacoeliidae is considered to consist of two subfamilies, the Accacoeliinae and
the Paraccacladiinae subfam. nov. The latter differs from the former in the configuration of
the gut, the absence of a uroproct, the post-ovarian situation of Mehlis' gland and the structure
of the vitellarium. Keys and original descriptions are presented for all of the accacoeliids so
far recorded from the north-east Atlantic region together with full host-records, locality-data
and comments on some aspects of their biology. The species described are : Accacoelium
contortum, Accacladium serpentulum, Accacladocoelium nigroflavum, A. alveolatum, A. macroco-
tyle, A. petasiporum, Rhynchopharynx paradoxa and Odhnerium calyptrocotyle, all from Mola
mola ; Tetrochetus raynerii from Luvarus imperialis ; Paraccacladium jamiesoni gen. et sp. nov.
from Coryphaenoides rupestris and P. sp. (immature) from alepocephalid and centrolophid fishes.

INTRODUCTION

THE HELMINTHS of marine fishes from the north-east Atlantic region have been
studied for about 200 years by numerous workers, notably Olsson, van Beneden,
Odhner, Nicoll, Lebour, Dollfus, Baylis, Punt, Sproston, Rees, Brinkmann, Williams
and Berland. Except, however, for the works of Nicoll (1915), Dawes (1947) and
Dollfus (1953) little has been done to compile lists or detailed descriptions of these
parasites into comprehensive works. Our aim is to produce complete descriptions,

54 R- A. BRAY AND D. I. GIBSON

figures and keys based, wherever possible, on original material, as we feel that this
is much more satisfactory than depending entirely upon existing descriptions which
are often inadequate. This is the first of a projected series of papers covering all
groups of parasitic helminths. The region covered by this series is enclosed by a
line drawn from Gibraltar to Cape Farewell in Greenland. It includes the North
Sea, Skagerrak, Kattegat, Irish Sea and English Channel, but not the Baltic or the
Barents Seas.

MATERIALS AND METHODS

Wherever possible original material has been studied. Some of this material
has been deposited in the British Museum (Natural History) over the years, and its
method of preservation is often unknown and varies considerably. The material
collected by ourselves was obtained during two visits to Plymouth in 1972 (by
D. I. G.), four trips to the North Atlantic aboard the R.V. Cirolana in 1971, 1973,
1974 and 1975 (by R. A. B.) and one visit to Aberdeen in 1973 (by both authors).
The fish from Plymouth and Aberdeen were obtained from catches made by vessels
of the Marine Biological Association and commercial fishing vessels, respectively.
All fish were dissected shortly after capture, usually being freshly killed. The worms,
except cestodes, were fixed in glacial acetic acid for about I minute and preserved in
80% alcohol. Cestodes were fixed using formol-acetic-alcohol, 4% formalin or hot
alcohol, usually under pressure. Whole-mounts were stained in Mayer's paracarmine
and mounted in Canada balsam. Wherever possible, examples of the platyhelminth
species were sectioned and stained with Mayer's haemalum and eosin (H. and E.).
Our studies indicate that, whereas it is usually essential to section platyhelminth-
material for the purpose of accurate determination, it is imperative for their
description.

The descriptions, in which certain key words are italicized, are, where possible,
original, and the references of previous recognizable descriptions are given prior to
our own.

SYSTEMATIC SECTION

The classification of the family Accacoeliidae is a matter of some contention.
Three main proposals have been put forward : (i) Dollfus (1935, I96oa) accepts two
subfamilies, the Accacoeliinae and the Tetrochetinae ; (2) Skrjabin & Guschanskaja
(1959) accept five subfamilies, the Accacoeliinae, Guschanskianinae, Orophocoty-
linae, Rhynchopharynginae and Tetrochetinae ; and (3) Yamaguti (1958) accepts
four, the Accacoeliinae, Accacladiinae, Orophocotylinae and Rhynchopharynginae,
and later (1971) five subfamilies, including the Guschanskianinae. Dollfus (i96oa)
indicates that the systems presented by Yamaguti and by Skrjabin & Guschanskaja
are, in neither case, satisfactory. Dollfus' detailed criticism has, however, produced
no reaction from Yamaguti (1971). We believe that Dollfus' classification is by far
the most acceptable of the three, but prefer to place all of the genera known hitherto
into one subfamily, the Accacoeliinae, as the two subfamilies accepted by Dollfus
clearly grade into one another in the form of Odhnerium Yamaguti, 1934. This will
be discussed in more detail elsewhere (p. 86). We do, however, consider it necessary
to erect a second subfamily, the Paraccacladiinae, for Paraccadadium, a new genus

ACCACOELIIDAE FROM N.E. ATLANTIC 55

to be described in this paper. It differs from the remainder of the accacoeliids in the
structure of the gut, the vitellarium and in the post-ovarian position of Mehlis' gland.
The latter subfamily appears to have a morphology somewhat intermediate between
the accacoeliine accacoeliids and the sclerodistomids. Full details of classification
and definitions of these taxa will be presented elsewhere (Gibson & Bray, in prep.).
The classification of the accacoeliids is based primarily on the structure of the
reproductive system. Although most of the terminology relating to the reproductive
system is well established, there are two features of the terminal genitalia which are
less familiar, as they tend to be restricted to hemiuroids. The sinus-organ is an
intromittent copulatory process lying within the genital atrium and through which
runs the hermaphroditic duct. The sinus-sac (hermaphroditic-sac) is a muscular
structure which surrounds the base of the genital atrium, and into which both male
and female ducts pass. This structure appears to be partly analogous with the
cirrus-sac of many other digeneans in that it usually contains an intromittent organ.
These two features can only be studied satisfactorily in serial sections.

Aspects of Biology

Adults of all but two of the ten species of accacoeliids from the north-east Atlantic
are recorded from the sun-fish Mola mola. One of the other species, which we
believe represents a new subfamily, is recorded from the macrourid Coryphaenoides
rupestris. Further immature forms of the same genus are described from alepoce-
phalid and centrolophid fishes. The parasites of Mola mola represented in the region
are all widespread, and are probably present wherever their wide-ranging host is
found. Mola mola is described as a 'high-ocean circum-global warm-water fish' by
Ekman (1953).

The first intermediate host of accacoeliids is not known, but numerous records
indicate that the metacercariae are found in planktonic animals. Metacercaria
owreae Hutton, 1954, was thought by Dawes (1959) to be the cercaria of a species of
Accacladocoelium Odhner, 1928. This larva has been recorded in both Atlantic
and Indian Oceans from many species of planktonic chaetognaths. The evidence
that this larva is in fact an accacoeliid is not unequivocal although the configuration
of the gut is somewhat like that of the accacoeliines. More certain identifications of
accacoeliid metacercariae have been made on specimens from siphonophores,
scyphomedusae, ctenophores and chaetognaths. These records are summarized by
Dollfus (1963) and Rebecq (1965). In fact, the original description of Odhnerium
calyptrocotyle (Monticelli, 1893) is based on specimens from the ctenophore Beroe
ovata. Nikolaeva (1968) has recorded various immature accacoeliids from pelagic
fishes in the Mediterranean Sea, and considers that these represent 'cystless' meta-
cercariae. The pelagic teleosts may, therefore, act as paratenic hosts. Presumably
this is how such predatory fishes as Coryphaena and Coryphaenoides, the usual final
hosts of Tetrochetus coryphaenae Yamaguti, 1934, and Paraccacladium, respectively,
become infested. It is worthy of note that, according to Bigelow & Schroeder
(I953)» Mola mola, as a rule, feeds upon coelenterates, ctenophores and salps, and
that all four species of teleost from which we have recorded immature specimens of

56 R. A. BRAY AND D. I. GIBSON

Paraccacladium sp. are known to feed extensively on medusae. Furthermore,
the stomach of one of the latter teleosts, Alepocephalus bairdii, was found by us to
be filled with the bathypelagic scyphomedusa Atolla wyvillei. Luvarus imperialis,
the host of Tetrochetus raynerii, also 'feeds primarily upon jellyfish and similar
gelatinous planktonic forms' (Fitch & Lavenberg, 1971). Other species of Tetrochetus
and Paraccacladium jamiesoni have been recorded as adults in predatory fishes, and,
therefore, it seems likely that the suggested paratenic teleost-host may, in fact, be an
obligatory intermediate host. Within the Accacoeliidae it is possible, therefore,
that there are two types of life-history : (i) direct from the coelenterate, etc., when
sun-fishes and Luvarus are the final hosts ; and (2) via a coelenterate-feeding teleost
(?) third intermediate host when predatory teleosts are the final hosts.

Family ACCACOELIIDAE Odhner, 1911

DIAGNOSTIC FEATURES. Body large or small, commonly elongate. Body-surface
smooth, but forebody may be papillate. Ventral sucker in anterior half of body, may
be pedunculate. Prepharynx absent. Pharynx well developed, with narrow
extension into base of oral sucker. Gut-caeca with or without long, anteriorly
directed diverticula ; terminate blindly or form uroproct. Testes two, usually
oblique but may be symmetrical or in tandem, normally close to middle of body ;
pre-ovarian. Seminal vesicle thin walled, tubular, sinuous or convoluted, commonly
reaching into hindbody. Pars prostatica tubular. Hermaphroditic duct, sinus-
sac and sinus-organ usually present. Genital pore mid- ventral in anterior region
of forebody. Genital atrium present. Ovary oval ; post-testicular Laurer's canal
and uterine seminal receptacle present. Uterus extensive, usually reaching to
posterior extremity before looping forward. Eggs small, very numerous, non-
filamented. Vitellarium composed of numerous filamentous tubules ; with or
without left side of system reduced to a rudiment. Excretory vesicle Y-shaped ;
arms initially in dorsal and ventral fields, unite dorsally to pharynx or oral sucker.

FIG. i.

Diagrams of intestinal and vitelline configuration in (a) Accacoeliinae and
(b) Paraccacladiinae.

ACCACOELIIDAE FROM N.E. ATLANTIC 57

KEY TO THE SUBFAMILIES OF THE ACCACOELIIDAE IN
THE NORTH-EAST ATLANTIC REGION

i. A. Gut distinctly H-shaped ; uroproct present ; oesophagus very long and narrow ;
vitellarium with single main collecting duct and system of branching tubules ;
Mehlis' gland pre-ovarian ..... Accacoeliinae (Fig. la) (p. 57)

B. Gut not clearly H-shaped ; uroproct absent ; oesophagus relatively short and
stout ; vitellarium with symmetrical pair of collecting ducts and systems of
branching tubules ; Mehlis' gland post-ovarian . Paraccacladiinae (Fig. ib) (p. 90)

Subfamily ACCACOELIINAE Odhner, 1911

Tetrochetinae Looss, 1912, emend. Dollfus, 1935.
Accacladiinae Yamaguti, 1958.
Orophocotylinae Yamaguti, 1958.
Rhynchopharynginae Yamaguti, 1958.
Guschanskianinae Skrjabin, 1959.

DIAGNOSTIC FEATURES. Body usually elongate. Oesophagus long and narrow.
Gut H-shaped. Uroproct present. Mehlis' gland pre-ovarian. Vitellarium fila-
mentous, with single main collecting duct and single system of ramifying branches
in fore- or hindbody ; left side of system reduced.

KEY TO THE GENERA OF THE ACCACOELIINAE IN
THE NORTH-EAST ATLANTIC REGION

1. A. Well-defined sinus-sac and sinus-organ absent ....... 2

B. Well-defined sinus-organ and sinus-sac present (i.e. clearly visible in sections) . 3

2. A. No sinus-organ, but small male papilla may be present ; sinus-sac rudimentary ;

sucker-like muscular pads present dorsally in forebody ; two muscular flanges on

ventral sucker Odhnerium Yamaguti, 1934 (?• 8l)

B. Sinus-organ and sinus-sac absent ; antero-dorsal muscular pads and flanges on

ventral sucker absent ..... Tetrochetus Looss, 1912 (p. 86)

3. A. Ectoparasitic on gills ; strongly developed ventral musculature in hindbody ;

very long proboscis-like sinus-organ ; enormous pars prostatica occupying much of
forebody ; vitellarium posterior to anterior testis Accacoeliurn Monticelli, 1893 (p. 57)
B. Endoparasitic in gut ; short, cylindrical or dome-shaped sinus-organ ; vitellarium

not usually extending posteriorly to ovary ....... 4

4. A. Pharynx pyriform with anterior extension into base of oral sucker ... 5
B. Pharynx consists of two bulbs and an anterior, elongate portion ensheathed in

glandular posterior process of oral sucker ; glandular oesophageal bulb immediately
posterior to pharynx ; large glandular structure of unknown function at base of
peduncle of ventral sucker . . . Rhynchopharynx Odhner, 1928 (p. 77)

5. A. Vitellarium confined to hindbody ; ventral sucker on extensible peduncle ; pars

prostatica relatively short .... Accacladium. Odhner, 1928 (p. 61)
B. Vitellarium wholly or partly in forebody ; ventral sucker sessile or nearly so ; pars

prostatica relatively long . . . Accacladocoelium Odhner, 1928 (p. 65)

ACCACOELIUM Monticelli, 1893

DIAGNOSTIC FEATURES. Forebody papillate. Oesophagus reaches to ventral
sucker. Ventral sucker on short peduncle. Thick muscular layer in ventral hind-
body. Sinus-sac present, surrounding base of genital atrium. Sinus-organ long

58 R. A. BRAY AND D. I. GIBSON

and strongly muscular, frequently extending through genital pore. Enormous pars
prostatica and associated gland-cells occupy much of forebody. Vitellarium
posterior to anterior testis. Parasitic on gills of marine teleosts (Mola).

TYPE-SPECIES. Accacoelium contortum (Rudolphi, 1819) (by subsequent desig-
nation : Looss, 1899, 631).

Accacoelium contortum (Rudolphi, 1819) Looss, 1899
(Figs 2 and 3)

Distoma contortum Rudolphi, 1819.
Podocotyle contortum (Rud.) Stossich, 1898.

TYPE-HOST AND LOCALITY. Mola mola, Naples, Italy.

RECORDS.
(i) Material studied.

Mola mola [gills] Skagen, Denmark (58°N, n°E) (Oct. 1923). Natural History
Museum of Gothenburg Reg. No. 23-3991.

Mola mola [gills] Locality unknown. BM(NH) Reg. No. 1973.1.25.17-20.
(ii) From the literature.

Mola mola [gills] Ireland (July 1839). Bellingham (1844 : 427).

Mola mola [gills] Varberg, Sweden (Nov.). Olsson (1868 : 39).

PREVIOUS DESCRIPTIONS. Olsson (1868) ; Monticelli (1893) ; Linton (1898, 1940) ;
Timon-David & Musso (1971).

DESCRIPTION. Fig. 2 represents the shape assumed by all the specimens, although
in some cases the hindbody is longer in proportion to the forebody. The body-surface
is not spined, but very wrinkled due to contraction, and numerous papillae ornament
the forebody. The ventral sucker is borne on a short peduncle. Ventrally in the
parenchyma of the hindbody there is a thick layer of longitudinal muscle.

The* measurements given in Table I are taken from laterally mounted specimens.
Anteriorly, the pharynx extends into the base of the oral sucker forming a conical
projection (Fig. 3), the main body of the pharynx measuring 0-3 mm in diameter.
There is a long, narrow oesophagus, which measures about 2 mm and possesses a
cuticular lining. It bifurcates dorsally to the ventral sucker to form two caeca
which give off anteriorly-directed blind extensions. The wide, laterally undulating
caeca reach close to the posterior extremity, where they unite with the excretory
vesicle to form a uroproct.

The terminal excretory pore leads into an excretory vesicle which is essentially
Y-shaped, the basal region being very short. Closely anterior to the union with the
gut-caeca, the vesicle divides forming a dorsal and a ventral arm, which are con-
voluted and pass anteriorly. They run laterally in the forebody and unite behind
the pharynx.

The testes are irregularly globular or oval, situated in tandem and quite close
together in the anterior half of the hindbody. The seminal vesicle is a relatively
narrow and greatly convoluted tube, lying mainly dorsally to the ventral sucker. At
about the level of the anterior margin of this sucker the seminal vesicle narrows and

ACCACOELIIDAE FROM N.E. ATLANTIC

59

FIG. 2. Accacoelium contortum: lateral
view (gut omitted) . Scale -i mm.

FIG. 3. Accacoelium contortum : diagrammatic
longitudinal section of terminal genitalia.

60

R. A. BRAY AND D. I. GIBSON

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ACCACOELIIDAE FROM N.E. ATLANTIC 61

leads into an enormous, convoluted and tubular pars prostatica, which occupies
much of the forebody of the worm. The duct is surrounded by a deep layer of large
glandular cells which are delimited by either a membrane or a local compaction of
parenchyma. A very short ejaculatory duct leads from the pars prostatica, through
the wall of the sinus-sac, into the base of the hermaphroditic duct. The latter is
long, running as it does through the centre of a long, muscular sinus-organ (Fig. 3).
The sinus-organ is frequently found extended through the genital pore, which occurs
mid-ventrally close behind the oral sucker. When withdrawn, the sinus-organ does
not appear to contract as much as is seen in some other hemiuroids, but lies folded
within the deep genital atrium. The sinus-sac envelops the proximal extremity of
the genital atrium, and consists of a relatively diffuse musculature.

The ovary is transversely oval and of a similar size to the testes. It is post-
testicular, lying just inside the posterior half of the hindbody. The oviduct, on
leaving the ovary, passes into a compact, but well-developed, Mehlis' gland which
lies directly anterior to the ovary. Before entering Mehlis' gland, however, it
receives the common vitelline duct and Laurer's canal. The latter passes antero-
dorsally and opens to the exterior dorsally to the posterior testis. The first coils of
the uterus contain large amounts of sperm, and thus form a uterine seminal receptacle
which lies between the ovary and the posterior testis. The remainder of the uterus
fills much of the hindbody, is greatly convoluted and contains numerous small
operculate eggs. The uterus becomes much less convoluted as it passes dorsally to
the ventral sucker, and then runs forward, undulating gently, to its union with the
ejaculatory duct at the base of the sinus-sac. This latter part of the uterus within
the forebody, which may be referred to as the metraterm, is lined by a narrow epi-
thelial layer, possesses a muscular wall and is invested by a thin layer of glandular
cells. The vitellarium consists of a complicated ramifying system of tubules, lying
laterally between the level of the anterior testis and the posterior end of the body.
This system is drained by a single (right) main collecting duct, the left-hand system
being reduced to a small blind 'vitelline reservoir' which lies ventrally to Mehlis' gland.

DISCUSSION. This is the only known species of the genus and is unusual because
of its ectoparasitic habitat on the gills. Morphologically it is rather similar to
Accadadium, which normally occurs in the intestine of Mola mola. It is possible
that the well-developed ventral musculature of the hindbody is associated with a
prehensile function, as the animal must be continually exposed to the stream of
water over the gills.

ACCACLADIUM Odhner, 1928

DIAGNOSTIC FEATURES. Body-surface smooth. Oesophagus reaches to ventral
sucker. Ventral sucker on short peduncle. Pars prostatica fairly well developed.
Sinus-sac surrounding base of genital atrium. Sinus-organ short, cylindrical.
Vitellarium between ventral sucker and ovary. Parasitic in intestine of marine
teleosts (Mola).

TYPE-SPECIES. Accadadium serpentulum Odhner, 1928 (by original designation
and monotypy).

62

R. A. BRAY AND D. I. GIBSON

Accacladium serpentulum Odhner, 1928
(Fig. 4)

Distomum nigroflavum of Monticelli, 1893, and of Linton, 1898, nee Rudolphi, 1819.
Accacladium nematulum Noble & Noble, 1937.

TYPE-HOST AND LOCALITY. Mola mola, Bergen, Norway.

RECORDS.
(i) Material studied.

Mola mola [intestine] Kattegat, Sweden (58°N, n°E) (Dec. 1929). Natural

History Museum of Gothenburg Reg. No. 29-5153.
(ii) From the literature.

Mola mola [intestine] Bergen, Norway. Odhner (1928 : 173).

a

FIG. 4. Accacladium serpentulum : (a) lateral
view (gut omitted) ; (b) forebody showing
peduncle extended. Scale - 2 mm.

ACCACOELIIDAE FROM N.E. ATLANTIC 63

PREVIOUS DESCRIPTIONS. Monticelli (1893) ; Odhner (1928) ; Linton (1898,
1940) ; Yamaguti (1934) ; Noble & Noble (1937) ; Timon-David & Musso (1971).

DESCRIPTION. Two whole worms were available for study, the dimensions of
which are given in Table 2.

The body of this form is elongate, with the ventral sucker surmounted upon a
peduncle of variable length (Fig. 4a and b). The body-surface is smooth, lacking
papillae. The oral sucker is globular with a prominent conical projection at its base,
caused by the elongate anterior portion of the pharynx. A long, narrow, oesophagus
leads from the pharynx and reaches back at least to the level of the ventral sucker.
The H -shaped gut-caeca reach forward, from the region of the ventral sucker, to the
pharyngeal region and backward to the posterior extremity, where they connect
with the excretory vesicle to form a uroproct.

The excretory vesicle is Y-shaped, the arms being dorsal and ventral in the hindbody,
lateral in the forebody, and uniting dorsally in the region of the pharynx. The basal
region of the vesicle is very short.

The elongate oval testes may have slightly indented margins and occur in the
middle-region of the hindbody, with the posterior testis lying ventrally and the
anterior testis dorsally. The seminal vesicle is long, tubular, coiled and reaches
from just in front of the anterior testis to a region dorsal to the ventral sucker. The
pars prostatica is also coiled and is surrounded by numerous gland-cells. It passes
anteriorly from the level of the ventral sucker to the base of the sinus-sac. A very
short ejaculatory duct joins with the metraterm just inside the sinus-sac to form the
hermaphroditic duct, which runs through a prominent, muscular sinus-organ. The
latter organ is not as large as that in Accacoelium, but may protrude through the
genital pore, and lies in a fairly deep genital atrium. The genital pore is mid- ventral
in the forebody, slightly posterior to the oral sucker. The sinus-sac, which surrounds
the base of the genital atrium, is composed of diffuse muscle and is somewhat
difficult to distinguish in whole-mounts.

The post-testicular ovary is more or less globular, and Mehlis' gland is pre-ovarian.
Laurer's canal has been described by other authors, and a uterine seminal receptacle
is present. The uterus itself reaches fairly close to the posterior extremity and fills
much of the hindbody. The metraterm may be distinguished by its thick wall.
The vitellarium consists of numerous branching tubules which extend from just
posterior to the peduncle of the ventral sucker to the anterior margin of the ovary.

DISCUSSION. Odhner's (1928) is the only previous record of this species from the
north-east Atlantic, but it has been recorded in the Mediterranean Sea as well as on
the Atlantic and Pacific coasts of North America and off Japan. Noble & Noble
(1937) described a new species, A. nematulum, from off California, but its validity
does seem somewhat questionable. Very similar specimens under the same name
have been described from the Mediterranean Sea by Timon-David & Musso (1971),
and this description suggests that the Calif ornian form may not be distinct. Noble
& Noble comment on five characters which they consider may separate nematulum
and serpentulum. As far as the length, length -width ratio and elongation of hind-
body is concerned, there is little doubt that this species is variable enough to in-
validate these distinctions (see Table 2). The size and shape of the ventral sucker

64 R. A. BRAY AND D. I. GIBSON

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ACCACOELIIDAE FROM N.E. ATLANTIC 65

probably vary with fixation, and the egg-size, as recorded by various authors, has a
continuous range of 27-37 fjun x 17-27 /u,m. Finally, the size of the sinus-organ
should not be considered as a taxonomic criterion, as it is a protrusible muscular
organ and can therefore be expected to be found in a number of conditions, and
indeed may be influenced by fixation techniques. We, therefore, consider these
species to be identical.

ACCACLADOCOELIUM Odhner, 1928

Guschanskiana Skrjabin, 1959.

DIAGNOSTIC FEATURES. Body smooth. Lateral flanges on ventral sucker
occasionally present. Oesophagus reaches to ventral sucker. Pars prostatica long.
Sinus-sac surrounding base of genital atrium. Sinus-organ short, cylindrical.
Vitellarium reaches anteriorly to oral sucker, may extend posteriorly just past ovary
but usually not beyond anterior testis. Parasitic in intestine of marine teleosts
(Mola).

TYPE-SPECIES. Accacladocoelium nigroflavum (Rudolphi, 1819) (by original
designation).

KEY TO SPECIES IN NORTH-EAST ATLANTIC REGION

1. A. Muscular flanges on ventral sucker absent ....... 2

B. Muscular flanges on ventral sucker present . A. petasiporurn Odhner, 1928 (p. 75)

2. A. Vitellarium reaching to anterior margin of anterior testis ..... 3
B. Vitellarium reaching just posterior to ovary . A. macrocotyle (Diesing, 1858) (p. 72)

3. A. Numerous small papilla-like extensions at posterior end

A. alveolatum Robinson, 1934 (P- 68)
B. Normal body-surface at posterior end . A. nigroflavum (Rudolphi, 1819) (p. 65)

Accacladocoelium nigroflavum (Rudolphi, 1819) Robinson, 1934

(Fig. 5)

Distoma nigroflavum Rudolphi, 1819.

Accacoelium nigroflavum (Rud.) Liihe, 1901.

Echinostoma nigroflavum (Rud.) Barbagallo & Drago, 1903.

Distoma megnini Poirier, 1885.

Accacladocoelium megnini (Poirier) Dollfus, 1946.

TYPE-HOST AND LOCALITY. Mola mola, Naples, Italy.

RECORDS.

(i) Material studied.
Mola mola [intestine] Kattegat, Sweden (58°N, n°E) (Dec. 1929). Natural

History Museum of Gothenburg Reg. No. 29-5153.
Mola mola [intestine] Rockall, N Atlantic (Aug. 1956). BM(NH) Reg. No.

1964.7.16.1-20.
Mola mola [intestine] Porlock, Somerset, England (Aug. 1966). BM(NH) Reg.

No. 1967.10.2.1-10.

66

R. A. BRAY AND D. I. GIBSON

(ii) From the literature.

Mola mola [intestine] Ireland (July 1839). Bellingham (1844 : 427).
Mola mola [intestine] Atlantic WSW of Great Britain (48°N ig0W) (Sept. 1886).
Guiart (1938 : 41).

PREVIOUS DESCRIPTIONS. Poirier (1885) ; Timon-David & Musso (1971).

LIFE-HISTORY. Not known, but Dawes (1959) claims that Metacercaria owreae
Hutton, 1954, is the cercaria of a species of Accadadocoelium (see above p. 55).
This larval form was recorded from chaetognaths, but it seems likely that Accada-
docoelium spp., like other accacoeliine genera, also use coelenterates and ctenophores
as intermediate hosts.

DESCRIPTION. This description is based on six whole-mounts and one sectioned
specimen. The worms have an elongate, roughly cylindrical shape, with a promi-
nent ventral sucker situated close to the anterior end (Fig. 5) . Maximum breadth is
at the level of the ventral sucker. The dimensions of this worm are given in Table 3.
The body-surface is irregularly rugate, but bears no spines, papillae or regular
plications.

Name used

TABLE 3

Dimensions of Accadadocoelium nigroflavum as recorded by

Timon-David & Musso,
1971

A ccacladocoelium
nigroflavum

Poirier, 1885
Distoma megnini

Host
Locality
Length, mm
Max. width, mm
Oral sucker, mm
Ventral sucker, mm
Pharynx, mm
Testes, mm
Ovary, mm
Eggs, /im

unknown fish

10

28 X 21

Mola mola
Gulf of Marseilles
8-9 -10-7

'•5 - i'55

0-40- 0-42 x 0-48-0-50
0-81- 1-04x0-74-0-84
0-19- 0-20x0-15
1-03- 1-15x0-76-1-02
0-82- 0-91x0-72-0-78
25x15

various authors

Present Study

A ccacladocoelium

nigroflavum

Mola mola

see text

6-5 -25

1-04- 1-86

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0-31 x 0-54

0-50- 2-16 x 0-30-0-86

O-22- 0-68x0-43-0-89

24-36 x 16-25

An anterior extension of the pharynx protrudes into the base of the oral sucker.
A very narrow (0-025 mm) and long (about 2 mm) oesophagus reaches back to about
the posterior margin of the ventral sucker, where it leads into wide gut-caeca which
have anterior extensions reaching to the level of the pharynx. The caeca reach
nearly to the posterior end of the body where they open into the excretory vesicle,
forming a uroproct.

The terminal excretory pore leads into a Y-shaped excretory vesicle, the arms of
which pass dorsally and ventrally in the hindbody, but could not be traced in the
forebody.

The large oval testes lie in tandem, near the ventral surface, or with the anterior
testis more dorsal than the posterior, and fairly close behind the ventral sucker.
The seminal vesicle is a relatively narrow, convoluted tube, lying dorsally to the

ACCACOELIIDAE FROM N.E. ATLANTIC

67

FIG. 5. Accacladocoelium nigroflavum : lateral
view (most of gut omitted). Scale - i mm.

ventral sucker. At about the level of the anterior margin of the ventral sucker, the
seminal vesicle passes, via a short, aglandular duct, into a long gently sinuous pars
prostatica which reaches to the base of the sinus-sac. The pars prostatica is sur-
rounded by numerous gland-cells lying free in the parenchyma. The sinus-sac is a
small, rather weakly-developed muscular structure surrounding the base of the
genital atrium. Just within the wall of the sac a very short ejaculatory duct unites
with the metraterm to form a hermaphroditic duct, which passes through the centre
of a small, cylindrical sinus-organ. The latter usually almost fills the genital atrium
and may extend slightly through the genital pore, which is situated ventrally to the
pharynx.

68 R. A. BRAY AND D. I. GIBSON

The globular ovary is situated posteriorly to the testes in the middle of the body.
Immediately anterior to the ovary lies a large Mehlis' gland. Laurer's canal leaves
the oviduct, passes close to the level of the posterior margin of the ovary before
running across the body and opening dorsally at the level of the anterior margin of
the ovary. The initial coils of the uterus contain much spermatozoa, forming a
fairly compact uterine seminal receptacle. The remainder of the uterus, containing
numerous eggs, fills the bulk of the hindbody, and passes well forward into the fore-
body with one loop reaching almost to the pharynx. The metraterm reaches from
the posterior margin of the ventral sucker to just inside the sinus-sac and is sur-
rounded by gland-cells. The vitellarium consists of a ramifying system of tubules
which, while mostly confined to the forebody of the worm, may reach to the anterior
testis. Ventrally to the ovary lies a large 'vitelline reservoir', which communicates
with Mehlis' gland by means of a short duct. This 'reservoir' and duct is, apparently,
all that remains of the left side of the vitelline system.

DISCUSSION. Poirier (1885) describes and figures this species very well under the
name Distoma megnini. Odhner (1928), after examining about sixty of Rudolphi's
original specimens of Distomum nigroflavum, considered them to be synonymous with
Poirier's form. The specimen of D. nigroflavum figured by Olsson (1868) lacks a
sinus-organ and apparently has the sucker-like pads on the antero-dorsal surface
which are so characteristic of Odhnerium calyptrocotyle. The record of this latter
species from Varberg by Dollfus (1935) suggests that he considered that Olsson's
specimens belong to 0. calyptrocotyle. We agree with this and have, therefore,
included D. nigroflavum of Olsson as a synonym of 0. calyptrocotyle.

Accacladocoelium alveolatum Robinson, 1934
(Fig. 6)

Guschanskiana alveolatum (Robinson) Skrjabin, 1959.

TYPE-HOST AND LOCALITY. Mola mola, Devon, England.
RECORDS.

(i) Material studied.
Mola mola [intestine] Salcombe, Devon, England. BM(NH) Reg. No.

1968.12.2.103-104. [Two type-specimens which were originally deposited in

the Molteno Institute, Cambridge, and were presented to the British Museum

(Natural History) in 1968.]
Mola mola [intestine] Kattegat at Gothenburg, Sweden. Material lent by J.

Thulin.
Mola mola [intestine] Whitby, Yorkshire, England (Aug. 1973). BM(NH) Reg.

No. 1975.10.31.25.
(ii) From the literature.
Mola mola [intestine] Salcombe, Devon, England (June, 1933). Robinson

(1934 : 346).
Mola mola [intestine] Kattegat at Gothenburg, Sweden (Nov. 1971). Thulin

(1973 : 83).

ACCACOELIIDAE FROM N.E. ATLANTIC

69

FIG. 6. Accacladocoelium alveolatum : lateral view (most of gut omitted). Scale - i mm.

PREVIOUS DESCRIPTIONS. Robinson (1934) ; Thulin (1973).

DESCRIPTION. Based upon two type-specimens, one of which we sectioned, four
flattened specimens from Yorkshire and on two flattened and one sectioned speci-
men from Thulin. The original specimens are contracted and the tegument is
thrown into numerous annular rugae, which are much less evident in Thulin's
material. As pointed out by Robinson (1934), the peculiar distribution of minute
papillae at the posterior end of the body gives the superficial appearance of a large
terminal sucker (Fig. 6). The oral sucker and sessile ventral sucker were measured
from sections in lateral view. The various dimensions of these worms are included
in Table 4.

The pharynx is relatively small and extends forward slightly into the base of the
oral sucker. A long, narrow, oesophagus passes back to the level of the posterior
margin of the ventral sucker, where it divides to form two somewhat convoluted gut-
caeca, which extend forward to the level of the pharynx and backward close to the
posterior extremity, where they join with the excretory vesicle to form a uroproct.

70

R. A. BRAY AND D. I. GIBSON

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ACCACOELIIDAE FROM N.E. ATLANTIC 71

The wide, terminal excretory pore leads into a Y-shaped excretory vesicle, with
dorsal and ventral arms. The basal region is so short that the vesicle appears almost
V-shaped. In the forebody the arms run laterally and unite dorsally to the ventral
sucker.

The testes lie close to the ventral sucker, in tandem and just anteriorly to the middle
of the body. The seminal vesicle is fairly narrow, being a convoluted tube lying
dorsally to the ventral sucker. It passes into a sinuous pars prostatica, which is
invested with a deep layer of gland-cells. The sinus-sac is small, but relatively well
developed and surrounds the base of the genital atrium. Just within the wall of the
sinus-sac the male and female ducts unite, and the narrow hermaphroditic duct
passes through the middle of the conical sinus-organ, which lies within the genital
atrium. The sinus-organ measures o-o8-o-i4 mm x 0-05-0-08 mm. The genital
pore occurs mid-ventrally, just posteriorly to the oral sucker.

The ovary is smaller than the testes and lies close behind them. The oviduct leaves
the ovary anteriorly and Mehlis' gland is situated anteriorly (or in contracted speci-
mens antero-laterally) to the ovary. Laurer's canal opens dorsally at the level of
the posterior margin of the ovary. The uterus, which forms a uterine seminal
receptacle proximally, occupies the bulk of the hindbody, with individual loops
reaching into the forebody. The metraterm extends from the ventral sucker to the
sinus-sac, and is surrounded by gland-cells. A fairly large 'vitelline reservoir' lies
ventrally to Mehlis' gland : this is probably the reduced half of the usual two-sided
vitellarium, as suggested by Robinson (1934). The remainder of the vitellarium is
a ramifying tubular system, occupying much of the dorsal region of the worm's
forebody and feeding a single duct passing into Mehlis' gland.

DISCUSSION. This worm appears to be virtually identical to A. nigroflavum. The
only significant difference between these two species seems to be the nature of the
body-surface at the posterior extremity. Although some doubt must be expressed
as to the importance of this feature, it has been described by both Robinson and
Thulin and has not been reported in specimens described under the appellation A.
nigroflavum or its synonyms.

Robinson (1934) states that in this species one half of the vitellarium is reduced
to a small single follicle lying ventrally to Mehlis' gland, whilst the bulk of the
vitellarium, situated in the forebody of the worm, represents the other half of the
normally more symmetrical two-sided system. It is difficult to be absolutely
certain, from the specimens of this species which we have available, whether
Robinson has interpreted this system correctly ; but it does not appear to differ from
that of A. nigroflavum. In a flattened specimen of A. petasiporum, however, this
'one-sided' asymmetrical system can be plainly seen (Fig. 8), and it seems almost
certain that this is a characteristic not only of the genus, but also of the sub-
family.

The structures discussed above, which were used by Robinson to distinguish this
species, have been used by Skrjabin (1959) to erect a new genus Guschanskiana. He
also distinguishes this genus as having a sinus-sac. From the descriptions given
above and from the literature, it would appear that the presence of a sinus-sac is a
feature common to all species of Accacladocoelium, We consider, therefore, in view

72 R. A. BRAY AND D. I. GIBSON

of this and in the light of the results discussed above, that there is no valid reason
for maintaining Guschanskiana as a distinct genus.

Accacladocoelium macrocotyle (Diesing, 1858) Robinson, 1934

(Fig- 7)

Distoma sp. of Bellingham, 1844.
Distomum macrocotyle Diesing, 1858.
Podocotyle macrocotyle (Diesing) Stossich, 1898.
Accacoelium macrocotyle (Diesing) Liihe, 1901.

TYPE-HOST AND LOCALITY. Mola mola, Ireland.

RECORDS.
(i) Material studied.

Mola mola [intestine] Skagerrak, Sweden (58°N, n°E) (Dec. 1929). Natural
History Museum of Gothenburg Reg. No. 29-5153.

Mola mola [intestine] Aberdeen, Scotland. BM(NH) Reg. No. 1966.9.6.54.
(ii) From the literature.

Mola mola [intestine] Ireland (July, 1839). Bellingham (1844 : 429 ; as Distoma
sp.).

Mola mola [intestine] Kattegat at Varberg, Sweden. (Nov.) Olsson (1868 : 24).

Mola mola [intestine] Atlantic WSW of Great Britain (48°N, I9°W) (Sept. 1886).
Guiart (1938 : 40).

PREVIOUS DESCRIPTIONS. Monticelli (1893) ; Linton (1940) ; Timon-David &
Musso (1971).

DESCRIPTION. Numerous whole specimens, one bisected specimen and one sec-
tioned specimen were available for study. The measurements are summarized in
Table 5.

The body-surface may be partly rugate, but bears no plications or spines. The
oral sucker is globular, with a fairly long anterior process of the small pharynx
encroaching into its base. The pharynx leads into an elongate and very narrow
oesophagus, reaching to the level of the ventral sucker before it passes into the caeca,
which have an H -shaped configuration. They communicate with the excretory
vesicle close to the posterior end of the worm, forming a uroproct. The ventral
sucker is longitudinally extended, but does not appear to bear any flanges or similar
modifications.

The excretory pore is terminal, and leads to a short vesicle, which divides into dorsal
and ventral arms that run anteriorly to about the level of the ventral sucker before
passing laterally. They unite dorsally to the pharynx.

The two almost globular testes lie in tandem near the ventral surface of the worm,
closely posterior to the ventral sucker. A convoluted, tubular seminal vesicle lies
dorsally to the ventral sucker, and leads into a sinuous and tubular pars prostatica
with a well-developed external covering of gland-cells. A very small, but distinct,
sinus-organ lies inside a fairly deep genital atrium, and receives both the ejaculatory
duct and the metraterm, which join at its base. It appears to be capable of partly or

ACCACOELIIDAE FROM N.E. ATLANTIC

73

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R. A. BRAY AND D. I. GIBSON

completely invaginating itself so that it projects backward into the hermaphroditic
duct (Fig. 70). The sinus sac is also small and distinct. The genital pore is mid-
ventral, close behind the oral sucker.

The globular ovary lies just posteriorly to the testes. Mehlis' gland is directly
anterior to the ovary and Laurer's canal opens dorsally at the level of the anterior
margin of the ovary. The proximal part of the uterus constitutes a uterine seminal
receptacle whilst the remainder of the uterus occupies much of the hindbody and
contains numerous relatively large eggs. The vitelline duct bifurcates close to
Mehlis' gland. Numerous tubular branches of the mtellarium, which extend as far
as the pharynx, arise from the anterior branch, whilst the other branch passes

FIG. 7. Accacladocoelium macrocotyle : (a)
semi-diagrammatic lateral view (gut omitted).
Scale - 2 mm. (b) Semi-diagrammatic view
of terminal genitalia showing sinus-organ
projecting backward into hermaphroditic
duct.

ACCACOELIIDAE FROM N.E. ATLANTIC 75

posteriorly, being fed by a small number of very short tubules lying in the region
just posterior to the ovary. A compact 'vitelline reservoir' was not seen.

DISCUSSION. This species differs from other species of the genus in having a more
extensive vitellarium which reaches just posteriorly to the ovary. In this species
the reduced half of the vitellarium does not form a saccular reservoir, but is repre-
sented by a few very short tubules. This is similar to the condition in Tetrochetus
raynerii as described by Looss (1912) and below.

Accacladocoelium petasiporum Odhner, 1928
(Fig. 8)

Distoma calyptrocotyle Monticelli of Odhner, 1911.

TYPE-HOST AND LOCALITY. Mola mola, Mediterranean Sea.

RECORDS.

(i) Material studied.
Mola mola [intestine] Kattegat at Gothenburg, Sweden. Material lent by J.

Thulin.

(ii) From the literature.

Mola mola [intestine] Kattegat at Gothenburg, Sweden (Nov. 1971). Thulin
(1973 : 83).

DESCRIPTION. This description is based upon two well-flattened and whole-
mounted specimens, one of which has been cut in two. Description from flattened
specimens is not to be recommended ; but, as it appears that this species has never
been adequately described, it seems warranted in this case. The morphology of the
specimens has, no doubt, been altered by the flattening process, and this should be
taken into account in considering this description. See Table 6 for the dimensions
of this worm.

TABLE 6

Dimensions of Accacladocoelium petasiporum from Mola mola,
Kattegat, Sweden

Thulin, 1973 Present Study

Length, mm 10-4 8-0 -10-2

Max. width, mm 1-2 0-94- 1-2

Oral sucker, mm °'45~ °'47 x °'45~°'52

Ventral sucker, mm 0-70- 0-84 x 0-89-1-04

Pharynx, mm o-io- 0-12x0-12-0-16

Testes, mm 0-62- 0-76x0-76-1-0

Ovary, mm 0-40- 0-43x0-41-0-54

Eggs, fan. 35 x 18 32-39x19-21

The body-surface of this elongate worm is smooth. The oral sucker is rounded and
ventro-terminal. The ventral sucker is larger than the oral sucker and is situated
on a distinct, but short, peduncle in the anterior half of the body. This sucker is

10

76 R. A. BRAY AND D. I. GIBSON

clearly modified in that it possesses wing-like lateral muscular extensions or flanges
(Fig. 8).

The morphology of much of the alimentary system could not be made out in these
preparations ; but the pharynx with an anterior extension into the base of the oral
sucker and the caeca with the anterior arms, both so characteristic of the accaco-
eliines, are present. A uroproct may be present, as the caeca extend close to the
excretory pore. Similarly, much of the excretory system could not be seen in these
preparations. All that can be distinguished is that the excretory pore is terminal,
and that the stem of the Y-shaped vesicle is short.

The large oval testes are in tandem in the anterior half of the hindbody. The
seminal vesicle is narrow, convoluted and very long, reaching back in one specimen
to the anterior testis, and nearly so in the other (Fig. 8). The tubular pars prostatica
is enveloped by a wide region of gland-cells and extends forward sinuously from about
the middle of the forebody to the junction with the hermaphroditic duct. Although
the details of the musculature of the sinus-sac and sinus-organ are not visible, a
definite, if short, sinus-organ is visible surrounding the distal part of the herma-
phroditic duct. The proximal region of this duct is dilated in one specimen, and
appears to contain an accumulation of sperm. The sinus-organ is present within a
small genital atrium, and the genital pore occurs mid-ventrally, close to the posterior
margin of the oral sucker.

FIG. 8. Accacladocoelium petasiporum : flattened specimen (gut omitted).

Scale - i mm.

ACCACOELIIDAE FROM N.E. ATLANTIC 77

The globular ovary lies posteriorly to the testes, and Mehlis' gland is present be-
tween the ovary and the testes. Laurer's canal is present, and can be seen in one
specimen to open to the exterior at about the level of the hind margin of the posterior
testis. The uterus leads posteriorly from the ovary almost to the posterior extremity
of the worm, where it turns and runs forward directly past the gonads. It then
follows a convoluted course up into the anterior region of the forebody, returns to
the pre-testicular region of the hindbody and finally passes anteriorly again as a
long, narrow and relatively straight metraterm (Fig. 8), uniting with the pars prosta-
tica near the base of the sinus-organ. The eggs are rather larger than occur in most
species of this genus. The vitellarium consists of a large ramifying system of tubules
lying in the forebody, and which are connected by a single, long collecting duct to
Mehlis' gland. The other half of the vitellarium is reduced to a small oval sac
(? reservoir), which measures o-i x 0-08 mm and is connected to Mehlis' gland by a
short duct.

DISCUSSION. This species was named by Odhner (1928) for certain specimens
which he found amongst Monticelli's and Stossich's material from the Mediterranean
and labelled Distoma macrocotyle. Earlier Odhner (1911), because of the presence of
a 'collar' on the ventral sucker, had considered these specimens to be the adults of
Distoma calyptrocotyle (now Odhnerium calyptrocotyle) , the immature forms of which
had been described by Monticelli (1893) from Beroe ovata. In 1928, however,
Odhner corrected this error, noting that the 'collar' of the ventral sucker is smaller
than that in D. calyptrocotyle. Nevertheless, A. petasiporum is not clearly recog-
nizable from Odhner's comments. Dollfus (1935), however, managed to obtain
some of Odhner's specimens and figured the external details of a specimen of about
3 mm in length. He also pointed out that this species possesses a copulatory organ
(sinus-organ), which is lacking in Odhnerium. Our best clues as to the morphology
of A. petasiporum come from Dollfus' key which states : 'L'acetabulum presente
des expansions musculaires. Organe copulateur present. Branches de 1'intestin
brievement ramifiees. Vitellogenes plus ou moins rassembles en petits cordons
sinueux entre le pedoncule acetabulaire et le pore genital.' He states that he hoped
to publish a comparative study of this species and 0. calyptrocotyle, but this does not
yet appear to have been carried out.

RHYNCHOPHARYNX Odhner, 1928

DIAGNOSTIC FEATURES. Forebody papillate. Ventral sucker pedunculate.
Pharynx consisting of two muscular bulbs and an extended anterior snout, which
may extend through the oral sucker. Oral sucker possesses posterior glandular
extension which envelops the snout. Glandular oesophageal bulb present im-
mediately posterior to pharynx. Oesophagus reaches to ventral sucker. Large
glandular organ of unknown function at base of peduncle. Pars prostatica long.
Sinus-sac surrounds base of genital atrium. Sinus-organ small. Vitellarium extends
from anterior region of ventral sucker to ovary. Parasitic in intestine of marine
teleosts (Mola).

TYPE-SPECIES. Rhynchopharynx paradoxa Odhner, 1928 (by original designation).

78 R. A. BRAY AND D. I. GIBSON

Rhynchopharynx paradoxa Odhner, 1928
(Figs 9 and 10)

TYPE-HOST AND LOCALITY. Mola mola, Trieste.

RECORDS.
(i) Material studied.

Mola mola [intestine] Vaderoarna Islands, Sweden (59°N, n°E) (Sept. 1941).
Natural History Museum of Gothenburg Reg. No. 41-7819.

Mola mola [intestine] Havstenssund, Sweden (59°N, n°E) (Aug. 1941). Natural
History Museum of Gothenburg Reg. No. 41-7805.

(?) Mola mola [intestine] Locality unknown. Donor : Zool. Museum, Amster-
dam. BM(NH) Reg. No. 1946.12.20.42.

FIG. 9. Rhynchopharynx paradoxa : lateral view (most of gut omitted).

Scale - i mm.

ACCACOELIIDAE FROM N.E. ATLANTIC

79

FIG. 10. Rhynchopharynx paradoxa : detail of forebody. Scale -i mm. c., caecum ; g.o.,
glandular organ ; o.b., oesophageal bulb ; p.p., pharynx proper ; r., 'Riissel' ; r.b.
'Riisselblase' ; r.s., 'Riisselscheide'.

(ii) From the literature.
None.

PREVIOUS DESCRIPTIONS. Odhner (1928) ; Yamaguti (1934).

DESCRIPTION. These worms (four whole-mounts and one sectioned) are very
elongate. The ventral sucker is situated close to the anterior end of the body on an
extensible peduncle, which may reach 3 mm in length (Fig. 9) . The body surface
may be rugate, and bears papillae on the forebody, but lacks spines or plications.
The aperture of the ventral sucker appears to be extended antero-posteriorly. The
oral sucker is roughly globular with a glandular posterior extension in the form of a
tube : the glands being enclosed in a thin muscular sheath, and similar gland-cells
may be seen in the musculature of the oral sucker (Fig. 10). The glandular tube
surrounds the anterior process, the 'Riissel' or snout, of the pharynx and is known
as the 'Riisselscheide' or snout-sheath. This snout portion of the pharynx is
muscular and elongate, up to about 1-5 mm in length by 0-03 mm at its distal end
and 0-09 mm near its proximal end, and it may extend as an elongate papilla into
the lumen of the oral sucker. It is recurved within the anterior region of the body
and widens out to form an anterior, muscular bulb, the 'Riisselblase' or snout-bladder,
of the pharynx, which measures 0-28-0-45 x 0-42-0-48 mm. A short, narrow,

8o R. A. BRAY AND D. I. GIBSON

TABLE 7

Dimensions of Rhynchopharynx paradoxa from Mola mola as recorded by various authors
Odhner, 1928 Yamaguti, 1934 Present Study

Locality Trieste Pacific Ocean see text

Length, mm 20-31-5 36 2O~35

Max. width, mm 1-5 -2-0 1-7 -2-5

Oral sucker, mm 0-48-0-57 x 0-34-0-45 0-37x0-32 0-38-0-45 x 0-46-0-52

Ventral sucker, mm 1-25-1-4 1-26x1-0 1-04-1-48 x 1-10-1-48

Pharynx, mm 0-50-0-65 0-37 0-46-0-56 x 0-56-0-78

Testes, mm 1-57x0-77 1-34-2-05 x 0-72-1-62

Ovary, mm 0-95x0-74 1-17-1-32 x 1-19-1-32

Eggs, /*m 27-30x19 29-32x18-21 32-36x21-25

muscular tube connects the latter region to a muscular posterior bulb, the pharynx-
proper, the measurements of which are given in Table 7. This leads into a small,
globular oesophageal bulb, which is a complex mixture of muscular and glandular
tissue of about 0-26 mm in diameter (Fig. 10). This in turn leads into a long and
very narrow (1-5 x 0-03 mm) oesophagus, which is surrounded by gland-cells. The
oesophagus reaches to a level just posteriorly to the peduncle of the ventral sucker,
where it bifurcates to form two gut-caeca which have blind anteriorly-directed
branches, in addition to posteriorly-directed branches that fuse terminally with the
excretory vesicle to form a uroproct.

There is a pair of large glandular organs at the base of the peduncle of the ventral
sucker, the function of which is not known. Each consists of numerous, tightly
packed gland-cells, arranged in irregular and branching lamina, and which stain
darker than the prostatic cells with H. and E.

The excretory pore is terminal and leads into a Y-shaped excretory vesicle with
ventral and dorsal arms. The hindbody is so packed with eggs that, even in serial
sections, it is not possible to trace the course of these arms. In the forebody,
however, they can be seen to be muscular and tubular, and pass laterally at the level
of the pars prostatica, joining dorsally to the pharyngeal complex.

The very large, oval or irregularly-oval testes appear in dorso-ventral view to be in
tandem, but the anterior testis lies nearer to the dorsal surface than the posterior
testis. They occur in the anterior part of the hindbody. The seminal vesicle is a
relatively narrow, convoluted tube lying at about the level of the base of the peduncle
of the ventral sucker. It passes forward to join a long, gently undulating pars
prostatica which is surrounded by numerous closely-packed gland-cells. The pars
prostatica extends forward to the base of a relatively small oval sinus-sac, which has
a very diffuse wall and measures 0-26 mm in length. There is a short ejaculatory
duct within the sinus-sac, which joins with the metraterm to form an hermaphroditic
duct. The latter runs through the sinus-sac and opens at the apex of a short sinus-
organ, measuring about 0-08 mm in serial section. It lies within a shallow genital
atrium, which opens to the surface through the genital pore situated mid-ventrally
at the level of the posterior margin of the oral sucker.

ACCACOELIIDAE FROM N.E. ATLANTIC 81

The sub-globular ovary is situated posteriorly to the testes, being separated from
the posterior testis by coils of the uterus. Mehlis' gland lies nearly contiguous with
the anterior margin of the ovary, and Laurer's canal can be seen to pass anteriorly
and dorsally to the ovary. The great extent of the uterus tends to hamper sectioning,
and thus the remainder of Laurer's canal and the pore were not observed. The first
coils of the uterus form a uterine seminal receptacle situated anteriorly to Mehlis'
gland. The uterus then passes posteriorly, almost to the posterior extremity,
before looping forward and filling much of the hindbody. It passes into the f orebody,
and, in the form of a wide tube, reaches the base of the sinus-sac. It narrows as it
passes through the wall of the sinus-sac, after which the short metraterm unites
with the ejaculatory duct to form the hermaphroditic duct. Ventrally to Mehlis'
gland there lies a fairly large, subglobular 'vitelline reservoir' of about o-i mm in
diameter. It is connected by a short duct to the oviduct. The remainder of the
vitellarium consists of numerous, complex, ramifying tubules, which reach from the
level of the peduncle of the ventral sucker to the anterior margin of the ovary, and
which feed a single collecting duct.

DISCUSSION. Although Rhynchopharynx has been recorded from the Mediter-
ranean Sea and from off Japan, it has not, hitherto, been recorded from the north-
east Atlantic. Rhynchopharynx is a monospecific genus, which is unmistakable,
owing to the complicated and unusual modifications of the pharynx.

ODHNERIUM Yamaguti, 1934

Mneiodhneria Dollfus, 1935.

Caballeriana Skrjabin & Guschanskaja, 1959.

DIAGNOSTIC FEATURES. Body-surface smooth, but with muscular sucker-like
pads on antero-dorsal surface. Flange-like muscular extensions on ventral sucker ;
ventral sucker pedunculate. Oesophagus reaches to ventral sucker. Pars prostatica
reaches half-way back to ventral sucker. Diffuse muscular region surrounds distal
part of metraterm and genital atrium (may be vestige of sinus-sac). Male duct enters
genital atrium from side through small papilla. Vitellarium tubular, extending from
pharynx to ovary. Parasitic in intestine of marine teleosts (Mola).

TYPE-SPECIES. Odhnerium calyptrocotyle (Monticelli, 1893) (by original desig-
nation).

Odhnerium calyptrocotyle (Monticelli, 1893) Yamaguti, 1934

(Figs ii and 12)

Distoma nigroflavum Rudolph! of Olsson, 1868.
Distoma calyptrocotyle Monticelli, 1893.
Orophocotyle calyptrocotyle (Montic.) Looss, 1902.
Accacoelium calyptrocotyle (Montic.) Stiles & Hassall, 1908.
Mneiodhneria calyptrocotyle (Montic.) Dollfus, 1935.
Distoma foliatum Linton, 1898.
Orophocotyle foliatum (Linton) Looss, 1902.

82 R. A. BRAY AND D. I. GIBSON

Accacoelium foliatum (Linton) Stafford, 1904.
Mneiodhneria foliata (Linton) Dollfus, 1935.
Caballeriana lagodovsky Skrjabin & Guschanskaja, 1959.

TYPE-HOST AND LOCALITY. Beroe ovata, Naples, Italy.

RECORDS.

(i) Material studied.
Mola mola [intestine] Skagerrak, Sweden (58°N, n°E) (Dec. 1929). Natural

History Museum of Gothenburg Reg. No. 29-5153.
Mola mola [intestine] Hunnebostrand, Skagerrak, Sweden (58°N, n°E).

Natural History Museum of Gothenburg Reg. No. 36-4970.
Mola mola [intestine] Havstenssund, Skagerrak, Sweden (59°N, n°E) (Aug.

1941). Natural History Museum of Gothenburg Reg. No. 41-7805.
Mola mola [intestine] Toras Ungar Islands (between Bonden and Hallo Islands),

Skagerrak, Sweden (58°N, n°E) (Sept. 1941). Natural History Museum of

Gothenburg Reg. No. 41-7813.
Mola mola [intestine] Rockall, North East Atlantic. BM(NH) Reg. No.

1964.7.16.21-35.
Mola mola [?] Locality unknown. Donor : Zool. Mus. Amsterdam. BM(NH)

Reg. No. 1946.12.20.27-41.
(ii) From the literature.
Mola mola [intestine] Varberg, Kattegat, Sweden (Nov.). Olsson (1868 : 25 ;

as Distoma nigroflavum).
Mola mola [intestine] Atlantic SW of Ireland (48°N, 09°W) (July 1921) ; Roscoff,

English Channel (Aug. 1921) ; South of Belle Isle, Bay of Biscay (Aug. 1922).

Dollfus (1935 : 209 footnote).
Mola mola [intestine] Atlantic WSW of Great Britain (48°N, ig°W) (Sept. 1886).

Guiart (1938 : 38 ; as Mneiodhneria foliata}.

PREVIOUS DESCRIPTIONS. Monticelli (1893) ; Linton (1898, 1940) ; Yamaguti
(1934) ; Lloyd (1938) ; Timon-David & Musso (1971).

LIFE-HISTORY. The life-history is not completely known ; but this species was
originally described from the ctenophore Beroe ovata, presumably as a metacercaria.

DESCRIPTION. Numerous whole-mounts and one serially sectioned specimen of
these elongate worms have been studied. See Table 8 for the dimensions of these
worms. The body-surface may be smooth or rugate, depending on contraction. A
peculiar feature is the presence of a series of sucker-like muscular pads on the antero-
dorsal region of the body-surface (Figs n and 12). The most anterior pads are the
largest, and they decrease in size posteriorly, reaching about half-way down the fore-
body. The ventral sucker is very large and protrudes from the ventral surface of the
body, sometimes apparently on a short peduncle. Two muscular flanges arise from
the anterior margin and pass down each side of this sucker (Figs n and 12). These
flanges are fused with the sucker dorsally and extend about three-quarters of the
sucker-length.

The pharynx extends slightly into the base of the oral sucker. It is muscular and
longer than wide, and leads, via a short muscular extension, into an elongate narrow

ACCACOELIIDAE FROM N.E. ATLANTIC 83

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FIG. ii. Odhnevium calyptrocotyle : lateral
view (gut omitted). Scale - i mm.

oesophagus, which reaches to the level of the base of the ventral sucker. It divides
into two caeca, which extend anteriorly to the level of the pharynx and posteriorly
almost to the posterior extremity, where they unite with the excretory vesicle via
narrow ducts to form a uroproct.

The excretory pore is terminal and leads into a fairly capacious excretory vesicle.
The uterus is so greatly developed in the hindbody that the course of the arms of the
excretory vesicle have not been traced, but a convoluted duct can be seen lying
dorsally to the pharynx.

The testes lie in the anterior half of the hindbody, with the anterior testis lying
close to the dorsal surface and the posterior testis, which is separated from it by a few
coils of the uterus, lying near the ventral surface. The seminal vesicle is a convoluted
tubular structure, lying at about the level of the ventral sucker, or slightly further

ACCACOELIIDAE FROM N.E. ATLANTIC

forward. It leads into a tubular sinuous pars prostatica, which is lined with cells
and surrounded by a thick covering of gland-cells. Although the pars prostatica is
fairly long and tubular, it is relatively short when compared with that of most other
accacoeliines. The sinus-sac, if it may be so termed, is modified to form a narrow,
fairly weakly-developed, muscular sheath which surrounds the genital atrium and
the distal part of the metraterm. The male duct passes through the dorsal wall of
this structure and opens at the base of the genital atrium, often through a very small
papilla (Fig. 12). There is no hermaphroditic duct or sinus-organ. It may, how-
ever, be possible that what we term the genital atrium is in fact an hermaphroditic
duct. If it is a genital atrium, however, the male papilla may be a derivation of the
sinus-organ.

The ovary lies directly posteriorly to the hind testis, with few coils of the uterus
separating them. A large Mehlis' gland lies immediately anterior to the ovary.
Laurer's canal passes over the antero-dorsal surface of the ovary and opens dorsally
at about the level of the posterior margin of the ovary. The proximal coils of the
uterus are distended with spermatozoa, forming a uterine seminal receptacle which
reaches as far forward as the anterior margin of the posterior testis. The remainder
of the uterus occupies the majority of the hindbody and passes anteriorly to about

FIG. 12. Odhnerium calyptrocotyle : longitudinal section of forebody. Scale - i mm. c.
caecum ; e.v., excretory vesicle ; m., metraterm ; m.p., male papilla ; o., oesophagus
p., pharynx ; p.p., pars prostatica ; s.p., sucker-like muscular pads ; s.s., sinus-sac
s.v., seminal vesicle ; u., uterus ; v., vitellarium.

11*

86 R. A. BRAY AND D. I. GIBSON

the level of the pharynx. It contains numerous small eggs which vary considerably
in size. The narrow muscular metraterm originates at about the level of the ventral
sucker, and passes anteriorly close to the ventral surface of the body and leads
directly into the genital atrium. As stated above, it is surrounded distally by a
muscular sheath, which appears to be homologous with the sinus-sac of other
accacoeliids. The vitellarium consists of numerous, ramifying tubules present on
both sides of the body, which extend from the level of the pharynx to the ovary.
The reduced left side of the vitelline system forms a 'reservoir' which lies ventral to
Mehlis' gland.

DISCUSSION. Odhner (1928), after studying some of Linton's specimens of
Distoma foliatum from Mola mola, considered them to be the mature form of D.
calyptrocotyle, which up to that time had only been described from the ctenophore
Beroe ovata. Linton's (1898) description is fairly meagre, but no essential difference
between it and D. calyptrocotyle has been found. The description of Odhnerium
calyptrocotyle by Lloyd (1938) from the Pacific coast of the USA was used by Skrjabin
& Guschanskaja (1959) to erect a new genus and species, Caballeriana lagodovsky.
The new genus was separated from Odhnerium by having 'ventral sucker pedunculate,
without appendages'. Dollfus (i96oa), after studying these characters, concluded
that there is no basis to them. Briefly, Skrjabin & Guschanskaja's differences can
be dismissed thus : (i) the ventral sucker often appears to be surmounted on a short
peduncle, but sometimes this is not immediately evident [indeed, when the peduncle
is as short as it is in Lloyd's figure, it could easily be reduced to nothing by a small
amount of contraction, and Monticelli's immature specimens, as figured by him,
could quite conceivably be interpreted as possessing a short peduncle] ; and (2) the
appendages on the ventral sucker of 0. calyptrocotyle are clearly shown in Lloyd's

fig, 35-

In some respects this genus falls in between Dollfus' (1935) conception of the
Accacoeliinae and the Tetrochetinae, which depends essentially only upon the pres-
ence or absence of a sinus-organ. In Odhnerium, whilst the sinus-organ is absent,
the sinus-sac is present, but in a greatly reduced form. Similarly the pars prostatica
is shorter than that generally found in the Accacoeliinae and longer than that of the
Tetrochetinae. As these two subfamilies are morphologically very similar and
apparently grade into one another, we consider them to be synonymous.

TETROCHETUS Looss, 1912

Paratetrochetus Hanson, 1955.

DIAGNOSTIC FEATURES. Body surface smooth. Ventral sucker pedunculate.
Oesophagus fairly long to long. Diverticula present at intestinal bifurcation. Pars
prostatica short, straight, narrow. Sinus-sac and sinus-organ absent. Male and
female ducts open together into shallow genital atrium. Vitellarium tubular, in
hindbody. Parasitic in intestine of marine teleosts with a diet of coelenterates,
ctenophores and chaetognaths as well as in carnivorous marine teleosts.

TYPE-SPECIES. Tetrochetus raynerii (Nardo, 1833) (by monotypy).

ACCACOELIIDAE FROM N.E. ATLANTIC 87

Tetrochetus raynerii (Nardo, 1833*) Looss, 1912, emend.

(Fig. 13)

Distoma raynerium Nardo, 1833.

Distomum raynerianum Nardo, 1833 of Diesing, 1850.

Accacoelium raynerianum (Nardo) Liihe, 1901.

Tetrochetus raynerianus (Nardo) of Dollfus, 1935, and Fukui & Ogata, 1935.

TYPE-HOST AND LOCALITY. Luvarus imperialis, Venice, Italy.

RECORDS.

(i) Material studied.
Luvarus imperialis [intestine] Vinnesvag, near Bergen, Norway (Sept. 1918).

University of Bergen Zoological Museum Reg. No. 43779.

Luvarus imperialis [intestine] Lepsoy, near Bergen, Norway (Aug. 1946). Univer-
sity of Bergen Zoological Museum No. 43780.
(ii) From the literature.
None.

PREVIOUS DESCRIPTIONS. Liihe (1901) ; Looss (1912).

LIFE-HISTORY. Metacercariae of this genus are known to occur in the siphono-
phore Physalia (Okada, 1932 ; Dollfus, 19602.) and probably also occur in chaetog-
naths (Dollfus et al., 1955 ; Dollfus, I96oa, b). Dollfus (19600.) records one free in
the plankton. As Luvarus feeds on medusae and similar gelatinous planktonic
animals, it appears that Tetrochetus raynerii is passed directly from its invertebrate
second intermediate host to its final host. Other species of Tetrochetus, on the other
hand, are found in predatory fishes and probably have a third intermediate stage in
their life-history, as they have also been recorded from pelagic fishes.

DESCRIPTION. Nine specimens of this worm were kindly lent to us by Professor
A. Brinkmann Jr, of the University of Bergen. Six specimens are flattened whole-
mounts, one is an unflattened whole-mount and two are serially sectioned. The
dimensions are given in Table 9.

These worms are large and elongate, with a prominent ventral sucker on a short
peduncle in the anterior quarter of the worm. The body-surface is smooth. The
globular oral sucker is invariably larger than the ventral sucker and opens sub-
terminally. A narrow anterior extension of the pharynx protrudes into the posterior
wall of the oral sucker. The pharynx widens posteriorly, and leads into a straight,
narrow oesophagus 0-62-2-16 mm in length, which is surrounded by a concentration
of gland-cells near its junction with the pharynx. At the intestinal bifurcation a
few small diverticula radiate. The overall configuration of the intestine is H -shaped,
and a uroproct is present. There are numerous gland-cells scattered in the paren-
chyma of the f orebody.

The stem of the excretory vesicle is short, dividing near the uroproct and giving rise
to dorsal and ventral arms. Wide convoluted tubules can be seen lying laterally in

* Contrary to Stiles & Hassall's (1908) catalogue we have been unable to find the use of the name
Distoma raynerianum in Nardo (1827), or (1828). We have emended the specific name, placing it in the
genitive case, as Nardo (1875) clearly states that he named the species after Prof. S. A. Renier.

