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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 73 BAND 73

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 73 BAND

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN_ KAAPSTAD

1977

# SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY

THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE

440

LIST OF CONTENTS

CLARK, A. M.

The South African Museum’s Meiring Naude cruises, Part 4. Echinoderms.

(Published June 1977.)

EASTWOOD, E. B.

Notes on the scorpion fauna of the Cape, Part 2. The Parabuthus capensis (Ehren-

berg) species-group; remarks on taxonomy and bionomics (Arachnida,

Scorpionida, Buthidae). (Published August 1977.)

EstTEs, R.

Relationships of the South African fossil frog Eoxenopoides reuningi (Anura,

Pipidae). (Published June 1977.) a on nf 53

GRIFFITHS, C. L.

Deep-sea amphipods from west of Cape Point, South Africa. (Published June 1977.)

KENNEDY, W. J. see KLINGER, H. C.

KENNEDY, W. J. & KLINGER, H. C.

Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family

Tetragonitidae Hyatt, 1900. (Published August 1977.) ..

KLINGER, H. C.

Cretaceous deposits near Bogenfels, South West Africa. (Published June 1977.) ..

KLINGER, H. C. & KENNEDY, W. J.

Cretaceous faunas from Zululand, South Africa, and southern Mozambique. The

Aptian Ancyloceratidae (Ammonoidea). (Published December 1977.)

MILLARD, N. A. H.

Hydroids from the Kerguelen and Crozet shelves, collected by the cruise MD. 03 of

the Marion- Dufresne. (Published April 1977.) : y hye &

MILLARD, N. A. H.

The South African Museum’s Meiring Naude cruises, Part 3. Hydroida. (Published

June 1977.) Ae is xe pt Ee Be ae De

WOOLDRIDGE, T.

A new species of Halicyclops (Copepoda, Cyclopoida) from estuaries in Transkei,

southern Africa. (Published November 1977.) :

Page

133

199

149

215

105

361

NEW GENERIC AND SUBGENERIC NAMES PROPOSED

IN THIS VOLUME

Page

Helicancyloceras Klinger & Kennedy, 1977 ae ot Le ae os @-o2)

Helicancyloceras (Helicancyloceras) Klinger & Kennedy, 1977 ee ca a we OST

Helicancyloceras (Nonyaniceras) Klinger & Bee 1977 ae see a rs P27 |

Uniscyphus Millard, 1977 oe ae ae fe a ae Fe bse Roe! 274

VOLUME 73 PART 1 APRIL 1977 | ISSN 0303-2515

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIscHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs

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Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume. 73 Band

April 1977 April

Part 1 Deel

HYDROIDS FROM THE KERGUELEN

PEC ROZET SHELVES, COLLECTED BY

THE CRUISE MD.03 OF THE MARION-DUFRESNE

N. A. H. MILLARD

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

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OUT OF PRINT/ UIT DRUK

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6(1, t.—p.i.), 711-4), 8, 91-2, 7), 10(1),

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ISBN 0 908407 12 2

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES,

COLLECTED BY THE CRUISE MD.03 OF THE MARION-DUFRESNE

N. A. H. MILLARD

South African Museum, Cape Town

(With 12 figures and 2 tables)

[MS. accepted 28 December 1976]

ABSTRACT

Thirty-four species of hydroids are described from the Kerguelen and Crozet groups of

islands, including two new species: Halecium dufresneae and Zygophylax crozetensis. Among

them are five cosmopolitan species. The rest show affinities partly with the cold-temperate

Magellan Region of South America, and partly with Antarctica. The fauna is considered to

be subantarctic in nature.

CONTENTS

PAGE

Introduction é : : : : 1

Station list . ; ; 3 : ’ 2

Systematic section . : : : 3

Discussion . : ; . : ; 44

Acknowledgements . : : : 46

References . : , ; : i 46

INTRODUCTION

The cruise MD.03 of the Marion- Dufresne during the period 18 March to

28 April 1974 to the Kerguelen and Crozet groups of islands was intended as

a preliminary reconnaissance of the area to serve as a basis for a more detailed

survey. The cruise was highly successful in the quantity and variety of benthic

invertebrates taken, and although the subsequent cruises MD.04 (1975) and

MD.08 (1976) have now been completed, it may be some time before the hydroid

material is all sorted and worked up. It was felt that the publication of the

present results should not be delayed.

The cruise was financially and logistically supported by the Terres Australes

et Antarctiques Frangaises, Paris. The bulk of the material and the types of the

new species will be deposited in the Muséum National d’Histoire Naturelle, and

a few duplicates are being retained at the South African Museum.

The cruise MD.03 was described by the Chef de la Campagne, J. C. Hureau

(1976), who also gave a complete station list and a chart of the voyage. The

positions of the stations yielding hydroid material are, however, repeated here

for convenience.

Ann. S. Afr. Mus. 73 (1), 1977: 1-47, 12 figs, 2 tables.

ANNALS OF THE SOUTH AFRICAN MUSEUM

The Kerguelen Shelf lies in the southern Indian Ocean at approximately

50°S and 70°E. Kerguelen is the largest island, Heard and MacDonald lying

about 644 km to its south-east. The depths from which hydroids were taken

range from 6 to 790 m, the abyssal dredgings from over the edge of the plateau

yielding no hydroid material.

The Crozet Shelf (including among others the Ile de la Crozet, Ile de la

Possession and Ile aux Cochon) lies some | 930 km to the west of Kerguelen,

and hydroids were dredged from depths of 110 to 400 m.

STATION LIST

2/6. E. Kerguelen; 4.4.74; 49°30,7’'S 70°44,7’E; 115 m: Zygophylax crozetensis, Sym-

plectoscyphus subarticulatus.

2/7. E. Kerguelen; 4.4.74; 49°33,2’S 70°47,1’E; 130 m: Symplectoscyphus subarticulatus.

3/10. E. Kerguelen; 5.4.74; 49°28,5’S 71°52,8’E; 650 m: Eudendrium tottoni, Modeeria

rotunda, Opercularella belgicae, Lafoea dumosa, Plumularia insignis.

3/11. E. Kerguelen; 5.4.74; 49°25,4’S 71°51,7’E; 620-650 m: Eudendrium tottoni, Modeeria

rotunda, Filellum serratum, Lafoea dumosa, Symplectoscyphus subdichotomus, Plumu-

laria insignis.

7/22. N.E. Heard; 7.4.74; 52°12,7’S 75°38,4’E; 525-560 m: Staurotheca dichotoma, Plumu-

laria insignis.

8/24. N. Heard; 8.4.74; 52°58,0’S 73°42,0’E; 123 m: Campanularia norvegiae, Staurotheca

dichotoma, Sertularella picta, Symplectoscyphus elongatus, S. mawsoni.

8/25. N.E. Heard; 8.4.74; 52°59,4’S 73°38,0’E; 90 m: Modeeria rotunda, Staurotheca —

dichotoma, Symplectoscyphus elongatus, S. subdichotomus, Schizotricha unifurcata.

9/26. Heard, Atlas Cove; 8.4.74; 15-20 m: Sertularella picta, Schizotricha unifurcata.

9/27. Heard, Atlas Cove; 8.4.74; 6 m: Sertularella picta.

10/30. S.S.E. MacDonald; 9.4.74; 53°06,7’S 72°50,1’E; 255 m: Modeeria rotunda, Symplecto-

scyphus mawsoni, Schizotricha unifurcata.

11/31. W. Heard; 9.4.74; 53°20,3’S 72°29,2’E; 790 m: Staurotheca dichotoma.

14/44. W. Kerguelen; 13.4.74; 49°48,4’S 64°57,9’E; 250 m: Symplectoscyphus plectilis,

S. subdichotomus.

14/45. W. Kerguelen; 13.4.74; 49°45,8’S 64°50,6’E; 262 m: Hydrodendron arborea, Stauro-

theca dichotoma, Symplectoscyphus subarticulatus.

17/50. N.W. Kerguelen; 14.4.74; 47°24,9’S 66°04,0’E; 585 m: Kirchenpaueria triangulata,

Plumularia insignis.

18/52. N.W. Kerguelen; 15.4.74; 47°42,2’S 68°07,1’E; 243 m: Modeeria rotunda, Plumu-

laria insignis.

21/57. N.E. Kerguelen; 15.4.74; 48°29,7’S 70°55,4’E; 345-360 m: Zygophylax crozetensis,

Staurotheca dichotoma, Symplectoscyphus subdichotomus, Plumularia insignis.

22/58. N.E. Kerguelen; 16.4.74; 48°58,5’S 70°51,1’E; 90-105 m: Eudendrium rameum,

Halecium tenellum, Zygophylax crozetensis, Staurotheca dichotoma, Sertularella picta,

Symplectoscyphus subdichotomus, Schizotricha unifurcata.

23/59. S.E. Kerguelen; 16.4.74; 49°59,2’S 70°01,9’E; 158 m: Eudendrium rameum, Stauro-

theca dichotoma, Plumularia insignis, Schizotricha unifurcata.

24/61. S.E. Kerguelen; 17.4.74; 50°10,7’S 69°48,7’E; 195 m: Tulpa diverticulata, Stauro-

theca dichotoma, Plumularia insignis.

26/63. Crozet, chenal des Orques; 20.4.74; 46°21,5’S 51°55’E; 230 m: Modeeria rotunda,

Phialel. chilensis, Halecium dufresneae, Filellum serratum, Zygophylax crozetensis,

Campanularia sp., Staurotheca dichotoma, Symplectoscyphus curvatus,

S. subdichotomus.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 3

26/64. Crozet, chenal des Orques; 20.4.74; 46°24’S 51°59’E; 180 m: Eudendrium rameum,

Modeeria rotunda, Phialella chilensis, Hydrodendron arborea, Grammaria abietina,

Halisiphonia ?nana, Hebella striata, Zygophylax crozetensis, Campanularia sp., Tulpa

diverticulata, Staurotheca antarctica, S. dichotoma, Sertularella picta, Symplecto-

scyphus curvatus, S. elongatus, S. subdichotomus, Oswaldella bifurca, Plumularia

insignis.

26/65. Crozet, chenal des Orques; 21.4.74; 46°23,3’S 51°58,3’E; 165 m: Symplectoscyphus

subdichotomus.

28/71. Entre Possession et Cochons; 22.4.74; 46°18,1’S 51°29’E; 400 m: Halecium deli-

catulum, H. jaederholmi, Sertularella picta, Symplectoscyphus subdichotomus.