R. A. BRAY AND D. I. GIBSON

TABLE 9

Dimensions of Tetrochetus raynerii from Luvarus imperialis as recorded

Locality
Length, mm
Max. width, mm
Oral sucker, mm

Ventral sucker, mm
Pharynx, mm
Testes, mm
Ovary, mm
Eggs, Mm

Liihe, 1901

Looss, 1912

Present Study

(flattened)

Trieste

Trieste

Norway

10

11-12

9-15

c. 1-5

1-1*3

1-6-3-5

c. 1-2 dia.

1-2-1-3 dia.

1-05-2-03 x

1-05-1-68

c. i-o dia.

0-9-1-0 dia.

0-67-1-15 x

0-33-1-08

-

0-42 (plus ant.

0-61-1-02 x

ext.) x 0-33

0-23-0-43

—

—

0-52-1-43 x

0-35-1-24

—

-

0-41-1-02 x

0-30-0-54

34X17

30-35x18

32-34x19-23

by various authors

Present Study
(unflattened)

Norway

1-76

0-94 x 0-84

0-73x0-51
0-61 x 0-34

0-56-0-74 x
0-48-0-52
0-52 x 0-48

32 x 19

the forebody, and they are prominent in the area ventral to the pharynx. A trans-
verse tubule is visible dorsally to the oral sucker, and it seems probable that this is
the union of the excretory arms (see Looss, 1912) ; but the complete system in this
region has not been traced.

The testes are relatively small, subglobular to oval, and lie in the anterior half of
the hindbody, obliquely in lateral view, with the anterior testis near to the dorsal
surface and the posterior near to the ventral surface. The long, narrow, tubular
seminal vesicle reaches almost to the anterior testis. It lies, for most of its length,
near to the dorsal surface of the worm ; but about half-way along the forebody it
passes ventrally and runs into a straight pars prostatica, 073-1-35 mm in length.
The pars prostatica, which is surrounded by a thin layer of gland-cells, enters the
base of a small, narrow, genital atrium. The genital pore is situated ventrally to the
oral sucker. There is no sign of a sinus-organ, sinus-sac or hermaphroditic duct.

The reniform ovary lies posteriorly to the testes, near to the ventral margin of the
body. In the concavity of the anterior surface of the ovary lies a globular Mehlis'
gland, 0-20-0-35 mm in diameter. Laurer's canal runs in a rather straight course
from Mehlis' gland to open dorsally at the level of the ovary. The initial coils of the
uterus constitute a uterine seminal receptacle. The uterus, which contains numerous
small eggs, is rather less extensive than is usually the case in accacoeliines. First
of all, it passes anteriorly as far as the level of the anterior margin of the ventral
sucker. It then turns and runs to a short distance from the posterior extremity,
where it turns again and passes anteriorly, gradually becoming wider. Finally, the
uterus narrows to form a metraterm at about the level of the intestinal bifurcation.
The latter duct is narrow, and runs in a straight course to the base of the genital
atrium, which it enters ventrally to the pars prostatica. The vitellarium consists of
numerous narrow, branching tubules extending from about the level of the ventral

ACCACOELIIDAE FROM N.E. ATLANTIC 89

sucker to the ovary. It feeds a single collecting duct. The other side of the vitel-
larium is reduced to a short blind tubule, which may bear several very short
branches.

DISCUSSION. This species is found exclusively in Luvarus imperialis [= Ausonia
cuvieri, = Proctostegus prototypus] and has previously been found only in the

FIG. 13. Tetrochetus raynerii : flattened
specimen (most of gut omitted) . Scale -
I mm.

90 R. A. BRAY AND D. I. GIBSON

Mediterranean basin, i.e. in the Adriatic Sea (Nardo, 1833 ; Lime, 1901 ; Looss,
1912) and in the Tyrrhenian Sea (Parona, 1902). This fish is rather rare, not only in
the north-east Atlantic, but throughout its almost world- wide range (Topp, 1973).

Of the other species of Tetrochetus, the one most likely to be found in our area in
the future is probably T. coryphaenae Yamaguti, 1934, which occurs in the dolphin-
fish Coryphaena elsewhere in the Atlantic Ocean. This species, which has recently
been described in detail from the south Atlantic by Gibson (1976), differs from T.
raynerii in having a ventral sucker distinctly larger than the oral sucker, and in
being a considerably smaller and narrower worm.

Subfamily PARACGAGLADIINAE subfam. nov.

DEFINITION. Accacoeliidae. Body slender. Ventral sucker borne on small
peduncle. Body-surface without spines or plications ; papillae on outer surface of
ventral sucker. Oral sucker with internal ridge. Prepharynx absent. Pharynx
protruding into base of oral sucker ; otherwise unmodified. Oesophagus short,
wide. Caeca with small 'shoulders', but without anterior extensions, terminate
blindly near posterior extremity. Testes symmetrical to tandem, in anterior half
of hindbody. Seminal vesicle tubular, convoluted ; dorsal to ventral sucker and in
forebody. Pars prostatica elongate, convoluted. Ejaculatory duct unites with
metraterm within sinus-sac. Sinus-sac surrounding base of genital atrium ; with
diffuse musculature. Sinus-organ short ; cylindrical. Hermaphroditic duct within
sinus-organ. Genital atrium relatively large ; contains sinus-organ. Genital pore
mid-ventral in anterior forebody. Ovary globular, post-testicular. Mehlis' gland
post-ovarian, linked to posterior region of ovary by oviduct. Laurer's canal with
dorsal pore. Uterine seminal receptacle present. Uterus extensive in hindbody.
Eggs fairly small ; numerous. Vitellarium with branching tubules ; symmetrical
with two main lateral collecting ducts ; in region mainly lateral and posterior to
ovary. Excretory vesicle Y-shaped ; with arms situated dorsally and ventrally in
hindbody and laterally in forebody, uniting dorsally to pharynx. Immature forms
in rectum of teleosts with diet of coelenterates, ctenophores and chaetognaths,
adults in rectum of carnivorous teleosts (Coryphaenoides).

TYPE AND ONLY GENUS. Pamccadadium gen. nov.

PARACCACLADIUM gen. nov.

DEFINITION. As subfamily.

TYPE SPECIES. Paraccadadium jamiesoni sp. nov.

DISCUSSION. The above detailed definition of the subfamily will serve, for the
present time, to define the genus, although both will, no doubt, need to be redefined
if further new genera are discovered. Paraccadadium appears to be essentially an
accacoeliid, but differs in the structure of its alimentary system, the structure of the
vitellarium and the position of Mehlis' gland, from all of the genera included in the
Accacoeliinae. The accacoeliines have a long, narrow, oesophagus, an H-shaped gut,

ACCACOELIIDAE FROM N.E. ATLANTIC 91

a uroproct, a one-sided vitellarium (i.e. with one main collecting duct plus one which
is usually rudimentary), and Mehlis' gland lying anteriorly, antero-laterally or antero-
dorsally to the ovary. Paraccacladium has a short, wide oesophagus, only small
caecal 'shoulders', no uroproct, a symmetrical vitellarium with two main collecting
ducts, and Mehlis' gland posterior to the ovary (Fig. i). Its pharynx, however, is
typically accacoeliid, as it protrudes into the base of the oral sucker, and the presence
of immature forms in teleosts which feed on coelenterates suggests that the meta-
cercariae occur in the same invertebrate groups as the accacoeliines.

Paraccacladium jamiesoni sp. nov.
(Figs 14 and I5a)

TYPE-HOST AND LOCALITY. Coryphaenoides rupestris, 55°N, n°W.

RECORDS.

(i) Material studied.

Coryphaenoides rupestris [rectum] Off west coast of Great Britain :
5o°N, I2°W ; depth 965-980 m ; April 1973 (2 specimens).
5i°N, I4°W ; depth 920-960 m ; April 1973 (i specimen).
55°N, n°W ; depth 800-820 m ; April 1973 (5 specimens).
56°N, 09°W ; depth 780-790 m ; April 1973 (i specimen).
57°N, 09°W ; depth 560-572 m ; April 1973 (4 specimens).
58°N, io°W ; depth 780-800 m ; April 1973 (4 specimens).
59°N, 09°W ; depth 720-860 m ; April 1973 (3 specimens).
59°N, o8°W ; depth 570-700 m ; June 1974 (i specimen).
59°N, o8°W ; depth 880-900 m ; June 1974 (3 specimens).
54°N, I4°W ; depth 1050 m ; June 1974 (7 specimens).
56°N, I7°W ; depth 640 m ; June 1974 (3 specimens).
59°N, io°W ; depth 1000 m ; June 1974 (2 specimens).
Holotype : BM(NH) Reg. No. 1975.10.31.1.
Paratypes : BM(NH) Reg. No. 1975.10.31.2-18.
(ii) From the literature.
None.

LIFE-HISTORY. Immature forms of this genus have been found in the rectum of
Alepocephalus bairdii, Schedophilus medusophagus, Centrolophus niger and Xenoder-
michthys copei (see below), all of which feed upon coelenterates and ctenophores,
especially medusae, the presumed second intermediate hosts of accacoeliids. These
fishes may act as paratenic hosts, or, more likely, they may be obligatory, par-
ticularly as the mature adult has only been found in the carnivorous macrourid
Coryphaenoides rupestris. It is worth noting that one infested specimen of Alepo-
cephalus bairdii was found with its stomach full of the bathypelagic scyphomedusa
Atolla wyvillei.

DESCRIPTION. Eleven mature specimens were present and all of the remainder were
immature, exhibiting varying degrees of immaturity. The dimensions of the main
features, including those f or Paraccacladium sp. (see below), are given in Table 10.

92 R. A. BRAY AND D. I. GIBSON

^araccacladium sp. immat.

tccacladium sp. immat.

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ACCACOELIIDAE FROM N.E. ATLANTIC

93

The worm is long, thin and cylindrical in the case of the mature adults (Fig. I4a),
but shorter and stouter in the immature forms (Fig. I5a). The body-surface is
smooth, except for numerous dome-shaped papillae, which are present on the outer
surface of the ventral sucker, and to a lesser degree on the forebody ; these latter
papillae are not always readily visible. The ventral sucker, which is larger than the
oral sucker, lies on a short peduncle just within the anterior third of the body, but
further posteriorly in immature forms. The oral sucker opens almost terminally,
and bears an internal ridge close to the base of the inside wall. This ridge surrounds
the dome produced by the extension of the pharynx into the base of the oral sucker.
The pyriform pharynx leads into a short wide oesophagus, which has a thick cuticular
lining. This extends back to about the level of the sinus-organ, where it bifurcates.
The two wide caeca, which are lined with a deeply staining epithelium, form small
'shoulders', but not long anteriorly-directed diverticula, and extend posteriorly,

a

FIG. 14. Paraccacladium jamiesoni : (a) lateral view. Scale - i mm. (b) Semi-diagram-
matic view of terminal genitalia. (c) Ventral view of gonads and vitellarium in an
immature specimen. Scale - 0-5 mm.

94 R- A. BRAY AND D. I. GIBSON

following a sinuous course, almost to the posterior extremity, where they end
blindly.

The exretory pore is situated at the posterior extremity and leads into a short
convoluted vesicle, which divides to form two anteriorly-directed arms, one ventral
and one dorsal. At about the level of the gonads the dorsal duct passes over towards
the ventral surface, and then both ducts pass forward ventro-laterally to the level of
the pharynx, where they unite dorsally.

The gonads lie in the anterior half of the hindbody, apparently moving further
from the ventral sucker as the worm develops, presumably as a result of allometric
growth. The smooth, oval testes are pre-ovarian, and, in mature specimens, usually
He in tandem, often with the anterior testis lying close to the dorsal surface. Im-
mature specimens show the testes in varying positions from tandem to symmetrical
(Fig. 15). The testes are larger than the ovary in immature specimens ; but it
appears that this species may be protandrous, as in the largest adults the ovary is
distinctly larger than the testes. In some very small specimens, although the testes
are well developed, no sign of an ovary could be seen. The seminal vesicle is tubular,
stout and sinuous, and it extends from about the mid-region of the ventral sucker to a
position just anterior to the ventral sucker (Fig. I4b). It leads into a relatively-long
tubular pars prostatica, which is surrounded by gland-cells and gently convoluted.
The ejaculatory duct unites with the metraterm at the base of the cylindrical
sinus-organ to form an hermaphroditic duct, which passes through the sinus-
organ and opens terminally. The base of the sinus-organ is enclosed in a diffusely
muscular sinus-sac, the walls of which are attached to the walls of the genital atrium
a little way from its base. The space within the sinus-sac contains numerous muscle-
fibres and gland-cells. The external wall of the sinus-organ appears to be a continu-
ation of the wall of the genital atrium, which in turn is similar to that of the external
tegument.

The oval ovary lies just posteriorly to the testes, near the ventral surface and close
to the middle of the body in the largest worms. Immediately behind the ovary is an
oval Mehlis' gland, which is rather smaller than the ovary in mature specimens.
The oviduct leads into this organ after leaving the ovary posteriorly and receiving
Laurer's canal and the common vitelline duct. Laurer's canal runs across the body
and opens to the exterior dorsally at the level of the ovary. A uterine seminal
receptacle is present and is the sole seminal storage apparatus in the female system.
The uterus runs posteriorly from the ovary to a point beyond the hindermost end of
the vitellarium, but well anteriorly to the posterior extremity of the worm. It then
passes anteriorly and winds in the region of the gonads, but then runs forward in a
relatively straight or slightly sinuous manner to join the hermaphroditic duct within
the sinus-sac in the form of a short metraterm. The uterus contains numerous
relatively small eggs, which are noticeably smaller at the proximal extremity of the
uterus. The vitellarium consists of a symmetrical branching tubular system,
possessing two main collecting ducts which pass medially and unite to form a common
duct at the level of Mehlis' gland (Fig. I4c). The tubular branches extend laterally
between about the level of the posterior testis and a position about half-way between
the ovary and the posterior extremity.

ACCACOELIIDAE FROM N.E. ATLANTIC

95

Paraccacladium sp. (immature)

(Fig. isb-e)
RECORDS.

(i) Material studied.
AUpocephalus bairdii [rectum] Off west coast of Great Britain :

5i°N, I4°W ; depth 920-960 m ; April 1973 (3 specimens).

55°N, I7°W ; depth 821-850 m ; June 1974 (5 specimens).

57°N, I3°W ; depth 1000-1060 m ; June 1974 (3 specimens).

BM(NH) Reg. No. 1975.10.31.19-21.
Xenodermichthys copei [rectum] Off north-west coast of Scotland :

59°N, n°W ; depth 970-1025 m ; June 1974 (i specimen).

BM(NH) Reg. No. 1975.10.31.22.
Schedophilus medusophagus [rectum] Off north-west coast of Scotland

58°N, io°W ; depth 780-800 m ; April 1973 (i specimen).

BM(NH) Reg. No. 1975.10.31.23.
Centrolophus niger [rectum] Off north-west coast of Scotland :

57°N, I3°W ; depth 800-820 m ; June 1974 (i specimen).

BM(NH) Reg. No. 1975.10.31.24.
(ii) From the literature.
None.

FIG. 15. Paraccacladium jamiesoni : (a) immature specimen from Coryphaenoides
rupestris. Paraccacladium sp. immature : (b) from Alepocephalus bairdii ; (c) from
Schedophilus medusophagus ; (d) from Xenodermichthys copei ; (e) from Centrolophus
niger. Scale - i mm.

12

96 R. A. BRAY AND D. I. GIBSON

DESCRIPTION. These specimens are morphologically similar to the immature forms
of P. jamiesoni. Their measurements are given in Table 10, and specimens from
each of the four hosts are illustrated in Fig. 150-6.

DISCUSSION. These immature specimens probably belong to P. jamiesoni, as
they do not differ significantly from the immature specimens of this species from
Coryphaenoides rupestris. A complete series from immature to fully mature was
found in the latter fish. The sinus-organ is clearly visible in all of the immature
specimens, so that they cannot be confused with Tetrochetus Looss, 1912, which also
appears to utilize teleosts as intermediate hosts (see Gibson, 1976).

Accacladocoelium leontjevae Korotaeva, 1976, appears to belong to this genus,
although some important details are omitted from the original description.
Korotaeva found it in the intestine of six diverse species of teleost off the south-
east coast of New Zealand. It possesses eggs which are considerably smaller than
those of P. jamiesoni, and apparently also differs in lacking an oesophagus, but
this would be a very unusual feature and needs confirmation. We tentatively con-
sider it, as Paraccacladium leontjevae (Korotaeva, 1976) n. comb., to represent the
second species of this new genus.

The differences between Paraccacladium and the accacoeliines are discussed above.
It may, however, be worth drawing attention to the similarity between this species
and Sclerodistomoides pacificus Kamegai, 1971, from the gall-bladder of Cypselurus
pinnatibarbatus japonicus from the Pacific coast of Japan. According to Kamegai
(1971), however, Sclerodistomoides is a much more robust worm and lacks the definite
accacoeliid type of pharynx possessed by Paraccacladium, and also apparently the
presence of dorsal and ventral excretory arms in the hindbody.

HOST-PARASITE LIST*

Order Salmoniformes

Alepocephalus bairdii Goode and Beane : Paraccacladium sp. (immature).
Xenodermichthys copei (Gill) : Paraccacladium sp. (immature).

Order Gadiformes
Coryphaenoides rupestris Gunnerus : Paraccacladium jamiesoni gen. et sp. nov.

Order Perciformes

Centrolophus niger (Gmelin) : Paraccacladium sp. (immature).
Luvarus imperialis Rafinesque : Tetrochetus raynerii (Nardo).
Schedophilus medusophagus (Cocco) : Paraccacladium sp. (immature).

Order Tetraodontiformes
Mola mola (L.) : Accacoelium contortum (Rud.).

Accacladium serpentulum Odhner.
Accacladocoelium nigroflavum (Rud.).
Accacladocoelium macrocotyle (Dies.).
Accacladocoelium petasiporum Odhner.

* We have consulted Greenwood et al. (1966) for the ordinal names in this checklist.

ACCACOELIIDAE FROM N.E. ATLANTIC 97

Accacladocoelium alveolatum Robinson.
Rhyncho pharynx paradoxa Odhner.
Odhnerium calyptrocotyle (Monticelli) .

ACKNOWLEDGEMENTS

The authors are indebted to the following : Professor A. Brinkmann Jr, University
of Bergen, for the loan of specimens of Tetrochetus raynerii ; Mr J. Thulin, Univer-
sity of Gothenburg, for the loan of his material, and for his efforts in obtaining
material from the Gothenburg Natural History Museum ; M.A.F.F. staff and crew
of R.V. Cirolana, especially Dr A. Jamieson, Mr R. J. Turner and Mr J. H. Nichols,
for their help to one of us (R. A. B.) on various cruises ; Mr D. W. Cooper and Mr
S. J. Moore, British Museum (Natural History), for the preparation of the serial
sections ; Miss M. A. Bray, for typing parts of the manuscript ; Mr S. Prudhoe,
British Museum (Natural History), for his continual advice and encouragement.

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R. A. BRAY
D. I. GIBSON

Department of Zoology

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THE PLANKTONIC COPEPODS OF
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OCEAN: SOME TAXONOMIC
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ONCAEIDAE (CYCLOPOIDA)

G. A. BOXSHALL

BULLETIN OF

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ONCAEIDAE (CYCLOPOIDA)

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Issued 28 April, 1977 Price £4.75

THE PLANKTONIC COPEPODS OF THE

NORTHEASTERN ATLANTIC OCEAN: SOME

TAXONOMIC OBSERVATIONS ON THE

ONCAEIDAE (CYCLOPOIDA)

By G. A. BOXSHALL

SYNOPSIS

A new genus belonging to the family Oncaeidae is described. It differs from all other genera
in the family by the possession of a 2-segmented endopod in legs 1-4 and a 2-segmented second
antenna. Descriptions of three new species and the hitherto undescribed males of two species
are also given. The other thirteen species found in the samples are briefly described and figured.

INTRODUCTION

ALL THE important families of planktonic cyclopoid copepods, except the Oncaeidae
have been the subject of a major revision ; the Corycaeidae by Dahl (1912), most of
the Sapphirinidae by Lehnhofer (1926 & 1929) and the Oithonidae by Rosendorn
(1917). It is not surprising, therefore, to find that it is primarily within the Oncaei-
dae, which contains numerous small and often difncult-to-separate species, that
taxonomic problems occur. Also many new taxa are being discovered which have
previously escaped detection either by their exceedingly small size or by the remote-
ness of their bathypelagic habitat. Thus, when a series of plankton hauls from the
Atlantic was analysed priority was given to taxonomic studies on the members of
the family Oncaeidae present in the samples. In the following account the new taxa
and rare species are described in detail and, as a basis for future comparative studies,
brief descriptions of the commoner species are given because modern descriptions of
these are often not available and older descriptions are incomplete.

MATERIALS AND METHODS

The material described below was first lent and then donated to the British Museum
(Natural History) by the Institute of Oceanographic Sciences at Wormley. It
consists of a day and night series of vertical hauls made with a horizontally towed,
opening and closing, net system which was acoustically controlled. The copepods
were caught in a rectangular midwater trawl (RMT i) of I m2 mouth area and a
mesh of 0-32 mm. The net system and method of fishing are described by Baker
et al. (1973). Twenty-eight hauls were made in November 1969 at station 7089
(i8°N, 25°W) in the Atlantic near the Cape Verde Islands. The depth of each haul
is given below together with the fraction of the sample examined.

The copepods were stained in chlorazol black in lactophenol, dissected and ex-
amined in lactophenol and mounted as permanent preparations in polyvinyl lacto-
phenol. The preparations were examined by phase contrast microscopy and
drawings were made with the aid of a camera lucida.

104 G. A. BOXSHALL

SAMPLE (HAUL) NUMBER DEPTH (m) FRACTION

35 10-0 1/8

33 10-0 1/64

31 20-10 1/32

28 25-10 1/64
30 50-20 1/32
27 60-25 J/32
26 100-49 1/64
20 100-55 I/32

23 200-110 1/16
19 194-112 1/16
25 290-210 1/16

3 300-210 1/16

11 400-300 1/8

29 40°-3°5 J/8

24 500-410 1/8
22 500-410 1/8
17 600-505 1/8

13 600-515 1/8
16 700-610 1/8

9 700-610 1/8

15 785-700 1/8

5 790-700 1/8

4 890-800 1/8

12 900-800 1/8
8 1010-900 1/8

14 1020-910 1/8

34 1220- 1000 1/8

32 1250-1000 1/8

DESCRIPTIONS OF SPECIES

PARALUBBOCKIA gen. nov.

DIAGNOSIS. Body elongate ; urosome 5-segmented in <j>, 6-segmented in <$.
First antenna 7-segmented in $, 5-segmented in £. Second antenna 2-segmented
with 4 setae. Mandible a curved process with 2 strong spines and a hirsute seta.
First maxilla bilobed ; outer lobe with 3 ($) or 4 (^) apical setae, inner lobe with
i ($} or 2 ($) lateral setae. Second maxilla ($) with elongate basal segment and
terminal segment bearing 5 elements ; (<£) squat, indistinctly 2-segmented with 2

subapical elements. Maxilliped prehensile, 4-segmented. Legs 1-4 with 3-seg-
mented exopods and 2-segmented endopods ; armature formula :

Coxa Basis Endopod Exopod

Leg i o-o i-o o - i ; o, I, 5 o - o ; o - i ; I, I, 4

Leg 2 o-o i-o o - i ; o, II, 5 I - o ; o - i ; I, I, 5

Leg 3 o-o i-o o - i ; o, II, 4 I - o ; o - i ; I, I, 5

Leg 4 o-o i-o o - i ; o, II, 2 I - o ; I - i ; I, I, 5

Leg 5 a free segment with 2 terminal elements.
TYPE-SPECIES. Paralubbockia longipedia sp. nov.

OBSERVATIONS ON THE ONCAEIDAE 105

ETYMOLOGY. The generic name refers to the close resemblance in general facies
between the new genus and Lubbockia Claus, 1863.

Paralubbockia longipedia gen. nov. et sp. nov.

DESCRIPTION. Female. Body elongate, pointed anteriorly (Fig. la) ; prosome
longer than urosome ; rostrum small and rounded ; relative lengths urosome
somites and caudal rami 15 : 33 : 15 : 12 : 14 : n. Caudal rami about 1-5 times
longer than broad.

First antenna (Fig. ic) extending posteriorly beyond margin of cephalosome ;
fourth segment providing about half the length of the appendage ; armature elements
on segment I - i, II - 3, III - 3, IV - 5, V - 3, VI - 2, VII - 6.

Second antenna (Fig. ib) 2-segmented; long proximal segment with a hirsute
seta on lateral margin ; distal segment with 3 hirsute setae increasing in size
towards the apex with 2 rows of fine setules on the lateral surface.

Mandible (Fig. id) a curved tapering process with 2 stout spines on the convex
margin, a proximal hirsute seta and a row of bristles.

First maxilla (Fig. le) comprising 2 unequal lobes ; the larger inner lobe with 2
setae, the small outer lobe with 3 hirsute setae around the apex.

Second maxilla (Fig. if) with an elongate basal segment and a small terminal
segment bearing 2 curved apical spines, both with rows of spinules on their concave
margins, and 3 hirsute setae on the inner margin.

Maxilliped (Fig. ig) 4-segmented with a long claw-like distal segment ; second
segment with a continuous row of spinules along inner margin and 2 naked setae ;
claw with a spinule row on proximal half of concave margin, spinules increasing in
size distally.

Legs 1-4 (Figs 2a-d) with 3-segmented exopods and 2-segmented endopods ;
armature formula as in generic diagnosis. Spines on both rami bilaterally serrate
except for apical spine of exopod which is externaUy serrate and internally plumose ;
rows of setules present on lateral margins of endopod segments and inner margin of
exopod segment i ; 5 long spinules located on inner margin of basis of leg 4. Left
member of leg 2 pair aberrant in holotype with 2 inner setae on exopod segment 2
and only 4 inner setae on exopod segment 3.

Leg 5 (Fig. ih) a free segment with 2 naked apical setae and a lateral seta located
on surface of somite near base.

Leg 6 probably represented by unarmed cuticularized plate located near genital
aperture.

Body lengths of 4 complete female specimens 1-43, 1-26, 1*19 and 1-37 mm.

Male. Similar to female in general appearance ; urosome (Fig. 3a) 6-segmented ;
posterior margins of urosome somites 3 to 5 provided with a continuous dentate
hyaline frill on ventral surface ; anal somite with a row of fine spinules posteriorly.
Caudal rami shorter than anal somite.

First antenna (Fig. 3b) 5-segmented with the distal segment representing the fused
fifth to seventh segments of the female ; fourth segment extremely long as in female ;
armature elements on segment I - i, II - 5, III - 2, IV - 4, V - 10.

io6

G. A. BOXSHALL

Second antenna (Fig. 30) as in female.

Mandible a slightly curved blunt process without armature elements but this may
possibly be due to damage during dissection.

First maxilla (Fig. 3d) similar in form to that of female but with 4 setae on outer
lobe and with only i seta on inner lobe.

FIG. i. Paralubbockia longipedia gen. et sp. nov. a, female entire, dorsal ; b, second
antenna, anterior ; c, first antenna, ventral ; d, mandible, posterior ; e, first maxilla,
posterior ; f, second maxilla, posterior ; g, maxilliped, anterior ; h, anterior of urosome,
ventral.

OBSERVATIONS ON THE ONCAEIDAE

107

FIG. 2. Paralubbockia longipedia gen. et sp. nov. a, leg i, posterior ; b, leg 3,
posterior ; c, leg 2, posterior ; d, leg 4, posterior.

io8

G. A. BOXSHALL

FIG. 3. Paralubbockia longipedia gen. et sp. nov. a, male urosome, ventral ; b, first
antenna, ventral ; c, second antenna, posterior ; d, first maxilla, posterior ; e, second
maxilla, posterior ; f, maxilliped, anterior.

Second maxilla (Fig. 30) a squat indistinctly 2-segmented process with 2 small
setae on the inner surface of the distal portion.

Maxilliped (Fig. 3f) as in female but less robust and without setae on second
segment ; spinules on second segment very fine ; terminal claw with proximal
spinule armed with minute spinules.

OBSERVATIONS ON THE ONCAEIDAE 109

Legs 1-4 as in female.

Leg 5 (Fig. 3a) a free segment with a lateral plumose seta near its base and with
2 apical setae ; inner apical seta extending posteriorly beyond margin of anal somite,
outer seta half length of inner.

Leg 6 probably represented by small cuticularized area with a single spinule,
located postero-laterally on genital somite.

Body length of single male specimen 1-19 mm.

MATERIAL EXAMINED. 6 $$ and i $ ; holotype $ from sample 32, i $ from samples
8, 9 and 22, and 2 $ and i $ from sample 16. BM(NH) registration numbers : holo-
type 1976.8, paratype ^ 1976.9 and paratype $$ 1976.10-13.

REMARKS. Placing Paralubbockia in any of the existing families of the Cyclopoida
poecilostoma raises certain problems because of the unusual structure of the second
antenna and the reduced segmentation of the thoracic legs. However, it has many
characters in common with other genera of the family Oncaeidae (general facies,
structure of the first antennae, mandibles, maxillae and maxillipeds) and its true
relationships appear to lie with this family. It is proposed to place Paralubbockia
in the Oncaeidae and to emend the familial diagnosis as follows :

ONCAEIDAE Giesbrecht, 1891 emend.

Body cyclopiform. Prosome 5-segmented, urosome 4- or 5-segmented in $ and
5- or 6-segmented in <£. Caudal ramus with 6 setae. Female apertures usually
dorsal, sometimes laterally displaced. First antenna 4- to 8-segmented, never
geniculate in males. Second antenna usually 3- or 4-segmented, sometimes 2-
segmented. Mandible usually with 4 armature elements, sometimes reduced to an
unarmed lash. First maxilla bilobed, each lobe bearing at least i seta. Second
maxilla 2-segmented with a number of terminal elements. Maxilliped 3- or 4-seg-
mented, with distal segment claw-like in both sexes. Legs 1-4 with 3 segmented
exopod and usually 3-segmented endopod, rarely 2-segmented.

In the reduced condition of the second antennae and thoracic legs Paralubbockia
exhibits advanced characters which are presumably derived from the typical oncaeid
condition. The affinities of the new genus appear to lie with the genus Lubbockia as
incomplete separation of the segments in the rami of the thoracic legs of Lubbockia
has been reported (Rose, 1933) and the 2-segmented endopod of Paralubbockia could
have evolved from this condition.

LUBBOCKIA Claus, 1863

DIAGNOSIS. Oncaeidae. Urosome 5-segmented in <j>, 6-segmented in $. First
antenna 4- to 7-segmented, typically with a long terminal aesthetasc in £. Second
antenna 3-segmented, terminal segment with about 6 apical elements. Mandible a
curved lash with 2 or 3 proximal elements. First maxilla bilobed, outer lobe with
3 setae, inner lobe usually with 2 setae. Second maxilla 2-segmented, distal
segment with 3 or 4 apical elements. Maxilliped large with claw-like distal segment.

no G. A. BOXSHALL

Legs 1-4 with 3-segmented rami ; exopod segment 3 with 2 or 3 external margin
spines in legs i and 2. Leg 5 a long free segment with 2 apical setae.

TYPE-SPECIES. Lubbockia squillimana Claus, 1863.

REMARKS. The genus Lubbockia contains 7 described species ; L. squillimana ,
L. aculeata Giesbrecht, L. glacialis Sars, L. brevis Farran, L. minuta Wolfenden,
L. marukawai Mori and L. wilsonae Heron & Damkaer. The last 3 species are known
only from the females. A new species is described below, from both sexes, and the
following key will enable all known species to be distinguished.

KEY TO FEMALES OF THE GENUS Lubbockia

1 . 2 spines on external margin of exopod segment 3 of legs i and 2 .... 4
3 spines on external margin of exopod segment 3 of legs i and 2 .... 2

2. Urosome longer than prosome ........ L. minuta

Urosome shorter than prosome .......... 3

3. Endopod segment 3 of leg 4 with i inner margin seta .... L. glacialis
Endopod segment 3 of leg 4 with 2 inner margin setae ..... L. brevis

4. Maxilliped with large denticles on second segment ...... 5

Maxilliped without denticles on second segment . . . . L. extenuata sp. nov.

5. First antenna 5 -segmented ; epimeral plates of fourth thoracic somite not forming a

long spiniform process ........... 6

First antenna 6-segmented ; epimeral plates of fourth thoracic somite forming a long

spiniform process ............ 7

6. Genital complex nearly as long as succeeding somite . . . L. marukawai
Genital complex much longer than succeeding somite ... L. squillimana

7. Urosome longer than prosome ........ L. wilsonae

Urosome shorter than prosome ........ L. aculeata

KEY TO MALES OF THE GENUS Lubbockia

1. 3 spines on external margin of exopod segment 3 of legs i and 2 ... 4
2 spines on external margin of exopod segment 3 of legs i and 2 .... 2

2. Maxilliped claw longer than rest of appendage . . . . L. extenuata sp. nov.
Maxilliped claw shorter than rest of appendage ....... 3

3. Epimeral plates of fourth thoracic somite rounded ; anal somite with conspicuous

constriction ........... L. squillimana

Epimeral plates of fourth thoracic somite with spiniform processes ; anal somite

without conspicuous constrictions ....... L. aculeata

4. Urosome longer than prosome . . . . . . . . . L. glacialis

Urosome shorter than prosome ......... L. brevis

Lubbockia aculeata Giesbrecht, 1891
L. aculeata Giesbrecht 1891 : 477 ; 1892 : 611, pi. 48, figs 3, 9, n, 13, 16, 20.

DESCRIPTION. Female. Body elongate (Fig. 4a), prosome longer than urosome ;
postero-lateral angles of fourth thoracic somite drawn out into slender processes ;
relative lengths of urosome somites and caudal rami 12 : 24 : 17 : 17 : 16 : 14.
Posterior margins of abdominal somites provided with dentate hyaline frill (Fig. 4f )
and all urosome somites with minute denticles scattered over surface.

OBSERVATIONS ON THE ONCAEIDAE

FIG. 4. Lubbockia aculeata. a, female entire, dorsal ; b, first antenna, ventral ; c, second
antenna, anterior ; d, mandible, anterior ; e, second maxilla, posterior ; f, urosome,
ventral ; g, maxilliped, anterior.

112 G. A. BOXSHALL

First antenna (Fig. 4b) 6-segmented ; armature elements per segment 1-3,
II-8, III -4, IV-3, V-2, VI-5.

Second antenna (Fig. 40). Second segment with single short spine ; elongate
terminal segment with an inner curved seta plumose at apex, a stout curved seta and
a group of 4 setae on distal margin, 2 setae on inner margin and 2 patches of fine
setules near outer margin.

Mandible (Fig. 4d) becoming a curved terminal lash with a denticle row on convex
margin, a row of bristles on inner margin, 2 hirsute setae and a robust proximal
spine bearing 4 spinules.

First maxilla bilobed ; outer lobe with 3, inner lobe with 2 setae.

Second maxilla (Fig. 4e). First segment with 2 patches of setules on outer margin ;
second segment with inner hirsute seta, outer naked seta and 2 spinulate terminal
elements.

Maxilliped (Fig. 4g). Basal segment with i inner denticle ; second segment with
5 denticles on inner surface ; third segment unarmed ; terminal segment claw-like
with a row of between 18 and 25 small denticles distally on concave margin and a
small proximal spinule.

Armature formula of legs 1-4 :

Coxa Basis Endopod Exopod

Leg i o-i i-I o - i ; o - i ; I, i, 4 I - o ; I - i ; II, I, 4

Leg 2 o - i i - o o - i ; o - 2 ; I, II, 3 I - o ; I - i ; II, I, 5

Leg 3 o-i i-o o- i ; 0-2; I, II, 2 I -o ; I - i ; II, I, 5

Leg 4 o-i i-o o - i ; o - 2 ; I, II, i I - o ; I - i ; II, I, 5

A single setule row present on outer margin of endopod segments 1-3 and inner
margin of exopod segment i in all legs.

Leg 5 (Fig. 4f ) elongate, produced at outer distal angle into a spinous process ;
with 2 setae, the longer not extending as far as posterior margin of genital complex.

Leg 6 a flat plate with a single naked setule.

Mean body length of female 2-21 mm, with a range of 2-06-2-35 mm.

Male. Postero-lateral angles of fourth thoracic somite spiniform as in female
(Fig. 5a) ; relative lengths of urosome somites 10 : 10 : 16 : 15 : 14 : 17 : 18.

Appendages as in female except for both first and second antennae and the
maxilliped.

First antenna (Fig. 5b) 3-segmented with extremely long terminal aesthetasc.

Second antenna (Fig. 5c). Basal segment unarmed ; middle segment with single
inner spinule ; long terminal segment with 3 curved naked setae apically, a spinulate
seta at inner, and 2 naked setae at outer distal angle, 2 patches of fine setules on
outer surface and 2 inner margin setae.

Maxilliped (Fig. 5d). Second segment with an irregular row of spinules and a
distal tubercle on the inner margin ; third segment unarmed ; fourth segment claw-
like, highly recurved and with a single proximal spinule.

Mean body length of male 2-59 mm, with a range of 2-40-2-74 mm.

OBSERVATIONS ON THE ONCAEIDAE 113

MATERIAL EXAMINED. 92 $$, 17 33 and 21 developmental stages ; from samples
3, n, 12, 16, 17, 19, 20, 22-27, 29 and 31. BM(NH) registration numbers : £$
1976.14-23, 33 1976- 24-33 and developmental stages 1976.34-43.

REMARKS. The smallest specimen of L. aculeata recovered was an immature
female with a body length of i-oo mm (Fig. 5f ). The diagnostic characters of the
species ; the 6-segmented first antennae, the armature of the maxilliped and the

FIG. 5. Lubbockia aculeata. a, male entire, dorsal ; b, first antenna (segmentation only),
ventral ; c, second antenna, anterior ; d, maxilliped, anterior ; e, female copepodite,
maxilliped, anterior ; f, female copepodite, entire, dorsal.

U4

G. A. BOXSHALL

epimeral plates of the fourth thoracic somite are all distinct at this stage. The
armature of the maxilliped (Fig. 5e) is almost identical to that of the adult female
with i denticle on the basal segment, 4 denticles on the second segment (as compared
with 5 in the adult) and a row of 18 small denticles on the concave margin of the
claw. The possession of these characters early in development permits the positive
identification of juvenile stages.

0-25mm

FIG. 6. Lubbockia squillimana. a, female entire, dorsal ; b, urosome, ventral ; c, first
antenna (segmentation only), ventral ; d, second antenna, anterior ; e, maxilliped,
anterior ; f, male maxilliped, anterior ; g, male urosome, ventral.

OBSERVATIONS ON THE ONCAEIDAE 115

Lubbockia squillimana Claus, 1863

L. squillimana Claus, 1863 : 164, pi. XXV, figs 1-5.

L. squillimana : Giesbrecht, 1892 : 611, pi. 48, figs i, 2, 4-8, 10, 12, 14, 15, 17-19, 21.

DESCRIPTION. Female. Body elongate (Fig. 6a) ; prosome longer than urosome ;
relative lengths of urosome somites n : 29 : 16 : 17 : 13 : 14 (Fig. 6b) ; epimeral
plates of fourth thoracic somite not produced posteriorly.

First antenna (Fig. 6c) 5-segmented.

Maxilliped (Fig. 6e). First segment unarmed ; second segment with 5 large
denticles on inner surface ; third segment unarmed ; terminal segment claw-like
with smooth concave margin and small proximal spinule.

Leg 5 similar in basic structure to that of L. aculeata but with the large inner seta
extending to beyond the posterior margin of the genital complex.

All other appendages as in L. aculeata.

Body length of female ranging from 1-35 to 1-50 mm, with a mean of 1-42 mm.

Male. Prosome longer than urosome ; fourth thoracic somite with rounded
posterior angles ; relative lengths of urosome somites (Fig. 6g) 10 : 12 : 14 : 16 : 18 :
13 : 17. Anal somite with conspicuous constriction in middle of somite.

Second antenna (Fig. 6d) similar to that of L. aculeata but with only i small spine
on inner margin of distal segment.

Maxilliped (Fig. 6f ). Unarmed basal segment ; second segment with a row of fine
spinules and a distal tubercle on the inner margin ; third segment unarmed ; ter-
minal claw with a proximal spinule.

Body length of single male specimen 2-40 mm.

MATERIAL EXAMINED. 20 $$, i $ and 8 developmental stages ; from samples 12,
14, 19, 20, 22, 26-29 and 31. BM(NH) registration numbers : $$ 1976.44-53,
$ 1976.54 and developmental stages 1976.55-62.

Lubbockia extenuata sp. nov.

DIAGNOSIS. Body elongate, prosome longer than urosome ; epimeral plates of
fourth thoracic somite rounded. Caudal rami longer than anal somite and about
5 times longer than broad. First antenna ($) 6-segmented. First maxilla ($) with 3
long setae on outer lobe and 2 setae and a spinule on inner lobe. Second maxilla ($)
with 2 rows of fine setules on basal segment. Maxilliped ($) second segment un-
armed ; terminal claw with a row of small denticles on concave margin. Maxilliped
(^) second segment with a row of fine spinules ; terminal claw longer than rest of
appendage. Legs 1-4 armature formula as in L. aculeata. Leg 5 with longer inner
apical seta not extending to posterior margin of genital complex.

DESCRIPTION. Female. Prosome rounded anteriorly and longer than urosome
(Fig. 7a) ; epimeral plates of fourth thoracic somite rounded ; relative lengths of
urosome somites and caudal ramus 13 : 31 : 17 : 15 : 12 : 12. Posterior margins of
abdominal somites smooth.

First antenna (Fig. 7b) 6-segmented, with aesthetascs on segments 2, 4 and 6.

n6

G. A. BOXSHALL

a

FIG. 7. Lubbockia extenuata sp. nov. a, female entire, dorsal ; b, first antenna, ventral ;
c, second antenna, anterior ; d, mandible, anterior ; e, first maxilla, posterior ; f, second
maxilla, anterior ; g, maxilliped, anterior ; h, male second antenna, anterior ; i, maxil-
liped, anterior ; j, female caudal ramus, dorsal ; k, male entire, dorsal.

OBSERVATIONS ON THE ONCAEIDAE 117

Second antenna (Fig. 70) comprising an unarmed basal segment, middle segment
with single spinule and terminal segment with 2 rows of fine setules, 2 spinules on
lateral margin and with 6 apical elements ; I straight seta, i curved spinulate seta
and 4 curved spines.

Mandible (Fig. yd) similar to that of L. aculeata.

First maxilla (Fig. ye) with 3 long setae on outer lobe and 2 unequal setae and a
small spinule on inner lobe.

Second maxilla (Fig. yi ) segments well defined ; basal segment with 2 rows of
setules, apical segment with 4 armature elements as in other members of the genus.

Maxilliped (Fig. 7g). First segment with a patch of minute rounded denticles on
inner surface ; second segment elongate and unarmed ; third segment short ; ter-
minal claw with a row of 18 small denticles on the concave margin and a proximal
spinule.

Legs 1-4 armature as in L. aculeata.

Leg 5 with 2 apical setae, the longer inner seta not extending as far as the posterior
margin of the genital complex.

Leg 6 represented by a chitinous plate with a single spinule.

Body length of 2 female specimens 1-72 and 1-74 mm.

Male. Prosome longer than urosome (Fig. 7k) ; epimeral plates of fourth thoracic
somite rounded ; relative lengths of urosome somites 14 : 15 : 19 : 16 : n : n : 14.
Caudal ramus (Fig. 73).

First antenna 3-segmented.

Second antenna (Fig. 7h) basal segment unarmed ; second segment with inner
setule ; terminal segment with a long curved seta and a short seta on the distal
margin, a curved plumose seta and 3 straight setae on the inner distal margin, a
setule on the inner margin and 2 patches of fine setules on the outer surface.

Mandibles, first and second maxillae not dissected.

Maxilliped (Fig. 7!) with unarmed basal segment ; second segment with a regularly
convex inner margin provided with fine spinules ; third segment unarmed ; terminal
claw longer than rest of appendage and with a very fine serrated membrane along its
concave margin.

Legs 1-6 as in the female.

Body length of male 1-34 mm.

MATERIAL EXAMINED. 2 $$ and i $ ; the 2 $$ from sample 24 and i £ from sample
ii. BM(NH) registration numbers : holotype £ 1976.63, paratype <$ 1976.64 and
paratype $ 1976.65.

REMARKS. The new species is closely related to both L. aculeata and L. squillimana
and it possesses a number of characters in common with each of these species. It
has, for example, rounded epimeral plates on the fourth thoracic somite, as in L.
squillimana but the inner seta on leg 5 does not extend as far as the posterior margin
of the genital complex, a character exhibited by L. aculeata. The most distinctive
character of the new species is the reduced armature of the maxilliped. In the female
the second segment is unarmed whereas both L. aculeata and L. squillimana have
large denticles on this segment. It was thought possible that the absence of denticles

14

n8 G. A. BOXSHALL

merely indicated that the specimens were immature and that the denticles were
formed late in development but juvenile L. aculeata were examined (see p. 114) and
found to possess these denticles at an early stage in development. In the absence
of denticles on the second maxilliped segment the new species resembles L. glacialis
but the presence of 3 spines on the external margin of exopod segment 3 in legs i
and 2 in L. glacialis is a significant difference between them.

In the male of the new species the terminal claw of the maxilliped is longer than
the rest of the appendage whereas in the species of Lubbockia for which the male has
been described the terminal claw only just extends as far as the proximal portion of
the second segment. The specific name refers to the long slender form of the
maxilliped in both sexes of the new species.

Lubbockia glacialis G. O. Sars, 1900
L. glacialis G. O. Sars, 1900 : 114-8, pi. XXXIII.

DESCRIPTION. Male. Elongate body with urosome longer than prosome
(Fig. 8a) ; relative lengths of urosome somites 8 : 12 : 16 : 23 : 28 : 4 : 9.

First antenna (Fig. 8b) as described by Heron & Damkaer (1969) but with an
additional apical aesthetasc, approximately 7 times longer than the appendage
itself.

Second antenna (Fig. 8c) with 6 apical setae ; i longer than the distal segment, I
short and hirsute and 4 short and naked ; outer margin of distal segment with i
proximal setule and 2 distal setae.

Mandible damaged in dissection.

First maxilla (Fig. 8d) bilobed with 3 setae on outer lobe and 2 on inner lobe as in
other members of the genus.

Second maxilla (Fig. 8e), maxilliped (Fig. 8f) and legs 5 and 6 as described by
Heron & Damkaer. Legs 1-4 (Figs 8g, h) as figured by Sars (1900) for the female.

Body length of 2 male specimens 2*08 and 1-91 mm.

MATERIAL EXAMINED. 2 <&£ ; from sample 25. BM(NH) registration numbers :
1976.66-67.

REMARKS. Sars (1900) originally described L. glacialis from the female only but
the male has recently been described by Heron & Damkaer (1969). As this is the
first record of L. glacialis outside the North Polar region, except for that of Olson
(1949) which Heron & Damkaer regarded as a misidentification of L. minuta, the
diagnostic characters of the species are figured. Some small differences were ob-
served between the present specimens and those described by Heron & Damkaer
(1969). The presence of a normal first maxilla in the present specimens suggests
that the anterior mouthparts of Heron & Damkaer's specimen may have been dam-
aged during collection or dissection as were the mandibles of the present specimen.
The other differences between the specimens, such as the position of the armature
elements on the terminal segment of the second antenna, are not significant and
probably result from the difficulty in determining the exact point of insertion of a
small seta.

OBSERVATIONS ON THE ONCAEIDAE

119

FIG. 8. Lubbockia glacialis. a, male entire, dorsal ; b, first antenna, ventral ; c, second
antenna, anterior ; d, first maxilla, posterior ; e, second maxilla, posterior ; f, maxil-
liped, anterior ; g, leg i, posterior ; h, leg 4, posterior.

G. A. BOXSHALL

FIG. 9. Lubbockia brevis. a, male entire, dorsal ; b, first antenna, ventral ; c, second
antenna, anterior ; d, urosome, ventral ; e, mandible, anterior ; f, first maxilla, posterior ;
g, second maxilla, anterior ; h, maxilliped, anterior.

OBSERVATIONS ON THE ONCAEIDAE 121

Lubbockia brevis Farran, 1908
L. brevis Farran, 1908 : 96-7, pi. XI, figs 1-9.

DESCRIPTION. Male. Prosome very broad and twice as long as the urosome
(Fig. ga) ; genital complex short ; caudal rami just longer than anal somite.

First antenna (Fig. 90) y-segmented ; armature elements per segment 1-3,
II - 6, III - 3, IV - 4, V - 2, VI - i, VII - 6.

Second antenna (Fig. gc) 3-segmented ; second segment small, unarmed ; third
segment with 6 apical elements, a long curved seta, 3 shorter curved spines, i straight
seta and a small hirsute seta, and with 3 slender setae on the inner margin. Two
patches of fine setules present on outer surface of third segment.

Mandible (Fig. 96) armature comprising a curved apical lash with a row of denticles
on the convex margin and a row of fine spinules on the concave margin, 2 hirsute
setae lateral to lash, an inner and an outer proximal seta and a row of bristles.

First maxilla (Fig. gi ) bilobed ; inner lobe with 2, outer lobe with 3 setae.

Second maxilla (Fig. gg). Basal segment robust, terminal segment with 2 apical
spines, i with spinules, and 2 hirsute setae.

MaxiUiped (Fig. gh) 3-segmented ; middle segment with a row of fine spinules on
inner margin ; terminal claw unarmed.

Legs 1-4 (Figs zoa-d) armature formula as follows :

Coxa Basis Endopod Exopod

Leg i o-i i-I o - i ; o - i ; I, I, 4 I - o ; I - i ; III, I, 4

Leg 2 o-i i-o o-i ; 0-2 ; I, II, 3 I-o;I-i; 111,1,5

Leg 3 o-i i-o o-i;o-2; I, II, 1 + 2 I - o ; I - i ; II, I, 5

Leg 4 o-i i-o o - i ; o - 2 ; I, II, I + i I - o ; I - i ; II, I, 5

Apical spine on exopod of legs 2 and 3 about as long as whole exopod, on leg 4 longer
than exopod.

Leg 5 (Fig. gd) a free segment with a long apical seta and a short lateral seta.

Leg 6 represented on genital complex by a short postero-lateral projection with a
apical spinule.

Body length of male specimens 1-43 and 1-18 mm.

MATERIAL EXAMINED. 2 JcJ ; from samples 8 and 32. BM(NH) registration
numbers : 1976.68-69.

REMARKS. The presence of 3 external spines on exopod segment 3 of legs i and 2
is a character exhibited by only 3 species of Lubbockia, L. minuta, L. glacialis and
L. brevis. These species are closely related but differ in the proportional lengths of
prosome and urosome. The male of L. minuta is unknown but the urosome is longer
than the prosome in female L. minuta (Wolfenden, 1905) whereas it is shorter than
the prosome in female L. brevis. L. brevis differs from L. glacialis in the possession
of 2 setae on the inner margin of endopod segment 3 of leg 4 as compared with a
single seta in the latter species. The present specimens are, therefore, referred to
L. brevis on the basis of their body proportions and leg 4 armature. The relatively
short genital complex, the position of leg 6 and the short maxilliped claw suggest

122

G. A. BOXSHALL

FIG. 10. Lubbockia brevis. a, leg i, posterior ; b, leg, 2, posterior ; c, leg 3, posterior

d, leg 4, posterior.

OBSERVATIONS ON THE ONCAEIDAE 123

that these specimens are males and the segmentation of the urosome indicates that
they are immature.

L. brevis is uncommon but has a widespread geographical distribution with records
from the northeastern Atlantic Ocean (Farran, 1908 ; present account), the south-
eastern Pacific Ocean (Bjornberg, 1973) and the northwestern Pacific Ocean (Wilson,
1950).

ONCAEA Philippi, ±843

Oncaea Philippi, 1843 : 63.
Antaria Dana, 1846 : 229.

DIAGNOSIS. Oncaeidae. Urosome 5-segmented in $ and 6-segmented in £.
First antenna 6-segmented ($) or 4-segmented (<^) with a long third segment ; second
antenna 3-segmented usually with a proximal armature group of 4 setae and a distal
group of 6 or 7 setae on the terminal segment. Mandible usually with 2 apical
blades and 2 hirsute setae. First maxilla bilobed, inner lobe with 3 setae, outer
with 4 setae. Second maxilla 2-segmented, distal segment with 3 or 4 elements.
Maxilliped 4-segmented, second segment with 2 setae on inner surface, fourth seg-
ment claw-like. Legs 1-4 with 3-segmented rami. Leg 5 usually a free segment
with 2 apical setae.

TYPE-SPECIES. Oncaea venusta Philippi, 1843.

REMARKS. The main characters used to separate the species of Oncaea are the
proportional lengths of the body somites, the relative dimensions of the caudal rami,
the relative lengths of the second and third segments of the second antenna, the
armature of the maxilliped and the structure of the fourth thoracic leg. Other
characters such as the armature of the terminal segment of the second antenna, of the
mandible and of the first maxilla have rarely been used. The proportional lengths
of the urosome somites and caudal rami can be affected significantly by fixation which
often causes telescoping of the somites and it is important, therefore, to use such
measurements only in conjunction with other characters based on the armature of the
appendages.

Variation in size and in body proportion is common in species of Oncaea. It has
been reported for several species ; 0. venusta (Farran, 1929 ; Sewell, 1947), 0.
conifer a Giesbrecht (Farran, 1936 ; Moulton, 1973), 0. mediterranea (Claus) (Farran,
1929 ; Ferrari, 1975) and 0. media Giesbrecht (Sewell, 1947). Similar variation
was encountered in the present material, notably in 0. venusta, 0. conifera and 0.
ornata Giesbrecht. It is important that study should be concentrated upon the
extent of variation in closely related species especially before the establishment of a
new species from a small number of specimens.

During the last decade 14 new species of Oncaea have been described : 0. ivlevi
Shmeleva, 1966, 0. prendeli Shmeleva, 1966, 0. zernovi Shmeleva, 1966, 0. ovalis
Shmeleva, 1966, 0. atlantica Shmeleva, 1967, 0. vodjanitskii Shmeleva, 1969, 0.
tregoubovi Shmeleva, 1969, 0. bathyalis Shmeleva, 1969, 0. longiseta Shmeleva, 1969,
0. brodskii Shmeleva, 1969, 0. longipes Shmeleva, 1969, 0. minima Shmeleva, 1969,

124 G. A. BOXSHALL

0. neobscura Razouls, 1969 and 0. latimana Gordeyeva, 1975. The discovery of
such a large number of new species mostly from well-studied areas is surprising but
10 of these species are less than 0-5 mm in body length and may not have been taken
by earlier workers using larger meshed plankton nets. 0. minima is the smallest
species with a body length of 0-18 mm but, from the description (Shmeleva, 1969),
it appears to be a mature form. Unfortunately many of the characters of taxonomic
importance cannot be discerned from the original descriptions of these species and
they are in need of redescription.

Eight species of the genus Oncaea were found in the present samples. Four of
them, 0. mediterranea, 0. venusta, 0. conifera and 0. media, are the commonest
representatives of the genus and their descriptions are generally limited to diagnostic
features. Three species, 0. notopus Giesbrecht, 0. ornata and 0. curta Sars, are less
common and are described in detail. The remaining species is new to science.

Oncaea venusta Philippi, 1843

Oncaea venusta Philippi, 1843 : 63, pi. Ill, fig. 3.
Antaria obtusa Dana, 1852 : 1230, pi. LXXXVI, figs i3a-c.
Oncaea obtusa : Brady, 1883 : 120, pi. LI, figs i-n.

Oncaea venusta : Giesbrecht, 1892 : 590, pi. II, fig. 5, pi. Ill, fig. 7, pi. XLVII, figs 5, 13, 19, 39,
44, 48, 50, 54, 58.

DESCRIPTION. Female. Prosome vaulted, giving an arched appearance from the
lateral aspect (Fig. na) ; prosome provided with minute tubercles regularly dis-
tributed over the dorsal and lateral surfaces (Figs nc, e, f ). Genital complex about
equal to or slightly longer than rest of urosome. Caudal rami about twice as long as
anal somite and 3-2 to 4-5 times longer than broad.

First antenna 6-segmented.

Second antenna with large basal segment bearing a single distal seta ; second
segment with smooth inner margin and some fine setules near the outer margin ;
third segment smaller than second, with a proximal group of 4 setae and a distal
group of 7 setae.

Mandible with 2 terminal blades, i with spinules, and 2 hirsute setae.

First maxilla bilobed, outer lobe with 4 apical setae, inner lobe with i lateral and
2 apical setae.

Second maxilla with 2 spinulate terminal elements and a slender subapical seta.

Maxilliped (Fig. nh). Second segment robust with a spinule row and 2 setae on
inner surface, both setae with a single row of spinules ; third segment unarmed ;
terminal claw with a proximal spinule and a spinule row on the concave margin.

Legs 1-4 armature formula as follows :

Coxa Basis Endopod Exopod

Leg i o-o i-i o-i;o-i;o, I, 5 I - o ; I - i ; III, I, 4

Leg 2 o-o i-o o - i ; o - 2 ; II, I, 3 I - o ; I - i ; III, I, 5

Leg 3 o-o i-o o - i ; o - 2 ; II, I, 2 I - o ; I - i ; II, I, 5

Leg 4 o-o i-o o - i ; o - 2 ; I, II, i I - o ; I - i ; II, I, 5

OBSERVATIONS ON THE ONCAEIDAE

125

Apical spine on exopod segment 3 of all legs shorter than the segment. Exopod
segments i and 2 with a fine serrated membrane along the external margin in legs
1-4.

Leg 5 (Fig. nk) a free segment about 1-5 to 1-7 times longer than broad and with
2 apical setae of similar lengths.

a

FIG. n. Oncaea venusta. a, female, lateral ; c, female forma typica, dorsal ; e, female
robust form, dorsal ; f, female forma venella, dorsal ; g, maxilliped of robust form,
posterior ; h, maxilliped of forma typica ; k, leg 5, dorsal. Oncaea mediterranea. b,
female, lateral ; d, female, dorsal ; i, maxilliped, posterior ; j, leg 5, dorsal.

126

G. A. BOXSHALL

FIG. 12. Oncaea venusta. a, male urosome, ventral ; b, second antenna, anterior ; c,
first maxilla, posterior ; d, maxilliped, anterior. Oncaea mediterranea. e, male urosome,
ventral ; f, second antenna, anterior ; g, first maxilla, posterior ; h, maxilliped, anterior.

Male. Genital lappets produced into small postero-lateral processes (Fig. I2a) ;
caudal rami about 1-8 times longer than anal somite and about 2-2 times longer than
broad. Appendages as in $ except for antennae and maxilliped.

First antenna 4-segmented, with distal segment representing 3 terminal segments
of female.

OBSERVATIONS ON THE ONCAEIDAE

127

Second antenna (Fig. I2b) with 2 robust spines and 2 setae in proximal armature
group on third segment, and with 7 setae in distal group.

First maxilla (Fig. 120) bilobed ; outer lobe with 4 setae, the outermost seta is
slender and unarmed ; inner lobe with 3 setae.

Maxilliped (Fig. I2d) with single row of large blunt spinules on inner margin of
second segment ; terminal claw with a proximal spinule, concave margin of claw
smooth.

Body length of male ranging from 0-96 to 1-08 mm, with a mean of i-oi mm.

MATERIAL EXAMINED. 1551 $$ and 32 $$ ; from samples 3, 4, 5, 8, 9, 11-17, Z9»
20 and 22-25. BM(NH) registration numbers : 488 $$ (forma typica} 1976.70-79,
1052 $$ (forma venella} 1976.80-89, n $$ (robust form) 1976.90-99 and 32 <&J
1976.100-109.

REMARKS. Three forms of female 0. venusta were found in the series of samples,
two of which differed only in size (Figs lie, f ) with no other detectable differences.
A size frequency analysis of 333 female specimens of 0. venusta reveals a distinct
bimodal pattern with one mode around a length of 0-98 mm and the other around
1-13 mm (Fig. 13). The small form corresponds to 0. venusta f. venella as described
by Farran (1929) and the large form corresponds to 0. venusta f. typica. Some
individuals intermediate in size between f . typica and f . venella were recorded. The
existence of these two forms of 0. venusta has been noted by several authors (Farran,
1929 & 1936 ; Sewell, 1947 ; Tanaka, 1960 ; Ferrari, 1975) although the actual
range of body length recorded varied with locality. The distribution of the two
forms is summarized by Sewell (1947).

The third form of 0. venusta was present in small numbers in a few of the samples
and is referred to as the robust form. It is very squat in general appearance

0-83

091

6-98 105 1-13

body length (mm)

1-20

1-27

FIG. 13. Frequency distribution of body length in 333 specimens of Oncaea venusta
taken from samples 19 and 27.

128 G. A. BOXSHALL

(Fig. lie) and its exoskeleton is highly sclerotized at the posterior end of the urosome.
The appendages are identical to those of f. typica (see Fig. ng for example) but the
caudal rami are relatively more slender being about 4-0 to 4-5 times longer than
broad. The length of the robust form ranged from 0-88 to 1-40 mm.

Oncaea mediterranea (Glaus, 1863)

Antaria mediterranea Claus, 1863 : 159, pi. XXX, figs 1-7.

Oncaea mediterranea : Giesbrecht, 1892 : 591, pi. IV, figs 4, 16, pi. XL VII, figs 8-10 and 47.

DESCRIPTION. Female. This species is closely related to 0. venusta but can be
separated from it by the following characters.

Prosome not strongly vaulted ; body not giving an arched appearance from the
lateral aspect (Fig. lib) ; genital complex about 1-2 times longer than rest of urosome
(Fig. i id) and with sides tapering posteriorly. Caudal rami 2-4 times longer than
anal somite and 4-0 to 4-5 times longer than broad.

Maxilliped (Fig. ni). Setae on second segment both short and hirsute ; terminal
claw with a short row of denticles on concave margin.

Leg 5 (Fig. ii j) more elongate than in 0. venusta, 2-3 to 2-5 times longer than broad
and with i long apical seta and I short subapical seta.

Mean body length of female 1-25 mm, with a range from 0-98 to 1-35 mm.

Male. Genital lappets (Fig. I2e) produced into large postero-lateral points ;
caudal rami 1-4 times longer than the anal somite and about 2-8 times longer than
broad.

Second antenna (Fig. I2f ) with i robust seta in the proximal armature group on
the terminal segment.

First maxilla (Fig. I2g) with the outermost seta on the outer lobe well developed
and bearing a row of fine spinules.

Maxilliped (Fig. i2h) similar to that of male 0. venusta.

Body length of male ranging from 0-96 to 1-08 mm, with a mean of i-oi mm.

MATERIAL EXAMINED. 1881 $$ and 138 &? ; from samples 3-5, 8, 9, 11-17, I9»
20, 22-25, 32 and 34- BM(NH) registration numbers : $$ 1976.110-119 and <&£
1976.120-129.

REMARKS. Only one form of 0. mediterranea was found in the present sample
series although both Farran (1929) and Ferrari (1975) have reported the existence of
up to three distinct size groups in this species.

Oncaea rotundata sp. nov.

DIAGNOSIS. Female prosome rounded anteriorly, twice as long as urosome ;
genital complex about as long as rest of urosome ; caudal rami just shorter than anal
somite and 2-0 to 2-5 times longer than broad. Second antenna second segment
longer than third and with a row of denticles along inner margin. Segment 3 of
endopod of leg 4 about equal to or just longer than segments i and 2 combined,

OBSERVATIONS ON THE ONCAEIDAE

129

swollen proximally and tapering towards apex ; inner margin with I seta, outer
margin unarmed, apex with 2 unequal spines.

DESCRIPTION. Female. Prosome 5-segmented, rounded anteriorly and twice as
long as urosome (Fig. 143.) ; genital complex (Fig. I4f) as long as rest of urosome.
Caudal rami just shorter than anal somite, about 2-0 to 2-5 times longer than broad
and with 6 setae.

a

9

FIG. 14. Oncaea rotundata sp. nov. a, female entire, dorsal ; b, first antenna (segmen-
tation only), ventral ; c, second antenna, anterior ; d, mandible, anterior ; e, first maxilla,
posterior ; f, urosome, dorsal ; g, maxilliped, anterior ; h, second maxilla, posterior.

130 G. A. BOXSHALL

First antenna (Fig. I4b) 6-segmented, armature elements per segment ; 1-2,
II -5, III -4, IV -4, V-2 and VI -5.

Second antenna (Fig. 140). First segment with a long hirsute seta at postero-
lateral angle and a fine spinule row on margin proximal to seta ; second segment
with a denticle row on inner margin and a spinule row on ventral surface ; third
segment shorter than second, with proximal group of 4 hirsute setae and distal
group of 7 setae.

Mandible (Fig. I4d) with 2 apical blades, i spinulate and i naked, a long hirsute
seta on the inner margin and an outer spinulate seta.

First maxilla (Fig. 146) bilobed ; inner lobe with i lateral and 2 apical hirsute
setae ; outer lobe with i unarmed and 3 hirsute setae and a short row of fine spinules.

Second maxilla (Fig. I4h) with 4 spinulate terminal elements.

Maxilliped (Fig. I4g). Robust second segment with 2 spinule rows on inner
surface, a proximal naked seta and a distal stout serrate seta ; terminal claw with
a short inner spinule and a denticle row along concave margin.

Legs 1-4 (Figs I5a-d). Both rami 3-segmented ; armature formula as follows :

Coxa Basis Endopod Exopod

Leg i o-o i-I o-i;o-i;o, I, 5 I-o;I-i; III, I, 4

Leg 2 o-o i-o o - i ; o - 2 ; II, I, 3 I - o ; I - i ; III, I, 5

Leg 3 o-o i-o o - i ; o - 2 ; II, I, 2 I - o ; I - i ; II, I, 5

Leg 4 o-o i-o o - i ; o - 2 ; o, II, i I - o ; I - i ; II, I, 5

Apical spine on exopod of all legs externally serrate, internally plumose ; external
margin spines serrate ; serrations also present on lateral margins of exopod segments.
Rows of setules present on lateral margins of endopod segments and on inner margin
of exopod segment i. Endopod longer than exopod in legs 1-3, shorter than exopod
in leg 4 ; endopod segment 3 of leg 4 equal to or just longer than segments i and 2
combined, swollen proximally and tapering distally.

Leg 5 (Fig. I4f ) with lateral seta and free segment bearing 2 apical setae.

Body length of female specimens 1-27 and 1-05 mm.

MATERIAL EXAMINED. 2 $$ ; holotype from sample 32, paratype from sample 5.
BM(NH) registration numbers : holotype $ 1976.130 and paratype $ 1976.131.

REMARKS. The new species is closely related to 0. obscura Farran, 1908 but it can
be distinguished from 0. obscura by the structure and armature of leg 4. A spine
is present on the lateral margin of endopod segment 3 of leg 4 in 0. obscura (armature
formula of endopod o - i ; o - 2 ; I, II, i) whereas in the new species the lateral
margin is unarmed. Endopod segment 3 of leg 4 is about as long as segments i and
2 combined whereas in 0. obscura segment 3 is 1-5 to 2-0 times longer than segments
i and 2 combined. The shape of endopod segment 3 is different in the two species.
The setae of the first maxilla are hirsute and generally longer in the present specimens
than in 0. obscura. Another difference is that the urosome of 0. obscura is more
slender than that of the new species although this may possibly be related to the
state of maturity of Farran's specimens which was not given in the original descrip-
tion (Farran, 1908).

FIG. 15. Oncaea rotundata sp. nov. a, leg i, posterior ; b,

posterior ; d, leg 4, posterior.

2, posterior ; c, leg 3,

132

G. A. BOXSHALL

Oncaea notopus Giesbrecht, 1891
Oncaea notopus Giesbrecht, 1891 : 477 ; 1892 : 603, pi. XLVTI, figs 12, 15 & 45.

DESCRIPTION. Female. Genital complex (Fig. i6a) longer than rest of urosome ;
anal somite 1-5 to i«8 times longer than caudal rami. Caudal rami less than twice
as long as broad.

FIG. 1 6. Oncaea notopus. a, female urosome, dorsal ; b, second antenna, anterior ; c,
mandible, anterior ; d, first maxilla, posterior ; e, maxilliped, anterior. Oncaea media.
f, female urosome, dorsal ; g, second antenna, anterior ; h, mandible, anterior ; i, first
maxilla, anterior ; j, maxilliped, anterior ; k, leg 4, posterior.

OBSERVATIONS ON THE ONCAEIDAE 133

Second segment of second antenna (Fig. i6b) longer than third, with smooth
inner margin ; third segment with proximal and distal armature groups, with i seta
of distal group much longer than the others.

Inner blade of mandible (Fig. i6c) simple.

Outer lobe of first maxilla (Fig. i6d) with i naked and 3 hirsute setae ; inner lobe
with a spinulate seta and 2 setules.

Second maxilla with 3 terminal elements.

Maxilliped (Fig. i6e). Second segment with 2 setae situated close together near
the middle of the segment, distal seta serrate, proximal seta naked ; terminal claw
short, stout and armed with a row of denticles along the concave margin and a
proximal spinule.

Legs 1-4 armature formula as follows :

Coxa Basis Endopod Exopod

Leg i o-+ i-I o-i;o-i;o, I, 5 I - o ; I - i ; III, I, 4

Leg 2 o-+ i-o o - i ; o - 2 ; II, I, 3 I-o;I-i; III, I, 5

Leg 3 o-+ i-o o - i ; o - 2 ; II, I, 2 I - o ; I - i ; II, I, 5

Leg 4 o - + i-o o - i ; o - 2 ; I, II, i I - o ; I - i ; II, I, 5

( + small spinule present.)

Apical seta on exopod segment 3 shorter than the segment in all legs ; endopod of
leg 4 with apical seta about half as long as segment 3.

Leg 5 (Fig. i6a) free segment elongate, extending posteriorly to level of genital
apertures, with 2 apical setae.

Body lengths of the female specimens i-oi and 0-98 mm.

MATERIAL EXAMINED. 2 $$ from samples 4 and 5. BM(NH) registration
numbers : 1976.172 and 1976.173.

REMARKS. This species is readily identifiable on the basis of its elongate leg 5
which extends posteriorly to the level of the genital apertures on the genital complex.
Some minor differences in appendage armature were found between the present
specimens and those described by Sars (1900). The second antennae of the present
specimens possessed i long and 5 shorter setae in the distal armature group whereas
in Sars' specimens these setae were ah1 about equal in length except for the relatively
short apical seta. A spinule row on the second maxilliped segment was noted above
and by Giesbrecht (1892) but was not found by Sars (1900).

Oncaea media Giesbrecht, 1891

O. media Giesbrecht, 1891 : 477 ; 1892 : 602, pi. II, fig. 12, pi. XLVII, figs i, n, 29-33 & 40.
O. curia : Estrada & Genicio, 1970 : 30, pi. XII, figs 1-5.

DESCRIPTION. Female. Prosome not markedly vaulted ; genital complex
(Fig. i6f ) about 1-5 times longer than rest of urosome. Caudal rami about 1-8 times
longer than anal somite and 2-0 to 2-2 times longer than broad.

15

134

G. A. BOXSHALL

Second segment of second antenna (Fig. i6g) with smooth inner margin and longer
than third segment ; distal armature group on third segment comprising 7 setae.

Mandible (Fig. i6h) with tridentate inner blade.

First maxilla (Fig. i6i) with i naked and 3 hirsute setae on outer lobe ; inner lobe
with i naked and i spinulate apical setae and a hirsute lateral seta.

Maxilliped (Fig. i6j) with 2 very long naked setae and a short distal row of fine
spinules on the second segment ; terminal claw with a row of spinules along the
concave margin.

Legs 1-4 (Fig. i6k) armature formula as in 0. venusta. Apical setae on exopod
segment 3 shorter than the segment in all legs.

Leg 5 (Fig. i6f ) a short free segment with 2 similar apical setae.

Mean body length of female 0-81 mm, with a range of 0-78 - 0-86 mm.

MATERIAL EXAMINED. 58 $$ ; from samples 3, 5, 9, n, 13-17, 19, 22-29 and
32-34. BM(NH) registration numbers : 1976.174-183.

REMARKS. 0. media is a widespread and abundant species and in common with
many other species of Oncaea it exhibits a considerable degree of variability in the
body length of the females (Sewell, 1947 ; Tanaka, 1960 ; Ferrari, 1975). On the
basis of the range in body length observed the present material would be referrable
to the major form of Sewell (1947).

0. media and 0. curia are closely related species and are usually separated primarily
on the basis of the proportional lengths of the urosome somites in the females.
Recently numerous specimens have been found which possess body proportions inter-
mediate between 0. media and 0. curta (Olson, 1949 ; Estrada & Genicio, 1970 ;
Ferrari, 1975) and some of these have been referred to 0. curta. The wide range of
body form exhibited by female 0. curta (see p. 143) includes forms that approach
typical 0. media and it is necessary now to reappraise the criteria used to separate
these two species. A comparison of the characters of 0. media and 0. curta is given
in Table i using data derived from Giesbrecht (1892), Sars (1916), Sewell (1947),
Tanaka (1960), Shmeleva (1969), Estrada & Genicio (1970), Razouls (1974), Ferrari
(1975) and the present account.

TABLE i

A comparison between Oncaea media and O. curta

O. media O. curta-f

Length of genital complex :

length of rest of urosome 1-34-1-63 : i 1-10-1-30 : i
Length of caudal rami : length

of anal somite 1-33-2-00 : i 1-00-1-43 : i

Armature of setae on second setae usually long and setae usually of moderate

maxilliped segment naked length and armed with

hairs or serrated membrane

Lengths of second and third third segment markedly third segment equal to or

segments of second antenna shorter than second just shorter than second

Body length 0-55-0-93 mm 0-42-0-75 mm

fSee Table 2 for full derivation of these data.

OBSERVATIONS ON THE ONCAEIDAE 135

The data presented in Table i suggest that although the ranges in the relative
lengths of the urosome somites and caudal rami for 0. media and for 0. curia overlap
it is still possible to use these characters to discriminate between the species as long
as others, such as the structure of the second antennae and the armature of the
maxilliped, are also considered. On the basis of the criteria established in Table i
the 0. curta of Estrada & Genicio (1970) are referred to 0. media.

Oncaea ornata Giesbrecht, 1891
Oncaea ornata Giesbrecht, 1891 : 477 ; 1892 : 604, pi. XLVII, figs 20, 24, 49 & 53.

DESCRIPTION. Female. Form i. Prosome about 2-1 times longer than urosome
(Fig. I7a) ; genital complex about 2-5 times longer than broad and tapering pos-
teriorly into a narrow region about one fifth the length of the complex ; anal somite
approximately equal in length to the caudal rami (Fig. 170).

Second antenna (Fig. I7h) elongate, second segment with a denticulate inner mar-
gin ; terminal segment with i robust seta in the proximal armature group.

Mandible (Fig. 176) similar to that of 0. venusta.

First maxilla (Fig. i7f ) with i hirsute and 3 naked setae on outer lobe.

Maxilliped (Fig. I7g) large, second segment bearing a spinule row on the inner
margin ; distal seta 1-5 times longer than proximal seta and armed with serrate
flanges ; terminal claw with row of robust spinules on concave margin increasing in
size distally.

Legs 1-4 (Figs i8a, b) armature formula as follows :

Coxa Basis Endopod Exopod

Leg i o-+ i-I o-i;o-i;o, I, 5 I - o ; I - i ; III, I, 4

Leg 2 o-+ i-o o-i;o-2;I, I, 3 I - o ; I - i ; III, I, 5

Leg 3 o-o i-o o-i;o-2;I, I, 2 I - o ; I - i ; II, I, 5

Leg 4 o-o i-o o-i;o-2; I, I, i I - o ; I - i ; II, I, 5

(+ small spinule present (see Fig. i8a).)

Apical spine on exopod much longer than exopod segment 3 in all legs. Apical
spine on endopod of legs 2-4 longer than endopod segment 3 ; in leg i spine equal to
length of segment.

Leg 5 small, with i apical seta and i lateral seta.

Mean body length of form i female 0-99 mm, with a range from 0-91 to 1-05 mm.

Form 2. Similar to form i in all characters except : prosome (Fig. I7b) about
1-9 times longer than urosome including the caudal rami ; genital complex (Fig. I7d)
oval in outline and without a conspicuous narrow posterior portion.

Maxilliped (Fig. 17!). Distal seta on second segment about 3-5 times longer than
proximal seta ; spinules on concave margin of terminal claw not increasing in size
distally.

Mean body length of form 2 female 0-86 mm, with a range from 0-76 to 0-93 mm.

136

FIG. 17. Oncaea ornata. a, female form i entire, dorsal ; b, female form 2 entire dorsal ;
c, urosome form i, dorsal ; d, urosome form 2, dorsal ; e, mandible, anterior ; f, first
maxilla, anterior ; g, maxilliped form i, anterior ; h, second antenna, anterior ; i,
maxilliped form 2, anterior.

Male. Form i. Genital complex elongate, oval in outline (Fig. i8c) ; genital
lappets extending posteriorly, rounded. Caudal rami 1-2 times longer than the anal
somite.

Appendages as in the female except for the second antenna and maxilliped.
Second antenna (Fig. i8f) with an additional row of spinules on outer margin of

OBSERVATIONS ON THE ONCAEIDAE

137

basal segment. MaxiUiped (Fig. i8d) with unarmed basal segment ; second segment
with 2 rows of short blunt spinules on the inner surface and 2 short setae ; terminal
claw with single proximal spinule.

Mean body length of form I male 0-89 mm, with a range from 0-86 to 0-93 mm.

Form 2. Body and appendages as in form I except for the maxilliped.

FIG. 18. Oncaea ornata. a, female leg i, posterior ; b, leg 4, posterior ; c, male urosome,
ventral ; d, maxilliped form i, anterior ; e, maxilliped form 2, anterior ; f, second
antenna, anterior.

138 G. A. BOXSHALL

Maxilliped (Fig. i8e) with unarmed basal segment ; second segment with i short
row and I long row of long blunt spinules on the inner surface ; terminal claw with
a single proximal spinule.

Mean body length of form 2 male 0-77 mm, with a range from 0-74 to 0-78 mm.

MATERIAL EXAMINED. 380 ?$ and 26 <£<£ form i ; from samples 4, 5, 8, 9, 12-17,
23, 25, 32 & 34. BM(NH) registration numbers : $? 1976.132-141, $$ 1976.142-
151. 467 $$ and 22 <$<$ form 2 ; from samples 4, 5, 8, 9, 12, 13, 15-17, 22-24, 25>
32 & 34. BM(NH) registration numbers : $$ 1976.152-161 and <$<$ 1976.162-171.

REMARKS. Two readily distinguishable forms were observed in both sexes of
0. ornata. The females differed in size, body proportions, in the shape of the genital
complex and in maxilliped armature. The males differed only in size and in maxil-
liped armature. Neither form of the female, however, corresponds exactly to the
typical form described by Giesbrecht (1891 & 1892) which appears to exhibit an
intermediate condition in certain characters, such as the shape of the genital complex.
Although differences between the two forms are present in both sexes the magnitude
of these differences is not great enough to justify the separation of the forms into
distinct species especially when the variability displayed by other members of the
genus is considered.

The male of 0. ornata has not previously been described to my knowledge but the
similarities in appendage structure and armature between the males and the females
described above strongly indicate that they are conspecific. Infraspecific variation
has rarely been reported for male Oncaea except for notes on possible size variation
in 0. media (Sewell, 1947) and in 0. conifera (Ferrari, 1975) but it is probable that as
larger samples of male specimens become available this kind of variation will be
recognized more frequently.

Oncaea conifera Giesbrecht, 1891

O. conifera Giesbrecht, 1891 : 477 ; 1892 : 603, pi. XLVII, figs 4, 16, 21, 23, 28, 34-38, 42, 55
& 56.

DESCRIPTION. Female. Body form variable ; stocky (Fig. 2oa), long (Fig. 2ob)
and bumped forms present in sample series (using terminology of Moulton (1973)).
Bumped form with a dorsal projection on the cephalosome (Fig. 2oc). Second
thoracic somite of all forms with a conspicuous dorsal projection visible from the
lateral aspect (Figs iga & 2oc) ; genital complex (Fig. igb) 1-3 to 1-5 times longer
than the rest of the urosome. Caudal rami about equal to anal somite in length and
2-0 to 2-4 times longer than broad.

Second segment of second antenna with smooth margin in long and stocky forms
(Fig. 2od) but with a row of minute denticles in bumped form (Fig. 2oe) ; second
segment longer than third ; distal armature group on third segment comprising
7 setae.

Inner blade of mandible (Fig. ige) tridentate.

Outer lobe of first maxilla (Fig. igf ) with i hirsute and 3 naked setae.

OBSERVATIONS ON THE ONCAEIDAE

139

Maxilliped (Fig. igd). Second segment with 2 setae and a distal row of fine
spinules on the inner surface, distal seta serrate, proximal naked ; terminal claw
with a proximal spinule and a row of spinules along the concave margin.

Legs 1-4 armature formula as in 0. venusta. Apical spine on exopod shorter
than exopod segment 3 in all legs ; third segment of endopod of leg 4 with a conical
projection at its apex (Fig. igc).

FIG. 19. Oncaea conifera. a, female stocky form, lateral ; b, urosome, ventral ; c, leg 4,
posterior ; d, maxilliped, anterior ; e, mandible, anterior ; f, first maxilla, anterior ; g,
male urosome, ventral ; h, maxilliped, posterior ; i, second antenna, anterior.

1 4o

G. A. BOXSHALL

Leg 5 a free segment about 2-5 times longer than broad bearing a long and a short
apical seta.

Mean body length of female 1-13 mm, with a range from 0-98 to 1-29 mm.

Male. Genital lappets large and postero-laterally directed ; caudal rami shorter
than anal somite (Fig. igg) and about twice as long as broad.

FIG. 20. Oncaea conifera. a, female stocky form entire, dorsal ; b, long form entire,
dorsal ; c, bumped form, lateral ; d, second antenna stocky form, anterior ; e, second
antenna bumped form, anterior. Oncaea spp. copepodite. f, copepodite entire, dorsal ;
g, urosome, dorsal ; h, leg 4, posterior.

OBSERVATIONS ON THE ONCAEIDAE 141

Appendages as in female except for the maxilliped. Second antenna (Fig. 191).

Maxilliped (Fig. igh). Second segment with a double row of large blunt spinules
and 2 slender setae on inner surface ; terminal claw with serrate spinule proximally
and with a smooth concave margin.

Mean body length of male 0-82 mm, with a range from 0-76 to 0-88 mm.

MATERIAL EXAMINED. 7031 $$ and 34 <$<$ ; from samples 3-5, 8, 9, 11-17, I9»
20, 22-30, 32 & 34. BM(NH) registration numbers : ?$ 1976.186-195 and $$
1976.196-205.

REMARKS. The variability in body form within 0, conifera has been analysed by
Moulton (1973) using principal component analysis. The three forms of female
0. conifera which were found in the present series of samples are similar to the groups
designated stocky, long and bumped by Moulton (1973) and the stocky and long
forms also correspond to form a and form b of Farran (1936) respectively. The
taxonomic status of these well-documented forms cannot be determined without
data on the sexual isolation of the forms. Such data can only be obtained by direct
observation in experimental situations.

Oncaea curia Sars, 1916

O. curta Sars, 1916 : n, pi. IV.

O. ovalis Shmeleva, 1966 : 935, pi. 4 ; 1969 : n, figs 8 & 9.

O. longiseta Shmeleva, 1969 : 18, figs 14 & 15.

O. latimana Gordeyeva, 1975 : 1397, figs 1-14.

Oncaea sp. i Ferrari, 1975 : 228, figs 6E, F & 7A-D.

Oncaea sp. 2 Ferrari, 1975 : 22&> &8S 6G, H & 7E-H.

DESCRIPTION. Female. Prosome 1-7 to 1-9 times longer than urosome (Fig. 2ia) ;
genital complex 1-2 to 1-3 times longer than rest of urosome (Fig. 2ic) ; caudal rami
about equal to or i-i times longer than anal somite.

First antenna 6-segmented, segmental armature elements ; segment I - 2, II - 7,
III-4, IV-2, V-2, VI-5.

Second antenna (Fig. 2ib). First segment with distal outer seta ; second segment
with a short row of setules near the lateral margin ; third segment just shorter than
second and bearing a proximal group of 4 setae and a distal group comprising i long
hirsute seta and 5 shorter naked setae.

Mandible (Fig. 2 id) with tridentate inner apical blade.

First maxilla (Fig. 2ie) bilobed ; outer lobe with 4 naked setae ; inner lobe with
i spinulate seta and i naked seta apically and a hirsute seta laterally.

Second maxilla (Fig. 2 if ) with 3 apical elements ; a strong spine with a row of
large spinules, a small plumose seta and a slender outer seta.

Maxilliped (Fig. 2ig) . Second segment robust with a row of fine spinules along the
inner margin and 2 setae, the proximal naked and the distal hirsute ; terminal claw
with a row of spinules along proximal portion of concave margin, spinules increasing
in size distally.

16

I42

G. A. BOXSHALL

FIG. 21. Oncaea curia, a, female entire, dorsal ; b, second antenna, anterior ; c, urosome,
ventral ; d, mandible, anterior ; e, first maxilla, posterior ; f, second maxilla, anterior ;
g, maxilliped, anterior ; h, leg i ; i, leg 2, posterior ; j, leg 3, posterior ; k, leg 4,
posterior.

OBSERVATIONS ON THE ONCAEIDAE 143

Legs 1-4 (Figs 2ih-k) armature formula as follows :

Coxa Basis Endopod Exopod

Leg i o-o i-I o-i;o-i;o, I, 5 l_O;I-i; III, I, 4

Leg 2 o-o i-o o- 1 ; 0-2; II, I, 3 I-o; I-i; 111,1,5

Leg 3 o-o i-o o - i ; o - 2 ; II, I, 2 I - o ; I - i ; II, I, 5

Leg 4 o-o i-o o - i ; o - 2 ; II, I, i I - o ; I - i ; II, I, 5

Endopod longer than exopod in all legs ; digitiform projections present on endopod
segment 3 of legs 1-3 on both sides of apical spine and proximal to both lateral
spines on legs 2-4. Terminal spine of exopod longer than exopod segment 3 in all
legs ; inner apical spine on endopod of leg 4 approximately twice as long outer and
0-8 times as long as endopod segment 3.

Leg 5 (Fig. 2ic) a free segment with 2 equal apical setae.

Body length of female specimens 0-73 and 0-71 mm.

MATERIAL EXAMINED. 2 $$ ; from samples 16 and 24. BM(NH) registration
numbers : 1976.184 and 1976.185.

REMARKS. Within the genus Oncaea there exists a readily distinguishable group
of described species all closely related to 0. curta. This group comprises 0. curia,
0. ovalis, 0. longiseta, 0. latimana and Oncaea sp. i and 2 of Ferrari (1975). A
comparison of the diagnostic characters of these species and of the material described
above as 0. curta is given in Table 2. Infraspecific variation in size and body pro-
portions is common in species of Oncaea and is well documented (see p. 123). The
variation in the proportional lengths of the body regions and the overlap in these
measurements between the species of the 0. curta group indicate that these characters
cannot be used to separate species within the group. The details of appendage
armature given in Table 2 have been taken from the original descriptions and figures
but many important characters are either not given or are figured too poorly to be
used. The small differences in appendage armature between these species are not
sufficient to justify their separation especially when some of the characters are ex-
tremely difficult to observe, for example, the presence or absence of serrations on the
small subapical spines on the endopod of leg 4. It is proposed therefore to regard
0. ovalis, 0. longiseta, 0. latimana and Oncaea sp. i and 2 of Ferrari (1975) as junior
subjective synonyms of 0. curta.

0. curta is variable in body proportions, size and, to a smaller extent, in the arma-
ture of its appendages and in order to define the limits of this variability it will be
necessary to describe all the appendages in detail in any future work. It would
then be possible to assess the taxonomic significance of minute differences in arma-
ture, such as the presence of a single long hirsute seta in the distal armature group
at the apex of the second antenna in the form described by Shmeleva (1969) as 0.
longiseta and in the present material. The existence of infraspecific variation in the
armature of the second antenna in a well-defined species, such as 0. notopus, suggests
however that differences of this nature may not be significant.

The differences between 0. curta and 0. media have already been discussed (see
P- 134)-

144

G. A. BOXSHALL

CSt

1

y within the species O. curta

1

M

1 ? '„
"Si M

1 S. A

1 * >

<o
•«*

<3 '-^ M

§ c>

0 O f^ M
. O\ Tf N
OW . .
O M

of

e ^J
•S ^3 >o

>a r< r^-

s>| ?

?OT, ' '

o — N r^

Q 6 H

CO
p, 10

6 -i
|^6 2

M IH H

en LL, O M
^- |

1 °2 ^

. .

g 0 M

O

H ^_)

« S CD
9 rO 0

H . ^
M I?"1

h

CD

CO O

M . ~

CD

M CO

CO

a

N D
M en

IH

CD

o -e

^ fO 0

H . rfi

.. bo w

M CO
IH

2 «1

N . ^

CD

IH CO

o

H

bo
C

n..
I

ho

13

-3
t

1

3

en
-3

4->
tH

CO

0

o

i row hairs i row hairs dentate ? hirsute

i row hairs i row hairs dentate ? naked

en

13
1

-H

|t

— 4

to
CD

13

ft

•^

CO

-H .S

en
en

3

|t

ormal range of O. curta and it has been regarded as an aberrant form and omii

B

Variabilil

1 >
1 |S

3^ M ° 3-

dd o -

ro

M

.. bo

W CO

shorter

O

bo

G

hirsute

hirsute

en

1*

1

'J3

4->
'%

<1<— ^

13

1

Q "H

0

to

•g

"s 2^

OO

1

'-3

0>

i

j§

7*" <"•">

0

. »

^

-4->

o

IB lO w

M

A

IH

o

•

O

C

C/5 O H

M S^

•f.

«^

•*•

§

1 -a

^3

C

o

SH

O
g

I

*o

K rt

CD

'bo

B

~

S

D O
fl 0)

to

c3
&

0

'3

s

1

1

|i i

H M o

" 4J <D W
-M 'H S CS

bo CD O fl
C bo g cd

CO
T3

9

(S
en

-4-J

g
S" CD
g
rt hv

IH OJO

c^

antenna

Spinules on inner

-l->

d
B

i

bot3

CD CD

m p,

T3 3

§'x
« 1

en C

Distal seta on ma:
segment 2

Proximal seta on

M

-t->

C
0

&
O

M

Armature of claw

1

H

jo!

OBSERVATIONS ON THE ONCAEIDAE 145

Oncaea sp.

Many immature female specimens belonging to the genus Oncaea were recovered
from the samples. These specimens, which were not identified to the specific level,
can be recognized as developmental stages by the possession of a 4-segmented uro-
some (Fig. 2of ). Only i somite is present between the genital complex and the anal
somite as compared with 2 somites in mature forms. Another juvenile character is
the lateral position on the genital complex of the chitinous plates and spinules which
mark the position of the genital apertures in the adult (Fig. 20g). The apertures
are dorsally situated in the mature females. The appendages of the same individual
(as illustrated in Figs 2of, g) are almost indistinguishable from those of the adult.
Leg 4, for example, displays normal adult segmentation and possesses a full comple-
ment of armature elements (see Fig. 2oh).

MATERIAL EXAMINED. 68 specimens ; from samples 4, 5, 9, 11-13, 15, 16, 23, 24,
27 & 29. BM(NH) registration numbers : 1976.229-238.

CONAEA Giesbrecht, 1891

DIAGNOSIS. Oncaeidae. Urosome 5 ($) or 6 (<$) segmented. First antenna
6-segmented in $, 4-segmented in $. Second antenna 3-segmented ; third segment
with a single proximal seta and a distal group of 4 setae. First maxilla bilobed, both
lobes with 3 setae. Maxilliped 4-segmented, with distal segment claw-like ; second
segment with 2 inner setae. Legs 1-4 with 3-segmented rami, armature formula :

Leg i
Leg 2

Leg 3
Leg 4

Leg 5 reduced to a single seta.

TYPE-SPECIES. Conaea gracilis (Dana, 1852).

REMARKS. This genus is very closely related to Oncaea. The characters that
distinguish between these two genera are the armature of the second antennae and
the armature of legs 1-4. The transformation of the armature elements of both
armature groups on the third segment of the second antenna in Conaea is a character
not exhibited by any species of Oncaea, similarly the armature of the exopods of
legs 1-4 of Conaea is a feature unique to this genus.

Conaea gracilis (Dana, 1852)

Antaria gracilis Dana, 1852 : 1229, pi. 86, fig. na-d.

Conaea rapax Giesbrecht, 1891 : 477 ; 1892 : 605-6, pi. 48, figs 50-59.

Oncaea gracilis : T. Scott, 1894 : "6-7, pi. XIII, figs 4-12.

DESCRIPTION. Female. Prosome about 1-6 times longer than urosome (Fig. 22a) ;
genital complex about as long as rest of urosome including caudal rami. Caudal
rami i-i to 1-2 times longer than anal somite and 1-6 times longer than broad.

Coxa

Basis

Endopod

Exopod

0-0

i

-I

o

-x;

o-

i ; o,

1,5

I

-o;

i-i;

HI, I, 4

0-0

i

-0

0

-i;

0-

2 ; o,

1,3

I

-o;

i-i;

II, I, 5

0-0

i

-0

o

-x;

o-

2 ; o,

I, 2

I

-o;

i-i;

II, I, 5

0-0

i

-o

0

-i;

0-

2; o,

I, I

I

-o;

o-i;

1,1,5

I46

G. A. BOXSHALL

FIG. 22. Conaea gracilis. a, female entire, dorsal ; b, first antenna, ventral ; c, mandible,
anterior ; d, maxilliped, anterior ; e, leg i, posterior ; f, first maxilla, posterior ; g,
leg 2, posterior.

First antenna (Fig. 22b) 6-segmented, third segment constituting about half the
length of the appendage ; number of armature elements per segment 1-2, II - 8,

III -2, IV -2, V-2, VI -5.

Second antenna (Fig. 23d) 3-segmented ; first segment armed with an inner seta
and a row of spinules on the inner margin ; second segment with a proximal row of
spinules on the inner margin and a distal row of setules on the outer margin ; third

OBSERVATIONS ON THE ONCAEIDAE 147

segment elongate bearing a large seta and 2 spinules proximally, a row of denticles
on the anterior surface and an apical group of 4 setae and a spinule.

General form of mandible (Fig. 22c) as in Oncaea ; armature comprising 2 terminal
blades each with a row of spinules, an outer spinulate seta and an inner hirsute seta.

First maxilla (Fig. 22f ) bilobed ; outer lobe bearing 3 setae apically, inner lobe
with 2 apical and i lateral seta.

FIG. 23. Conaea gracilis. a, female leg 3, posterior ; b, leg 4, posterior ; c, male urosome,
ventral ; d, female second antenna, anterior ; e, second maxilla, anterior ; f, male
maxilliped, anterior.

148 G. A. BOXSHALL

Second maxilla (Fig. 236) 2-segmented ; distal segment terminating in a large
spinulate process and bearing a hirsute seta, a spine armed with 2 rows of spinules
and a small setule.

Maxilliped (Fig. 22d). Second segment with a short naked proximal seta, a long
distal seta with 4 spinules and a row of 8 robust spinules near the inner margin ;
terminal segment claw-like with an inner and an outer basal seta and a row of fine
spinules on its concave margin.

Legs 1-4 (Figs 22e, g & 23a, b). Armature formula as given in generic diagnosis.
Rows of setules located on the lateral margins of endopod segments i to 3 and on the
inner margin of exopod segment I on all legs. Legs 2 and 3 with strips of serrated
membrane on the lateral margins of exopod segments i to 3.

Leg 5 reduced to a single plumose seta on the surface of the first urosome segment.

Body length of female ranging from 0-88 to 1-05 mm, with a mean of 0-96 mm.

Male. Urosome (Fig. 23c) 6-segmented ; genital complex about 1-7 times longer
than rest of urosome ; caudal rami similar in length to anal somite and about 1-7
times longer than broad.

Appendages as in $ except for first antenna and maxilliped.

First antenna 4-segmented with the 3 distal segments of the female first antenna
fused, armature elements per segment 1-2, II - 8, III - 2, IV - 9.

Maxilliped (Fig. 23f ). Second segment with a row of fine spinules along the inner
margin divided into 2 rows proximally and with 2 slender unarmed setae ; terminal
segment forming an unarmed claw.

Body length of male ranging from 0-76 to 0-86 mm, with a mean of 0-81 mm.

MATERIAL EXAMINED. 3912 $$ and 99 <ft£ ; from samples 4, 5, 8, 9, 11-17, 22-
25, 29, 32 & 34. BM(NH) registration numbers : $$ 1976.206-215 and $$
1976.216-225.

REMARKS. C. gracilis is the only species of the genus and is readily distinguished
from species of Oncaea by the greater length of the urosome somites as well as by
differences in the appendages.

PACHOS Stebbing, 1910
Pachysoma Claus, 1863 : 162. (preocc.)

DIAGNOSIS. Oncaeidae. Body globular with very inflated prosome and short
4 (?) or 5 (<£) segmented urosome. Rostrum prominent. Second antenna 4-seg-
mented. Mandible reduced to a curved lash. First maxilla bilobed ; outer lobe
with 3 setae. Second maxilla with 2 apical elements. Maxilliped 3- or 4-seg-
mented ; terminal segment claw-like especially in <$. Legs 1-4 with 3-segmented
rami, armature formula :

Coxa Basis Endopod Exopod

Leg i o-i i-o o-i;o-i;o, I, 5 I - o ; I - i ; III, I, 4

Leg 2 o-i i-o o - i ; o - 2 ; I, II, 3 I - o ; I - i ; III, I, 5

Leg 3 o-i i-o o - i ; o - 2 ; I, II, 2 I-o;I-i; III, I, 5

Leg 4 o-i i-o o - i ; o - i ; o, II, o I - o ; I - i ; II, I, 5

OBSERVATIONS ON THE ONCAEIDAE 149

Leg 5 a conical process with 2 short apical setae. Leg 6 represented by 2 long
plumose setae.

TYPE-SPECIES. Pachos punctatum (Claus, 1863).

The genus Pachos comprises 3 species ; P. punctatum, P. tuberosum (Giesbrecht,
1891) and P. dentatum (Mori, 1932). The species can be separated using the following
key :

KEY TO FEMALES OF THE GENUS Pachos

1. Dorsal surface of carapace ornamented with minute denticles .... 2
Dorsal surface of carapace ornamented with a reticulate pattern of minute chitinous

ridges ............ P. tuberosum

2. Maxilliped 3 -segmented ......... P. punctatum

Maxilliped distinctly 4-segmented ....... P. dentatum

KEY TO MALES OF THE GENUS Pachos^

Maxilliped claw with broad spatulate tip ...... P. punctatum

Maxilliped claw with a narrow pointed tip ...... P. tuberosum

Pachos differs markedly from the other genera of the Oncaeidae and Sars (1918)
regarded its systematic position in this family as rather questionable. The aberrant
structure of the mouthparts in Pachos is the prime factor isolating this genus from
the rest of the family. However, detailed comparison with Lubbockia, for example,
demonstrates that the basic structure of each limb is the same. The mandible in
Pachos is reduced to a curved lash with denticles on the outer surface and a row of
bristles on the inner surface. This condition could readily be derived from that in
Lubbockia (cf. Fig. 4d) by the loss of the additional proximal armature elements.
The first maxilla is bilobed in both genera. The second maxilla is 2-segmented in
both although again there is a reduction in the number of armature elements in
Pachos. The structure of the maxillipeds is very similar with 2 setae present on the
second segment and with the distal segment forming a claw bearing at least I proxi-
mal spinule. The second antenna is 4-segmented in Pachos but only 3-segmented in
Lubbockia ; however, the distal segment of the second antenna in Lubbockia probably
represents the fused third and fourth segments of this appendage in Pachos and the
outer margin setae halfway along the third segment in Lubbockia represent the
vestiges of the armature elements found on the third segment in Pachos. Such
detailed comparisons show that the relationships of Pachos probably lie with the
family Oncaeidae.

Pachos punctatum (Claus, 1863)

Pachysoma punctata Claus, 1863 : 163, pi. XXV, figs 6-n.

P achy soma punctatum : Giesbrecht, 1892 : 612-615, pi. 48, figs 22-36, 38, 39.

DESCRIPTION. Female. Body globular, with inflated prosome and short uro-
some ; surface of carapace covered with minute denticles ; a pair of sensory setules

f The male described by Mori (1932 & 1937) as the male of P. dentatum does not differ significantly
from P. tuberosum (see p. 154) and is not included in the key.

FIG. 24. Pachos punctatum. a, female urosome, ventral ; b, first antenna (segmentation
only), ventral ; c, second antenna, anterior ; d, mandible, anterior ; e, first maxilla,
anterior ; f, second maxilla, anterior ; g, maxilliped, an tero- ventral ; h, leg i, posterior ;
i, leg 2, posterior ; j, leg 3, posterior ; k, leg 4, posterior.

OBSERVATIONS ON THE ONCAEIDAE 151

present on carapace and each of remaining 3 prosome somites. Rostrum prominent.
Postero-lateral corners of last prosome somite rounded ; 2 urosome somites posterior
to genital complex. Caudal rami about 2-2 times longer than broad, armed with a
lateral seta just proximal to the mid point of the ramus.

First antenna (Fig. 24b) y-segmented ; armature elements per segment 1-8,
II-I2, III-4, IV-3, V-4, VI-3, VII-7.

Second antenna (Fig. 24c). First segment with 2 minute spinules at inner distal
angle ; second segment with a single spinule on inner margin ; third segment short
with 3 slender setae at inner distal angle ; fourth segment with 4 long apical setae
and 2 shorter subapical setae. Segments 2 to 4 with dense patches of minute denticles
on their inner surface.

Mandible (Fig. 24d) produced into a slender lash with wide base ; armature
reduced to a single denticle on the outer margin and a short row of fine bristles near
the inner margin.

First maxilla (Fig. 246) bilobed ; inner lobe with i curved seta, outer with i short
and 2 long apical setae.

Second maxilla (Fig. 24f ) 2-segmented ; terminal segment forming a curved
element distally and with a proximal spine.

Maxilliped (Fig. 24g) 3-segmented, comprising an unarmed basal segment, a
robust middle segment with 2 stout denticulate setae, and a claw-like terminal
segment bearing a short seta proximally.

Legs 1-4 (Figs 24h-k) with 3-segmented rami ; endopod larger than exopod in all
legs ; armature formula as in generic diagnosis. Rows of setules present on the
inner margins of the basis and exopod segment i and on the lateral margins of endo-
pod segments i to 3.

Leg 5 a short conical process with 2 apical setae and a lateral seta located near its
base.

Leg 6 represented by 2 long plumose setae situated on the lateral margin of the
genital complex.

Body length of single female specimen 2-25 mm.

Male. Body similar to $ in general facies (Fig. 25a) ; urosome with 3 somites
posterior to genital complex ; minute denticles and sensory setules present on cara-
pace and prosome as in $. Anal somite with a discontinuous row of spinules on
ventral surface near the posterior margin ; caudal rami (Fig. 25c) as in <j> but pos-
sessing several spinules located on the distal portion of the ventral surface.

Appendages as in $ except for first and second antennae and maxilliped.

First antenna (Fig. 25b) y-segmented ; armature elements per segment 1-6,
II - 12, III - 5, IV - 3, V - 3, VI - 2, VII - 7 (armature incomplete).

Second antenna differing from that of $ in having only 3 apical setae on the ter-
minal segment instead of 4.

Maxilliped (Fig. 25d) 4-segmented ; comprising a broad basal segment, a second
segment with an inner proximal swelling armed with 2 setae and covered with fine
spinules, a small unarmed third segment and a fourth segment forming a curved claw
with a spatulate tip and armed with 2 unequal setae at its base.

Body length of single male specimen 2-20 mm.

O5mm

FIG. 25. Pachos punctatum. a, male entire, dorsal ; b, first antenna, ventral ; c, urosome,
ventral ; d, maxilliped, anterior. Pachos tuberosum. e, male entire, dorsal ; f, urosome,
ventral ; g, second antenna, anterior ; h, maxilliped, anterior ; i, mandible, anterior ;
j, first maxilla, posterior ; k, second maxilla, ventral.

OBSERVATIONS ON THE ONCAEIDAE 153

MATERIAL EXAMINED, i $ and i $ : from samples 25 and 23 respectively ;
BM(NH) registration numbers : 1976.226 and 1976.227.

REMARKS. The segmentation of the first antenna appears to be rather variable
in this species. Claus (1863) recorded 8 segments in the female whereas Giesbrecht
(1892) found 5 in the female and 7 in the male. This apparent variation is probably
due in part to the difficulty of determining the precise segmentation of the densely
setate proximal portion of the first antenna. Seven distinct segments were observed
in both sexes of P. punctatum although some degree of fusion was observed between
segments 3 and 4 in the female.

Pachos tuberosum (Giesbrecht, 1891)

Pachysoma tuberosum Giesbrecht, 1891 : 478 ; 1892 : 612, pi. 48, fig. 37.
Pachysoma dentatum Mori, 1932 : 172, 176, pi. V, figs 1-6 (male only).

DESCRIPTION. Male. Body globular (Fig. 256) as in P. punctatum ; prosome
inflated but more elongate than in $ P. punctatum ; rostrum very conspicuous from
dorsal aspect. Dorsal surface of carapace covered with a reticulate pattern of
minute chitinous ridges and with a pair of sensory setules. Posterior angles of last
prosome somite produced into slender processes ; anal somite bearing a row of
spinules on the ventral surface (Fig. 25f ). Caudal rami about 2-2 times longer than
broad ; lateral seta situated approximately a quarter of the distance along the
ramus ; several spinules present on the distal portion of the ventral surface.

First antenna 7-segmented, armature similar to that of <$ P. punctatum.

Second antenna (Fig. 25g) comprising a basal segment with an inner distal
spinule, second segment with a mid inner margin spinule and an outer patch of
minute denticles, third segment with 2 setae at the inner distal corner and denticles
on the outer surface, fourth segment with i stout and 3 slender apical setae, 2 sub-
apical setae and a zone of minute denticles on the outer surface.

Mandible (Fig. 25!) produced into a slender lash with 4 denticles on the outer
margin and a row of fine bristles near the inner margin.

First maxilla (Fig. 25]) bilobed ; inner lobe apparently unarmed, outer lobe with
3 setae.

Second maxilla (Fig. 25k) 2-segmented ; basal segment with a large lobe pos-
teriorly ; terminal segment tapering into a claw and armed with a single proximal
spine.

Maxilliped (Fig. 25h). Second segment with an inner proximal swelling covered
with fine spinules and bearing 2 setae ; terminal segment forming a claw with a
pointed tip and with a short straight seta and a curved seta proximally.

Legs 1-6 as in P. punctatum.

Body length of single male specimen 2-00 mm.

MATERIAL EXAMINED, i <J from sample 27 ; BM(NH) registration number :
1976.228.

REMARKS. There are no significant differences between the male of P. dentatum
described by Mori (1932 & 1937) and the male of P. tuberosum. Mori found only i

I54 G. A. BOXSHALL

stout and 4 slender setae around the apex of the second antenna of male P. dentatum
but the absence of a small seta from the subapical group (cf. Fig. 25g) does not
justify specific recognition especially when the seta is easily overlooked. In the
armature of the other appendages, in body proportions and in the surface orna-
mentation of the carapace male P. dentatum very closely resembles P. tuberosum.

ACKNOWLEDGEMENTS

I am most grateful to Dr H. S. J. Roe (I.O.S.) for arranging the original loan of
material and for supplying other relevant data. I would also like to thank Miss
A. R. Gurney for assistance with sorting and counting of the samples and both Dr
J. P. Harding and Dr R. J. Lincoln for reading and commenting on the manuscript.

REFERENCES

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1+8) and further developments of rectangular midwater trawls. /. mar. biol. Ass. U.K.

53 : 167-184.
BJORNBERG, T. K. S. 1973. The planktonic copepods of the Marchile I expedition and of the

'Eltanin' cruises 3-6 taken in the SE Pacific. Bol. Zool. e Biol. Mar. N.S. 30 : 245-394.
BRADY, G. S. 1883. Report on the Copepoda collected by H.M.S. 'Challenger' during the

years 1873-76. Rep. scient. Results Voy. Challenger, Zool. 8 : 1-142.
CLAUS, C. 1863. Die Frei Lebenden Copepoden. Leipzig. 230 pp.
DAHL, M. 1912. Die Copepoden der Plankton-Expedition. I. Die Corycaeinen. Ergebn.

Plankton-Exped. 2 : 1-135.
DANA, J. D. 1846. Notice of some genera of Cyclopacea. Am. J. Sci. ser. 2, 1 : 225-230.

1852. Crustacea. U.S. Explor. Exped. 13 : 1-1618.

ESTRADA, J. C. & GENICIO, M. F. 1970. Nota sobre el plancton de la costa noroccidental

Africana. Boln. Inst. esp. Oceanogr. 140 : 1-38.
FARRAN, G. P. 1908. Second report on the Copepoda of the Irish Atlantic slope. Scient.

Invest. Fish. Brch. Ire. 1906, 2 : 3-104.

1929. Copepoda. Nat. Hist. Rep. Br. antarct. Terra Nova Exped. (Zool.), 8 : 203-306.

— 1936. Copepoda. Scient. Rep. Gt. Barrier Reef Exped. 6(3) : 73-142.
FERRARI, F. D. 1975. Taxonomic notes of the genus Oncaea (Copepoda : Cyclopoida) from the

Gulf of Mexico and northern Caribbean Sea. Proc. biol. Soc. Wash. 88, No. 21 : 217-232.
GIESBRECHT, W. 1891. Elenco dei Copepodi pelagici raccolti dal tenente di vascello Gaetano

Chierchia durante il viaggio della R. Corvetta 'Vettor Pisani' negli anni 1882-1885, e dal

tenente di vascello Francesco Orsini nel Mar Rosso, nel 1884. Alt Accad. naz. Lincei Re.

7(io) : 474-481.
- 1892. Systematik und Faunistik des pelagischen Copepoden des Golfes von Neapel und

der angrenzenden Meeresabschnitte. Fauna Flora Golfo Napoli, 19 : 1-831.
GORDEYEVA, K. T. 1975- A new species of the genus Oncaea (Copepoda) from the bathypela-

gial in the tropic zone of the Atlantic and Gulf of Mexico. Zool. Zh. 54(g) : 1397- 1399-

(In Russian.)
HERON, G. A. & DAMKAER, D. M. 1969. Five species of Deep-water Cyclopoid Copepods from

the Plankton of the Gulf of Alaska. Smithsonian Contr. Zool. 20 : 1-24.
LEHNHOFER, K. 1926. Copilia Dana, 1849 der deutschen Tiefsee Expedition. Wiss. Ergebn.

dt. Tiefsee-Exped. 'Valdivia', 23(3) : II3~I77-
1929. Copepoda 5 : Sapphirina J. V. Thompson 1829. Wiss. Ergebn. dt. Tiefsee-Exped.

'Valdivia', 22(5) : 269-347.

OBSERVATIONS ON THE ONCAEIDAE 155

MORI, T. 1932. Honpo Nankai ni oite saishusita Fuyu-Tokyakurui no shinshu. Dobutsugaku

Zasshi, 44, No. 523.
1937- The pelagic Copepoda from the neighbouring waters of Japan. Tokyo. 145 pp.

(Reprinted 1964.)
MOULTON, T. 1973. Principal component analysis of variation in form within Oncaea conifer a

Giesbrecht, 1891, a species of copepod (Crustacea). Syst. Zool. 22(2) : 141-156.
OLSON, F. 1949. The pelagic cyclopoid copepods of the coastal waters of Oregon, California

and Lower California. Ph.D. Thesis, University of California. 208 pp.
PHILIPPI, A. 1843. Fernere Beobachtungen iiber die Copepoden des Mittelmeeres. Arch.

Naturgesch. 9(i) : 54-71.
RAZOULS, C. 1969. Description d'une espece nouvelle du genre Oncaea (Copepoda, Cyclo-

poida). Vie Milieu, 20(2B) : 317-324.
1974- Les Oncaeidae (Copepoda, Cyclopoidea) de la region de Banyuls (Golfe du Lion).

Vie Milieu, 24(2 A) : 235-264.

ROSE, M. 1933. Copepodes pelagiques. Faune Fr. 26 : 1-374.
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23: i -58.

SARS, G. O. 1900. Crustacea. Scient. Results Norw. N. polar Exped. 1893-1896, 1(5) : 1-141.
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pendant les campagnes des S.A.S. le Prince Albert de Monaco, avec descriptions et figures

des especes nouvelles. Bull. Inst. oceanogr. Monaco, 323 : 1-15.
1918. An account of the Crustacea of Norway. VI. Copepoda Cyclopoida. Parts 13- 14:

173-225. Bergen Museum.
SCOTT, T. 1894. Report on Entomostraca from the Gulf of Guinea, collected by John Rattray,

B.Sc. Trans. Linn. Soc. Lond. Zool. ser. 2, 6(1) : i — 161.
SEWELL, R. B. SEYMOUR. 1947. The free-swimming planktonic Copepoda. Systematic

account. Scient. Rep. John Murray Exped. (Zool.), 8(1) : 1-303.
SHMELEVA, A. A. 1966. New species of the genus Oncaea (Copepoda, Cyclopoida) from the

Adriatic Sea. Zool. Zh. 45(6) : 932-936. (In Russian.)
1967. New Oncaea species (Copepoda, Cyclopoida) from south-western part of the

Atlantic Ocean. Zool. Zh. 46(4) : 621-622. (In Russian.)
1969. Especes nouvelles du genre Oncaea (Copepoda, Cyclopoida) de la mer Adriatique.

Bull. Inst. oceanogr. Monaco, 68, No. 1393 : 1-28.

TANAKA, O. 1960. Pelagic Copepoda. Spec. Publs. Seto mar. biol. Lab. 10 : 1-95.
WILSON, C. B. 1950. Copepods gathered by the United States fisheries steamer 'Albatross'

from 1887 to 1909, chiefly in the Pacific Ocean. Bull. U.S. natn. Mus. 100, 14 : 141-441.
WOLFENDEN, R. N. 1905. Plankton Studies, preliminary notes upon new or interesting species.

Part i. Copepoda. London & New York. 24 pp.

DR G. A. BOXSHALL

Department of Zoology

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NOTES ON DEEP-WATER ATLANTIC

CRINOIDEA

A. M. CLARK

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 31 No. 4

LONDON: 1977

<«/,

NOTES ON DEEP-WATER ATLANTIC ^ ^

CRINOIDEA

BY

AILSA M. CLARK

Pp. 157-186 ; 5 Text-figures

BULLETIN OF

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NOTES ON DEEP-WATER ATLANTIC CRINOIDEA

By AILSA M. CLARK

SYNOPSIS

Notes are given on some little-known species of the stalked crinoid family Bathycrinidae
from the north, equatorial and south-west Atlantic, including Bathycrinus aldrichianus, B.
australis and the type-species, B. gracilis, collected by the U.S. research vessels Atlantis II and
Chain, and Democrinus parfaiti, collected by the R.R.S. Shackleton. In addition a corrected
type citation for Bathycrinus carpenteri is given and notes on the type material of Democrinus
brevis in the British Museum collections. A post-pentacrinoid of Atelecrinus is figured and new
records cited for a few other species of unstalked crinoids, including the deepest to date for the
Mediterranean species Leptometra phalangium.

INTRODUCTION

THE small but interesting collection of crinoids which forms the basis for much of this
paper was collected by the Wood's Hole Oceanographic Institution. It was for-
warded to the British Museum by Dr D. L. Pawson of the Smithsonian Institution
who received it from Dr H. Sanders of Wood's Hole. The Shackleton material of
Democrinus parfaiti was presented by Dr J. D. George of this museum. I am
indebted to all of them.

About two-thirds of the American material is being returned to the Smithsonian,
the remainder being retained for the British Museum collections.

Unfortunately the small size and vulnerability of most of the crinoids concerned
prevented positive specific identifications of some specimens. However, the
material did allow some studies on the early growth stages. Most important, it
includes two mature crowns of Bathycrinus gracilis Wyville Thomson - the type-
species of the genus, described from an immature specimen which has been lost.

DESCRIPTION OF SPECIES

Order MILLERICRINIDA

Family BATHYCRINIDAE

Bathycrinus aldrichianus Wyville Thomson

(Fig- ig)

Bathycrinus aldrichianus Wyville Thomson, 1876 : 47-51, fig. i ; 1877 : 92-95, fig. 23 ; Gisl6n,
1938 : 15-16; A. H. Clark, 1949: 377; Gisl6n, 1951: 51-53; Macurda & Meyer, 1976:
647-666.*

Bathycrinus campbellianus P. H. Carpenter, 1884 : 238-240, fig. 15.
Bathycrinus serratus A. H. Clark, 1908 : 205-207, fig. i.

MATERIAL. Challenger st. 106, oi°47'N, 24°26'W (about midway between Cape
Verde and the NE. corner of Brazil), 3382 metres ; the holotype (crown only)
BM (NH) reg. no. 85.3.30.34.

* Thanks to Dr Macurda I have seen a typescript of this paper, which records new material of B.
aldrichianus from oo°i2'N, o5°u'E, in the Gulf of Guinea at 3595 metres, including several adult speci-
mens with the basal ring and upper part of the stalk still attached to the crown. The authors give
a detailed description, mainly of the fine structure of the articulations, as studied by SEM.

160 AILSA M. CLARK

Atlantis II cruise 60, st. 25gA & D, 37°i3'S, 52°45'W (off Argentina, SE. of Rio
de la Plata), 3305-3317 metres ; 2 crowns.

Chain cruise 50, st. 85, 37°59.2'N, 6g°26.2'W (SE. from New York), 3834 metres ;
i small armless specimen.

The specimen with basal ring and at least the proximal part of the stalk (Fig. ig)
is very small and has broken above the radials which themselves appear to be only
partially regenerated. It has the five uppermost columnals discoidal (height not
more than a sixth of the breadth). Macurda & Meyer (1976) note that there are
'at least 5 but usually 12 or more short proximal columnals . . . with a synostosial
articulation between them.' Clearly, Carpenter (1884) was right to reject as genuine
the long-segmented column drawn juxtaposed to the crown of the holotype by
Wyville Thomson's artist.

Like the holotype, both crowns from the Atlantis II have strongly grooved sur-
faces to the division series and brachials and all these ossicles are markedly flared
at their distal ends producing a very serrated profile. The surface of the radials
has a fingerprint-like texture with much finer grooving, similar to that of B. gracilis.
Even the syzygial joints have their proximal elements flared so the syzygies (or
cryptosynarthries according to Macurda & Meyer) are not easy to distinguish. The
first three brachial ones are usually at 1 + 2, 4 + 5 and 6 + 7 but a total of four
(possibly five) arms out of twenty have the second syzygy at 3 + 4, as in Monacho-
crinus. Conversely, the distal joint of Br6 is muscular on six arms. Because of
this variation, the position and identity of the first pinnule vary considerably. On
one specimen it is P3 in every case but on six arms it arises on Br9 and on three on
Br8. The second specimen has P3 present only on three arms, once on Br9 and twice
on Br10 (when five syzygies precede it) ; otherwise the first pinnule is Pc, usually on
Br10 (preceded by only four syzygies) but once on Bru. On both specimens the
first pinnules have 10 or n segments and measure 4*2 -4-5 mm. The longer pin-
nules, such as P5, have 15 or 16 segments and measure 6-0-6-5 mm-

Other numerical data for these specimens are given in Table i with the values for
the holotype, which is of similar size and has the first pinnule Pc or P3 on Br9 or Br8.
It is therefore surprising that Gislen's much larger Swedish Deep-Sea Expedition
specimen (1951) with the crown 56 mm long has the first pinnule on Bru. (This is
P4 since Br9 as well as Br3 and Br6 have muscular joints at both ends.) It is
possible that there is not a progressive development of the more proximal pinnules
during growth in this species, as there appears to be in Democrinus parfaiti.

AFFINITIES. As mentioned under B. gracilis, both that species and B. aldrichianus
have very serrated profiles to the crowns. The ornamentation of the ossicles of
B. gracilis is more restricted to a central keel, with only traces of other fluting, but
this could be correlated with the somewhat smaller size. Possibly further material
may indicate that only one species can be recognized.

DISTRIBUTION. The Atlantis II record provides an extension of range to the south-
west Atlantic off northern Argentina from the equatorial Atlantic and overlaps with
the new extension to the range of the much larger smooth species of southern
Bathycrinus, B. australis A. H. Clark. Other records are off the United States,

DEEP ATLANTIC CRINOIDEA 161

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Atlantis II st. 247
Maximum heights c
'Br' signifies brachi;

162 AILSA M. CLARK

between there and the Azores and off the Cape Verde Islands. Bathymetric dis-
tribution 3317-5600 metres.

Bathycrinus australis A. H. Clark

Bathycrinus aldrichianus : P. H. Carpenter, 1884:241-243, pi. 7, pi. ya, figs 1-21, pi. yb,
pi. 8a, figs 4, 5 ; Gislen, 1928 : 14. [Non B. aldrichianus Wyville Thomson, 1876.]

Bathycrinus australis A. H. Clark, 1907 : 553-554 ; Doderlein, 1912 : 9-10, pi. 5, fig. i, pi. 6,
fig. 7 ; Gislen, 1938 : 16 ; 1956 : 61-62, pi. i, figs 1-6.

Ilycrinus australis : A. H. Clark, 19155 : 154-155.

MATERIAL. Challenger st. 146, 46°i6'S, 48°27'E (W. of the Crozet Islands,
Southern Ocean), 2515 metres ; 2 syntypes, BM(NH) reg. no. 85.2.30.3, 4.

Atlantis II cruise 60, st. 245A & B, 36°557'S, 53°oi-4'W (off Rio de la Plata,
Argentina), 2707 metres ; i specimen lacking most of stalk, i crown and a length of
stalk.

Atlantis II cruise 60, st. 247A & B, 43°33'S, 48°58-i'W, 5208-5223 metres, 8
specimens with stalk and arms more or less incomplete, 19 crowns, 2 lengths of stalk.

Some numerical data from the more nearly intact of these specimens is given in
Table i. No Atlantis II specimens retain the distal end of the stalk. The syntypes
retain stumps of the main branches, as shown in Carpenter's pi. 7 (1884) and Gislen's
pi. i (1956) with stout cylindrical segments and no radicular cirri arising above the
main dichotomy. One syntype has several successive very short (but not quite
discoidal) columnals just above the dichotomy. The form of 'rooting' is similar to
that in the northern species Bathycrinus carpenteri (Danielssen & Koren).

The longest detached piece of stalk measures c. 100 mm and has 44 columnals, the
uppermost ones short but not discoidal so that the break probably came between
the fifteenth and twentieth segments. The breadth at the top is 1-6 mm. As the
columnals elongate distally they also become slightly constricted medially (a dice-
box-shape according to Carpenter). The longest columnal measures 3-3/1-7 mm
length/median breadth (1-9/1) with the maximum breadth at the joints up to 2-0 mm.
A more distal piece of another stalk 80 mm long with 37 columnals, similar to the
first in the proportions of its middle segments, has the distalmost columnals remaining
(probably to close above the main division) much stouter and shorter, the measure-
ments 2-75/2-2/3-0 mm ; their successive joints are 'screwed'.

Gislen (1956) gives the proportions of the longer columnals in a detached piece of
stalk as 3/0-8/1 mm, which is relatively longer than in these specimens. In the
syntype measured the columnals are also relatively slender with length/median
breadth up to 2-7/1.

Specimen 14 in Table i from st. 245 is somewhat smaller than those from the more
southern station 247 as regards the stalk and calyx but has relatively large division
series. This is also true of the broken crown from st. 245, where the division series
are c. 6-0 mm long at a radial height of only 1-8 mm and upper breadth of 4-8 mm.
Another difference in the first specimen is the more numerous discoidal proximal
columnals, about 18 as opposed to 9-13, with the first columnal to be higher than
broad not until about the thirty-fifth, rather than about the twenty-seventh.

DEEP ATLANTIC CRINOIDEA 163

There is also one much smaller crown from st. 247 with the height of the radials
only 1-35 mm (as opposed to 2-8-3-8 mm in the others from that station measured).
The lower and upper breadths of its radial ring are 0-7 and 1-9 mm, so the height is
relatively greater in comparison with the larger specimens, 71% of the upper breadth
as opposed to 48-61% (mean 55%). This small specimen, like the syntypes, also
differs in having the lateral flange on the division series and proximal brachials (in
this case restricted to the first four or five) relatively thin and the arms are markedly
convex medially, from Br5 with a blunt keel. The larger specimens have the arms
almost flat on top proximally, though beyond the bases they become somewhat
compressed laterally. Their lateral flanges are blunter and extend as far as about
Brn. As usual, syzygies in the small specimen are proximally at 1 + 2, 4 + 5 and
7 + 8 ; all are broken by Br10. Two arms at least have the first pinnule (Pb) on
Br6 ; on the other arms it is P3 on Br8 (as in the larger specimens from this station)
and measures 6-5-7-0 mm in length. Only the specimen from st. 245 agrees with
the syntypes in having the first pinnule usually on Brn or 12. According to Gislen
(1956) pinnules begin distal to Brn in his material from the Kermadec Trench.

An intact arm of specimen 6 in the table is 60 mm long and has 108 brachials.
The frequency of syzygies on this arm decreases distally so that beyond c, Br40 only
one out of every five joints is usually a syzygy. This compares with B. aldrichianus
where syzygies tend to alternate with muscular joints throughout the arm beyond the
base.

A broken calyx shows four vertical cracks in the basal ring, of which three corre-
spond to the interbasal (BB) sutures ; these are otherwise not distinct.

The suture between the basal ring and the uppermost columnal is indistinct in all
the specimens. Only in the syntype shown in Carpenter's pi. 7, fig. 2 is the basal
ring so abruptly marked off from the radial ring by slightly swollen contours ad-
joining the suture ; otherwise the transition is fairly smooth.

AFFINITIES. This relatively large and smooth species of Bathycrinus closely
resembles B. carpenteri (Danielssen & Koren) from the deep basin of the Norwegian
Sea (see A. M. Clark, 1970), though considerably exceeding the northern species in
size - the largest B. carpenteri described having crowns up to c. 35 mm in length,
compared with nearly 70 mm in the present material. The division series and proxi-
mal parts of the arms in B. carpenteri are more convex but this may be correlated
with the smaller size.

Bathycrinus australocrucis McKnight (1973) from New Zealand is another rela-
tively smooth species, but has median keels on the arms from Br4 and beyond the
arm bases these are flared distally so as to produce a serrated profile, though much
less so than in B. aldrichianus. It also differs in the fewer short proximal columnals,
of which the twelfth is already longer than broad. The crown in the holotype is
probably about 20 mm long.

B. australis, with its flared radial ring, also has some resemblance to Zeuctocrinus,
from the North Atlantic, which I described and referred to the Phrynocrinidae in
1973, particularly to the young specimen shown in fig. 5c (Clark, 1973) where the
uppermost columnals are discoidal, unlike the condition in larger specimens. Phryno-
crinids have the calyx more integrated and compact and the bases of the division

164 AILSA M. CLARK

series are well separated laterally. In addition their stalk attachment is by an
expanded disc.

DISTRIBUTION. The present records extend the range of B. australis to the deep
Argentinian plateau of the SW. Atlantic. The other records are widely scattered
around the southern hemisphere from the vicinity of the Crozet Islands (c. 46°S,
48°E - the type locality), off Enderby Land, Antarctica (c. 63°S, 58°E) and around
New Zealand (c. 35°S, i7g°W and 38°S, i7i°E). The depth range is 1748 (?i73o) -
8210 (P8300) metres.

Bathycrinus carpenteri (Danielssen & Koren)

Ilycrinus carpenteri Danielssen & Koren, 1877 : 45-56, pis i, 2.

Bathycrinus carpenteri : Danielssen, 1892 : 1-28, 5 pis ; A. M. Clark, 1970 : 14-18, fig. 3.

Bathycrinus carpenterii : GislSn, 1938 : 16-17.

TYPE DESIGNATION. Due to incorrect labelling as Types' of two specimens in the
Oslo Museum collection from Norwegian North- Atlantic Expedition st. 295, my
citation of these as syntypes (1970 : 14) has to be emended. Dr Holthuis has
pointed out to Dr Marit Christiansen of the Oslo Museum that this sample dates
from 1878 and only material from N.N.E. stations 35, 40, 51 and 53 was available to
Danielssen & Koren in 1877. Since that paper was in Norwegian I had relied upon
Danielssen's of 1892 with its English translation.

Although the entire holdings of crinoids in the Oslo, Bergen, Trondheim and
Troms0 Museums were sent to me (as I understand), from these four stations only
the crown and stalk from st. 35, Bergen Museum no. 1159, were included, so that the
fate of the best eligible specimen for selection as lectotype, from st. 51 is unknown.
The other material from these early stations consisted of stalks or broken specimens.
I therefore designate as lectotype the crown and stalk from st. 35, 63°22'N, i°2o'W
(between Norway and Iceland, NE. from the Faeroes), 1920 metres, in the Bergen
Museum.

Bathycrinus gracilis Wyville Thomson
(Fig. za-f)

Bathycrinus gracilis Wyville Thomson, 1872 : 772-773 ; 1873 : 450-454, fig. 73 ; P. H. Car-
penter, 1884 : 243-245, fig. 16, pi. 8a, figs 1-3 ; Koehler, 1909 : 254 ; Gisl6n, 1938 : 18.

MATERIAL. Chain cruise 106, st. 328, 5o°04'7'N, i5°44-8'W (c. 200 miles (320 km)
WSW. of Cape Clear, SW. Ireland), 4426-4435 metres ; 2 crowns, I calyx with costals
and proximal part of stalk, 3 juveniles, 8 pentacrinoids.

Porcupine 1869, st. 37, 47°38'N, i2°o8'W (c. 200 miles (320 km) S. of Cape Clear),
4450 metres ; incomplete paratype, BM(NH) reg. no. 85.3.30.33.

Only three specimens of this, the type species of Bathycrinus, have so far been
recorded (Perrier's from off Morocco having been referred to a distinct species,
B. perrieri, by Koehler & Vaney in 1910). According to Carpenter (1884), the
immature holotype had the whole crown only 8 mm high with just 12 brachials and
only rudiments of pinnules developing at the tips. Unfortunately its whereabouts

DEEP ATLANTIC CRINOIDEA

165

H

FIG. i. a-f. Bathycrinus gracilis Wyville Thomson, a. Paratype, basal ring and proximal
end of stalk [badly decalcified], b-f. Chain st. 328. b. Crown, c. Detail of pinnule,
d. Specimen broken after first post-radial ossicle (IBr^, which itself with the radials is
probably incompletely regenerated, e. Very young specimen with 'integrated' calyx
and only two brachials of each arm yet developed, f. Pentacrinoid at 'cystoid' stage
with rudimentary radials not yet obscuring the oral plates, g. Bathycrinus aldrichianus
Wyville Thomson. Chain st. 85, juvenile specimen apparently broken above the radials
revealing the orals. All viewed radially. [The scales measure i mm ; the small one
applies to b and c and the large one to the remainder.]

are unknown. A crownless paratype in the British Museum collection consists of
c. 60 mm length of stalk topped by the basal ring ; it is now badly decalcified and
broken into three pieces. Both specimens were taken at a single Porcupine station
c. 200 miles (320 km) S. of Cape Clear, SW. Ireland. The third known specimen, a
stalkless crown only slightly larger than the holotype, measuring n mm, was col-
lected by the Princesse Alice between the Azores and Portugal and was described by
Koehler (1909) simply as conforming with the holotype.

The paratype (Fig. la) has the upper side of the basal ring slightly domed ; the
height of the exposed part is 0-35 mm, the maximum (upper) diameter 0-60 mm and
the lower diameter 0-45 mm. The individual basals are slightly convex above and
faint interbasal sutures can just be discerned when partially dried. The top 13
columnals can be described as discoidal (i.e. with 1/br less than 1/6). The stalk
diameter tapers to a minimum of 0-3 mm at about the fifteenth columnal. The
seventeenth is the first with length exceeding breadth and the following columnals
become very attenuated, up to 3 mm long, with I/median br c. 6/1 and the joints
markedly broadened, the successive ones elliptical in almost alternating planes.
The columnals have a very fine fingerprint-like texture of fine longitudinal grooves,
while the basal ring appears superficially granular but these effects are likely to be

166 AILSA M. CLARK

inherent in the plate structure and a reflection only of the considerable magnification
used, and as such probably without taxonomic significance.