30/73. Entre Possession et Cochons; 22.4.74; 46°02,3’S 50°50,2’E; 187 m: Tulpa diverticu-

lata, Staurotheca antarctica, S. dichotoma, Symplectoscyphus subdichotomus, Plumu-

laria insignis.

31/74. Entre Possession et Cochons; 22.4.74; 45°57,2’S 50°32,8’E; 110 m: Modeeria rotunda,

Phialella chilensis, Halecium jaederholmi, Hydrodendron arborea, Grammaria abietina,

Zygophylax crozetensis, Silicularia rosea, Tulpa diverticulata, Staurotheca antarctica,

S. dichotoma, Sertularella geodiae, S. picta, Symplectoscyphus subdichotomus.

SYSTEMATIC SECTION

Family Eudendriidae

Eudendrium rameum (Pallas, 1766)

Eudendrium rameum: Allman, 1888: 4, pl. 2 (figs 1-2). Vervoort, 1946: 150, figs 60-61.

Vervoort, 1972a: 21.

Stations

22/58-C; 23/59-E; 26/64-P. All fertile.

Description

Stiff, bushy colonies with thick, fascicled stems reaching a maximum height

of 120 mm and a diameter of 5 mm at base, branching irregularly and in all

planes. Maximum diameter of unfascicled stems 0,26 mm. Perisarc annulated

on origin of branches, partly or completely on hydranth pedicels, and at other

irregular intervals, brown in colour. Hydranth with about 20 tentacles, with

abundant large nematocysts on hypostome and in nettle ring.

Male gonophores borne on non-atrophied hydranths, one- to two-

chambered. Female gonophores borne on non-atrophied hydranths, young

ones with unbranched spadix.

Large nematocysts: microbasic euryteles very similar to those illustrated

by Weill (1934, fig. 66); 19,8 x 9,0 — 25,8 x 10,8 um, discharged shaft 20,4-24,0

pm.

Remarks

Totton (1930) considered that all Antarctic material of Eudendrium should

be included in one species: E. antarcticum Totton = E. tottoni Stechow.

Totton included Jaderholm’s (1905) material from South Georgia doubtfully

in this species; and it presumably follows that Allman’s (1888) material from

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kerguelen Island should be included too, since Jaderholm said that the two

samples were very similar.

In the present collection, however, there are almost certainly two species

of Eudendrium, a delicate form with a flexuous stem (E. fottoni) and a stiff

bushy form which the author has assigned to E. rameum. Unfortunately the

former is not well enough preserved for examination of nematocysts, which

must provide the final evidence, but the latter possesses abundant large micro-

basic euryteles, a type which according to J. Bouillon (personal communication)

is characteristic of E. rameum. The bushy colony is also very similar to Ver-

voort’s (1946) illustration of E. rameum.

Eudendrium tottoni Stechow, 1932

Eudendrium ramosum: Vanhoffen, 1910: 288, fig. 13.

Eudendrium antarcticum Totton, 1930: 140.

Eudendrium tottoni: Rees & Thursfield, 1965: 62.

Stations

3/10-E; 3/11—A. (Both fertile.)

Description

Delicate colonies reaching a height of about 16 mm. Stems unfascicled

or weakly fascicled at base, flexuous, difficult to distinguish from hydrorhiza,

branching irregularly, about 0,08 mm in diameter. Groups of annulations —

present on origins of branches and rarely at other points. Hydranths poorly

preserved.

Female gonophores with unbranched spadix arching over a single egg.

Male gonophores one- to two-chambered.

Remarks

The hydranths in this species are too poorly preserved to distinguish

structure or number of tentacles, though the gonophores have survived. The

tentacles of the fertile hydranths appear to be persistent. E. tottoni is known

from a number of localities in the Antarctic.

Family Campanulinidae

Modeeria rotunda (Quoy & Gaimard, 1827)

Stegopoma fastigiatum: Vervoort, 1972a: 42.

Modeeria rotunda: Edwards, 1973: 573, figs 1-3. Millard, 1975: 137, fig. 45A.

Stations

3/10—-C; 3/11—G; 8/25-D; 10/30-C; 18/52-B; 26/63-B; 26/64—G; 31/74-M.

Description

Many infertile colonies epizootic on other hydroids. Hydrotheca and

pedicel extremely variable in size.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 5

Remarks

This is a new record for this area. The species has, however, been recorded

from the south-western Atlantic at a very similar latitude (Vervoort 1972a).

Opercularella belgicae (Hartlaub, 1904)

Figure 1B

Campanulina belgicae Hartlaub, 1904: 10, pl. 1 (figs 8-9). Vanhoffen, 1910: 308, fig. 28.

Opercularella belgicae: Leloup, 1974: 4, fig. 2.

Opercularella sp. Vervoort, 1972a: 42, fig. 11b, c.

Station

3/10-F.

Description

A sparse colony without gonothecae growing on a polyzoan. Colony

stolonic. Pedicels of very variable length, with 2-4 spiral annulations at base,

merging smoothly into hydrotheca. Hydrotheca deep, widest at opercular base,

with about eight opercular segments and a delicate diaphragm.

Measurements (mm)

Total height, pedicel + hydrotheca .. - 1,00-6,90

Hydrotheca, height from diaphragm to tip oF eseetge a 0,46-0,57

maximum diameter As ie + Ae re 0,16—-0,20

Remarks

Vervoort’s material (1972a: Opercularella sp.) from Magellan is very

similar to the present material, though with shorter pedicels. It appears to fall

within the range of variation of O. belgicae.

The generic name of this species is still provisional pending knowledge of

the gonophore.

Phialella chilensis (Hartlaub, 1905)

Figure 1A

Campanulina chilensis Hartlaub, 1905: 589, fig. L?, M?, N?. Naumov & Stepaniants, 1962:

76, fig. 3. Leloup, 1974: 3, fig. 1.

Phialella chilensis: Vervoort 1972a: 38, fig. 10.

Stations

26/63—H; 26/64-Q; 31/74_N.

Description

Slender stems, a few lightly fascicled at base, branching sympodially and

reaching a maximum height of 10,8 mm. Stem closely annulated in basal half

of each internode and smooth in distal half, as illustrated by Vervoort (1972a)

rather than by Hartlaub (1905). Hydrothecal pedicels completely annulated

or with a smooth area in the centre of the longer ones. Hydrotheca as in previous

descriptions. Gonothecae absent.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1.

Phialella chilensis (Hartlaub). A. Part of stem with hydrothecae.

Opercularella belgicae (Hartlaub). B. hydrothecae.

Halecium delicatulum Coughtrey. C. Hydrophore. D. Gonophore.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES a

Measurements (mm)

Hydrotheca, height from diaphragm to tip of operculum .. 0,21-0,29

maximum diameter oF a ae B ae 0,11-0,14

Remarks

This species is well known from the Antarctic and Subantarctic, especially

from the South American area. This is the first record from the Kerguelen

region.

Family Haleciidae

Halecium delicatulum Coughtrey, 1876

Figure 1C-—D

Halecium delicatulum: Ralph, 1958: 334, figs 1le, h-n, 12a—p. Naumov & Stepaniants, 1962:

94, fig. 16. Vervoort, 19725: 341, fig. 2a.

Halecium antarcticum Vanhoffen, 1910: 317, fig. 34. Billard, 1914: 7, fig. 5. Totton, 1930:

144, fig. 4.

Station

28/71-B.

Description

A number of stems growing on a gorgonian and reaching 32 mm in height.

Stem fascicled and stiff, branching irregularly. Internodes separated by oblique

nodes sloping in alternate directions, with one or two annulations at the base

of each. Primary hydrophores free from stem, smooth, sometimes containing

a pseudodiaphragm. Secondary hydrophores annulated, at least in basal region.

Hydrothecal margin strongly flared outward.

Female gonothecae present, compressed, oval in broad view, with terminal

aperture surrounded by an internal collar of perisarc, containing 6-10 eggs.

Measurements (mm)

Internode length .. a i ed a br se 0,53-0,72

Hydrotheca, depth to diaphragm m oe . 0,03-0,06

diameter at margin ee a sia sis sat 0,17-0,20

Gonotheca, length ae 7. a vs ae 0,71-1,12

maximum diameter ee oe es aes as 0,35—0,52

Remarks

The author has bowed to the opinion of Naumov & Stepaniants (1962)

and Vervoort (19725) in including H. antarcticum in the synonymy of H. deli-

catulum, although she feels that there is a distinct Antarctic form in which the

female gonotheca is smaller and more slender and without the ‘ears’ of typical

H. delicatulum as illustrated by Millard (1975). The gonothecae in the present

material are similar to those illustrated by Totton (1930) and Vervoort (19725).

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

H. delicatulum is well known from the Antarctic, particularly in the area

south of America, and was reported from Marion Island by Allman (1888,

as H. flexile).

Halecium dufresneae sp. nov.

Figure 2A—D

Material

Holotype: from Station 26/63-A. Part in Muséum National d’Histoire

Naturelle, Paris, and part in the South African Museum (SAM-H2785).

Description of holotype

One colony, with a thick rootstock flattened below and a thick fascicled

stem 245 mm in height. Stem branching and rebranching irregularly but mainly

in one plane. Final branches unfascicled, divided into internodes by oblique

nodes sloping in alternate directions, each internode bearing a hydrotheca

from an apophysis at distal end.

Primary hydrophore not clearly demarcated from apophysis, with a stout

pseudodiaphragm in base, adnate to internode almost to diaphragm. Secondary

hydrophores rather rare in this material, smooth and not constricted after

origin, asymmetrical with adcauline wall longer than abcauline, with or without

a pseudodiaphragm. Hydrotheca shallow, with adcauline surface free from _

stem, with margin usually everted.

Gonothecae borne on sides of hydrophores, the two series together forming

a single row on the anterior surface of the stem. Gonotheca flattened, in broad

view widening to the truncated distal end, with one or two large embryos

discharged into a marsupium which may be as large as, or larger than, itself.

Measurements (mm)

Internode, length Ag ce cP <. = ie 0,51-—1,00

Hydrotheca, depth from diaphragm .. Pn i se 0,03-0,07

diameter at margin ae os ch es a 0,19-0,23

Gonotheca, depth =f = ue tA - a 0,94-1,14

maximum diameter ae es si if te 0,38—0,64

Marsupium, depth co se ie . =e 52 0,66-1,14

Remarks

Female gonothecae with external marsupia are known from two other

species of Halecium, namely H. pallens Jaderholm, 1904, from South Georgia,

and H. marsupiale Bergh, 1887, from the Arctic.