The two crowns taken by the Chain probably had slightly larger basal rings since
the lower diameter of the radial ring is 0-75 and 0-80 mm. Some measurements of
them are given in Table i. One specimen (Fig. ib) has the radials distinctly concave
in profile, whereas in the other they are straight. The individual radials are convex
in cross-section but not keeled ; their surface is finely 'fingerprinted' with longitudinal
grooves. This texture is not continued on the division series which have a sharp
median hyaline crest and thin, sharp-edged lateral flanges. On the IBr2 (axillary)
the median crest is Y-shaped, matching up distally with the crests on the proximal
brachials. From about Br3 additional crests appear at the distal end of each brachial
beside the median one, producing a fluted median area. The distal ends of the
brachials from Br2 onwards are more or less flared so as to make the profile distinctly
serrated. Lateral flanges extend on the arms to c. Br10. Syzygies are present at
i + 2, 4 + 5, 7 + 8, 10 + ii and then alternately (i.e. only brachials 3, 6 and often also 9
have muscular joints at both ends). The first pinnule, Pc, is on Br10 (or Bru). The
gonads are visible by transparency. The longer pinnules have six or sometimes seven
segments.

Clearly Bathycrinus gracilis shares the vulnerability of the calyx to break above
the basal ring shown by other species of the genus. Regeneration of the crown is
probably common and seems to have occurred in the specimen with the stalk and
basal ring (Fig. id) since its radials are abnormally small and their lower ends are
inset so that the diameter of the lower end of the radial ring is less than that of the
upper end of the basal ring (0-65 mm) . The basals lower diameter is 0-60 mm and the
height 0-55 mm, so the relative height is greater but the expansion of the top of the
ring less than in the paratype. No interbasal sutures can be seen. The maximum
upper diameter of the radial ring is only 0-85 mm and above this only one ossicle,
the IBrx, remains in each radius ; these costals are relatively short and are spaced
laterally. Both radials and costals have a sharp median crest. The basal ring has a
granular texture but the radials show a suggestion of 'fingerprinting'. The diameter
at the top of the stalk is 0-60 mm (compared with 0-45 mm in the paratype) and there
are 16 discoidal proximal columnals. The first columnal to be longer than broad
is the twenty-first. The stalk diameter reduces to 0-45 mm at c. 10 mm from the
top. The succeeding columnals become very elongated with 1/med. br c. 2-0/0-35 mm
(5-5/1). The joints are expanded elliptically in almost alternating planes.
Columnals 21-29 have a slightly projecting median belt around them. The stalk is
broken at the forty-fifth columnal and the total length measures 42 mm ; at a thick-
ness of less then 0-5 mm the whole specimen is thread-like.

It is to be expected that very young specimens of Bathycrinus would have the
individual basals still separate, as in the related genus Monachocrinus, which is
neotenous in this respect. This theory is borne out by two little specimens from the
same Chain station with only two brachials on each arm yet developed (Fig. le). In
these the basal and radial rings are integrated into a cone and all the sutures between
them are distinct. The radials are distinguished by their hyaline median crest-like
keels matching those on the division series and first brachials. Unfortunately only

DEEP ATLANTIC CRINOIDEA 167

a few discoidal proximal columnals remain attached to both these specimens. There
are also present several smaller ones which may be classed as pentacrinoid larvae in
the 'cystoid' stage, with radials undeveloped and the orals (of similar size to the
basals) exposed. Yet other specimens have narrow radials and laterally separated
post-radial series developing.

Also included in this sample is a relatively large very zig-zag segmented fragment
of the distal end of a stalk, with radicular cirri branching off it, often at acute reflexed
angles. This resembles the stalk attachments of Bathycrinus nodipes Doderlein,
from the East Indies (1907, pi. 5, figs 2, 3) and the size is commensurate with the two
crowns, to one of which it presumably belonged ; sadly the intervening column and
basal ring are missing.

The other Atlantic species of Bathycrinus currently recognized are B. carpenteri
(Danielssen & Koren) from the deep basin of the Norwegian Sea and B. aldrichianus
Wyville Thomson recorded from the Gulf of Guinea, equatorial mid-Atlantic and
east of the U.S.A. (B. perrieri Koehler & Vaney having been referred to
Monachocrinus since it has the interbasal sutures distinct) . In addition, the present
collection extends the range of B. australis A. H. Clark, previously known from
various localities in the Southern Ocean and South Pacific, to the South Atlantic off
Argentina. Both B. carpenteri and B. australis have notably smooth rounded ossicles
and profiles but B. aldrichianus resembles these two crowns of B. gracilis in being
similarly very serrated in profile. However, the holotype of B. aldrichianus has the
median parts of the ossicles more rounded, except on the middle part of each division
series where the convexity is constricted to a narrow median blunt ridge. Gislen
(1951) described a crown of B. aldrichianus as much as 56 mm long from off the Cape
Verde Islands in which the division series have a 'rounded, narrow and indistinct
median dorsal ridge'. It is possible that the sharpness of the crests in B. gracilis is
correlated with the relatively small size.

DISTRIBUTION. SW. of Ireland to NE. of the Azores (c. 39°N, 2i°W), 4435 (?4426)
- 5005 metres.

Democrinus Perrier

Democrinus Perrier, 1883 : 450 ; A. H. Clark, 1917 : 392 ; Gislen, 1938 : 25. Type species

D. parfaiti Perrier, 1883.

Rhizocrinus (pt) : Carpenter, 1884 : 245 ; A. H. Clark, 1909 : 673-676.
Rhizocrinus (Bythocrinus) Doderlein, 1912 : n [No type designated for subgenus.]
Bythocrinus : A. H. Clark, 1917 : 392.

Seven nominal species of five-armed Atlantic bathycrinids with at least the inter-
basal sutures of the calyx distinguishable have been referred to Democrinus, namely :

Democrinus parfaiti Perrier, 1883, from Morocco

Rhizocrinus rawsoni Pourtales, 1874, Barbados

R. conifer A. H. Clark, 1909, Brazil

R. brevis A. H. Clark, 1909, Atlantic side of Panama [and ? Leeward Is]

R. sabae A. H. Clark, 1909, Leeward Is

168 AILSA M. CLARK

R. robustus A. H. Clark, 1909, W. of Florida

Bythocrinus intermedius A. H. Clark, 1915 (a), Gulf of Mexico

The first two were treated as synonymous by Carpenter (1884) but D. parfaiti was
revived by A. H. Clark (1909) when he named and partially described four other
related nominal species. Of these, Gislen (1938) referred R. sabae to the synonymy
of D. rawsoni and R. robustus to the synonymy of D. conifer, adding to the latter
(at A. H. Clark's own suggestion) also B. intermedius (a name based only on a drawing
of part of the stalk and a locality given in the caption to this). The calyx of D.
conifer was figured by A. H. Clark (igisa) and those of the holotype and a paratype
of D. brevis had been given by Carpenter (1884) but no representations of the calyces
of R. sabae, R. robustus or B. intermedius are extant. A. H. Clark's (1909) descrip-
tions of the first two are hardly explicit but diagrammatic reconstructions as shown
in Fig. 2e and f can be made from them. The resultant shapes support Gislen's
conclusions. Following study of a new sample of D. parfaiti, described below, I
think that calyx shape may be used as a specific character for some species of
Democrinus, provided the growth stage is taken into consideration. This sample
also includes distal ends of some stalks showing a slender branching root-like form of
cylindrical radicular cirri (Fig. 3b), as in Bathycrinus carpenteri (Danielssen &
Koren). This contrasts with the attachment found in D. brevis (Fig. 2d) by an
expanded, irregularly lobed plate more or less appressed to and fused with the
substrate, there being only a few radicular cirri arising from the lower columnals.
In 1973 I established a new family Porphyrocrinidae for Porphyrocrinus Gislen and
Naumachocrinus A. H. Clark, which resemble bathycrinids in most respects but are
attached by a simple expansion of the distal end of the stalk moulded to a solid
piece of substrate. The attachment in Democrinus brevis and probably also in
D. rawsoni appears transitional to this and casts doubt on the one hand on the
wisdom of making a family distinction on this character alone and on the other of
regarding D. brevis and D. rawsoni as congeneric with D. parfaiti. Knowledge of the
stalk attachments of a wider range of species of bathycrinids than we now have is
needed to settle this problem.

Unfortunately, from the above list material only of D. parfaiti and Z). brevis is
available to me at present and a revision of the Atlantic Democrinus specimens in the
U.S. National Museum is badly needed, particularly the types of D. rawsoni, about
which only some comments derived from Pourtales (1874) can now be given.

Democrinus brevis (A. H. Clark)
(Fig. 2a-d)

Rhizocrinus rawsoni : Carpenter, 1884 (pt) : 263 (Investigator records), 266-267, fig- 19 '• A.

Agassiz, 1892 : 62 (footnote).
Rhizocrinus brevis A. H. Clark, 1909 : 675.
Bythocrinus brevis : A. H. Clark, 1917 : 392.
Democrinus rawsoni : Gislen, 1928 : 14-15.
Democrinus brevis : Gislen, 1938 : 26.

MATERIAL. Investigator, Capt. Cole : 'Off Colon' [altered by Bell from 'off
Panama' following query by Agassiz], '15 miles N. by E. of Panama [i.e. Colon on

DEEP ATLANTIC CRINOIDEA 169

the Atlantic side], 'Saba Bank, 15 miles N. by E. of Panama' and 'Saba Bank,
Panama', 550 metres ; 4 specimens, BM(NH) reg. nos 84.6.20.8 and 9, 1935.3.15.1
and 1949.4.5.1. The two last specimens were transferred to the Zoology Department
from the Geology Department about 1935 but one was put on exhibit until 1948.
Carpenter cited the Investigator records as 'Saba Bank, 200 fathoms. 15 miles N. by
E. of Panama, 300 fathoms' but the two appear to have been confused in labelling
and relabelling the last two specimens. I can find no Saba Bank in the vicinity of
Colon and it must presumably refer to the one in the Leeward Islands near St Kitts.
The first specimen is the one shown in Carpenter's fig. igA and was designated as
(holo)type by A. H. Clark ; the rest may be treated as paratypes since all four were
evidently studied by him.

Cuba (no details), M. H. Gray ; 8 specimens, no. 1900.5.17.1-3.

DESCRIPTION. Some numerical details of these specimens are given in Table 2.

The holotype is badly broken. Four columnals are still attached to the calyx ;
the uppermost one only is discoidal, the length increasing so that the fourth is longer
than broad (Carpenter's figure being incorrect in showing two non-existent discoidal
columnals in place of the lower end of the basals below the constriction in the calyx).
The arms also have broken off, mostly at the 3 + 4 syzygy. They are slightly
reduced in size beyond the first brachial (Br^ and may have incompletely regenerated.
The detached distal end of the stalk is still present with three very squat 'screwed'
columnals above a flared expansion of branching irregular lobes attached to a piece
of black substrate resembling solidified mud with a fibrous backing. Two cylindrical
radicular cirri appear to have broken off from one side of the upper end of the
distalmost columnal. Two parts of the stalk contribute to a total of only 38
columnals so there are probably other parts missing. The calyx (Fig. 2a) expands
fairly abruptly between a quarter and a third of its height above the topmost
columnal and there is a second but lesser expansion in the radials, so as to create a
slight hollowing in the vicinity of the basiradial (BR) suture, which forms a very
shallow zig-zag around the calyx and is clearly distinguishable, as are the interbasal
(BB) and interradial (RR) sutures.

The paratype, no. 84.6.20.9, is in much better condition, though the distal end of
the stalk is lost. As Carpenter's fig. 196 shows, the calyx is more evenly conical
than in the holotype. Also the proximal brachials are rather irregular, some ossicles
being unnaturally long, possibly fused, and others apparently subdivided. There
may also be two successive muscular or syzygial joints instead of the normal alternation
of joints that occurs for the rest of the arms. The position of the first pinnule is also,
perhaps consequently, rather irregular. One arm has a pinnule on both sides of Br8,
another on both sides of Br9 ; otherwise it is unduplicated and twice on Br7 and
once on Br8. A complete arm measures 80 mm and has 113 brachials, with about
26 pinnules each side. A proximal pinnule has 16 segments and measures c. 7-5 mm
in length. The first six brachials have sharp lateral flanges after which the arm
becomes constricted laterally.

No. 1935.3.15.1 again has retained a detached distal end of the stalk with lobed
extensions fused to the substrate. The calyx is regularly conical except for a slight
hollowing near the BR suture. The sutures are less distinct than in the other

170

AILSA M. CLARK

FIG. 2. a-d. Democrinus brevis (A. H. Clark), a. Holotype, calyx and adjacent ossicles
viewed radially, b-d. B.M. no. 1900.5.17.1, Cuba. b. Calyx and adjacent ossicles
viewed interradially. c. Tenth columnal. d. Distal end of stalk (attached substrate
omitted), e and f. Reconstructions from descriptions and measurements of holotypes of
Rhizocrinus sabae A. H. Clark and R. robustus A. H. Clark, synonymized by Gislen with
Democrinus rawsoni and D. conifer respectively. [The scale measures c. 2 mm.]

specimens (but still more easily seen than those of Democrinus parfaiti}. The arms
have become detached at the first joint but remain in a group ; the first pinnule is on
Br9, Br10 or Br12 on three arms, has not yet appeared by the last remaining brachial,
Br13, on the fourth, while the last arm is broken earlier. Lateral flanges extend
to Br7.

The last paratype, 1949.4.5.1, is badly broken. The top four columnals are
abnormal in all being discoidal, not gradually increasing in length, so that it is not
until the seventh that the length exceeds the breadth. Only the bases of three arms
remain attached to the calyx.

Carpenter (1884) gave the maximum length of these specimens (presumably
inducting the arms) as 190 mm with arm length 75 mm, making a stalk and calyx
length of 115 mm. This approximates to my measurements of the first paratype,
lacking the distal end of the stalk.

The specimens from Cuba also show irregular distal expansions of the stalk
(Fig. 2d) attached to pieces of the substrate, with relatively few radicular cirri
arising from the distalmost columnals. The arms, or parts of them, are present in
five individuals and are broad for about the first five brachials. The calyx is dis-
tinctly conical, sometimes with a small angle in the profile of the lower half if the
broadening increases abruptly and usually a slight hollowing around the BR suture.

DEEP ATLANTIC CRINOIDEA 171

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Democrinus brevis
Holotype
Paratypes
Cuban specimens
Aberrant Cuban

Democrinus parfai
Smallest
Largest
Remainder

The length of tl
measured across 1
minimum breadth
measured in D. PC

172 AILSA M. CLARK

Again there is no sign of any sudden constriction near the top of the calyx, unlike
D. parfaiti. The sutures are relatively easily distinguished and mostly fairly straight,
though the long BB ones may be partially sinuous (Fig. 2b).

Like three of the types of D. brevis, these Cuban specimens were hitherto labelled
as Democrinus rawsoni, under which name Gislen (1928) discussed them, treating
the two lots as conspecific. Although the calyx is more squat in the types (maximum
breadth 81-100% of the height, compared with 64-83% in the Cuban specimens),
the similarity in stalk attachment, shape and proportions of columnals* and arms
and conical form of the calyx, is so marked that I must agree with Gislen.

It remains to be determined whether D. brevis is specifically distinct from Z).
rawsoni, the type material of which evidently has the calyx much more nearly
cylindrical in shape, at a comparable height, and not broadest at the top of the radial
ring, unlike the specimens just discussed. The consistency of form of the calyx in
the sample of 24 specimens of D. parfaiti discussed below indicates that the shape
does provide a specific character in some cases, though it must be considered in the
light of growth changes. Gislen (1927) has demonstrated that growth in the calyx
of his Democrinus japonicus changes the shape from conical in the young to almost
cylindrical in the larger specimens and I have also found this to be true in Demo-
crinus chuni (Doderlein) in south-east Africa (1972 : 146-150). In D. parfaiti too,
growth of the calyx is stronger vertically than horizontally. However, it is perhaps
not impossible that in D. rawsoni the growth gradients are reversed or much more
variable. Until more precise details are available this problem must go unsolved.

DISTRIBUTION. Atlantic side of Panama ; northern Leeward Islands ; Cuba ;
360-550 metres.

Democrinus parfaiti Perrier

(Fig- 3)

Democrinus parfaiti Perrier, 1883 : 450-451 ; 1886 : fig. 192 ; A. H. Clark, 1923 : 46 ; Mortensen,
1927 : 20, fig. 8 ; Gislen, 1938 : 28-29 ', *947 : 7~9» fig- 3 > A. H. Clark, 1949 : 376 ; Gislen,

1951 : 56, 57-

Rhizocrinus rawsoni : Carpenter, 1884 (pt) : 263, 265, 268, ? pi. 9, figs 3-5, pi. 10, figs 3-5 and
8—14 only, ? pi. 53, figs 7, 8 ; Koehler, 1909 : 255-256 ; Koehler & Vaney, 1910 : 31 ; Doder-
lein, 1912 : 12-13, fig- 4» pl- 4» fig- 7» pl- 9> fig- 2- Non R. rawsoni Pourtales, 1874].

Rhizocrinus parfaiti : A. H. Clark, 1909 : 676.

MATERIAL. Shackleton cruise 3, 1973 : st. 152, 36°35'7'N, ii°O4-i'W-36°37-2'N,
II°°5'3/W, Ormonde Sea Mount (SW. from Cape St Vincent, Portugal), 2292-1691
metres, muddy ; 5 specimens. St. 153, 36°39-oN, n007-9'W-36°397'N, ii°o8-i'W,
871-650 metres ; i specimen. St. 154, 36°33-8'N, ii°o6-5'W-36035-4'N, ii°o6-9'W,
2959-1946 metres, limestone and mud ; 2 specimens. St. 175, 36°20'9'N, I2°53'3'W
-36°22-8'N, i2°53'i'W, 3010-2210 metres, rocky and mud gravel ; 14 specimens.

* As Gislen noted, one Cuban specimen - the aberrant one in Table 2 - has relatively longer columnals
than the others, length/median breadth up to 3-3/1, otherwise not more than 2-7/1. It has the longest
stalk, 210 mm plus c. 5 mm since a few of the distalmost columnals are lost. Even so, the maximum
distal breadth is as much as 3-0 mm. In other respects it agrees with the remaining seven specimens
and I think they are conspecific.

DEEP ATLANTIC CRINOIDEA 173

St. 176, 36°26-4'N, i2°55-8'W-36°27-6'N, i2°56-o'W, Cromer Sea Mount, 1143-
1130 metres, muddy ; 2 specimens. BM(NH) reg. nos 1976.1.12.3-5, 6, 7, 8-20 and
21.

Challenger st. 76, 38°n'N, 2^°og'W (S. of Terceira, Azores), 1645 metres ; 2
specimens, no. 85.3.30.1.

Porcupine st. 43, 1869, 5o°oi'N, i2°26'W (off Cape Clear, SW. Ireland), 2210 metres,
globigerina ooze ; 2 young specimens, no. 85.3.30.2.

NOMENCLATURE. Only a year after being first described, Democrinus parfaiti
from Morocco was referred by Carpenter (1884) to the synonymy of Rhizocrinus
rawsoni Pourtales, 1874, type locality off Barbados. In 1909 A. H. Clark restored it
to specific rank, at the same time describing no less than four nominal species of
Rhizocrinus from the western Atlantic, one of them, R. brevis, split off from R.
rawsoni sensu Carpenter. In 1917 A. H. Clark revived the genus Democrinus for the
species previously referred to Rhizocrinus with distinct sutures within the calyx,
unlike the type species, Rhizocrinus lofotensis M. Sars. This treatment was followed
by Mortensen and Gislen. In 1947 Gislen published drawings by Cherbonnier of the
calyces of two of Perrier's Travailleur (syn) types from Morocco, with correct measure-
ments, unlike the misleading ones given by Perrier to Carpenter. However, D.
parfaiti has never been adequately described and information about the growth
changes and variation is needed for comparison with the other Atlantic species of
the genus.

DESCRIPTION. In 12 specimens with the distalmost columnals preserved, the
stalk terminates in an abrupt division into two or three major branches of similar
magnitude, which themselves divide further into a number of radicular cirri with
long, slender, cylindrical segments, forming an extensive ramifying root system
similar to that in Bathycrinus. On one well-preserved stalk c. 100 mm long the total
length of the 'rooted' portion exceeds 35 mm. Sometimes there may be some
additional small radicular cirri arising near the ends of several of the distalmost
columnals.

These lower columnals are 'screwed', that is elliptical at the joints but twisted so
that the successive joints have their long axes almost at right angles, facilitating
multi-directional flexure. Unlike the squat distalmost columnals of Democrinus
brevis, in this species they are appreciably longer than the maximum breadth. After
about the ten lowest ossicles, the following ones become more nearly round in cross-
section throughout their length and are less constricted medially, sometimes almost
perfectly cylindrical and, in larger specimens, the more proximal ones may be
slightly barrel shaped. They retain about the same length until close below the
calyx, where the length rapidly decreases so that the third or fourth columnal from
the top is the last to be longer than broad, except in two specimens (including the
smallest one) where it is the fifth. The stalk length to the main distal division ranges
from 70 to c. 200 mm and the number of columnals from 39 to at least 67.

The calyx is relatively high and narrow and in every case has a more or less deep
constriction near its upper end, which is emphasized by shrinkage of the thickened
integument when specimens are dried. As noted by Carpenter, the constriction does

174

AILSA M. CLARK

FIG. 3. Democrinus parfaiti Perrier. a. Shackleton st. 175, calyx and adjacent ossicles
viewed interradially. b. st. 152, distal end of stalk with 'roots'. [The scale measures
2 mm.]

not coincide with the basiradial (BR) suture but 'crosses the radials at the level of
the highest angles of the basals' as shown in a drawing sent to him by Perrier and as
he observed himself in one of the small Porcupine specimens (1884, pi. 53, fig. 8).
However, Carpenter's artist incorrectly drew the BB suture as extending right up to
the constriction in a Challenger Azores specimen (pi. 10, fig. 3), an impression also
given by Cherbonnier's drawings of two syntypes (in Gislen, 1947, fig. 3), he having
probably been misled by Perrier's description of the constriction separating the basals
and radials. Certainly the upper end of this suture is usually very faint. In some
of the Shackleton specimens even the upper interradial angles of the basals come below
the constriction (Fig. 3a) which falls entirely within the radials, as shown in a type of
D. conifer by A. H. Clark (igi5a, fig. 131) and in a Caribbean specimen by Carpenter
(1884, pi. 9, fig. 3). Below the constriction the calyx is almost cylindrical or narrowly
conical, with a slight increase in the degree of taper in the lower half. The diameter
at the lower end is as much as 69% of the maximum diameter across the radials,

DEEP ATLANTIC CRINOIDEA 175

decreasing to c. 45% in the largest few specimens (calyx height >6 mm), the mean
over the whole size range being 54%. The shape of the calyx is therefore not more
openly conical in younger specimens in contrast to Democrinus chuni (Doderlein) and
D. japonicus Gislen (see A. M. Clark, 1972 and Gislen, 1927). The sutures in the
calyx are very indistinct ; though the interbasal (BB) ones can usually be found by
partially drying, the BR and RR sutures almost invariably need also to have been
wetted with xylol or clove oil to show up. The BR sutures are rarely completely
straight and may be quite devious, while the BB ones are often partially sinuous.

One specimen shows an extra horizontal suture about half way up the basals,
while another has only four interbasal sutures (but the usual five radials).

The first brachial (Brx) is rectangular or slightly tapering, usually just broader
than long. Only four specimens retain parts of some arms beyond the first brachial
joint, which appears to be more vulnerable than that of D. brevis (though the greater
depth of dredging might be responsible for this). The largest of the four (calyx
height 5-3 mm) has a complete arm c. 23 mm long but may be incompletely re-
generated from Brx judging by the slight diminution of breadth at the first joint
(Fig. 3a). It has pinnules up to 6-0 mm long with up to 15 segments. The first
pinnule is usually on Br8 (i.e. = Pb). A second specimen, calyx height 4-7 mm, has
the first pinnule (P3) on Br10 on the single arm remaining, while the smallest, calyx
height 2-5 mm, has the first pinnule (Pc) on Br12. All have syzygies alternating
regularly with muscular joints throughout the arms remaining. There is a slight
lateral flange on Br2 and 3 but the arms are distinctly compressed laterally from the
syzygy at 3 + 4.

GROWTH CHANGES. In the stalk the maximum distal breadth ranges from 1-2 to
2-5 mm in the present material, tending to increase with growth much more than the
proximal breadth, which is 1-0-1-6 mm. Although the longer columnals are rela-
tively longer in smaller specimens and have length/median breadth up to 3-2/1 (at
calyx height 2-5 mm), above a calyx height of 4 mm the ratio is fairly restricted,
ranging from 1-6 to 2-3/1, with a mean of 1-9/1.

Over most of the size range there is not much change of shape in the calyx ;
excepting the largest specimen (see Table 2), the maximum breadth shows a mean of
50% of the height in nine specimens with height 5 mm or more, compared with a
mean of 57% in six specimens with height up to 4 mm. However, this largest
specimen (height 7-7 mm) is distinctly narrower with a breadth of only 36% and the
larger syntype drawn by Cherbonnier (in Gislen, 1947, fig. 3) at a height of 7-4 mm
has the breadth c. 42%. Beyond c. 6 mm height it is possible that the breadth
increases more slowly.

Judging from the few specimens with arms, the more proximal pinnules are
developed successively, as noted in some other bathycrinids.

AFFINITIES. In spite of its restoration from the synonymy of Democrinus
rawsoni (Pourtales) by A. H. Clark in 1909, no proper comparison between D.
parfaiti and that Caribbean species has been made. A. H. Clark grouped the two as
both having the calyx approximately cylindrical but distinguished them by the
relative size of the bottom diameter of the calyx in comparison with the diameter of

176 AILSA M. CLARK

the radials (c. 50% in D. parfaiti but more than 66% in D. rawsoni from his estimates).
However, his concept of D. parfaiti was based on Perrier's fig. 192 (1886) (reproduced
by Mortensen, 1927 : 20) which gives c. 58% from my measurements). Gislen (1938)
in his key again emphasized the more conical form of the calyx in D. parfaiti and also
separated them on the more attenuated form of the columnals in that species
(length/breadth 2-3/1, with breadth rarely exceeding i mm, as opposed to 1-5-2/1
with breadth >2 mm in D. rawsoni}. However, out of the present sample of D.
parfaiti, even the smallest one has the minimum (top) diameter of the stalk just over
i mm and it may be up to 1-6 mm in the larger ones with the maximum distal
diameter up to 2*5 mm.

Unfortunately, out of six samples in the British Museum collections hitherto
labelled as D. rawsoni, two are probably or certainly D. parfaiti, three form part of
the type series of D. brevis and the last, from Cuba, is also conspecific with D. brevis.
It is therefore necessary to fall back on the published information concerning D.
rawsoni, which is rather inadequate. Even so, several significant features can be
extracted from it.

Firstly, the form of the single distal stalk attachment from Barbados seen by
Pourtales (1874) is described as having 'the appearance of having been partly attached
to a solid body by enlarged surfaces and to have had relatively few radicular cirri',
possibly much as in D. brevis illustrated here (Fig. 2d), though his fig. 2 shows only
some ill-defined truncated branches. He contrasts this attachment with the
'ramifying rootlets' of Rhizocrinus lofotensis which, he says, are adapted for anchorage
in a loose substrate, as are those of the Shackleton specimens of D. parfaiti (though
the initial division into major branches resembles more closely the root form of
Bathycrinus than of Rhizocrinus}. Carpenter (1884, pi. 10, fig. 15) illustrates a
peculiar stalk termination of a specimen from off Havana, Cuba, with about eight
short, splayed radicular cirri each ending in an irregular expanded disc fixed to bits
of substrate (the whole resembling a spider with big flat feet).

Secondly, the columnals do appear relatively broader in Pourtales' figs i and 2 of
D. rawsoni than they are in D. parfaiti, even if the breadth is of similar magnitude
(see below under D. rawsoni}, so Gislen's distinction by this character probably holds
good.

Thirdly, Pourtales' figs i and 4 and A. H. Clark's fig. 133 (igiSa) of calyces of
type material of D. rawsoni show not only a nearly cylindrical form of calyx but
also no sign of the abrupt constriction near the top found so consistently in the
Shackleton material and the types of D. parfaiti. Also Pourtales describes the sutures
of the calyx as 'plainly visible with a lens of moderate power', those of D. parfaiti
being only visible with difficulty (sometimes not at all in the case of the BR and RR
sutures) .

It is also possible that there is a difference in the shape of the arm bases. Pour-
tales shows them as broad for about the first six brachials before tapering, as in
D. brevis, whereas in D. parfaiti only Br2 and Br3 have a slight lateral flange, the
arm apparently narrowing from Br4.

Of the other Atlantic species of Democrinus, D. brevis, like D. rawsoni, has the
stalk attached by a terminal expansion of irregular flattened lobes, relatively stouter

DEEP ATLANTIC CRINOIDEA 177

columnals, distinct sutures in the calyx and no sharp constriction around the
radials. In addition, its calyx is relatively broad and squat.

A much closer resemblance is shown by Democrinus conifer (A. H. Clark, 1909)
from off Brazil, particularly in the elongate conical form of the calyx, with an abrupt
constriction around the radials. Gislen (1938) was misled by Perrier's description
of D. parfaiti into distinguishing it from D. conifer by having a furrow (constriction)
between the radials and basals, not above, as it really is, sometimes markedly so.
The calyx height in the holotype of D. conifer is as much as n mm. Carpenter
(1884, pi. 10, figs 3-5) shows a Caribbean specimen (presumably from the Blake
collections), also with an abrupt constriction in the radials.

Gislen (1938) lists numerous references under the heading of D. conifer and
synonymizes with it Rhizocrinus robustus A. H. Clark, 1909* and Bythocrinus
intermedius A. H. Clark, 1915. In 1923 A. H. Clark cited records of (Bythocrinus)
robustus from Brazil as well as the Gulf of Mexico (the type locality) and also from
the vicinity of Portugal. At least the last record I believe must be referable to
D. parfaiti. If the stalk attachment of D. conifer proves to be root-like, then it will
be difficult to maintain it as a valid species distinct from D. parfaiti.

Finally, Gislen (1947) has described Rhizocrinus magnus from off Tangier (c. 36°N,
8°W) in 2150-2300 metres, the holotype lacking the arms and the distal part of the
stalk. The shape is very similar to D. parfaiti, even to a constriction at the top of
the conical calyx, which is 5 mm high, but the sutures of the calyx are evidently so
closely fused as to be invisible - hence the inclusion in Rhizocrinus. In spite of this,
the approximation of locality and the resemblance in shape throw some doubt on the
validity of R. magnus as distinct from D. parfaiti in my opinion.

DISTRIBUTION. Re-examination of two of Carpenter's Challenger specimens from
the Azores, named by him Rhizocrinus rawsoni, shows that they are conspecific with
D. parfaiti, which A. H. Clark (1949) has also recorded from the vicinity of these
islands. Doderlein's Valdivia specimens from south of the Canary Is are also
referable to D. parfaiti, judging from his illustrations. Unfortunately the only
Porcupine specimens from off Cape Clear, Ireland, retained in the BM collections are
the two very small ones figured by Carpenter (1884, pi. 53, figs 7 and 8) ; with their
distinct sutures in the calyx and constriction or hollow around the radials they are
most likely also to belong to this species. However, Doderlein's record based on a
stalk fragment from the Faeroe Channel is very dubious. The known distribution
may therefore be limited to the NE. Atlantic from the Azores eastward between
latitudes 24° and 5o°N in depths of 870 (?65o)-250o (?30io) metres.

Democrinus rawsoni (Pourtales)

Rhizocrinus rawsoni Pourtales, 1874 : 27-31, pi. 5, figs 1-7, 9-12, 14 ; Carpenter, 1884 (pt) : 263

(Hassler and probably most Blake records), ? pi. 10, figs 6, 7, 15-20.
? Rhizocrinus sabae A. H. Clark, 1909 : 675.
Democrinus rawsonii : A. H. Clark, igi5a : 203, 205, figs 133, 138 ; H. L. Clark, 1918 : 16 ;

A. H. Clark, 1921 : 26 ; 1923 : 45 ; Gislen, 1938 : 29-30 ; H. L. Clark, 1941 : 7-8. [Non

D. rawsoni : Gislen, 1928.]

* In listing the echinoderms from the Gulf of Mexico in 1954, A. H. Clark still cites robustus as a valid
species (of Democrinus).

178 AILSA M. CLARK

MATERIAL. None. As explained above, all the specimens referred to this species
by Carpenter (1884) in the British Museum collections have since been referred to
other species of Democrinus.

REMARKS. Pourtales (1874) described this species from several specimens taken
west of Barbados. His figure of a calyx (1874, pi. 5, fig. 4) shows an almost cylindrical
shape, broadest at about half its height and maintaining a similar breadth to the
top of the radials. The maximum breadth is only about 50% of the height (if the
drawing is accurate). A. H. Clark's camera lucida drawing (igi^, fig. 133) of a
syntype shows a calyx of similarly almost cylindrical but even more elongate form,
the maximum breadth only 33% of the height. Although both figures show a
slight hollowing in the vicinity of the BR suture, there is no sign in either of an abrupt
constriction such as in D. parfaiti.

Neither Pourtales (1874) nor A. H. Clark (iQJ-Sa) gives any measurements for the
calyx but the largest syntype measures 170 mm in length without the arms and the
distal extremity of the stalk, which are missing ; the remaining part of the stalk has
63 columnals and is 1-5-2 mm in breadth. Judging from the drawings, if the upper
end of the stalk is 1-5 mm broad then the calyx height would be c. 4 mm in the one
shown by Pourtales and 6 mm in Clark's. This is a similar magnitude to the
specimens of D. brevis and D. parfaiti described above. In 1921 A. H. Clark
described a specimen from the Barbados - Antigua Expedition, closely resembling
the type material, having calyx height 4-7 mm (breadth not given) and a stalk of
58 columnals measuring 155 mm in length. H. L. Clark (1918) recorded a specimen
from off Havana, Cuba, with calyx 4 mm high and 1-85 mm in (presumably maximum)
diameter (46% of the height), the stalk having 60 columnals and measuring 145 mm.
Carpenter (1884) gives the calyx height and maximum breadth of a Blake specimen
as 6 and 2-5 mm, making a relative breadth of 42% at a stalk length of 180 mm, with
68 columnals present. Most of the Blake specimens he records were from the southern
Caribbean in the vicinity of the type locality but one was from Cuba.

Carpenter treated this species as having a very wide range of form of the calyx
but subsequent workers, notably A. H. Clark (1923), have emphasized that it is
nearly cylindrical. It is surprising then that A. H. Clark in 1923, when listing
the Atlantic crinoids with their ranges, omits D. brevis but cites its type locality
(off Colon) under the heading of D. rawsoniand presumably at that time thought them
to be synonymous, in spite of the squat conical calyces in the type material of D. brevis.
In view of this conflict and the other Caribbean species of Democrinus which have
been described, it is desirable that the material from this area should be reexamined.

DISTRIBUTION. Owing to the confusion of records it is uncertain whether this
species extends beyond the Caribbean ; A. H. Clark (1923) gives the depth range as
66-652 metres.

Democrinus sp.
(Fig. 4)

MATERIAL. Challenger st. I22C, O9°io'S, 34^9 'W (off NE. Brazil), 730 metres ;
i specimen, BM(NH) reg. no. 85.3.30.29. [Named R. lofotensis by Carpenter (1884).]

DEEP ATLANTIC CRINOIDEA 179

Atlantis II cruise 31, st. i62A, o8°02'S, 34°03'W-07°56'S, 34°09'W (off NE.
Brazil), 1493 metres ; 6 calyces, 2 distal ends of stalks.

Atlantis II cruise 31, st. 167, 07°58'S, 34°i7'W-o8°5o'S ; 943-1007 metres ; 2
calyces.

Atlantis II cruise 31, st. iGgA, o8°03'S, 34°23'W-o8°02'S, 34°25'W, 587 metres ;
5 calyces and 2 pieces of 'root'.

Atlantis II cruise 60, st. 262A, 36°o5-2'S, 520i7'9'W (off Rio de la Plata), 2440-
2480 metres ; I calyx and pentacrinoid.

The Challenger specimen is badly decalcified. Numerical data from the other 13
similarly small specimens are given in Table 3.

breadth = calyx height breadth Br. as

Station minimum breadth (maximum) (maximum) % of Ht.

i6aA 0-33 2-17 1-58 73

262A 0-45 2-10 1-64 78

167 0-54 2-08 1-50 72

i62A 0-33 2*00 1-42 71

i6gA 0-29 1-92 1-54 80

i6gA 0-42 1-83 i'25 68

i6gA 0-33 1-82 i'20 66

i62A* 0-42 i'75 1-17 67

i62A 0-30 1-58 1-29 82

i62A 0-27 1-46 1*25 86

167 0'37 1-42 I-21 85

i62A 0-25 1-42 1-17 82

i6gA 0-33 1-33 1-17 88

* This specimen is the six-armed one.

DESCRIPTION. One calyx from st. i62A (Fig. 4b, the eighth in Table 3) is six-
rayed and also differs from the rest in having the interbasal (BB) sutures distinct
and regular, as in Democrinus chuni. In the last and smallest specimen in Table 3
(Fig. 4c) three very irregular BB sutures can just be discerned when partially dried
after wetting with xylol and in a few other specimens there are also traces of these
sutures. Otherwise in the remaining specimens only the BR and RR sutures are
distinguishable (Fig. 4a) and even these sometimes only appear after xylol treatment.
This contrasts with Rhizocrinus lofotensis M. Sars, ranging from Norway to NE.
America, which has the BR and RR sutures distinguishable in only a minority of
specimens and the BB ones not at all. Gislen (1938) diagnosed Rhizocrinus Sars as
having the BB sutures fused and the radials Very closely connected with and most
often coalesced with' the basals, as opposed to Democrinus Perrier with 'basals
separated by distinct sutures' and radials 'always separated from each other and the
basal cup by distinct sutures'. The six-rayed specimen therefore runs down to
Democrinus but not the rest ; the visibility of its sutures could be correlated with

i8o

AILSA M. CLARK

FIG. 4. Democrinus sp. a-c. Calyces and adjacent ossicles of the first, eighth and last
specimens in Table 3. d. Thirteenth columnal of c. e and f. Distal ends of two stalks
(possibly of another species). [The scale measures i mm.]

the additional one. As noted above, the sutures in the calyx of Democrinus parfaiti
Perrier-the type-species of the genus -may be very hard to distinguish so the
character is not altogether reliable.

The shape of the normal calyces is more or less conical (fig. 4a), though the largest
one is slightly swollen near the top of the radials. However, about five specimens
have somewhat deformed calyces with irregular bulges or constrictions. Two from
st. i62A have perforations in the calyx and the Challenger specimen has two large
round holes with a relatively huge parasitic gastropod still hanging from one of them.
Possibly the other specimens have also been parasitized. The maximum breadth of
the calyx ranges from 66 to 88% of the maximum height in the 13 specimens, with a
mean of 77%. There appears to be a relative decrease in breadth compared to
height during growth, as in R. lofotensis and in the similarly small South and East
African species Democrinus chuni (Doderlein). The height of the basal ring forms
c. 65% of the maximum calyx height.

In all the specimens the arms are broken after the first brachial, which ossicle is
relatively narrow, even in the largest specimen only slightly broader proximally than
long. It should be noted that, although Gislen (1938) also distinguished Rhizocrinus
by the relatively long Br^ as here, I found in 1972 that D. chuni has length/breadth
of the first brachial ranging from 0-6 to 1-7/1.

As for the stalks, only a few proximal columnals are short, the fourth one being
usually the first to have length exceeding breadth. The length increases markedly
and by the tenth the length/median breadth ratio is usually 3'5-4-o/i. No complete
stalk remains. Some of the broken middle or distal pieces have very swollen joints
(Fig. 46) while others are only slightly so (Fig. 4f ) and it is possible that two species

DEEP ATLANTIC CRINOIDEA 181

might be represented. It is significant that the distal ends of the stalks present
bifurcate or trifurcate into relatively large branches, with few, if any, small radicular
cirri arising from the columnals immediately above the division, in contrast to
Rhizocrinus lofotensis where the column usually ends abruptly without producing
any large branches and numerous fine radicular cirri arise on most columnals of the
distal half of the stalk (see Sars, 1868, pi. i, figs I, 6, 7, 17 and 19, though the speci-
men in fig. 1 6 does have a large branch from the penultimate columnal). This is
much more like the rooting form of Democrinus chuni and D. parfaiti.

AFFINITIES. Two other species remain in Rhizocrinus in addition to R. lofotensis,
namely R. minimus (Doderlein, 1907) from the East Indies and R. magnus Gislen,
1947 from off Morocco. Both are known only from single specimens lacking the
distal end of the stalk but both agree with R. lofotensis in lacking any distinct sutures
in the calyx. Since the present specimens have at least the RR and BR sutures
visible (if not some or all of the BB ones) and branching distal ends to the stalks,
they clearly have greater affinity with Democrinus chuni than with Rhizocrinus
lofotensis. In spite of the wording of Gislen's (1938) diagnoses I think that they
should be referred to Democrinus. The distinction of these two genera needs further
study when more material is available.

Order COMATULIDA

Family ATELEGRINIDAE

Atelecrinus sp. juv.

(Fig. 5)

MATERIAL. Atlantis II cruise 60, st. 242, 38°i6-9'S, 5i°56-i'W (SE. of Rio de la
Plata, Argentina), 4382-4402 metres ; i pentacrinoid.

Atlantis II, cruise 60, st. 256, 37°40-9'S, 52°i9'3'W, 3906-3917 metres ; i post-
pentacrinoid and i pentacrinoid.

Atlantis II cruise 60, st. 262A, 36°05-2'S, 52°i7'9'W, 2440-2480 metres ; i
pentacrinoid.

[Additionally some of the 25 pentacrinoids from st. 247A may belong here rather
than in Bathycrinus australis.]

The post-pentacrinoid (Fig. 5a) has probably only recently broken free of its stalk,
though the perfectly flat broad apex of the centrodorsal shows no perforation. The
breadth is 0-6 mm, whereas the height is less at this stage. There were probably
X cirri arranged in a zig-zag row around the side ; between the five adapical ones the
stereom extends into somewhat irregular lobes.

Two intact cirri remain, both with 14 segments, all but the first one longer than
broad. The larger one measures 6-8 mm and its longest segment has length/median
breadth 0-83/0-12 mm (c. 7/1). The terminal claw is almost straight and has a
blunted tip.

The calyx is hyocrinid-like, the postradial series being distinctly separated
laterally so that the upper ends of the radials extend between them. The maximum

I 82

AILSA M. CLARK

FIG. 5. Atelecrinus sp. a. Atlantis II st. 256, post-pentacrinoid, viewed radially.
b. st. 242, pentacrinoid, viewed interradially. [The scale measures i mm.]

breadth across the top of the radials is 1-3 mm and the height interradially 0-7 mm.
The basals still form a complete ring and make about a third of the height of the
calyx.

The two arms of each pair are slightly spaced, corresponding to the condition of the
division series. The arm length was probably c. 8 mm but all are broken by the
sixteenth brachial. Proximal syzygies are at 3 + 4 and 9 + 10. The breadth at
3 + 4 is 0-25 mm. No pinnules are developed before Br14 (P6).

The pentacrinoid from the same station (256) has the calyx very similar except
that the radials are relatively longer. Only the IBrx have so far developed from the
radials. The uppermost columnal is slightly swollen.

The pentacrinoid from st. 242 (Fig. 5b) is a little more advanced with rudiments of
the IBr2 forming and the uppermost columnal markedly swollen in each interradius,
heralding the first cirri. Only eight columnals remain, of which the last three are
longer than broad, the eighth with length 3-5 x breadth. The three lower segments
each have a slightly raised belt, usually medially. The lower ends of the basals are
distinctly constricted. Possibly the centrodorsal is liable to grow up over this area.

The generic position of these specimens is indicated by several factors, the com-
pleteness of the basal ring, even at the post-pentacrinoid stage, the very attenuated
form of the cirri and the retarded development of the pinnules, in comparison with
young Antedonidae.

DEEP ATLANTIC CRINOIDEA 183

Atelecrinus balanoides P. H. Carpenter (see Clark & Clark, 1967 : 823) was taken by
the Challenger off Pernambuco, Brazil (c. g°S), and may well extend south to
these latitudes.

Family ANTEDONIDAE
? Hybometra sent a A. H. Clark

Hybometra senta A. H. Clark, 1913 : 54 ; A. H. & A. M. Clark, 1967 : 550-552, fig. 31.

MATERIAL. Atlantis II cruise 42, st. 202, o8°56'S, i2°i5'E-o8°46'S, i2°47'E (off
Angola), 1427-1643 metres ; c. 50 poorly preserved specimens.

The holotype and only recorded specimen of Hybometra senta was from Brazil in
only 42 metres. It is considerably larger than these specimens, arm breadth at
Br3+4 2-0 mm and arm length 90 + mm, whereas the largest of these have the breadth
c. 1-25 mm and arm length c. 55 mm.

There is agreement between them in the spinose and flared division series and
brachials and in the arrangement of the cirri in fairly regular vertical columns
(especially interradially) on the conical centrodorsal, though the vertical height
usually just exceeds the maximum breadth in these specimens, whereas in the holo-
type it is relatively lower, height/breadth 3-2/4-1 mm. The cirri of the holotype are
unfortunately known. In the present specimens they number up to XC, forming
four columns of four (sometimes five) sockets in each radius. The cirri have up to
36 segments, all except the first one longer than broad, the longest (about the fifth)
with length/median breadth 1-5/0-35 mm = 4-28/1. All the segments are very
expanded at their distal ends ; the opposing spine is well developed and the terminal
claw slightly curved. The total length is up to 22 mm.

The division series are slightly flared and spinose at their distal edges but the
brachials soon develop long spiny tufts. The IB^ and Brt are markedly constricted
laterally, the succeeding ossicle markedly projecting.

Px has well over 20 segments and measures 6 + mm ; the segments are elongated
and spinose. P2 and the genital pinnules were probably smaller and with even
more slender segments.

Of the few crinoids recorded from West Africa, Leptometra celtica has a similarly
often conical centrodorsal and its cirri and pinnules have similar proportions ;
however, the curved terminal claw on the cirri of these specimens and the spinose
flared ossicles serve to distinguish them.

Leptometra phalangium (J. Miiller)

Alecto phalangium J. Muller, 1841 : 182.

Leptometra phalangium : A. H. & A. M. Clark, 1967 : 553-564, fig. 32a, b.

MATERIAL. Atlantis II cruise 59, st. 211, 34°4i-5'N, i7°28'E (Mediterranean, N.
from Libya), 2834-2789 metres ; I poorly preserved specimen.

The previous deepest record in the Mediterranean for Leptometra phalangium is
1292 metres, which is considerably exceeded by the present one.

184 AILSA M. CLARK

The centrodorsal is relatively low with only about X cirri in one irregular ring
Possibly this is correlated with the depth and quieter water.
The smooth attenuated cirrus and pinnule segments agree with L. phalangium.

Indeterminable Antedonidae

Some small, more or less broken, Antedonidae belonging to three different species,
probably all of Bathymetrinae, were taken respectively at the following stations :

1. Atlantis II cruise 60, st. 256, 37°40'9'S, 52°i9'3W (off Rio de la Plata), 3906-
3917 metres; 7 specimens. St. 259A, 37°i3'3'S, 52°45'W, 3305-3317 metres ; 6
specimens.

2. Atlantis II cruise 31, st. i62A, o8°02'S, 34°03'W-07°56'S, 34°09'W (off NE.
Brazil), 1493 metres ; I specimen.

3. Chain cruise 106, st. 328, 50°047'N, i5°44-8'W (SW. from Cape Clear, Ireland),
4426-4435 metres ; 5 specimens.

SPECIES i. The largest specimen has arms 15+ (possibly c. 25) mm long ; the
arm breadth at 3 + 4 is 0-9 mm and the length from the IBrx to Br9 (inclusive) is
7-2 mm. The centrodorsal is low blunt conical with cirrus sockets irregular in
position. The longest cirri have up to 23 fairly long segments, total length up to
n-2 mm with the longest segments having length/median breadth 2-5/1 ; they are
flared at the joints. The division series and arms are laterally constricted and
rounded, the division series finely rugose, the brachials soon developing a small tuft
of spines at the flared distal end. All the proximal pinnules are broken ; the first
three probably had at least 12 segments, P3 is the first genital pinnule and much larger
than the preceding ones with segments 4-6 relatively longer and broader, rather as
in Isometra angustipinna (P. H. Carpenter) from off Uruguay. However, that species
at a similar size has only five or six segments in the first two pinnules and a large
flange on the second and third segments of the genital pinnules. There is also some
resemblance to the holotype of Phrixometra longipinna (P. H. Carpenter), also known
from off Uruguay.

SPECIES 2. The single specimen has the arm breadth at 3 + 4 only c. 0-5 mm. It
has a low conical centrodorsal with irregularly arranged sockets ; the dorsal pole
is crowned with truncated hyaline papillae. The cirri are very attenuated with up
to 19 segments, which are long, compressed and distally flared. The opposing spine
is sharp and the terminal claw slender and curved.

SPECIES 3. The three larger specimens have the arm breadth at 3 + 4 c. 0-7 mm.
The centrodorsal is low conical and some of the cirrus sockets are arranged in vertical
columns (more like the Zenometrinae) . The cirri have up to 15 relatively short
segments, not more than twice as long as the median breadth, markedly flared dor-
sally and keeled distally. Most of the ossicles are almost smooth, only very slightly
rugose. The pinnules are very attentuated ; the genital pinnules, starting with
P3, have particularly slender segments. The specimens have some affinity with
Orthometra hibernica (A. H. Clark) (see A. M. Clark, 1970 : 34) though the ossicles
are more attenuated, as well as with Trichometra delicata A. H. Clark (see A. M. Clark,

DEEP ATLANTIC CRINOIDEA 185

1970 : 48) though the ossicles are smoother and the latter species has P2 the first
genital pinnule.

REFERENCES

AGASSIZ, A. 1892. Calamocrinus diomedae, a new stalked crinoid. Mem. Mus. comp. Zool.

Harv. 17(2) : 1-95, 32 pis.
CARPENTER, P. H. 1884. Crinoidea. i. Rep. scient. Results Voy. Challenger. Zool. 11(32) :

1-440, 62 pis.
CLARK, A. H. 1907. A new species of crinoid (Ptilocrinus pinnatus) from the Pacific coast

with a note on Bathycrinus. Proc. U.S. natn. Mus. 32 : 551-554.
1908. New stalked crinoids from the eastern coast of North America. Proc. U.S. natn.

Mus. 34 : 205-208, 3 figs.
1909. Four new species of the crinoid genus Rhizocrinus. Proc. U.S. natn. Mus. 36 : 673-

676.
igi5a. A monograph of the existing crinoids. l(i). Bull. U.S. natn. Mus. 82(i) : 1-406,

513 figs, 17 pis.

19150. Die Crinoiden der Antarktis. Dt. Sudpol. Exped. 16. Zool. 8 : 101-209, 9 pis.

1917. A revision of the recent genera of the crinoid family Bourgueticrinidae with the

description of a new genus. /. Wash. Acad. Sci. 7 : 388-392.

1921. Report on the Crinoids. In : Clark, A. H., Rathbun, M. J., Boone, P. L., Shoe-
maker, C. R. & Clark, H. L. Reports on the Crinoids, Ophiurans, Brachyura, Tanidacea

and Isopoda, Amphipods and Echinoidea of the Barbados -Antigua expedition of 1918.

Stud. nat. Hist. Iowa Univ. 9(5) : 1-28.

— 1923. Crinoidea. Dan. Ingolf-Exped. 4(5) : 1-58, 58 figs.
— 1949. Echinoderms from the mid- Atlantic dredged by the 'Atlantis' in the summer of

1948. /. Wash. Acad. Sci. 39 : 371-377.
1954- Echinoderms (other than holothurians) of the Gulf of Mexico. Bull. U.S. Fish.

Commn 55 : 373-379.
& CLARK, A. M. 1967. A monograph of the existing crinoids. 1(5). Suborders

Oligophreata (concluded) and Macrophreata. Bull. U.S. natn. Mus. 82(5) : 1-860, 53 figs.
CLARK, A. M. 1970. Echinodermata : Crinoidea. Marine Invertebrates of Scandinavia.

No. 3. Universitetsforlaget, Oslo. 55 pp., 19 figs.
1972. Some crinoids from the Indian Ocean. Bull. Br. Mus. nat. Hist. (Zool.) 24(2) : 73-

156, 17 figs.
1973- Some new taxa of recent stalked Crinoidea. Bull. Br. Mus. nat. Hist. (Zool.)

25(7) : 265-288, 6 figs, 2 pis.
CLARK, H. L. 1918. Report on the Crinoidea and Echinoidea collected by the Bahama

Expedition from the University of Iowa in 1893. Bull. Labs nat. Hist. St. Univ. la 7(5) : I -

38.
1941. Reports on the scientific results of the "Atlantis" Expeditions to the West Indies.

The Echinoderms. Mem. Soc. cubana Hist. nat. 15(i) : 1-54, 10 pis.
DANIELSSEN, D. C. 1892. Crinoida. The Norwegian North- Atlantic Expedition, 1876-1878.

Zoology. Christiania. 28 pp., 5 pis.

& KOREN, J. 1877. Fra den norske Nordhavsexpedition : Echinodermer. Nyt Mag.

Naturvidensk. 23(3) '• 45-^3, 6 pis.

DODERLEIN, L. 1907. Die gestielten Crinoiden der Siboga-Expedition. Siboga Exped.

42a : 1-52, 12 figs, 23 pis.
1912. Die gestielten Crinoiden der deutschen Tiefsee-Expedition. Wiss. Ergebn. dt.

Tiefsee-Exped. "Valdivia" 17(i) : 1-34, 9 figs, 12 pis.
GISLEN, T. 1927. Japanese crinoids. Vidensk.Meddrdansk.naturh.Foren.S3:i-6g,8ofigs.

1928. Notes on some crinoids in the British Natural History Museum. Ark. Zool.

19(32): 1-15.

i86 AILSA M. CLARK

GISLEN, T. 1938. A revision of the recent Bathycrinidae, with a study of their phylogeny

and geographical distribution. Ada Univ. lund. N.F. 34(io) : 1-30, 18 figs.
1947- The echinoderms collected by the "Skagerak" Expedition in the eastern Atlantic.

1946. 2. Crinoidea. Goteborgs K. Vetensk.-o. vitterhSamh. Handl. B5(io) : 1-9, 3 figs.
1951. Crinoidea, with a survey of the bathy metric distribution of the deep-sea crinoids.

Rep. Swedish Deep-Sea Exped. 1947-8. 2(i) : 49-59, 2 figs, i pi.

— 1956. Crinoids from depths exceeding 6000 meters. Galathea Rep. 2 : 61-62, i pi.
KOEHLER, R. 1909. Echinodermes provenant des campagnes du yacht Princesse- Alice.

Result. Camp, scient. Prince Albert 34 : 1-317, 32 pis.
— & VANEY, C. 1910. Note pr&iminaire sur les Crinoides du Travailleur et du Talisman.

Bull. Mus. natn. Hist. nat. Paris 1910 : 23-32, 3 figs.
MCKNIGHT, D. G. 1973. Stalked crinoids from the New Zealand region. N.Z.O.I. Rec.

1(14) : 199-210, fig. 3.
MACURDA, D. B., JR. & MEYER, D. L. 1976. The morphology and life habits of the abyssal

crinoid Bathycrinus aldrichianus Wyville Thomson and its paleontological implications.

/. Paleont. 50 (4) : 647-667, 5 figs, 5 pis.
MORTENSEN, T. 1927. Handbook of the Echinoderms of the British Isles. London. ix + 47i

pp., 269 figs.
MULLER, J. 1841. tlber die Gattungen und Arten der Comatulen. Mber. K. preuss. Akad.

wiss. 1841 : 179-189.
PERRIER, E. 1883. Sur un nouveau crinoide fixe", le Democrinus parfaiti, provenant des

dragages du 'Travailleur'. C. r. hebd. Seanc. Acad. Sci., Paris 96 : 450-452.

1886. Les Explorations sous-marins. Paris. 352 pp., figs.

POURTALES, L. F. DE. 1874. Crinoids. In : Agassiz, A. & Pourtales, L. F. de. Zoological

results of the Hassler Expedition, i. ///. Cat. Mus. comp. Zool. Harv. No. 8 : 27-31, pi. 5.
SARS, M. 1868. Memoires pour servir a la connaissance des Crinoides vivants. Christiania.

65 pp., 6 pis.
THOMSON, C. WYVILLE. 1872. On the crinoids of the 'Porcupine' deep sea dredging expedition.

Proc. R. Soc. Edinb. 7 : 764-773.

1873. The Depths of the Sea. London, xx + 527 pp., 84 figs.

1876. Notice of new living crinoids belonging to the Apiocrinidae. /. Linn. Soc. Lond.

Zool. 13 : 47-55, 4 figs.
1877. The Atlantic: a preliminary account of the general results of the exploring voyage of

HMS 'Challenger' during the year 1873 and the early part of the year 1876. London. 2 :

xiv + 396 pp., 62 figs, pis 15-42.

Miss A. M. CLARK

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

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A REVISION OF THE LIZARD GENUS ^
SCINCUS (REPTILIA : SCINCIDAE)

E. N. ARNOLD

AND

A. E. LEVITON

f BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 31 No. 5

LONDON: 1977

^

A REVISION OF THE LIZARD GENUS SCINCUS
(REPTILIA : SCINCIDAE)

BY

EDWIN NICHOLAS ARNOLD

AND

ALAN E. LEVITON

Pp. 187-248 ; 3 Plates, 7 Text-figures, 6 Tables

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 31, No. 5

LONDON: 1977

THE BULLETIN OF THE BRITISH MUSEUM

(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Scientific
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. 31, No. 5, of the Zoology series.
The abbreviated titles of periodicals cited follow those
of the World List of Scientific Periodicals.

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

ISSN 0007 -i 498

Trustees of the British Museum (Natural History), 1977

BRITISH MUSEUM (NATURAL HISTORY)

Issued 30 June, 1977 Price £5.25

A REVISION OF THE LIZARD GENUS SCINCUS
(REPTILIA : SCINCIDAE)

By E. N. ARNOLD AND A. E. LEVITON

CONTENTS

Page

SYNOPSIS ........... 189

INTRODUCTION .......... 190

MATERIAL EXAMINED ......... 191

CHARACTERS ALREADY USED OR POTENTIALLY USEFUL IN CLASSIFYING

Scincus ........... 192

Ear and associated structures . . . . . . . 192

Premaxillary rostrum ......... 196

Eyes 197

Variations in scaling ......... 19?

Size and shape of head ........ 199

Colouring 199

SYSTEMATIC ANALYSIS ......... 200

Scincus Laurenti .......... 201

Key to Scincus .......... 205

The Scincits scincus complex ........ 206

North African populations of the Scincus scincus complex . . . 209

Scincus scincus meccensis Wiegmann . . . . . . 219

Scincus scincus conirostris Blanford . . . . . . 221

Scincus mitranus J. Anderson . . . . . . . 224

Scincus hemprichii Wiegmann . . . . . . . 233

SPECULATIONS ON THE HISTORY OF THE GENUS Scincus . . . 235

Relationship of Scincus to other skinks . . . . . . 235

Interrelationships of the species of Scincus ..... 238

Origin of species distribution pattern ...... 242

SUMMARY OF SYSTEMATIC CONCLUSIONS ...... 245

ACKNOWLEDGEMENTS ......... 245

REFERENCES ........... 246

SYNOPSIS

Until recently, up to thirteen species of the scincid genus, Scincus, were recognized, but
examination of some 590 individuals from a wide range of localities suggests that only three or
four are valid. Of these, 5. mitranus is confined to eastern and southern Arabia and 5. hemprichii
probably to southwest Arabia. The remaining forms constitute the S. scincus complex, which
may consist in North Africa of two largely allopatric species, S. scincus and S. albifasciatus,
although evidence for this is not conclusive. The 5. scincus complex is represented in southwest
Asia by two forms : S. scincus meccensis in southern Jordan, northwest and west Arabia and
S. s. conirostris in southern and eastern Arabia, Iraq and southwest Iran.

Scincus appears to have evolved from a primitive scincine, very similar to members of the
Eumeces schneideri group, especially E. (schneideri) algeriensis; it does not seem to be directly
related to the sympatric genus Scincopus. Within Scincus, the 5. scincus complex is the least
specialized component of the genus and both S. mitranus and S. hemprichii may have been
independently derived from it, or from a closely related form. Possibly the whole range of the

190 E. N. ARNOLD AND A. E. LEVITON

genus was once occupied by a S. scincus-like species and its distribution was subsequently
restricted by the onset of less desertic conditions leaving reduced populations in North Africa,
southwest Arabia and southeast Arabia that gave rise to the 5. scincus complex, S. hemprichii
and 5. mitranus respectively. A renewed expansion of arid areas could then have enabled the
S. scincus complex to invade southwest Asia. Some of the characters of its most eastern
subspecies, S. 5. conirostris, may have arisen, or been maintained, by character displacement
through contact with S. mitranus.

INTRODUCTION

THE scincid lizard genus Scincus Laurenti, 1768 is a clearly-defined group of species
adapted to life in areas of loose, usually wind-blown sand. It is distributed from the
western Sahara Desert and its borders, eastwards to Arabia, southern Iraq and
southwestern Iran.

Scincus falls clearly within the assemblage of relatively primitive skinks referred
to the subfamily Scincinae by Greer (1970). Like the majority of these, Scincus
has a more or less unmodified lacertilian palate in which the palatine bones do not
meet on its mid-line, although they are opposed above it. The members of most
scincine genera have elongate bodies and reduced limbs and often the nostril is situ-
ated in, or broadly bordered by, the rostral scale (instead of being pierced in a separate
nasal scale or between two such scales). Scincus has none of these specialized
features and is thus similar to the Saharan genus Scincopus Peters, 1864 and the
Holarctic Eumeces Wiegmann, 1834. It seems likely that these three genera are
among the more primitive living scincines.

Scincus can be distinguished from Eumeces and Scincopus by its single premaxilla,
its postorbital and postf rental bones which are completely fused from earliest life,*
and by possessing a wide range of morphological adaptations to living in loose sand.
These include : fringed digits to increase purchase ; streamlined body shape in-
cluding a spatulate snout with bony rostral support ; powerful body musculature to
facilitate sand-swimming ; mechanisms to allow breathing beneath the sand ;
protected orifices to keep sand out, e.g. very small ear apertures, valvular nostrils,
countersunk jaw ; and often some modifications of the ear that may well facilitate
subterranean hearing and increase the sensitivity of spatial orientation. These
features will be discussed more fully elsewhere.

Numerous taxa of Scincus have been described and some thirteen species, plus a
further two subspecies, are either recognized or have only recently been called into
question. Mertens (1972), the last worker to discuss the genus, lists the following
forms : Scincus albifasciatus Boulenger, 1890 ; 5. arabicus Schmidt, 1939 ; S.
arenarius Murray, 1884 ; 5. conirostris Blanford, 1881 ; 5. deserti Haas, 1957 ; 5.
gasperetti Haas, 1957 ; S. hemprichii Wiegmann, 1837 ; S. meccensis Wiegmann,

* Checked on numerous Scincus and on at least one specimen of each of the following species;
Scincopus fasciatus Peters, 1864 ; Eumeces blythianus (Anderson, 1871) ; E. brevilineatus Cope, 1880 ;
E. brevirostris (Gunther, 1860) ; E. chinensis (Gray, 1838) ; E. dugesii Thominot, 1883 ; E. egregius
Baird, 1858 ; E. elegans Boulenger, 1887 ; E. fasciatus (Linnaeus, 1758) ; E. inexpectatus Taylor, 1932 ;
E. latiscutatus (Hallowell, 1860) ; E. longirostris (Cope, 1861) ; E. lynxe (Wiegmann, 1834) ; E. marginatus
(Hallowell, 1860) ; E. obsoletus (Baird & Girard, 1852) ; E. oshimensis Thompson, 1912 ; E.quadrilineatus
(Blyth, 1853) ; E. schneideri (Daudin, 1802) ; E. septentrionalis (Baird, 1858) ; E. skiltonianus (Baird
& Girard, 1852) ; E. sumichrasti (Cope, 1866) ; E. taeniolatus (Blyth, 1854). In Eumeces the postorbital
and postfrontal bones are at least partly separate in young animals and often in adults as well.

REVISION OF SCINCUS 191

1837 ; 5. mitranus J. Anderson, 1871 ; 5. muscatensis Murray, 1886 ; S. philbyi
Schmidt, 1941 ; S. richmondi Haas, 1961 ; S. scincus scincus Linnaeus, 1758 ; 5. s.
cucullatus Werner, 1914 ; S. s. laterimaculatm Werner, 1914.

Many of these taxa have been described from very small samples, and sometimes
on the basis of single specimens. Furthermore, the characters used in defining
species often only involve minor differences in colour and scaling, some of which are
now known to be quite variable even within populations. It therefore seems
desirable to reassess the validity of the numerous taxa of Scincus. The situation is
particularly acute in Arabia where no less than ten species have been described.

The main aim of this account is the recognition of biological species within the
genus Scincus. Infraspecific geographical variation is also described, but a conserva-
tive attitude has been taken towards the use of formal subspecies names. These
have only been employed in cases where geographical groups of populations differ
fairly consistently from neighbouring groups in at least three independent features.
Even when this is so, formal names have not been applied unless the apparent infra-
specific unit concerned is known from a reasonable number of examples. It will
become obvious that, if a less stringent working definition is used, far more sub-
species of Scincus could be recognized, but this seems undesirable. Many widespread
species have complex patterns of variation that do not really lend themselves to
description by the formal naming of numerous, often arbitrary, segments of the range.
It seems best in such cases to describe geographical variation as such and to reserve
subspecific names for entities that are capable of very clear definition. It is of
course sometimes possible, using multivariate techniques, to define objective infra-
specific entities, even although there are no consistent individual character differences
between the forms concerned (see, for instance, Thorpe's (1975) treatment of the
east and west populations of Natrix natrix), but as no sophisticated numerical
methods have been used in the study of Scincus, no attempt to discover such hidden
entities has been made here.

MATERIAL EXAMINED

Material looked at in the course of this study is listed at the end of each species
description. Data in brackets were not originally attached to the specimens, but
have been added by the present authors. Localities for which no co-ordinates are
given can be found in The Times Atlas of the World, Mid Century Edition. Where
the spelling differs markedly from that attached to the specimens the Times Atlas
spelling is given in parentheses. Abbreviations used are as follows :

BM British Museum (Natural History), London

AMNH American Museum of Natural History, New York

CAS California Academy of Sciences, San Francisco

CM Carnegie Museum, Pittsburgh

EBD Estacion Biologica de Donana, Seville

FMNH Field Museum of Natural History, Chicago

HUJ Hebrew University of Jerusalem

IFAN Institut Fondamental d'Afrique Noire, Dakar

MCZ Museum of Comparative Zoology, Harvard

192 E. N. ARNOLD AND A. E. LEVITON

MF Museo Zoologico dell'Universita di Firenze

MHNP Museum Nationale d'Histoire Naturelle, Paris

MSNG Museo Civico di Storia Naturale 'Giacomo Doria', Genoa

NMW Naturhistorisches Museum, Vienna

SMF Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt a.M.

USNM United States National Museum, Washington

ZMB Zoologisches Museum an der Humboldt-Universitat zu Berlin

ZSM Zoologische Staatssammlung, Munich

ad. Adult of undetermined sex

juv. Juvenile (arbitrarily denned as any Scincus under 70 mm)

* Ear dissected

As the manuscript was almost entirely written at the British Museum (Natural His-
tory), the greater part of the material, including that from most other institutions, was
examined there. However, California Academy of Sciences specimens and a few
individuals belonging to the Field Museum of Natural History, Chicago, were checked
by the junior author at the former institution ; unlike the rest of the material, these
were not sexed.

Specimens used for checking osteological features are in the collection of the British
Museum (Natural History) ; radiographs and a list of the individuals employed are
also deposited there.

The number preceding each sample correlates with those indicating localities on
the distribution maps (Figs 5-7).

CHARACTERS ALREADY USED OR POTENTIALLY USEFUL IN
CLASSIFYING SCINCUS

It will become apparent that Scincus can best be regarded as three well-defined
entities, as follows :

1. 5. hemprichii.

2. 5. mitranus (including 5. arabicus, S. arenarius, S. muscatensis, S. philbyi and
5. richmondi).

3. The S. scincus complex, which consists of two apparently well-defined subspecies
east of the Red Sea and a less easily analysed series of populations in northern Africa
(for which the following names are presently in use : 5. s. scincus, S. s. cucullatus,
5. s. laterimaculatus and S. albifasciatus). The two eastern subspecies are S. s.
meccensis (including 5. deserti) and 5. s. conirostris (including S. gasperetti).

To simplify description of characters, the forms listed inparentheses will be in-
cluded in the taxa recognized above unless otherwise stated. For the present,
North African populations of the S. scincus complex will be referred to informally as
such.

The ear and associated structures

The size and position of the external ear opening have long been used in the clas-
sification of Scincus. Differences in these features turn out to correlate with more
profound modifications of the ear and surrounding parts of the skull. These may

REVISION OF SCINCUS 193

well represent varying degrees of adaptation to the problems of subterranean hearing
and spatial orientation.

The ear of Scincus corresponds fairly closely to the typical lacertilian pattern
described by Baird (1970). However, it is slightly anomalous in having a relatively
long external auditory meatus that runs forwards and downwards to a small exterior
aperture. Features of the ear and associated structures that exhibit variation
within Scincus are described below and illustrated in Figs 1-4.

EXTERNAL ORIFICE OF THE EAR. 5. scincus complex. The external orifice of the
ear is relatively large and its upper margin is situated just above, or almost level
with, a continuation of the line made by the free edges of the upper labial scales. It
is nearly always covered by two flat, closely connected scales that have their posterior
borders strongly serrated to produce a fringe ; in most cases the free upper edge of
the lower scale overlaps the lower edge of the upper one. Serration is very variable,
but often two principal lobes can be discerned on each scale.

In a few instances, the upper part of a third scale is involved in the lowermost
region of the fringe and, very rarely, there is only a single large scale covering the ear
orifice.

S. mitranus. Here the ear orifice is considerably smaller than in the 5. scincus
complex, and its upper margin is nearly always well below the level of the line made
by the free edges of the upper labial scales. The orifice is also often displaced
forwards when compared with that of S. scincus. As in this species, it is covered by
two scales in most cases, although since the orifice is very small, it is only the con-
tiguous parts of the two scales that lie over it. The overlying area is again usually
serrated although the denticulation is typically far less than that found in the 5.
scincus complex. The two scales covering the ear sometimes coalesce and this
fusion may be so complete that they appear as a single scale, which may have only
very reduced serration on its posterior margin.

S. hemprichii. In this species, the ear orifice is minute and situated well below the
level of the line made by the free edges of the upper labial scales. It is placed deep
beneath a single large, unserrated scale which may have arisen by fusion of two smaller
scales (as in some S. mitranus, see above).

EXTERNAL AUDITORY MEATUS. S. scincus complex. The meatus is relatively
short and, distally, is quite thick-walled and lined by obvious, although unossified,
well-spaced scales, the overlapping edges of which are directed towards the external
orifice. More proximally, the meatal wall becomes membranous, except for a
cartilaginous plate that is incorporated into its lateral surface and originates from the
superior outer border of the quadrate. This plate tapers distally to a point and is
relatively short. The more proximal part of the meatus is usually well covered with
muscle and clearly separated from contact with the overlying skin.

S. mitranus. The meatus is considerably longer than in the 5. scincus complex.
As in this species, its distal portion is lined with scales, but these are less well
developed and the meatal wall in this area is thinner. The proximal part of the
meatus is again membranous, except for the cartilage in its outer (lateral) wall which
is much longer and more pointed than in the 5. scincus complex. The lateral wall of

194

E. N. ARNOLD AND A. E. LEVITON

the proximal part of the meatus lies close to the skin and is only separated from it by
a thin layer of muscle.

S. hemprichii. The meatus is fairly long (although rather shorter than in S.
mitranus). The lower section is very thin-walled and either lacks scales or has only
scattered traces of them. The cartilaginous plate in the membranous proximal part
of the meatus is relatively short and broad. This area lies close to the skin.

m

FIG. i. S. scincus: ear and associated region of skull, (a) Head showing position of
external orifice of ear. (b) Diagrammatic exposure of outer ear and hind part of skull.
Abbreviations (reading clockwise from top) are as follows : p = supratemporal process
of parietal ; s = squamosal ; o = otic capsule ; q = quadrate ; m = external auditory
meatus (lateral wall removed) ; t = tympanum ; e = extracolumella ; st = supratemporal.
(c) External auditory meatus showing cartilage (c) in lateral wall. (Based on BM
1909.7.28.15 ; El Oued, Algeria ; (J.)

FIG. 2. 5. mitranus: ear and associated region of skull : compare with Fig. I.
(Based on BM 1970.1594 ; Sharjah, United Arab Emirates ; <J.)

REVISION OF SCINCUS

195

FIG. 3. 5. hemprichii: ear and associated region of skull : compare with Fig. I.
(Based on BM 95.5.23.77 ; Haithalhim, Aden ; $.)

QUADRATE AND TYMPANUM (Fig. 4). The quadrate bone and the tympanum are
largest (when compared with skull length) in the 5. scincus complex. In S. mitranus,
they are both slightly smaller and the tympanic crest of the quadrate is better devel-
oped. 5. hemprichii has a smaller quadrate still and the tympanum is further reduced
so that it is smaller than in the other two species, both relative to skull length and to
quadrate size. The quadrate in S. hemprichii usually has a more prominent antero-
superior projection than the other species.

FIG. 4.

a b c

Right quadrates of (a) S. scincus, (b) S. mitranus and (c) 5. hemprichii
lateral view. (Based on specimens used for Figs 1-3.)

EXTRASTAPES AND STAPES. S. scincus complex. The extrastapes has a distinct
pars superior and pars inferior embedded in the tympanic membrane. Together,
these partes form a clearly defined, slender bar running gently forwards and often
downwards from the posterior border of the tympanum (where the pars superior is
attached) to beyond its centre. The stapes is slender and terminates in a relatively
small footplate at the foramen ovale.

S. mitranus. The pars superior and pars inferior are expanded to form an elongate
cartilaginous plate that involves a comparatively large area of the tympanum. The
size of the plate varies somewhat but it is always relatively larger than the combined
pars superior and pars inferior of the S. scincus complex. Also, the stapes of 5.
mitranus is slightly more robust and has a larger foot-plate.

5. hemprichii. The combined pars superior and pars inferior is larger still and
forms an often irregular, ovoid, cartilaginous plate that may occupy all but the

IQ6 E. N. ARNOLD AND A. E. LEVITON

periphery of the tympanum. The stapes of 5. hemprichii is much more robust and
has a larger foot-plate than other species of Scincus.

OTIC CAPSULE. 5. scincus complex. The otic capsule is fairly large in young
animals but decreases in relative size with growth. In adults, its posterior border is
not, or only slightly, visible in lateral view.

5. mitranus. Here the otic capsule is about the same length or only slightly larger
than that of equivalent-sized members of the S. scincus complex, but its posterior
border is often clearly visible in lateral view, partly because the squamosal and the
supratemporal process of the parietal usually descend at a more abrupt angle in this
species.

S. hemprichii. This species has an immense otic capsule that may occupy a
substantial proportion of the space beneath the supratemporal process of the parietal
which is more usually occupied by parts of the jaw musculature. The capsule
extends about as far forwards as the epipterygoid bone and projects backwards well
beyond the confines of the skull.

Otic enlargement in Scincus mainly involves the upper part of the capsule con-
taining the semicircular canals. The lower section shows much less variation in size.
In animals with a skull length of about 20 mm (excluding rostrum) the otic capsule
is about a quarter of this length in the S. scincus complex and in S. mitranus and well
over a third in S. hemprichii.

SQUAMOSAL BONE. 5. scincus. In adults, this bone curves gently forwards and
upwards to meet the jugal.

S. mitranus. Here, the upper border of the squamosal is much more sharply
angled than in S. scincus. Where it contacts the supratemporal process of the
parietal, the upper border may sometimes turn upwards to form a small spur.

S. hemprichii. In this species, squamosal flexure is even more pronounced and an
upward spur bordering the supratemporal process of the parietal is present and
better developed than in any S. mitranus.

These differences are best marked in adult animals but juveniles also show them
to some extent.

SUPRATEMPORAL BONE AND SUPRATEMPORAL PROCESS OF THE PARIETAL. The
supratemporal bone is relatively small in the S. scincus complex and largest in S.
hemprichii.

The supratemporal process of the parietal extends roughly to the level of the
suture between the supraoccipital and exoccipital bones. In the 5. scincus complex,
its ventral tip is almost level with that of the squamosal, while in S. mitranus it is well
above this. In S. hemprichii the surface exposure of the process becomes acutely
narrowed above the level of the upper edge of the squamosal, but continues down-
wards for some way as a narrow spur, although this is not always visible in lateral
view.

Premaxillary rostrum

In Scincus, the premaxilla projects forwards as a bony rostrum that supports the
spatulate snout tip. The rostral part of the premaxilla is strengthened on each side

REVISION OF SCINCUS 197

by a flange arising from the maxilla. The extent of the rostrum is not easy to assess
accurately without dissecting the snout, but on the basis of radiographs and a
limited number of skull preparations, it appears to vary as follows :

5. scincus complex : rostrum moderately well developed (least so in 5. s. coni-
rostris}.

S. mitranus : rostrum extremely well developed.
S. hemprichii : rostrum poorly developed.

Eyes

The eyes of most Scincus have an approximately round pupil and a relatively
small cornea (horizontal diameter about 32-39% of total diameter of eye). In S.
hemprichii, however, the contracted pupil is a vertical slit and the cornea is con-
siderably larger (horizontal diameter about 47-58% of total diameter of eye).

Variations in scaling

Terms used in the literature for particular head scales of Scincus have varied. The
scale immediately behind the nasals has been called either the first loreal or the post-
nasal and both internasal and supranasal have been employed for those scales that
wholly or partly separate the rostral from the frontonasal. In this account, first
loreal and internasal will be used.

ROSTRAL-FRONTONASAL CONTACT. In most cases, the rostral scale contacts the
frontonasal, thus separating the two internasals. The width of such contact varies.
Contact is absent in S. hemprichii, in nearly all 5. s. conirostris examined and, very
rarely, in other populations of the S. scincus complex as well.

PREFRONTAL FUSION. Most individuals of Scincus have two separate prefrontal
scales, fusion usually being rare. However, in northern populations of S. mitranus,
fusion is by far the commonest condition.

NUMBER OF SUPRAOCULAR SCALES. In nearly all cases, members of the genus
Scincus have six supraocular scales on each side. Infrequently, two of the scales,
usually the second and third, fuse and reduce the supraocular count to five, either
unilaterally or on both sides. This is a rare variant in many populations but it is
more common in Arabian S. scincus : it occurs in 15 out of 28 5. s. meccensis and in
eight out of 28 5. s. conirostris. The number of supraocular scales has been used in
defining nominal species of Scincus, for instance 5. gasperetti is distinguished by this
feature, but the fact that animals with five or six supraoculars over each eye may
occur together in the same populations suggest that this character is not of great
importance in itself.

FRONTOPARIETAL CONTACT. In most Scincus, the frontoparietal scales are nearly
always in contact, usually broadly so, but in 5. hemprichii this contact is reduced or
even absent.

SIZE OF PARIETAL SCALES. The parietal area of the skull in Scincus is covered by a
series of paired scales bordering the interparietal on each side ; two, three or more

ig8 E. N. ARNOLD AND A. E. LEVITON

pairs are usually involved. Schmidt (1939), in describing his S. arabicus (= S.
mitranus}, refers to all these as parietals : 'parietals broken into small transverse
scales, three on the left, two on the right'. However, all but the first pair, which is
often enlarged, are not easily distinguishable from the paired mid-dorsal scales on the
neck and it seems most convenient to refer to all these as 'nuchals' and confine
'parietal' to the first pair, which contacts the frontoparietals.

In the S. scincus complex and S. hemprichii, the parietal scales are usually con-
spicuously larger than the contiguous nuchals, but in S. mitranus they are relatively
small.

LOREALS. Loreal scales exist in three main conditions in Scincus : (a) three
loreals increasing in size posteriorly, the first of which is separated from the fronto-
nasal by contact between the internasal and the second loreal ; (b) three loreals
increasing in size posteriorly, the first of which contacts the frontonasal scale ;
(c) two loreals, the first scale in conditions (a) and (b) being absent.

Condition (a) is widespread. Condition (b) occurs as a relatively rare variant in
many populations but is the common condition in the 5. scincus complex in parts of
western Africa, etc., in S. s. meccensis and in S. mitranus from 'Muscat' and Masirah
Island. Condition (c) is also widespread as a rare variation but is the common con-
dition in northern populations of 5. mitranus.

DEPTH OF UPPER LABIALS RELATIVE TO LOREALS. If the depth of the third upper
labial is compared with that of the penultimate loreal (both depths being taken
perpendicular to their common suture) considerable variation is encountered. The
labial is only about 1-3 to 2-2 times as deep as the loreal in the 5. scincus complex
whereas it may be 2-3 to 3-0 times as deep in S. hemprichii. In 5. mitranus it is
about 1-5 to 2-8 times as deep.

MID-DORSAL SERIES OF SCALES ON TAIL. There are two mid-dorsal series of scales
on the base of the tail, but these are replaced by a single, laterally expanded, mid-
dorsal series distally. In S. hemprichii, this midline series extends far forwards,
always well beyond the extremities of the posteriorly adpressed hind limbs. In the
5. scincus complex, it is often reduced to a few scales on the tail-tip and only occasion-
ally extends to the level of the extremity of the adpressed limbs. In S. mitranus, it
is variable but tends to be intermediate in extent between S. hemprichii and S. scincus.

BODY SCALES. Various aspects of body scaling have been used in defining species
of Scincus. They include the number of scales around the mid-body, the size of the
mid-dorsal scales compared with the ventrals, and the presence or absence of longi-
tudinal ridging on the dorsals.

Mid-body counts range from 22 to 31. In most animals, the count is even, but a
substantial minority have odd counts (often about 10% of a population, but up to
20% may be involved). Counts vary in the S. scincus complex from 24 to 30, in
5. mitranus from (25) 26 to 30 (31) and in S. hemprichii from 22 to 24. In the 5.
scincus complex and 5. mitranus, there is some geographical variation and this is
summarized on pp. 210, 220 and 228.

Actually, the number of scale-rows across the mid-belly is constant at eight,
including the row covering the lateroventral ridge on either side. This means that

REVISION OF SCINCUS 199

variation in the number of scales around the mid-body is due only to variation in the
number of dorsal scales. As might be expected from this, animals that have high
mid-body counts tend to have mid-dorsals that are relatively small compared with
the mid-ventrals. Thus the largest mid-dorsals occur in 5. hemprichii and in some
western populations of the 5. scincus complex, both of which have low mid-body
counts. These are the only populations to have longitudinal ridging on the dorsal
scales. It is confined to adults and in the 5. scincus complex it is extremely weak.

Size and shape of head

SIZE OF HEAD. Head proportions are not easy to measure accurately in intact
specimens of Scincus. The back of the skull is usually well covered by soft tissue,
and this, together with the often thick osteodermal layer of the skin, makes the
precise location of reference points very difficult. However, some idea of the relative
differences in head proportion can be conveyed by making approximate measure-
ments.

In all forms, relative head length at first falls steadily with growth, but in larger
animals this trend sometimes reverses and the head begins to increase in relative
length. Such increase is much more pronounced in males, as these grow considerably
larger than females. It is very marked in 5. hemprichii but scarcely apparent in
S. mitranus. The S. scincus complex is rather variable, increase in head size being
marked in S. s. conirostris but weaker in other populations. As might be expected
from this, relative head lengths of adults are least in 5. mitranus (19-24% of snout
to vent length, in animals over 80 mm), greatest in S. hemprichii (24-30% in animals
over 80 mm), and intermediate in the S. scincus complex (about 21-26%).

Head width may increase relative to head length. This is most apparent in S.
mitranus, S. s. conirostris and S. hemprichii. As the last two forms are relatively
large-headed, this results in their having heads conspicuously wider than the body.

SHAPE OF SNOUT. Snout shape is distinctive in some forms although there is
often considerable variation within taxa (for instance, as Mertens, 1972, notes for
west African members of the S. scincus complex - under the name S. albifasciatus] .
In S. hemprichii the snout is very broad with no obvious canthi rostrales, and in
profile from above the sides are straight or slightly convex. In other forms the snout
is narrower, canthi are well developed and the sides are often slightly concave. The
snout tends to be rather short in 5. s. conirostris and overhang of the lower jaw by
the rostral is least in this form and in 5. hemprichii.

Colouring

DORSAL PATTERN OF YOUNG. With the exception of 5. hemprichii, populations of
Scincus have young with a fine, homogeneous dorsal pattern and no bold markings.
Usually, the ground colour is some pale shade of buff, grey, brown or cinnamon.
This may be virtually uniform, or the scales may have darker edges, or they may be
dappled with light and dark markings, which are sometimes confined to the scale-
edge (there often being one or two small, dark, spots bordered by light pigment), or
may involve a much greater area of the scale.

200 E. N. ARNOLD AND A. E. LEVITON

S. hemprichii is peculiar in that its young have a bold dorsal pattern. Most scales
are brownish, with a dark edge, but these are interspersed by transverse bands of
light-edged scales.

DORSAL PATTERN OF ADULTS. Adult animals may retain the juvenile pattern
without much change (at least as discernible in spirit material), apart from some
modification in contrast. For example, in S. hemprichii, there may be some fading,
especially of the dark scale-edges. A fine homogeneous pattern is retained by 5.
mitranus, some S. scincus conirostris, some 5. s. meccensis and some members of
the 5. scincus complex from North Africa (animals assigned to S. s. laterimaculaus,
and many females of populations usually assigned to 5. s. cucullatus). In other
members of the Scincus scincus complex, the juvenile colouring differentiates to
produce a bold pattern visible in spirit material. Differentiation involves either
the spread of dark pigment in restricted areas of the dorsum or loss of dark pigment
to form partly translucent areas that are often brightly coloured in life, or both. In
all cases, the pattern so produced consists of often irregular transverse bands.
In North Africa, with the exceptions listed above, dark transverse bands are
prominent in the pattern.

BARS ON FLANKS. In many populations of Scincus, adults have prominent vertical
bars or spots on the flanks. These do not disappear in alcohol and vary in colour
from pink, through red and maroon, to a deep warm, blackish-brown. They are
absent in the young and develop first on the foreparts. Flank bars usually occur in
5. mitranus, 5. s. meccensis and in the African populations of the 5. scincus complex
to which the names albifasciatus and laterimaculatus are often assigned.

TAIL PATTERN. In most Scincus, the fine, dappled pattern on the body becomes
weaker on the dorsum of the tail, but in some northwest African populations (usually
assigned to S. s. laterimaculatus) it may become more pronounced in adults.

SYSTEMATIC ANALYSIS

Of the characters listed in the previous section, those involving the ear and
associated parts of the skull are very stable within populations. This character
complex exists in three clearly separable states and, as these often correlate with some
other morphological features, it seems certain that they indicate systematic entities.
Two of these are very clearly defined species, 5. mitranus and 5. hemprichii, both
confined to Arabia. The third covers virtually the whole range of the genus and
may consist of more than one species, so it will be referred to as the S. scincus
complex.

North Africa and Israel

All the Scincus of this area belong to the 5. scincus complex and are very similar
in general morphology, although there is considerable local variation in colouring
and some features of scaling. As will be shown, there is at least one area where
populations exhibiting consistent differences may possibly come into contact ; this
is discussed further on p. 215.

REVISION OF SCINCUS 201

Arabia, Jordan, Iraq and Iran

Northwestern Arabia and nearby southern Jordan are occupied by a form
generally similar to 5. scincus s. sir. of Egypt and Israel, with which it should be
regarded as conspecific. All its characteristic features are found in one or other of
the populations that constitute the North African 5. scincus complex, but it differs
from the nearest ones in the following features : adults lack dark transverse bands
on the back, dark bars or spots are present on the flanks of adults, the first loreal
almost always contacts the frontonasal and the supraoculars are often reduced from
six to five on each side. It seems reasonable to treat this form as a distinct sub-
species, 5. scincus meccensis. The range of this taxon approaches closest to Israeli
and Egyptian populations of Scincus in the region of Wadi Araba, which joins the
Gulf of Aqaba to the Dead Sea. This wadi often separates the ranges of related
reptiles that are distinct at the species or subspecies level (Werner, 1971).

In parts of southern and eastern Arabia, Iraq and southwestern Iran, another form
generally similar to 5. scincus occurs. This differs from neighbouring 5. s. meccensis
in lacking dark bars or spots on the flanks, in having the frontonasal separated from
both the rostral and the first loreal and in possessing a wider head, at least in adult
males. There is no evidence that this form is sympatric with 5. s. meccensis
(although it must be remembered that relatively few 5. scincus-\ike skinks have
been collected in Arabia to date), so it is regarded here as a further subspecies,
S. s. conirostris.

In southern and eastern Arabia, another form is found, for which the oldest name
is S. mitranus. There is no doubt that it is specifically distinct from S. scincus, for it
coexists with 5. s. conirostris at several localities (Kuwait, Abqaiq, Dhahran,
eastern United Arab Emirates, Hadhramaut) and differs from this form in many
features. Apart from those characters set out in Table 5 (p. 240) that distinguish the
5. scincus complex as a whole from 5. mitranus, the latter differs from the sympatric
5. s. conirostris in having the rostral scale in broad contact with the frontonasal, a
broader snout-tip, a relatively smaller head, at least in adult males, and dark bars or
spots on the flanks of mature animals.

The extreme southwest of the Arabian peninsula is occupied by a single form of
Scincus, S. hemprichii. It appears to be separated from 5. mitranus and 5. s.
conirostris by mountainous country unsuitable for Scincus, and there is a gap of over
500 km between its northernmost locality on the coast and the nearest known
populations of 5. s. meccensis. This hiatus contains geographical features that may
be important in isolating the two forms from each other (see p. 242). However,
although it may well be completely allopatric, S. hemprichii has so many distinctive
features that it must be regarded as a fully differentiated species (see Table 5, p. 240).

SCINCUS Laurenti

Scincus Laurenti, 1768 : 55.
Pedorychus Wiegmann, 1837 : I2^-

Scincus (Arenatius) listed by Haas & Battersby, (1959) without comment is a lapsus calami for
5. arenarius and was not intended as a subgenus.

TYPE SPECIES. Lacerta stincus (lapsus calami for scincus) Linnaeus, 1758.

202 E. N. ARNOLD AND A. E. LEVITON

DIAGNOSIS. A genus of skinks with a full scincomorph complement of skull
bones. Palatines and pterygoids widely separated on the midline of the palate,
premaxilla single, postorbital and postfrontal bone fused, nostril situated largely or
completely between two nasal scales. Limbs well developed. Numerous features
associated with life in loose sand present : a wedge-shaped head with spatulate,
overhanging snout supported by a premaxillary rostrum ; countersunk jaw ; angular
upper labial scales ; valvular, often crescentic nostrils ; small ear orifices with long
meatus ; Q-shaped body section ; powerful body musculature ; and depressed,
fringed digits. Dorsum uniform, with fine dappled pattern or with transverse
bands. Adult males considerably larger than adult females.

HABITUS. Rather robust, with poorly defined neck and well-developed limbs.
Head wedge-shaped in lateral view and, seen from above, tapers forward from broad,
temporal region. Snout spatulate, overlapping the lower jaw anteriorly and at the
sides, its tip forming a sharp, horizontal edge. Upper labials very strongly angled
in cross-section, the ridge so formed being continuous with the sharp edge of the
snout-tip. Front of the lower jaw is countersunk, so that the underside of head is
almost flat. Canthus rostralis usually present, although not always very distinct.
Eyes relatively small. Ear apertures small, or very small, their upper border
situated just above, on, or below a continuation of the line made by the lower edges
of the upper labial scales. Neck and body Q-shaped in cross-section, producing a
latero ventral ridge on each side that runs from beneath the jaw to the groin. Tail
relatively short, usually less than 80% of snout to vent distance, a tapering cylinder
that may be laterally expanded proximally and is often rather compressed distally.
Forelimb extends forwards to about level of eye. When fore- and hind-limbs
adpressed, fingers and toes overlap. Digits dorsoventrally compressed.

SCALING. Rostral broad, spatulate and sharp-edged, often bordering the approxi-
mately hexagonal or pentagonal frontonasal, but sometimes separated from it by
midline contact of the internasals. Paired prefrontals usually meet, but are oc-
casionally fused (very frequently so in northern populations of S. mitranus) or, more
rarely, separated by a small azygos scale. Frontal wider in front than behind,
often markedly so. Frontoparietals typically in broad contact (but contact limited
or even absent in many 5. hemprichii and in occasional specimens of other forms).
Interparietal large, more or less lozenge-shaped, with a clearly defined parietal 'eye' ;
it separates the typically quadrangular parietals, which are usually larger than the
contiguous nuchals (sometimes subequal in S. mitranus). Usually six supraoculars
(second and third sometimes fused). Supraciliaries typically four on each side, but
number may vary from two to five.

Nostril oval or crescentic, situated just above the sharp ridge formed by the rostral
and upper labials, placed between an upper and lower nasal scale and sometimes also
making narrow contact with the rostral anteriorly. Usually three loreals that in-
crease in size posteriorly : first may reach frontonasal, but separated from it, in
majority of cases, by contact between internasal and second loreal. First loreal
sometimes absent or fused with second. Upper labials usually eight on each side
(but number varies from seven to nine), typically very deep posteriorly where they

REVISION OF SCINCUS 203

may project well below tips of maxillary teeth. The anterior upper labials are
strongly angled, to form a sharp ridge continuous with that of the rostral. A single
row of sub- and postoculars borders eye. Lower eyelid covered by translucent
scales, some of them large. Temporal formula : 1 + 2.

Mental single. Typically eight lower labials on each side and two azygos post-
mentals followed by about three pairs of large scales, each pair usually being divided
on the midline by a smaller scale.

Small ear aperture often protected by two (rarely three) overlapping, denticulated
scales, or by a single, large, often smooth-edged scale.

Laterally expanded nuchal scales up to about twice the width of neighbouring
scales ; typically four or five pairs present, but number may vary from nil to ten on
each side. Scales on rest of neck and on body cycloid, either all smooth or dorsals
with two or three weak ridges.

Mid-dorsals about as large as mid-ventrals, although often slightly bigger or smaller.
22 to 30 (31) scale rows round mid-body, of which eight cover the belly, including the
lateroventral ridges. Even mid-body counts are commonest, but odd ones do occur
(often in about 10% of a population). Vent covered by two large, flat shields and
bordering it on each side of the tail-base is a small, raised, often sharp-edged scale.
Scales on tail mainly cycloid, but central row on underside is laterally expanded.
A less expanded row exists on the mid-dorsal line, but is confined to the more distal
parts of the tail and is sometimes restricted to the tip.

Scales on limbs cycloid and regular. Digits dorsoventrally flattened and covered
by an upper and a lower row of lamellae. On most digits, the upper row projects
posteriorly and the lower anteriorly, to produce fringes of pointed lamella tips, but
on the fifth toe of each forelimb, the upper row projects anteriorly and the lower
posteriorly. On all toes the posterior fringes are better developed than the anterior
ones. Ungual lamellae strongly expanded ; claws flattened and hollowed beneath.

COLOUR AND PATTERN. In one species (5. hemprichii), adults and young are
brown above with narrow, light, transverse bands. Other species have young that
are uniform (ivory, cream, grey, buff, warm brown or cinnamon) or finely dappled
with small, light and dark markings. This homogeneous pattern may be retained
and often matches the sand in which the animal lives. In other cases, the fine,
dappled pattern becomes overlaid by a series of sometimes irregular, transverse
bands, either dark or light, or both. There is considerable variation in the number,
extent and intensity of these bands ; they may be narrow or broad and do not always
extend right across the body. Flanks of adults may bear a series of usually dark,
vertical bars or spots that often diminish in size posteriorly and alternate with the
dark, dorsal bands, where present.

Often a rather obscure, dark bar in the area of the canthus rostralis, which may
continue backwards onto the cheek and neck and is sometimes strongly developed.
Dorsum of tail may be more, or less, heavily marked with dark pigment than back,
or, distally may be contrastingly coloured bluish or reddish, especially in young.

Underside uniformly pale ; white, cream or greyish. In life, many animals have
some yellow or pale orange pigment, especially on feet and flanks and often in the
dorsal area as well. This disappears in spirit.

204 E. N. ARNOLD AND A. E. LEVITON

Iris dark brown or blackish in life ; tongue apparently always pink.

EYE. Usually 14 scleral ossicles. Cornea about 30-60% of horizontal diameter
of eye. Pupil more or less round or vertical, when contracted.

EAR (Figs 1-3). External auditory meatus long or very long, sloping downwards
and forwards. Distal section usually with at least some traces of scaling, proximal
section membranous with a triangular cartilage in its outer wall. Extracolumella
varies in form from a relatively slender bar to a large, irregular plate. Stapes
slender or relatively robust.

HEMIPENIS. Symmetrically and shallowly bilobed, with at least some plication
on the lobes.

SKELETON. Skull. Full scincomorph complement of skull bones present.
Premaxilla single with three or four teeth, projecting forward to form a bony rostrum
that receives support on each side from a maxillary flange. Paired nasal bones
extend anteriorly only as relatively narrow spurs on each side of the posterior
process of the premaxilla. Nasal cavity uncovered anteriorly and front of septo-
maxilla clearly visible from above. Frontals paired, each contacting a maxilla
anteriorly, thus separating nasal and prefrontal bones. Lateral processes of frontal
not well developed. Parietal single with well-defined central foramen, its lateral
descending processes long and finger-like.

Supratemporal process of parietal may extend almost to level of ventral tip of
squamosal, or not. Supratemporal bone variable in size.

Postorbital and postfrontal bones fused even in young animals, not extending
backwards to cover the posterior part of the Supratemporal fossa. Squamosal varies
in shape and runs forward to contact jugal. Maxillae with pleurodont, simple
teeth, about 15-20 in adults, but counts are lower in juveniles.

Vomers separate, but in contact on midline, rather elongate. Palatines closely
opposed (although not touching) above air passage, widely separated on palate, not
usually contacting ectopterygoids along posterior edge of suborbital foramen
(Greer, 1970, states that they do so in material of S. scincus examined by him.)
Pterygoids with teeth. Otic capsule normal, or strongly expanded. Osteodermal
layer covers most of upper surface of skull and the supraorbital areas.

Mandible. Meckel's groove open. No fusion between the main elements. About
13-20 teeth present in adults, but counts are lower in juveniles.

Poster anial skeleton. Usually 27 procoelous presacral vertebrae (rarely 26 or 28),
with rather long neural spines that are expanded sagittally. Eight cervical ver-
tebrae, of which numbers one to five have long hypopophyses and numbers four to
eight bear ribs (first three short and flattened, last two long and cylindrical). All
other presacral vertebrae bear dorsal ribs, the last four to six being shorter than the
rest, although the reduction in length is not very abrupt.

Three sternal ribs and two xiphisternal ribs present. Sternum lozenge-shaped,
nearly always without a central fontanelle. Xiphisternal cartilages arise close
together, diverge to the origin of the first pair of xiphisternal ribs and then converge
to join at the origin of the second pair. A single pair of inscriptional ribs may occur
immediately posterior to the xiphisternum. It is present and attached to the dorsal

REVISION OF SCINCUS 205

ribs in S. hemprichii, whereas in S. mitranus it usually forms a free chevron on the
midline and is sometimes absent. In the 5. scincus complex, it may be attached,
free or lacking. Clavicle laminar, either expanded throughout its length or, more
usually, expansions confined to the region of contact with the lateral arm of the
interclavicle and to the medial extremity, where it is fenestrated. The posterior
margin may be very irregular with one or more projections. Interclavicle cruciform,
the lateral arms being almost at right angles to the longitudinal axis or, more fre-
quently, clearly directed forwards.

Hyoid apparatus consists of glossohyal, hypohyal and ceratohyal, first cerato-
branchial and a short second ceratobranchial.

OSTEODERMS. These have the usual scincid pattern of canals (see, e.g., Greer,
1970).

SIZE. Adult males considerably larger than females. Largest about 140 mm
snout to vent, but males in some populations not much exceeding 120 mm. Females
on average about 75-85% length of males. Smallest hatchlings are about 38 mm
snout to vent.

DISTRIBUTION. Sahara and neighbouring areas eastwards to east Arabia and
southwest Iran. Northward limit runs south of Atlas Mountains, along the coast of
Tunisia, Libya and Egypt, through southern Israel, southern Jordan, central Iraq
and southwest Iran. Southernmost parts of distribution include Senegal, Mali,
Niger, northwest Nigeria, southeast Libya, central Egypt and southern Arabia.
May possibly occur in Eritrea (p. 235) and has been erroneously stated to be found in
Pakistan (p. 229).

SPECIES REFERRED. 5. hemprichii, S. mitranus, S. scincus, complex (which may
possibly include two largely allopatric species, 5. scincus and S. albifasciatus) .

KEY TO Scincus

a. Ear opening clearly visible and relatively large, its upper border reaching the back-
ward continuation of the line made by the lower edges of the upper labial scales.
External meatus relatively short, meatal cartilage short, extracolumella a slender bar,
squamosal gently curved, otic capsule normal .... 5. scincus complex

b. Ear opening very small but usually visible, nearly always situated well below the line
made by the lower edges of the upper labial scales. Dorsum more or less uniform.
Vertical bars or spots usually present on flanks of adults and rostral in broad contact
with frontonasal scale (neither character apparently found in sympatric populations
of S. scincus). External meatus very long, meatal cartilage long, extracolumella
expanded, squamosal angled, otic capsule normal. South, central and eastern Arabia

only ............ 5. mitranus

c. Ear opening minute, not visible externally. Dorsum usually with light, transverse
bands. Vertical bars or spots absent from flanks and rostral separated from fronto-
nasal scale. External meatus long, meatal cartilage short, extracolumella greatly
expanded, squamosal angled, otic capsule huge. Extreme southwest Arabia only

S. hemprichii

206 E. N. ARNOLD AND A. E. LEVITON

The Scincus scincus complex

Lacerta stincus (lapsus calami for scincus) Linnaeus, 1758 : 205. Type locality : 'in montosis
Lybiae, Aegypti, Arabia petreae'. Location of type : Naturhistoriska Riksmuseet, Stockholm
(Registered number : I4ia).

Scincus officinalis Laurenti 1768 : 55 (nomen substitutum).

For other names applied to S. scincus, see discussion of geographical variation and subspecies

(p. 209 et seq.).

DIAGNOSIS. Head variable in size but tending to be intermediate between that
of S. mitranus and 5. hemprichii ; external ear orifice relatively large, its upper
margin reaching the continuation of the line made by the lower edges of the
upper labial scales and typically covered by two serrated scales. Eye with relatively
small cornea and rounded pupil. Rostral scale usually borders frontonasal, at least
narrowly (but not in 5. s. conirostris, or in rare examples of other populations) ;
frontoparietals in usually extensive contact ; parietals relatively large (usually 1-5
to 2-5 times length of adjoining nuchals). Dorsal scales usually smooth, but very
weakly ridged in some adults, mid-dorsals subequal to mid-ventrals, but sometimes
distinctly larger, or slightly smaller. 24 to 30 scales around mid-body. Young
either almost uniform or with fine, dappled pattern ; this may be retained by adults
or converted into a system of dark or light transverse bands, or both ; dark bars or
spots may be present on the flanks of adults in some populations.

Auditory meatus comparatively short ; its distal section thick-walled and lined
with well-developed scales ; cartilage in lateral wall short. Extrastapes a slender
bar. Premaxillary rostrum moderately developed. Squamosal gently curved, no
spur on its upper border. Distal section of supratemporal process of parietal not
constricted, terminating almost at level of ventral tip of squamosal.

EXTERNAL FEATURES. Head rather variable in size, often a little narrower than,
or as broad as, the neck and as the body at level of forelimbs, but considerably
broader in mature males from Egypt and in male S. s. conirostris ; length in animals
over 80 mm varies from about 21 to 26% of snout to vent distance, width is from
about 52 to 70% of head length. Top of head may rise gently in lateral profile, as
in S. mitranus, or more steeply and sometimes as much as in 5. hemprichii. Canthus
rostralis quite well marked ; loreal region distinctly concave. Tail broader than
deep proximally, laterally compressed - at least to some extent - distally. Up to
80% of snout to vent distance when undamaged.

Rostral overhangs mental strongly, sometimes as much as in 5. mitranus, least in
some S. s. conirostris ; in contact with frontonasal in most cases but not in nearly all
5. s. conirostris and rare individuals from other populations ; if present, such contact
is variable in extent, but may be as great as in 5. mitranus. Prefrontals in contact
(rarely fused, or separated by an azygos shield). Frontal broader in front than
behind. Frontoparietals usually in broad contact (but occasionally this is rather
restricted). Parietals relatively large, typically i| to 2\ times the length of adjoin-
ing nuchals but sometimes three times as long. Supraoculars usually 6 : 6, but five
supraoculars, either uni- or bi-laterally, occurs as a rare variant in North Africa and
more commonly in Arabian populations. Supraciliaries usually four on each side,
but number varies from three to five.

REVISION OF SCINCUS 207

Nostril laterally oval, or crescentic ; rostral completely excluded from its border
in the great majority of cases. Three (very rarely two) loreal scales on each side ;
first loreal does not contact frontonasal in nearly all individuals of many populations,
but reaches frontonasal in majority of animals from western North Africa etc. and
from northwest Arabia etc. Upper labials rather variable in depth when compared
with second loreal, usually 1-3 to 2-2 times as deep ; typically eight on each side,
but number varies from seven to nine. Ear orifice relatively large, its upper margin
situated slightly above, or level with, a continuation of the line made by the lower
edges of the upper labials ; covered by two (occasionally three) scales, the posterior
edges of which are strongly serrated to produce a fringe, often with four main lobes.

Typically three to five pairs of nuchal scales, but number on one side may vary from
one to ten. Dorsal scales usually smooth but may be very slightly ridged, at least
posteriorly, in some individuals from the extreme west of the range (Spanish Sahara
to Senegal) ; mid-dorsals often subequal to mid-ventrals but they may be rather
wider in western animals. 24 to 30 scales around mid-body. Row of laterally
expanded scales on dorsal midline of tail usually quite poorly developed, and in
great majority of cases does not extend to the level of the extremities of the pos-
teriorly adpressed hindlimbs, and only very rarely beyond.

The parts of the lamellae forming digital fringes are somewhat variable being
sometimes quite broad and sometimes mucronate. Claws long, and, like ungual
lamellae, often expanded laterally.

COLOUR AND PATTERN. Dorsum of young animals uniform, or with a dappling of
small dark or light spots, or both. Ground colour varies and may be ivory, pale
grey, straw, pale buff, pale brown or cinnamon. The dappled pattern typically
consists of one or two dark spots per scale, sometimes confined to the posterior edge,
where they may coalesce to form a dark border ; they are typically edged and
separated by pale pigment. These elements may sometimes extend forward to
cover much of the scale. A simple dappled dorsal pattern is retained by some
animals (e.g. northwest Algerian populations and some animals in Arabia) but it
often differentiates with maturity to produce a series of dark or light transverse
bands or both. Dark bands produced by spreading of the dark pigment are found
in many North African populations and vary considerably in width and number.
They may be much wider than the lighter interspaces or much narrower and they
vary in intensity. From the forelimbs to the vent, there may be five to ten bands ;
a further band is present on the posterior neck and this may or may not fuse with an
anterior nuchal patch that is usually visible. The dark transverse bands often
anastomose and are sometimes staggered on each side of the body. When poorly
developed, they may be most prominent in the dorsolateral region. Light bands
visible in spirit are produced most usually by loss of pigment from the scales and in
life are often yellow or deep orange. In North Africa they may alternate with the
dark bands, but in Arabia this is not so and they are often confined to the dorsolateral
region. Dappled pattern is often reduced on the tail but is intensified in northwest
Algerian animals. Flanks and limbs tend to be paler than the back and there may
be some dark (often greyish) pigment along the sides of the head and neck : this is very
marked in northwest Algerian animals where there is a distinct dark streak on each side.

208 E. N. ARNOLD AND A. E. LEVITON

In life, most animals have some yellow or orange pigment. Apart from in the
light transverse dorsal bands, when present, there is often a suffusion of yellow on the
feet, limbs and flanks.

Adults in some populations (western North Africa etc., northwest Arabia) have a
row of spots or short vertical bars on the flanks that vary from pale pink through red
to a very deep purple brown. They alternate with the dark dorsal bands, if any,
and begin in front of the forelimbs and may extend to the tail-base, although they
tend to diminish in size and intensity posteriorly. There is considerable local and
individual variation in their form, colour and arrangement. Typically there is one
in front of the forelimbs and from three to six between the two pairs of legs. These
markings are absent in hatchlings but may develop in animals as small as 62 mm.

Underside whitish or cream. Iris very dark brown or blackish.

EYE. Cornea relatively small, about 32-40% (n = 6) of horizontal diameter of
eye. Pupil rounded ; not a vertical slit when contracted.

EAR (Fig. i). Meatus relatively short, distal section quite thick- walled and lined
with scales. Cartilage in upper portion short and triangular. Tympanum relatively
large (compared with skull length) . Distal section of extrastapes forms a slender bar.
Stapes slender, foot-plate small.

SKULL (Fig. i). Premaxillary rostrum rather variable but better developed than
in S. hemprichii and less so than in S. mitranus. Squamosal gently curved, no upward
spur bordering supratemporal process of parietal. The latter is not constricted
ventrally and terminates at, or just above the level of, the ventral tip of the squa-
mosal. Supratemporal bone comparatively small. Otic capsule relatively small (in
animals with a skull length of 20 mm, it is about a quarter of that length) ; in lateral
view its posterior border is not, or only slightly, visible behind the supratemporal
bone and supratemporal process of the parietal. Maxillary teeth 16-20 in large
adults ; mandibular teeth about 15-20.

SIZE. Some geographical variation in maximum size. Largest animals are from
northwest Algerian Sahara (populations known as S. s. laterimaculatus) where males
may attain 140 mm (Werner, 1914). In other areas animals are smaller and in
Egypt and Arabia males do not exceed 120 mm. Females are about 80-85% snout
to vent length of males. See Tables 1-3 and subspecies texts for details of variation.
Smallest juveniles are about 40 mm snout to vent and animals up to at least 50 mm
may have a visible umbilical scar.

DISTRIBUTION (Figs 5 and 6). Widespread in the Sahara Desert and neighbouring
areas eastwards to southwest Iran and southeast Arabia. Northern border runs
south of Atlas Mountains, along the Mediterranean coast of Tunisia, Libya and
Egypt, through southern Israel, southern Jordan, central Iraq to southwest Iran
(Khuzistan and Gulf Coast). Southern border (so far as it is known, runs from
Senegal (Dakar area), through Mali (Timbouctou ; Adrar des Iforas - fide Angel &
Lhote, 1938), northwest Nigeria (Sokoto), Niger (Air ; Bilma-fide Angel & Lhote,
1938), southeast Libya (Kufra), central Egypt (Dakhla), north Sinai to southern
Arabia, but not extreme southwest, southernmost records being Democratic Republic
of Yemen (Hadhramaut) and eastern United Arab Emirates (Ra's al Khaymah etc).

REVISION OF SCINCUS 209

GEOGRAPHICAL VARIATION. Considerable geographical variation exists in the
5. scincus complex. East of the Red Sea there are two populations that differ quite
consistently in some aspects of colouring, scaling and proportion and these are
recognized here as subspecies : 5. s. meccensis (p. 219) and 5. s. conirostris (p. 221).
The situation in North Africa is less clear and is discussed below.

NORTH AFRICAN POPULATIONS OF THE Scincus scincus COMPLEX

Most of the characters known to vary geographically are displayed in Table I.
The position of the numbered localities can be found under MATERIAL EXAMINED
and in Fig. 5, which also shows the known distribution of pattern types. Numbers
of dark, transverse bands on the dorsum include the one situated between the fore-
limbs and all others as far as the vent region, but not bands on the neck.

Israel

Locality : 51.

Sample size : i.

Main features. This single specimen, which is a male, is generally very similar to

Egyptian material.

North and central Egypt

Applicable name. S. s. scincus (Linnaeus).

Synonyms. Lacerta (Edda) Donndorf, 1798 : 134. Type locality : 'in Abyssinia'.

Location of type : not traced.

Scincus officinalis var. lineolata Werner, 1914 : 343. Type locality : Cairo.

Location of type : Naturhistorisches Museum, Vienna (Registered number :

NMW 10390 : n).
Localities : 42 to 48, 50.
Sample size : 46.

Main features. Relatively small maximum body-size ; mature males often broad-
headed (head distinctly broader than neck). Typically 26 scales around mid-body,
or less ; frontonasal nearly always separated from first loreal. Dorsal colouring
consists of relatively numerous (usually eight or nine from forelimbs to vent), dark,
transverse bands that are about the same width as the interspaces, or a little broader
or narrower ; posterior neck band often separated from nuchal patch ; no sexual
dimorphism. Banding may develop at 76 mm snout to vent (largest animals without
bands are 78 mm). No dark bars on flanks.

Dakhla Oasis and 'Libyan Desert'

Locality : 49.

Sample size : 4.

Main features. Generally like other Egyptian material, but 28 or 30 scale-rows at

mid-body and transverse bands may be slightly narrower.

E. N. ARNOLD AND A. E. LEVITON

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UfM -4-*

M t^oo _ . M N rt ^

H ^ ^•'-'<^8S«5

•M c M^, is^;

Jf|M

•a ^ S

^ M ON O r^ 2 2
ON M « ON 03 C S

-*j j-> "-<

M OO 4)

fl *"* rt

M ON rC|

0) O o
O fl o

Tl

*O O t^ oo ON co

s

IH , — , CO rS

jj -. .

Dark transverse bai

11 la I 11 1
si fl ^s-g« ^

3n O^+Jfl wco

• • ^ .2 8 4 • o . &^

-gS^ u-S^E^^I

la* igii^a? i
ill IH^Si !

PH P c/3 (2

I'*!*

212 E. N. ARNOLD AND A. E. LEVITON

Northeastern Libya
Locality : 40.
Sample size : 6.

Main features. Similar to Egyptian material, but often 28 scale-rows at mid-body,
transverse bands tend to be narrower and fewer, seven or eight being present between
the forelimbs and the vent region.

Northwestern Libya, Tunisia and northeastern Algeria

Applicable name. 5. s. cucullatus Werner.

Scincus qfficinalis var. cucullata Werner, 1914 : 343. Type locality : Tripolis.

Location of type : Naturhistorisches Museum, Vienna (Registered number :

NMW 10386 : 3).
Localities : 21 to 29, 31 to 39.
Sample size : 72.

Main features. Moderate maximum body size, but some populations may be small.
Most commonly 28 scales around body (range 26-30) ; frontonasal usually separated
from first loreal. Dorsal colouring of most males and a few females consists of rela-
tively few (usually six or seven from forelimbs to vent region) transverse bands that
are much narrower than the interspaces ; posterior neck band is always fused with
nuchal patch. Many females and some males are without bands (of animals
70-90 mm snout to vent, three out of five males are banded and three out of twelve
females ; of animals over 90 mm snout to vent, 26 out of 26 males are banded and
three out of eleven females). Banding may develop at 77 mm snout to vent. No
dark bars present on flanks. See also p. 215.

Southeast Algeria (Djanet, Hoggar)

Locality : 30, 3oa.

Sample size : 32.

Main features. These animals are generally like north and central Egyptian

material, although contact between the frontonasal and first loreal scales is more

frequent and mature males do not appear to be very broad-headed. The single

example from Hoggar (3oa) has fewer, narrower transverse bars than the others and

is consequently more like northwest Algerian material.

Northwest Algeria

Applicable name. S. s. laterimaculatus Werner.

Scincus officinalis var. laterimaculata Werner, 1914 : 343. Type locality : Ain
Sefra, western Algeria. Location of type : Naturhistorisches Museum, Vienna
(Registered number : NMW 10377 : 4)-

Localities : 17 to 20.

Sample size : 28.

Main features. Large maximum body-size : even mature animals have relatively

small, narrow heads. Scales round mid-body usually 26 or 28 (rarely 30),

REVISION OF SCINCUS 213

frontonasal usually separated from anterior loreal. No sexual dimorphism in pat-
tern. Adults lack dorsal transverse banding and a dark nuchal patch, but head and
neck often bear a strong dark dorsolateral stripe on each side and the dorsum of the
tail may be speckled with dark pigment. Vertical bars or spots are present on the
flanks of adults.

A sample from Erg er Raoui (locality 18), west of Beni Abbes, consists of small
individuals (largest of four is only 95 mm snout to vent, although they may be
immature) with contact between the frontonasal and the anterior loreal and very
large flank spots.

Rio de Oro, Mauretania, Senegal, Mali (Timbouctou), Niger (Air) and southeast

Libya (Cufra)

Applicable name. S. s. albifasciatus Boulenger.

Scincus albifasciatus Boulenger, 1890 : 85. Type locality : Dakar. Location of
type : British Museum (Natural History), London (Registered number : BM
89.10.18.1/1946.8.13.60).

Boulenger states that, in the holotype, the scales covering the ear opening are
unfringed. This is not natural ; the lizard was apparently preserved some con-
siderable time after death and has lost many of its more delicate extremities,
including most of the digits.
Localities : i to 13, 41.
Sample size : 65.

Main features. Moderate maximum body-size. 24 to 26 scales around mid-body
(24 commonest in coastal areas of Mauretania and Senegal) ; frontonasal usually in
contact with first loreal. Dorsal colouring of adults consists of relatively few
(usually five to seven from forelimbs to vent), dark, transverse bands that are often
much wider than the interspaces, although sometimes about the same width ; pos-
terior neck band fused with nuchal patch ; no sexual dimorphism in colouring.
Faint banding may develop in animals as small as 62 mm snout to vent. Vertical
bars present on flanks of adults (and sometimes in individuals as small as 60 mm snout
to vent).

A sample from Chinguetti (locality 5) contains two animals patterned in the way
described above and others of similar size that are almost uniform above, with only
faint cross-bars or none at all. The two phases were collected on different dates, so
it is possible that they did not come from precisely the same collecting site. The
virtually unpatterned animals have at least a superficial resemblance to northwestern
Algerian individuals, especially those from Erg er Raoui. Pasteur & Bons (1960)
describe an apparently similar population from Taouz, southeast Morocco (locality 16),
that also contains both banded and unbanded individuals with dark bars on the flanks.

Northwestern Nigeria
Locality : 15.
Sample size : 5.

Main features. Apparently small (largest is only 95 mm snout to vent, but might be
immature). 26 or 28 scales round mid-body ; frontonasal separated from first

214 E. N. ARNOLD AND A. E. LEVITON

loreal scale. Dorsal pattern consists of relatively few (six to eight from forelimbs to
vent area), dark, transverse bands that are much narrower than interspaces ; pos-
terior neck band fused with nuchal patch ; no sexual dimorphism in colouring. No
dark bars or spots present on flanks.

Interpretation of the Scincus scincus complex in North Africa

At present, it is difficult to decide how to treat the North African specimens of
the S. scincus complex. There are four comparatively well-investigated peripheral
areas, in each of which the Scincus population has been named. These are : Egypt
and nearby areas (5. s. scincus) ; northwestern Libya, Tunisia and northeastern
Algeria (5. s. cucullatus] ; northwestern Algeria (S. s. laterimaculatus] ; Spanish
Sahara, Mauritania and Senegal (S. (s.) albifasciatus). In all these regions, the
Scincus are relatively uniform and show differences from those in the others, but
these areas cover only a fraction of the apparent range of the S. scincus complex in
North Africa. The more central parts of the Sahara and its southern borders
include many sandy regions apparently suitable for Scincus, yet only a few scattered
specimens are available from this vast and undercollected area. Thus, any decisions
about the systematic treatment of the complex must inevitably rest on an unsatis-
factory sample base. Certainly, a simple division into four subspecies is not tenable,
for the Scincus available from the poorly known central area do not necessarily
closely resemble those from the nearest well-investigated peripheral region. For
instance, southeast Algerian Scincus from Djanet (locality 30) are superficially more
like Egyptian animals than those from northern Algeria, and Nigerian specimens
look more similar to examples from northeastern Algeria, Tunisia and northwestern
Libya than to intervening populations.

The loose sand habitat, to which Scincus is more or less confined, is widespread in
North Africa, but it is by no means continuous and dune areas are often separated
by substrates unsuitable for Scincus thus producing an intricate system of at least
partial isolates. Mobile dunes travelling across unsuitable areas in the direction of
the prevailing strong winds could act as propagators of gene flow between some other-
wise isolated Scincus populations, but the pattern of connexion between sandy areas
and its extent have undoubtedly varied through time, so at least some geographical
units of the S. scincus complex have probably been subjected to a complicated
sequence of contact and separation. In such circumstances, it might be expected
that the degree of differentiation between populations would vary greatly, depending
on their length of separation and differences in the selective for cesacting in the
areas concerned. Such a situation would be difficult to reflect adequately by de-
lineating subspecies and there is the additional problem that some of the populations
may have differentiated enough to act as good species. In a closely knit population
complex, the best evidence for this is sympatry of distinguishable populations.
However, in the stringent desert conditions of the areas occupied by Scincus, the
number of ecological niches that a given environment can support is apparently
quite strictly limited. So, two species adapted to occupy more or less the same
ecological space are unlikely to be able to divide their common niche to produce two

REVISION OF SCINCUS 215

viable subniches. In such cases, abutment of ranges is therefore more usual than
extensive sympatry. Abutment occurs in other species groups in the Saharo-
Sindian desert region. For instance, in southeastern Arabia, Stenodactylus slevini
Haas, 1957 and S. leptocosymbotes Leviton & Anderson, 1967 fill very similar niches
and, although their ranges are in contact and not separated by any obvious barrier,
they seem to be virtually allopatric south of the Arabian Gulf (Arnold, 1977).
Similarly, Scincus mitranus and 5. scincus conirostris are much better differentiated
morphologically than any of the populations forming the S. scincus complex in
North Africa and are sometimes found at the same locality, yet their area of real
sympatry is probably small (see p. 243).

In a poorly sampled assemblage with a huge range, like the North African 5.
scincus complex, the chances of locating the very limited areas of sympatry or abut-
ment that might be expected are not great, but there is one situation that could be
interpreted as species contact. This is in the El Golea region of northern Algeria
(localities 20 to 23). At El Golea itself (locality 21) and at Ouargla (23), the Scincus
are generally similar to those of northeastern Algeria, Tunisia and northwestern
Libya (cucullatus) , except that they are rather small. In contrast, a sample stated
to come from northeast of El Golea (locality 20), collected by Hartert, is very like
northwestern Algerian animals. Differences between the two populations are listed
in Table 2. There is nothing in Hartert's account (1913) to suggest that his general
locality is erroneous. Unfortunately he does not state precisely where his material
came from, but it is unlikely to be more than 100 km from El Golea itself. If this
situation really indicates the abutment or overlap of specifically distinct populations,
the whole of the North African 5. scincus complex might be interpretable as two
closely related but largely allopatric species. Animals from El Golea, northeastern
Algeria, Tunisia and northwestern Libya (cucullatus} are generally similar to those of
Egypt and northeastern Libya (scincus s. str.), especially to the more western samples
which tend to have higher mid-body scale counts and fewer narrower dark dorsal
bands than the rest.

The southeastern Algerian sample, particularly the specimens from Djanet
(locality 30), is also quite like Egyptian material, so all the above populations may

TABLE 2

Differences between two populations of the Scincus scincus complex in northern Algeria

El Golea, Ouargla etc. Northeast of El Golea

Sample size 13 5

Size in mm (snout to vent

length of largest two <$<$) 102,96 134,124

Mid-body scale count 28 26

Bands on back frequent none

Dark spots or bars on

flanks no yes

Dark dorsolateral streaks

on sides of head and neck no yes

Tail more strongly speckled

than back no yes

2i6 E. N. ARNOLD AND A. E. LEVITON

form a single association. Similarly, skinks from northeast of El Golea, and north-
western Algeria (laterimaculatus) , are apparently connected to those of Spanish
Sahara, Mauretania and Senegal (albifasciatus) by intermediate populations in eastern
Mauretania (Chinguetti, locality 5) and in southeast Morocco (Taouz, locality 16, see
Pasteur & Bons, 1960). Specimens from Niger, Mali and southeastern Libya are
also essentially like the albifasciatus populations. Such a division would leave only
the Nigerian sample to be considered. This is much more like cucullatus type
animals from the north Sahara and may be closer to these than to the geographically
nearer albifasciatus populations. If the two groups really represent good species,
the correct names for them are 5. scincus for the mainly northern and eastern one
and S. albifasciatus for the other. Although juveniles do not seem to be easily
separable, adults could be keyed as follows.

a. No bars on flanks : dorsal bands, when present, not much wider and often much
narrower than interspaces, six to nine from forelimbs to vent region

Scincus scincus

b. Bars present on flanks : dorsal bands, when present, usually much wider than
interspaces, five to seven from forelimbs to vent region

Scincus albifasciatus

However, before such a division of North African Scincus can be accepted, con-
siderably more information will be necessary. The possible abutment near El
Golea needs confirming and other areas of contact or sympatry should be looked for.
For the present, it seems best to refer to all these animals as the North African S.
scincus complex and to recognize within this two groups of populations which can be
informally subdivided as follows :

scincus population group

a. Egypt and Israel (scincus s. str.).

b. Northeastern Algeria, Tunisia, northwestern Libya (cucullatus).

c. Southeastern Algeria.

d. Northwestern Nigeria.
albifasciatus population group

a. Rio de Oro to Senegal, eastwards through Mali and Niger to southeastern

Libya (albifasciatus s. str.).

b. Northwestern Algeria (laterimaculatus).

MATERIAL EXAMINED (n = 260). For southwestern Asian material, see S. s.
meccensis and S. s. conirostris (pp. 221 and 223). Abbreviations etc. are explained
on p. 191 and localities are shown in Fig. 5.

Rio DE ORO

1. Neighbourhood of El Aaiun ; EBD 2411, 6190-2, 6665 ; ?.$$, 3?°..

MAURITANIA

2. Adrar Choum (2i°2o'N I2°59'W) ; MHNP 67.556 ; <J.

3. Tasiat (= Tassiast, 2i°o2'N I5°I5/W) ; MHNP 1937.10; $.

4. Atar ; IFAN 55.7.147-9, 152, 155, 157-60, 163, 165, 168, 170-6, 181, 183, plus two
unregistered specimens ; g$$, 6$?, 7 juv.

REVISION OF SCINCUS

217

2i8 E. N. ARNOLD AND A. E. LEVITON

5. Chinguetti ; IFAN 55.7.150, 153-4, 156, 162, 166, 167, 169, 178-80, 182 ; 36*6*. 6$$, 4 juv.

6. Legleitat (= El Gleitat, i9°59'N I3°I7'W), Adrar ; MHNP 53.21 ; <?.

7. Nouakchott ; IFAN 55.7.177 ; 6*-

8. Trarza country ; BM 1913.5.9.41*, 1913.5.9.42-4 ; 26*6*, 2$$.

SENEGAL

9. Lompoul (i5°27'N i6°2i'W) ; IFAN unregistered ; 26*6*. $, 2 juv.

10. Thiaroye (i4°44'N i7°23/W) ; IFAN 55.18.315-17 ; 26*6*. ?•

11. Dakar; BM 89.10.18.1/1946.8.13.60 ; 6* (holotype of 5. albifasciatus) .

MALI

12. Timbouctou ; MHNP 17.86-8 ; 6*. 2??-

1 2a. Adrar des Iforas. No material examined, but occurrence reported by Angel & Lhote ( 1 938) .

NIGER

13. Oued Ifeinarem ( = ? Iferouane) ; MHNP 00.501 ; 6*-

14. Bilma. No specimens examined, but occurrence reported by Angel & Lhote (1938).

NIGERIA

15. Sokoto ; BM 1966.263* ; ad. AMNH 84300-3 ; 26*6*. 2??.

MOROCCO

1 6. Tafilalt, Taouz. No specimens examined, but occurrence reported by Pasteur & Bons
(1960).

ALGERIA

17. Ain Sefra, Prov. Oran ; BM 1913.7.3.78*, 1913.7.3.79-84, 1920.1.20.3711, MHNP

1974-313-17: "(JcJ. ?•

18. Erg er Raoui, Beni-Abbes ; MHNP 53.20, 53. A. 20, 53.6.20, 53.6.20 ; 26*6*. 2?$.

19. Grand Erg Occidental near Beni-Abbes ; BM 1961.1636, MHNP 46.305-6, FMNH 121765 ;
AMNH 91852 ; 6", 4??-

20. NE of El Golea ; BM 1912.11.9.103*, 1912.11.9.104-6, 1969.2162 ; 36*6*. 2°-$.

21. El Golea ; BM 1912.11.9.102 ; FMNH 62161-6; 36*6*. 3??, juv.

22. Wargla (= Ouargla) to El Golea ; BM 1912.11.9.101 ; $.

23. Ouargla; BM 1920.1.20.1191, 1920.1.20.1512 ; 36*6*. ?» Juv-

24. Bou Saada ; BM 1920.1.20.3037, MCZ 4615 ; 6*. Juv-

25. Biskra ; MHNP 20.48, 2O.A.48 ; 2 juv.

26. Toggourt ; BM 91.5.4.110-14, 94.7.25.5, 1969.2163-9 ; 46*6*. 8$$.

27. Sif el Menadi (33°58'N 6°22'E) ; BM 1920.1. 2O.3848a ; ?.

28. El Wed (= El Oued) ; BM 1909.7.28.15-18*, 1909.7.28.19-33^ 1909.7.28.34, 1920.1.
20.3848 ; 136*6*. 9??. 2 juv., ad. skeleton.

29. Dunes of Debala (= Debila, 20 km north-northeast of El Oued) ; BM 91.5.4.110 ; 6*-

30. Oued Imihrou, near Djanet (= Fort Charlet) ; MHNP 36.94, 36.95, 36.A.95, 36.96,
36. A. 96, 36.97, 36. A. 97, 36.98, 36. A. 98 ; 26*6*. 6?$, juv.

30. 10 km east of Djanet, near Efri ; CAS 138557-68.

3oa. Western border of Hoggar ; specimen lent by J. Bons ; 6*-
'Algiers' ; BM 69.6.22.1 ; $.
'Sahara' ; BM 59.11.30.3 ; °-.

TUNISIA

31. Nefta ; BM 1920.1.20.3046, MHNP 85.228-9 ; 6*. 2??.

31. 30 km west of Nefta ; CAS 138850, 138852, 138998-9005, 139506.

REVISION OF SCINCUS 219

32. 36 km southeast of Nefta ; CAS 132786-99, 135134-6.

32. Tozeur ; MHNP 85.227 ; $.

33. Sfax ; ZSM 26/1952 ; $.

34. Gabes ; MCZ 22353 '> 6*-

35. 50 km southwest of Douz ; FMNH 81745-6 ; $, $.

36. Duirat (= Douirat) ; BM 91.5.4.115-18 ; 2^, $, juv.

37. Djebel Sanrhar ; 40 km southwest of Bordj le Bceuf (32°n'N io°O4'E) ; FMNH 81747 ; <J.

LIBYA

38. Tripoli ; MCZ22354 ; <J. CAS 12751.

39. Hascian (= Qabilat al Hashshan, 32°46'N I2°55'E) ; BM 1955.1.8.70-3 ; 3^, <j>.

40. Gialo (= Jalo), Cyrenaica ; BM 1932.3.6.18*, MSNG 31578 ; 5??, juv.

41. Kufra, Cyrenaica ; MSNG 31577, MF 863 ; <J, ?.

EGYPT

42. Minyet Salamant (3O°23'N 3i°25'E) ; FMNH 79018-19, 79021, 79023, 79027, 79029 ;

43. Cairo ; BM 90.6.21.8-10 ; <$, $, juv.

43. Giza ; BM 97.10.28.440*, 97.10.28.441-9, 1936.9.1.17-18, FMNH 69268 ; 5^, 5^9, 3 juv.

44. El Mansuriya ; FMNH 69274 ; $.

45. Abu Roash, Abu Ruwash (= Abu Ruwaysh, 29°4o'N 3i°i4'E) ; BM 97.10.28.439, FMNH
66312 ; 2<J<J.

45. Zawyet (= Zawirat) Abu Musallem (29°56'N 3i°io'E) ; FMNH 69277 ; <J.

46. Wadi Muwellih (= Muweilih), Faiyum ; FMNH 164665-70 ; 3^$, 2$$, juv.

47. Manfalut ; FMNH 81184-5 ; <J, ?.

48. 3 km northwest of Durunka, Asyut ; FMNH 81179 ; $.

49. 35 km N of Mount Dakhla - sandy area ; FMNH 171834 ; $.

49. Dachel (= Dakhla) oasis ; ZSM 840/0.

50. El Arish, northeastern Sinai ; FMNH 58555 ; juv.
'Libyan Desert' ; ZSM 3/1974 ;

ISRAEL

51. Revivim ; BM 1961.690 ; $.

Scincus scincus meccensis Wiegmann

Scincus meccensis Wiegmann, 1837 : I27- Type locality : 'Arabia'. Location of type : Zoo-
logisches Museum der Humboldt Universitat zu Berlin (Registered number : ZMB 1178).