The author feels that in these two, as in the present species, the marsupium

is an important diagnostic character, and she cannot agree with Naumov &

Stepaniants (1962) and Vervoort (19725) who include H. pallens in the synonymy

of H. delicatulum. A marsupium has never been seen in the latter; moreover

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 9

Fig.2:

Halecium dufresneae sp. nov. from the holotype. A-—C. Parts of stem with hydrophores:

C with secondary hydrophore. D. Gonophore with two embryos in marsupium.

Hydrodendron arborea (Allman). E. part of glomulus with female gonophores. F. Part of stem

with hydrophores and a giant nematotheca on bottom left. G. normal nematothecae.

Scale in mm/10.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

the shape of the mature female gonotheca is different, being wider in broad

view and ‘eared’ in H. delicatulum.

It follows that it is not possible to identify most of the species of Halecium

from sterile material alone.

The present species differs from H. pallens in the shorter pedicel of the

primary hydrophore and in the marsupium, which is almost as large as, or

larger than, the gonotheca and contains only one or two eggs.

Halecium jaederholmi Vervoort, 1972

Figure 3A—B

Halecium jaederholmi Vervoort, 1972a: 21, fig. 2 (synonymy).

Stations

28/71-A; 31/74-A.

Description

The first colony (28/71—A) is medium-brown in colour, with thick fascicled

stems 4 mm in diameter at the base and 160 mm in maximum height, branching

irregularly and in all planes. The second colony (31/74-A) is dark brown and

has an even thicker main stem (8 mm diameter at base) and is more stiff and "

rigid in habit. It reaches a height of 140 mm.

In both colonies the unfascicled parts together with their internodes and

hydrophores have a structure exactly like that illustrated by Vervoort (1972a)

except that no pseudodiaphragmata are present nor any secondary hydrophores.

The hydrothecae are very shallow and adnate to the internodes.

Typical female gonothecae are present in the first colony and contain up

to six larvae; some of the gonothecae are damaged and broken off transversely

just beyond the aperture. In the second colony all the gonothecae (female) are

damaged in this way.

Measurements (mm)

Internode, length ae he a ae Ea 0,50-0,94

Hydrotheca, depth to diaphragm a o is aoe 0,03-0,04

diameter at margin on < a os a 0,17-0,23

Gonotheca, length Re i ae ac sa ae 1,49-1,85

maximum diameter. . a ae 3 fs a 0,59-0,71

Remarks

The distribution of this species has been summarized by Vervoort; it occurs

in the Antarctic and Subantarctic, but this is the first record from the Kerguelen

area.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 11

Halecium tenellum Hincks, 1861

Halecium tenellum: Millard, 1975: 156, fig. 5SOF—L. Cornelius, 1975: 409, fig. 12.

Station

22/58—D.

Description

Colony epizootic on a polyzoan, with sparsely branched stems reaching

8 mm in height, and with female gonophores.

Remarks

This cosmopolitan species has been reported from the south-western

Atlantic and from Antarctica. This is the first record from the Kerguelen area.

Hydrodendron arborea (Allman, 1888)

Figure 2E-G

Halecium robustum Allman, 1888: 10. Vervoort, 1972a: 23.

Halecium arboreum Allman, 1888: pl. 4. Hickson & Gravely, 1907: 27, pl. 4 (figs 27-29).

Ophiodissa arborea: Totton, 1930: 142, fig. 2a. Vervoort, 1972a: 25.

Stations

14/45-B; 26/64-N; 31/7-J. All colonies fertile.

Description

Shrubby colonies with thick, fascicled stems reaching a maximum height

of 105 mm. Branching very irregular; final branches divided into internodes

by oblique nodes slanting in alternate directions. Each internode bearing one

hydrophore on an apophysis at distal end. Primary hydrophore not distinctly

demarcated from apophysis, adnate to internode almost, or completely, to level

of diaphragm.

Primary hydrotheca with free adcauline wall, with margin sometimes

everted on adcauline side. Secondary hydrophores usually quite symmetrical,

with no constriction after origin.

Nematothecae goblet-shaped, normally one on each internode on side

opposite to hydrotheca, but often broken off leaving only the aperture, and

sometimes absent without trace; also present on tubes of gonosome.

Occasionally giant nematothecae present, about five times the size of normal

ones.

Gonothecae borne in glomulus (term from Naumov 1960: 442), strongly

curved, with tubular neck, containing eggs or planulae. Glomuli forming large

masses about 15 mm in diameter.

Measurements (mm)

Internode, length “a fie 43 a! . is 0,42-0,93

Hydrotheca, depth to diaphragm of ae ae d's 0,04—0,07

diameter at margin we ae a Re ot 0,21-0,28

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm)

Normal nematotheca, depth .. a ie ae i 0,06-0,10

Giant nematotheca, depth i o ee ue a 0,44-0,48

Gonotheca, maximum length .. a : oe a 0,78-1,02

maximum diameter nts oe 5A . os 0,47—0,60

Remarks

This species is widely distributed in the Antarctic; its type locality is

Kerguelen.

As other authors have remarked, the nematothecae are variable in occur-

rence and not invariably present on each internode. The’ giant nematothecae

have not been recorded before. The regenerated (secondary) hydrophores are

very distinctive and clearly distinguish the species from forms such as H. beanii.

Family Lafoeidae

Filellum serratum (Clarke, 1879)

Filellum serratum: Vervoort, 1972a: 51, fig. 14a—b. Millard, 1975: 178, fig. S9YA—C.

Stations

3/11-F; 26/63-J: infertile colonies epizootic on polyzoa and other hydroids.

Remarks

This cosmopolitan species has not yet been reported from the Kerguelen

area, though it is known from New Zealand and South America.

Grammaria abietina (Sars, 1850)

Figure 3D-E

Grammaria abietina: Cornelius, 1975: 382, fig. 3 (synonymy).

Stations

26/64-K; 31/74-L.

Description

Two colonies with thick, fascicled stems reaching a maximum height of

83 mm and branching in an alternate, subopposite or opposite manner in one

plane. Hydrothecae forming six longitudinal rows on stem and branches, but

not strictly regular in arrangement.

Hydrotheca curved outwards to a varying degree, with margin everted

and either parallel or oblique to axis of stem; free part 0,34-0,51 mm in length;

0,28-0,35 mm in diameter at margin.

A single coppinia present, about 15 mm iong and 5 mm wide. Gonothecae

tightly adpressed, flask-shaped, with terminal aperture on a short tubular

neck. Accessory tubes about three times length of gonothecae, very strongly

curved.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 13

Bue

’

ee ee ee ee ee oe |

Fig. 3.

Halecium jaederholmi Vervoort. A. Part of stem with two hydrophores. B. Gonophore.

Halisiphonia ?nana Stechow. C. Hydrothecae.

Grammaria abietina (Sars). D. Part of stem with six rows of hydrothecae. E. Part of t.s. of

coppinia, showing gonothecae, protective structures, and arrangement of thecal bases in

three’s in centre of stem.

Scale in mm/10.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The synonymy of this species was recently revised by Cornelius (1975),

who has included as synonyms G. stentor Allman, 1888, G. magellanica Allman,

1888, and G. insignis Allman, 1888.

The species has a bipolar distribution and is widely known from the Arctic

and Subarctic as well as from the Antarctic and Subantarctic. The type locality

of G. stentor is Kerguelen Island.

Halisiphonia ?nana Stechow, 1921

Figure 3C

Halisiphonia nana: Stechow, 1925: 452, fig. 22.

Station

26/64-S.

Description

An infertile colony epizootic on Sertularella picta. Pedicel arising from

hydrorhiza and merging smoothly into hydrotheca which widens evenly to

margin. No diaphragm or annular thecal thickening. Margin sometimes slightly

everted.

Measurements (mm)

Pedicel + hydrotheca, height .. ab iby Es ba 0,64—0,95

Pedicel, diameter at base we ee a me ae 0,04—0,12

Hydrotheca, diameter at margin ne ae Ee - 0,14-0,23

Remarks

This material is assigned with some doubt to H. nana. The dimensions are

more or less in agreement with those of Stechow, and are too small for H. mega-

lotheca Allman, but the pedicel is often rather wide at the base and the thecal

margin may be everted. H. nana has been reported only once, from east of

Bouvet Island; the gonothecae are unknown.

Hebella striata Allman, 1888

Hebella striata Allman, 1888: 30, pl. 15 (figs 3, 3a). Vanh6offen, 1910: 313. Vervoort, 1972a:

62, fig. 17b-c.

Station

26/64-R.

Description

An infertile colony epizootic on Grammaria abietina. Details exactly as in

Vervoort’s material, but dimensions a little larger.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES iS

Measurements (mm)

Pedicel, length .. me cS ce i a es 0,44-0,87

Hydrotheca, depth we os ce ee na eS 0,77-1,10

diameter a Ra ae ba a os 0,24-0,31

aks

This species is known mainly from the Subantarctic near South America,

but was reported from Kerguelen by Vanhdffen (1910).

Lafoea dumosa (Fleming, 1820)

Lafoea fruticosa: Millard, 1975: 187, fig. 61A—F.

Lafoea dumosa: Cornelius, 1975: 385, fig. 4 (synonymy).

Stations

3/10—B; 3/11-B. Both colonies infertile.

Description

Stems flexuous, reaching a maximum height of 78 mm, with many

fragments.

Remarks

The author has followed Cornelius (1975) in uniting L. fruticosa (M. Sars,

1851), L. gracillima (Alder, 1856) and L. dumosa under the last name.

Zygophylax crozetensis sp. nov.

Figure 4

Material

Holotype: from station 26/64-B. Part in Muséum National d’Histoire

Naturelle, Paris, and part in the South African Museum (SAM-H2779).

Other stations: 2/6-A (infertile); 21/56—-C (infertile); 22/58—B (male);

26/23-D (fertile); 31/74-F (fertile).

Description of holotype

A large, branching colony 220 mm in height. Rootstock a large mass of

interwoven fibres flattened below and reaching 45 mm in diameter. Stem

fascicled and thick, giving off thick primary branches in an irregular fashion

and mainly in one plane, these branches giving off secondary and tertiary

branches which are strictly in one plane. Final branches (hydrocladia) sub-

opposite in arrangement, lightly fascicled or unfascicled. Hydrocladia and

final terminations of other branches segmented where exposed, with one or two

hydrothecae to an internode, but with many irregular regenerative nodes.