Scincus deserti Haas, 1957 : 76- Type locality : 23 miles [37 km] North of Hail, Saudi Arabia.
Location of type : Hebrew University of Jerusalem (Registered number : HUJ 2674).

DIAGNOSIS. A subspecies of 5. scincus in which the head is apparently not
markedly expanded in large animals and which has the frontonasal in contact with
both the rostral and the anterior loreal in nearly all cases. Adults have fine dappled
dorsal pattern of lighter and darker spots, often interspersed with light, sometimes
translucent scales that are yellow or orange in life and are frequently arranged in
short transverse bars. Dark vertical bars or spots are present on flanks of adults.

EXTERNAL FEATURES. Head fairly large ; in mature males not more than slightly
broader than neck and body at level of forelimbs ; length in animals of over 70 mm
varies from about 23 to 25% of snout to vent distance, width is about 55-63% of
head length. Top of head rises gently in lateral profile as in many 5. mitranus.

220 E. N. ARNOLD AND A. E. LEVITON

Rostral typically contacts frontonasal at least narrowly (two exceptions out of 17
animals checked). Supraoculars usually 5 : 5 or 6 : 6 (5 : 5 in 12, 5 : 6 in three, 6 : 6
in 13).

Nostril oval or slightly crescentic ; rostral usually excluded from its border (one
exception). Three loreal scales on each side ; first in contact with frontonasal.

Usually four or five nuchals on each side (although unilateral counts may vary
from three to five). Dorsal scales smooth in adults and juveniles. Mid-dorsals more
or less subequal to mid-ventrals. 24 to 28 scales around mid-body (see Table 3).

TABLE 3

Southwest Asian S. scincus: geographical variation in mid-body scale count and in number of
nuchals

Mid-body scale count Number of nuchals (higher uni-

lateral count)

24 25 26 27 28 29 30 I 2 3 4 5 6 7 8

Iran 25 5

Iraq 121 4

NE. Arabia 62 34

SE. Arabia 2 2

S. and SW. Arabia i 6 I 4 I i

S. Jordan, NW. and

W. Arabia 6714 7 10

All listed populations are 5. s. conirostris, except the last which is S. s. meccensis

COLOUR AND PATTERN. Dorsal colouring generally very similar to that of 5. s.
conirostris (p. 222), both in living and preserved animals. Differs mainly in presence
of vertical bars or blotches on flanks of non-juveniles. These vary in colour from
pale pink or orange-brown to deep purple-brown. They may develop in animals as
small as 62 mm. Iris blackish in life.

SIZE. A relatively small race in which, as in other Scincus, males are larger than
females. Largest males examined are 105 mm, 91 mm and 91 mm snout to vent ;
largest females 76 mm, 75 mm and 75 mm. Animals up to 51 mm long have
umbilical scars.

GEOGRAPHICAL VARIATION. Small sample size restricts comment, but there are
no obvious trends discernible, except that reduction of the supraoculars to 5 : 5 is
commoner in the south and mid-body scale counts are higher in Jordan (28 or 27
instead of 24 to 26).

SYNONYMIZED FORMS. 5. deserti Haas, 1957 falls clearly within the range of
variation of S. s. meccensis. Haas states that this form has four fringed scales cover-
ing the ear, but the paratype of deserti, and all other S. s. meccensis examined, have
the ear covered by two scales, each with two main lobes, as in most other members of
the 5. scincus complex.

DISTRIBUTION (Fig. 6). Known only from northwest Arabia and south Jordan.
It extends south on the Red Sea coast at least to the Jiddah area and eastwards at

REVISION OF SCINCUS 221

least to a point 37 km north of Hail (i.e. 27°35'N 4i°45'E), the type locality of 5.
deserti.

MATERIAL EXAMINED (n = 28). Abbreviations etc. are explained on p. 191 and
localities are shown in Fig. 6.

JORDAN

i. Guweira (= El Quweira) to Aqaba ; HUJ 1360 ; juv.

1. Wadi Rum ; MF, BM 1976.1715 ; <£, 3 juv.

2. Mudawara (= Al Mudawwarah), 'Central Arabia' ; HUJ 2673 ; juv.

SAUDI ARABIA

3. 27°2o'N 38°3o'E ; BM 1964.147 ; ?.

4. 26°5o'N 38°i5'E ; BM 1964.146 ; 6*-

5. 30 km NE of Yanbu al Bahr, 24°I3'N 38°I9'E ; CAS 119213.

6. Jiddah ; BM 1935.2.1.20, 1975.992 ; <J. $.

6a. Wadi Resifa or Wadi Fatma; BM 1976.1860 ; juv.

7. 19 miles [30-5 km] south of Jiddah, 2i°i2'N 39°i2'E ; BM 1964.302*, 1964.303 ; 2 juv.

8. Royal Saudi Naval Reservation, from coast to 6 km inland between 2i°o7'N and 2i°2i'N ;
CAS 136485 - 136494.

9. Burayman, 2i°39'N 39°i3'E ; BM, unregistered live specimen.

IMPRECISE LOCALITIES

Arabia ; ZMB 1178 ; $ (type of S. meccensis).

Arabia ; BM 1964.148* ; $.

'Central Arabia' ; HUJ 2752 ; Q* (paratype of S. deserti).

Scincus scincus conirostris Blanford

Scincus conirostris Blanford, 1881 : 677, fig. I. Type locality : Tangyak, 7 miles [n km] south of

Bushire (Iran). Location of types : British Museum (Natural History) (Registered numbers :

BM 79.8.15.1-3/1946.8.20.55-7).
Scincus gasperettii Haas, 1957 : 75, fig. 10. Type locality : Dhahran (Al Hasa district, Saudi

Arabia). Location of type: California Academy of Sciences (Registered number: CAS

84519).

DIAGNOSIS. A subspecies of S. scincus differing from other populations in the
following combination of features : snout often rather short and not as spatulate as
in other forms, head frequently quite broad in large animals, rostral and loreals
nearly always separated from frontonasal. Adults have fine, dappled dorsal pattern
of lighter and darker spots, often interspersed with light, sometimes translucent
scales that are yellow or orange in life and are frequently arranged in short irregular
transverse bars. No dark vertical bars or spots present on flanks.

EXTERNAL FEATURES. Head fairly big, and in large animals is often distinctly
broader than the neck and the body at the level of the forelimbs ; length in animals
over 80 mm may be about 20-26% of snout to vent distance, and width about
55-70% of head length. Snout is frequently rather short and this combined with
the often wide temporal region gives a characteristic profile in old animals (see
PI. 2a).

222

E. N. ARNOLD AND A. E. LEVITON

FIG. 6. S. SCWCMS : localities in southwest Asia. Figures refer to entries in lists of material
examined.
B - 5. 5. wecc£«sis.

D - Type locality of .S. deserti (= S. s. meccensis).
• - S. s. conirostris.

Rostral separated from frontonasal by the internasals (one exception out of 29
animals examined). Supraoculars typically 6 : 6 (5 : 6 in one case and 5 : 5 in seven) ;
supraciliaries usually 4 : 4 (but vary from 3 to 5).

Nostril oval ; three loreals on each side, the first nearly always separated from the
frontonasal (one exception out of 29). Ear covered by a single scale in one
instance.

Usually three to five nuchals on each side of neck but number can vary from
two to eight. Dorsal scales smooth in adults and juveniles ; mid-dorsals subequal
to mid-ventrals. 24 to 30 scales around mid-body.

COLOUR AND PATTERN. Dorsal ground colour (in spirit) varies from pale cream,
straw or grey to warm greyish-buff-brown. Young animals are finely dappled with
light and dark markings. Dappled pattern is retained by adults, but often becomes

REVISION OF SCINCUS 223

interspersed by paler, often at least partly translucent scales on the body that may
be scattered, but are usually arranged in irregular transverse rows ; sometimes these
go right across the back or they may be confined to dorsolateral areas. Dappled
dorsal pattern varies in intensity and is usually absent from the tail and head, and
often also from the neck and even from the forebody. There are no vertical bars
or spots on the sides.

In life (animals from Bahrain and United Arab Emirates), tail may be lighter than
back, the translucent scales are orange or deep yellow and the feet are pale yellow
this colour may also occur in irregular blotches on the flanks. Iris dark brown with
gold reflections.

SIZE. Largest males examined are 112 mm, 109 mm, 105 mm, 98 mm and 96 mm
snout to vent ; largest females are 98 mm, 82 mm, 77 mm and 69 mm. Two ani-
mals, respectively 37-5 mm and 46 mm snout to vent, have umbilical scars.

GEOGRAPHICAL VARIATION. The small sample available makes assessment of
geographical variation difficult. The largest animals come from the south (Hadh-
ramaut), but there is no clear evidence of size trends. Number of scales around the
body is highest in the north and may be lowest in the southeast. In contrast,
number of nuchals is lowest in Iran (see Table 3). Animals from around Dhahran,
Bahrain and the eastern United Arab Emirates have only five pairs of supraoculars
instead of six.

SYNONYMIZED FORMS. 5. gasperetti Haas, 1957 is based on animals with five
instead of six supraoculars on each side, otherwise it is indistinguishable from 5. s.
conirostris.

DISTRIBUTION. Known from south and east Arabia, Iraq and southwest Iran. In
southern Arabia, it occurs in the eastern United Arab Emirates, Hadhramaut
and Wadi Dawasir, but does not appear to traverse the highlands separating the
interior of the peninsular from the southwestern coast. In northern Arabia, 5. s.
conirostris extends westwards at least as far as 45°E. In Iraq, it is known from the
Baghdad area and in Iran it is confined to the provinces of Khuzistan and Fars.

REMARKS. 5. s. conirostris is known only from rather scattered localities. In
Arabia, it is far less common, and less often encountered than the partly sympatric 5.
mitranus.

MATERIAL EXAMINED (n = 29). Abbreviations etc. are explained on p. 191 and
localities are shown in Fig. 6.

IRAQ

1. Baghdad ; FMNH 20863, 20865.

2. Hindia (= Hindiyah) Dam ; SMF 55583-4 ; ad., juv.

IRAN

3. Agha Jari, Pars Province ; CAS 86479-80.

4. Bushire ; BM 87.9.22.40 ; ?.

5. Tangyak, 7 miles [11 km] south of Bushire ; BM 79.8.15.1-3/1946.8.20.55-7 ; 2<31<J, juv.
(types of 5. conirostris) .

6. Tangistan (= coastal area south of Bushire) ; BM 87.9.22.41 ; ?.

224 E. N. ARNOLD AND A. E. LEVITON

KUWAIT

7. Near Kuwait town ; BM 1972.1311* ; ad.

SAUDIA ARABIA

8. Jarrab (= Jirab), 26°45'N 45°oo'E ; BM 1964.145 ; 6*.

9. Dhahran ; CAS 84519 (holotype of S. gasperettii).

10. Abqaiq ; FMNH 73986 (paratype of 5. gasperettii).

11. Wadi Dawasir ; BM 1963.801, 1963.802* ; <£, $.

BAHRAIN

12. Ra's Jazayir, 25°58'N 5o°28'E ; BM 1971.116; $.

13. South of Amar ; BM 1975.989, unregistered live specimen ; 0*0*.

14. South Bahrain ; MCZ 52064 ; Q*.

UNITED ARAB EMIRATES

15. Near Jazirat al Hamra, 25°38'N 55°44'E, between Umm al Qaywayn and Ra's al Khaymah ;
BM 1972.1365 ; juv.

16. Ra's al Khaymah ; BM 1975.988* ; $.

DEMOCRATIC REPUBLIC OF YEMEN

17. Hadhramaut (collected on T. Bent's 1893-4 expedition to Hadhramaut, these specimens
may have been taken in Wadi Hadhramaut itself between Haynin and Shibam, or possibly
on the route from Mukkala, or on the way back to the coast at Shihr) ; BM 97.3.11.101-2 ;
6*6*.

18. Husn al Abr ; BM 1956.1.7.74 ; <£•

19. Ataq ; BM 1956.1.7.73* ; Q*.

IMPRECISE LOCALITY

'S. Hejaz' (these two specimens, which are without further locality, are part of a collection made
by H. StJ. Philby and stated to come from southern Hejaz. However, other material in the
collection is known to come from as far south as the western Democratic Republic of Yemen.
All precise localities are east of the coastal mountains of western Arabia) ; BM 1938.2.1.67-8 ;

6*6*.

Scincus mitranus J. Anderson

Scincus mitranus ]. Anderson, 1871:115. Type locality: 'Arabia'. Location of type:

Zoological Survey of India, Calcutta (Registered number : unavailable).
Scincus arenarius Murray 1884 : 353. Type locality : Hubb (= Hab) Plains, Sind, Pakistan.

Location of types : two of the seven types are in the British Museum (Natural History)

(Registered numbers : BM 87.9.22.42/1946.8.20.59 ; BM 86.6.30.52/1946.8.20.58) ; the

others may be in Karachi, but it has not been possible to confirm this.
Scincus muscatensis Murray 1886 : 67. Type locality : 'Muscat*. Location of types : three of

the 13 types are in the British Museum (Natural History) (Registered numbers : BM 85.7.11.

16-18/1946.8.20.68-70) ; the others may be in Karachi.
Scincus arabicus Schmidt, 1939: Type locality: 'near Shabwa, Hadhramaut' (= Shabwah,

Democratic Republic of Yemen). Location of type : Field Museum of Natural History,

Chicago (Registered number: FMNH 18460).
Scincus philbyi Schmidt 1941:162. Type locality: 'Anaiza' (= Unayzah), Saudi Arabia.

Location of type : Field Museum of Natural History, Chicago (Registered number : FMNH

31664).

REVISION OF SCINCUS 225

Scincus richmondi Haas, 1961 : 24. Type locality : Southeastern Jafura Desert, Al Hasa
District, Saudi Arabia. Location of type : Carnegie Museum, Pittsburgh (Registered
number CM 33535).

DIAGNOSIS. Head relatively small ; external ear orifice very small but visible,
usually situated well below level of line made by lower edges of upper labial scales
and typically covered by two scales, the hind edges of which are usually serrated over
the orifice (rarely covered by a single, less serrated scale). Eye with relatively small
cornea and a rounded pupil. Rostral scale broadly borders frontonasal ; fronto-
parietals in usually extensive contact (rarely fused or separated) ; parietals rela-
tively small (usually less than twice the length of adjoining nuchals). Dorsal scales
smooth, mid-dorsals subequal to mid-ventrals (rarely more than slightly larger or
smaller). (25) 26 to 30 (31) scales around mid-body. Adults and young without
bold dorsal pattern, either almost uniform or with fine dappled pattern of lighter
and darker spots ; dark bars or spots present on flanks of nearly all adults.

Auditory meatus long with some fairly well-developed scales distally and a long
cartilage in its lateral wall. Extrastapes an elongate plate. Premaxillary rostrum
long. Otic capsule normal ; squamosal quite sharply flexed, its upper border often
forming a small spur. Distal section of supratemporal process of parietal not con-
stricted, terminating well above level of ventral tip of squamosal.

EXTERNAL FEATURES. Head small, a little narrower or at most slightly broader
than neck and body at level of forelimbs ; length in animals over 80 mm may be
about 19-24% of snout to vent distance ; width about 55-65% of head length.
Some variation exists in snout proportions. Sides of snout usually at least slightly
concave when viewed from above. Top of head rises relatively gently from snout-tip.
Canthus rostralis quite well marked ; loreal region distinctly concave. Tail
broader than deep proximally, clearly laterally compressed towards tip ; up to 80%
of snout to vent distance when undamaged.

Rostral overhangs mental strongly, more so than in sympatric 5. scincus ; pre-
f rentals in contact , or fused (in north of range) . Frontal often much wider in front than
behind, often more so than in the 5. scincus complex. Frontoparietals in usually
extensive contact (rarely fused). Parietals relatively small, in most cases once
to twice the length of adjoining nuchals. Supraoculars nearly always 6 : 6
(very rarely 5 : 6 and said to be 5 : 5 in type) ; supraciliaries usually four on each
side (but up to seven may be present).

Nostril oval or quite strongly crescentic, often in narrow contact with rostral shield.
Often three loreals on each side (but first absent in north of range) ; if present, first
loreal usually separated from frontonasal (but not in animals from Masirah Island
and from 'Muscat'). Upper labials fairly deep (third is about 1-5 to 2-8 times depth
of second loreal) ; typically eight on each side, but number varies from seven to
nine. Ear orifice very small but visible, situated usually well below level of line
made by edges of upper labials ; typically covered by the contiguous parts of two
scales, the posterior edges of which are nearly always at least slightly serrated over
actual orifice (in nine out of 101 animals checked for this feature, scales covering ear
coalesce and may appear as a single plate, often with reduced serration).

226 E. N. ARNOLD AND A. E. LEVITON

Typically three to five pairs of nuchals, but sometimes less and nuchals may even
be absent. Dorsal scales smooth in adults and juveniles ; mid-dorsals subequal to
mid-ventrals (rarely more than slightly wider or narrower). (25) 26 to 30 (31) scales
around mid-body. Row of laterally expanded scales on dorsal midline of tail
usually quite developed, typically extending proximally almost to the level of
extremities of posteriorly adpressed hindlimbs, or a little beyond.

Parts of scales forming posterior digital fringes often relatively broad and less
acutely pointed than in 5. hemprichii, and the outer border of each is more curved.
Claws and ungual lamellae often strongly expanded laterally.

COLOUR AND PATTERN. Dorsal ground colour varies from pale buff or grey to
warm brown or deep cinnamon. This colour may change somewhat in spirit, but
in life, in combination with other pattern elements, generally matches the sand on
which the individual is found, although there is some variation within populations.
This colouring does not change, persisting for at least four years in captivity, even
when animals are kept on contrasting backgrounds. Young animals may be almost
uniform above or finely dappled with dark or light spots, or more usually with both.
Adults possess a similar dappled dorsal pattern. Usually, this consists of one (or
two) dark spots, sometimes confined to the posterior edge of the scale and bordered
and separated by light, often whitish pigment. Both these elements may eventually
extend forwards to cover a greater part of the scale. There is considerable variation
in intensity and extent of the markings, even within populations ; in animals from
'Muscat' and Masirah Island, the light spots are often large and form broken longi-
tudinal lines along the back. The head usually lacks dappled patterning. The distal
part of the dorsal surface of the tail is often reddish or light bluish, especially in
young animals. The flanks and limbs are paler than the back and there may be
some grey pigment, especially on the sides of the head and neck.

In life, most animals (at least non- juveniles) have some deep yellow or orange pig-
ment, although this rapidly disappears in alcohol. It is found on the feet where it is
intense, on the upper surfaces of the limbs and as a series of short transverse bars
situated dorsolaterally on the body and tail-base. At least some of those on the
body extend upwards from the dark bars or spots on the flanks. There may also
be yellow blotches between the dark bars or spots.

Adults usually have a row of spots or short vertical bars on the flanks that vary in
colour from pale pink through red to a very deep warm brown. They begin in front
of the forelimbs and often extend to the groin and sometimes onto the tail-base as
well. Often these flank bars decrease in size and intensity posteriorly. The number
and spacing of bars or spots present and their size, shape and colour vary and there
are often both individual and local differences. There is usually one or more rarely
two bars in front of the forelimbs and from four to ten between the two pairs of legs.
These markings are absent in hatchlings but may develop in animals as small as
57 mm snout to vent and, by the time they reach 70-80 mm, the majority have them.
Above this size they are only occasionally absent (largest animal encountered without
bars is a male, 105 mm snout to vent).

Underside whitish or cream. Iris very dark brown.

REVISION OF SCINCUS 227

EYE. Cornea relatively small, about 33-38% (n = 4) of the horizontal diameter
of the eye. Pupil roundish, not a vertical slit when contracted.

EAR (see Fig. 2). Meatus long, its distal portion lined with scales, but these are
less well developed and the meatus is less thick- walled than in the S. scincus complex.
The cartilage in the lateral wall of the upper section of the meatus is long and pointed.
Tympanum somewhat smaller (compared with skull length) than in the 5. scincus
complex. Distal section of the extrastapes forms an anteroposteriorly expanded
cartilaginous plate that occupies a considerable area of the tympanum. Stapes
slightly more robust than in the S. scincus complex, with a larger foot-plate.

SKULL (see Fig. 2). Premaxillary rostrum strongly developed. Squamosal quite
strongly flexed and there may be a small upward spur bordering the supratemporal
process of the parietal. Surface exposure of the latter is not constricted ventrally,
but it terminates well above the level of the ventral tip of the squamosal. Supra-
temporal bone larger than in the 5. scincus complex, smaller than in 5. hemprichii.
Otic capsule not, or not much, bigger than in the 5. scincus complex and much smaller
than in S. hemprichii ; in lateral view, its posterior border is often clearly visible
behind the supratemporal bone and the supratemporal process of the parietal.
Maxillary teeth about 15-20 in adults. Mandibular teeth about 13-18.

SIZE. There are some indications of local variation in size, but these do not seem
to follow any consistent pattern. Adult males are considerably larger than adult
females and the majority are over 100 mm from snout to vent, whereas most females
are less than this. Largest five males examined are 134 mm, 126 mm, 123 mm, and
121 mm and the largest five females 105 mm, 104 mm, 104 mm, 94 mm and 92 mm.
The smallest juvenile encountered is 38 mm, and umbilical scars are visible, although
fading, on animals up to 50 mm.

GEOGRAPHICAL VARIATION. Snouts tend to be rather short and broad in the north
and northwest of the range and are longest and narrowest on Masirah Island. Some
other features of S. mitranus that vary geographically are displayed in Table 4.
From this it will be apparent that mid-body scale counts are lowest in the southeast
but often show considerable local variation. Northern animals almost always have
the pref rentals fused and two loreals on each side. In the Al Hasa region (localities
5 to 9, 13, 14) various combinations of prefrontal arrangement and loreal number
occur. Animals from the western United Arab Emirates (localities 23 to 29) have
unfused prefrontals ; most have three loreals on each side, but some (9 out of 45)
have only two. Elsewhere, three loreals are almost always found, but fused pre-
frontals may occur as a minority condition.

Apart from their low mid-body scale counts, animals from 'Muscat' and Masirah
are peculiar in consistently having the first loreal in contact with the frontonasal.
Also, the light spots in the dappled dorsal pattern are often large and tend to form
broken longitudinal stripes.

The name 5. philbyi Schmidt 1941 is available for the northern populations with
rather broad snouts, fused prefrontals and two loreals on each side, but it seems
unwise, without a more searching analysis, to recognize a subspecies largely on two
minor scale characters, especially when there is a large area where one or both of

228 E. N. ARNOLD AND A. E. LEVITON

TABLE 4

Geographical variation in Scincus mitranus

First loreal,

if present, Loreal scales
Region Mid-body scale count contacts on each side Prefrontals

25 26 27 28 29 30 31 frontonasal 3 2 separate fused

Northern Arabia
(localities i to 4, 10,

ii) i 10 8 3 19 — 19

Dhahran etc.

(localities 5 to 9) 3 3 38 2 7 18 32 n 40

Jafurah etc.

(localities 13, 14) 2 11 i 11 3 14 —

Qatar and western
United Arab Emir-
ates
(localities 23 to 29) i 10 4 32 36 9 46 —

Eastern United Arab
Emirates
(localities 30 to 34) 7 4 25 i 39 — 38 2

Central Arabia

(localities 12,

15 to 17) 3 i 9 13 — 11 2

Southwest Arabia

(localities 18 to 21,

39, 41, 42) 9 i 8 17 i 15 3

Southern Rub-al- Khali
(localities 22, 37,
38) 12 321

'Muscat* and Masirah

(localities 35, 36) i 7 i +10 — 10 —

these exist alongside more typical arrangements. The 'Muscat' and Masirah popu-
lations, for which the name S. muscatensis Murray, 1886 is available, may be largely
separated from the rest of the range of S. mitranus by unsuitable, non-sandy country,
so they could well be subspecifically distinct. However, only ten specimens are
presently available to us and most of these come from the probably imprecise
locality of 'Muscat' (see p. 231). Until southeast Arabia is better collected, it seems
safer to reserve judgement about the status of these animals.

SYNONYMIZED FORMS. S. philbyi and S. muscatensis have already been discussed
above and S. arenarius is dealt with under DISTRIBUTION. S. arabicus Schmidt,
1939 is based on two juvenile specimens, 48 mm and 46 mm snout to vent. The
former is the holotype and comes from Shabwa (= Shabwah), and the paratype is
from Wadi Beihan, both localities in the the Democratic Republic of Yemen. Both
agree very closely with S. mitranus. The absence of dark bars on the flanks men-
tioned by Schmidt is to be expected in animals of this size, and the digital fringes,

REVISION OF SCINCUS 229

said to be poorly developed, are well within the range of variation for this species.
The author also states that the parietals are broken into small transverse scales,
but apart from the left frontoparietal of the holotype being divided in two, the
head scaling is quite normal for S. mitranus.

S. richmondi Haas, 1961 also falls well within the range of variation encountered
in S. mitranus.

DISTRIBUTION. (Fig. 7). Restricted to south and east Arabia. In the south, it
occurs from southeastern Sultanate of Oman to the lowlands of southwestern Saudi
Arabia, east Yemen and the western Democratic Republic of Yemen, but it does not
appear to cross the high mountains that separate the interior of Arabia from the
southwestern coast of the peninsula, nor is it known from the southern coast west
of Masirah Island, except for seven specimens reported from Gischin ( = Qishn) by
Steindachner (1903). In the north, it reaches Kuwait at about latitude 29°. Its
northwestern limits are uncertain, but they extend at least as far as Unayzah
(26°o6'N 43°58'E) and Jabal al Alam (23°36'N 43°3i'E).

Pakistan records. S. mitranus was recorded from Pakistan by Murray (1884)
under the name S. arenarius. He described this taxon on the basis of seven in-
dividuals said to come from Hubb ( = Hab) Plains, Sind, but there is considerable
circumstantial evidence suggesting that this locality is erroneous. Firstly, no
Scincus appear to have been collected in Pakistan since Murray's description and,
as Minton (1966) says, 'it is somewhat surprising that a rather large and distinctive
reptile should not be rediscovered in the relatively well-collected lower valley of the
Hab River'. Secondly, Murray's specimens seem to be well within the range of
variation of the Arabian 5. mitranus. Two of the types of S. arenarius are in the
British Museum, and these differ slightly in their head scaling : one has the pre-
frontals fused and three labials on each side, and the other, prefrontals separate and
two labials. As has been pointed out (p. 227), these two combinations of loreal and
prefrontal arrangement are relatively common only in a small area of the total range
of S. mitranus, namely the Al Hasa region of eastern Arabia, which is more or less
opposite Bahrain Island. Murray's material also agrees with the Al Hasa skinks
in other respects. It would be curious if an isolated population of Scincus, separated
(over land) by a distance of almost 2000 km from its nearest relative, should resemble
the Al Hasa population so closely. Thirdly, there are good reasons for believing that
specimens of an Arabian Scincus could turn up in Pakistan for lizards of this genus
have a long history of medical use and were frequently items of trade. Scincus was
long valued as an antidote to poisons in Europe, while in the East it had more
hedonistic uses. Anderson (1898) states that 'among Arabs it was in high repute
as a remedy for reanimating the powers exhausted by age or debauchery, and in
eastern countries fables are still extant regarding its potency in this respect'. This
author (1871) confirms the use of Scincus in the Indian subcontinent, particularly as
an aphrodisiac amongst the Muslim population, and it is perhaps significant that the
type locality of 5. arenarius is in a Muslim area and near a large sea-port (Karachi,
where Murray worked) . It seems very likely that eastern Arabia was the main source
of Scincus for the Pakistan pharmaceutical market, for it is the closest area (by sea)

230 E. N. ARNOLD AND A. E. LEVITON

that is certainly within the range of the genus and there is a long history of trade
between the Irano-Arabian Gulf and Pakistan and western India. Trade in Scincus
was certainly still going on in Murray's time. The type of S. mitranus was purchased
in about 1871 as a dried specimen in India or Pakistan from a merchant who said
that he had brought it from Arabia (Anderson, 1871). Indeed, over 30 years later
in 1904 the British Museum received a 5. mitranus (BM 1904.3.25.1) from a London
trading company which had obtained it through a Bombay merchant who, in turn,
had imported it from Arabia. He told the trading company that the Scincus had
medicinal properties. Remarks of Cheesman (1926) suggest that the east Arabian
Scincus trade continued for some time after this.

Another line of evidence concerns the way in which Murray's specimens had been
treated after capture. From Anderson (1898) and examination of BM 1904.3.25.1
it is apparent that Arabian skinks prepared for commerce were roughly cut along the
belly and all or part of the guts removed ; the carcase was then dried, presumably
in the sun, when it sometimes became fly-blown. The two Murray specimens in the
British Museum show all of these features.

It seems, therefore, that Murray was probably misinformed about the locality of
his S. arenarius and that these animals really came from Arabia.

SEX RATIO. Of the animals over 70 mm that were sexed, 69 are males and 33
females. Of 22 adults collected near Shajah, 16 are males. This disparity may be
more apparent than real, possibly males are less retiring than females, at least at
certain times of the year.

MATERIAL EXAMINED (n = 244). Abbreviations etc. are explained on p. 191 and
most localities are shown in Fig. 7.

KUWAIT

1. Kuwait; BM 1976.1859 ; Q*. Unregistered live specimen.

SAUDI ARABIA

2. Anaiza (= Unayzah) ; MCZ 53930 ; <J (paratype of S. philbyi). FMNH 31664 (holotype of
S. philbyi) ; FMNH 28599 (n = 6, paratypes of 5. philbyi).

3. Zilfi ; BM 1963.803-4 ; ?.

4. Shaib Atj (= Wadi al Atk, 26°oo'N 46°3o'E) ; BM I935.io.8.i6*-2i ; 4<JCJ, ?. juv.

5. Jubail (= Al Jubayl) ; BM 1 953-1 -8-53 ; ?•

5. Ras Tamura (= Ra's Tannurah) ; AMNH 66602.

6. Qatif ; CAS 84414.
6. El Alat ; CAS 84435.

6. Dhahran ; FMNH 74003 ; $. CAS 84535, 84580-7.
6. Shimal ; CAS 84427, 84429.

6. Tomayiah ; CAS 84626.

7. Abqaiq ; FMNH 74004, HUJ 7087, MCZ 80925 ; 6*. 2??. CAS 84296, 84303, 84305-6,
84308-10, 84313, 84315-17, 84320, 84324-9, 84487-9, 84494.

8. Jurdah (= Judah), NW of Hufuf ; BM 1947. 3. 2. 23* -29 ; 6<J6\ 3??-

9. Zanuza, Hufuf ; BM 1924.11.18.16 ; <J.
9. Near Hufuf; BM 1953.1.8.54 ; 6*.

10. Dahna, 100 km east of Riyadh, 25°o8'N 47044'E ; CAS 119202-3.

10. Dahna, 25°oo'N 47°3o'E ; CAS 134138-40.

11. Riyadh; SMF 55014 ; ad.

REVISION OF SCINCUS 231

12. Northwestern fringes of Jabal al Alam, 23°36'N 43°3i'E ; CAS 113717.

13. Wadi Sahaba (= Wadi as Sahba) ; BM 1924.11.18.17 ; ?.

14. Jafurah ; BM 1932.10. i.83*-gi, 1924.11.18.15 ; 8^,2$$.

14. Southeastern Jafurah desert ; CM 33513, 33517, HUJ 7920 (originally CM 33516) ; 2^,
$. CAS 93034 (originally CM 33518) (all are paratypes of S. richmondi).

15. Irq Minttar, in Uruq Rumaylah, 2o°i5'N 46°45'E ; CAS 119216-19.

16. Near Irq Abu Faqr, 75 km south, 69 km west of Rayda well, 2O°i5'N 47°i5'E ; CAS
119249-51.

17. Irq Kabdan, 2i°oo'N 48°4o'E ; CAS 119244-8.

18. Bir Husayniyah, edge of Wasi Habawnah, I7°49'N 44°27'E ; CAS 119207-9.

19. Khobash, I7°33'N 44°49'E ; CAS 119210, 119241.

20. i7°i6'N 46°i6'E ; CAS 119235-8.

21. Near Ash Sharawrah, I7°29'N 47°O7'E ; CAS 119239-40.

22. Shanna, 1000 ft [300 m], i8°55'N 5i°o5'E ; BM 1931.7.16.55* ; 6*-

23. 22°io'N 54°2i'E ; BM 1971.1348-9 ; <j>, juv.

QATAR

24. Qatar; BM 1950.1.5.12-13 ;

UNITED ARAB EMIRATES

25. Al Hamra (24°oi'N 52°25'E) ; BM 1971.758-9, 1972.840 ; <J, 2 juv.

26. Between Al Hamra and Bada Haza (23°4o'N 52°36'E) ; BM 1972.841-2 ; 2 juv.

27. 23°45'N 53°33'E ; MCZ 85562 ; <J, CAS 97768-75, 97777-9, 97786-90, 97792-96, 98091.
27. Habshan, 23°5o'N 53°37'E ; CAS 97780-5.

27. Near 23°4i'N 53°28'E ; CAS 97776.

28. Near Bada Zaid, 23°35'N 53°5o'E ; BM 1972.1260-2 ; 6*, 2$?.

29. North edge of Liwa ; BM 1971.1747-8 ; $, $.

29. Al Liwa area ; BM 1971.542-5 ; 2<J<J, 2 juv.

30. Between Abu Dhabi and Al Aga ; BM 1971.1654 ; juv.

31. Bereimi (= Buraimi) ; BM 1950.1.5.11 ; $.

33. Sharjah ; BM 1970.1594*, 1971.1023-4, 1975.1305, 1972.1366-7, 1973-734-5, 1973-
2088-103, J975-985-6, 1975.990, HUJ 2671 ; i6<Jo\ 6$$, 5 juv.

34. South of Jazirat al Hamra, 25° 38'N 54°44'E ; BM 1973.2087, 1975.980-1 ; <$, 2$$.

34. Ra's al Khaymah ; BM 1975.982-3, 1975.987 ; 3^, juv.

32. Jebel Fayah (= Jebel al Fajah), 25°o6'N 55°5o'E ; BM 1971.1304, 1975.991 ; i$$t ?•

SULTANATE OF OMAN

35. 'Muscat'. It is unlikely that the locality attached to these individuals indicates the town
of Muscat. This is situated in a largely rocky area not obviously suitable for Scincus and
none appears to have been taken there since the listed specimens were obtained in the i88os.
'Muscat' has sometimes been used in a more general way for the eastern section of the
Sultanate of Oman. Certainly, some of the other reptile material of A. S. G. Jayakar, who
obtained a proportion of the S. mitranus listed here, may not have come from the immediate
environs of Muscat town itself. Mrs F. E. Warr (personal communication) points out that
Jayakar collected fossils in the sandy Ash Sharqiyah region southwest of Muscat and some
of his bird material labelled 'Muscat' is likely to have come from here. So this may also
be the origin of some of his reptiles.) BM 85.7.11.16-18/1946.8.20.68-70 (types of 5.
muscatensis) , 85.11.7.9-10, MCZ 112246, AMNH 17821 : 46*6*. ?• 2 juv-

36. Masirah ; BM 1971.1025, 1973.405, 1975.984 ; 2^, $.

37. Ramlat Shu'ait, 1000 ft [300 m], i8°4o'N 52°io'E ; BM 1931.7.16.54 ; ?.

38. Kharaiyim Fasad, 850 ft [255m], i8°3o'N 53°io'E ; BM 1931.7.16.56; juv.

232

E. N. ARNOLD AND A. E. LEVITON

FIG. 7. S. mitranus and 5. hemprichii: localities,
material examined.

• - S. mitranus.

• - S. hemprichii.

Figures refer to entries in lists of

DEMOCRATIC REPUBLIC OF YEMEN

39. Zamakh, north Jol, Hadhramaut ; BM 1953.1. 7. i8.i9*-23 ; <J, 2$$, 3 juv.

41. Shabwa (= Shabwah), Hadhramaut ; FMNH 18460 ; juv. (type of S. arabicus).

42. Wadi Beihan ; FMNH 18459 ; juv. (paratype of S. arabicus).

IMPRECISE OR ERRONEOUS LOCALITIES

Oman ; BM 1950.1.4.75-77 ; <$, 2$$.

Rub-al-Khali desert (collected by H. StJ. Philby on his traverse of the desert ; see Philby

(1933) for his route) ; BM 1932.10.1.92*. 93*-9 ; 46*6*. 2??. Juv-

S. Hejaz (see comment on this locality under S. s. conirostris, p. 224) ; BM i938.2.i.65*-6 ;

26*6*.

Arabia ; BM 1904.3.25.1 ; ad.

Hubb ( = Hab) Plains, Sind (see comment on p. 229) ; BM 86.6.30.52/1946.8.20.58,

87.9.22.42/1946.8.20.59 ; 2 ad. (two of the types of S. arenarius).

REVISION OF SCINCUS 233

Scincus hemprichii Wiegmann

Scincus hemprichii Wiegmann, 1837 : 128. Type locality : 'Abyssinia' (aus Massaua in Abys-
sinien' [= Massawa, Ethiopia] - fide Peters, 1865). Location of type : Zoologisches Museum
der Humboldt Universitat zu Berlin (Registered number : ZMB 1179).

Scincus (Pedorychus) hemprichii Peters, 1865 : 44.

DIAGNOSIS. Head large and broad ; external ear orifice minute, placed well below
the level of the line made by the lower edges of the upper labial scales, and completely
hidden by a large, smooth-edged scale ; eye with large cornea and a pupil that is
vertically slit-shaped when contracted. Rostral scale separated from frontonasal by
internasals ; frontoparietals in limited contact or separated ; parietals large (usually
about twice the length of adjoining nuchals, or more). Dorsal scales ridged in
adults ; mid-dorsals often distinctly larger than mid-ventrals ; 22-24 scales around
mid-body. Dorsal pattern of adults and young consists of narrow, pale transverse
bands on brownish ground ; no dark bars or spots present on flanks.

Auditory meatus quite long with at most a few scale vestiges distally and, proxi-
mally, a short triangular cartilage in its lateral wall. Extrastapes large and ovoid.
Premaxillary rostrum short. Otic capsule huge, squamosal sharply flexed, its
upper border forming a distinct spur posteriorly. Distal section of supratemporal
process of parietal narrowed, terminating well above level of ventral tip of squamosal.

EXTERNAL FEATURES. Head large, distinctly broader than neck and much
broader than body at level of forelimbs ; length in mature animals may be about
24-30% of snout to vent distance ; width about 55-70% of head length. Snout
often broad in adults, its sides at least slightly convex when viewed from above.
Top of head rises relatively steeply from snout-tip, more so than in S. mitranus.
Canthus rostralis very weak or absent ; loreal region only slightly concave. Tail
approximately cylindrical in cross-section, slightly compressed laterally towards
tip ; about 50-60% of snout to vent distance in adults, up to 70% in juveniles.

Rostral overhangs mental less than in most other Scincus, and is separated from
frontonasal by internasals. Prefrontals in contact, unfused. Frontal rather wider
in front than behind. Frontoparietals either meeting narrowly or separated (in
one case, fused to frontal). Interparietal kite-shaped, often rather elongate,
occasionally separating first pair of nuchals. Parietals relatively large, about
i -7 to 3 times as long as first pair of nuchals. Supraoculars usually 6 : 6 (sometimes
reduced to 5, at least unilaterally) ; supraciliaries typically four on each side (but
three to seven may be present).

Nostril round, oval or slightly crescentic ; rostral excluded from its border.
Usually three loreals on each side (but only two in one individual) ; if present, first
loreal is separated from frontonasal. Upper labials deep (third is 2-3 to 3 times
depth of second loreal) ; typically eight on each side but number varies from seven
to nine. Ear orifice minute and situated well below level of line made by lower
edges of upper labials ; completely hidden by smooth-edged scale that is larger than
neighbouring ones.

Typically four or five pairs of nuchals (although counts may vary from two to five
on each side). Dorsal scales more or less smooth in juveniles but ridged in older

234 E- N. ARNOLD AND A. E. LEVITON

animals (two or three ridges on each scale, the ridging being strongest towards the
posterior edge) ; mid-dorsals distinctly wider than mid-ventrals. 22 to 24 scales
around mid-body. Row of laterally expanded scales on dorsal midline of tail well
developed, extending proximally beyond level of extremities of posteriorly adpressed
hindlimbs.

Parts of scales forming posterior digital fringes relatively narrow and pointed.
Claws and ungual lamellae not strongly expanded laterally.

COLOUR AND PATTERN. In young animals, most dorsal scales are some
shade of brown with a darker edge. These are interspersed by irregular transverse
bands of paler scales, usually bearing opaque, whitish pigment, especially towards
their posterior borders. This basic coloration is retained by many adults, but the
dark scale edges tend to disappear and the whole pattern may sometimes fade.
Nuchal area tends to be darker than the rest of back, and ground colour and intensity
of pattern vary as does the number of light transverse bands. There may be one on
the neck, from six to ten on the body and at least indications of more on the proxi-
mal part of the tail. Northern populations tend to have fewer bands on the body
than southern ones (six or nine as against eight to ten). Dorsal pattern fades on
flanks, which lack dark vertical bars or spots. Underside whitish or cream. Iris
dark brown.

In life, there is often some pale yellow on the flanks, legs and belly. Snout and
extremities of limbs are usually pinkish.

EYE. Cornea large, about 47-58% of horizontal diameter of eye (n = 3).
Contracted pupil a vertical slit.

EAR (Fig. 3). Meatus quite long with only a few vestiges of scales in its distal
section, which is membranous ; cartilage in lateral wall of upper portion is short
and bluntly triangular. Tympanum relatively small (compared with skull length) ;
distal section on extrastapes large and ovoid, occupying much of the tympanic area.
Stapes robust, with a large foot-plate.

SKULL (Fig. 3). Premaxillary rostrum present, but short. Squamosal strongly
flexed and there is a well-developed upward spur bordering the supratemporal
process of the parietal. The surface exposure of the latter is abruptly reduced at
about this point and it continues downwards as a narrow rod to terminate well above
the level of the ventral tip of the squamosal. Supratemporal bone relatively large.
Otic capsule huge, extending forwards almost to epipterygoid bone and backwards
well beyond the supratemporal process of the parietal. About 20 teeth in each
maxilla and mandible.

SIZE. Adult males considerably larger than adult females. Five largest males
examined, 138 mm, 138 mm, 131 mm, 130-5 mm and 126 mm snout to vent ; the
three largest females 108 mm, 95 mm and 95 mm. Smallest juvenile examined is
44 mm.

GEOGRAPHICAL VARIATION. Apparently some in markings ; see above.

DISTRIBUTION (Fig. 7). Known with certainty only from the extreme southwest
of the Arabian Peninsula. All localities are in the coastal lowlands, from Sabya

REVISION OF SCINCUS 235

(Asir Province, Saudi Arabia) through western Yemen to the Aden region of the
Democratic Republic of Yemen.

There is also a slight possibility that 5. hemprichii occurs on the western coast of
the Red Sea. The type was obtained by Ehrenburg and Hemprich during their
travels in northeast Africa and Arabia in the early 18205. It was said to come from
the Ethiopian area ('habitat in Abyssinia' - Wiegmann, 1837) and more specifically
from Massawa, Eritrea ('aus Massaua' - Peters, 1865), but in the 150 years since the
type was collected, no further Scincus appear to have been found in the Horn of
Africa (i.e. Ethiopia including Eritrea, French Territory of Afars and Issas and the
Somali Republic). As Scincus are often fairly conspicuous where they occur, this
may indicate that the type locality of 5. hemprichii is incorrect. Ehrenburg and
Hemprich also collected in southwest Arabia, so there might have been some con-
fusion over the origin of the type. It seems best, for the present, to treat the
northeast African occurrence of S. hemprichii with caution.

MATERIAL EXAMINED (n = 31). Abbreviations are explained on p. 191 and
localities are shown in Fig. 7.

SAUDI ARABIA

i. Sabya, I7°O7'N 42°35'E ; BM 1964.301 ; $.

i. Near Sabya, i70og'N 42°36'E ; CAS 134131-4.

i. Hakiwah, i7°oi'N 42°5o'E, 6om ; BM 1976.1858 ; (J.

1. Khasawiyah, Wadi Damad, i7°oo'N 42°o7'E, 3om ; 2 ads., juv. Unregistered live specimens.

2. Ras-al-Bourge, Jizan, i6°53'N 42°32'E ; CAS 134154-6, 134161, 134164—5.
2. Matri Wells, 22 km east of Jizan, i6°58'N 42°42'E ; CAS 119221.

2. South of Matri Wells, i6°58'N 42°44'E ; CAS 119222.

YEMEN

3. Near Zohra (= Az Zuhra) ; BM 1963.799-800*, MCZ 112245 ; 2$$, juv.

4. i mile [1.6 km] east of Hodeida ; FMNH 66311 ; $.

DEMOCRATIC REPUBLIC OF YEMEN

5. Haithalhim ; BM 95.5.23.77* ; °-.

5. Laheg (= Lahej), Aden ; BM 95.5.23.78 ; Q*-

5. Lahej and south to Sheikh Othman ; BM 99.12.13.76-7 ; 2<$<$.

6. Sheikh Othman ; BM 99.12.13.78, 79*-8i ; (J, 3 juv.
6. Aden ; SMF 21657 ; 6*-

DOUBTFUL LOCALITY

'Abyssinia' ('aus Massaua' -fide Peters, 1865) ; ZBM 1179 ; <$.

SPECULATIONS ON THE HISTORY OF THE GENUS SCINCUS

Relationship of Scincus to other skinks

Scincus is primarily an active forager on the surfaces of desert dunes and their
perimeters. As such, it occupies a unique adaptive niche amongst the skinks.
Some other groups, e.g. Sphenops Wagler 1830 and some members of Ophiomorus
Dumeril & Bibron, 1839 and Typhlosaurus Wiegmann, 1834, also inhabit sandy
desert environments, but they tend to be long-bodied and more or less cryptic,

236 E. N. ARNOLD AND A. E. LEVITON

spending a great part of their life below the sand surface rather than submerging
intermittently, mainly as an antipredator device, as Scincus does.

Scincus has a number of features that appear to be almost certainly primitive
(plesiomorphic in Hennig's terminology) within the Scincidae. These include an
unmodified palate, in which the palatine bones are not in contact, unfused frontal
bones, a full complement of skull elements, pterygoid teeth, an open Meckel's groove,
an unelongate body, well developed limbs, an unmodified postcranial skeleton, and
nostrils set between nasal scales and not broadly bordered by, or incorporated in,
the rostral. Furthermore, at least one member is egg-laying rather than live-
bearing (S. scincus}. Greer (1970) provides arguments for the primitiveness of
these characters. The first two features place Scincus within Greer's subfamily
Scincinae. Some of the modern members of this assemblage appear to be close to
the stock from which the other three subfamilies of the skinks recognized by this
author were derived. The only other genera that have all these unspecialized
features are Eumeces and Scincopus. Like Scincus, they have a high number of
supraocular scales (often six) ; from its correlation with numerous other features
believed to be primitive, it seems possible that this is also an unspecialized feature.

The characters of Scincus that appear to be apomorphic within the skinks include
fusion of the postorbital and postfrontal bones, fusion of the originally paired
premaxillae and a wide range of features showing clear functional association with
life in loose sand, such as fringed digits, spatulate snout supported by a bony pre-
maxillary rostrum, modified ears, valvular nostrils, countersunk jaw etc. The
bone-fusion characters occur sporadically in a number of disparate scincid groups
and may well have been developed several times. Some other skink genera that
live in dune habitats also have sand modifications, but their resemblance to Scincus
in these features seems to be superficial and is likely to be a result of convergence or
parallelism. A number of factors suggest this, (i) Sand modifications tend to
differ in detailed structure from group to group indicating that they are probably
not a result of recent common ancestry. Thus, although sand-dwelling Sphenops
and Ophiomoms have spatulate snouts, they lack the bony supporting rostrum of
Scincus. Similarly, while nostrils tend to be crescentic and valvular, the arrange-
ment of the scales surrounding them varies considerably and, although all the groups
have the ear structure modified to protect the tympanum and probably to improve
the reception of subterranean vibrations, the way this is done is not precisely the
same. (2) Sand habitats are known to be powerful producers of striking convergences
in lizards and a number of lizard families (e.g. Agamidae, Iguanidae, Lacertidae,
Scincidae, Gerrhosauridae) have developed a range of analogous structures in this
environment. So, parallelism or convergence within a single family, such as the
skinks, would not be unexpected. (3) When sand-related characters are set aside,
the other sand-dwelling skinks do not have a close resemblance to Scincus in other
features.

Because of the strong possibility of convergence in many apomorphic characters,
it is not possible, on present evidence, to relate Scincus certainly to any other
scincid genus on the basis of these features. Nor is Scincus likely to be identical
with, or very similar to, a stock that has given rise to any other modern genus, for

REVISION OF SCINCUS 237

some of its apparently apomorphic features (e.g. fringed digits, and bony pre-
maxillary rostrum) seem to be unique in the family. But it appears worth enquiring
whether either of the genera with which Scincus shares numerous plesiomorphic
features could be considered close to the line that produced it. As will be shown,
there are reasons for disqualifying Scincopus (see below), leaving only Eumeces to be
considered. It has already been suggested (Greer, 1970) that its members are among
the most primitive of living skinks. The genus is Holarctic in distribution and the
American and east Asian species appear to be closely related to each other, but the
rest of the genus, which is separated by a hiatus in central Asia, seems to stand apart.
It comprises only E. taeniolatus and the E. schneideri group. E. taeniolatus is rather
aberrant in having an elongate habitus and a row of expanded scales on the vertebral
midline, but the E. schneideri complex, especially E. (s.) algeriensis, has more overall
similarity to Scincus than the rest of the genus does. A distinct, although small
meatal cartilage is present, at least in the young (also found in some other species of
Eumeces and in some other scincines, e.g. Chalcides] and the ear opening is often
partly protected by two scales, the upper overlapping the lower, as in Scincus.
Whereas the dorsal pattern of most Eumeces consists of longitudinal stripes in the
young, which are often retained by adults, E. (s.) algeriensis is peculiar in having
irregular transverse banding, consisting of areas of scales bearing dark and light
spots separated by regions of light scales that are orange or yellow in life. Although
the bands are more numerous than in any Scincus, this pattern is generally reminis-
cent of that found in banded Scincus and it is likely that the latter could be derived
from it. Also, while the diploid chromosome number of all nine east Asian and
American species of Eumeces investigated to date is 26, that of E. (s.) schneideri is
32, the same number that occurs in 5. scincus, the only Scincus that has been karyo-
typed so far (Gorman, 1973). Although E. (s.) algeriensis has not yet been checked,
it seems likely that it will have a similar count to E. (s.) schneideri. Other factors
that enhance the case for E. (s.) algeriensis-like skinks being ancestral to Scincus
include the following, (i) This form and other west Palaearctic Eumeces seem to be
better adapted to life in arid areas than most other members of the genus. (2) The
distribution of the E. schneideri complex borders that of Scincus, running along the
northern edge of the range of that genus. (3) There is palaeontological evidence
that skinks generally similar to the members of the E. schneideri complex may have
been at least intermittently present in northwest Africa for a long time, possibly
since at least the Miocene (Hoffstetter, 1961). So, while it would be foolhardy to state
categorically that E. (s.) algeriensis is identical with, or very closely related to, the
precursor of Scincus, it is at least apparent that the Eumeces schneideri complex has
been capable of giving rise to a form with the qualifications to be expected in a
Scincus ancestor.

Scincopus occupies similarly arid habitats to Scincus (but apparently not very
mobile sand systems) and is sympatric with it in northern Africa, but the following
specialized features exclude it as a forefunner of the latter genus.
i. As in Scincus, the ear opening is well protected, but this is brought about not
by lengthening the meatus but by the great enlargement of the two scales that
overlap the opening. Furthermore, no meatal cartilage appears to be present.

238 E. N. ARNOLD AND A. E. LEVITON

2. Scincopus is nocturnal and has a vertically elliptical pupil and a large cornea.
These features are found in Scincus hemprichii, but are absent in the most primitive
constituent of the genus, the 5. scincus complex (see below), and in the remaining
species, S. mitranus, as well. This suggests that their presence in S. hemprichii is a
secondary development and that the precursor of Scincus lacked them.

3. Scincopus lacks any obvious sexual dimorphism in size, yet this feature is present
both in Scincus and Eumeces.

4. Like many Scincus, Scincopus has dark transverse bars on the dorsum, but their
arrangement differs. The first body band is behind the level of the forelegs and not
roughly coincident with them ; there are only four bands between the forelegs and
the vent region (minimum of five in Scincus) and they extend to the tail tip.

All four features of Scincopus seem to be derivable from conditions found in
E. (s.) algeriensis, as are equivalent characters of primitive Scincus. However, the
specialized characters found in the latter do not appear to be derivable from those
occurring in Scincopus, or vice versa. In view of this, it is best to regard
both Scincus and Scincopus as independent derivatives of a E. (s.) algeriensis-like
stock.

Interrelationships of the species of Scincus

In this discussion, the 5. scincus complex will be treated as a single unit. The
main characters known to vary between the members of the genus Scincus are set
out in Table 5 and compared with the conditions present in Eumeces (s.) algeriensis
which, it has been suggested, may well be similar to the stock that gave rise to
Scincus. In most cases, the character states found in E. (s.) algeriensis are quite
widespread in the Scincidae as a whole and often in other scincomorph groups as well.
This, together with the presumed similarity of E. (s.) algeriensis to the ancestor of
Scincus, suggests that alternative states in the latter genus are apomorphic and that
states identical with, or similar to, those of E. (s.) algeriensis are plesiomorphic. It
will be seen that the S. scincus complex has by far the most characters that are like
their counterparts in Eumeces, or that approach closer to these counterparts than do
those of other Scincus species (see section A of Table 5 for summary) . This is true
for 21 out of 23 characters in the 5. scincus complex, but for only five in 5. mitranus
and eight in 5. hemprichii, which indicates that the S. scincus complex is the most
primitive constituent of the genus.

It seems likely that the two more specialized species were derived from a form
generally similar to S. scincus and very doubtful that they had a separate origin
from a very much more primitive form. Of the characters tabulated (see section B
of Table 5), they have a number that are more similar to those of S. scincus than to
those of Eumeces, but very few, or none, where the reverse is true. In the case of

5. mitranus, n out of 23 characters are about equally similar to the conditions found
in S. scincus and Eumeces, 12 are more similar to S. scincus and none is more similar
to Eumeces. In the case of S. hemprichii, 16 out of 23 are equally similar, six are
more similar to S. scincus and only one is more similar to Eumeces. However, to
gain a real idea of the greater relative similarity of these species to the 5. scincus
complex rather than to Eumeces, the characters common to all Scincus, but absent in

REVISION OF SCINCUS 239

Eumeces, must be included, viz. fuzed premaxillae, fused postfrontal and post-
orbital bones and the extensive range of adaptations to life in loose sand.

Having assumed an origin from a S. scincus-like form, it remains to inquire how
S. mitranus and 5. hemprichii are related to each other. Two extreme possibilities
exist : (i) they arose as a common stock that evolved a substantial number of
apomorphic characters before they diverged, or (2) they arose as independent
derivatives of a S. scincus-like form. If the first possibility were true, it would be
expected that S. mitranus and 5. hemprichii would share a high proportion of the
apomorphic characters that they possess relative to 5. scincus. Also, it is likely
that their overall resemblance to each other would be greater than the overall
resemblance of each to the 5. scincus complex. When apomorphic characters are
considered, it will be seen that both 5. mitranus and S. hemprichii show similar
apomorphic trends in characters I, 2, 4, 5, 6, 7, 9, 10, n and 19, but in characters
3, 12 and 18 only S. mitranus is apomorphic and in characters 8, 13, 14, (15), 17 and
21 this is true only of 5. hemprichii. So shared trends make up only just over half
of the total apomorphic features shown by S. mitranus and 5. hemprichii.

It could be argued that these shared apomorphic trends developed in common
before divergence. But, apart from character 19 (extension of the mid-dorsal scale
series of the tail), all are confined to one organ system, the ear and associated struc-
tures. As has been pointed out, a number of skink stocks show similar adaptations
of the ear to sandy habitats, so parallel development in S. mitranus and 5. hemprichii
would not be impossible. Although both species show similar apomorphic trends
in the ear, these are developed to different extents. Thus, S. mitranus is more
specialized in characters 2 and 3 and 5. hemprichii in i, 4, 5, 6, 7, 8, 9, 10 and n.
Therefore the amount of development that could have occurred in a common stock
is quite restricted.

Estimation of overall resemblance also indicates that any common origin that
5. mitranus and S. hemprichii had from 5. scincus-like. skinks must have been limited.
Comparisons of the three species pairs within Scincus are given below.

Features most similar in
Identical features given pair

S. scincus -S. mitranus 10 n

S. scincus - S. hemprichii 6 3

S. mitranus - S. hemprichii o g

Thus, on the basis of identical characters, there is little resemblance between
S. mitranus and 5. hemprichii, whereas both of these have considerable affinity to
5. scincus. If the (less important) features of comparative similarity are also taken
into account, the clarity of the arrangement is somewhat reduced, but there are still
no grounds for thinking of S. mitranus and 5. hemprichii as being closer to each other
than to the 5. scincus complex.

In summary, it seems likely that S. mitranus and S. hemprichii either had a very
restricted common evolution from the S. scincus complex or none at all, and that
there was parallel development of some ear characters.

240

E. N. ARNOLD AND A. E. LEVITON

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REVISION OF SCINCUS

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242 E. N. ARNOLD AND A. E. LEVITON

Origin of species distribution pattern

It has been suggested above that the widespread S. scincus complex is the most
primitive component of the genus and that the Arabian species, 5. hemprichii and
S. mitranus t either are each independent derivatives of a S. scincus-like stock, or had
only a restricted common history. This pattern of speciation and distribution may
partly stem from changes in the map of aridity during the Quaternary. Just as
fluctuations in the extent of the South American rain forest are thought to have
promoted speciation in the iguanid Anolis chrysolepis group (Vanzolini & Williams,
1970), so the contraction and fragmentation of the desert areas of the Saharan -
Arabian region, followed by their renewed expansion, could be seen as a crucial
factor in the differentiation of the species of Scincus. If the Saharan- Arabian
deserts were originally occupied by populations similar to the present 5. scincus
complex, change to more mesic conditions would have fragmented this continuum
into a series of isolates. The area of aridity would have undoubtably contracted
and this is likely to have severed contact between the African and Arabian deserts
through Sinai. It is also possible that the contraction of dry areas may have
separated the main southern Arabian desert from the arid southwestern coastal
strip of the peninsula. Today, these two dry areas are largely divided by high
mountains and a relatively modest extension of mesic conditions would have isolated
them completely. Another factor that may be important in separating the Scincus
of extreme southwestern Arabia from the rest is the presence of geophysical barriers
that interrupt the Tihama, the sandy plain running along the western seaboard of
Arabia. At the present time this is crossed by extensive gravel fans that probably
do not provide suitable habitats for Scincus and south of these, at about latitude
i8°3o'N, is the Harrat Habhab, a large lava flow. Although this is now largely
covered by sand, it could have been important in isolating the Scincus of south-
western Arabia in the past. Alternatively, if its origin predates their isolation, the
populations might have been continuous around its outer edge during a period of
lowered sea level but became isolated when this rose.

Whatever the details of the situation in southwest Arabia, it seems feasible that
the once more or less continuous Saharan -Arabian desert became divided into at
least three separate refuges (or complexes of refuges) in which the present 5. scincus
complex, 5. mitranus and 5. hemprichii differentiated. If arid conditions then again
became widespread, the three allopatric units so formed would have been inclined
to expand, although, as has been suggested, the southwest Arabian 5. hemprichii
was probably restricted by geomorphological features. It is to be expected that
such range extension would have been initially greatest among populations that were
not strictly adapted to very arid habitats, since the more mesic areas surrounding the
refuges would not dry out immediately.

On this premise, one might predict that S. scincus would be the first to spread, for
morphological evidence and the limited ecological data available suggest that it is
less strictly adapted to very arid conditions than 5. mitranus and 5. hemprichii.
This seems to have been the case, for S. scincus now occurs in both North Africa and
Arabia and it seems likely that it spread eastwards through the Sinai area, with the
onset of drier conditions, and invaded much of the Arabian peninsula.

REVISION OF SCINCUS 243

However, this advance into Arabia appears subsequently to have been partly
reversed by expansion of the endemic 5. mitranus. The most southern and eastern
populations of 5. scincus in the peninsula, 5. s. conirostris, occur very close to (or in
actual contact with) 5. mitranus at a number of places including Kuwait, Abqaiq,
Dhahran, eastern United Arab Emirates (Ra's al Khaymah and Jazirat al Hamra)
and Hadhramaut. All these localities are on the periphery of the range of 5.
mitranus and no 5. s. conirostris are presently known from the more central parts of
its distribution, nor do populations of S. mitranus seem to occur far within the range
of S. scincus. This suggests that the two species are not usually capable of existing
in close and stable sympatry. The areas where they do occur together appear to be
localities where really desertic areas, with extensive moving sand, border rather more
mesic territory with a more stable sandy substrate. In such places, S. mitranus
appears to occupy the former regions and S. s. conirostris the latter. This seems to
be the situation in the eastern United Arab Emirates. Here S. mitranus was the
sole form encountered in large dune systems and S. s. conirostris was seen only in an
area of mainly stabilized sand ridges. Even in this area, the few mobile sand slips
were occupied by S. mitranus. Similarly, in the Hadhramaut, S. mitranus is known
from the sand sea or its edges, whereas 5. s. conirostris appears to occur in the more
stable areas of Wadi Hadhramaut itself.

It might be thought that S. scincus was merely halted by ecological exclusion when
it reached the periphery of the range of S. mitranus. But at least one population of
S. scincus conirostris, that in the eastern United Arab Emirates, is now apparently
completely isolated, and in the comparatively well-worked area between it and the
nearest other known population (Abqaiq), only S. mitranus is known. It is possible
that some of the other S. s. conirostris localities such as those in the Hadhramaut are
also cut off in this way. Such relicts suggest that S. scincus has actually lost ground
to 5. mitranus in Arabia.

As stated, most records of 5. s. conirostris come from the periphery of the range of
5. mitranus and it may be that some of the characteristic features of this subspecies
have arisen from this contact by character displacement (in the sense of Brown &
Wilson, 1956) or at least have been maintained by it. Certainly 5. mitranus has
more characters in common with the allopatric 5. s. meccensis of west and north-
western Arabia than it does with S. s. conirostris. In contrast to the latter, it shares
with S. s. meccensis a narrow head in adults, a strongly spatulate snout with well-
developed rostral-frontonasal contact and dark bars on the flanks. As we have
seen, 5. mitranus appears to exclude 5. s. conirostris from the extensive loose-sand
habitats that are typical of the genus Scincus and some of the characteristic features
of 5. s. conirostris appear to reflect this. In most populations of Scincus, the head
is relatively narrow and this seems to be functionally important in sand-diving.
However, in areas with relatively limited loose sand this feature may well be at a
lower premium and the advantages of a bigger, broader head, typical of many
skinks, would assert themselves. A shorter, less spatulate snout which does not
involve well-developed rostral-frontonasal contact, can be 'explained' in a similar
way. The bars on the flanks of Scincus seem to be important sign-stimuli and
captive 5. mitranus often appear to assume a position where they are conspicuously

244 E- N- ARNOLD AND A. E. LEVITON

displayed to other individuals. In areas of sympatry, species recognition would be
enhanced if one species had bars and the other did not.

It might be argued that if the characters of 5. s. conirostris are a result of dis-
placement, then neighbouring populations of S. scincus not in contact with S.
mitranus should lack them. This is largely so since they are missing in the adjacent
5. s. meccensis. The only S. s. conirostris outside the known range of 5. mitranus
occur to the north in a relatively small area of Iran, Iraq and extreme northern
Arabia. These retain all the characters diagnostic of 5. s. conirostris, possibly partly
because the area contains no really extensive dune systems such as are occupied by
S. s. meccensis and S. mitranus so there is no strong pressure to develop the features
typical of these forms. This allopatric area may represent a relatively recent
extension of the range of 5. s. conirostris.

Although the basic pattern of speciation and distribution of Scincus can be
interpreted in terms of three refuges and two arid periods separated by a more mesic
one, it is likely that the history of the genus has been more complex. Certainly, the
climatic fluctuations must have been more complicated than suggested here. Palaeo-
climatologists no longer adhere to a simple scheme . of progressive low-latitude
desiccation following a 'pluvial' corresponding to high-latitude glaciation, and are
indeed reluctant to assume that any episode of moister conditions was felt throughout
the area under discussion. Nevertheless, the reality of such changes is accepted
(see, e.g., Butzer, 1972). Also, it seems possible that at least some of the main
desert refuges were divided further during the most mesic climatic phases. Such
subdivision would help explain the high level of variation in the North African S.
scincus complex.

If the above explanation of the origin of the pattern of speciation and distribution
in Scincus is valid, it would be expected that other animal groups with similar
general requirements should show a similar pattern. Unfortunately, most other
Saharan- Arabian lizard groups are in need of thorough revision, but analogies to
the Scincus situation can be found in the gekkonine genus Stenodactylus and in the
lacertid, Acanthodactylus scutellatus group (see Table 6). Both these assemblages
contain desert-adapted species largely associated with sandy habitats. In Steno-
dactylus, the species often show the same basic distributions as those of Scincus,
there being one occurring in both North Africa and Arabia, three confined to Arabia
excluding the southwest of the peninsula and two confined to the southwest. In

TABLE 6

Species distribution of some mainly sand-dwelling lizards in the Saharan Arabian desert area.

North Africa and Arabia only, but Southwestern coast
Genus north Arabia not southwestern coast only

Scincus S. scincus complex S. mitranus S. hemprichii

Stenodactylus S. sthenodactylus S. slevini Stenodactylus n. sp.

S. doriae S. pulcher

S. arabicus

Acanthodactylus A. scutellatus A. arabicus schmidtii A. arabicus arabicus

REVISION OF SCINCUS 245

the Acanthodactylus scutellatus species group, A. scutellatus is North African, with
one population, A. s. hardyi, entering northern Arabia, while at least one form,
A. arabicus* is confined to Arabia and has a distinctive subspecies in the southwest.

SUMMARY OF SYSTEMATIC CONCLUSIONS

Scincus can be divided into three clear entities as follows.

1. Scincus mitranus is confined to Arabia, but is absent from the west and south-
west of the peninsula. Its synonymy includes 5. arabicus, S. arenarius, S. musca-
tensis, S. philbyi and 5. richmondi. S. mitranus shows considerable geographical
variation : specimens from southeastern Oman and from the north of the range
have distinctive features, but judgement is reserved as to whether distinct subspecies
should be recognized. The Pakistan record of S. mitranus, under the name S.
arenarius, is considered to be erroneous.

2. Scincus hemprichii is probably limited to the coastal lowlands of southwestern
Arabia and its supposed occurrence in Eritrea is doubtful. It is a very cleaily
defined species that exhibits little obvious geographical variation.

3. The Scincus scincus complex includes all other members of Scincus and occurs
over nearly the whole known range of the genus, but is absent from southwestern
Arabia. In North Africa, the populations fall into two groups, which may represent
largely allopatric species for which the oldest names are S. scincus and S. albifasciatus.
However, evidence for specific status is not conclusive. The scincus group of
populations can be divided into scincus s. sir. in Egypt and Israel and cucullatus in
northeast Algeria, Tunisia and northwest Libya ; animals from northeast Libya are
intermediate. Isolated populations in southeast Algeria and northwest Nigeria
also seem to belong to the scincus group, but cannot be allocated to either of the
named forms. The albifasciatus group of populations consists of albifasciatus
s. sir. in West Africa, from Rio de Oro to Senegal and eastwards through Mali and
Niger to southeast Libya, and laterimaculatus in northwest Algeria ; the two forms
appear to be connected by intermediates in southeast Morocco and in Mauretania.

In southwest Asia, the Scincus scincus complex is represented by two quite
well-defined allopatric entities : 5. scincus meccensis (incorporating S. deserti) in
southern Jordan, northwest and west Arabia and 5. s. conirostris (including 5.
gasperettii) in south and east Arabia, Iraq and southwest Iran.

It is also suggested that lacertid lizards in Arabia usually assigned to Acantho-
dactylus cantoris represent a distinct species for which the oldest name is A . arabicus
Boulenger, 1919.

ACKNOWLEDGEMENTS

Any revision of a group that is reasonably abundant in museum collections is
inevitably very dependent on the aid and goodwill of workers in other institutions.
We would consequently like to acknowledge the willing help given to us by the

* The Arabian lizards assigned to Acanthodactylus cantoris seem not to be conspecific with this form and
have more affinity with A . scuttellatus. They are therefore recognized as a distinct species for which the
oldest available name is Acanthodactylus arabicus Boulenger, 1919.

246 E. N. ARNOLD AND A. E. LEVITON

following people, who either lent material or provided information : Mr J. C.
Battersby, Professor J. Bons (Laboratoire de Biogeographie et Ecologie, Mont-
pellier), Dr J. Castroviejo (Estacion Biologica de Donana, Seville), Dr C. Edelstam
(Naturhistoriska Riksmuseet, Stockholm), Dr J. Eiselt (Natur-historisches Museum,
Vienna), Dr G. W. Foley (American Museum of Natural History, New York), Dr U.
Griiber (Zoologische Sammlung des Bayerischen Staates, Munich), Professor J. Guibe
(Museum Nationale d'Histoire Naturelle, Paris), Dr S. W. R. Heyer (United States
National Museum, Washington), Mr P. W. Hopkins (Estacion Biologica de Donana,
Seville), Dr K. Klemmer (Natur-Museum und Forschungs-Institut Senckenberg,
Frankfurt a.M.), Dr H. Marx (Field Museum of Natural History, Chicago), Dr C. J.
McCoy (Carnegie Museum, Pittsburgh), Dr G. Peters (Zoologisches Museum, Berlin),
Dr Thys van den Audenaerde (Koninklijk Museum voor Midden-Afrika, Tervuren),
Professor E. Tortonese (Museo Civico di Storia Naturale, Genoa), Dr J. A. Valverde
(Estacion Biologica de Donana, Seville), Dr Y. Werner (Hebrew University of
Jerusalem), Professor E. E. Williams (Museum of Comparative Zoology, Harvard),
and the Director, Institut Fondamental d'Afrique Noire, Dakar.

Others, knowing of our interest in Scincus, went out of their way to collect fresh
material, often from critical areas. Amongst these are Major M. D. Gallagher,
Mr J. Gasperetti, Mr M. C. Jennings, Mr J. P. Mandaville, Mr and Mrs E. Morris and
Major D. J. G. Williams. Other important specimens were obtained by Mr G.
Popov.

Major Gallagher also arranged for the senior author to visit the United Arab
Emirates on two occasions, during both of which Scincus were observed and collected.

Dr C. Vita-Finzi (University College, London) gave very helpful advice on climatic
changes in the Quaternary and Mrs F. E. Warr provided information on obscure
Arabian localities.

Skeletal material and radiographs were prepared by Mr C. J. McCarthy, Figs 1-4
were drawn by Mr D. W. Ovenden, photographs were taken by Mr F. Greenaway,
and drafts of the manuscript were typed by Mrs S. C. Trigonopoulus and Mrs
Renate Arnold.

REFERENCES

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115-116.
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Batrachians of Egypt. London, 122 pp.
1898. Zoology of Egypt: Reptilia and Batrachia. London, lxiv + 3yi pp.

ANGEL, F. & LHOTE, H. 1938. Reptiles et Amphibiens du Sahara central et du Soudan.

Bull. Com. Etud. hist, scient. Afr. occid.fr. 21 (3) : 345-84.
ANON. (= ANDERSON, J. ?) 1871. (Gift) from Babu Rajendralala Mitra - a dried specimen of

a new species of Scincus. Proc. Asiat. Soc. Beng. 1871 : 96.

ARNOLD, E. N. (in preparation). A review of the genus Stenodactylus (Reptilia Gekkonidae).
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the Sultanate of Oman. Results of the Oman Flora and Fauna Survey, 1975.
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reproduction of Chalcides ocellatus (FORSK.) and Scincus scincus (LINN.) Bull. Fac. Sci.

Egypt. Univ. 39 : 179-185.

REVISION OF SCINCUS 247

BAIRD, I.L. 1970. The anatomy of the reptilian ear. In Cans, C. et al. Biology of the Reptilia,

vol. 2. London & New York, xv + 374 PP-

BENT, T. & BENT, M. V. A. 1900. Southern Arabia. London, xii + 455 pp.
BLANFORD, W. T. 1881. On a collection of Persian reptiles recently added to the British

Museum. Proc. zool. Soc. Lond. 1881 : 677-682.
BOULENGER, G. A. 1890. First report on additions to the lizard collection in the British

Museum (Natural History). Proc. zool. Soc. Lond. 1890 : 77-86.

BROWN, W. L. & WILSON, E. O. 1956. Character displacement. Syst. Zool. 5 (2) : 49-64.
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CHEESMAN, R. E. 1926 In Unknown Arabia. London, xvi + 447 pp.
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systems, vol. 3. Leipzig, 980 pp.
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(continued and completed). Niger. Fid 38 (2) : 54-80.
EL-TOUBI, M. R. 1938. The osteology of the lizard Scincus scincus (Linn.). Bull. Fac. Sci.

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In Chirelli, A. B. & Capanna, E. Cytotaxonomy and vertebrate evolution. New York.
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248

E. N. ARNOLD AND A. E. LEVITON

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VANZOLINI, P. E. & WILLIAMS, E. E. 1970. South American anoles : the geographic dif-
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WERNER, Y. L. 1971. Lizards and snakes from Transjordan, recently acquired by the

British Museum (Natural History). Bull. Br. Mus. nat. Hist. (Zool.) 21 (6) : 213-256.
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— 1837. Herpetologische Notizen. Arch. Naturgesch. 3 : 123-136.

E. N. ARNOLD

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7 5BD

A. E. LEVITON
Department of Herpetology
CALIFORNIA ACADEMY OF SCIENCES
SAN FRANCISCO 94118

PLATE i

Scincus scincus complex from North Africa : variations in pattern.

a. 'cucullatus' : few, narrow bands ; no side-bars. BM 1920.2.20.3848 ; El Oued, Algeria ;

6*.

b. 'scincus': numerous fairly broad bands ; no side-bars. BM 97. 10.28. 439 ; Abu Ruwaysh,

Egypt; 6*-

c. ' later imaculatus' : no bands, but dark dorsolateral streaks on head and neck ; side-bars
present. MHNP 46.305 ; Grand Erg Occidental, near Beni Abbes, Algeria ; c?.

d. ' albifasciatus' : few, broad bands ; side-bars present. EBD 6190 ; El Aaiun, Spanish
Sahara ; $.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 5

PLATE i

PLATE 2

Scincus from Arabia : dorsal patterning and body proportions.

a. 5. scincus conirostris : BM 97.3.11.101 ; Hadhramaut, Democratic Republic of Yemen ; <$.

b. S. mitranus : BM 1971.1023 ; Sharjah, United Arab Emirates ; <J.

c. 5. hemprichii : BM 95.5.23.78 ; Lahej, Democratic Republic of Yemen ; £.

Bull. Br. Mus. nat. Hist. (Zool.) 31,5

PLATE 2

PLATE 3
Scincus from Arabia : live animals.

a. S. scincus meccensis : Burayman, Saudi Arabia (collected by Mr J. Gasperetti).

b. 5. scincus conirostris : S. of Amar, Bahrain (collected by Major M. D. Gallagher

c. S. mitranus : Kuwait (lent by Mr D. Maclntyre).

d. 5. mitranus : south of Jazirat al Hamra, United Arab Emirates.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 5

PLATE 3

•*"' -.;-'; •- ;--%N%J<"'! v'-. \f": *i ,.. *,\-.-X. '/';>x'' i'-«Y

A LIST OF SUPPLEMENTS
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1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
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1965. (Out of Print.)

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and Mineralogy
of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ; 29 Plates,
77 Text-figures. 1969. £4.50.

4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £10.80.

5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72
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6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the
Biology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures.
1974. £3 -75. Hardback edition £6.

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Department.

This paper is Vol. 31, No. 6, of the Zoology series.
The abbreviated titles of periodicals cited follow those
of the World List of Scientific Periodicals.

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

ISSN 0007-1498

Trustees of the British Museum (Natural History), 1977

BRITISH MUSEUM (NATURAL HISTORY)

Issued 30 June, 1977 Price £5.50

CONTENTS

Page

The morphology of a common soil flagellate, Heteromita globosa Stein (Masti-
gophorea : Protozoa). By C. M. MACDONALD, J. F. DARBYSHIRE & C. G.
OGDEN 255

Notes on the morphology and nomenclature of three members of the Euploti-

dae (Protozoa : Ciliatea). By C. R. CURDS 267

Notes on the Alpheids in White's List of the specimens of Crustacea in the
collections of the British Museum (1847). By A. H. BANNER & D. M.
BANNER ........... 279

Establishment of a new genus in the family Phoxocephalidae (Crustacea :
Amphipoda) and a description of a new species from North Island, New
Zealand. By A. A. FINCHAM 285

A new species of the synaptid genus Rynkatorpa Rowe & Pawson, 1967 (Holo-

thurioidea : Apodida) from the Great Barrier Reef. By F. W. E. ROWE . 293

A review of the family Centropomidae (Pisces, Perciformes) : an appendix.
By P. H. GREENWOOD 297

A revision of the synonymy of three species of Leiolopismid skinks from New
Zealand. By. J. ROBB ......... 303

THE MORPHOLOGY OF A COMMON SOIL

FLAGELLATE, HETEROMITA GLOBOSA STEIN

(MASTIGOPHOREA : PROTOZOA)

By C. M. MACDONALD, J. F. DARBYSHIRE & C. G. OGDEN

CONTENTS

Page
SYNOPSIS ........... 255

INTRODUCTION .......... 255

MATERIALS AND METHODS ........ 255

KEY TO ABBREVIATIONS USED IN THE PLATES AND FIGURES . . . 257
OBSERVATIONS .......... 257

Light microscopy . . . . . . . . . .257

Electron microscopy . . . . . . . . .260

DISCUSSION ........... 262

ACKNOWLEDGEMENTS ......... 263

REFERENCES ........... 263

SYNOPSIS

Heteromita globosa, a colourless flagellate, was established in clonal culture. The ultrastructure
is described for the first time. Two organelles, hitherto undetected in soil flagellates, were
observed, but many diagnostic features of Bodonid flagellates were absent. The affinities of
H. globosa are discussed, and it is suggested that Heteromita is a valid genus, clearly distinct
from Bodonid flagellates.

INTRODUCTION

ALTHOUGH colourless flagellates of the genus Heteromita are common in soils, their
ultrastructure is unknown, and their morphology has not been studied with modern
microscopes. The last morphological study was by Robertson (1928). Recent
systematic accounts do not recognize the genus Heteromita Dujardin, 1841 and place
these flagellates in the genus Bodo Ehrenberg, 1830 (Hollande, 1952 ; Lepinis et al.t
1973). Nevertheless, observations on the external morphology and ultrastructure
of a Scottish isolate of Heteromita globosa Stein, reported in this paper, suggest that
Heteromita is a valid genus.

MATERIALS AND METHODS

The flagellate was separated in December 1971 from other protozoa by diluting a
soil sample (plough or S layer, Tarves Soil Series, Tarves Soil Association at Tilly-
corthie Farm, near Udny, Aberdeenshire) to extinction with sterile liquid soil extract
and microdiluters (Darbyshire et al. , 1974) . Five clones of H. globosa were isolated by

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6 Issued 30 June, 1977

256 C. M. MACDONALD, J. F. DARBYSHIRE & C. G. OGDEN

diluting the original pure culture to extinction with the same method, and by choos-
ing individual cultures in the dilution series with single flagellates. Another clone
was isolated by pipetting minute drops from this second dilution series with a fine
capillary on to agar squares (about 5 mm2 of i% w/v Oxoid agar No. 2) inside
sterile Petri dishes (12 agar squares/Petri dish of 10 cm diam.). A small piece of a
coverslip was placed on each agar square, and single cell isolates were selected under a
light microscope. Subsequently, one agar square with a single cyst was transferred
to sterile soil extract and the clonal culture was derived from this cyst. The 6 clonal
cultures were subcultured into sterile soil extract at weekly intervals and incubated
at either 25 or 10 °C.

Living flagellates were observed in liquid mounts and filmed in agar chambers of
Heunert (1962). Eastmann Plus X negative film was used for 16 mm cinephoto-
micrography. The flagellates were preserved with either the fixative of Nissenbaum
(1953) or with a i : 6 mixture of saturated HgQ2 and OsO4 (2% w/v). After the
latter fixative, the flagellates were dehydrated in a series of acetone/water mixtures
and absolute acetone, before they were mounted in epoxy resin (Spurr, 1969) on
microscope slides. Three permanent slides have been deposited in the British
Museum (Natural History) and registered with the following numbers : 1976:
1:20:1, 1976:1:20:2, and 1976:1:20:3. After Nissenbaum's fixative, specimens were
stained with acridine orange (io~4 M in citric acid-Na2HPO4 buffer, pH7), or
Giemsa (5% v/v in S0rensen's buffer, pH 7) or toluidine blue (0-5% w/v) in borax
(0-5% w/v). Some flagellates were drawn (Fig. i) with the aid of a Wild M2O
microscope and a Treffenberg drawing tube. The dimensions of fixed samples of the
clonal cultures are shown in Table i.

Flagellates, from a 24 hr culture grown at 25 °C, were fixed for scanning electron
microscopy in a i : i mixture of saturated HgCl2 solution and OsO4 (2% w/v) solu-
tion. They were then pipetted on to polycarbonate membrane filters (0.2 /*m pore
diam. Nuclepore) inside a specimen carrier (Marchant, 1973) and dehydrated in a
series of ethanol/water mixtures, absolute ethanol, 'Freon H3'/ethanol mixtures and
finally absolute 'Freon 113'. The specimens were dried in a critical point apparatus
(Cohen et al., 1968). The polycarbonate membranes were mounted on scanning
electron microscope stubs with conductive paint (Silver Dag) and coated evenly with
carbon and gold/palladium alloy in an evaporating unit. The stubs were examined
in a Cambridge Stereoscan 84 at 30 kV and the results were recorded on Kodak
Tri-X Pan film.

Four different fixation methods for transmission electron microscopy were tested
involving acrolein, glutaraldehyde and OsO4 ; the most satisfactory results were
obtained when the flagellates were fixed for 15 min in Os04 (2% w/v) in cacodylate
buffer (o-i M, pH 7-1, containing 0-25% w/v CaCl2 and 0-2% w/v NaCl) (Tamm &
Tamm, 1973).

For the cytochemical demonstration of catalase, cells were fixed with glutaralde-
hyde in soil extract (final vol. 2% v/v) for 30 min, washed in half -strength soil
extract and incubated in the enzyme medium (10 mg 3, 3'-diaminobenzidine tetra-
hydrochloride (DAB), 10 ml tris buffer at pH 7-2 and o-i ml 0-3% v/v H2O2) for
i hr at a room temperature of 18-20 °C. The control medium lacked DAB but was

MORPHOLOGY OF HETEROMITA GLOBOSA

257

otherwise identical to the enzyme medium. The cells were subsequently washed for
i hr in tris buffer and post-fixed with OsO4 (i% w/v) in half -strength soil extract for
30 min (Norris & Pearson, 1975).

All the fixed cells were concentrated by centrifugation (150 g for 10 min). The
pellets were dehydrated in acetone and embedded in a low viscosity epoxy resin
(Spurr, 1969). Sections were cut with glass knives on a Reichert OMU 3 ultramicro-
tome and stained in either a saturated solution of uranyl acetate in ethanol (50% v/v)
or uranyl acetate followed by lead citrate (Reynolds, 1963). The sections were
examined in an AEI EM6 electron microscope at 75 kV.

For shadow cast preparations, live flagellates were fixed in OsO4 vapour, on
formvar coated grids. The grids were shadowed with nickel/palladium at an angle
of 30° in a coating unit.

KEY TO ABBREVIATIONS USED IN THE PLATES AND FIGURES

af anterior flagellum

b bacterium

bb basal body

cv contractile vacuole

cw cyst wall

ed electron dense deposits

er endoplasmic reticulum

f flagellum

fi striated filament

fv food vacuole

G Golgi apparatus

k kinetocyst-like organelle

lipid droplet

mitochondrion

microbody-like organelle

microtubules

multivesicular area

nucleus

nucleolus

pore

posterior flagellum

semicircular ridge of cytoplasm

tubular elements

vacuole

OBSERVATIONS

LIGHT MICROSCOPY. In the logarithmic phase of growth, the flagellate is either
ovoid (5-8 //,m long, 3-6 /mi wide) or pyroid with the anterior half of the body
slightly wider than the posterior region (Fig. i J ; Table i). Two heterodynamic

TABLE i

Mean dimensions (jttm) of H. globosa from i mixed and 6 clonal cultures
CULTURE TROPHOZOITE THIN- WALLED CYST

Mixed
Clone i
Clone 2
Clone 3
Clone 4
Clone 5
Clone 6

Body
length

6-7
6-6
7-0
6-9
6-8
6-8
7*3

Body

Body

Flagellum length

width

depth

Anterior Posterior

4-0

4'9

8-5 12-4

4*o

4-6

8-1 12-8

4-0

4'7

9*2 13*9

5'3

5'0

7-9 12-6

4'3

4'7

8*7 I3'3

4'5

4'7

8-4 12-8

4-6

4-8

8-4 13*4

Diameter
4-8
4'7
4*7
4'8
4-6

4'7
4-8

Fixed in HgQ2/OsO4 and mounted in epoxy resin; 50 cells/clonal culture and 100 cells/mixed culture
measured.

258 C. M. MACDONALD, J. F. DARBYSHIRE & C. G. OGDEN

FIG. i. Camera lucida drawings of H.
globosa. A & B are thin- walled resting
stages, C to G are stages in excystment,
H to K are trophozoites (I side view,
J dorsal view, K ventral view), L is a
mature thick-walled cyst.

10pm

acronematic flagella are inserted close together at the anterior extremity of the body
(Fig. iK). The anterior flagellum is slightly longer than the body and beats in an
antero-lateral direction in a helical manner from the proximal region to the tip.
The posterior flagellum is about twice as long as the other flagellum (Table i) and
often trails passively behind the body. Usually, the flagellate moves forward in a
vibratory manner with the anterior extremity of the body and the posterior flagellum
in contact with the substratum. The longitudinal axis of the body is maintained
at an angle of about 45° to the substratum and at an acute angle to the posterior
flagellum (Fig. ij). As the body moves forward, the longitudinal axis tends to
swing from side to side, pivoting on its anterior extremity. When the flagellate is
dislodged from the substratum and becomes free-swimming, both flagella move
haphazardly. If the body or flagella become attached to organic debris or bacteria,
then the movements become more irregular and violent.

The nucleus with its conspicuous central karyosome is anterior and on the longi-
tudinal axis of the body (Fig. i J ; PI. i). No kinetoplast was observed near the
nucleus, either in living cells or in fixed preparations stained with acridine orange.
The single contractile vacuole usually lies slightly posterior to the nucleus near the
ventral surface (PI. i). Bacteria are rapidly ingested by means of short transient
pseudopodia at any point of the body, even during cytokinesis. Sometimes, the
bacteria are first entangled with the anterior flagellum and gradually move towards
the body, where they are engulfed by pseudopodia (PI. i). After ingestion, the
bacteria are rapidly transferred to the posterior half of the body.

Reproduction can occur 10-24 nr after the addition of fresh soil extract to an
encysted culture at 25 °C and follows the stages described by Robertson (1928). The
mother cell becomes quiescent and elongates, as the flagella migrate to opposite ends
of the body. A second flagellum develops beside each 'mother' flagellum, the body
becomes constricted in the middle, and the two daughter cells pull apart until only a
thread of cytoplasm remains between them. This cytoplasmic thread normally

MORPHOLOGY OF HETEROMITA GLOBOSA

259

disintegrates ; sometimes the daughter cells coalesce, and a binucleate flagellate is
formed. Trinucleate individuals were observed ; these are formed when only one
of the daughter cells from the division of a double individual is released. Binucleate
cysts were occasionally found in the cultures and were probably derived from binu-
cleate trophozoites. A bicellular (Fig. lA) cyst was once observed in the cultures.

In addition to a thick-walled cyst (Fig. iL), H. globosa possesses a thin- walled
resting stage (Fig. iB), which was originally described by Sandon (1927). Both
resting stages developed in liquid cultures but dried cultures invariably formed thick-
walled cysts.

Encystment and excystment of Heteromita have been described in detail by
Robertson (1928). Encystment in H. globosa begins 24-48 hr after inoculation of a
batch culture of soil extract at 25 °C. During the initial stages of encystment, the
contractile vacuole is very active. Later, the flagellar movements become sluggish,
the body becomes spherical and the food vacuoles congregate around the nucleus.
The flagella decrease in length and gradually disappear.

FIG. 2. Diagram of Heteromita globosa
showing the arrangement of the main
organelles.

fv

260 C. M. MACDONALD, J. F. DARBYSHIRE & C. G. OGDEN

Excystment of the thin-walled resting stages can be readily induced by the addition
of fresh soil extract. Within 30-60 min at 25 °C, the contractile vacuole becomes
active and a small translucent pseudopodium, which gradually increases in size, soon
emerges from the cyst (Fig. iC, D, E). The anterior flagellum can be seen outside
the cyst before the posterior flagellum (Fig. iF), although they are both eventually
situated close to each other on the pseudopodium. The body is gradually extricated
from the cyst by vigorous movement of the anterior flagellum. Sometimes, the
trophozoite swims around for several minutes still attached to the empty cyst wall
(Fig. iG). Usually, the recently excysted flagellate is ovoid (Fig. iH). The whole
process of excystment extends over i-2hr at 25 °C. Excystment of the thick-
walled cysts is a similar process, but 24-48 hr elapses before it is complete at 25 °C.

ELECTRON MICROSCOPY. Trophozoite. The origin of the anterior flagellum is
partly surrounded by a semicircular ridge of cytoplasm at the anterior end of the
body. The longer posterior flagellum appears to be continuous with one end of this
ridge and is set at right angles to the anterior flagellum (PI. 2, figs A, B, C). Both
flagellar axenomes have the typical 9 + 2 arrangement of tubules ; the central pair
of tubules originates just above the peripherally thickened distal basal plates (PI. 2,
fig. B). No paraxial rods or mastigonemes were observed. Although scanning elec-
tron microscope and shadow-cast preparations show that the flagella taper distally, we
could not determine if this is due to the absence of tubules. Striated rootlets connect
the basal bodies, and several other rootlets run from the basal bodies to the sub-
pellicular region of the body (PI. 2, figs B, D, E). In the extracytoplasmic portion
of the basal bodies and proximal to the basal plate, the 9 peripheral doublets are
connected to the flagellar walls by strands (PL 3, fig. F) ; in the intracytoplasmic
portion the peripheral tubules are arranged in triplets.

The semicircular ridge, near the origins of the flagella (PI. 2, figs A, C), contains a
row of small organelles immediately below the plasma membrane (PI. 2, figs A, D ;
PI. 3, fig. A). Each of these organelles (about 170 nm long and 120 nm wide) has a
central conical element with an electron dense core and is surrounded by a unit
membrane. These organelles are present beneath the plasma membrane on other
parts of the body and the flagella (PI. 4, figs A, B, C) . Similar structures, kinetocysts,
have been described in Heliozoa (Bardele, 1972, 1975).

The nucleus is usually ovoid (about 1-7 /urn long, 1-3 /urn wide) and is enclosed by
a double unit membrane (PI. 3, figs B, E). The nuclear matrix is granular with
occasional concentrations of electron-dense chromatin attached to the inner nuclear
membrane and a dense central nucleolus (PI. 2, fig. B ; PI. 3, figs B, E). Cisternae
of the endoplasmic reticulum arise from the outer membrane of the nuclear envelope
and ramify in the cytoplasm (PI. 3, fig. E). Ovoid or spherical mitochondria, with
dense matrices and plate-like cristae, are distributed throughout the cytoplasm (PI. 3,
figs B, E, F). A single Golgi apparatus lies directly beneath the flagellar basal bodies
and consists of 6-8 saccules, which have both smooth and coated vesicles associated
with their margins (PI. 2, fig. B ; PI. 3, fig. E). The contractile vacuole is slightly
posterior to the nucleus (PI. 3, fig. C). Numerous food vacuoles are usually present
in the posterior half of the cytoplasm. If partly digested food is present, the single
unit membranes of these vacuoles are covered with small vesicles. An intermediate

MORPHOLOGY OF HETEROMITA GLOBOSA 261

stage in phagocytosis, with a bacterium partly engulfed by pseudopodia (PI. 3, fig. D),
was observed in thin sections of one flagellate.

In the cytoplasm adjacent to the posterior end of the nucleus, there is a microbody-
like organelle (160-260 nm wide). This organelle is surrounded by a unit membrane
and has a finely granular matrix with tubular elements (PI. 3, fig. B). Unstained
sections of cells incubated in DAB showed no reaction inside this organelle, although
there was a strong positive reaction in mitochondria.

In a few sections, a group of 50-60 cross-linked vesicles was observed close to the
plasma membrane in a posterior-lateral region of the trophozoite (PI. 3, fig. E) . Each
vesicle was about 20 nm in diameter. A similar array of small virus-like particles
was reported by Swale & Belcher (1973) in the nucleus of one specimen of the colour-
less flagellate, Aulacomonas submarina.

Cysts. The thick walls of mature cysts appear to be impermeable to acrolein,
glutaraldehyde and OsO4. All the observations on the ultrastructure of the resting
stages relate to the thin-walled cysts. During the first stages of encystment, the
body assumes a spherical shape and the flagella are resorbed into the cytoplasm
(PI. 6, fig. B). The flagella lie close to the plasma membrane inside the cell but are
seldom adjacent to each other (PI. 5, fig. A ; PI. 6, fig. B). Food vacuoles are still
present at this stage, although they are usually empty or only contain waste material
(PI. 6, fig. B). Vacuoles with single electron-dense spherical particles are present in
most cysts (PI. 5, fig. B ). The Golgi saccules are often swollen and appear to be
producing smooth and coated vesicles from their margins. Numerous small lacunae
are present in the peripheral cytoplasm of the cyst ; these structures are enclosed by
unit membranes and are either translucent or contain small central granules (PL 5,
figs A, B). Bardele (1972) found similar lacunae in Heliozoa and suggested they
were repositories for reserve plasma membrane, which can be utilized during the
rapid transformation of flagellate to heliozoon. Plasma-membrane material may be
stored in H. globosa in the same fashion and may become significant during excyst-
ment. Some of the lacunae with granular material in H. globosa may represent
degeneration stages of the kinetocyst-like organelles. Certainly, definite examples
of this organelle were not observed in cysts. The mitochondrial cristae in the cysts
are similar to those in trophozoites apart from occasional electron-dense particles
(PL 5, fig. A ; PL 6, fig. B). The nucleus in the cyst seems unaltered, but the adjac-
ent microbody-like organelle appears to be reduced. The contractile vacuole col-
lapses in the cyst, although the tubular spongiome usually remains visible. The
young cyst wall is a single filamentous layer (PL 6, fig. B) in close contact with the
cell contents. As the cysts mature, other layers are added to the wall and the points
of contact between the wall and cell contents become greatly reduced. Brooker &
Ogden (1972) described a similar type of encystment in Bodo caudatus with the cyst
wall consisting of several filamentous layers. The flagella in the cyst of B. caudatus,
however, lie in a groove of the cytoplasm close to the cyst wall.

Excystment starts with the emergence of one flagellum from the cytoplasm and
with further shrinkage of the cell contents from the cyst wall (PL 6, fig. C). The
invaginations gradually become more pronounced but fewer in number until there
are two deep grooves on opposite sides of the cell. Later, the second flagellum

262 C. M. MACDONALD, J. F. DARBYSHIRE & C. G. OGDEN

emerges from the cytoplasm, and both flagella become wrapped around the cell (PI. 6,
fig. A). The flagellate emerges from the cyst by extruding a pseudopodium through
a small pore (about 200 nm). Initially, the organelles remain within the cyst (PI. 6,
fig. D) and only later migrate to the pseudopodium (PI. 6, fig. D). Lipid droplets
persist in the cytoplasm for a short period after excystment (PI. 5, fig. C).

DISCUSSION

The present observations with the light microscope confirm most of Sandon's and
Robertson's accounts of the morphology and reproduction of Heteromita spp. No
perforations in some cyst walls (Sandon, 1927) or rhizoplasts (Robertson, 1928),
however, were observed. Although the dimensions of the trophozoites of the
Scottish clones in Table I are slightly smaller than those reported by Sandon for
H. globosa, the mean diameters of the resting stages lie within the same range
(4.2-5.0 /mi). Robertson (1928) identified two isolates of Heteromita as H. globosa
and it is clear from her description that she was studying a flagellate very similar
to the Scottish clones. We therefore suggest that the Scottish clones belong to
H. globosa.

Two organelles, which closely resemble microbodies and kinetocysts, were observed
by electron microscopy. These organelles have not previously been reported in
soil flagellates. The negative reaction of the microbody-like organelle to the DAB/
peroxide test suggests that this organelle is not analogous to the microbodies of
Hartmannella culbertsoni and Paramecium aurelia. A more likely analogy is with
those Phytomastigophorean microbodies, which also show a negative reaction for
catalase (Miiller, 1975). As the trophozoites of H. globosa have larger microbody-
like organelles than the resting stages, these organelles may be glyoxysomes with
the enzymes responsible for the conversion of lipids into carbohydrates. Certainly,
large lipid deposits were only observed in the resting forms and recently excysted
trophozoites. The positive reaction in the mitochondria with DAB may be caused
in Heteromita by cytochrome C oxidase. Bardele (1975) suggested that the kineto-
cysts of centrohelidians may be involved in the selection and transportation of
particles in the axopodia. In H . globosa, the flagella and pseudopodia are involved
in feeding, and the kinetocyst-like organelles just beneath the plasma membrane may
play an important role in the capture and ingestion of food. Bardele (1972, 1975)
believed that the kinetocysts originate from vesicles close to the dictyosomes and
later migrate to the plasma membrane. In H. globosa the dictyosome lies close to
the base of the flagella, and it may be significant that the majority of kinetocyst-like
organelles are found nearby.

The taxonomic position of Heteromita within the Zoomastigophorea is uncertain ;
the only suborder, which contains zooflagellates with 2 heterodynamic flagella in
modern classifications (Hollande, 1952 ; Honigberg et al., 1964), is the Bodonina of
the order Kinetoplastida. It is believed that some species in this suborder have lost
their kinetoplasts secondarily, and it is possible that such a loss has occurred in
Heteromita spp. Nuclear division in Heteromita, however, is unlike the promitotic
division of Bodonids (Robertson, 1927 ; Hollande, 1952). Robertson (1928) showed

MORPHOLOGY OF HETEROMITA GLOBOSA 263

that the blepharoplasts in Heteromita acted as centrosomes and that the karyosome
disintegrated during nuclear division. Recent studies of Bodo spp. (Pitelka, 1961 ;
Brooker, 1971 ; Burzell, 1975) and Rhynchomonas spp. (Burzell, 1973 ; Swale, 1973)
have also shown that the Bodonina have several characteristic ultrastructural
features, for example, a flagellar pocket, paraxial flagellar rods, pellicular micro-
tubules, a buccal cavity, a cytopharynx and a single coiled mitochondrion. As
many of these features were not observed in H. globosa, we consider Heteromita is a
valid genus and not a synonym of Bodo as proposed by Hollande (1952). Similar
investigations of other Bodo-like flagellates without kinetoplasts, such as Cyathobodo
(Swale & Belcher, 1975), should clarify the taxonomic affinities of Heteromita.

ACKNOWLEDGEMENTS

The time lapse photographs in PI. I are reproduced from a film sequence taken by
Mr H-H. Heunert during a short visit by J. F. D. to the Institut fur den Wissen-
schaftlichen Film (I.W.F.), Gottingen. The director of I.W.F., Professor G. Wolf,
is thanked for his permission to publish these photographs and several members of
the staff for their advice on cinephotomicrography, in particular Dr H. K. Galle,
Mr H-H. Heunert and Miss M. Milthaler. Mr J. Mitchell of the Macaulay Institute
provided the final prints for PL I.

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PITELKA, D. R. 1961. Observations on the kinetoplast-mitochondrion and the cytostome of

Bodo. Expl. Cell Res. 25 : 87-93.
REYNOLDS, E. S. 1963. The use of lead citrate at high pH as an electron-opaque stain in

electron microscopy. /. Cell. Biol. 17 : 208-212.
ROBERTSON, M. 1927. Notes on certain points in the cytology of Trypanosoma raiae and Bodo

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C. M. MACDONALD

Dr J. F. DARBYSHIRE

Department of Microbiology

THE MACAULAY INSTITUTE FOR SOIL RESEARCH

CRAIGIEBUCKLER

ABERDEEN ABg 2QJ

C. G. OGDEN

Department of Central Services

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7 5BD

PLATE i

Heteromita globosa. Phagocytosis of a bacterium. Nucleus and contractile vacuole are
visible, x 1270 phase contrast. Time lapse at i sec intervals.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6

PLATE i

K..SI

PLATE 2
H. globosa trophozoite

FIG. A. Section through the anterior end showing the basal body of the anterior flagellum,
the semicircular ridge with the kinetocyst-like organelles, the posterior flagellum and micro-
tubules near the kinetocyst-like organelles. x 38 ooo.

FIG. B. Section showing the position of the Golgi apparatus in relation to the nucleus and
flagella. Microtubules run from the base of the posterior flagellum into the cytoplasm, x 24 885.

FIG. C. Scanning electron micrograph showing the posterior flagellum continuous with the
semicircular ridge of cytoplasm and the origin of the anterior flagellum inside this ridge, x 3500.

FIG. D. Transverse section through the basal bodies, which are linked by striated filaments.
Microtubular rootlets run from the basal bodies to the sub-pellicular region, x 36 ooo.

FIG. E. Section showing microtubular rootlets near the base of the flagellum. x 39 300.

Bull. Br. Mus. mat. Hist. (Zool.) 31, 6

PLATE 2

PLATE 3
H. globosa trophozoite

FIG. A. Longitudinal section through the anterior end showing the Idnetocyst-like organelles
and the associated microtubules. x 37 100

FIG. B. Longitudinal section through the posterior end of the nucleus. An electron-dense
septum (arrow) lies across a pore in the nuclear membrane. Tubular elements can be seen in
the granular matrix of the microbody-like organelle. x 36 800.

FIG. C. Section showing the contractile vacuole lying on the ventral side of the nucleus.
Note food vacuoles, kinetocyst-like organelles, Golgi apparatus and microbody-like organelle.

XII 2OO.

FIG. D. Oblique section towards the posterior end showing pseudopodia around a bacterium,
x 21 ooo.

FIG. E. Longitudinal section showing the continuity between the outer layer of the nuclear
membrane and the endoplasmic reticulum (arrow). The multivesicular area is posterior to the
nucleus, the Golgi apparatus is anterior to the nucleus, x 17 625.

FIG. F. Longitudinal section showing the extracytoplasmic portion of the basal body
without the two central tubules. Note plate-like cristae in the mitochondrion, x 17 183.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6

PLATE 3

B

ifebfrrn *

cv

PLATE 4
H. globosa trophozoite

FIG. A. Kinetocyst-like organelles are present in the semicircular ridge of cytoplasm and
elsewhere on the body, below the plasma membrane, x 31 ooo.

FIGS B & C. Longitudinal sections showing kinetocyst-like organelles below the flagellar
membrane. Fig. B x 42 ooo, Fig. C x 58 ooo.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6

PLATE 4

PLATE 5
H. globosa

FIG. A. Transverse section of a resting stage. The mitochondria have electron dense
deposits. The flagella are seen in transverse section, x 43 260.

FIG. B. Transverse section of a more mature resting stage with a thicker cyst wall. Small
membrane-bound lacunae (arrows) lie around the periphery. Note lipid droplets in cytoplasm,
and in a vacuole. x 27 090.

FIG. C. An oblique section through the anterior end of a newly excysted trophozoite with
lipid droplets, x 14 100.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6

PLATE 5

B

PLATE 6
H. globosa resting stages

FIG. A. Transverse section through an excysting cell. The contents have shrunk from the
cyst wall and the flagella are lying around the body. x 18 682.

FIG. B. Transverse section of an early resting stage with intracytoplasmic flagella lying
around the periphery. Note one vacuole with food remnants and cyst wall with a single fila-
mentous layer (arrow), x 21 ooo.

FIG. C. Section showing initial stages of excystment. One flagellum has emerged from the
cytoplasm, x 26 670.

FIG. D. Section of an excysting cell with the cytoplasm extruding through a pore in the cyst
wall. The flagella are inside the cyst, x 14 128.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6

PLATE 6

NOTES ON THE MORPHOLOGY

AND NOMENCLATURE OF THREE MEMBERS

OF THE EUPLOTIDAE (PROTOZOA: CILIATEA)

By C. R. CURDS

INTRODUCTION

ALTHOUGH members of the genera Euplotes Ehrenberg, 1830 and Aspidisca Ehren-
berg, 1831 are commonly observed in marine and freshwater samples, they are often
difficult to identify to the species level. In recent years a considerable effort has
been devoted to the species problem in these two genera particularly in the case of
the genus Euplotes. Tuffrau (1960) introduced the application of the silver-line
system or argyrome as a taxonomic feature for the latter genus which made it
possible for the first time to determine the identities of some Euplotes species and
recently these ideas have been extended by Curds (1975). Tuffrau (1964) further
applied silver-impregnation methods as a means of identifying species of Aspidisca
but few have followed his lead.

The revisions of Borror (1972) and Curds (1975) concerning the order Hypotrichida
and genus Euplotes respectively contain lists of synonyms for species of Euplotes and
Aspidisca with which other authorities do not always agree. In several cases,
however, these disagreements are little more than matters of opinion based on
sparse and inadequate evidence. Nevertheless these works have provided a stimulus
to gather more data about these organisms in an attempt to ascertain their specific
identities.

The aim of the present paper is to provide further information on three previously
known members of the Euplotidae Ehrenberg, 1838. In particular, the nomen-
clature of Aspidisca cicada (Muller, 1786) Claparede & Lachmann, 1858 is discussed
in detail and its silver-line system described for the first time. A new form of
Euplotes aediculatus Pierson, 1943 is described and Euplotes daidaleos Diller &
Kounaris, 1966, a species hitherto unknown outside the United States of America,
is redescribed.

MATERIALS AND METHODS

SOURCE AND CULTIVATION. Samples of water, associated weed and debris were
collected from two different freshwater sources. Samples containing Aspidisca
cicada and Euplotes daidaleos were collected from an artificial pond located in the
grounds of the University of Copenhagen, Denmark, in September 1974 and these
were airmailed back to London in 5 ml aliquots. Samples of water containing
Euplotes aediculatus were collected from an indoor cold freshwater aquarium in the

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6 Issued 30 June, 1977

268 C. R. CURDS

British Museum (Natural History). In all cases the wild material was placed in
Petri dishes of sterile freshwater Erdschreiber solution (Medium i, Committee on
Cultures, Society of Protozoologists, 1958). Several hypotrichous ciliates grew in
these cultures which were then isolated, cloned and maintained in the same culture
medium. It was found that a thin layer of Musgrave & Clegg's agar (2-5% agar,
0-5% sodium chloride and 0-5% Liebig's beef extract in distilled water) covered
with Erdschreiber solution supported the growth of many more organisms than the
latter solution alone. This method was therefore routinely adopted when large
populations of organisms were required for staining procedures.

STAINING PROCEDURES. Silver-line preparations to display the dorsal and ventral
argyrome patterns were made following the 'dry' method of Klein (1958). This
method proved to be far more reliable and rapid than the conventional 'wet' method
(Corliss, 1953) or the protargol method (Tuffrau, 1967). Nuclei were stained using
Dippell & Chao's modification of DeLamater's basic fuchsin as described by Sonne-
born (1950). The nuclei of the cells were stained after fixation by air-drying rather
than using chemical fixation methods.

RESULTS

Aspidisca cicada (Miiller, 1786) Claparede & Lachmann, 1858

NOMENCLATURE. The nomenclature of the organism commonly, but erroneously,
referred to as Aspidisca costata (Dujardin, 1841) Stein, 1859 is a lengtny series of
synonyms, mixed identities and general taxonomic confusion. Since a complete
historical account has already been given by Brown (1968) only the major points
and recent developments will be mentioned here to support the case made by the
latter worker.

Brown (1968) correctly pointed out that Miiller (1786) was the first to describe
Trichoda cicada Miiller, 1786, an organism with several longitudinal ribs, and
presented figures that are clearly recognizable as the organism under consideration.
Bory (1826) later emended the generic name to Coccudina Bory, 1826 and specifically
refers to Miiller's (1786) descriptions in his own description of Coccudina cicada
(Miiller) Bory.

In 1831, Ehrenberg erected the genus Aspidisca Ehrenberg, 1831 and later the
same author described an organism with crenated or toothed longitudinal ribs which
he called Oxytricha cicada Ehrenberg, 1838. Dujardin (1841) also described an
organism with five or six tuberculated dorsal ribs which he recognized as being
identical to 0. cicada Ehrenberg, but considered that it should be included in the
genus Coccudina. Clearly Dujardin (1841) could not simply emend the generic
name of the species since the specific name was already preoccupied by C. cicada
(Miiller), a species with smooth longitudinal dorsal ribs. Dujardin (1841) therefore
named his species Coccudina costata Dujardin, 1841, having 0. cicada Ehrenberg as a
synonym. Thus at this point in time there were thought to be two distinct
organisms, one with smooth ribs - C. cicada (Miiller) and one with crenated ribs -
C. costata Dujardin.

EUPLOTIDAE 269

In 1858 Claparede & Lachmann transferred several species of the genera Coccudina
and Oxytricha to Ehrenberg's genus Aspidisca. Among the species they transferred
was Aspidisca cicada (Miiller, 1786) Claparede & Lachmann, 1858, an organism with
smooth dorsal ribs and furthermore concluded that 'it is not impossible that A.
cicada (Miiller) is identical to 0. cicada Ehrenberg'. Claparede & Lachmann (1858)
also briefly mentioned C. costata Du jar din and said that in their opinion it was
probably a member of the genus Aspidisca. The confusion really began with Stein
(1859) who was the first to use the combination Aspidisca costata (Dujardin) which he
considered to be identical to C. costata Dujardin even though the figures of this
author (Stein, 1859) did not indicate the presence of teeth nor tubercles along the
dorsal ridges. Nevertheless, he did not regard this species to be identical to 0.
cicada Ehrenberg. Since that time, authorities such as Kent (1880-1882) and Kahl
(1932) have referred to Stein's (1859) description so that organisms with smooth
dorsal ribs have been called Aspidisca costata (Dujardin) Stein instead of Aspidisca
cicada (Miiller) Claparede & Lachmann. The name A. costata should correctly only
be used in connection with an organism with crenated or tuberculated dorsal ribs
as was originally intended by Ehrenberg (1830) and Dujardin (1841).

Although Brown (1968) indicated most of these points in great detail, Borror
(1972) listed Aspidisca costata (Dujardin) as a major synonym and considered
Aspidisca cicada (Miiller) to be a minor synonym of Aspidisca lynceus (Miiller, 1773)
Ehrenberg, 1831 in his review of the hypotrichs. More recently Diller (1975) agreed
with the conclusion of Borror (1972) on the grounds that Miiller (1786) : 'Had
assigned it to a different genus, Trichoda' '. However, following the International
Code of Zoological Nomenclature (Stoll et al., 1964), Article 23 concerning the Law of
Priority states : The valid name of a taxon is the oldest available name applied to
it provided that the name is not invalidated by any provision of this code'. In this
particular case, Aspidisca cicada (Miiller) is the oldest available name for the taxon.
Since the generic name Aspidisca Ehrenberg did not exist in 1786, Diller's (1975)
comments are inappropriate.

It is suggested therefore that the evidence of Brown (1968) be accepted and the
organisms often erroneously referred to as Aspidisca costata (Dujardin) Stein be
correctly referred to as Aspidisca cicada (Miiller) Claparede & Lachmann. A list of
synonyms is given below :

Aspidisca cicada (Miiller, 1786) Claparede & Lachmann, 1858 : syn. Trichoda cicada Miiller,
1786 : Coccudina cicada Bory, 1826 : Coccudina costata Dujardin, 1841 : Coccudina cicada
Dujardin, 1841 : Aspidisca costata Stein, 1859 : Aspidiscopsis bengalensis Ghosh, 1921 :
Aspidisca sulcata Kahl, 1932 : Aspidisca costata f. tetracirrata Tucolesco, 1962.

The original drawings of these organisms are often poor by modern standards and
it would appear highly likely, particularly from Dujardin's (1841) figures, that A.
costata (Dujardin) is a junior synonym of A. cicada as is shown above, whereas
Borror (1972) attributed C. cicada Bory an organism with dorsal ridges as a junior
synonym of A. lynceus an organism without dorsal ridges.

In spite of the historical taxonomic problems of Aspidisca cicada this organism is
usually simple to identify since there are few freshwater species of the genus and

270

C. R. CURDS

A. cicada is the only one with six prominent longitudinal dorsal ridges. The number
of dorsal ridges in this species does vary, for example, Diller (1975) reported that
although his specimens usually had 6-7 ridges the number actually ranged from
3 to 10. From this information it seems likely that Aspidisca sulcata Kahl, 1932 is a
synonym of A. cicada since the difference lies only in the number of dorsal ridges.

DESCRIPTION. When one considers the genus as a whole the problems of identi-
fication and synonymy become acute. Such a situation is also found in the closely
related genus Euplotes where the problem has been greatly alleviated, but not solved,
by the application of silver-impregnation methods (Tuffrau, 1960 ; Curds, 1975).
The silver-line system or argyrome of some species of Aspidisca has been published
(Tuffrau, 1964). A. cicada has been the subject of several morphological papers
(Hamm, 1964 ; Brown, 1966 ; Diller, 1975) but none of these has included a descrip-
tion of the dorsal and ventral argyrome, even though Diller (1975) employed silver-
impregnation methods in his study. In Euplotes (Tuffrau, 1960 ; Curds, 1975), the
dorsal silver-line system has been of considerable value in identification and although
the detail of the argyrome, such as numbers of kineties, is not as stable as was
orginally hoped (Genermont et al., 1976) the gross geometry has yet to be shown to
vary within species of that genus. It is hoped that the gross patterns of the dorsal

FIG. i. External morphology and nuclear features of Aspidisca cicada, (a) ventral surface
showing cirri, adoral zone and nuclei ; (b) posterior aspect showing dorsal ridges ; (c) lateral
aspect.

EUPLOTIDAE

271

FIG. 2. Silver-line system of Aspidisca cicada, (a) ventral argyrome ; (b) dorsal argyrome.

silver-line system of Aspidisca will prove to be of similar diagnostic value. In any
event an extra character is always to be welcomed for identification purposes.

The external morphology of A . cicada has been described on several occasions and
the descriptions by Hamm (1964), Brown (1966) and Diller (1975) are the most com-
prehensive. Their accounts differ little from each other and the specimens examined
in the present study conform with their descriptions unless stated otherwise. The
diagrams of the external morphology and nuclear structure (Fig. i) are given so that
the descriptions of the argyrome can be linked to other details of the animal for
future reference.

The dorsal and ventral silver-line systems of Aspidisca cicada are shown in Fig. 2
and are very similar to those illustrated by Klein (1929) and von Gelei (1939) for
Aspidisca lynceus. However, a comparison between these two species shows that
A. lynceus has more kinetics and more dorsal bristles than A. cicada. The ventral
argyrome and the positions of the seven frontoventral cirri, the five transverse cirri
and the adoral zone of membranelles are easily seen (Fig. 2a). Although hundreds
of specimens were stained and examined, the anterior ventral ciliary structure
referred to by various names in the literature was never displayed by the dry silver
method used, but it could easily be seen beating in the anterior notch in living speci-
mens. Neither Klein (1929) nor Diller (1975) seemed to have had difficulties staining
this ciliary structure even though Klein used the same silver-impregnation method.

The dorsal argyrome of A. cicada (Fig. 2b) is a simple structure by comparison with
those in species of the genus Euplotes and is composed of a total of five dorsolateral
kinetics. There are two outer kinetics that encircle the perimeter of the animal to
join anteriorly, two kinetics that travel longitudinally either side of the longitudinal
axis of the body and a single short kinety on the right that occurs only in the
posterior third of the body. Silver impregnation clearly displays the positions of the
dorsal cilia or bristles and numbers of these were found to be constant in the clone

272 C. R. CURDS

examined. There were 3, 6, 6, 3 and 2 bristles respectively in the kinetics from left
to right. Brown (1966) stated that A. cicada had no dorsal bristles and indeed they
are extremely difficult to see in living specimens. Diller (1975) also disagreed with
Brown (1966) on this point and stated that he had seen indications of dorsal bristles
similar to those found in Euplotes. Other workers (Tuffrau, 1964 ; Deroux &
Tuffrau, 1965) have shown the presence of dorsal cilia in other species of Aspidisca.
Slides showing silver-line systems and nuclei have been deposited in the slide
collection of the BM(NH), reg. nos 1975:7:4:5-7.

Euplotes aediculatus Pierson, 1943

Euplotes aediculatus Pierson, 1943 has been the subject of several papers on
Euplotes taxonomy (Pierson, 1943 ; Tuffrau, 1960 ; Pierson et al., 1968 ; Carter,
1972) and a complete account of the confusion and misidentifications recorded in the
literature concerning this species has already been given by Curds (1975).

At least four species of Euplotes have been described that have varieties with
wing-like extensions to their lateral margins or extensions to their dorsal ridges.
Kahl (1932) stated that Euplotes patella (Miiller, 1773) Ehrenberg, 1838 has five
varieties and based four of these on the size and location of the dorsal ridges. Simi-
larly Kahl (1932) listed three varieties of Euplotes muscicola Kahl, 1932 on the basis
of dorsolateral ridging. Two species, Euplotes carinatus Stokes, 1885 and Euplotes
novemcarinata Wang, 1930, have been described as species mainly on the basis of
possessing large dorsolateral ridges. Kahl (1932) and subsequent workers (Borror,
1972 ; Curds, 1975) have tended to consider E. carinatus to be a synonym for E.
patella, but E. novemcarinata is still regarded as being a possibly valid species. Other
hypotrichs such as Aspidisca cicada have also been noted to have several different
heights in their dorsal ridges (Hamm, 1964).

An organism collected from an indoor unheated freshwater aquarium was noted to
possess large wing-like processes on the dorsal surface and left lateral border. On
the basis of these structures and general morphology the organism was suspected to
be a variety of Euplotes patella. An individual with ridge extensions was isolated
and the initial descriptions given here refer to that clonal culture.

The external morphology and nuclear structure of the organism mentioned above
is illustrated in Fig. 3. Two of the six dorsal ridges were extended, the central
ridge was large and there was a slightly smaller one on its right. In addition, the
left border of the animal projected out and curved ventrally (Fig. 3b, c). On the
ventral surface (Fig. 3a) there were nine frontoventral, five transverse and four
caudal cirri. The nuclear features were the first indication that this species was a
variety of Euplotes aediculatus rather than E. patella. The macronucleus of both
species is C-shaped but in the case of E. aediculatus there is much more of a distinct
flattening of the anterior portion.

Silver-line preparations (Fig. 4) showed that the organism was indeed Euplotes
aediculatus since the dorsal argyrome was of the double-eurystomus type (Curds,
1975) rather than the double-patella type. There were eight dorsolateral kinetics
(Fig. 4), but the collapse of the extended central ridge, during air-drying, made it

EUPLOTIDAE

273

FIG. 3. External morphology and nuclear features of Euplotes aediculatus. (a) ventral aspect
showing cirri and nuclei ; (b) transverse section through centre to show extended ridges
and lateral border ; (c) dorsal aspect showing extended lateral border.

difficult to count accurately the numbers of cilia in the central kinetics - however
they were estimated to be about 25. On the basis of the nuclear and argyrome fea-
tures this species was identified to be E. aediculatus. This appears to be the first
record of E. aediculatus having extensions to the ridges and furthermore it is the
first time that a variety such as this has been traced to its parent species by the use
of its silver-line system.

After a few months, examination of the clonal culture showed that some of the
cells had lost their ridge and lateral extensions. Unfortunately, soon after this
observation the clonal culture was lost due to a malfunction of the cooled incubator
in which it was kept. Samples taken from the same aquarium showed the presence
of a Euplotes species of the same size and general appearance but without projections

274 C. R. CURDS

FIG. 4. Dorsal argyrome of Euplotes aediculatus specimen with extended dorsal ridges.

and large ridges. Silver impregnation and nuclear stains of these organisms showed
that this was a 'normal' Euplotes aediculatus with a dorsal argyrome similar to that
shown in Fig. 4. Although it is possible that the first organism had been replaced by
a second it seems unlikely and in view of the observations on the original clone it
would appear that the 'abnormal' form had reverted to the 'normal' shape. Why
'abnormal' forms like this should be produced remains a mystery. There seems little
advantage in possessing large ridges and lateral borders and to the author's know-
ledge no theories have yet been published in the literature concerning this pheno-
menon.

Slides of both 'normal' and 'abnormal' forms of Euplotes aediculatus showing the
silver-line systems and nuclear features have been deposited in the slide collection
of the BM(NH), reg. nos 1975:7:4:3-4 ('abnormal') and 1975:11:19:1-2 ('normal').

Euplotes daidaleos Diller & Kounaris, 1966

Diller & Kounaris (1966) described the species Euplotes daidaleos which they
obtained from an artificial pond in Philadelphia, U.S.A. This organism contains
symbiotic zoochlorellae and appears to be the only record of a species of Euplotes
containing a green symbiont that has been identified using modern methods. There
are many other records of zoochlorellae-bearing Euplotes in the literature (Curds,
1975) but in all other cases these identifications are open to considerable doubt since
the host cells were not stained with silver.

The organism described in the present paper was isolated from an artificial pond
situated in the grounds of the University of Copenhagen, Denmark. On arrival in
our laboratory the organisms were maintained in a dark cooled incubator at 18 °C.
On the basis of its size and general shape it was provisionally thought to be Euplotes
patella. However, after staining the nucleus and silver-line system it soon became
apparent that this provisional identification was incorrect. The results of these

EUPLOTIDAE

275

FIG. 5. Euplotes daidaleos. (a) cell cultivated in dark ; (b) cell cultivated in light ; (c) nuclei

(d) dorsal argyrome.

276 C. R. CURDS

staining techniques are shown in Figs 50 and d. The dorsal argyrome was of the
double-patella type with nine dorsolateral kinetics containing 15-17 cilia in the mid-
dorsal rows. The macronucleus was found to be an open C-shape with the micro-
nucleus deeply embedded in a depression of the macronucleus. The ventral aspect
of the animal is shown in Fig. 5a which illustrates the positions of the nine fronto-
ventral, five transverse and four caudal cirri. Fig. 5a also shows the large numbers
of membranelles (about 60 in silver-line preparations) in the adoral zone. From
this information the organism was tentatively identified as E. daidaleos even though
zoochlorellae were not apparent. Closer examination of the cytoplasm revealed the
presence of some yellow to light green inclusions although they were too few in number
to impart any coloration to the animal as a whole. Some of the cultures were then
exposed to light which led to an immediate increase in the numbers of these cyto-
plasmic bodies and to the depth of colour. Within a week of constant exposure to
light the zoochlorellae reached sufficiently high numbers in the cytoplasm to impart
a bright green colour to the host cells. Figs 5a and b illustrate the numbers of
zoochlorellae in this species of Euplotes when cultured in the dark and light respect-
ively. The presence or absence of light did not have any obvious effects upon the
rate of growth, nor the peak population, of Euplotes supported in the cultures.

On the basis of the dorsal argyrome (Fig. 5d), numbers of membranelles in the
adoral zone, nuclear features and presence of symbionts this organism was identified
as Euplotes daidaleos. This is the first record outside America. The only difference
that could be visually detected between the Danish and American strains was the
precise shape of the macronucleus. In the case of the American strain (Diller &
Kounaris, 1966) the macronucleus was rather more angular than that described
here (Fig. 5c). The fact that an organism such as this can be easily recognized from
silver-impregnation preparations is encouraging particularly when the sources of
material are so remote from each other both in terms of geography and time.

Slides of the Danish strain of Euplotes daidaleos showing silver-line systems and
nuclear features have been deposited in the slide collection of the BM(NH), reg. nos
1975:7:4:1-2.

ACKNOWLEDGEMENT

The author is indebted to Dr Cicily Chapman-Andresen of the Department of
Zoology, University of Copenhagen, Denmark, for the supply of organisms during a
visit in September 1974.

REFERENCES

BORROR, A. C. 1972. Revision of the order Hypotrichida (Ciliophora, Protozoa). /. Proto-

zool. 19 : 1-23.
BORY, DE ST VINCENT, J. B. G. M. 1826. Essai d'une classification des animaux microscopiques.

104 pp. Paris.
BROWN, T. J. 1966. Observations on the morphology and reproduction of Aspidisca cicada

Muller. N.Z. Jl Sci. 9 : 65-76.
1968. A reconsideration of the nomenclature and taxonomy of Aspidisca costata (Dujardin,

1842) (Ciliata). Ada Protozool. 5 : 245-252.

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CARTER, H. P. 1972. Infraciliaturc of eleven species of the genus Euplotes. Trans. Am.

microsc. Soc. 91 : 466-492.
CLAPAREDE, E. & LACHMANN, J. 1858. Etudes sur les infusoires et les rhizopodes. M6m.

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DR C. R. CURDS

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7

NOTES ON THE ALPHEIDS IN WHITE'S
LIST OF THE SPECIMENS OF CRUSTACEA IN
THE COLLECTIONS OF THE BRITISH MUSEUM

(184?)

By ALBERT H. BANNER & DORA M. BANNER

IN 1847 the British Museum published anonymously a catalogue of the specimens of
crustaceans then found in the Museum's collections ; the authorship was that of
Adam White (see Miers, 1874 14, 5). The catalogue was a mere listing of generic
and specific names, without descriptions or figures, but giving locality and donor or
collector. Under the family name of Alpheidae he listed 16 species of the genus
Alpheus, one species of the genus Athanas as well as species under the generic names
of Alope, Atya and Nika ; these notes concern themselves only with the species
listed under Alpheus. In this genus, 10 of the 16 species were given new names,
but as they were without 'a description, definition or indication' they are nomina
nuda. In 1874 Miers published the section of 'Crustacea' in The Zoology of the
Voyage of the HM.S. Erebus & Terror . . . and used the lithographic plates prepared
many years before under the direction of White for the 7 species that he indicated
would be described in the publications of the voyage.* For these species Miers
added his own notes and at times a short description. Kingsley (1882) listed all
species mentioned by White and for some he also gave additional descriptions, but
often did not cite White's collection record. Bate (1888 : 543-544) also referred to
two of White's names and Miers' determinations, but it is difficult to determine what
he did except to confuse and reverse Miers' determinations.

During a recent visit to the British Museum (Natural History) we had made
available to us both the publications of White and Miers and the collections upon
which they were based. All of White's specimens were preserved as dried specimens
upon pins, similar to specimens in standard insect collections. During the course of
almost a century and a half many of the desiccated specimens had fallen apart, lost
their legs, fallen off their pins and were in general in a condition that would make
exact identification difficult or impossible. Moreover, in many instances the label
cards were also loose. However, in some cases exact identification was possible
because the specimens were surprisingly intact, and some specimens were identifiable
because of the possession of a unique characteristic such as the dactylus of the third
legs in Alpheus lottini Guerin, 1830.

* According to the narrative of the Voyage of the Erebus & Terror (vol. i) while the ships did visit
Tasmania and New Zealand, and 'a short visit to Sydney', they did not visit tropical Australian waters.
It is not known why White chose to include tropical Australian specimens given by various donors under
the reports of the voyage.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6 Issued 25 August, 1977

28o A. H. BANNER & D. M. BANNER

As the publication of White is rare and not available to most workers, we have
decided to publish Miers' (1874) and our own determinations of the species listed by
White, together with some additional notes. The quotations below are from White
in the order of his listing.

'Alpheus ventrosus, Edw. Crust, ii. 352.

a-g. Indian Ocean. From Mr. Gardiner's collection.'

Three specimens remaining in fair condition, but all lacking small chela. They are certainly
not A. ventrosus Milne-Edwards, 1837 (= A. lottini Guerin) which is in the Macrochirus Group.
They are in the Edwardsi Group, but the species cannot be identified without the small chelae.
Kingsley (1882) does not cite White's record.

'Alpheus Neptunus, n. s. Zoo/. Erebus and Terror, t. f.
a. Port Essington. Presented by the Earl of Derby.'

Specimen lacking both chelipeds. Miers (1874 : 4) reported this to be A. edwardsi (Audouin),
1827 and Kingsley (1882 : 120) accepts this synonymy. In the present fragmentary condition
of this specimen we cannot confirm or deny this identification. The specimen certainly has no
relationship to Alpheus (now Synalpheus) neptunus Dana, 1852, but as White had merely created,
a nomen nudum, Dana's use of the name may stand. Port Essington is in Northern Territories,
Australia.

'Alpheus chiragricus, Edw. Crust, ii. 354.
a, b. China. Presented by Gen. Th. Hardwicke.'

A pair of complete specimens. We find these are definitely not A . chiragricus Milne-Edwards,
1837, but A. hoplocheles Coutiere, 1897 who also reported the species from Japan, China and the
Indian Archipelago, but gave no specific locations.

'Alpheus Triton, n. s.

a. Indian Ocean. From Col. Whitehill's collection.'

Specimen complete except for small chela. This specimen is either A. pacificus Dana, 1852
or a closely related form. A. pacificus does occur in the Indian Ocean. The name is merely
listed as a manuscript name by Kingsley (1882 : 126).

'Alpheus Rhode, n. s.

a. Philippine Islands. From the collection of Mr. Cuming.'

Specimens complete except for small chela. Miers (1874 : 5) determined this to be A . strenuus
Dana, 1852 and the synonymy was accepted by Kingsley (1822 : 121). Without the small chela
we can neither confirm nor deny Miers' identification. A . strenuus does occur in the Philippines.

'Alpheus Amphitrite, n. s.

a. Philippine Islands. From the collection of Mr. Cuming.'

Specimen badly broken, lacking thoracic legs and small chela. In the remaining character-
istics, this specimen appears to be close to or identical with A. euphrosyne de Man, 1896 ; A.

ALPHEIDS IN WHITE'S LIST . . . 1847 281

euphrosyne has not been recorded from the Philippines, but it could be expected to occur in the
mangrove swamps of the archipelago. Kingsley (1882 : 125) listed this species as too imper-
fectly characterized to be recognized.

'Alpheus Doris, n. s. Zool. Erebus and Terror, t. f.
a, b. Torres Straits. Presented by J. B. Jukes, Esq.'

Specimen complete. This was determined by Miers to be A. strenuus Dana, 1852 which we
confirm. Kingsley (1882 : 121) accepted this synonymy. Bate (1888 : 544) lists this species as a
synonym of A. avarus Fabricius, 1798. This species is known from the Torres Straits and other
Australian waters.

'Alpheus heterochelis, Say, Journ. Acad. Sc. Phil. i. 243? Edw. Crust, ii.
356?

a, b. . . . From the collection of Dr. Leach.'
Specimens lost.

'Alpheus forceps, n. s.

a. Pondichery [sic]. From the collection of the French Museum.'

Specimen lacking only a few thoracic legs. Henderson (1893 : 434). when he revived
Fabricius' name A. malabaricus, stated 'In the British Museum is a specimen of our species from
Pondicherry bearing a MS. name "A. forceps", White.' Coutiere (1899 : 46, 49, 238) accepted
this synonymy. From the parts remaining, we also agree with Henderson. Kingsley
(1882 : 126) merely lists this as a manuscript name. A. malabaricus has a wide distribution in
brackish water in South and Southeastern Asia.

'Alpheus spinifrons, Edw. Crust, ii. 355.
a. Chili. Presented by the Rev. Mr. Hennah.'

Specimen almost complete, lacking only several thoracic legs. This specimen is not A.
spinifrons (now Synalpheus spinifrons) Milne-Edwards, 1837 that was described from the west
coast of South America. In spite of the distortion of the orbital teeth and small chela from
desiccation this specimen appears to be identical with a specimen in the spirit collection of the
British Museum bearing the label 'Portland Bay, West Patagonia' (= Chile) taken by the Alert
in 1870. This specimen was left unnamed by Miers (1881 : 74) but subsequently identified by
Coutiere (1899 : 34) and Holthuis (1952 : 48) as A. dentipes Guerin, 1832. As Holthuis (1952 : 48)
has remarked, it is improbable that A . dentipes, known from the Mediterranean and Cape Verde
Islands, would be found in the cold waters of Chile. Holthuis suggested that the locality label
of the specimen in spirit might be in error. However, this dried specimen was 'presented by
Rev. Mr. Hennah' sometime before 1847 also is clearly labelled as coming from Chile. It is
almost too much to expect that both labels, written over 30 years apart, were equally in error.
We suggest that a more careful study of new specimens from Chile is warranted.