Hydrothecae normally alternate in arrangement with the two rows more or less

in one plane, and this arrangement characteristic of unfascicled hydrocladia

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

a is B ‘<a

Fig. 4.

Z. ygophylax crozetensis sp. nov. from the holotype. A. Part of stem showing origins of hydro-

cladia. B-C. Hydrothecae and nematothecae. D. Surface view of coppinia showing hoods

of gonothecae and protective structures. E. Gonothecae from t.s.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES lg

and smaller branches, but in fascicled branches supplementary hydrothecae

may arise from peripheral tubes in any plane and in any position imparting a

very irregular appearance to the whole.

Hydrothecae tubular, widening to margin which is slightly everted, curved

outwards, those in axils of hydrocladia less curved and practically straight.

Diaphragm distinct and oblique.

Nematothecae two-chambered, with a short basal chamber and a long

tubular distal chamber, one or two on each hydrotheca-bearing apophysis,

that/those of the axillary hydrotheca shifted on to the apophysis next to it,

various irregular supplementary ones present on peripheral tubes of stem and

branches.

Coppiniae numerous and clothing most of the larger branches completely

to a width of 3-5 mm, so that it is not possible to distinguish one from another.

Coppinia consisting of adpressed gonothecae with between them irregular

branching structures bearing nematothecae and rarely hydrothecae as well.

Gonotheca slender, widening distally to top of adnate part, then narrowing

to a free, pointed, and sometimes curved, hood or horn bearing an aperture on

one side. Sex not determinable.

Measurements of holotype (mm)

Hydrothecal pedicel, height .. e se ie is 0,06-0,13

Hydrotheca, height abcauline .. m, oe ve : 0,32-0,38

diameter at margin ye os fe os S 0,16—0,20

Pedicel + hydrotheca, height .. in ¥ Ms bes 0,40-0,47

Nematotheca, height .. = oe & ye — 0,09-0,19

diameter at margin m we a Ee ce 0,05-0,08

Remarks

In the four fertile samples all the gonothecae are of the same shape. Most,

however, are empty. Contents are present in 22/58—B only, and these are male.

The hydrothecae of this species are similar to those of Z. africana Stechow,

1923, and a number of related species, and particularly to ‘Lafoea’ halecioides

Allman, 1874 (= ‘Lafoea’ pinnata Sars, 1873). The last-mentioned, however,

has an open coppinia with unfused gonothecae.

Nutting (1905) described and figured a closed coppinia with one-horned

gonothecae for a species which he called Lictorella halecioides. This is apparently

a synonym for Zygophylax antipathes (Lamarck, 1816), and not for Z. pinnata.

It is probably this account which prompted Totton (1930: 166) to mention

‘hooded apertures’ for Z. antipathes and not the account of Trebilcock (1928)

who does not mention the species. However, Z. antipathes has somewhat

broader hydrothecae than the present material, and the gonothecae as depicted

by Nutting have terminal apertures.

It appears that a number of southern hemisphere species of Zygophylax

have similar trophosomes but different gonosomes. The gonothecae of the

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

present species are almost exactly like those of Cryptolaria pectinata (Allman,

1888).

Family Campanulariidae

Campanularia norvegiae Broch, 1948

Figure 5A-E

Campanularia norvegiae Broch, 1948: 16, fig. 5.

Station

8/24-E.

Description

A dense fertile colony growing on an empty tube. Hydrorhiza reticular.

Pedicel sometimes spirally grooved throughout, but very variable and often

smooth in certain areas, always with a segment of lesser diameter at distal end.

Hydrotheca funnel-shaped, oval in section and thickened, more so at the narrow

ends; thickened at, or just below, margin.

Gonotheca irregularly bottle-shaped, with a short annulated pedicel,

narrowing distally to terminal aperture, often twisted or irregularly corrugated,

circular in section, empty or containing planulae.

Measurements (mm)

Pedicel + hydrotheca, length .. xn ne a a 1,70—8,50

Hydrotheca, depth - oe oa ; 0,48-0,72

maximum diameter (broad view) .. oe = ee 0,38-0,56

Gonothecal pedicel, depth pe oe Le 3 7 0,24-0,32

Gonotheca, depth - ae as leg 2 ss 1,58—2,28

maximum diameter a > Le 433 sa 0,52-0,65

Remarks

This species is distinguished from the closely related C. integra Mac-

Gillivray, 1842, by the shape of the gonotheca. Since the gonothecae contain

planulae it is obvious that no medusoid is released.

C. norvegiae was originally described from South Georgia.

Campanularia sp.

Figure 5F

Stations

26/63-G; 26/64-R.

Description

Infertile colonies epizootic on other hydroids. Pedicels of variable length,

with one terminal spherule of lesser diameter, generally roughly corrugated

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 19

Fig. 5.

Campanularia norvegiae Broch. A-C. Hydrothecae. D-E. Gonophores.

Campanularia sp. F. Hydrothecae.

Tulpa diverticulata Totton. G. Gonotheca. H. Hydrotheca.

Scale in mm/10.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

immediately below this and at base. Hydrotheca deep, with parallel sides, with

an annular thickening near base, with 8-10 slender, bluntly rounded marginal

teeth.

Measurements (mm)

Pedicel, length .. oe Ate on * we is 0,48-1,07

Hydrotheca, depth fs ie ss a ah we 0,60—-0,90

diameter at margin Ke eS bs ave eo: 0,25—0,33

diameter/depth a: oe - e. i. mc 0,34-0,51

Remarks

In the absence of gonophores this species cannot be definitely diagnosed.

The two most commonly reported species of Campanularia from the Antarctic

are C. hicksoni Totton, 1930, and C. tincta Hincks, 1861. (C. cylindrica Allman,

1876, from Kerguelen is probably a synonym of the former.) These two species

are distinguished mainly by their gonothecae—smooth in the former, annulated

in the latter; however, none of the Antarctic material attributed to C. tincta has

yet been found with gonothecae and possibly only one species is involved.

Silicularia rosea Meyen, 1834

Silicularia rosea: Ralph, 1956: 293. Millard, 1968: 259. Millard, 1971: 405.

Silicularia bilabiata: Ralph, 1956: 285, figs 2-3. Ralph, 1957: 842.

Station

31/74-C.

Description

A rich, fertile colony on brown alga.

Remarks

S. rosea is known from Crozet and Kerguelen Islands and many other

localities in the Antarctic and Subantarctic. The distribution was summarized

by Ralph (1957).

Tulpa diverticulata Totton, 1930

Figure 5G-H

Tulpa diverticulata Totton, 1930: 145, fig. 5. Ralph, 1957: 844, fig. 7l-n.

Campanularia diverticulata: Naumov & Stepaniants, 1962: 72.

Stations

24/61-C; 26/64-H; 30/73-A; 31/74-B.

Description

Hydrorhiza creeping on other hydroids, polyzoa and worm tubes, but

often becoming free to form a tangled mass. Hydrothecal pedicels of very

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 21

variable length, unsegmented, though often with regeneration nodes. Hydro-

thecae as in previous descriptions; regenerated margins sometimes present.

Gonothecae arising from hydrorhiza on very short pedicels, irregularly oval,

with terminal aperture on long narrow neck.

Measurements (mm)

Hydrothecal pedicel, length .. A a i i! 0,60—2,80

Hydrotheca, length 7: ne a a Se nfs 3,00-4,10

diameter at margin ee a eS ie - 0,95-1,35

Gonotheca, length including pedicel .. i sd ef 1,85—3,45

maximum diameter oe ne oe s a 0,89-1,50

Remarks

Some of these colonies are very rich, and several stolons may run together

or twine round each other simulating a fascicled stem. However, the pedicels

always arise at right angles, and there is never any question of an erect branching

stem with oblique pedicels as in 7. tulipifera (Allman, 1888). It is clear that

these are two separate species.

Very fine cross-striations may be present on some of the hydrothecae and

gonothecae in certain colonies.

Young gonothecae are cone-shaped and similar to those described by

Fraser (1944) for T. speciosa (Clarke).

T. diverticulata is so far known only from New Zealand and to the south

of it. It is a new record for Kerguelen and Crozet Islands. The closely related

T. tulipifera occurs in Heard Island (Allman 1888) and in the south-western

Atlantic (Vervoort 1972a).

Family Syntheciidae

Staurotheca antarctica Hartlaub, 1904

Staurotheca antarctica Hartlaub, 1904: 16, pl. 1 (fig. 4), pl. 2 (fig. 4). Totton, 1930: 176, 178,

fig. 28, pl. 2 (fig. 6). Vervoort, 1972a: 198, figs 67, 68b.

Stations

26/64-M (fertile); 30/73-B (fertile); 31/74-D.

Description

Luxurious colonies with unfascicled stems which branch dichotomously

and reunite to form an elaborate reticulum in one plane. Nodes irregular.

Hydrothecae generally arranged in decussate pairs, but rarely in groups of three,

and with many irregularities.

Male and female gonothecae present, as in previous descriptions.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm, all without regenerations)

Hydrotheca, depth abcauline

depth adcauline, adnate part

depth adcauline, free part ..

adnate part/total adcauline depth

diameter at mouth

Gonotheca, male, length

maximum diameter

Gonotheca, female, length

maximum diameter (including spur)

Remarks

0,34—0,44

0,38-0,50

0,10—0,20

0,66-0,83

0,17-0,20

0,84-1,12

0,55-0,66

1,30-1,48

0,98-1,21

There is little to add to the description of this well-known Antarctic and

Subantarctic species. Vervoort (1972a) has summarized the distribution.

Staurotheca dichotoma Allman, 1888

Staurotheca dichotoma Allman, 1888: 76, pl. 36 (fig. 1). Billard, 1910: 27, figs lus 12. Billard,

1914 >-15, ne. 9 votton, 1930: 175, 178.tie. 27;

Stations

7/22-B; 8/24-A (fertile: female gonophores); 8/25-B (fertile: male and

female gonophores); 11/31-A; 14/45—A; 21/57-B (fertile: female gonophores); —

22/58—G; 23/59-A (fertile: male gonophores); 24/61-B;

-C; 26/64-L;

30/73-C (fertile: female gonophores); 31/74—-E (fertile: female gonophores).

Description

Luxuriant colonies. Stems thick, fascicled at base in larger colonies,

branching in a subdichotomous manner, reaching a maximum height of 130 mm.

Hydrothecae generally 3 to a whorl (and forming 6 longitudinal rows), but

some of the thickest stems have 4 to a whorl (and 8 longitudinal rows) and the

thinner terminal branches only 2 (4 longitudinal rows). Hydrothecae adnate

for almost their entire length. Male and female gonothecae as described by

Billard (1910) and Totton (1930).

Measurements 8/25—B (mm)

Hydrotheca, depth abcauline .. ay ah ae er 0,59-0,67

depth, adcauline, adnate part ri a » me 0,78—-0,96

depth adcauline, free part . ; Me i bo 0,03-0,08

adnate part/total adcauline tent os as Bo 0,91-0,97

diameter at margin i: a me e es 0,27-0,32

Gonotheca, female, length a Ay a% Fe oa 1,49-2,17

maximum diameter a oe a ay %y 1,11-1,29

Gonotheca, male, length oe so oe a ae 1,23-1,60

maximum diameter ae 9 ai m1 i 0,78—1,02

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 23

Remarks

As previous authors have remarked, S. dichotoma is clearly distinguished

from S. antarctica by the structure of the female gonotheca and by its greater

dimensions. The species is widely distributed in the Antarctic and Subantarctic.

Its type locality is Marion Island, not far from the present localities.

Family Sertulariidae

Sertularella geodiae Totton, 1930

Figure 6E-F

Sertularella geodiae Totton, 1930: 196, fig. 43, pl. 3 (figs 7-8). E. A. Briggs, 1939: 37. Ralph,

1961: 831, fig. 24c, g. Vervoort, 1972a: 120, fig. 37.

Station

31/74-K.

Description

A number of infertile stems and fragments, reaching a maximum height

of 40 mm. Stem stiff, fascicled and straight at base, geniculate in terminal

regions only, branching rather sparsely and in one plane. Branches unfascicled,

arising below hydrothecae and, when several are present, below every third

hydrotheca.

Hydrotheca adnate for about half adcauline wall, bent outwards (a perpen-

dicular dropped through centre of margin passes through adcauline wall), with

a few undulations on free part of adcauline wall, with wide mouth. No internal

teeth.

Measurements (mm)

Internode length .. mn re io e ie ode 1,01-1,52

diameter at node .. xe yd ee oe ae 0,20-0,32

Hydrotheca, length abcauline .. ser o Se ais 0,66—0,77

length adcauline, adnate part ne te on “i 0,49-0,61

length adcauline, free part a Se £e ae 0,46-0,60

adnate part/total adcauline length Bi se is 0,46-0,55

diameter at margin ay $3 ae ie ee 0,35-0,40

Remarks

This material is in many ways intermediate between S. geodiae and S. conica

Allman, 1877, considering especially Vervoort’s description of the latter (1972a:

123). However, S. conica has an unfascicled stem and a hydrotheca which

narrows more markedly to the mouth; its distribution is mainly tropical.

The author has therefore assigned this material to S. geodiae in spite of the

absence of gonothecae; this is a species known from New Zealand, Tasmania

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

A, DF eet

B,C le

“TI

Fig. 6.

Sertularella picta (Meyen). A. Part of stem showing origin of a hydrocladium. B—C. Hydro-

thecae. D. Gonotheca.

Sertularella geodiae Totton. E. Hydrotheca. F. Part of stem showing origins of two

hydrocladia.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 25

and the area around the southern tip of South America, but which has not been

reported from Kerguelen.

Sertularella picta (Meyen, 1834)

Figure 6A—D

Sertularella picta: Hartlaub, 1905: 645, fig. L*. Billard, 1922: 106, fig. 2B. Stechow, 1923:

187, fig. B’. Millard, 1971: 405, fig. 6A, B. Vervoort, 1972a: 111, figs 34, 35.

Stations

8/24-D; 9/26-A; 9/27-A; 22/58-E; 26/64-F; 28/71-D; 31/74-H. The

last three fertile.

Description

Stems slender and straggling, branching repeatedly in an irregular fashion

to produce tangled colonies often intertwined with polyzoans and with Symplec-

toscyphus subdichotomus. Stem normally unfascicled, but rarely with one or

two supplementary tubes in basal region, usually with one or two annulations

at base and on origin of branches, geniculate, reaching a maximum height of

38 mm, often reuniting by stolons with other stems. On most stems the two

rows of hydrothecae are in the same plane, but in other stems, or even in other

parts of the same stem, they may be shifted on to the anterior surface so that

the two rows subtend an obtuse angle between them. Branches arising below

hydrothecae, with one or two annulations at origin and a long first internode,

otherwise similar to stem and of same diameter. Internodes slender, very

variable in length, separated by oblique nodes sloping in alternate directions,

usually with an annulation immediately above each node.

Hydrotheca fusiform, adnate for less than half adcauline height, narrowed

below margin. In most hydrothecae the marginal teeth are equally developed

and the margin is perpendicular to the axis (a perpendicular dropped through

centre of margin passes through basal thickening of adcauline wall), but in

some the abcauline marginal tooth is produced so that the margin is tilted

towards the stem (and a perpendicular dropped through centre of margin

passes through the hydrophore or base of the abcauline wall). All intergradations

may occur in one colony. The free part of the adcauline wall usually has two to

four indistinct annulations, but the latter are of variable development—they

may form distinct striations which pass round the sides of the hydrotheca

almost to the abcauline wall, or they may be almost obsolete. Small internal

teeth are present in most hydrothecae, but again the degree of development is

variable; the full complement appears to be three (one median abcauline and

two latero-adcauline), but one or two, or even all three, may be absent. In a

few stems there is no trace of internal teeth.

Gonothecae borne on front of stem immediately below hydrothecae,

obovate, with three to four distinct annulations in distal half, with three or four

blunt marginal spines, with external marsupium. Sex not determinable.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm)

Internode, length a - ot . a ay 0,77-1,34

diameter at node .. As ae R Me a 0,12-0,18

Hydrotheca, length abcauline .. - i vit a 0,55-0,70

length adcauline, adnate part ie a a i 0,24—0,35

length adcauline, free part i op ae ae 0,40-0,51

adnate part/total adcauline length * us fi 0,33-0,44

diameter at margin 2 ae os se e 0,20-0,26

Gonotheca, length a mn ne Se a mY 1,50-2,15

maximum diameter ve ee oF Ms Le 1,30—1,52

Remarks

The identification of this species has been based mainly on Vervoort’s

work (1972a), who also found considerable variability in structure. The form

of the colony, the shape of the hydrotheca and the gonotheca are very similar

to his material.

The species is known from the Falkland/Tierra del Fuego region, and was

also reported from Marion Island (Millard 1971). The Marion Island material

differs in the smooth hydrothecal wall and the greater displacement of the two

rows of hydrothecae.

Symplectoscyphus curvatus (Jaderholm, 1917)

Figure 7A—C

Sertularella curvata Jaderholm, 1917: 9, pl. 1 (figs 11-12).

Symplectoscyphus curvatus: Totton, 1930: 192, fig. 40, pl. 2 (figs 1-3). E. A. Briggs, 1939: 31.

Sertularella curvatus: Naumov & Stepaniants, 1972: 45, fig. 7.

Stations

26/63—-E; 26/64—-C. Both infertile.

Description

Many unrooted, unfascicled stems and fragments yellow-brown in colour

and reaching a maximum length of 44 mm. Most stems pinnate, geniculate,

giving off alternate branches at each elbow and immediately below every third

hydrotheca. On the proximal part of the stem the nodes are not clearly demar-

cated and the hydrothecae are well spaced and do not overlap, but on the

distal part and on the branches the nodes are distinct and the tip on one hydro-

theca overlaps the base of the next. The two rows of hydrothecae and branches

in one plane.

Hydrotheca tubular, large, adnate for less than half adcauline length,

curved outwards, not narrowing to mouth and with diameter in centre region

approximately equal to that at margin, marginal teeth well developed, no

internal teeth. No gonothecae.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 27

4 <p a

cA Vy

Pes te

| Dd

Fig. 7.

Symplectoscyphus curvatus (Jaderholm). A. Distal part of hydrocladium. B. Hydrotheca.

C. Part of stem showing origin of hydrocladium.

Symplectoscyphus elongatus (Jaderholm). D. Part of stem showing origins of hydrocladia.

E. Hydrotheca.

Scale in mm/10.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Symplecto-

scyphus

exsertus

Measurements (mm) 26/64-C (Allman)

Internode length .. si bs Ms o 0,80—1,32 0,43-0,60

Hydrotheca, length abcauline .. ne ao 0,62-0,70 0,28-0,32

length adcauline, adnate part a ms 0,36-0,44 0,14-0,17

length adcauline, free part a oe 0,59-0,68 0,22-0,28

adnate part/total adcauline length ie 0,35-0,41 0,35-0,44

diameter at mouth ee a re 0,41-0,49 0,12-0,14

Remarks

The measurements of the hydrothecae agree with those of Jaderholm and

Totton; they are a little greater than those of Briggs and Naumov & Stepaniants.

S. curvatus shows resemblances to S. exserta (Allman, 1888) from Heard

Island. However, the author has examined type material of the latter species

and finds that it is much smaller in all its dimensions. The measurements are

included above for comparison.

Symplectoscyphus elongatus (Jaderholm, 1904)

Figure 7D-E

Sertularia articulata Allman, 1888: 61, pl. 29 (figs 3, 3a).

Sertularella articulata: Jaderholm, 1905: 29, pl. 11 (fig. 4), pl. 12 (figs 1-3). Naumov & Stepani-

ants, 1972: 42, fig. 5A.

?Sertularella spiralis Hickson & Gravely, 1907: 19, pl. 3 (figs 19-20).

Sertularella elongata: Naumov & Stepaniants, 1962: 80, fig. 5. Naumov & Stepaniants, 1972:

42, fig. 5B.

Symplectoscyphus articulatus: Rees & Thursfield, 1965: 127.

Symplectoscyphus elongatus: Vervoort, 1972a: 136, figs 43, 44a (synonymy).

Stations

8/25—C; 26/64-D; 8/24-C.

Description

Three samples, the first with several infertile stems reaching 67 mm, the

second with numerous infertile stems and fragments reaching a maximum

height of 52 mm, and the third with numerous stems and fragments reaching a

maximum height of 58 mm. None of the stems rooted. Stems golden brown in

colour, unfascicled, geniculate. Internodes of stem very long, up to 3,3 mm,

each bearing a prominent apophysis on distal end on which is seated one

hydrotheca and on each side of it a hydrocladium, the apophyses with their

paired hydrocladia arranged in a loose spiral. Nodes very distinct.

Hydrocladia bearing alternate hydrothecae, and commonly alternate sub-

branches arising opposite every third hydrotheca, the two rows of hydrothecae

and branches usually in one plane but occasionally very slightly displaced

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 29

towards one surface. Internodes of variable length, those of hydrocladium

bearing one or two hydrothecae, those of its sub-branches one.

Hydrothecae tubular, adnate for about half adcauline wall, abcauline wall

straight or curved slightly outwards, with three prominent marginal teeth, but

no internal teeth.

A few empty and damaged gonothecae present in 8/24-C, of the same

shape and size as described by Hickson & Gravely (1907).

Measurements (mm)

Hydrotheca, length abcauline .. bs mn ne im 0,39-0,52

length adcauline, adnate part a: se ns a 0,21-0,35

length adcauline, free part ie se = ce 0,18-0,36

adnate part/total adcauline length sd ae oe 0,37-0,65

diameter at mouth .. oe id i a ne 0,17-0,23

Remarks

This species has been reported many times from Antarctic and Subantarctic

waters; Kerguelen Island is the type locality. The synonymy of the species is

fully discussed by Vervoort (1972a). This synonymy is accepted here, though

with some reservations about the inclusion of Sertularella spiralis Hickson &

Gravely, 1907, a form with internal hydrothecal teeth.

Symplectoscyphus mawsoni Briggs, 1939

Figures 8—9

Non Sertularella biformis Jaderholm, 1905: 28, pl. 11 (figs 1-3).

Sertularella biformis: Totton, 1930: 199, fig. 45, pl. 2 (fig. 8). Naumov & Stepaniants, 1972:

47, fig. 9.

Symplectoscyphus mawsoni E. A. Briggs, 1939: 35, fig. 2, pl. 16 (figs 1-2). Naumov & Stepani-

ants, 1972: 46, fig. 8.

Stations

10/30-A; 8/24-B.

Description

The first colony (10/30—A) consists of two stems 56 mm in maximum

length, and a number of fragments. Basal part of stem lightly fascicled for

16-30 mm, the rest unfascicled and straight. Stem divided into internodes by

distinct oblique nodes sloping in alternate directions. Each internode with a

distal apophysis, alternately on the left and right, which bears a hydrotheca

and two hydrocladia, one on each side of the hydrotheca, the two rows of

hydrothecae in one plane. Many of the hydrocladia and hydrothecae of the

lower half of the stem missing or damaged, those in the upper half better

preserved. Each pair of hydrocladia forming an angle of about 40° with the

stem and about 90° with each other. Hydrocladium divided into regular inter-

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8.

Symplectoscyphus mawsoni Briggs. A. Thick part of stem in side view showing origins of

hydrocladia. B. Hydrotheca. C. Gonotheca with external marsupium. D. Part of hydro-

cladium showing bifurcation. E. A hydrotheca from Totton’s material (BM 1929.10.28.158)

attributed by him to Sertularella biformis, here included in S. mawsoni.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES Sif

nodes by straight nodes, each bearing a hydrotheca near the distal end; dividing

dichotomously at the distal end of the fourth internode, one limb of the dicho-

tomy being slightly thicker than the other; both limbs sometimes dividing

dichotomously once or twice more. The two rows of hydrothecae on the hydro-

cladium not in one plane but forming an acute angle between them. Hydrotheca

completely free from internode, very delicate and often crumpled in basal part,

curved outwards, widening towards aperture, with three well-developed mar-

ginal teeth, one adcauline and two latero-abcauline. No internal teeth. Oper-

culum of three valves. Floor similar to that of Sertularella (Fig. 8A, B, D;

Fig. 9 left).

The second colony (8/24—-B) is in a very fragmentary condition, consisting

mainly of small pieces of hydrocladia. There is, however, a stem of 110 mm

with a few stumps of hydrocladia left; it is fascicled for the first 45 mm. Another

portion of stem 46 mm long bears regular pairs of hydrocladia as in 10/30-A.

The hydrocladia branch up to five times. This colony is fertile and bears

Fig. 9.

Symplectoscyphus mawsoni Briggs. A stem from the Marion-Dufresne collection (Station

10/30—A) on the left, compared with Totton’s specimen attributed to Sertularella biformis

on the right (x 2,5). Photo: D. Gerneke.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

numerous gonothecae on the hydrocladial internodes. Female gonotheca

elongated oboval, with external marsupium, containing one planula (Fig. 8C).

Measurements (mm) 10/30-A 8/24-B

Stem, internode length .. ns a a 2,20-3,95 2,50—4,10

diameter oy = we ‘- - 0,52-0,70 0,50-0,62

Hydrocladium, internode length a Ae 0,57—1,09 0,67-1,15

Hydrotheca, depth abcauline .. a be 0,71-0,82 0,74-0,83

depth adcauline.... Y Re a 0,87—1,00 0,92-1,01

diameter at margin ae oe a 0,24-0,32 0,29-0,32

Remarks

Several species have been recorded from the Antarctic and Subantarctic

which resemble one another in their method of branching, aptly described by

Naumov & Stepaniants (1972) as ‘panicle-shaped’. These can be divided into

two groups:

1. Those where the hydrothecae are adnate for half or more of their height

and where the ‘panicle’ is flexuous: included under the synonymy of Symplecto-

scyphus elongatus by Vervoort (1972a) and in this paper.

2. Those where the hydrothecae have little or no adnate part and where

the ‘panicle’ is stiff.

It is the second group that concerns us here and it includes material recorded

under the names of Symplectoscyphus (or Sertularella) biformis and mawsoni.

Previous and present records of these species, together with the more important

measurements, are listed in Table 1 (the depth of the hydrotheca is not included

because of doubt as to exactly how it was measured by different authors).

The type material of Sertularella biformis Jaderholm, 1905, was examined

(Fig. 1OE-F), and Jaderholm’s description is confirmed. The material differs

from all other records in Table 1 in the absence of hydrocladia or any vestiges

thereof, in the absence of nodes in most of the stem, in the grouping of the

distal hydrothecae in three’s, and in the shape of the hydrotheca which narrows

towards the margin. Totton (1930) assigned material to this species (S. biformis)

assuming that the specimens ‘would correspond with the apical part missing

from Jaderholm’s type . . .’, and that the stem progresses in complexity from

base to apex, both assumptions being unwarranted. Totton’s material, on

examination, was found to differ from Jaderholm’s in all the characters listed

near the beginning of this paragraph (Fig. 8E, Fig. 9 right). It is clearly a different

species altogether. Jaderholm’s species has not been rediscovered to date, and

the type material remains the sole specimen.

The remaining records in Table 1 seem to be conspecific and to form a

series in the order shown, starting with the type material of Symplectoscyphus

mawsoni Briggs and ending with Totton’s material. In this series the complexity

of branching in the hydrocladia increases, and the diameter of the hydrotheca

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES

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decreases. The stem internodes increase in length except for Totton’s material

where they are relatively short and wide so that the hydrocladia are closer

together and give a dense ‘bottle-brush’ appearance to the colony. The material

of Naumov & Stepaniants (1972) attributed with a query to Sertularella biformis

differs from the rest in the absence of hydrothecae on the stem, but it is very

possible that these have been damaged or eroded away. (Totton omitted to

state that his material possesses cauline hydrothecae.)

The present material from the Kerguelen area lies about mid-way in the

series resembling Briggs’ material in the appearance of the colony and the

structure of the stem internodes, but resembling Totton’s material in the shape

of the hydrothecae.

It seems that the first five samples in Table 1 cannot be separated on any

definite characters and it is proposed to unite them under the name Symplecto-

scyphus mawsoni Briggs, 1939. The gonothecae were first described by Naumov

& Stepaniants (1972, fig. 9).

Symplectoscyphus plectilis (Hickson & Gravely, 1907)

Figure 11A—C

Sertularella plectilis Hickson & Gravely, 1907: 20, pl. 3 (fig. 21). Ritchie, 1913: 30, figs 8, 9, 11.

Broch, 1948: 11, fig. 2f-g.

Symplectoscyphus plectilis: Vervoort, 1972b: 354, fig. 8b—c.

Station

14/44-C.

Description

A single, small infertile colony of about 12 stems growing on Halecium sp.

Stem unfascicled, reaching a maximum height of 9 mm, slender, slightly geni-

culate, branching sparingly and in one plane. Branches arising from apophyses

immediately below hydrothecae at a wide angle (95-125°). Nodes usually indi-

cated by indentations of perisarc only. The two rows of hydrothecae in one

plane.

Hydrotheca very small, tubular, curved outwards, adnate for about one-

third adcauline length, narrowing to margin. Marginal teeth well developed.

No internal teeth.

Measurements (mm)

Internode, length R. Se ie ast ie oy 0,39-0,63

Hydrotheca, length abcauline .. ae 4 ie a 0,23-0,29

length adcauline, adnate part ie: oR a8 <i 0,09-0,14

length adcauline, free part 2 se a a 0,22-0,28

adnate part/total adcauline length a a a 0,26-0,36

diameter at margin 2 Me ey 8 es 0,08-0,12

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 35

Remarks

The hydrothecae in this material are of a similar size to those described by

Ritchie (1913); they are a little longer than those described by most other

authorities.

S. plectilis is known from a number of localities in the Antarctic and

Subantarctic, the nearest to the present locality being Bouvet Island (Broch

1948). The distribution and synonymy are summarized by Vervoort (19725).

Symplectoscyphus subarticulatus (Coughtrey, 1875)

Figure 10OA—D

Thuiaria sub-articulata Coughtrey, 1875: 287, pl. 20 (figs 32-34).

Symplectoscyphus subarticulatus: Ralph, 1961: 801, 811, figs 14g-h, 15a—c (synonymy).

Blanco, 1968: 213, pl. 4 (figs 1-3). Leloup, 1974: 41, fig. 38.

Stations

2/6—B (fertile); 2/7—A; 14/45-C.

Description

Fascicled, pinnate stems reaching a maximum height of 105 mm and a

maximum diameter at the base of 1,5 mm. Hydrothecae completely obscured

by the peripheral tubes in the proximal part of the stem, with only the margins

visible in the central region, uncovered in the distal unfascicled region. In the

distal region nodes fairly clearly marked and sloping in alternate directions,

each internode bearing three hydrothecae and a hydrocladium arising imme-

diately below the third hydrotheca. The two rows of hydrothecae and hydro-

cladia in one plane.

Hydrocladium separated from stem apophysis by a distinct node, narrower

than stem. Nodes scarce, oblique, not well marked, sometimes not visible

for the entire length. Hydrothecae closely set, more so in the distal region where

the margin of one overlaps the base of the next, most hydrothecae with a

fenestra immediately below the base on one side.

Hydrotheca tubular, curved outwards, adnate for about two-thirds

adcauline height.

Gonothecae borne on hydrocladia below hydrothecae, long and carrot-

shaped, widest at distal end, strongly annulated with 15-17 circular crests (not

spiral), with terminal aperture on a raised collar.

Measurements (mm)

Hydrotheca, length abcauline .. a ve ba fe 0,40-0,47

length adcauline, adnate part - “ed a mS 0,38—0,60

length adcauline, free part on zi sh ye 0,23-0,31

adnate part/total adcauline length ue af ae 0,57—0,70

diameter at margin - i Le ry - 0,31-0,33

Gonotheca, length re wt as se iis “a 2,35-3,10

maximum diameter i a e7 ap isk 0,86—-1,24

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10.

Symplectoscyphus subarticulatus (Coughtrey). A-B. Hydrothecae. C. Distal part of stem

showing origins of hydrocladia. D. Gonotheca.

Symplectoscyphus biformis (Jaderholm). E. A single hydrotheca. F. A group of three hydro-

thecae. Drawn from the holotype.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 37

Remarks

The hydrothecae of this material are larger than those of Ralph but smaller

than those of Blanco. The details of structure and the method of branching

agree exactly with those described by Ralph. The only hesitation in the identi-

fication lies in the gonotheca, which is about twice the length of that described

by Ralph and has more annulations (15-17 as against 5-8). However, the

shape of the gonotheca is exactly like that depicted by Coughtrey (he does not

give the size, but the diagram shows 10 annulations).

S. subarticulatus is best known from New Zealand, but recent reports

show that it also occurs in the southern Argentine (Blanco) and in Chile (Leloup).

It is a new record from the Kerguelen area.

Symplectoscyphus subdichotomus (Kirchenpauer, 1884)

Figure 11D-—F

Sertularella subdichotoma Kirchenpauer, 1884: 46, pl. 16 (fig. 1).

Symplectoscyphus subdichotomus: Blanco, 1969: 49, figs 1-18. Vervoort, 1972a: 140, figs 44b—d,

45 (synonymy). Leloup, 1974: 42, fig. 40.

Stations

Small form: 8/25—E; 26/64-E (fertile); 26/65—A (fertile); 28/71-C (fertile);

30/73-D; 31/74-G.

Large form: 3/11—D; 14/44-B; 21/57-D; 22/58-J; 26/63-F.

Description

Colonies flexuous and straggling, growing profusely with polyzoans and

Sertularella picta, and often intertwined to form a bushy mat. Stem unfascicled,

reaching 33 mm in height (small form) or 58 mm (large form), generally geni-

culate and giving off a branch at each elbow immediately below every third

hydrotheca, but many irregularities present. The two rows of hydrothecae and

branches in one plane. Branches of same diameter as stem, forming an angle

of 85—-108° with it, and often rebranching producing a subdichotomous effect.

Most of the nodes not clearly defined and recognizable only by an indentation

of the perisarc, but those at the origin of each branch, and on the stem imme-

diately above each axillary hydrotheca, more clearly defined and usually

forming a definite septum. Internodes always longer than the hydrothecae,

which do not overlap each other.

Hydrotheca adnate for over half adcauline height, curved outwards, widest

in centre and narrowing to margin, with three well-developed marginal teeth

and no internal teeth.

Gonotheca obovate (female), with 10-11 strong and crested circular

annulations (not spiral) and a long flaring mouth-funnel, containing two or

three large planulae.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11.

Symplectoscyphus plectilis (Hickson & Gravely). A. Part of stem showing origin of hydro-

cladium. B—C. Hydrothecae.

Symplectoscyphus subdichotomus (Kirchenpauer). D. Stem of small form showing origins

of hydrocladia and a gonotheca. E. Hydrotheca of small form. F. Hydrotheca of large form.

Scale in mm/10.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES

Measurements (mm)

Small form Large form

Internode length .. 0,47-0,74 0,74-1,27

Hydrotheca, length abcauline 0,26-0,31 0,28-0,38

length adcauline, adnate part 0,25—0,30 0,25—0,39

length adcauline, free part 0,13-0,21 0,14-0,28

adnate part/total adcauline length 0,58—0,68 0,48-0,71

diameter at mouth 0,13-0,17 0,20—-0,30

Gonotheca, length 1,06—1,55

maximum diameter 0,58-0,84

Remarks

As far as the detailed measurements are concerned this material includes

two size-ranges—a small and a large—the smaller fitting best with the dimensions

given by Vervoort, Blanco and earlier workers. The growth-form and structure

of the two are the same, but there are no gonothecae to confirm the identifica-

tion of the larger form. |

The synonymy of S. swbdichotomus has been reviewed by Vervoort (1972a),

who also gave some excellent diagrams. There is no doubt at all that the present

material (at any rate the smaller form) belongs to the same species. Because

of the presence of a number of closely allied species of dubious validity, the

geographical range is uncertain—the species is certainly common in the Falk-

lands/South American region, but has apparently not been reported from

Kerguelen.

Family Plumulariidae

Kirchenpaueria triangulata (Totton, 1930)

Plumularia triangulata Totton, 1930: 225, fig. 61.

Kirchenpaueria triangulata: Millard, 1975: 375, fig. 119E—-H.

Station

17/50-B.

Description

A rich, fertile colony epizootic on Plumularia insignis. Stems reaching

18 mm in height, unfascicled, with nodes absent or very indistinct, otherwise

details as in previous descriptions. Gonophores female, reaching 2 mm in

length, triangular in section and containing a single layer of eggs over a central

spadix.

Remarks and distribution

This is a new record for the Kerguelen area. The species was previously

known from South Africa (in depths greater than 100 m) and from New Zealand.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Oswaldella bifurca (Hartlaub, 1904)

Figure 12A—C

Schizotricha bifurca Hartlaub, 1904: 16, pl. 3 (figs 4-8).

Oswaldella bifurca: Totton, 1930: 208, fig. 50. Naumov & Stepaniants, 1962: 98.

Station

26/64-J.

Description

Several infertile stems reaching a maximum height of 22 mm. Stem divided

by transverse nodes into internodes which usually bear one hydrocladium

each, but rarely two. Cauline nematothecae very difficult to distinguish and

sometimes missing; at the maximum development there is one mamelon on

the hydrocladial apophysis and one nematotheca in the axil, with another

nematotheca (or a naked sarcophore) on the main stem above the axil.

Hydrocladia unbranched near the proximal and distal ends of colony,

branched in the central region. Branches arising from below hydrothecae and

to one side, as many as four from one hydrocladium, and these may be all on

the same side, on alternate sides or in opposite pairs. One secondary hydro-

cladium has a tertiary branch.

The two rows of primary hydrocladia not in the same plane but displaced —

towards anterior surface of stem. Secondary hydrocladia similarly displaced,

so that the whole arrangement is in at least six different planes and presents a

stiff and spiky appearance. Since the hydrothecae are borne on the anterior

surface of the hydrocladia, those of primary and secondary hydrocladia do not

face one another.

Hydrocladia generally heteromerous with alternating athecate and thecate

internodes, but many variations; the athecate internodes may be absent or

duplicated, long or short, and may or may not bear a median nematotheca.

Thecate internodes usually with one median inferior and one median superior

nematotheca, but the nematotheca is often missing leaving only a naked sarco-

phore, and this is especially common in the first thecate internode of the hydro-

cladium. Nematothecae, when present, curved and scoop-shaped. Median

superior nematotheca situated behind distal part of adcauline thecal wall

which is not completely adnate.

Hydrotheca wider than deep, with more or less straight abcauline wall

and margin at least slightly oblique.

Measurements (mm)

Thecate internode, length an i ds oe a 0,42-0,96

Hydrotheca, depth abcauline .. ¥ - ng ad 0,12-0,19

diameter at margin oe oe es 7 a 0,17-0,22

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 4]

A-C

Dh le

Fig. 12.

Oswaldella bifurca (Hartlaub). A-B. Normal parts of hydrocladia. C. Proximal end of

hydrocladium showing origin from stem and first thecate internode which is without a

nematotheca.

Plumularia insignis Allman. D. Part of stem in thinner region showing origins of two branches

and their preceding hydrocladia. E. Gonothecae. F. Part of hydrocladium.

Scale in mm/10.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

This material is assigned to O. bifurca with some hesitance, the decision

being based mainly on the shape of the hydrotheca, which is wider than deep,

and on the method of branching. As regards the latter feature Totton’s words

(1930) are pertinent, when he says for O. bifurca ‘Hydrothecae of hydroclade

and branch not strictly opposed, but facing slightly upwards’. On the other

hand the features used by Totton as diagnostic for O. bifurca, namely ‘once-

branched hydrocladia’ and the absence of an inferior nematotheca on the first

hydrocladial internode, are found to be variable and of dubious value. The

presence of athecate internodes in the hydrocladium and the number of hydro-

cladia to a stem internode are also variable characters. This species is known

from various localities in the Antarctic; it has not been reported before from

Kerguelen.

Plumularia insignis Allman, 1883

Figure 12D-F

Plumularia flabellum* Allman, 1883: 19, pl. 1 (figs 1-4).

Plumularia insignis Allman, 1883: 21, pl. 2. Billard, 1910: 32, fig. 14.

Plumularia abietina Allman, 1883: 21, pl. 3.

Plumularia insignis v. flabellum: Billard, 1910: 34, fig. 15.

Plumularia insignis v. abietina: Billard, 1910: 35.

Plumularia sp. Naumov & Stepaniants, 1962: 99, fig. 19.

Stations

3/10—-A; 3/11-—C; 7/22—A; 17/50—A; 18/52—A; 23/59-D; 24/61-A; 26/64-A;

30/73-E.

Description

Many magnificent colonies, the tallest reaching 940 mm. Rootstock a

mass of interwoven fibres suitable for anchoring in mud. Stem long, flexuous

or fairly stiff, fascicled for the greater part, giving off branches in all planes,

unsegmented. Immediately below each branch a single hydrocladium arises

from the stem and from the same component tube, otherwise cauline hydro-

cladia occur only rarely. Component tubes of stem bearing longitudinal rows

of nematothecae.

Branches fascicled or unfascicled, pinnate, divided by transverse nodes at

irregular intervals into internodes bearing one to four alternate hydrocladia.

Hydrocladial apophysis with two axillary nematothecae (one anterior and one

posterior) with a mamelon between them, and one (or rarely two) nematothecae

on distal end. Nematothecae on main axis rather irregular, but at least one

between every two successive hydrocladia and several below the first

hydrocladium.

* Although P. flabellum has page priority over P. insignis, from Article 24(a) of the Code of

Zoological Nomenclature, Billard (1908: 759) counts as the ‘first reviser’ and he has clearly

chosen P. insignis as the name of the composite species.

een aise

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 43

In some colonies the hydrocladia bear practically only thecate internodes,

each with one hydrotheca and three nematothecae (one median inferior and

one pair of laterals), athecate internodes occurring only sporadically and

obviously resulting from regeneration after injury. In other colonies intermediate

athecate internodes are present throughout, each with one median nematotheca

and two internodal septa. Thecate internodes very rarely have two median

inferior nematothecae. Internodal septa usually not so well developed as in

the type material: 1-2 below hydrotheca, 0-3 behind it, and 0-1 above it.

Gonothecae arising from branches next to hydrocladia, elongated, widening

distally, with a terminal oblique opening, as illustrated by Allman (1883) for

P. abietina.

Measurements (mm)

Thecate internode, length v = Ne < ae: 0,54-0,80

Hydrotheca, height abcauline .. ot: . He Pa 0,21-0,34

diameter at margin A 2 on =? fa 0,14-0,21

Gonotheca, length 43 ie a a ne A 1,40-1,95

maximum diameter ie ae = Ae ue 0,35-0,58

Remarks

This species occurs in several varieties, of which three (described by Allman

as P. insignis, P. flabellum and P. abietina) are known from the Subantarctic.

In the general appearance of the colony and in the detailed measurements,

most of the present material closely resembles the nominal variety illustrated

by Allman (1883: pl. 2). One colony, however (26/64—A), is of stouter build

and more irregular branching and is more like Allman’s diagram of P. flabellum

(1883: pl. 1). Of the three, Allman showed hydrocladia on the main stem only

in P. flabellum, Nowhere has the singular character of one hydrocladium below

each branch been described, and if this character is indeed peculiar to the

Kerguelen material it would be worth naming a separate variety.

Most of this material is unusual in possessing as the ‘normal’ condition

only thecate internodes in the hydrocladia, athecate internodes being rare and

sporadic. One colony alone (26/64-A, mentioned also above) has regular

intermediate athecate internodes. Billard (1910) has commented on the irregu-

larity of occurrence of athecate internodes which are often the result of regenera-

tion after injury. Naumov & Stepaniants (1962) described some infertile material

from Kerguelen and Heard Islands without athecate internodes as Plumularia sp.

This is without doubt the same species.

The species is known from Prince Edward Island, Marion Island, Kerguelen

and Heard Islands, and there are two varieties from the East Indies.

Schizotricha unifurcata Allman, 1883

Schizotricha unifurcata Allman, 1883: 28, pl. 7 (figs 1-3). Stechow, 1925: 498. Totton, 1930:

231, fig. 65, pl. 3 (fig. 4). Naumov & Stepaniants, 1972: 54. Blanco & De Miralles, 1972:

21, pl. 5 (figs 3440).

Polyplumaria unifurcata: Billard, 1910: 41, fig. 18.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Stations

8/25-A; 9/26-B (fragments); 10/30-B; 22/58-A; 23/59-C.

Description

Stems with tangled rootstock for mud-penetration and reaching 175 mm

in height. Hydrocladia branching one to three times, 10-18 mm in length.

Details of hydrocladia as in type material.

Male gonothecae present on first colony, arising from thecate internodes

below hydrothecae, pear-shaped, with oblique distal aperture, bearing two to

four nematothecae on basal region.

Remarks

Stechow (1925) combined the species S. unifurcata Allman, 1883; S. tur-

queti Billard, 1906; S. anderssoni Jaderholm, 1904; and S. glacialis (Hickson

& Gravely, 1907) under one name. Totton (1930) added S. multifurcata Allman,

1883. All of these species are Antarctic or Subantarctic in distribution.

This material is very similar to the type material and indeed, two of the

colonies come from the type locality (Kerguelen Island).

DISCUSSION

Of the 33 species here recorded (omitting Campanularia sp.) 5 are cos-

mopolitan, namely Modeeria rotunda, Halecium delicatulum, Halecium tenellum, -

Filellum serratum and Lafoea dumosa (Table 2).

The remaining 28 species show only a very slight affinity with the nearest

continental masses, for only 1 (3,6%) also occurs in South Africa (Kirchen-

paueria triangulata) and only 8 (28,6°%) also occur in the Australasian region.

The affinities of these 28 species are in fact mainly with the South American

region (or Magellan Province, including the Falklands and South Georgia)

and with the Antarctic Continent, for 19 of the species (67,9%) also occur in

the former and 14 (50,0°%) in the latter.

Surprisingly, there is little obvious affinity with Bouvet Island (3 species:

10,7°%) or Marion and Prince Edward Islands (5 species: 17,9%) which lie

at much the same latitude, but this is probably merely a reflection of the inade-

quate knowledge of these islands.

This analysis thus supports the classification of the Kerguelen and Crozet

groups of islands as Subantarctic (J. C. Briggs 1974), with a fauna intermediate

between that of the Cold Temperate South American Region and the Antarctic

Region. As has been remarked by Naumov & Stepaniants (1962), it is not

possible to draw a sharp line of demarcation between the fauna of the Ant-

arctic and that of the Subantarctic.

There is at this stage a little evidence of a divergence between the fauna

of the Kerguelen group on the one hand and the Crozet group on the other.

Although they share 12 (42,9°%%) of the non-cosmopolitan species, 8 occur only

in Kerguelen and 8 only in Crozet.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 45

TABLE 2

The distribution of the 33 species recorded in this paper.

—

s 3 E

bie Sige Tee = <

S -h 8: « oe

ee a ay ie ee eS

aos Peete gs ee

Gig Wa Ei ee wae gs Se

o 5 w = oS oS = = 5

O wo A a = Se Co 2

Eudendrium rameum x x a

E. tottoni x x x

Modeeria rotunda x x x x (cosmop.)

Opercularella belgicae x x SA

Phialella chilensis * ys sé

Halecium delicatulum xX x x x x x x x (cosmop.)

H. dufresneae WA

H.jaederholmi . x x x

H. tenellum : x x oe * x (cosmop.)

Hydrodendron arborea xX NA x Se Me

Filellum serratum x x x x x (cosmop.)

Grammaria abietina. xX mK x

Halisiphonia ?nana x x

Hebella striata . % x x

Lafoea dumosa . : x ~ 4 x K x (cosmop.)

Zygophylax crozetensis X

Campanularia

norvegiae : ; Xx Xx

Silicularia rosea x Me Sc x <

Tulpa diverticulata x rm x

Staurotheca antarctica x x < we

S. dichotoma Done x x x x

Sertularella geodiae . x S x

S. picta . x * x x

Symplectocyphus

curvatus . s< x

S. elongatus . xX x x o<

S. mawsoni < x

S. plectilis . x * x x

S. subarticulatus x SS x

S. subdichotomus . xX x x x x

Kirchenpaueria

triangulata . ; x x x

Oswaldella bifurca . x x

Plumulariainsignis . x x x

Schizotricha

mayurcata” tt S< Se oe x

Total (less cosmop.) 20 19 19 3 5 1 14 1 8

map icss.cosmop.) 71:4 “67,9 67,9 - 10,7 17,9 ° 3,6 50:0 3,6 28,6

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

The author wishes to thank Dr J. C. Hureau of the Muséum National

d’Histoire Naturelle, Paris, and Dr P. M. Arnaud of the Station Marine

d’Endoume, Marseille, for the privilege of examining and reporting on this

collection.

The British Museum (Natural History) granted permission for the exami-

nation of Challenger material of Sertularia (Symplectoscyphus) exserta and

Terra Nova material of ‘Sertularella biformis’, while the Naturhistoriska Riks-

museum, Stockholm, permitted examination of type material of Sertularella

biformis from the Swedish Southpolar Expedition. This help is gratefully

acknowledged.

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Hickson, S. J. & GRAVELY, F. H. 1907. Coelenterata. I]. Hydroid zoophytes. Nat. Antarct.

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HureEau, J. C. 1976. La campagne d’océanographie biologique MD 03/ICHTYO a bord du

‘Marion-Dufresne’ (18 Mars—28 Avril 1974). Résultats préliminaires et liste des stations.

CNFRA 39: 3-25.

JADERHOLM, E. 1905. Hydroiden aus antarktischen und subantarktischen Meeren gesammelt

von der schwedischen Siidpolarexpedition. Wiss. Ergebn. schwed. Siidpolarexped. 5: 1-41.

HYDROIDS FROM THE KERGUELEN AND CROZET SHELVES 47

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LeLoup, E. 1974. Hydropolypes calyptoblastiques du Chili. Report no. 48 of the Lund Uni-

versity Chile Expedition 1948-1949. Sarsia 55: 1-61.

MILLarp, N. A. H. 1968. South African hydroids from Dr Th. Mortensen’s Java—South Africa

Expedition, 1929-1930. Vidensk. Meddr dansk naturh. Foren. 131: 251-288.

MILLARD, N. A. H. 1971. Hydrozoa. In: ZINDEREN BAKKER, E. M. VAN, WINTERBOTTOM, J. M. &

Dyer, R. A., eds. Marion and Prince Edward Islands: 396-408. Cape Town: A. A. Balkema.

MILLARD, N. A. H. 1975. Monograph on the Hydroida of southern Africa. Ann. S. Afr. Mus.

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6. SYSTEMATIC papers must conform with the International code of zoological nomenclature

(particularly Articles 22 and 51)

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.

nov., Syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes should be listed separately. The complete data (registration number,

depository, description of specimen, locality, collector, date) of the holotype and paratypes

must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,

Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g. *... the Figure depicting C. namacolus...’; ‘. . . in C. namacolus (Fig. 10)...”

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded

by initials or full names

e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a

book or article, such as

“Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation

to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the

abstract, counter to Recommendation 23 of the Code, to meet the requirements of

Biological Abstracts.

N. A. H. MILLARD

HYDROIDS FROM THE KERGUELEN J

AND CROZET SHELVES, COLLECTED BY

THE CRUISE MD.03 OF THE MARION-DUFRESNE

JME 73 PART 2 JUNE 1977 ISSN 0303-2515

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