'Alpheus Alope, n. s. Zool. Erebus and Tenor, t. f.

a. N. Holland (Port Stephen). Presented by the Earl of Derby.'

Specimen lacking large and small chelae and most thoracic legs. Miers (1874 : 5) stated that
the specimen was 'in too bad a condition to be distinctively characterized.' In spite of the

282 A. H. BANNER & D. M. BANNER

figure (1874 : Tab. 4, fig. 6) we agree. Kingsley (1882 : 126) states that this is one of those
species whose 'descriptions are in works at present inaccessible to me'.

'Alpheus Thetis, n. s. Zool. Erebus and Terror, t. f.

a. N. Holland. Presented by the Entomological Club of London.'

Specimen lacking only large and small chela. Miers (1874 : 5) states that the specimens 'are
in too imperfect a condition to be well distinguished . . .' but that 'A thetis resembles . . . A.
laevis Randall . . .'. We found that the dactyli of the thoracic legs are of the unique develop-
ment of A. lottini (previously A. ventrosus Milne- Edwards and A. laevis Randall), and other
distinguishable characteristics remaining on the specimen are also in agreement with this species.
A . lottini is well known from the coasts of Australia (Banner & Banner, in press) . The picture
is further complicated by Kingsley (1882 : no) who stated that he had examined two speci-
mens from New Zealand 'in the museum of the Academy of Natural Sciences of Philadelphia,
presented by Dr. T. B. Wilson and labelled in Adam White's handwriting'. He then proceeds
to give additional notes on some species that plainly is not A . lottini. For example, the palm
of the large chela is sculptured and the first carpal article of the second leg is as long as the two
following. We have been unable to find these specimens. From the description we cannot
determine which New Zealand species that Kingsley is describing, but the matter is of no
importance as the specimens are not the same as White's in the British Museum (Natural
History).

'Alpheus Galathea, n. s. Zool. Erebus and Terror, t. f.
a. Port Essington. From the collection of Mr. Gould.'

Specimen completely lacking all thoracic appendages except the maxillipeds. Miers (1874 : 5)
stated that this specimen also was too broken for identification and in spite of the illustration
(1874 : Tab. 4, fig. 4) we agree. Kingsley (1882 : 126) lists this species as one where the descrip-
tions are not available to him.

'Alpheus Doto, n. s. Zool. Erebus and Terror, t. f.

a -e. N. Holland, Sir C. Hardy's Island ; dredged in n fathoms. Presented by
J. B. Jukes, Esq.'

Specimens presently lacking large and small chelipeds, but these were figured, albeit poorly, in
Miers (1874 : Tab. 4, fig. 5). Miers (1874 : 5) gave a brief description of this species, so the name
would be acceptable with Miers as the author. Coutiere (1899 : 25) stated that 'A . doto White . . .
est sans doubte synonyme de A . socialis Heller . . .'. We agree with Coutiere on the basis of the
unique lobe on the inferior margin of the second article of the third maxillipeds, the configuration
of the rostral front, a long tooth on the basicerite, etc. We will discuss these characteristics,
and the range of this and closely related new species in Part III of our Australian study. How-
ever, there are additional complications for Sir C. Hardy's Island is in the far northern portion
of the Great Barrier Reef and none of the specimens of A . socialis in the Australian collections
that we have examined were from tropical Australian waters.

'Alpheus minus, Say, Journ. Acad. Sc. Phil. i. 245.
a-c. East Florida. Presented by Thomas Say, Esq.'

Three specimens, all lacking antennules and antennae. The anterior regions and the telsons
are typical of Synalpheus minus (Say), 1817. Since these specimens were given to the Museum
by Say and identified by Say, we feel it is undoubtedly his species. Kingsley (1882 : 114) does
not cite White's record.

ALPHEIDS IN WHITE'S LIST . . . 1847 283

'Alpheusfrontalis, Edw. Crust, ii. 356. Cuv. R. A. (Crock), t. 53, f. 2.
a. Torres Straits. Presented by J. B. Jukes, Esq.'

Specimen lacking only small chela. This specimen is unmistakably A. frontalis Milne-
Edwards, 1837 and the species is well known from the Torres Straits and other tropical Australian
waters (Banner & Banner, in press). Kingsley (1882 : 123) does not cite White's record.

In conclusion, none of the new names proposed by White are valid on the basis of
White's publication and those two subsequently described by Miers are junior
synonyms ; of the six species of which White applied previously published and valid
names, four were misidentified.

ACKNOWLEDGEMENTS

We wish to acknowledge the cooperation of the staff of the British Museum
(Natural History), especially the aid of Dr Anthony Fincham and Mr Paul Clark.
This work was supported in part by United States National Science Foundation
Grant BMS74-n844.

REFERENCES

ANONYMOUS [White, A.] 1847. List of specimens of Crustacea in the collection of the British

Museum, pp. i-viii, 1-143. London.
BANNER, D. M. & BANNER, A. H. In press. The Alpheid shrimp of Australia. Part III.

The genus Alpheus, remaining genera and the family Ogyrididae. Rec. Aust. Mus.
BATE, C. SPENCE, 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger

during the years 1873-76. Rep. Voy. Challenger Zool. 24 (i) : 1-942. [150 pis separate

in 24 (2).] London.
COUTIERE, H. 1897. Note sur quelques especes du genre Alpheus du Musee de Leyde. Notes

Leyden Mus. 19 (23) : 195-207.
1899. Les 'Alpheidae' morphologie externe et interne, formes larvaires, bionomie. Annls

Sci. nat., Zool. 9 : 1-560.
DANA, J. D. 1852. Crustacea. In United States Exploring Expedition during the years 1838,

1839, 1840, 1841, 1842 under the command of Charles Wilkes, U.S.N. 13 : pp. 1620. C.

Sherman, Philadelphia.

FABRICIUS, J. C. 1798. Supplementum entomologiae systematicae. 572 pp. Hafniae.
HELLER, C. 1865. Crustacea. In Reise der osterreichischen Fregatte Novara um die Erde in den

Jahren 1857, 1858, 1859 unter den Befehlen des Commondore B. von Wullerstorf-Urbair.

Zool. 2 (3) : 1-280. Wien.
HENDERSON, J. R. 1893. A contribution to Indian carcinology. Trans. Linn. Soc. Land.

II, 5 (10) : 325-458.
HOLTHUIS, L. 1952. The Crustacea Decapoda Macrura of Chile. Reports of the Lund

University Chile Expedition 1948-49. K. fysiogr. Sdllsk. Lund Fork. 62 (10) : 1-109.
KINGSLEY, J. S. 1882. Carcinological notes : Number V. Bull. Essex Inst., Salem, Mass.

14 : 105-132.
MIERS, E. J. 1874. Crustacea. In S. Richardson & S. E. Gray, eds, The Zoology of the

Voyage of the H.M.S. Erebus and Terror, under the command of Captain Sir James Clark

Ross, R.N., F.R.S., during the years 1839 to 1843 . . . 2 : 1-5, pis 1-4. London.
1881. Crustacea. In Account of the Crustacea collected during the survey of H.M.S. 'Alert'

in the Straits of Magellan and on the Coast of Patagonia. Proc. zool. Soc. Land. 1881 : 61-79.

284 A. H. BANNER & D. M. BANNER

MIERS, E. J. 1884. Crustacea. In Report of the zoological collections made in the Indo-Pacific
Oceans during the voyage of H.M.S. 'Alert'. 1881-2, pp. 178-322, 513-575, pis 18-35,
46-52. London.

MILNE-EDWARDS, H. 1837. Histoire naturelle des Crustaces, comprenant I'anatomie, la physio-
logic et la classification de ces animaux 2 : 1-532, atlas pp. 1-32, pis 1-42. Paris.

ALBERT H. BANNER, PH.D.

DORA M. BANNER, B.A.

Hawaii Institute of Marine Biology

UNIVERSITY OF HAWAII

P.O. Box 1364

KANEOHE

HAWAII 96744

USA

ESTABLISHMENT OF A NEW GENUS IN

THE FAMILY PHOXOCEPHALIDAE

(CRUSTACEA: AMPHIPODA) AND

A DESCRIPTION OF A NEW SPECIES FROM

NORTH ISLAND, NEW ZEALAND

By ANTHONY A. FINCHAM

SYNOPSIS

A new genus, Waitangi, of phoxocephalid amphipod is diagnosed with Paraphoxus rakiura
Cooper & Fincham, 1974 as type-species. Waitangi brevirostris n. sp. is described ; the new
material was collected from sandy beaches in North Island, New Zealand.

INTRODUCTION

WITH THE notable exception of the Talitridae (Hurley, 1956, 1957) few of the inter-
tidal sand-dwelling species of amphipods have been described from beaches in New
Zealand. Hurley (1954) listed five genera and six species of Phoxocephalidae
recorded from New Zealand. More recently algae-dwelling species have been
collected and figured by Barnard (1972), and Cooper (1974) has described infaunal
species from Wellington Harbour, some of which also live intertidally (Fincham
1974). Four new species of intertidal, infaunal amphipods have been described
recently (Cooper & Fincham 1974) from Stewart Island and from beaches in the
Auckland area ; the present paper adds a further species belonging to the Phoxo-
cephalidae.

Fourteen beaches were visited in North Island from March 1972 to November
1973. Collections were made as part of a general ecological survey of Crustacea
Peracarida of the intertidal sand biotope of New Zealand (Fincham 1974 ; in press).
Beach sediments examined ranged from black ironsand at Onaero on the west coast
to the pale pumice sand at Mt Maunganui. The new phoxocephalid was found in
conditions varying from very exposed (Baylys Beach) to partially protected (Para-
paraumu Beach).

Abbreviations used throughout : MLWN - mean low water neaps, LMB - low
mid beach, HMB - high mid beach. Where measurements are given, articles were
measured along the midline (length) and at right angles to the midline at the widest
part (breadth). Total length of the amphipod was measured from the tip of the
rostrum to the tip of the telson after straightening the body.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6 Issued 30 June, 1977

286 A. A. FINCHAM

DESCRIPTION OF GENUS AND SPECIES

FAMILY PHOXOCEPHALIDAE
Waitangi n. g.

DIAGNOSIS. Rostrum abruptly narrowed or absent ; article 2 of gnathopods
narrowing distally ; article 2 of pereiopod 7 with ventral and lateral setae ; molar of
mandible semitriturative, bearing four spines ; rami of uropods i, 2 with apical
spines (not a nail).

TYPE-SPECIES. Paraphoxus rakiura Cooper & Fincham, 1974 : 168-73, figs 7a,
8-10.

ETYMOLOGY. Waitangi is derived from the place name of the Treaty of 1840 at
which Maori chiefs ceded land to Queen Victoria.

REMARKS. Bearing in mind the combination of the four phoxocephalid genera
Pontharpinia Stebbing, 1897, Parharpinia Stebbing, 1899, Protophoxus K. H.
Barnard, 1930 and Trichophoxus K. H. Barnard, 1930 as subjective synonyms of
Paraphoxus Sars, 1895 by J. L. Barnard (1960) a new species of New Zealand
phoxocephalid was designated Paraphoxus rakiura by Cooper & Fincham (1974).
In a recent paper, however, by Barnard & Drummond (1976) these four genera,
together with Metharpinia Schellenberg, 1931, were reinstated following an examin-
ation of Australian phoxocephalid material. In the same paper Barnard & Drum-
mond suggest that the distinctive New Zealand form Paraphoxus rakiura Cooper &
Fincham should be established in a new genus '. . . to differ from Trichophoxus in
the presence of setae on the peduncle of uropod i, the pattern of the various orna-
ments on uropods 1-2, and the giant tooth of epimeron 3 . . .'. In addition the
extreme abbreviation of the rostrum, shape of gnathopod article 6, setation of
article 2 pereiopod 7, together with the pattern of spines on uropods i, 2 have been
used in the generic diagnosis. However, the shape of the third epimeral plate has
not been included because of the interspecific variability of this character, not
uncommon in amphipods, that is demonstrated clearly by the new species described
below.

Waitangi brevirostris n. sp.

(Figs 1-3)

DIAGNOSIS. Rostrum absent ; gnathopod article 6 broad, shorter than article 5,
narrowing distally for about half its length to a weakly chelate palm ; pereiopod 7
with article 2 reaching just beyond distal margin of article 5, with a well-defined field
of long setae medially and scattered short setae ; uropods i, 2 with spines along the
length of the peduncle ; epimeral plate 3 truncated with small notch at junction
with somite.

DESCRIPTION. Length 5-5-7-7 mm (female), body fusiform, broad, dorsally
flattened, laterally expanded (Fig. i) ; cream in colour.

Coxal plates 1-5 rather large ; coxal plate i (Fig. 2n) produced slightly forward,
group of short setae at antero-distal corner, longer setae at postero-distal corner ;

NEW GENUS OF PHOXOCEPHALIDAE

287

FIG. i. Waitangi brevirostris n. sp. adult female. Bar scale = i-o mm.

marginal setules between these two groups ; plates 2, 3 (Figs 2q, s) with group of
setae at postero-distal corner, 2-3 setules on anterior distal edge ; plate 4 (Fig. 30)
group of setae at anterior distal corner, smaller setae along ventral margin ; plates
5-7 (Figs 3C, d, e) small, lobed, with marginal setules.

Epimera plate i (Fig. 4d) group of fine plumose setae on anterior distal margin,
simple setae in postero-distal corner and postero-laterally ; plate 2 (Fig. 46) with
simple setae in postero-distal corner, distal margin and postero-laterally ; plate 3
(Fig. 4f ) truncated in postero-distal corner, small notch on posterior edge at junction
with somite, marginal postero-ventral setae, group of setae dorsal and ventral to
notch.

Head (Fig. i) wide, rostrum absent, lateral lobes moderately produced ; eyes
moderate in size, black visual elements bleaching somewhat in spirit.

Antennae (Figs 2b, c) short, less than % length of body, antenna 2 a little longer
than i ; antenna i peduncle article i large, ventral distal margin armed with plumose
seta and group of small spines, articles 2, 3 with arrays of moderate to long setae ;
flagellum and accessory flagellum sub-equal in length, each of 12 segments, 8 distal
segments bearing small setae ; antenna 2 with peduncle article 2 robust heavily
setose and spinose, flagellum shorter than peduncle, with 14 segments, moderate in
length, all bearing short setae.

Upper lip (Fig. 2e) entire, margin rounded.

Mandible bulbous, molar bearing four spines (Fig. 2f, g), palp of three articles,
terminal article with group of distal setae.

Maxilla i (Fig. 2h) inner plate bearing three sparsely plumose setae distally,
outer plate with a row of spines distally, palp biarticulate, article 2 with an apical
group of setae.

288

A. A. FINCHAM

FIG. 2. Waitangi brevirostris n. sp. adult female, (a) dorsal view head ; (b) antenna i ;
(c) antenna 2 ; (d) epistome (sketch) ; (e) upper lip ; (f) mandible ; (g) mandibular
molar ; (h) maxilla i ; (i) maxilla 2 ; (j) maxilliped ; (k) maxilliped inner plate ; (1) gnatho-
pod i ; (m) gnathopod i palm ; (n) coxal plate i ; (o) gnathopod 2 ; (p) gnatho-
pod 2 palm ; (q) coxal plate 2 ; (r) pereiopod 3 ; (s) coxal plate 3. Bar scale : a = 0-5 mm,
b, c, f, j, 1, o, q, r, s = 0-4 mm, e, h, i = 0-2 mm, g, k, m, p = o'i mm.

NEW GENUS OF PHOXOCEPHALIDAE 289

Maxilla 2 (Fig. 21) inner and outer plates rounded distally with groups of sparsely
plumose setae distally.

Maxilliped (Fig. 2J, k) inner plate with plumose setae and two stout, distal spines ;
outer plate normal ; palp article 4 simple.

Gnathopods I, 2 (Figs 2!, m, o, p, Table i) densely setose, dactyl of gnathopod 2
with setae facing palm.

TABLE i
Ratio of length to breadth of articles 5 and 6 in gnathopods i and 2

Article 5 6

L : B L : B

Gnathopod i 50 : 25 43 : 18

2 49 : 28 42 : 18

TABLE 2

Ratio of length to breadth of articles 2-6 in pereiopods 5-7

Article

2

L:B

3
L:B

4
L:B

5
L:B

6
L:B

Pereiopod 5
6
7

103 : 72
104 : 96
92 :68

21 : 20
14:32
21 : 22

59:74
68:95
12 : 16

72:58
66:47
33: 14

66 : 14
67: 16
33: 5

Pereiopods 5, 6 (Figs 3c, d, Table 2) heavily spinose, article 2 with row of spines
medio-laterally.

Pereiopod 7 (Fig. 36, Table 2) miniaturized.

Uropods 1-3 (Fig. 4a-c) peduncle of uropod i with row of stout setae, inner ramus
with lateral and apical spines, outer ramus lateral spines only ; peduncle of uropod 2
with stout setae, rami short with stout apical setae ; peduncle of uropod 3 with
group of stout distal setae, rami long, sub-equal in length, outer ramus two-seg-
mented, with plumose setae and a row of stout setae on outer edge.

Telson (Fig. 4g) cleft to base, each lobe with 6 stout apical setae.

Male. Length 5-4-7-1 mm. Sexual dimorphism typical.

MATERIAL EXAMINED.

Holotype, adult female, length 7-7 mm, carrying 7 eggs, collected at MLWN from
intertidal sand on 13 April 1973, Baylys Beach, North Island, New Zealand.
BM(NH) reg. no. 1976:166:1.

Paratypes, adults and juveniles collected at MLWN from intertidal sand on 13
April 1973, Baylys Beach, North Island, New Zealand. BM(NH) reg. nos. 1976:167:2
(adult females dissected, mounted in polyvinyl lactophenol with lignin pink on
slides) ; 1976:168:10 ; adults and juveniles collected at HMB from intertidal sand
on 19 November 1973, Paraparaumu Beach, North Island, New Zealand. BM(NH)
reg. no. 1976:169:5.

290

A. A. FINCHAM

FIG. 3. Waitangi brevirostris n. sp. adult female, (a) pereiopod 4 ; (b) coxal plate 4 ;
(c) pereiopod 5 and coxal plate 5 ; (d) pereiopod 6 and coxal plate 6 ; (e) pereiopod 7
and coxal plate 7. Bar scale = 0*4 mm.

NEW GENUS OF PHOXOCEPH ALIDAE

291

Other material from intertidal sand HMB and MLWN Mt Maunganui, LMB Ohope
Beach, North Island, New Zealand.

ETYMOLOGY. The species is named brevirostris in view of the extreme fore-
shortening of the rostrum.

REMARKS. In his paper on the Phoxocephalidae, Barnard (1960) describes
evolutionary trends shown by the known genera and in diagrammatic form demon-
strates progression from the 'basic' phoxocephalid form with broad rostral hood,
narrow pereiopod 3 and obtuse palm of gnathopod I to the 'trichophoxid' form with
rostrum narrowing abruptly, broad pereiopod 3 and right-angled palm of gnathopod
i. In the present new species these trends can be seen to have developed further ;
the rostrum has virtually disappeared, pereiopod 3 is very broad and the palmar
angle acute (the latter characters foreshadowed in the other, closely related New

I 1

FIG. 4. Waitangi brevirostris n. sp. adult female, (a-c) epimeral plates 1-3 ; (d) uropod
i ; (e) uropod 2 ; (f) uropod 3 ; (g) telson. Bar scale = 0-4 mm.

292 A. A. FINCHAM

Zealand genus Trichophoxus) . At first glance the present new species is much more
'haustoriid-like' than phoxocephalid. Barnard (1960) suggests that the genus
Paraphoxus (now divided into several genera) offers the zoologist an opportunity
to study '. . . evolution and the correspondence of morphology to specific environ-
ments'. It would appear that haustoriids dominate intertidal sand in the northern
hemisphere but are replaced by phoxocephalids occupying the equivalent niche in
the southern hemisphere. In both, however, the families co-exist.

On the basis of preliminary diagnoses, two further species of New Zealand phoxo-
cephalids Paraphoxus spinibasus Cooper, 1974 and P. chelatus Cooper, 1974 should
be transferred to different genera, tentatively Trichophoxus and Waitangi n. g.
respectively but this can only be confirmed on publication of figures supporting the
diagnoses. In common with Paraphoxus chelatus Cooper and Waitangi rakiura
(Cooper & Fincham), W. brevirostris n. sp. has the distinctive narrowing of article 6
of the gnathopods, together with setae on article 2 of pereiopod 7 and the develop-
ment of apical spines on the rami of uropods i, 2. The closely related genus Tri-
chophoxus has a terminal nail instead of these apical spines. Waitangi brevirostris
n. sp. differs from the one other known species (and one surmised - P. chelatus} in
the extreme abbreviation of the rostrum.

REFERENCES

BARNARD, J. L. 1960. The amphipod family Phoxocephalidae in the eastern Pacific Ocean,

with analyses of other species and notes for a revision of the family. Allan Hancock Pacif.

Exped. 18 (3) : 175-368.
1972. The marine fauna of New Zealand : algae-living littoral Gammaridea (Crustacea :

Amphipoda). Mem. N.Z. oceanogr. Inst. No. 62 : 1-216.
& DRUMMOND, M. M. 1976. Clarification of five genera of Phoxocephalidae (marine

Amphipoda). Proc. biol. Soc. Wash. 88 (49) : 515-548.
COOPER, R. D. 1974. Preliminary diagnoses of three amphipod species from Wellington

Harbour (Note). N.Z. Jl mar. freshw. Res. 8 (i) : 239-241.
& FINCHAM, A. A. 1974. New species of Haustoriidae, Phoxocephalidae, and Oedicero-

tidae (Crustacea : Amphipoda) from northern and southern New Zealand. Rec. Dom. Mus.

Wellington 8 (12) : 159-179.
FINCHAM, A. A. 1974. Intertidal sand-dwelling peracarid fauna of Stewart Island. N.Z.

Jl mar. freshw. Res. 8 (i) : 1—14.
In press. Intertidal sand-dwelling peracarid fauna of North Island. N.Z. Jl mar. freshw.

Res.
HURLEY, D. E. 1954. Studies on the New Zealand amphipodan fauna. No. 3. The Family

Phoxocephalidae. Trans. R. Soc. N.Z. 81 (4) : 579~599-
1956. Studies on the New Zealand amphipodan fauna. No. 13. Sandhoppers of the

genus Talorchestia. Trans. R. Soc. N.Z. 84 (2) : 359-389.
1957- Terrestrial and littoral amphipods of the genus Orchestia, family Talitridae. Trans.

R. Soc. N.Z. 85 (i) : 149-199.

DR A. A. FINCHAM

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7 5BD

A NEW SPECIES OF THE SYNAPTID GENUS

RYNKATORPA ROWE & PAWSON, 1967

(HOLOTHURIOIDEA : APODIDA) FROM THE

GREAT BARRIER REEF

By F. W. E. ROWE

INTRODUCTION

THE specimens described in this paper form part of a collection of echinoderms from
off northern Queensland (i5°43-2' S, 14^27-4 E), in the vicinity of Lizard Island,
Great Barrier Reef, collected by Dr P. E. Gibbs during the Royal Society of London/
Universities of Queensland Expedition to the Great Barrier Reef in 1973. The
majority of echinoderm material has been identified and described by Gibbs,
Clark & Clark (1976). However, three pieces of a synaptid holothurian were sent
to me. These proved to represent a new species of the genus Rynkatorpa Rowe and
Pawson.

DESCRIPTION OF NEW SPECIES

Rynkatorpa gibbsi sp. nov.

MATERIAL. Station 033, i5°43-2' S, i45°27'4' E, 16 m, sandy silt with few
foraminifera, collected by Dr P. E. Gibbs on 31 August 1973. Holotype (anterior
and posterior ends separated), British Museum (Natural History) no. 1974.12.3.60.
Paratype (posterior end), the Australian Museum no. J9743-

DESCRIPTION. The two ends of the holotype and the posterior end of the paratype
each measures 10 mm in length, with a diameter between 1-5 and 2*5 mm. The
anterior end shows the animal to have 12 tentacles, each with a distal median knob
and two pairs of lateral digits more or less equal in size. The most ventral tentacles
are the smallest, the others increasing gradually in size dorsally. The tentacles
bear 3-5 sensory cups in each of two vertical rows on the oral surface. There are
12 eye-spots, each at the base of and between adjacent tentacles. The calcareous
ring has the usual form of the genus. The gonad is present, small and about 2 mm
long. It is a dichotomous, antler-like structure with 3-4 lateral projections from
each of the two branches (Fig. i) and a single, median gonoduct. There is a single,
ventral polian vesicle and the ciliated funnels occur along the mid-line of the left
dorsal-lateral interradius. The stone canal is minute, single and sigmoid in form.

The calcareous spicules of the tentacles comprise C- or S-shaped rods (25-40 ^m x
3-4 /mi) and dumb-bell-shaped miliary granules (21-30 /mix 9-12 /mi) (Fig.2).

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6 Issued 30 June, 1977

2Q4

F. W. E. ROWE

FIG. i. Gonad, scale = 2 mm. FIG. 2. Rods and miliary granules from tentacle.
FIG. 3. Miliary granules from body wall. FIG. 4. Anterior anchor. FIG. 5. Posterior
anchor. FIG. 6. Anterior anchor plate (similar plates occur posteriorly). FIG. 7.
Larger posterior anchor plate. Scale = 100 /um for figs 2-7.

In the body wall dumb-bell-shaped miliary granules (21-30 /u,m x 9-12/^1) are
present in the anterior portion but absent from the posterior portions of the body

(Fig. 3).

The anchors are of one size only in the anterior portion of the body (125-135 /u,m
long x 105 p,m across the arm tips) but anchors in the posterior region are of two
distinct sizes (150 /nm long x 112 /x,m across the arm tips and 225-285 /urn long x 112-
135 /Lcm across the arm tips). The stock of the anchors is unbranched and the arms
bear 2-3 teeth on the small anchors and 8-n on the larger anchors (Figs 4 and 5).
The associated anchor plates correspond in size with the anchors. The anterior
plates measure 127-150 /urn in length x 90-112 /u,m in width and the posterior plates
measure 135-143 /u,m x 90-120 /urn and 180-225 /u,m x 105-127-5 /*m. The plates
are distinctive. The anterior and smaller posterior plates are more or less regularly
pear-shaped, with the pair of larger central perforations similarly pear-shaped but
aligned at 90° to the long axis of the plate and with the narrow end directed towards
the outer edge of the plate. Anterior to these two perforations are three smaller
ones in a horizontal line. The central one of these is usually the largest. The rest
of the anterior part of the plate is perforated by 10-12 small holes. Posterior to the
central perforations there are many small holes, but the posterior region is not
produced in a 'handle'. A weakly formed bridge is present (Fig. 6). The larger

A NEW SPECIES OF RYNKATORPA 295

posterior plates are elongate, more or less rectangular and with the central pair of
holes very much larger than the other perforations in the plate. There is a suggestion
of a posterior 'handle' and a weak bridge is developed (Fig. 7).

DISCUSSION

The genus Rynkatorpa was established by Rowe & Pawson (1967) to accommodate
a new species (R. hickmani} together with seven species that Clark (1907) included in
the genus Protankyra Ostergren, 1898. Since that time one new species (R. pawsoni)
has been described by Martin (1969) from four specimens which had been attached
to the posterior end of a fish host. Spicules were not found in these specimens, so
there must remain some doubt whether R. pawsoni should be referred to Rynkatorpa,
at least until further material is collected and the presence or absence of spicules is
firmly established. Martin comments that the spicules may have been lost during
destaining procedures in the preparation of mounted serial sections of the animals.
'R'. pawsoni is the first reported ectocommensal holothurian.

'R. pawsoni' is distinguished from R. gibbsi, as is R. bicornis (Sluiter) and R.
duodactyla (H. L. Clark), by the possession of only one pair of digits on the tentacles.
R. gibbsi is most readily distinguished from the other species in the genus by the pear-
shaped smaller anchor plates and the arrangement of the perforations in them.
Additionally, R. gibbsi differs from R. sluiteri (Fisher) in the smaller size of the
anchors which do not have a branched stock ; from R. timida (Koehler) by having
a smooth and not a spinous margin to the perforations of the anchor plates ; from
R. challenge* i (Theel) in having smaller anchors and plates and only a single instead
of two polian vesicles ; from R. bisperforata (H. L. Clark) in the form of the plates
with a weak but definite posterior bridge and only one instead of three polian vesicles ;
from R. uncinata (Hutton) in the form of the anchor plate perforations. R. gibbsi is
most closely related to R. hickmani Rowe and Pawson from Tasmania and more
recently collected from off Cape Howe, New South Wales (National Museum of
Victoria, no. H302 ; an extension of range to the Australian mainland) but the form
of the smaller anchor plates and the higher number of serrations on the arms of the
anchors (8-n in gibbsi ; up to 7 in hickmani) and the shape of the miliary granules
separate the two species.

Rowe (in Rowe & Pawson, 1967) mistakenly described the young, branched gonad
in the paratypes of R. hickmani as a stone canal. Gonads are obvious in specimens
of this genus which are as small as 20 mm in length.

The discovery of R. gibbsi increases the number of species of Rynkatorpa to three
occurring around the coasts of Australia. Interestingly the genus is now represented
on the north-west coast (R. bisperforata), off the north-east coast (R. gibbsi) and from
Tasmania and the south-eastern tip of Australia (R. hickmani). It is possible that
the genus circumscribes Australia but has yet to be discovered on the south or south-
western coasts. The other species have been recorded from New Zealand (R.
uncinata) ; Indonesia (bicornis and sluiteri) ; Indonesia, Fiji and Hawaii (challen-
geri) ; Aleutian Islands, North Pacific (duodactyla) and Andaman Islands, Indian
Ocean (timida).

2g6 F. W. E. ROWE

ACKNOWLEDGEMENTS

I would like to thank Miss A. M. Clark (British Museum (Natural History), London)
for sending the material to me for description and critically reading the manuscript,
and Dr P. E. Gibbs (Marine Laboratory, Plymouth, England) for agreeing to the
donation of the paratype to The Australian Museum collections.

REFERENCES

CLARK, H. L. 1907. The apodous holothurians. Smithson. Contr. Knowl. 35 : 1-206, 13 pis.

1946. The echinoderm fauna of Australia ; its composition and origin. Carnegie Inst.

Washington Publ. 556 : 1-567.

GIBBS, P. E., CLARK, A. M. & CLARK, C. M. 1976. Echinoderms from the northern region of
the Great Barrier Reef, Australia. Bull. Br. Mus. nat. Hist. (Zool.) 30, 4 : 101-144, 3 figs,
i pi.

MARTIN, W. E. 1969. Rynkatorpa pawsoni n. sp. (Echinodermata : Holothuroidea) a commen-
sal sea-cucumber. Biol. Bull. mar. Biol. Lab., Woods Hole 137 : 332-337, 9 figs.

ROWE, F. W. E. & PAWSON, D. L. 1967. A new genus in the holothurian family Synaptidae,
with a new species from Tasmania. Pap. Proc. R. Soc. Tasm. 101 : 31-35, 15 figs.

F. W. E. ROWE

Department of Marine Invertebrates

THE AUSTRALIAN MUSEUM

6-8 COLLEGE STREET

SYDNEY

NEW SOUTH WALES

AUSTRALIA 2000

A REVIEW OF THE FAMILY CENTROPOMIDAE
(PISCES, PERCIFORMES) : AN APPENDIX

By PETER HUMPHRY GREENWOOD

NIPHON SPINOSUS

WHEN preparing my recent review of the centropomid fishes (Greenwood, 1976) I
overlooked a relevant paper by Rivas & Cook (1968) on the relationships of the
'percichthyid' fish Niphon spinosus Cuv. These authors disagree with Gosline's
(1966) placement of this species in the Percichthyidae and consider that it is more
closely related to the centropomid genus Lates, as represented by the species L.
calcarifer. Indeed, they go so far as to suggest that the monotypic genus Niphon
should be placed in the Centropomidae rather than in the Percichthyidae.

Rivas & Cook (1968) reached this conclusion after comparing 22 characters (9
external osteological features, 7 characters of the squamation and 6 miscellaneous
ones) in Lates calcarifer (Bloch), Niphon spinosus Cuv. and Percichthys melanops
Girard. The degree of relationship among the three species is expressed as an index
for each character, and for each of the three groups of characters the indices are
summed to give an overall level of similarity. Thus, for the external osteological
features the index for Lates and Niphon is 13 and that for Niphon and Percichthys
is i ; for the scale characters the indices are n and 2 respectively, and for the
miscellaneous characters 8 and i respectively.

Unfortunately Rivas & Cook do not seem aware that 20 of the 22 characters, or
character states, employed in their analysis are shared either with several members
of the family Serranidae or with other members of the Percichthyidae (both families
sensu Gosline, 1966) in such a way that the indices of relationship are rendered
meaningless. That is to say, on these characters Niphon could just as well be related
to members of the Serranidae or to percichthyids other than Percichthys. (It is
clear that Rivas & Cook (1968 : 202) do not consider Gosline's Percichthyidae to be a
natural assemblage, but they offer no suggestions as to the affinities of the several
genera embodied in Gosline's (1966) concept of the family.) To further complicate
the issue, some of the characters present in Lates are absent or differently developed
in non-latine tribes of the Centropomidae (sensu Greenwood, 1976), and these
departures from the latine 'type' are not taken into account by Rivas & Cook.
For example, there are no ventral spines on the horizontal arm of the preoperculum
in Psammoperca, no opercular spine is present in any Centropomus species, and some
members of that genus are without a noticeably enlarged spine at the posterior angle
of the preoperculum (or the spine may be absent altogether, as in C. undecimalis
(Bloch), see fig. ice in Eraser, 1968). Again, in Centropomus the anterior end of the
swimbladder is not bilobed (see Eraser, 1968 ; Greenwood, 1976), although in several
species a pair of horn-like projections, sometimes of considerable length, are developed

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6 Issued 30 June, 1977

298

P. H. GREENWOOD

at the anterior end (Greenwood, 1976). Rivas & Cook (1968 : 203), however, state
that the bladder is bilobed in Centropomus (no species named) and thus it is like that
in Lates and Niphon but not Percichthys, an observation with which I would disagree.

In brief, 20 of the 22 characters considered by Rivas & Cook (1968) cannot be
used to establish a closer relationship between Niphon spinosus and the centropomids
than between Niphon and the taxa placed by Gosline (1966) in the Serranidae,
Grammistidae and the Percichthyidae. Most of the characters used by Rivas &
Cook seem to be symplesiomorph ones, and the five apomorphic ones are too widely
distributed amongst related taxa in the basal percoid radiation to be of value in this
context.

Rivas & Cook (1968 : 203) also think that most of the characters used by Gosline
(1966) in his table contrasting the Serranidae and Percichthyidae (few of which
characters were used by Rivas & Cook) '. . . indicate closer relationship of Niphon
with the Centropomidae than with the Percichthyidae'. On the basis of my analysis
of centropomid features (Greenwood, 1976) I would consider these characters to have
about the same value as those used by Rivas & Cook (1968), and thus certainly not
indicative of a close phyletic relationship between Niphon and the centropomids.

The caudal fin skeleton (Fig. i) in Niphon spinosus (a feature not used by Rivas &
Cook) provides few characters of value in determining relationships. There are 3
epurals (as in some centropomids), 2 uroneurals very closely applied to one another
and apparently fused basally (some centropomids have 2 uroneurals but these are
clearly distinct, see Greenwood, 1976), and the anterior uroneural is drawn out and
expanded basally so that, with its partner of the opposite side, it forms a steeply

FIG. i. Caudal fin skeleton of Niphon spinosus (specimen 156 mm standard length, BMNH
reg. no. 1879.5.14:144, Yokohama), x : point at which the two uroneurals apparently
fuse.

NIPHON AND THE CENTROPOMIDAE 299

pitched roof over the neural groove on the fused first ural and preural centra. In
one specimen (c. 135 mm standard length) the expanded portion of the uroneurals is
closely applied ventrally to the underlying centrum, but in a larger fish (156 mm
S.L.) there is a slight gap between the bones. The neural spine of the second ural
centrum (U2) is short but greatly expanded ; posteriorly it articulates with the front
margin of the expanded uroneurals. In all these features the caudal skeleton is
virtually identical with that in the 'percichthyids' I have examined (Percolates,
Lateolabrax, Ctenolabrax) or which were mentioned by Gosline (1966) ; it also
approaches the condition found in Centropomus amongst the Centropomidae (see
Fraser, 1968 ; also personal observations) although in that genus the first uroneural
is not so markedly expanded. However, the presence of 3 epurals and 2 uroneurals
is certainly a plesiomorphous condition, and an expanded antero-basal part of the
first uroneural is likewise primitive, judging from its widespread occurrence amongst
basal neoteleostean fishes as well as in the basal percoids (see figures in Rosen, 1973
and Monod, 1968 respectively). The caudal skeleton in centropomids, other than
Centropomus, shows more derived features.

The two exceptional characters used in Rivas & Cook's (1966) analysis (see above)
which are not shared by the Centropomidae on the one hand and the Serranidae and
Percichthyidae on the other, are the extension of the lateral line beyond the caudal
base, and the presence of an expanded, blade-like neural spine on the second ab-
dominal vertebra.

Rivas & Cook (1968 : table 3) give an affinity index to Lates and Niphon because,
according to them, in these species but not in Percichthys the lateral line '. . . extends
beyond the caudal base'. I would disagree with their decision because in Niphon
only about 3 or 4 pore-bearing scales extend posteriorly beyond the level of the
hypural-caudal fin ray junction, and even then none extends onto the caudal fin
membrane. In all centropomids bar one (Lates (Luciolates] stappersi (Blgr.)) the
lateral line scales extend to the posterior margin of the caudal fin (see Greenwood,
1976 : 48). The condition of the lateral line in Lates stappersi is clearly a derivative
of the usual tripartite Lates type and cannot thus be used to invalidate the argument
that the lateral line extends to the caudal fin margin in centropomids. The condition
of the lateral line in Niphon spinosus is, in fact, exactly like that in Percalates
colonorum (Gunth.) and Lateolabrax japonicus (Cuv.), two species which Gosline
(1966) included in the Percichthyidae, and also like that in at least some serranids as
well (e.g. in the genus Epinephalus) . Niphon spinosus then certainly does not have
a centropomid-like caudal extension of the lateral line, although the lateral line does
extend 2 or 3 scales further posteriorly than it does in Percichthys trucha (Val.), and
6 scales further than in the specimens of P. melanops I have examined (where the
lateral line pore-scales do not even extend to the level of the hypural-fin ray joints).

Rivas & Cook (1968) do not consider the blade-like expansion of the second neural
spine in Centropomidae (Gosline, 1966 ; Greenwood, 1976) to be of diagnostic value
since in their opinion '. . . the expansion is only a matter of degree . . .'. Be that as
it may (and several of the characters used in their tables are, as they admit, also ones
of degree), there is still a trenchant difference between the shape of this neural
spine in all centropomids and its shape in the Serranidae and other basal percoids

300 P. H. GREENWOOD

(personal observations, see also Gosline, 1966, and Greenwood, 1976). Thus, I do
not consider that Rivas & Cook (1966) have provided any evidence to weaken or
destroy the value of an extended lateral line and an expanded second neural spine,
when taken in combination, as diagnostic autapomorphous characters for the family
Centropomidae (see also discussion in Greenwood, 1976 : 10-11 & 62-71).

Since Niphon shows neither of the principal diagnostic features of the Centro-
pomidae, and because none of the characters adduced by Rivas & Cook (1968) to
show its closer affinity with that family than with the Percichthyidae is in fact a
true shared specialization, there are no grounds for placing Niphon spinosus in the
Centropomidae.

The difficulties of establishing phyletic relationships amongst the basal percoids is
well demonstrated by Rivas & Cook's paper, as it was by Gosline's (1966) attempt to
clarify serranid-percichthyid interrelationships. These difficulties are not much
reduced when phyletic rather than phenetic principles of classification are applied
to the task, although some progress has been made (see Rosen, 1973 ; Greenwood,
1976). For the moment, Niphon will have to be retained in the heterogeneous and
aphyletic assemblage of serranid-percichthyid species. If certain characters
apparent in this generic complex do later prove to be reliable phyletic indicators,
then I suspect that, contrary to the views of Rivas & Cook (1968), Niphon will
prove to have as its nearest relatives at least some, if not all of those taxa included
by Gosline (1966) in his Percichthyidae.

CENTROPOMIDAE

Since my review of the Centropomidae was published (Greenwood, 1976), I have
found in it one error of fact and two lapsi that can now be corrected :

DORSAL AND ANAL FIN SKELETONS. On page 45, when describing the dorsal and
anal fin skeleton in Lates, it is said that except for Lates (Luciolates) stappersi, no
Lates species has distinct medial radials supporting the branched dorsal fin rays.
This latter statement is wrong, as recently prepared dissections and alizarin trans-
parencies have shown. In all Lates species discrete medial radials are present in the
posterior 5 to 8 pterygiophores. In this respect both Lates and Psammoperca differ
from Centropomus, where all the medial radials are fused with their respective
proximal pterygiophores. The condition in Centropomus is the derived one, that in
Lates and Psammoperca the plesiomorphous one (which, incidentally, is also the
condition in at least the four genera of Gosline's (1966) Percichthyidae which I have
examined (Percichthys, Percilia, Roccus and Niphon) ; the Serranidae, on the other
hand, have the fused, apomorphic, condition seen in Centropomus).

BRANCHIAL SKELETON OF Lates (page 37). When describing the upper pharyngeal
dentition it is said that there is a cup-shaped tooth-plate fitting closely around the
fourth epibranchial ; the latter bone is, of course, the fourth pharyngobranchial.

DEFINITION OF THE SUBFAMILY CENTROPOMINAE (page 76), ornamentation of the
opercular series. It is the preoperculum and not the operculum that has three or
four enlarged spines at its posterior angle.

NIPHON AND THE CENTROPOMIDAE 301

ACKNOWLEDGEMENTS

I am very grateful to my colleague Gordon Howes for drawing my attention to
Rivas & Cook's paper, and to Dr Victor Springer of the U.S. National Museum, whose
pertinent questioning led to my reinvestigating the condition of the posterior ptery-
giophores in Lates.

REFERENCES

FRASER, T. H. 1968. Comparative osteology of the Atlantic Snooks (Pisces, Centropomus) .

Copeia 1968 : 433-460.
GOSLINE, W. A. 1966. The limits of the fish family Serranidae with notes on other lower

percoids. Proc. Calif. Acad. Sci. 33 : 91 — 111.
GREENWOOD, P. H. 1976. A review of the family Centropomidae (Pisces, Perciformes).

Bull. Br. Mus. nat. Hist. (Zool.) 29 : 1-81.

MONOD, T. 1968. Le complexe urophore des teleost6ens. Mem. Inst.fr. afr. noire 81 : 1-705.
RIVAS, L. R. & COOK, B. A. 1968. Relationships of the western Pacific 'percichthyid' fish

Niphon spinosus, with the family Centropomidae. Wassman J. Biol. 26 : 201-208.
ROSEN, D. E. 1973. Interrelationships of higher euteleostean fishes. Zool. J. Linn. Soc.

53 (Suppl. i) : 397-5*3-

P. H. GREENWOOD, D.Sc.

Department of Zoology

BRITISH MUSEUM (NATURAL HISTORY)

CROMWELL ROAD

LONDON SW7 5BD

A REVISION OF THE SYNONYMY OF

THREE SPECIES OF LEIOLOPISMID SKINKS

FROM NEW ZEALAND

By JOAN ROBB

CONTENTS

Page
SYNOPSIS ........... 303

INTRODUCTION .......... 303

DESCRIPTION OF SPECIES ......... 304

Leiolopisma ornatum . . . . . . . . .304

Leiolopisma zelandicum . . . . . . . . .306

Leiolopisma moco .......... 308

ACKNOWLEDGEMENTS ......... 309

REFERENCES ........... 310

SYNOPSIS

The synonymies of three species of New Zealand skinks are reviewed and clarified. Their
valid names are Leiolopisma ornatum (Gray), L. zelandicum (Gray) and L. moco (Dumeril &
Bibron).

INTRODUCTION

FOR MANY years there has been considerable confusion over the identity of some
species of New Zealand skinks ; and it has been evident that some basic errors must
have occurred in their complex synonymies.

Historically, it is easy to see how this could have happened. The initial collection
of biological specimens in New Zealand was done by scientists from the northern
hemisphere, some of whom accompanied various expeditions (U.S. Exploring
Expedition, 1838-1842 ; Erebus & Terror, 1839-1843 ; Novara, 1857-1859 ;
Challenger, 1873-1876 ; and a few (such as Dr Dieffenbach, naturalist to the New
Zealand Company) who travelled independently. In all cases, the material collected
was sent back to various European and American museums, and thus many reptilian
(and other) type specimens came to be lodged in London, Paris, Hamburg and
certain American cities. Furthermore, as several expeditions were operating in the
Pacific during the period 1835-1870, a few species were described almost simul-
taneously in different reports, resulting in a taxon being given different specific, or
even different generic names. Another source of error was the fact that recorded
details of the localities from which specimens were taken were often meagre in the
extreme. Thus not only are many specimens merely labelled 'New Zealand', but
in some cases there is doubt about whether they were in fact taken in New Zealand,
Tasmania or Australia.

Bull. Br. Mus. nat. Hist. (Zool.) 31, 6 Issued 30 June, 1977

3o4 JOAN ROBB

For some years after the settlement of New Zealand much material was still
despatched to the British Museum for identification and lodgement. Gradually, as
museums became established in New Zealand, more and more material was deposited
locally. Thus, by the time McCann (1955) made his notable revision of the New
Zealand lizards, he was faced with the situation where many type-specimens, being
deposited in European or American institutions, were unavailable to him and he had
to rely on the frequently meagre descriptions already published and any further
information that could be supplied by the Curators of the day from the by then
often badly faded type-material. It is not surprising that in some cases McCann
made mistakes in relating unseen type-specimens to particular series held in New
Zealand museums.

The present revision has been made in the British Museum (Natural History)
(BM(NH)), using material held there, and comparing it directly with recently col-
lected series brought from New Zealand, and with other material kindly lent by the
Paris Museum of Natural History (MNHP), and the Leiden Museum of Natural
History.

DESCRIPTION OF SPECIES

Leiolopisma ornatum (Gray)

Tiliqua ornata Gray, 1843 : 202.

Hinulia ornata (part) Gray, i845a : 77.

Hinulia ornata (part) Gray, i845b : 7.

Cyclodina aenea Girard, 1857 : 196.

Euprepes ornatus Steindachner, 1867 : 49.

Lygosoma aeneum Boulenger, 1887 : 275.

Lygosoma ornatum Boulenger, 1887 : 317, PI. 26, Figs i, la.

Liolepisma aeneum Lucas & Frost, 1897 : 278.

Homolepida ornatum Lucas & Frost, 1897 : 279.

Leiolopisma aeneum McCann, 1955 : 102.

Sphenomorphus pseudornatus McCann, 1955 : 125.

DESCRIPTION. Habit lacertiform ; body rather stout ; limbs well developed,
pentadactyl ; snout short, obtuse ; lower eyelid with a divided or undivided
transparent disc ; nostril pierced in the nasal ; no supranasal ; frontonasal
broader than long, forming a suture with the rostral and the frontal ; prefrontals
small, clearly separated ; frontal equal to or a little longer than the frontoparietals,
and shorter than the frontoparietals and interparietal together, in contact with the
first and second supraoculars ; 4 supraoculars, the second the largest, the third
almost equal to it ; 7 or 8 supraciliaries ; parietals bordered by a pair of nuchals
and a pair of temporals ; one pair of nuchals, sometimes followed by several pairs
of enlarged dorsal scales ; 6-8 supralabials ; 5 or 6 infralabials ; anterior and pos-
terior loreals contact up to \ (occasionally all) of the upper border of the second
supralabial ; subocular scales row complete, excluding any supralabials from the
orbit ; earhole circular, smooth or with a few anterior lobules ; 26-30 mid-body
scale-rows, scales more or less striated, laterals smallest, dorsals and ventrals almost
equal in size ; preanal scales somewhat enlarged ; limbs short, and when adpressed

NEW ZEALAND SKINKS 305

just meet or are distant ; digits subcylindrical, 15-21 smooth lamellae below the
fourth toe, and 43-60 in total below toes two to five inclusive (in general the higher
numbers occur in specimens from the northern part of the species' range) ; tail
slightly longer than snout-vent length, and tapering finely.

MEASUREMENTS (of larger specimen in type bottle BM 1946.8.19.38). End of
snout - forelimb 20 mm ; axilla - groin 30 mm ; snout - vent 56-5 mm ; tail
61 mm ; nostril - anterior of eye 2-25 mm ; nostril - anterior of ear 9 mm ; pos-
terior of ear - axilla 10-5 mm.

COLOUR. Mottled coppery brown dorsally ; somewhat paler laterally with dark
brown markings ; and creamy-white ventrally, often with scattered dark spots, and
chin and throat often heavily speckled with black. An irregular pale line bordered
below by dark brown or black extends from above the eye to the root of the tail ;
and below the eye there is a pale spot edged with dark brown - this gives a 'tear-
drop' effect.

DISTRIBUTION. New Zealand, endemic. Common in many parts of the North
Island and on off-shore islands.

TYPE LOCALITY. Cook's Straits, New Zealand. Types in BM(NH).

SYNONYMY. This species has the most confused synonymy of any of the three
under consideration. The original description of Tiliqua ornata given by Gray
(1843), specifically mentioned that he had only a single specimen. Two years later
in his Catalogue of Lizards, Gray (i845a) transferred the species to Hinulia, and
listed a second specimen (presumably in a separate bottle), of which he gave a some-
what different description. The second is probably BM XV.n.b, which has in the
past been listed as a syntype of H. ornata, but is in fact a specimen of Leiolopisma
moco. Specimen (a) of Gray's (i845a) Catalogue of Lizards must be the type of
T. ornata. The accession number on the bottle is 42.8.19.2, and the corresponding
entry in the accession book reads 'Tiliqua ornata Gray. N.Z. Presented by R. Owen,
Esq. from specimens sent by Dr. Dieffenbach to the College of Surgeons'. Boulenger
(1887 : 317) evidently found two specimens in bottle no. 42.8.19.2, as he listed an
adult and a half grown animal as types. These two specimens are still extant, and
still in one bottle ; they are listed by their post-war numbers, 1946.8.19.38 and
1946.8.19.39. There is no way of telling which is the original specimen, or when or
from whence the second specimen came to the Museum. However, as both belong
to the same species there is no question of confusion of specific identity.

The diagram of H. ornata referred to by Gray (i845b : pi. n, fig. i) clearly was
not drawn from either of these specimens, and is probably of BM XV.ii.b (Leiolopisma
moco). Although Gray (i845b) referred to this figure on p. 7 where he listed it as
pi. n, fig. 2, it was not published until 1867, and was later included by Giinther

(1875).

One of the best diagnostic characters of L. ornatum, taken in conjunction with
size and colour, is the complete subocular scale row, which excludes any of the upper
labials from the orbit. This feature is clearly seen on the two specimens now known
as types, but was not mentioned by Gray in either of his descriptions.

306 JOAN ROBB

In 1857 Girard published a description of Cydodina aenea from New Zealand and
from the details he gave, including reference to the complete subocular row, it is
clear he was dealing with the species named H. ornata by Gray. Boulenger (1887)
recognized that Gray had included specimens of two species in his description of H.
ornata, and referred part of Gray's H. ornata to Lygosoma ornatum, and part to L.
moco. Boulenger gave a much more detailed description of L. ornatum than had
Gray, and referred specifically to the scaly lower eyelid (mentioned by Gray in his
diagnosis of the genus Hinulia}, and the yellow, dark-edged spot below the eye. He
did not mention the complete subocular scale row, though this feature is very
clearly shown in his illustrations of the species (Boulenger, 1887, pi. xxvi, figs I, ia).

Lucas & Frost (1897) transferred the species to the genus Homolepida Gray, and
did not attempt a redescription, but quoted verbatim from Boulenger. They added
this comment : 'In habit and coloration this lizard has a strong resemblance to
Leiolopisma aeneum' and gave the distribution of H. ornatum as The neighbourhood
of Auckland'. (Steindachner, 1867, referred to 'numerous examples from Auck-
land'.)

McCann (1955) also recognized that Gray had included members of two species
in his description of H. ornata, but erred in stating that Gray regarded his Tiliqua
ornata and Hinulia ornata as conspecific. In fact the name H. ornata was a subse-
quent usage of T. ornata and so the same type specimen relates to both names.
McCann also believed Gray's type(s) to be lost.

On the correct premise that Gray had included two species under one description,
and the erroneous premise that he regarded T. ornata and H. ornata as conspecific,
and must therefore have had two type specimens, McCann tried to deduce which
were the two species concerned. He pointed out that while Gray (i845a) had men-
tioned the scaly lower eyelid in defining the genus Hinulia, he had made no mention
of it in describing T. ornata (1843). McCann therefore assumed that Gray had con-
fused two species, which could in fact be separated on the basis of presence or absence
of a scaly lower eyelid, and that as this was 'an error of determination' the name
H. ornata was invalid (Rules of Nomenclature : Art. 49).* McCann accordingly
retained the specific name 'ornata' for the species he assumed to have been Gray's
Tiliqua ornata, and renamed Gray's Hinulia ornata as Sphenomorphus pseudornatus.
To complicate the issue further, McCann did not recognize L. aeneum as conspecific,
nor that the scaly character of the lower eyelid is extremely variable, and not a good
character for this species. From examination of many specimens it is clear that
while many individuals do show this feature, many more may lack it, or even possess
a scaly eyelid on one side, and an undivided transparent disc on the other.

Leiolopisma zelandicum (Gray)

Tiliqua zelandica Gray, 1843 : 202.
Mocoa zelandica (part) Gray, i845a : 82.
Lygosoma moco (part) Boulenger, 1887 : 272.
Leiolopisma moco (part) Lucas & Frost, 1897 : 276.
Leiolopisma ornata McCann, 1955 : 109.

* Quoted as Art. 31 by McCann.

NEW ZEALAND SKINKS 307

DESCRIPTION. Habit lacertiform ; body moderately elongate ; limbs well
developed, pentadactyl ; snout short, subacute ; lower eyelid with an undivided
transparent or opaque disc ; nostril pierced towards the centre of the nasal ; no
supranasal ; rostral almost twice as wide as deep ; frontonasal domed posteriorly,
forming a suture with the rostral and the frontal, prefrontals widely separated ;
frontal longer than the frontoparietals, but shorter than the frontoparietals and
interparietal together, in contact with the first and second supraoculars ; 4 supra-
oculars, the second the largest, and the first the smallest ; 7 or 8 (occasionally 9)
supraciliaries ; parietals bordered by a pair of nuchals and a pair of temporals ;
1-3 pairs of nuchals, sometimes followed by up to 3 pairs of enlarged dorsal scales ;
7 supralabials ; 6 or 7 infralabials ; anterior and posterior loreals contact the entire
upper border of the second supralabial, the posterior loreal sometimes extending
slightly above the third supralabial ; subocular scale row interrupted by the fifth
supralabial ; earhole oval or circular, with few or many projecting lobules ; 32-34
mid-body scale rows, the dorsal scales largest and faintly striate ; preanal scales
not or slightly enlarged ; adpressed limbs just meet or fail to meet ; digits moderately
long, subcylindrical, 16-21 smooth lamellae below the fourth toe, and 50-62 in total
below toes two to five inclusive ; tail stout, tapering, somewhat longer than snout-
vent length.

MEASUREMENTS OF TYPE (BM 1946.8.16.19). End of snout - forelimb 16-00 mm ;
axilla - groin 27-00 mm, snout - vent 48-00 mm, tail 67-00 mm, nostril - anterior of
eye 2-5 mm, nostril - anterior of ear 8-00 mm, posterior of ear - axilla 9-00 mm.
(The type specimen is not a fully mature animal.)

COLOUR. Dorsal surface olive brown with a variable pattern of dark flecks or
blotches ; the head may be unmarked, or heavily marked with dark areas. A
large dark spot is normally discernible on the outer edge of each parietal, though
this feature is not unique to this species. A pale dorso-lateral line extends posteriorly
from behind the eye, breaks up into disjointed specks behind the insertion of the fore-
limb and continues so on to the tail. This line is bordered below by black. A
lateral line extends from the ear to the top of the forelimb and becomes broken up
into an irregular pattern of markings along the flanks. These two pale lines are
separated by a variable wide brown band. There is a pale line on the dorsal surface
of the forelimb and a similar, but more broken line on the dorsal surface of the hind-
limb. Ventral surface pale creamy- white ; the chin and throat or the entire under-
surface may be speckled with black or dark brown.

DISTRIBUTION. New Zealand, endemic. Mainly found in the Wellington district
and on the islands in Cook's Strait.

TYPE LOCALITY. Cook's Straits. Type in BM(NH), BM 1946.8.16.19.

SYNONYMY. In 1843 Gray described a new species of skink from New Zealand as
Tiliqua zelandica, based on only one specimen. In i845a, b, he transferred T.
zelandica to Mocoa zelandica and Boulenger (1887) included both names in the
synonymy of Lygosoma moco (Dumeril & Bibron) while Lucas & Frost (1897)
followed Boulenger's lead. McCann (1955) concluded, correctly, that there was no

308 JOAN ROBB

evidence to suggest that Dumeril & Bibron's (1839) Lygosoma moco and Gray's
(1843) T. zelandica were conspecific, and that Lygosoma moco as defined by Boulenger
(1887) consisted of two species, L. moco and L. zelandica. Unfortunately, McCann
was not able to examine the type specimen of L. zelandica, and on the information
available to him, related it to a different species, and not to the correct one, which
he regarded as L. ornata.

Leiolopisma moco (Dumeril & Bibron)

Lygosoma moco Dumeril & Bibron, 1839 : 718.

Hinulia ornata (part) Gray, i845a : 77.

Mocoa zelandica (part) Gray, i845a : 82.

Mocoa owenii (part) Gray, i845a : 272.

Mocoa zelandica (part) Gray, 18455 : 8.

Euprepes moco (part) Steindachner, 1867 : 47.

Mocoa zealandica (part) Giinther, 1875 : 13, PL 7, Fig 4.

Lygosoma moco (part) Boulenger, 1887 : 272.

Leiolopisma moco (part) Lucas & Frost, 1897 : 2?6-

Leiolopisma moco McCann, 1955 : 104.

Leiolopisma zelandica (part) McCann, 1955 : 106.

DESCRIPTION. Habit lacertiform ; body moderately elongate ; limbs well
developed, pentadactyl ; snout short, obtuse ; lower eyelid with an undivided
transparent disc ; nostril pierced more or less in the centre of the nasal ; no supra-
nasal ; rostral less than twice as wide as deep ; frontonasal broader than long, often
pointed posteriorly, forming a suture with the rostral and the frontal ; prefrontals
distant or nearly meeting ; frontal shorter than the frontoparietals and interparietal
together, in contact with the first and second supraoculars ; 4 supraoculars, the
second largest and the first smallest ; 6-8 supraciliaries ; parietals bordered by a
pair of nuchals and a pair of temporals ; 2-5 pairs of nuchals, sometimes followed
by enlarged dorsals ; 7 or 8 supralabials ; 6 or 7 infralabials ; anterior and posterior
loreals contact upper border of second supralabial and up to \ of third supralabial ;
subocular row interrupted by the fifth (rarely sixth) supralabial ; earhole circular,
with few to many anterior lobules ; 28-34 mid-body scale rows, the dorsal scales
largest, smooth or faintly striate ; preanal scales enlarged or not enlarged ; adpressed
limbs meet or fail to meet ; digits moderately long, subcylindrical, 18-27 smooth
lamellae below the fourth toe, and 60-69 (rarely 53-73) in total below toes two to
five inclusive ; tail relatively long and tapering, at least i£ times the snout-vent
length.

MEASUREMENTS OF TYPE (MHNP3OI9). End of snout - forelimb 23mm;
axilla - groin 33 mm ; snout - vent 66 mm ; tail (broken) ; nostril - anterior of eye
3 mm ; nostril - anterior of ear 10 mm ; posterior of ear - axilla n mm.

COLOUR. In this species colour and pattern are very variable. Generally
coppery or olive brown, usually with a (sometimes double), mid-dorsal dark line ; a
narrow pale dorso-lateral stripe extending from the snout to the base of the tail,
and a similar lateral stripe from behind the ear to the groin, both these pale stripes
are edged above and below by a narrow dark line. Between the two is a wider

NEW ZEALAND SKINKS 309

mid-brown stripe, approximately two scales wide, and may be plain or speckled or
broken up into dark blotches. The linear markings may be straight-edged or
irregular. The ventral surface is very pale grey or cream-coloured, sometimes with
dark specks in longitudinal rows. In some specimens there is an X-shaped mark
on the dorsal surface of the head, between the two series of supraoculars ; and
there may be a dark spot on each labial scale.

DISTRIBUTION. New Zealand, endemic.

TYPE LOCALITY. 'New Zealand'. Type in the National Museum of Natural
History, Paris.

SYNONYMY. The synonymy of this species is interwoven with that of L. zelandi-
cum, and to a much lesser extent, with that of L. ornatum. Dumeril & Bibron (1839)
based their description of the species partly on the type in the National Museum of
Natural History in Paris, and partly on specimens held in London. They comment
'. . . we have seen several specimens in the collection of the Zoological Society of
London, labelled Tiliqua moco by Mr. Gray'.* These latter specimens cannot now be
traced. However, the name Tiliqua moco Gray, quoted by Dumeril & Bibron
(1839) was never published, and Gray himself lists the name in the synonymy of
Mocoa zelandica as 'Tiliqua moco Gray M SS.' (i845a, b). Thus the first published
name of the species was Lygosoma moco Dum. & Bibr., 1839.

In 1843 Gray described Tiliqua zelandica, and in 1845 he described Mocoa zelandica
and referred T. zelandica to the synonymy of the species, as well as giving Dumeril &
Bibron's (1839) reference. This mistaken assumption that T. zelandica and L. moco
were conspecific was perpetuated by Steindachner (1867), Giinther (1875) and Bou-
lenger (1887), although the latter was responsible for recognizing that Gray (i845a, b)
had confused a specimen of L. moco with Hinulia ornata. Lucas & Frost (1897)
followed Boulenger's synonymy for L. moco. As has already been stated McCann
(1955) recognized that L. moco and T. zelandica were not conspecific, but unfortunately
associated the latter name with the wrong species.

In passing, it is appropriate to note that McCann was probably wrong in including
Hinulia variegata Buller (1870) and Mocoa striata Buller (1870) in the synonymy of
L. zelandica ; they are more likely to be conspecific with L. moco. However, as the
specimens are no longer available, and the descriptions given are meagre, it is not
possible to be sure of their identity.

ACKNOWLEDGEMENTS

I wish to express my sincere thanks to Miss A. G. C. Grandison, Curator of Her-
petology, British Museum (Natural History), and the members of her staff for their
hospitality within the Section, and access to the collection, and for many forms of
help while I was carrying out this work. I am also very grateful for the loan of
material from the National Museum of Natural History, Paris, and from the Museum
of Natural History, Leiden.

* Author's translation.

310 JOAN ROBB

REFERENCES

BOULENGER, G. A. 1887. Catalogue of the lizards in the British Museum (Natural History),

vol. 3, ed. 2. xii + 436 pp. London.
BULLER, W. 1870. A list of the lizards inhabiting New Zealand, with descriptions. Trans.

N.Z. Inst. 3: 4-11.
DUMERIL, A. M. C. & BIBRON, G. 1839. Erpdtologie ginirale ou histoire naturelle complete des

Reptiles, 5. 854 pp. Paris.

GIRARD, C. 1857. Descriptions of some new reptiles, collected by the U.S. Exploring Ex-
pedition. Proc. Acad. nat. Sci. Philad. 8 : 195-199.
GRAY, J. E. 1843. Descriptions of Reptilia and Amphibia. In E. Dieffenbach, ed., Travels

in New Zealand, 2 The Fauna of New Zealand.
i845a. Catalogue of the specimens of lizards in the collection of the British Museum, xxviii +

289 pp. London.
i845b. The reptiles of Australia. In J. Richardson & J. E. Gray, eds, The Zoology of the

Voyage of H.M.S. Erebus and Terror, under the command of Captain Sir James Clark Ross,

R.N., F.R.S., during the years 1839 to 2^43> 2 (i) : 1-8.
1867. The Lizards of Australia and New Zealand in the collection of the British Museum.

7 pp. + 20 pis. London.
GUNTHER, A. 1875. A list of the saurians of Australia and New Zealand. In J. Richardson

and J. E. Gray, eds, The Zoology of the Voyage of H.M.S. Erebus and Terror, under the

command of Captain Sir James Clark Ross, R.N., F.R.S., during the years 1839 to 1843,

2 (i) -.9-19.
LUCAS, A. H. S. & FROST, C. 1897. On the New Zealand lizards. Trans. N.Z. Inst. 29 :

264-280.

McCANN, C. 1955. The lizards of New Zealand. Dom. Mus. Bull. 17 : 126 pp.
STEINDACHNER, F. 1867. Reise der Frigatte Novara 1857-1859, Zoologischer Theil, 1 (3 and

4) : Reptilien - Amphibien.

J. ROBB

UNIVERSITY OF AUCKLAND,

AUCKLAND,

NEW ZEALAND

A LIST OF SUPPLEMENTS
TO THE ZOOLOGICAL SERIES

OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

1. KAY, E. ALISON. Marine Molluscs in the Cuming Collection British Museum
(Natural History) described by William Harper Pease. Pp. 96 ; 14 Plates.
1965. (Out of Print.)

2. WHITEHEAD, P. J. P. The Clupeoid Fishes described by Lacepede, Cuvier and
Valenciennes. Pp. 180 ; n Plates, 15 Text-figures. 1967. £4-80.

3. TAYLOR, J. D., KENNEDY, W. J. & HALL, A. The Shell Structure and
Mineralogy of the Bivalvia. Introduction. Nuculacea-Trigonacea. Pp. 125 ;
29 Plates, 77 Text-figures. 1969. £5-40.

4. HAYNES, J. R. Cardigan Bay Recent Foraminifera (Cruises of the R.V. Antur)
1962-1964. Pp. 245 ; 33 Plates, 47 Text-figures. 1973. £11-90.

5. WHITEHEAD, P. J. P. The Clupeoid Fishes of the Guianas. Pp. 227 ; 72
Text-figures. 1973. £10.65.

6. GREENWOOD, P. H. The Cichlid Fishes of Lake Victoria, East Africa : the
Biology and Evolution of a Species Flock. Pp. 134 ; i Plate, 77 Text-figures.
1974. £3-75. Hardback edition £6.

Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU