THE HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH

AFRICA. PART III. THE GYMNOBLASTEA AND SMALL FAMILIES

OF CALYPTOBLASTEA

N. A. H. Miniarp

XKoology Department, University of Cape Town

(With 15 text-figures and 1 plate)

CONTENTS

PAGE

Introduction : : é : ed Oy,

Station List . : : : : . 428

List of Species. ‘ : i =) 433

Tubulariidae é 3 : 6 - 434

Myriothelidae . : : : » 435

Corynidae . : F ; : - 440

Solanderiidae 3 é , : - 444

Bougainvilliidae . : 3 : - 449

Clavidae . é 3 : 5 - 452

Eudendriidae : : . : - 454

Hydractiniidae . i : : - 457

Pandeidae . : : 3 : 4 Or

Aequoreidae : : : : . 461

Lovenellidae : : : : . 464

Haleciidae . 6 ; : : . 464

Campanulariidae . : : 6 ye 47t

Summary . : ; ; : . 484

Acknowledgements : , : . 484

References . 3 é : é AGA

INTRODUCTION

This paper represents the third and final part of a systematic account of

the hydroids from that part of the South African coast stretching from South

West Africa on the west to the southern border of Natal on the east. For

details on the origin of the collections the reader is referred to the introduction

to part I of the series (Millard, 1962).

When examining Gymnoblast hydroids one must bear in mind that a

considerable quantity of the more delicate material brought up by dredging

becomes destroyed by rough handling or is so badly damaged that identification

beyond the genus level is impossible. Moreover most genera require the presence

of gonophores, usually female, for identification to species level. This applies

particularly to such genera as Eudendrium and Tubularia. It is felt that no useful

Ann. S. Afr. Mus. 48 (18), 1966: 427-487, 15 figs., 1 pl.

427

428 ANNALS OF THE SOUTH AFRICAN MUSEUM

purpose would be served by including dubious records and, at the risk of

giving a false impression of the abundance, poorly preserved material and

sterile material of doubtful specific identification has generally been omitted.

The author wishes to express thanks to the following: the South African

Museum for permission to examine the collection of the s.s. Pieter Faure, the

Zoology Department of the British Museum of Natural History for accommoda-

tion and permission to examine their collection in 1960, Dr. W. Engelhardt

of the Munich Museum and Dr. J. S. Pringle of the Natal Museum for the

loan of type material, Dr. M. E. Thiel of the Hamburg Museum for identifica-

tion of certain medusae and finally all past and present members of the Zoology

and Oceanography Departments of the University of Cape Town who have

helped to accumulate material in the now very extensive University collection.

Type specimens of new species have been deposited in the South African

Museum and have been given a Museum registered number in addition to the

University catalogue number.

STATION LIST

A. Littoral material from Oudekraal on the west coast of the Cape Peninsula.

Position: 33°58°5’'S/18°22-2’E.

Date

A 40 16.1.34

A 116-122) -15.3.34

A 139 17-3-34

A 353

A 383 13-5.34

A 384 25.8.34

A 405 18.10.34

AFR. Material dredged by the government research vessel, R.S. Africana.

Date Position Depth (m.) Bottom

AFR 736 17.8.4.7 30°42°4’S/15°59°2’E 201 co gn S, Sh

AFR 801 7.10.47 32°34°4/S/17°52°2’E 71 gn M,R

AFR 866 9.1.48 34°36-8’S/19°16-4’E 38 R,S

AFR 967 23.3.48 35°07’S/20°49’E gI {S$

AFR 985 5.4.48 34.°4.7°4’S/20°19’E 80 gn M

AFR 994 19 4.48 34°34°5'S/21°22°5’E 68 co S, Sh

AFR 0002 6.6.49 33°10'S/17°57'E 73

B. Littoral material from Lambert’s Bay on the west coast. Date: July 1938.

Position: 32°05'S/18°18’E.

BMR. Bushman’s River Estuary, south coast, on sand and muddy banks.

Date: 14.9.50. Position: 33°41'S/26°40’E. Depth: 2-4°5 m.

BRE. Breede River Estuary, south coast, littoral. Date: 3.2.52. Position:

34°25'S/20°51°5’E.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 429

CP. Littoral material from various localities on the west coast of the Cape

Peninsula.

Date Locality Position

CP 325 20.9.48 Sea Point 33°55°2°S/18°22-6’E

CRi927 16.12.48 Blaauwberg Strand 33°48'S/18°27-5’E

CP 331 8.8.49 Kommetje 34°08°-5'S/18°19:4’E

CP 336 12.5.49 Oudekraal 33°58°5'S/18°22-2’E

CP 378 3.6.52 Cape Town docks (on 33°54°5'S/18°25°5’E

Squalus fernandinus)

CP 646 5-12.60 Oudekraal 33°58°5'S/18°22-2’E

CPR. Material from various localities in the Cape Province.

Date Locality Position Depth (m.)

CPR 9 30.4.50 Glentana Strand 34°04’S/22°20’E littoral

CPR 46 20.6.59 Umgazi Bay 31°4.3'S/29°26’E 27

HAM. Keiskama River Estuary, Hamburg, on the south coast. Date: 9.1.50.

Position: 33°17'S/27°32’E.

HB. Littoral material from Hondeklip Bay on the west coast. Date: 8.2.40.

Position: 30°19'S/17°16’E.

KNY. Knysna Estuary on the south coast. Position: 34°05'S/23°04’E (average).

Date Depth (m.) Bottom

KNY 30 16.7.47 5-7 M,S

KNY 70 15.7.47 2-6 S

KNY 164 9.7.50 0-3 M

KNY 165 9.7.50 I-13 S

KNY 176 11.7.50 Floating buoy

KNY 212 7.7.60 o-1 S

KNY 270 14.2.64 o-I M

L. Littoral material from East London on the south coast. Position: 33°01'S/

27°54'E.

Date

L 44 6.3.37

L 56-177 ty

LAM. Dredged in Lambert’s Bay, west coast.

Date Position Depth (m.) Bottom

LAM 24 16.1.57 32°04°6’S/18°18-15’E 17 R

LAM 30 19.1.57 32°05-1'S/18°17-7’E 20 R

LAM 35 19.1.57 32°05'5'S/18°17-7’E 27 R, Sh

LAM 46 22.1.57 32°04°4'8/18°17-7’E 23 R

LAM 50 23.1.57 32°08°5'S/18°17°7’E 16 R,S

LAM 52 21.1.57 32°04°7'S/18°18-2’E 17 S

LAM 59 22°05 7, 32°09S/18°18’E 16 R,S

LB. Langebaan Lagoon, west coast.

Date Position Depth (m.) Bottom

LB 127 26.4.48 33°05'6’S/18°01-6’E Littoral R

LB 166 15.7.46 33°09'0'S/18°03°4’E

LB 266 3.5.51 33°07°4’S/18°02-1’E Littoral S

430 ANNALS OF THE SOUTH AFRICAN MUSEUM

Date Position Depth (m.) Bottom

LB 296 55-51 33°04'5/18°00’E Ship’s hull

LB 314 4.5.51 33°05°7'S/18°01°5’E Littoral R

LB 371 7.5.53 33°05°7'S/18°01°5’E Littoral R

LB 378 7.5.53 33°05:9'5/18°01-9’E O-I Wooden piling

LB 380 75-53 33°06-7’S/18°01-0’E 4 S, Sh

LB 386 8.5.53 33°06'5’S/18°02’E Ship’s hull

LB 398 24.7.53 33°07'1'S/18°02-9’E O-1 S)

LB 403 5-12.53 33°10'S/18°03°5’E Littoral M

LB 542 4.5.60 Littoral

LB 556 22.7.63 33°07°3'S/18°03:0’E Littoral Wooden piling

LIZ. Dredged in Algoa Bay, Port Elizabeth, south coast.

Date Position Depth (m.) Bottom

LIZ 2 5-4-54 33°55°7 9/24°37°2 E 9 M

LIZ 3 5.4.54 33°56-1'S/25°40’E 17 S

LIZ 7 6.4.54 33°58°1'S/25°38-9’/E 9 R, St

LIZ 11 6.4.54 33°57°2'S/25°38-0'E 9 Clay, R

LIZ 13 6.4.54 33°58-2'S/25°38-8’E 5] S

LIZ 16 7.4.54 33°58-4'S/25°40°5’E 14 St

LIZ 24-25 11.4.54 34.°00°4'S/25°44°5'E 39 co S, Sh

LIZ 40 11.4.54 34.°00°8/S/25°4.2°4’E 6 R

LU. Littoral material from Luderitz Bay, South West Africa.

Date Position

LU 59 24.2.63 26°38/S/15°05'5'E

LU 113-118 =. 22.2.63 26°38’S/15°09°3’E

MB. Dredged in Mossel Bay, south coast.

Date Position Depth (m.) Bottom

MB 8-12 12.1.56 34.°04°3/S/22°13°9’E 1g R

MB 19 13.1.56 34.°08-7'S/22°07-2’E 13 R, S, Sh

MB 25 13.1.56 34°09°3'S/22°10°1’E 29 S)

MB 37 16.1.56 34°09°3'S/22°10°0’E 31 S

MB 47 17.1.56 34°11°3/S/22°10-0’E 10 R

MB 52 17.1.56 34°11°0'S/22°09'9’E 14 R,S

MB 55 17.1.56 34.°10°7'S/22°09°6’E 9 R

MB 60 18.1.56 34.°04°3'S/22°14°2’E 18 R, co 8, Sh

MB 64 18.1.56 34.°04°8'S8/22°13°1’E 26 R, co S, Sh

MB 69 19.1.56 34°08-6’S/22°07°3’E 13 R,S

MB 70 19.1.56 34°08-9’S/22°07°9’E 18 S

MB 81 20.1.56 34°06-2’S/22°10°9’E 27 M

MB 84 Q1.1.56 34°11°4’S/22°10°1’E 29 R

MB 88 18.1.56 34°04°8'8/22°13°1’E 26 R, co S, Sh

OLF. Olifant’s River Estuary, west coast. Date: 23.1.55. Position: 31°42’S/

TG mn5 jh.

PP. Littoral material from Paternoster, west coast. Date: 24.9.57. Position:

32°43'S/17°55 E.

SAMH. Material from the collection of the South African Museum. Specimens

147-352 and 361-383 were dredged by the s.s. Pieter Faure. Their positions

were given in the original records as compass bearings off salient points

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 431

on the coast, and were probably not very accurate. These have been

converted into latitude and longitude and are given to the nearest minute.

SAMH 147

SAMH 157-162

SAMH 166-170

SAMH 174-177

SAMH 179-180

SAMH 189

SAMH 202-203

SAMH e214

SAMH 221-227

SAMH 230-233

SAMH 235

SAMH 239-247

SAMH 250-253

SAMH 269-274

SAMH 283

SAMH 296

SAMH 315

SAMH 327-328

SAMH 335-338

SAMH 341-343

SAMH 351-352

SAMH 357

SAMH 361

SAMH 380-383

SAMH 404

SAMH 405-408

Date

23.6.1898

15.7.1898

11.11.1898

19.11.1898

22.12.1898

28.12.1898

7-3-1899

24..3.1899

19.6.1899

20.9.1899

5-7-1900

11.10.1900

15.7-190I

17.7.1QOI

25.7.1QOI

13.8.1901

10.9.1QOI

22.9.1904.

4.10.1904

19.10.1904

22.8.1905

19.6.1914

11.11.1898

15.3.1899

—.4.1962

—.7.1962

Position

South of Mossel Bay

34°08'S/22°16’E

33°49'S/25°56’E

33°45 '9/26°44’E

32°52'S/28°12’E

33°09'S/28°03’E

33°59'S/25°51'E

33°50'S/26°35’E

34°26'S/21°42’E

34°15'S/22°10°5’E

34°27'S/20°58’E

34°08’S/22°59°5'E

33°13°5'S/27°58’E

33°07'S/27°47°5'E

32°50'S/28°18-5’E

32°4.7'8/28°28’E

33°54'9/26°51’E

34°12’S/22°15:5’E

34°12'S/22°15°5’E

34°15°5'S/22°14’E

33°52'S/26°09’E

33°55°9/18°27’E

33°49'S/25°56’E

33°47'5/26°19’E

28°30’S/16°10’E

28°30'S/16°10’E

SB. Saldanha Bay, west coast.

SB 132

SB 153-168

SB 174

SB 178

SB 196

SB 231

SB 235

SB 267

SB 269

SCD. Dredged off the

SCD 5

SCD 20

SCD 22

SCD 26

SCD 29

SCD 37

SCD 50

SCD 52

SCD 56

SCD 60

SCD 61

SCD 75

SCD 79-81

Date

26.3.53

—:9-57

27-4-59

28.4.59

1.5.59

4.5.60

25.4.62

Date

19.4.58

26.5.58

235.58

22.6.58

19.5.58

18.5.58

20.8.58

19.8.58

16.8.58

15.8.58

16.7.59

Position

33°04S/17°59°3E

33°02°5'S/18°02’E

33°02:8’S/18°00-6’E

33°03°6’S/18°00-4’E

3304°4'S/17°56-4’E

33°00-6’S/17°59:6’E

33°04°1'S/17°59°7'E

33°02'S/17°57°2’E

33°02°1'S/17°58’E

south coast.

Position

34°15/S/25°05’E

34°07°3'S/23°23°8'E

34°26-7'S/23°26-0’E

3347°5/26°04’E

33 38°6'S/26°54-7'E

32°15°2'S/28°57-7’E

31°38-8'S/29°34°4’E

34°01'S/25°45°5'E

33°37'5/26°56-6’E

33°02'S/27°56:2’E

32°17°7'S/28°54°5'E

32°33'S/28°38’E

32°43’S/28°28’E

Depth (m.)

73-78'5

24527

Depth (m.)

Littoral

Bottom

S, Sh, St

Bottom

432 ANNALS OF THE SOUTH AFRICAN MUSEUM

Date Position Depth (m.) Bottom

SCD 84-85 17-7-59 33°03/S/27°55'E 27 R

SCD 94 20.7.59 33°55°5'9/25°51’E 46 bk M, S$

SCD 111 23.7.59 34°35/S/21°11’E 75 co S, Sh, St

SCD 112 20.7.59 33°55°5'S/25°51E 46 bk M, S

SCD 113 26.11.59 34°24’S/21°45’E 27 S; £Sh

SCD 114 26.11.59 34°29'8/21°49°5’E 73 M

SCD 117-118 14.2.60 34°24’8/21°46’E 18 R

SCD 119 14.2.60 34°33'S/21°52’E 77 kh S

SCD 126 3.6.60 34°26°5'S/21°48’E 67 bk M

SCD 129-133 3.6.60 34°48/S/22°06’E 100 kh S

SCD 154 25.11.60 34.°03'S/25°59’E 84 R

SCD 169 24.11.60 33°58-9'S/25°41°4’E 7 R

SCD 175 30.11.60 34°208/23°31’E 110 R,khS

SCD 179 24.11.60 33°58-9'S/25°41°4’E 7 R

SCD 188 30.11.60 34°10'S/23°32’E 97 gen M

SCD 190 29.11.60 34°05'8/S/23°23°2’E 10 {S$

SCD 206 30.11.60 34°51'S/23°41’E 182 kh §

SCD 215 25.11.60 34°03'S/25°58’E 78 S, Sh

SCD 239 29.11.60 34°02'0'S/23°28-4’E 49 M,R,S

SCD 258 14.7.61 33°53°8'S/25°42°5'E 32 £8

SCD 265 14.7.61 33°48'S/25°47’E 27 bl M, R

SCD 276 14.7.61 33°53°8'S/25°42°5'E 32 {Ss

SCD 281-283 11.2.62 34°04'S/23°23’E 22 £S, Sh

SCD 284 6.2.62 33°01'S/27°55'E 7 £{S

SCD 312 9.2.62 33°58'8/25°47'E 48 {S$

SCD 314 9.2.62 33°58S/25°43'E 36 fM

SCD 328 10.2.62 34°43'S/25°40’E Floating buoy

SCD 330 11.2.62 34°03°5'S/23°23’E 15 {S$

SCD 333 11.2.62 34°02'S/23°27’E 42 M

SCD 347 12.2.62 34°10'S/22°15’E 54 M

SCD 354 11.10.62 32°08’S/290°12’E 210

SCD 387 8.12.62 34°04°4'S/23°25-6’E 46 R

SCD 394 2.12.62 33°50°7'S/25°47°5'E 36 R,S

SH. Cape Town docks, from pylons and cable below tug jetty.

Date

SH 433 14.4.61

SH 436 3.4.62

STJ. St. John’s River Estuary, eastern coast of Cape Province. Date: 20.1.50

Position: 31°37’S/29°37’E.

SUN. Sunday’s River Estuary, south coast. Date: 7.1.50. Position:

33°42'S/25°53’E.

SWD. Dredged off the coast of South West Africa.

Date Position Depth (m.) Bottom

SWD 12 10.2.63 26°35’S/15°o1’E 71 R

SWD 39 12.2.63 26°37°5'S/15°04°5’E 40 R

SWD 42 13.2.63 26°38-0’S/15°00-2’E 71 R

TB. Dredged from Table Bay, Cape Town.

Date Position Depth (m.) Bottom

TB6 4.8.46 33°4.9°5'S/18°27-5’E 13 S, Sh

TB 7-8 25.10.46 33°52°5'S/18°27-5’E 17 S Sh

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 433

TB 9g

TB 13

TB 14

TBI5

TB 16

BET 7

TB 19

TB a1

Date

25.10.46

4.8.46

25.10.46

11.2.4.7

25.10.46

26.6.4.7

3-7-47

15.12.57

TRA. Material collected by commercial trawlers.

TRA 32

TRA 33

TRA 35

TRA 38

TRA 42

TRA 56

TRA 57

TRA 59

TRA 86

TRA 92

TRA 99

TRA 150

TRA 156

TRA 159

WCD. Dredged

WCD 7

WCD 12

WCD 18

WCD 20

WCD 25

WCD 81

WCD 125

WCD 134

WCD 145

WCD 156

WCD 158-160

WCD 164

Tubularia solitaria Warren, 1906.

Date

Q.11.47

20.7.49

21.1.50

—.7.50

—.7.51

28.11.52

27.11.52

26.11.52

23-3-53

—.1.54

18.1.56

6.3.58

15.10.58

6.7.58

off the west coast of the Cape Province.

Date

243-59

24.3-59

29-4-59

30-4-59

1.5-59

15-9-49

23.4.62

25.4.62

29.8.63

22.10.63

21.10.63

7.10.63

Monocoryne minor n. sp.

Myriothela capensis Manton, 1940.

Bicorona elegans, n.g., n.sp.

Sarsia eximia (Allman, 1859).

Position Depth (m.)

33°52'S/18°28’E 15

33°49°5'S/18°27-5’E 13

33°52°5/8/18°27°5'E 17

33-47°5'5/18°24-3’E 19

33°52'5/18°28’E 15

33°52°7'S/18°28-7’E 9

33°51°2'S/18°27°3’E 23

33°48-6’S/18°24°6’E 15

Position Depth (m.)

34.°49'S/20°21-5’E ¢. QI

34.°55'S/21°10’E c. gO

34°34S/20°50’E 70

34°30'S/20°56’E 73

34°30'5/20°55’E ¢. 70

34°40'S/21°35’E 73

34°24'S/21°55'E 59

34°28'S/21°45’E 70

32°41°7'S/17°58'5/E 9

35°03/S/21°50’E TIO

34°25°5'S/21°50°2’E 60

34°42°2’S/20°25:0’E gI

34°12’S/18°22’E 22

33°56'S/25°36’E

Position Depth (m.)

34°09°3/S/18°17°5’E 43

34°09°4'S/18°16-5’E 75

33°05°6/S/17°54°5/E 64,

33°07°5'S/17°52°5'E 86

33°06°5’S/17°55°4E 86

34°05’S/18°21’E II

33°08/S/17°46’E 157

33°07°3/S/17°57°5'E 26

33°50°3'S/18°23-2’E 15

34°01°7'S/18°14-7’E 100

33°55°8'S/18°21-3’E 37

32°52’S/18°25’E 29

LisT OF SPECIES

Family Tubulariidae

Tubularia warrent Ewer, 1953.

Family Myriothelidae

Bottom

S, Sh

, Sh, St

prot

R,S

S,S

Myriothela tentaculata n. sp.

Family Corynidae

Staurocladia vallentini (Browne, 1902).

434 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Solanderiidae

Solanderia procumbens (Carter, 1873).

Family Bougainvilliidae

Bimeria vestita Wright, 1859. Dicoryne conferta (Alder, 1856).

Bougainvillia macloviana (Lesson, 1836). Rhizorhagium robustum (Warren, 1907).

Bougainvillia sp.

Family Clavidae

Clava sp. Merona cornucopiae (Norman, 1864).

Family Eudendriidae

Eudendrium ?capillare Alder, 1856. Eudendrium deciduum Millard, 1957.

Eudendrium ?carneum Clarke, 1882. Eudendrium ramosum (Linn., 1758).

Family Hydractiniidae

Hydractinia altispina Millard, 1955. Hydrocorella africana Stechow, 1921.

Hydractinia kaffraria Millard, 1955. Podocoryne carnea M. Sars, 1846.

Family Pandeidae

Leuckartiara octona (Fleming, 1823).

Family Aequoreidae

Aequorea africana n. sp.

Family Lovenellidae

Lovenella chiquitita Millard, 1957.

Family Haleciidae

Halecium beanii (Johnston, 1838). Halecium halecinum (Linn., 1758).

Halecium delicatulum Coughtrey, 1876. Halecium ?muricatum (Ellis & Sol.,

1786).

Halecium dichotomum Allman, 1888. Halecium tenellum Hincks, 1861.

Family Campanulariidae

Campanularia hincksi Alder, 1856. Clytia hemisphaerica (Linn , 1767).

Campanularia integra MacGill., 1842. Clytia hummelincki (Leloup, 1935).

Campanularia laminacarpa n. sp. Clytia paulensis (Vanh6ffen, 1910).

Campanularia ?mollis (Stechow, 1919). Obelia dichotoma (Linn., 1758).

Campanularia morgansi Millard, 1957. Obelia geniculata (Linn., 1758).

Family Tubulariidae

Tubularia solitaria Warren, 1906.

Tubularia solitaria Warren, 1906: 83, pl. 10, 11. Millard, 1957: 179.

Records. West coast: A 122. LB 166 (recorded by Day, 1959). SB 153U.

South coast: L 172.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 435

Tubularia warreni Ewer, 1953

Tubularia warreni Ewer, 1953: 351, fig. 1-4. Millard, 1959a: 299. Millard, 1959): 240.

Records. West coast: LB 296A, 386A (reported by Day, 1959). South

coast: CPR 9G. KNY 176A (reported by Day, Millard and Harrison, 1952,

as T. ?betheris).

Description. This material conforms to previous descriptions of the species

with the exception of the CPR sample which differs in its paler coloration.

The general effect, when alive, was a creamy colour, with pink tinges in the

manubrium and gonophores. It is obviously a young colony, reaching a

maximum height of 1-75 cm., and the largest hydranths measuring only about

2°5 mm. in length and 1 mm. in basal diameter. The blastostyles are unbranched

and few in number (5-9) though fully mature gonophores are present. Apart

from the colour there is nothing to distinguish it from T. warrent.

Family Myriothelidae

Monocoryne minor n. sp.

Bigs

Material. The holotype, SCD 215C, is a single specimen detached from

its substratum and the only specimen known. Most of the colour has gone,

though signs of a dark-red pigmentation are visible on the gonophores. South

African Museum registered number: SAMH 41o.

Description. Polyp measuring approximately 5 mm. in length, though

somewhat contracted and curled up. Basal part of body (1 mm.) covered in

transparent perisarc which is continued proximally in a number of threads and

filaments, some of which appear to be filled with living coenosarc. Remainder

of body cylindrical, bearing about 110 capitate tentacles. Tentacles solitary

or arranged in groups of 2, 3, or 4 united at their bases. Length of tentacles

variable: sometimes the middle one of a group of three is the largest as in

M. gigantea, but sometimes all members of the group are equal in length and

sometimes the first of a group of four is longest.

Eleven gonophores, apparently male, borne irregularly on the body of

the polyp, the largest about 0-3 mm. in diameter. Gonophores pear-shaped,

not distinctly demarcated from pedicel.

Nematocysts. At least three kinds visible in smears of preserved material:

(1) Desmonemes, the most abundant type. Capsule oval, thread with

about 2 complete coils, on which a spiral ridging is visible. Size

variable: 9:0-13°5 x 6-3—-10°8 p.

(ii) Stenoteles, fairly common. Capsule egg-shaped, with shaft occupying

about half length. 15:3-18-0 X 13:0-15°3 yp.

(iii) Undetermined heteronemes, rare. Capsule elongated, with shaft

occupying about 2 length. 16-2-18-9 x 6-3-7°6 pu.

Remarks. As there is only one specimen available no sections were cut and

accurate measurements could not be made without damage. A few detached

436 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 1. Monocoryne minor n. sp.

A. The whole animal (holotype).

B. A typical group of 4 tentacles.

C-F. Nematocysts. (C, stenotele. D, undetermined heteroneme. E—F, desmonemes).

tentacles were used for examination of nematocysts. The specimen is fairly

well preserved, but there is no knowing how many tentacles may have been

broken off during handling, or whether the base was originally attached to a

firm substratum. Since the tip of the body is curled over, the mouth could not

be observed.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 437

The arrangement of the tentacles differs from that in the only two known

species of Monocoryne, namely M. gigantea (Bonnevie, 1898) and M. bracteata

(Fraser, 1941), in both of which the middle one of a group is the largest. It is

also much smaller than either of them. The gonophores are not seated in the

axils of the tentacles but distributed at random over the body.

Myriothela capensis Manton, 1940

Mbpriothela capensis Manton, 1940: 276, pl. 1 (figs. 12, 13), pl. 3 (fig. 27), figs. 7, 8b, 9. Millard,

1957: 186.

Records. West coast: CP 331 (one almost mature male specimen attached

to weed). LAM 52F (one mature female specimen attached to weed).

LU 118Y (two mature male specimens torn from their base and two young

specimens on crustacean appendage).

Description. The material from Luderitz Bay (LU 118Y) had retained its

colour after about four months in spirit. The whole distal end of the body

was a vivid splash of colour—the capita of the body tentacles and tips of the

gonophores bright magenta, shading to pink on the stalks and colourless on the

base of the polyp. Specimens preserved for a longer period are pale pink or

completely colourless.

Largest specimen 1-6 cm. in length. Structure and nematocysts as described

by Manton. One mature female specimen with actinulae larvae (one gonophore

on opening contained three larvae). Female gonophores reaching 0-9 x 1:0

mm.

Myriothela tentaculata n. sp.

Pige2:

Material. WCD 7Q: 5 specimens from west coast attached to encrusting

polyzoan, two of them rather badly damaged and the others in various states

of preservation. The best preserved was selected as the holotype and another

was used for sectioning. South African Museum registered number of holotype:

SAMH 411.

Description of holotype. Total length 2-5 cm. No colour left in specimen except

for a circle of dark-red spots round the distal end of each gonophore.

Basal region of hydranth (8 mm.) naked and drawn out into an irregular

shape, bearing about g chitinoid, flattened, adhesive discs attached to the

substratum.

Above this a single whorl of 17 long, tapering blastostyles reaching a

maximum length of about 2 cm. These are generally somewhat coiled and bear

a superficial resemblance to the tentacles of an octopus. They are the most

distinctive feature of the species. Blastostyle unbranched, bearing 4-6 gono-

phores in the proximal 5 mm., of which the most distal is the oldest and the

others in various stages of development. Capitate tentacles present amongst

the gonophores and others (about 25) scattered irregularly over the distal

region of the blastostyle. Tentacles rather poorly developed and resembling

boot-buttons. Gonophores male, spherical, reaching a diameter of about

438 ANNALS OF THE SOUTH AFRICAN MUSEUM

2 mm., each with a very short pedicel and 8-10 pigmented spots around distal

end.

Distal region of hydranth (about 2 of length) covered with densely packed

capitate tentacles and bearing a terminal mouth. Tentacles well-developed,

with slender stalks and large capituli, but more poorly developed in the region

of the blastostyles.

Nematocysts. Four types distinguishable from smears and sections:

(i) ‘Heteronemes’ (fig. 2D), elongated and banana-shaped, with an

axia] body which stains redin Mallory’s stain and a fine much-coiled

thread. 44°1-45:9 X 8-1 p.

(ii) Desmonemes of two sizes (fig. 2E—H), the larger 17-1-18-0 X 12-6—

14°4 », the smaller 10-2-10°8 x 7-2—7-7 w. In both, part of the thread

stains red in Mallory’s stain, the rest remains unstained. It bears a

distinct spiral marking which may indicate the presence of barbs.

The larger form contains a thread in 3 complete longitudinal coils

and the smaller a thread in 2 coils.

Steroteles (fig. 2C). Oval capsules containing a butt which stains red

in Mallory’s stain and increases in diameter towards the base. The

thread is coiled transversely in the lower half of the capsule and forms

an opaque mass in this region. A few were found exploded and showed

a characteristic butt, though the thread and spines had been broken

off. 14°4 X 10°8 p.

(iv) Atrichous isorhizas (fig. 2B). Elongated capsules with the contents

difficult to discern, but apparently containing a long tangled thread

which stains faintly blue in Mallory’s stain. 15:3-19°8 X 4:0—-7:2 p.

(iii

Histology. The hydranth body wall (fig. 2M) is very similar in structure to

that of M. capensis. It is 110-180 p thick, with a fairly thin layer of mesogloea

10-20 2 thick from which slender lamellae bearing muscle fibres project into

the ectoderm. The lamellae arise at intervals of approximately 14 » and are

40-80 » deep. From the endoderm arise villi anything up to 1 mm. in depth,

each containing a very thin supporting lamella of mesogloea. The apical cells

of the villi contain dark-staining granules, but the remainder are clear and

vacuolated. 85 villi were counted in one section. In the region of the mouth the

granular cells are more numerous and many cells contain droplets of a yellowish

material.

The body tentacles (fig. 2K) are exactly like those of M. capensis as

described by Manton (1940), with an apical pad of mesogloea fibrils about

50 w in thickness and a central cavity which passes into the stalk without

constriction. The endoderm of the stalk is separated from that of the hydranth

body by a thin layer of mesogloea. The capita of the tentacles are richly armed

with nematocysts of all 4 types, of which desmonemes are the most numerous.

The blastostyle is the most distinctive structure in the species. In the

proximal region where the gonophores arise the endoderm bears villi which

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 439

Fic. 2. Myriothela tentaculata n. sp.

A. The whole animal (holotype).

B-H. Nematocysts. (B, atrichous isorhiza. C, stenotele. D, ‘heteroneme’. E and F, large

desmoneme in side view and end-on view. G and H, small desmoneme in end-on view and

side view.)

t.s. blastostyle in proximal region with young male gonophore arising from it.

l.s. body tentacle.

l.s. blastostyle tentacle.

t.s. through body-wall of hydranth in distal region including the origin of 2 tentacles.

(e, ectoderm. enb, endoderm of bell. m, mesogloea. ml, mesogloeal lamella. mp, apical pad

of mesogloeal fibrils. n, nematocysts. se, subumbrella ectoderm. sp, spermatogenic cells.

v, endodermal villi).

east ate

440 ANNALS OF THE SOUTH AFRICAN MUSEUM

project into the central cavity and fill it almost completely (fig. 2J). The cells

of the villi are densely packed with granules which stain red in Mallory’s

stain, and many of the granules float freely in the interstices between the villi.

The mesogloea bears lamellae about 20 uw deep on its ectodermal surface.

The ectoderm contains many developing nematocysts which are particularly

abundant around the origins of the tentacles. In the slender distal part of the

blastostyle ectoderm and mesoderm are normal and well-preserved, but the

endoderm appears to be largely senile. Although remains of villi and their

mesogloeal lamellae can be recognised, most of the internal cavity is filled with

dead cells and fragments of cells most of which have an opaque yellowish

coloration.

The blastostyle tentacles (fig. 2L) are similar to those of the body, but each

has a much shorter stalk and a thinner pad of mesogloeal fibrils (about 20 p

thick in maximum). The lumen is very narrow and the endoderm separated

from that of the blastostyle by a basal lamella of mesogloea. The superficial

layer of the capitum is densely packed with nematocysts, including

‘heteronemes’ and atrichs, while desmonemes and stenoteles appear to be

entirely absent.

The structure and development of the gonophore is similar to that in

other species (Briggs, 1929, 1931; Manton, 1940). Only male gonophores are

present and the largest one sectioned is 1:24 mm. in diameter with a layer of

spermatogenic cells 0-24 mm. thick around the spadix. It is apparently almost

mature, for a velar invagination is present in the exumbrella ectoderm though

not yet perforate. The endoderm of the spadix bears rounded villi which

project into the central cavity which is continuous with that of the blastostyle.

Remarks. Of the six southern hemisphere species listed by Manton, (p. 288)

this species is most closely related to M. capensis. It differs from it in its larger

size, its adhesive structures, which are less obviously tentacular, and its long

and tendril-like blastostyles.

The nature of the nematocysts supports the establishment of a new species.

All are larger than the corresponding types in M. capensis, the banana-shaped

‘heteronemes’ being enormous, about 4 times those of M. capensis and over

twice those of M. penola and M. cocksi. The presence of atrichs in the hydranth

is unusual since in other species this type is confined to the actinula. The term

‘heteroneme’ has been used following Manton, although the only evidence

that this type possesses a distinct butt is that of Allman (1876), and needs

confirmation. No haplonemes could be identified in this material, and if

present cannot be distinguished from stenoteles.

Family Corynidae

Genus Bicorona nov. gen.

Diagnosis. Corynidae with firm perisarc and upright, monopodially

branched stems. Hydranths with two whorls of capitate tentacles. Gonophores

in the form of fixed sporosacs borne on the body of the hydranth.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 441

Bicorona elegans nov. sp.

Fie. 3

Types and records. Holotype: SB 161X (South African Museum registered

number: SAMH 412), from Saldanha Bay on the west coast. Other records:

A 116, 139, 353 (west coast); L 44, 56 (south coast).

Description of holotype. A luxuriant colony of upright branching stems

reaching a maximum height of 5-8 cm., growing in the lower intertidal region

of the shore.

Hydrorhiza embedded in sponge, branching and giving rise to upright

stems. Stem unfascicled, with monopodial growth and terminal hydranths.

Branches arising alternately, either bearing one terminal hydranth, or

rebranching in a manner similar to the stem. Stem and branches covered

with firm perisarc which is closely annulated throughout except for a smooth

area on the origin of each branch. Hard part of perisarc terminating just

below hydranth, but continued as a gelatinous layer onto the base of the

hydranth.

Hydranth Tubularia-like, with two whorls of capitate tentacles. Aboral

tentacles 10-21 in number in mature hydranths, usually with long and short

tentacles alternating. Oral tentacles short, 4-7 in number. ‘Neck’ region of

hydranth (below aboral tentacles) with indistinct longitudinal striations.

Gonophores in form of fixed sporosacs, borne on approximately 7 short

blastostyles which form a whorl just distal to the aboral tentacles. Gonophores

oval to spherical, 2-4 to a blastostyle, the oldest one terminal.

Histology (fig. 3K). Stem normal, with no endodermal canals, terminating

in a swollen region in the last perisarcal segment.

‘Neck’ region of hydranth with longitudinally ridged ectoderm, surrounded

by thick gelatinous sheath continuous with the perisarc of the stem.

Hydranth expanding at level of aboral tentacles, with spacious enteron.

Ectoderm of hydranth normal. Endoderm specialized: immediately below

aboral tentacles thickened and granular but with no indication of parenchyma,

in the region between the two whorls of tentacles folded inward around sup-

porting strands of mesogloea in the nature of endodermal villi, in region of

hypostome thickened.

Tentacles solid, with central core of ‘chordal’ endoderm. Aboral tentacles

with no ‘supporting lamella’ of mesogloea, and endoderm continuous with

that of hydranth. Oral tentacles attached obliquely to hydranth body, with a

‘supporting lamella’ of mesogloea which may be incomplete in its aboral

region.

Blastostyle hollow and tubular. Gonophores with a hollow spadix, without

subumbrella cavity or radial canals. Female gonophore containing many

small eggs arranged in a single series around the spadix (counts give numbers

varying from 29 to 84). Eggs 0:05-0:10 mm. in diameter. Male gonophore

containing a thick mass of spermatogenic cells.

Nematocysts (fig. 3H, J) all stenoteles, of 2 sizes: large ones reaching

442 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 3. Bicorona elegans n. sp.

A. Two complete stems from the holotype.

B-G. Stages in the development of the hydranth (B-F from A 139, G a mature hydranth

from the holotype).

Hand J. Small, and large, stenotele from A 116.

K._ 1.s. hydranth with female gonophores from L 44.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 443

26 X 17, small ones reaching 13:5 X 7 uw (measurements from preserved

material, undischarged).

Measurements (mm., preserved).

(NV.B. the holotype material is well expanded, A 116 less so.)

holotype A 116

Stem, diameter. : : : : : . 0°21-0°35 0:26-0°43

Hydranth, length from end of ‘hard’ perisarc to

hypostome . : : : . 0°96-1°95 1°29-1°73

Gonophores, length : eEKeAcnin gy Ordy7 0°50

maximum diameter : i ceaching 0-44 0:36

Remarks. The material chosen as the holotype is a well-developed, mature

colony, with presumably the maximum number of tentacles on the polyps.

One of the other samples (A 139) contains shorter stems with younger, well-

relaxed polyps, permitting observations on the origin of the tentacles (fig.

3B-F).

In the young polyp bud, two whorls of capitate tentacles appear simul-

taneously, 4-7 oral and approximately 7 aboral. These are at first short and

knob-like, but they lengthen as development proceeds, especially the aboral

ones. The number of oral tentacles is not increased, but before long a new

series of aboral tentacles appears, about 7 in number, alternating with those

of the first series and at a slightly Jower level on the polyp. Polyps at this stage

of development (fig. 3F) thus appear to have three whorls of tentacles, one

oral and two closely alternating aboral. When the second series of aboral

tentacles is about half the length of the first, young blastostyles make their

appearance, one in the axi] of each aboral tentacle of the first series. The oldest

polyps in this particular sample have reached this stage, but the appearance

of a third series of aboral tentacles would bring the number up to about 21

(the maximum number observed in the holotype). In the fully mature polyps

of the holotype and other samples, however, the aboral tentacles are arranged

in a single whorl and at one level (fig. 3G). Whether this is due to further

growth in circumference of the polyp or to differences in the state of contraction

can only be determined from living material.

Annandale (1915) established a new genus Dicyclocoryne for a species

previously described in 1907 as Syncoryne filamentata, in which the generic

diagnosis is the presence of two whorls cf capitate tentacles. But D. filamentata

bears free medusae and thus, following the practice adopted in these papers,

cannot be included in the same genus as forms with fixed sporosacs. There are

also other differences from the present species (e.g. unbranched stems, smooth

perisarc, different position of gonophores) which make the establishment of a

new genus for this material desirable, viz. Bicorona.

Dicyclocoryne and Bicorona appear to be more closely related to the Corynidae

than to any other family of Capitata, and this is supported by the presence

of only one category of nematocyst. Although stenoteles and desmonemes have

444. ANNALS OF THE SOUTH AFRICAN MUSEUM

been reported from medusae of this family, only stenoteles (large and small)

are known from the hydroids (Russell, 1938, and Millard, 1959b, for Sarsia

eximia). In the Pennariidae on the other hand, with which Bizcorona also has

certain affinities, 3 or 4 different categories of nematocyst occur (Weill, 1934,

and Millard, 1959a).

Sarsia eximia (Allman, 1859)

Coryne sp.: Millard, 1957: 179.

Sarsia eximia. Russell, 1953: 50, pl. 2 (fig. 3), figs. 17a, 18A, B. Millard, 1959d: 241. Kramp,

1959: 79, fig. 15. Kramp, 1961: 27.

Records. West coast: LB 556A. LU 113D. SH 433D, 436A. South coast:

CPR oF. KNY 165E (recorded by Day, Millard and Harrison, 1952, as

Syncoryne Peximia).

Remarks. Living material from Table Bay docks (SH 436A) and Langebaan

(LB 556A) was kept in the laboratory where it released medusae. At liberation

these measured 0-9 mm. in depth and o-8 mm. in diameter, with a hypostome

of 0-3 mm. in length. After 3 days the size had increased to 1:1 mm. depth and

I°2 mm. diameter, with a hypostome of 0-4 mm. At this stage the teniacles

reached a length of about 1-8 mm. when extended and bore about 12 clusters

of nematocysts. The structure was similar in every way to that of S. eximia.

The medusa of S. eximia has not previously been recorded from South Africa.

Staurocladia vallentini (Browne, 1902)

Cnidonema capensis Gilchrist, 1919: 509, pl. 30. :

Staurocladia vallentini: Browne and Kramp, 1939: 274, pl. 14 (figs. 3-4), pl. 15 (fig. 4), pl. 19

(fig. 2).

Cnidonema vallentini: Ralph, 194.7: 414, pl. 35 (figs. 1-6).

Records. West coast: LB 266A, 398A (recorded by Day, 1959, as Cnidonema

vallentini). A 40 (recorded by Bright, 1938, as Eleutheria vallentint).

Description. Material from Langebaan (LB) consists of numerous young

medusae found crawling on weed (Gracilaria confervoides), most of them under-

going asexual reproduction. Medusae reaching a maximum diameter of about

I mm., and bearing 20-30 tentacles. Aboral branch of tentacle with 4-5

nematocyst batteries, of which one is terminal and the rest dorsal. No gonads

present.

Material from Oudekraal (A) includes two larger medusae, reaching a

diameter of 3 mm. and bearing gonads. Specimens rather damaged, but

apparently bearing over 40 tentacles. Nematocyst batteries as in LB material.

Hydranths not as yet found.

Family: Solanderiidae

Solanderia procumbens (Carter, 1873)

Fig. 4 and Plate I

Ceratella procumbens Carter, 1873: 10.

Ceratella spinosa Carter, 1873: 12.

Solanderia atrorubens: Marshall, 1892: 12, pl. 5, pl. 7 (figs. 2-4).

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 445

Solanderia procumbens: Vervoort, 1962: 535.

Solanderia spinosa: Vervoort, 1962: 535.

Non Dehitella atrorubens Gray, 1868.

General. The material of this species is divisible into two forms on general

appearance (here designated as form A and form B), which do not seem to

be specifically distinct, but are kept separate in case they should later prove

to be so. In the description which follows the skeleton of the largest specimen

of each (WCD 158F and E respectively) is described and the soft parts of the

best preserved (SAMH 247).

Records

Form A: West coast: WCD 158F. South coast: SAMH 247. TRA 590A.

Form B: West coast: WCD 158E. South coast: SCD 154A.

Description, form A. The largest colony (Plate IA, left) a magnificent

fan-shaped specimen 33 cm. high with a spread of 38 cm. Hydrorhiza 5 cm.

broad, and flattened below for attachment. Main stem flattened in the plane

of branching and expanding from a base 2-4 cm. wide and 1-2 cm. thick to a

broad leather-like blade 5 cm. wide and 0:5 cm. thick; beyond this giving rise

to a number of large branches, which in their turn branch and rebranch.

Branching in one plane, with a tendency for the branches to bend to one side

and give off more sub-branches on this side, suggesting the influence of a

strong current during growth. Method of branching variable, often dichoto-

mous, sometimes alternate and sometimes unilateral. Smaller branches quite

round in section, comparatively thick and stumpy (about 2 mm. in diameter),

with rounded tips.

Fibrous meshwork of main stem reticulate, with rounded, rectangular,

or hexagonal apertures between the trabeculae, but no marked longitudinal

arrangement. Trabeculae of unworn areas bearing numerous spiny structures

0-3-0:4 mm. in height, some of them flattened and spatulate and resembling

abortive hydrophores with one or two supporting ribs (fig. 4D).

Fibrous meshwork of branches arranged in a more definite fashion, in

which the longitudinal trabeculae predominate. Groups of trabeculae often

raised up into prominent longitudinal ridges surmounted by blade-like crests

parallel to the long axis of the branch (Plate IB, lower right corner). Spines

reduced to tubercles on the branches and absent on the smallest subdivisions.

Hydrophores numerous and scattered irregularly over surface of branches.

A typical one consisting of a scoop-shaped bracket supported on its lower

surface by longitudinal ribs continuous with the trabeculae of the branch

(fig. 4B). Ribs normally not reaching margin, which is smooth and rounded.

Hydrophores, however, subject to much variation, some being bilobed (fig. 4C),

some divided into two halves (one on each side of the hydranth) and some

represented by a laterally seated hemihydrophore only. Badly eroded hydro-

phores with margin worn down between the supporting ribs (fig. 4H). Normal

hydrophores approximately 0-5 mm. in length, and 0-5 mm. in basal width

tapering to 0-3 mm. at margin.

446 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hydranths arising from coenosarc above hydrophores and, rarely, directly

from the stem with no protective supports whatever; bearing approximately

15 capitate tentacles (but difficult to count due to poor preservation).

Gonophores arising directly from coenosarc of stem, spherical, shortly

stalked. Immature male gonophores present on two specimens, the largest

measuring 0°34 mm. in diameter and 0-32 mm. in length; containing several

layers of spermatogenic cells around a central spadix, and 4 distinct radial

canals.

Colour: stem and larger branches dark brown, occasionally with a purplish

tinge, shading to yellowish-brown on the smaller branches.

Form B. Colonies smaller than form A, the largest reaching 19:5 cm. in

height and 15:5 cm. in spread, with more slender branches and lighter coloration

(Plate 1A, right). Main stem rounded in section. Hydrophores on smaller

branches restricted to two sides, but irregularly scattered on larger ones. The

‘double’ type of hydrophore predominant (fig. 4E, G), bilobed or single ones

rare (fig. 4F).

Nematocysts. Three types present in smears of preserved material:

(i) Large oval stenoteles (fig. 4R, S). Fairly common. Undischarged

capsules with a broad, central shaft and a long thread coiled trans-

versely in the lower half. Discharged capsules with a broad, slightly

tapering butt armed with 3 large spines and several spiral rows of

smaller spines. Thread broken off in all examples seen.

15°3-24°3 X 11°7-18-0 pw

(ii) Small oval heteronemes (fig. 4P). Abundant. Undischarged capsules

with a central shaft and a thread which appears to be irregularly

coiled. A single discharged and rather distorted capsule bore a short,

swollen butt and a number of spines. 8-1-8-7 « 6:3 »

(iii) Elongate-oval capsules of unknown category (fig. 4Q). Rare. Only

discharged capsules seen with no sign of butt or thread. 15:3 X 5:4 ph

Remarks. It is felt that form A and form B represent different growth-forms

of the same species, form B including younger colonies which have possibly

grown more rapidly. Their main claim to distinction is the nature of the

hydrophores which are predominantly ‘single’ in form A and ‘double’ in form

B, although both types can be found in all colonies. There is no difference in

the nematocysts. The ‘double’ hydrophore is in some ways similar to the

‘spines’ which flank the hydranth in S. secunda (Inaba). However, diagrams

by Stechow (1909, pl. 4, fig. 7) and Vervoort (1962, figs. 2b, 7) show that the

latter are little larger than the area of one mesh of the skeleton, whereas in

the present material they approximate to the area of 4 meshes.

Carter’s dried type material of Ceratella procumbens was available for com-

parison in the British Museum. It includes one specimen from the Cape of

Good Hope (reg. no. 1867.3.22.1) and three from Natal (reg. no. 1872.8.1.1).

The largest is one of the latter and measures 26-5 cm. in height with a spread

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 447

Fic. 4. Solanderia procumbens (Carter).

A-C. Hydrophores from form A, WCD 158F (A and B single, C bilobed).

D. A portion of the older part of the stem in WCD 158F showing spines.

E-G. Hydrophores from form B, SCD 154A (F single, E and G double).

H. A typical eroded hydrophore from False Bay.

J-N. MHydrophores from Carter’s type material (J and K single, L double, M and N eroded).

P-S. Nematocysts (P, a small heteroneme. Q, undetermined type. R and S, large stenotele).

448 ANNALS OF THE SOUTH AFRICAN MUSEUM

of 11 cm. The main stem is 1-8 cm. wide and 0-6 cm. thick. This is evidently

the specimen whose measurements were quoted by Carter in 1873. I select it

as the lectotype.

In spite of the fact that the type material is battered and obviously beach-

worn the resemblance to form A described above is unmistakable. Here too

the hydrophores are predominantly single (fig. 4J, K) though bilobed and

double ones also occur (fig. 4L). In the older and badly worn parts the

hydrophores are eroded so that the supporting ribs project and give a

spinous appearance to the margin resembling that in S. fusca (Gray)

(fig. 4M, N).

S. spinosa (Carter, 1873), is here considered to be a synonym for

S. procumbens. The dried holotype of this species from Port Natal is also present

in the British Museum (reg. no. 1872.8.1.17). It is less beach-worn than the

type material of S. procumbens and this would account for the minor differences

said to distinguish the species. Its general appearance is similar to form A

described above though most of the hydrophores are of the ‘bilobed’ type.

The ‘spines’, which are the main specific character, are present only on the

older part of the colony as in the present material, and are comparatively

delicate structures which would easily be rubbed off by sand erosion. This

would account for their absence in the type material of S. procumbens.

Another species which possesses well-developed hydrophores is S. fusca

(Gray, 1868), from Australia, described and figured by Spencer (1892). Gray’s

type material is present in the British Museum (reg. no. 1884.12.6.15-16; the

first of these designated as lectotype by Vervoort (1962: 533). The most

obvious characteristic of this species is the nature of the hydrophores which

have spiny margins due to the projection of the supporting ribs beyond the

edge. That this effect is not in this case the result of wear is evident from the

fact that spiny hydrophores are present in the youngest and thinnest branches

and, according to Spencer, also in the living animal.

S. atrorubens (Gray, 1868), is closely related to S. fusca and possibly a

synonym, although Vervoort, 1962, considers that it can be distinguished by

its method of growth. An examination of the type material in the British

Museum, probably from Australia (of which no. 1962.4.14.1 has since been

designated as lectotype by Vervoort, 1962, p. 535), showed that it has the

same spiny hydrophores.

The fact that hydrophores of S. procumbens may acquire a spiny appearance

as the result of erosion has probably led to confusion between this species and

S. fusca and S. atrorubens. Marshall’s record of S. atrorubens from Port Natal can

be included in the synonymy of S. procumbens as his figures show that the edges

of the hydrophores are smooth. Other species recorded from South Africa,

including S. labyrinthica (Hyatt, 1877), and S. rugosa Marshall, 1892, and also

Brazier’s record (1887) of S. atrorubens are insufficiently described but are

possibly all synonyms of S. procumbens.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 449

Family Bougainvilliidae

Bimeria vestita Wright, 1859

Fig. 5 A-F

Bimeria vestita: Hincks, 1868: 103, pl. 15 (fig. 2). Allman, 1872: 297, pl. 12 (fig. 1-3). Hamond,

1957: 297, figs. 3-4.

Leuckartiara vestita forma nana: Vervoort, 1946a: 294.

Records. South coast: MB 88L. SAMH 158, 177, 233, 235, 327, 337, 343;

361. SCD 85T, 118M, 347F.

Description. Colonies epizootic on other hydroids, and reaching a height

of 3-8 mm. Stem upright, bearing from 1 to 6 or 7 alternately arranged

hydranth pedicels. Pedicels occasionally rebranching. Perisarc annulated at

base of stem, on origin of pedicels and at other irregular intervals, covered

throughout with adherent particles. Stem and pedicels narrower at base than

at distal end. Perisarc continued over the hydranth, sheathing the tentacles

for part of their length and covering the greater part of the hypostome. Tentacles

varying in number from about 10 to 16.

Gonophores borne on the stem and hydranth pedicels, each on a short

annulated pedicel, completely covered by a thick coat of gelatinous perisarc

male and female on separate colonies. Male sporosac elongated-oval, with

branching spadix. Female sporosac oval to spherical, bearing a single terminal

ovum, which develops zm sztu into a planula larva (fig. 5 B—D).

Measurements (mm., preserved).

Stem, diameter. : : ‘ : . 0:04-0:09

Pseudohydrotheca, length d : : : . 0*21-0°49

diameter . : ; ; l : 1 Of12—0:20

Gonophore, female, length (without perisarc) . . reaching 0-16

diameter . : x : . reaching 0-16

male, length (without perisarc) : . reaching 0°33

diameter . 5 : : . reaching O12

Remarks. These colonies are very similar to those described by Hincks,

Allman and Hamond, although they are not so richly branched as some of

Allman’s specimens. The size of the hydranths and the thickness of the stem

appears to be less than in most descriptions, though it corresponds well with

Hamond’s material.

The only previous record of this species from southern Africa is that of

Vervoort, 1946a, from Inhaca in Portuguese East Africa. Female gonophores

are apparently described here for the first time.

Bougainvillia macloviana (Lesson, 1836)

Perigonimus maclovianus: Vanh6ffen, 1910: 284, fig. 10.

Bougainvillia macloviana: Jaderholm, 1923: 3. Millard, 1959): 242, fig. 1 A-C. Vannucci & Rees,

1961: 69.

Records. West coast: SB 178D.

Description. Colony epizootic on other hydroids. Stems only 2 mm. in

450 ANNALS OF THE SOUTH AFRICAN MUSEUM

yee |

Fic. 5.

A-F. Bimeria vestita Wright. A, a typical stem. B—D, stages in the development of the female

gonophore (C and D drawn without the perisarcal covering). E, a contracted hydranth.

F, a male gonophore. (A-E from SAMH 361, F from SAMH 177).

G. Clava sp. Two hydranths sketched from living material (CP 646A).

H-J. Merona cornucopiae (Norman) from SCD 119R. H, part of the colony showing hydranths,

a single blastostyle and nematothecae. J, three nematothecae on a larger scale.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 451

height, slender, flexuous, branching irregularly, bearing medusa buds, of which

the oldest has its tentacles unfurled.

Remarks. In its general appearance this colony strongly resembles

Vanhéffen’s description of the species and also the material previously described

(Millard, 19590). The identification is strongly supported by the discovery

of abundant mature medusae of this species from the same area in Saldanha Bay

(identification by M. E. Thiel of the Zoologisches Museum, Hamburg). It is

interesting also that the two ships on whose hulls this species was recorded in

19596 both came from Saldanha Bay.

Bougainvillia sp.

Records. West coast: LB 542A. South coast: KNY 165D (reported by Day,

Millard and Harrison, 1952, as B.ramosa). SAMH 170, 245. SCD 190A,

281G. TRA 33Z, 150F.

Description. Stems slender, upright, weakly fascicled at the base in the larger

colonies and branching profusely in a more-or-less alternate fashion. Maximum

height 5-3 cm. Larger stems generally smooth, smaller branches wrinkled or

corrugated, particularly over the origins. Smaller colonies unfascicled and less

profusely branched. Medusa buds with 4 unbranched oral tentacles and 4

marginal bulbs, each with 2 black ocelli and 2 marginal tentacles.

Remarks. It is felt that this material could not be assigned to a species with

any certainty. The general growth-form is very similar in all the colonies,

though some are obviously older than others. Well-developed medusa buds

may be present on stems of only 3 mm. in height.

The stems are stiffer and more profusely branched than those of

B. macloviana, yet not so sturdy as those described as B. ramosa (Millard, 19596)

from ships’ hulls.

The material might well be included in B. ramosa forma musca Allman,

1864, yet no mature medusae of B. ramosa have so far been recorded from this

country, whereas medusae of B. macloviana do occur on the west coast (see

above).

Dicoryne conferta (Alder, 1856).

Eudendrium confertum Alder, 1856: 354, pl. 12 (figs. 5-8)

Dicoryne conferta: Allman, 1872: 226, 293, pl. 8. Jaderholm, 1909: 47, pl. 3 (fig. 6).

Records. South coast: SCD 133C.

Description. A dense colony growing on a gastropod shell occupied by a

hermit, reaching a maximum height of 2:5 cm. Hydrorhiza reticulate. Stem

unfascicled, branching irregularly, increasing slightly in diameter towards

distal end. Perisarc roughly corrugated throughout, but more definitely

annulated on origin of stem, terminating below hydranth when the latter is

expanded and covering the base of the body as a ‘pseudohydrotheca’ when

contracted. Hydranth long, with a single whorl of tentacles near distal end.

‘Blastostyles’ in the form of tentacle-less hydranths borne on stem and

452 ANNALS OF THE SOUTH AFRICAN MUSEUM

hydrorhiza. Male gonophores present, borne on lower region of blastostyle

either as a tight cluster or distributed along its length. The two tentacles of the

swimming sporosac clearly visible within the perisarcal covering of the

gonophore.

Nematocysts all small, 5-4 x 2-7 ph

Remarks. This appears to be the first record of the species from the southern

hemisphere. The appearance of the colony is very characteristic and closely

resembles that illustrated by Jaderholm in 1909.

Rhizorhagium robustum (Warren, 1907)

Parawrightia robusta Warren, 1907: 187, pl. 33 (figs. 1-5), figs. 1-4.

Records. South coast: L 177.

Description. A sterile colony growing on a coralline alga. Most stems bear

a single terminal hydranth, but many 2 or 3. In some cases a number of stolons

or stems (it is impossible to determine which) are twisted together simulating

a fascicled stem which is quite free from the substratum and reaches a maximum

height of 17 mm. Perisarc well developed and forming a very distinct

‘pseudohydrotheca’ over the base of the hydranth. Tentacles 18-22.

Family Clavidae

Clava sp.

Fig. 5G.

Records. West coast: CP 646A.

Description. Colony growing on a stone just below low tide level. Hydrorhiza

reticulate, penetrating into calcareous matter on stone, covered with a thin

layer of perisarc which forms a very low collar round the base of each hydranth.

Hydranths reaching a maximum height of 7 mm., creamy pink in colour

when alive, with 22 to 30 scattered filiform tentacles which reach a maximum

length of 1 mm. when fully extended.

Gonophores absent.

Nematocysts of two kinds:

(1) Microbasic euryteles, 0-67—0:72 X 0°22 pm.

(ii) Desmonemes, 0°45 X 0°32 p

Remarks. This species cannot be definitely identified in the absence of

gonophores. The trophosome is similar in size and all other characters to

C. multicornis (Forskal, 1775), forma genuina Broch, 1916. The sizes of the two

categories of nematocyst are also close to those given by Weill, (1934) for

C. squamata (= C. multicornis).

Merona cornucopiae (Norman, 1864)

Fig. 5 H,, J.

Merona cornucopiae: Rees, 1956: 499, figs. 1-3.

Records. South coast: LIZ 25P. SCD 119R.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 453

Description. Colonies growing on the bivalve Crassatella capensis Lamy. Of the

two samples recorded above the first consists of a single bivalve bearing a male

colony of about 160 hydranths. The second consists of three bivalves bearing

colonies of between 30 and 50 hydranths each, two of these colonies bearing

female gonophores. In each case the colony is restricted to one corner of the

shell.

Hydrorhiza in the form of an open reticulum at the margins of the colony,

but consolidated into a mat in the denser regions.

Hydranths with 16 to 20 scattered filiform tentacles, surrounded by sturdy

perisarcal tubes into which they can be completely retracted. Tubes slender at

base, expanding distally, often regenerated.

Gonophores borne in clusters on blastostyles which arise separately from

hydrorhiza, male and female on separate colonies. Blastostyle surrounded at

base by short collar of perisarc.

The hydrorhiza of one colony only (SCD 119R, male) bearing numerous

minute nematothecae. Nematotheca in the form of an asymmetrical perisarcal

funnel borne on a long and slender pedicel. Containing a cluster of large

nematocysts.

Measurements (mm.)

SCDri9gR LIZ 25P

Perisarcal tube, height . : : : - 1£-8354-25 2:15 —2°79

maximum diameter : : : : . 0°30-0°38 0:32-0:35

Blastostyle, total height . 4 : . 1:°27-1°41 0:63

Gonophore, length ‘ : : : : . O'17—-0°25 O°1Q—-0°35

maximum diameter : : : : . O°12—-0:20 0-12-0:21

Nematotheca, height ; : ‘ : . 0:03-0:06

maximum diameter : : : : . 0:07-0:09

length of pedicel : : : : . 0°22-0°54.

Nematocysts. At least two types present:

(1) Microbasic euryteles. Elongated capsules measuring 16-2 X 4°5 u

undischarged. Abundant in nematophores, scarce on hydranth

tentacles. A single discharged and rather damaged capsule observed.

(i1) ?Desmonemes. Ovoid capsules measuring 7:2 X 2°7-3°6 » undis-

charged. Abundant on hydranth tentacles, scarce in nematophores.

Remarks. This species has so far been reported only from northern seas.

It is a new record for South Africa and its presence here is surprising. Even

more so is the discovery of nematophores on the hydrorhiza There can be

no doubt about the identification of the species thanks to the detailed description

of living material by Rees and the characteristic habitat of the animal.

The nematophores, when first noticed, were thought to be the hydrothecae

of an epizootic species, but the microscope showed that they arose from the

same hydrorhiza as the clavid ‘host’, a fact which was convincingly proved

when the same nematocysts of identical measurements were found in both. A

454 ANNALS OF THE SOUTH AFRICAN MUSEUM

noteworthy point is that nematophores occurred in only one colony (male)

out of four which were similar in every way other than sex.

Family Eudendriidae

Eudendrium annulatum Norman, 1864.

Remarks. The only record of this species from South Africa is that of

Ritchie (1909) from the entrance to Saldanha Bay, and that a doubtful one.

In 1960 the opportunity offered to examine two slides of Ritchie’s material

from Saldanha Bay and to compare them with Norman’s preserved type

material of E. annulatum. In Ritchie’s material the stem is not so densely

annulated as in the usual conception of E. annulatum, although in the type

material of the latter smooth areas do occur in some parts of the larger branches.

More important is the fact that in Ritchie’s material bifurcating spadices could

be distinctly seen on some of the female gonophores, whereas in FE. annulatum

the spadices are unbranched (Broch, 1916, p. 62). Ritchie’s record of FE. annula-

tum from South Africa should thus be discarded, and the material is provisionally

placed in E. carneum (see p. 455).

Eudendrium ?capillare Alder, 1856.

Eudendrium capillare Alder, 1856: 355, pl. 12 (figs. 9-12). Broch, 1916: 62. Stechow, 19252:

202. Leloup, 1952: 124, fig. 63. Picard, 1955: 183.

Eudendrium parvum Warren, 1908: 272, pl. 45 (figs. 1-4), fig. 1.

Eudendrium ?parvum: Millard, 1959a: 305, fig. 1G, H.

Records. South coast: SCD 154K. SH 433A.

Description. SCD 154K: Stems unbranched or sparsely branched. Perisarc

annulated at origin of stem and branches and often at other irregular intervals.

Old female gonophores present on ‘blastostyles’ which arise from stem or

hydrorhiza. Pedicel of blastostyle corrugated, bearing 3-5 gonophores or their

empty capsules irregularly distributed near distal end. Gonophores covered

by transparent capsule which has a warty appearance possibly due to shrinkage.

Soft parts too badly preserved for nematocyst examination or tentacle counts.

SH 433A: Living material kept in laboratory for several weeks. Stems

unfascicled, but profusely branched and reaching a maximum height of 1-7 cm.

Perisarc annulated on origins of stems and branches and at other irregular

intervals. Hydranths orange-pink in colour with white tentacles and hypostome;

with 23-28 tentacles held alternately elevated and depressed. Various stages

of female gonophores present on ‘blastostyles’ which arise from the stem or its

branches. Pedicel of blastostyle corrugated. Young blastostyle with fully-formed

hydranth and a ring of gonophores around its base; each gonophore with an

unbranched spadix arching over a single egg. Older blastostyles with the

hydranth showing signs of reduction and the gonophores more irregularly

distributed, each gonophore without a spadix and bearing a single embryo

surrounded by a transparent capsule. Nematocysts of two types: large isorhizas,

27 X 11-5 4, and small heteronemes, 8 x 2-4 « (undischarged).

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 455

Remarks. Warren’s E. parvum is now considered to be a synonym for the

cosmopolitan EF. capillare. This conclusion is based on examination of —

(i) material of E. capillare from Marseilles bearing female gonophores

and kindly supplied by J. Picard in 1958.

(ii) material of E. parvum from Knysna deposited by Warren in the

British Museum (slides 22.3.6.104-107). This is obviously not the

holotype, which came from Park Rynie, but is labelled co-type.

Both male and female gonophores present.

Warren in 1908 summarised the differences between the two species. One

of these was the presence of 3-chambered male gonophores in E. parvum as

against two in F. capillare. But the basal chamber in E. parvum is very small

indeed and its development may well be a matter of degree. Moreover, Alder

in his original description of E. capillare mentions 2- or 3-chambered gonophores

(‘two or three capsules in linear series on each pedicle’).

Another difference was the presence of a terminal tubercle on the male

gonophore in E. parvum. This appears to be a variable character. It is present

in only some of the gonophores in Warren’s material from Knysna, and is

apparently present or absent in FE. capillare (Stechow, 19252).

Further, Warren mentioned the extension of the perisarc over the base

of the hydranth in E. parvum. This perisarc is very delicate and may well have

been missed in earlier descriptions of E. capillare. It is visible in Picard’s material

of the latter from Marseilles.

Although Warren did not describe female gonophores in the holotype of

E. parvum, they are present in his material from Knysna and are exactly like

those of E. capillare.

Finally, the mention of small nematocysts only in E. parvum by Warren,

1908, and Millard, 1959q, is in agreement with Picard’s statement (1955) that

only small microbasic euryteles occur in FE. capillare.

The presence of both large and small nematocysts in the present material

(SH 433A) is the only reason for the query in the identification. One is loath

to create a new species on the basis of nematocysts only, but one is forced to the

conclusion that either there are two species with exactly similar female gono-

phores or that L£. capillare has the potentiality of producing two kinds of

nematocysts, which is not always realised.

Eudendrium ?carneum Clarke, 1882

?Eudendrium annulatum: Ritchie, 1909: 70.

Eudendrium carneum: Vannucci, 1954: 101, pl. 1 (figs. 1-9), pl. 2 (fig. 8), pl. 4 (figs. 2-5). Millard

1959a: 302, fig. 1A-F.

Records. West coast: A 118. CP 336A.

Description. Fascicled, branching stems reaching a maximum height of

about 5 cm. Colonies more heavily annulated than is usual for the species,

the groups of annulations on the main stem more common and more extensive,

often with about 15 rings; hydranth pedicels usually completely annulated,

456 ANNALS OF THE SOUTH AFRICAN MUSEUM

though some with smooth areas. Hydranths with 15 to 24 tentacles.

Young female gonophores with bifurcating spadices. Male gonophores

2-chambered.

Nematocysts of two types: large isorhizas, 20 x gp, and small

heteronemes, 7 X 4p.

Remarks. This material appears to be a smaller and more closely annulated

form of E. carneum. The young female gonophores are exactly the same, though

completely mature ones with basket-shaped capsules were not present. The

male gonophores are 2-chambered, whereas material from the east coast had a

minimum of 3 chambers (Millard, 1959a). However, Vannucci (1954) has

described 2-chambered male gonophores in the same species.

The nematocysts are similar to those of E. carneum, though both types are

slightly smaller. The presence of large nematocysts excludes the material from

E. racemosum which also has a forked spadix in the female gonophore.

This material is very similar to that reported from Saldanha Bay by

Ritchie (1909) as E. annulatum(?). Ritchie’s material also has bifurcating

spadices. (See also p. 454.)

Eudendrium deciduum Millard, 1957

Eudendrium deciduum Millard, 1957: 184, fig. 2.

Records. South coast: LIZ 7R. MB 19R, 47N.

Description. No large nematocysts have been observed in. this species.

Small heteronemes (probably microbasic euryteles) similar to those of E. carneum

(Millard, 1959, fig. 1B) present, measuring 6-3-8-1 x 2-7—-3°6 » undischarged.

Eudendrium ramosum (Linn., 1758)

Eudendrium ramosum: Hincks, 1868: 82, pl. 13. Stechow, 19234: 83. Weill, 1934: 388, fig. 237.

Leloup, 1952: 127, fig. 64.

Records. West coast: AFR ooo2E. South coast: SCD 37R_ (dubious

identification).

Description. The first specimen (AFR o002E) has long, graceful stems

reaching 17:5 cm. Main stem fascicled, branches unfascicled and given off in a

pinnate fashion either in one plane or twisted into a spiral. Female gonophores

present, with unbranched spadix. Gonophore-bearing hydranths with reduced

tentacles.

The second sample (SCD 37R) includes portions of a colony with a more

bushy and stiff appearance and with both main stem and principle branches

fascicled. Female gonophores present, with unbranched spadix.

Nematocysts (from AFR 0002E)

(i) Small heteronemes, 7-2 x 2:7 , present on both body and tentacles.

(ii) Large isorhizas, 18-0-18-g x 7:2—7-6 » present on body only.

Remarks. The first of these specimens agrees well with published descriptions

of E. ramosum and there can be little doubt as to the identification. The appear-

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 457

ance and measurements of the two categories of nematocyst in the undischarged

state are also close enough to those given for the species by Weill (1934), the

small heteronemes probably being microbasic euryteles.

The identification of the second specimen is more doubtful due to the

different appearance of the colony and the fact that the hydranths were too

poorly preserved for nematocyst examination.

This is the first record of the species from South Africa.

Family Hydractiniidae

Hydractinia altispina Millard, 1955

Hydractinia altispina Millard, 1955: 215, fig. 1. Millard, 1957: 179.

Records. West coast: B 62, 92 (reported by Millard, 1955). LAM 50W,

59C. LU 59Z. SAMH 407. SB 153T.

Hydractinia kaffraria Millard, 1955

Fig. 6

Hydractinia kaffraria Millard, 1955: 217, fig. 2. Millard, 19594: 307.

Records. South coast: BMR 23h. BRE 111A. HAM 3Q. KNY g3o0P, 70E,

164, 270J. SUN 3N. (All reported by Millard, 1955.)

Remarks on living material. Specimens from Knysna Estuary were kept alive

on their host snails for a period of two weeks in the laboratory in January, 1956,

when the release of the sexual products from mature sporosacs was observed.

The ripe male and female sporosacs have the structure of degenerate

medusae, with radial and circular canals and rudimentary marginal tentacles,

but the sexual products are discharged while still attached to the gonozooids

and there is no active medusoid generation.

The male sporosacs when mature are practically spherical and measure

approximately 0-65 mm. in length and 0-63 mm. in diameter. The four radial

canals are distinctly visible and around the opening are 4-6 rudimentary but

distinct marginal tentacles. The cavity is filled with spermatogenic cells

attached to the central spadix. While still attached to the gonozooid irregular

powerful contractions of the whole bell expel the active spermatozoa through

the aperture. A cloud of massed spermatozoa surrounds the sporosac for some

time until finally dispersed by the movements of the hydroids and their host.

Once the sporosacs are empty they fall off and lie loose on the bottom, but

show no further activity. At this stage they are more oval than spherical, the

length being greater than the diameter.

The mature female sporosacs are larger than the male, measuring

approximately 0-90 mm. in length and 0-86 mm. in diameter. Radial canals

are visible but the marginal tentacles are very indistinct and not so well

developed as in the male. The ripe eggs separate from the spadix and lie loose

the cavity of the bell. Fertilization apparently occurs inside the bell, since

eggs contain either one or two nuclei immediately after discharge. The area

458 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 6. Hydractinia kaffraria Millard.

A. A female sporosac in the process of releasing eggs.

B. A male sporosac.

(Both from living material.)

around the aperture of the sporosac performs irregular contractions which

constrict the opening, but no complete contractions of the bell have been

observed. These may possibly occur sporadically in the natural condition.

Under the microscope the pressure of the coverslip is sufficient to squeeze out

the eggs one by one, and the final fillip is provided by the contraction of the

margin. The diameter of the eggs is greater than that of the aperture, but their

elasticity allows them to be ejected with ease. Occasionally the sporosacs fall

off the gonozooids before all the eggs are discharged and continue to perform

gentle contractions of the margin, though no swimming movements have been

observed.

Hydrocorella africana Stechow, 1921

Fig. 7

HAydrocorella africana Stechow, 19256: 409. Millard, 1957: 183.

Records. West coast: A 405. AFR 801Q. CP 646B. HB 4D. SAMH 408.

WCD 12], 25C. South coast: AFR 866R, 967.0.V, 985E, 994M. SCD 114Q,

133B, 175U, 206R. TRA 33.00.D, 42K, 56C, 99E.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 459

Description. Common on shells of gastropods and hermits. Has been observed

on Turbo sarmaticus Linn., Argobuccinum argus (Gmelin), Fusus verruculatus Lam.,

Nassa speciosa A. Adams, and on shells occupied by the hermits Clibanarius sp.,

Pagurus arrosor (Herbst), Dzogenes costatus Hend. and Eupagurus placens Stebb.

Skeleton as described by Stechow (19255) with two grades of calcareous

ridged processes; the larger 5 mm. or more in height and covered with naked

coenosarc bearing gastrozooids and gonozooids; the smaller about 0-5-1 mm.

in height, scattered amongst the hydranths. Young colonies with smaller

processes only.

Living gastrozooids reaching 3 mm. in length when extended, with 5-12

extensile tentacles, of which 1 or 2 are usually much longer than the others.

Gonozooids reduced, reaching about 0-5 mm. in length in living material,

with about 6 rudimentary, knob-like tentacles, and each bearing 3 or 4 spherical

sporosacs in various stages of development. Male and female sporosacs on

separate colonies. Male sporosacs bearing the spermatogenic cells around a

central, hollow spadix, reaching 0-33 mm. in length and 0-33 mm. in maximum

diameter. Female sporosacs containing a single central egg surrounded by a

number of blind, hollow outgrowths from the basal spadix, reaching 0-36 mm.

in length and 0-56 mm. in maximum diameter. Developing gonozooid often

enclosed by curved laminar outgrowths of the skeleton, which form a sort of

basket-work imprisoning it.

Colour: skeleton and spines white, hydranths creamy white, female

sporosacs orange.

Nematocysts of two kinds: microbasic euryteles, 0:72 <x 0:27, and

desmonemes, 0°45 X 0°27 p.

Details of female sporosacs. Sections were cut to elucidate the rather unusual

structure of the female reproductive bodies.

Female gonozooids are normal in structure with the exception of the

tentacles which are reduced to knobs. In each gonozooid a number of young

ova are present in the endoderm of the central region of the body. The youngest

sporosac is nothing more than a bulge in the ectoderm into which one of the

enlarging ova has been pushed together with a few endoderm cells. In one

case such a bulge contains two ova, one large and one small. At a slightly later

stage the sporosac is well-defined and contains a single large ovum seated

on a low evagination of the gonozooid endoderm, which presumably represents

a reduced spadix. A few endoderm cells are flattened against the periphery

of the egg. There is no entocodon development and the ectoderm is

single-layered.

After this the ovum enlarges enormously and becomes packed with yolky

material. At the same time from the basal spadix several (usually 4) hollow,

finger-shaped processes grow out around the ovum until they partially surround

it. These may represent radial canals, but they do not communicate distally

and there is no sign of a circular canal or any other medusoid structure. This

is the oldest stage seen in sections, but presumably fertilization occurs in situ,

460 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 7. Hydrocorella africana Stechow.

A colony growing on a shell occupied by a hermit.

A section through colony and shell on a larger scale, showing many small processes, one

large one, and contracted hydranths.

Expanded gastrozooids sketched from a living colony.

Gonozooids bearing sporosacs, two female and one male.

Planula larva after release.

ts. 3 female sporosacs at different stages.

l.s. female gonozooid and sporosac. Young eggs visible in endoderm of gonozooid.

Grane he

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 461

for on several occasions gourd-shaped planulae have been found imprisoned

by the over-arching extensions of the skeleton.

Remarks. This species was described by Stechow (19250) with particular

emphasis on the skeleton, but it has never been illustrated. Stechow mentioned

male sporosacs only and failed to observe the gonozooids.

Podocoryne carnea M. Sars, 1846

Podocoryne inermis Allman, 1876: 255, pl. 10 (figs. 4-5).

Hydractinia carnea: Vervoort, 19460: 126, fig. 49. Millard, 1957: 181.

Records. West coast: LB 380C, 403C. SB 132N, 174M, 231Y, 267V,

2690B. TB 13, 14, 15, 21K. WCD 134C. South coast: KNY 212G. LIZ 3B,

24X. MB 25D. SCD 26E, 111C, 113E, 1261, 239G, 2585, 281E, 330F, 333F.

All on shells of the gastropod Nassa (Hinia) speciosa A. Adams.

Remarks. One of these colonies (SCD 26E) is without spines and would

correspond to the spineless form found in Europe (‘Podocoryne inermis’ of Allman).

Family Pandeidae

Leuckartiara octona (Fleming, 1823)

Leuckartiara octona: Rees, 1938: 12, figs. 3-5. Millard, 1957: 182.

Records. South coast: LIZ 3A. SCD 2oJ, 26D, 94B, 258T, 281F, 314C,

333G.

Remarks. To avoid misidentification, only those specimens with medusa

buds are included in the above records. SCD 281F was growing on the shell

of Nassa speciosa A. Adams, SCD 314C on Nassa analogica Sow. and all others on

Bullia annulata (Lam.).

Family Aequoreidae

Aequorea africana n. sp.

Fig. 8

Holotype: MB 70G from Mossel Bay on the south coast. South African

Museum catalogue number: SAMH 413.

Description. A colony with well-extended hydranths growing on an empty

snail-shell.

Stem unbranched or branching sympodially up to three times, reaching a

maximum height of 0-2 cm. Stem and branches annulated or corrugated,

increasing in diameter from base to distal end, bearing terminal hydrothecae.

Shorter stems closely and distinctly annulated throughout, longer ones closely

annulated in basal region, irregularly corrugated in more distal part.

Hydrotheca thin and membranous except for base which is somewhat

stouter and remains as a saucer-shaped Halecium-like structure in damaged or

regenerated specimens (fig. 8A), with distal region irregularly creased and

folded longitudinally to form an operculum below which the hydranth can be

462 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 8. Aequorea africana n. sp.

A. An empty stem surmounted by a saucer-shaped structure (the remains of the hydrotheca).

B-E. Hydrothecae containing hydranths in various stages of expansion, E with a gonotheca.

F and G. Gonothecae containing medusa buds, F arising from hydrorhiza, G from the stem.

H. An expanded hydranth showing the web between the tentacle-bases.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 463

withdrawn. Hydranth long and extensile, with conical hypostome and 11-17

tentacles united by a web at their bases (fig. 8H). Tentacles often moniliform

in appearance.

Gonophores borne on stem or hydrorhiza, each on an annulated

pedicel which increases in diameter towards the distal end. Gonotheca

not sharply demarcated from pedicel, pear-shaped, containing one young

medusa-bud.

Measurements (mm.)

Stem, length : ; : . 0°18—2°05

maximum diameter d : . 0°045—-0:07

Hydrotheca, length : - 0°32-0°55

maximum diameter : : ; ; : : Obl. 0-10

Hydranth, length to hypostome (preserved) » )) reaching \ 0-72

Gonotheca, length, with pedicel : : é : . 0°29-0:40

maximum diameter ‘ : : : . Or12-O'15

Remarks. This species is closely related to several members of the ‘Campanu-

linid’ group of hydroids, in particular to the following:

(1) Campomma hincksi (Hartlaub, 1897). This species was linked by

Hartlaub with the medusa Eucheilota maculata WHartlaub, 1894,

probably erroneously according to Rees (1939, p. 442) and Russell

(1953, P- 313). Campomma hincksi is the sole representative of its

genus, which is retained provisionally only until such time as the

medusa can be definitely placed. Both C. hinckst and E. maculata are

known only from Europe. Aequorea africana strongly resembles Leloup’s

material of C. hincks: from Belgium (1952, fig. 70).

Campanulina paracuminata Rees, 1938. This is almost certainly the

hydroid of Aequorea aequorea (Forskal, 1775) (= Aequorea forskalea

Péron and Lesueur, 1809; see Russell, 1953, p. 350, and Huvé, 1952,

p. 36). The medusa is known from Atlantic coasts including South

Africa (Kramp, 1961).

(iii) Campanulina acuminata (Alder, 1857). This is probably the hydroid

of Aequorea vitrina Gosse, 1853 (see Russell, 1953, p. 354) or of Aequorea

pensilis (Haeckel, 1879) (see Huvé, 1952, p. 37). A. vitrina occurs

in N.W. Europe and A. fensilis in the English Channel and

in the Pacific and Indian Oceans including South Africa (Kramp,

1961).

The present material may well be Campanulina paracuminata, whose medusa

is known from South Africa and material of which was examined in the British

Museum in 1960; but as there are slight structural differences (e.g. in the nature

of the operculum and the number of tentacles) and as there are other South

African species of Aequorea with unknown hydroids (e.g. A. coerulescens (Brandt,

1838) and A. macrodactyla (Brandt, 1838)), a separate species has been

established as a temporary measure.

464 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Lovenellidae

Lovenella chiquitita Millard, 1957

Lovenella chiquitita Millard, 1957: 198, fig. 7. Millard, 1959): 250, fig. 3.

Records. West coast: A 384D. SWD 39G. TRA 156C.

Remarks. The placing of this species in the genus Lovenella is a temporary

measure only and is based on the morphology of the hydroid generation,

which shows close affinity to Lovenella clausa (Lovén, 1836).

The medusa, in its absence of cirri at the time of liberation, is more

closely related to Phialella. ‘That it is not P. quadrata (Forbes, 1848) was estab-

lished by comparison with material of the hydroid generation of the latter in

the British Museum in 1960, for which privilege the author is indebted to

Dr. W. J. Rees. It might well be P. falklandica Browne, 1902, the medusa of

which has been found in South Africa at Saldanha Bay (identification by

Dr. M. E. Thiel, Zoologisches Museum, Hamburg), yet the youngest known

medusae of this species (o-8—1-0 mm. diameter) have ‘four perradial tentacles,

and the four interradial tentacles are just beginning to develop and are visible

as four minute bulbs’ (Brown and Kramp, 1939: 298), suggesting that they are

liberated with only 4 tentacles and not 8 as in the present species.

Family Haleciidae

Halecium beaniit (Johnston, 1838)

Fig. 9 A-F

Halecium beanii: Hincks, 1868: 224, pl. 43 (fig. 2). Broch, 1918: 38, fig. 13. Millard, 1957: 188.

Millard, 1958: 168. Ralph, 1958: 332, fig. 10 a, b, e-k. Vervoort, 1959: 224, fig. 6.

Records. West coast: CP 336C. SB 196M. SWD 12D, 42F. TB 17A, 21B.

WCD 125U, 145V. South coast: CPR 46L. LIZ 7S. MB 47T, 60Q. SAMH 157,

214, 273, 328, 335, 341. SCD 37M, 85M, 154C, 387G, 394B.

Description. Stiff, shrubby colonies, many of them epizootic on other

hydroids, the largest reaching a height of 7-4 cm.

Remarks. Since no criterion can be found for distinguishing between

sterile colonies of H. beanit and H. halecinum, only samples containing female

gonophores have been included above. The species is certainly more common

than is indicated by the records, as a further 26 sterile samples are present in

the collection, all, or most of which, probably belong to the same species.

Attempts have been made by various authors to distinguish H. beanu from

H. halecinum on the basis of the angle of the hydrothecal margin and the shape

of the basal part of the hydrophore, but, as the accompanying diagrams (fig.

g A-E, all chosen from fertile specimens) will show, both characters are so

variable that no reliability can be placed upon them.

Halecium delicatulum Coughtrey, 1876

Fig. 10L

Halecium parvulum: Millard, 1957: 189, fig. 4A. Vervoort, 1959: 227, fig. 7.

Halecium parvulum, var. magnum Millard, 1957: 190, fig. 4 B—O.

Halecium delicatulum: Ralph, 1958: 334 (synonymy), figs. 11e, A—n, 12 a—p.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA

465

Fic. 9. Halecium beanii (Johnston) (A-F), and H. halecinum (Linn.) (G-L).

A-E and G-K, hydrophores from various colonies.

F and L, gonothecae.

Records. West coast: AFR 736Z. CP 327C. LAM g30P, 35A. TBg, 17C,

21C. WCD 18T, 156A, 160P. South coast: LIZ 16F. MB 19Q, 84E, 88J.

SAMH 179, 189, 242, 250, 381. SCD 5F, 22A, 29H, 37N, 52J, 85K, 154B,

179H. TRA 92M.

466 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks. It has been pointed out by Ralph (1958) that the large variety

(var. magnum Millard) described in 1957 has similar dimensions to the type

material of H. flexile Allman, 1888, which is now recognized as a synonym for

H. parvulum and H. delicatulum. Since so great a variation of size is possible

it is not justifiable to retain a subspecies on this character alone.

Halecium dichotomum Allman, 1888.

Fig. 10 A-K

Halecium dichotomum Allman, 1888: 13, pl. 6. Billard, 1910: 4. Stechow, 1925): 419. Millard,

1957: 188 (excluding the male gonophores).

Records. West coast: WCD 20G, 164C. South coast: LIZ 7W. MB 8V,

12W. SAMH 162, 227, 243, 274, 352. SCD 5G, 20J, 37P, 81P, 85N, 154D,

1751, 1885, 239F, 265G, 333C, 387F, 394A. TRA 38], 92N.

Description. Colonies very variable in appearance and growth-form. Stem

fascicled, branching with the typical dichotomy described by Allman and

generally in all planes, usually geniculate, with a tendency for annulation in

the region of the nodes. Primary hydrophores usually sessile, secondary hydro-

phores usually symmetrical and annulated in the basal region. Hydrotheca

low and wide, with walls flared outwards, over 0-17 mm. in diameter at the

margin.

Among the variety of colonies examined two extremes of growth-form

occur:

(i) Large upright colonies reaching 11-12 cm. in height,. with strongly

fascicled stems and branching mainly in one plane. Main stem more

or less straight and dichotomy not obvious. Larger stems and branches

stiff in appearance and up to 2 mm. thick, though unable to support

themselves out of fluid. Smaller branches graceful and flexuous.

Stem and branches with long internodes with no annulation other

than a shallow constriction near the base. Secondary hydrophores

scarce. Closer examination shows that the typical dichotomy is in

fact present in this form, but that one limb is always short (the branch)

and the other long, contributing to the axis of the stem. The latter

limb is enveloped by peripheral tubes in fascicled regions, so obscuring

the dichotomy (fig. 10 A, B).

(ii) Low, scrubby colonies, often epizootic, reaching a height of 1-2 cm.

Stem usually weakly fascicled and strongly geniculate. Branching

profuse and in all planes, and stolonisation common, resulting in a

tangled mat which may cover large areas of the substratum and is

very easily recognised. The typical dichotomy is common in this form

and very obvious (fig. 10 C). The internodes of the stem and branches

tend to be shorter and more annulated, and secondary hydrophores

are abundant and often closely annulated in their basal regions.

In two specimens (MB 12W and WCD 164(C) of the low, scrubby form,

the colony is epizootic on a dead polyzoan and the hydrorhiza is provided with

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 407

2 a

Fic. 10. Halecium dichotomum Allman (A-K) and H. delicatulum Coughtrey (L). ©

AandB. Portions of stem from the distal and proximal ends respectively of the tall form, to

show the incorporation of one limb of the dichotomy into a main stem. Peripheral tubes

teased apart in B. (TRA g2N.)

Cand D. Portions of stem from TRA 38J, showing typical dichotomy in C and a unilateral

branch in D.

EandF. Female and male gonophores.

G-J. Details of secondary hydrophores.

K. Part of the hydrorhiza to show root-like structures (MB 12W).

L. Part of a branching stem in H. delicatulum for comparison.

468 ANNALS OF THE SOUTH AFRICAN MUSEUM

unusual rootlike projections which enter the pores of the host and anchor the

colony (fig. 10 K). Similar ‘roots’ have been observed on other mounted

specimens.

Female gonophores as described by Allman. Male gonophores on separate

colonies, slender, elongated and often curved, tapering distally to a small

rounded aperture, closely annulated throughout or smooth in distal region

(fig. 10 F). Reaching 1-24 mm. in height and 0-36 mm. in maximum diameter.

Remarks. The variation in the growth-form of this species is remarkable,

and at first glance the two extremes might be taken for different species.

However, female gonophores have been found in both, and intermediate

types exist. Comparison with Allman’s type material in the British Museum

(slide no. 88.11.13.9) confirms the identification, and shows that the hydro-

phores are similarly variable. Primary hydrophores, though usually sessile,

may be quite long. The hydrotheca may be strongly flared out, weakly flared

at the extreme distal edge only (the commonest type), or the everted part

may be quite worn off.

This variation may introduce difficulties in the identification of sterile

colonies and possible confusion with species such as H. delicatulum, though to one

familiar with the species no difficulty arises. Useful diagnostic characters are

the normally sessile primary hydrophores, the typical dichotomy in which 2 or

even 3 internodes arise from a previous one at the same level, and the character-

istic curved apophysis by which each internode arises from its predecessor.

Dichotomy may also occur in H. delicatulum, but the two limbs usually arise

at different levels and the apophyses are not curved (fig. 10 L). H. dichotomum

may be distinguished from H. tenellum by its fascicled stem and larger

hydrothecae.

With the abundant material available, it was possible to clear up the

confusion which previously existed over the nature of the male gonophores

of this species. A re-examination of the material described from False Bay

(Millard, 1957: 188) showed that the smooth gonophores from sample FAL 78Z

belonged in fact to an epizootic male colony of H. beanii inextricably tangled

with a female colony of H. dichotomum. The annulated gonophores without

lateral openings from sample FAL 64N and thought to be female, were in

fact male.

Halecium halecinum (Linn., 1758)

Fig. 9 G-L

Halecium halecinum: Hincks, 1868: 221, pl. 42. Broch, 1918: 36, fig. 11. Vervoort, 1946): 158,

figs. 63-64. Vervoort, 1959: 225.

Records. South coast: SCD 85P.

Description. A small epizootic, yet fertile, colony, reaching a maximum

height of 0-6 cm. Stems stunted and branching irregularly, but fascicled at base.

Internode length about twice width at distal end.

Secondary hydrophores very variable, most are asymmetrical, curving

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 469

towards abcauline side and with an obliquely set aperture. Some are quite

symmetrical in the basal region, though asymmetrical more distally. In only

a few is the aperture perpendicular to the axis.

Female gonothecae banana-shaped with concave adcauline side and

terminal aperture. Containing a single row of 3-4 large eggs.

Remarks. This species is possibly more abundant than is indicated by this

single record, as without gonophores it is impossible to distinguish from

H. beanii. This is the only fertile colony observed and the first fertile, and

therefore unquestionable, record from South Africa.

Halecium ?muricatum (Ell. & Sol., 1786)

Fig. 11 A, B

Halecium muricatum: Hincks, 1868: 223, pl. 13 (fig. 1). Broch, 1918: 43, fig. 17. Vervoort, 1946):

163, fig. 67.

Records. West coast: TB 19A. South coast: SCD 56U.

Description. Fascicled stems reaching a maximum height of 4:7 cm.,

branching in an irregularly pinnate fashion, straight for most of length though

sometimes weakly geniculate in distal regions, fairly rigid and able to support

themselves out of fluid, though more graceful in appearance than in H. beanii.

Nodes oblique and sloping alternately to left and right.

Primary hydrophore long; borne on broad apophysis at distal end of

each internode, from which it is separated by a distinct groove; usually asym-

metrical with adcauline side more convex than abcauline; with a well-marked

pseudodiaphragm in distal region below hydrotheca and usually a second one

in proximal region. Pseudodiaphragm better developed on adcauline side.

Secondary hydrophores similar, though not so long and usually with only one

pseudodiaphragm. Hydrotheca relatively deep, margin everted, more so on

adcauline side.

Gonophores absent.

Measurements (mm.)

TBi9gA SCD 56U

Stem, internode length . ; i : 8 . 0:70-0:86 0-61-0-77

diameter across node : : : . O°19-0'24 0:12-0:28

Hydrotheca, diameter at margin. : : . 0°24-0:31 0:24-0:29

depth (diaphragm to margin) . : : . 0:09-0:14. 0:08-0-12

Remarks. The identification of this species must remain dubious until the

gonophores have been discovered. The stem and hydrophores are remarkably

similar to those illustrated by Broch (1918) for H. muricatum, yet H. muricatum

is an arctic species and has not been reported from the Southern Hemisphere.

Another closely related species is H. filicula Allman, 1877, from the Gulf

Stream and N. Atlantic Ocean. This species has smaller dimensions (cf.

Billard, 1906: 163) and differs in the presence of 2 annulations at the base of

the hydrophore and in the absence of a pseudodiaphragm. The gonothecae

470 ANNALS OF THE SOUTH AFRIGAN MUSEUM

(Billard, 1906, fig. 2) appear to be similar to those of H. muricatum, and the

two species may prove to be conspecific.

Fic. 11. Halecium ?muricatum (Ell. & Sol.) (A-B) and H. tenellum Hincks (C-F).

Aand B. Portions of stem from TB 19A and SCD 56U respectively.

Cand D. Female gonothecae in side and front views respectively, D with 2 larvae (SCD 37Q).

E. Part of colony and female gonotheca with several larvae (SCD 60B).

F. Male gonotheca from SAMH 315.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 471

Halecium tenellum Hincks, 1861

Fig. 11 C-F

Halecium tenellum: Millard, 1957: 193, fig. 5. Hamond, 1957: 307, fig. 14. Ralph, 1958: 340.

Vervoort, 1959: 229, fig. 8. Naumov, 1960: 454, fig. 344.

Records. South coast: MB 8X. SAMH 315. SCD 37Q, 60B, 394C.

Description. A number of colonies, mostly epizootic, with unfascicled

stems reaching a maximum height of 0-4 cm. Stems geniculate and often

branching in a dichotomous manner, usually with many athecate internodes

as previously figured (Millard, 1957), but often of more normal appearance

as figured by Vervoort (1959) and Hamond (1957).

Male and female gonophores present, on separate colonies. Male

gonothecae compressed, broad and bluntly rounded at distal end when young,

pointed at distal end when mature, reaching a maximum of 1:07 mm. in

length and o-60 mm. in diameter. Female gonothecae borne on stem or

hydrorhiza, compressed, pear-shaped in anterior view with bluntly rounded

distal end, and a circular, terminal aperture blocked by a small papilla which

is shed on the escape of the contents, reaching a maximum of 1:07 mm. in

length, 0-51 mm. in diameter and 0:30 mm. in thickness; with no hydranths;

containing 2-7 eggs on a branching blastostyle, which develop into planulae

in situ.

Remarks. Vervoort was correct in assuming that the male gonophores

previously described (Millard, 1957) were young ones, as larger mature ones

in the process of shedding their contents have now come to light.

Family Campanulariidae

Campanularia hincksu Alder, 1856

Fig. 12 A-D

Campanularia Hincksii Alder, 1856: 360, pl. 13 (fig. 9). Hincks, 1868: 162, pl. 24 (fig. 3), fig. 18

Billard, 1906: 172, figs. 4—5 (incl. var. grandis).

Campanularia hincksii: Vervoort, 1959: 311, fig. 55a.

Records. South coast: SAMH 283. SCD 354G.

Description. Pedicels unbranched and smooth, except for occasional

regeneration lines, with a single spherule at distal end.

Hydrotheca inverted cone-shaped, with 8-12 broad, marginal teeth in

which the distal end is either hollowed out to form two points or (more rarely)

truncated. Hydrotheca polygonal in end-on view, the angles between the

teeth forming longitudinal striations visible in side view, which may continue

to the base or peter out half-way down. Diaphragm in form of annular thecal

thickening.

Female gonotheca with very short stalk, elongated, broad near base and

narrowing slightly to truncated distal end, with about 8 low, rounded annula-

tions. Only 2 gonothecae seen, one empty and one almost spent yet still

containing 2 eggs.

472 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm.) SAMH 283 SCD 354G

Pedicel length : : : : : : . 2:26-4:19 3°70-6°81

maximum diameter : : ; ; . 0°08.0°13 O°12—0°15

Hydrotheca, length : : : : : . 0°85-1°24 1°20—-1°55

maximum diameter ; : ; : . 0°47-0°77 0:60—0-88

length/diameter : 3 : : : . 1:36-1:97 1:48-2:27

Gonotheca, length . : ; ; ; : : 1°52

maximum diameter : : : 0°64

Remarks. This is the first ae of a species from South Africa. The

measurements correspond well with Billard’s var. grandis, but the material

resembles the typical form more in the double nature of the marginal teeth.

These teeth appear to become truncated as they wear down with age.

Campanularia integra MacGillivray, 1842

Fig. 13 A-D

Campanularia caliculata: Warren, 1908: 338, fig. 19.

Campanularia integra: Billard, 1907: 340. Jaderholm, 1917: 4. Broch, 1918: 159 (synonymy).

Millard, 1957: 193. Millard, 1958: 171.

Clytia compressa: Vanhoffen, 1910: 303, fig. 24.

?Campanularia gracilis: Stechow, 1925): 423, fig. 6.

Orthopyxis caliculata: Ralph, 1957: 838, fig. 6 a—.

Records. South coast: CPR gE. LIZ 7U, 13C. MB52L, 55J. SCD 84X,

179E.

Description. Colonies creeping on weeds and other hydroids, particularly

Lytocarpus filamentosus.

Remarks. In the identification of this species I have followed Broch (1918)

who considers C. integra, C. caliculata and C. compressa as synonymous, although

this opinion is not accepted by all recent authors. The difference between

these species is supposed to lie in the gonotheca, which is said to be spirally

annulated in C. integra, smooth and round in section in C. caliculata, and smooth

and compressed in C. compressa. Broch claims to have found intergrading forms

between C. integra and C. caliculata, while the South African material shows

intergrading forms between C. caliculata and C. compressa. Here the gonotheca

is usually compressed and smooth, but may also be round in section, and is

sometimes roughly corrugated, though not distinctly annulated.

Medusa systematists distinguish the ‘medusa’ of C. caliculata (Agastra mira

Hartlaub, 1897) from that of C. compressa (Agastra rubra Behner, 1914) on

minor points such as the arrangement of eggs. Of the two, the South African

material resembles more Agastra rubra, as the eggs are large and comparatively

few in number.

Campanularia laminacarpa n.sp.

Fig. 12 E-K

?Campanularia tincta: Jaderholm, 1923: 6

?Campanularia africana: Stechow, 19256: 420, 421 (material from Agulhas Bank, station 105).

Non Campanularia tincta: Warren, 1908: 337, fig. 18.

Non Campanularia africana Stechow, 19235: 104.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 473

(Lab e Aererrerey | fonvnnle LS.

Fic. 12. Campanularia hincksti Alder (A—D) and C. laminacarpa n. sp. (E-K).

A-C, hydrothecae, and D, gonotheca, from SAMH 283.

E-G, hydrothecae from TRA 38P, and K, from SCD 84U.

H. Male gonophore from SCD 84U.

J. Female gonophore from the holotype, TRA 32A.

474 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype: TRA 32A, a female colony from the Agulhas Bank growing on

Thyroscyphus aequalis Warren. South African Museum registered number:

SAMH 414.

Other records: South coast: SAMH 174, 269, 351. SCD 37V, 61F, 79M,

84U, 112A, 117L, 154E, 169Y, 265E, 394F. TRA 35Z, 38P, 56U, 92B.

Description. Colony stolonic and epizootic on the Sertulariid Thyroscyphus

aequalis. Hydrothecal pedicel upright, unbranched, smooth or corrugated, with

a spherule of smaller diameter at the distal end.

Hydrotheca tubular or slightly expanding to margin, smooth, with length

about twice diameter at margin, with a diaphragm in the form of a well-

developed annular thickening of the perisarc, with 10-14 rounded marginal

teeth.

Gonotheca erect, flat, smooth, generally held in a plane at right angles

to the stem of the host, broadening to distal end which is abruptly truncated,

with a wide, operculate aperture. Male and female similar. Female containing

a single sporosac bearing numerous small eggs. Male with a single sporosac

with the structure of a degenerate medusa, spermatogenic cells arranged in

4 longitudinal bands with indications of 4 radial canals.

Measurements (mm.)

Hydrothecal pedicel, length . : , ; : 2 . O-3I-1°34

maximum diameter : : . : : : . 0:06-0:11

Hydrotheca, height ‘ , : : : : ' _. 0°36—-0°71

diameter at margin . : : ‘ ‘ 5 : .0'225-0°34

diameter/height ; : ; : : : . 0°46—-0°74

Gonotheca, height . : ; : : . 1°53-2°01

maximum diameter : . : : : : . o-81—1-20

Remarks. This species has previously been confused with Campanularia

africana Stechow, 1923b. The discovery of gonothecae, which are larger and

quite different in appearance from those of the latter species, necessitates the

establishment of a new species. Unfortunately the two species have identical

trophosomes and there is no method of distinguishing sterile material.

Campanularia africana was originally described from Park Rynie, Natal, as

C. tincta by Warren in 1908 (p. 337, fig. 18). It has cylindrical to ovate gonothe-

cae, narrowing towards the distal end and then everted to form a short, circular

collar around the aperture. Although there are variations in shape the

gonothecae are never flattened, but always round or nearly so in section. I

have examined Warren’s material, which was kindly loaned to me by the

Director of the Natal Museum, and found his diagrams to be a faithful represen-

tation of the structure. The gonothecae contain a number of large planulae.

It was to this material that Stechow (1923b) gave the new name of Campanularia

africana. The species presumably also includes Pennycuik’s material from

Australia (1959: 169), which possessed male gonothecae, one with ‘5 shallow

annulations’.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 475

Fic. 13. Campanularia integra MacGill. (A-D) and C. ?mollis (Stechow) (E-J).

A and B, Gonothecae, and C and D, hydrothecae from CP 258.

E-J, various hydrothecae from CP 646C showing variation in thickness of walls.

476 ANNALS OF THE SOUTH AFRICAN MUSEUM

Stechow (1925) stated that he had examined material loaned to him by

Warren. This was obviously not the type material, and was not recorded by

Warren, for it came from Algoa Bay, and it was growing on Thyroscyphus

regularis (= T. aequalis) and presumably unfertile. This material was probably

C. laminacarpa, as was Stechow’s Valdivia material (unfertile) reported in the

same paper, at any rate that growing on T. aequalis. C. laminacarpa possibly

only occurs on 7. aequalis as I could only identify gonothecae in material

growing on this host, and I have only quoted records (above) of material

growing on Thyroscyphus. Of these, 7 have gonothecae of the flattened type.

Several other sterile colonies present in this collection, and growing on

other species of hydroids, cannot be identified with certainty, nor can unfertile

material in the literature.

Campanularia ?mollis (Stechow, 1919)

Fig. 13 E-J

Clytia mollis Stechow, 1919: 44, fig. L.

Campanularia mollis: Picard, 1951: 344, fig. 3.

Orthopyxis mollis: Ralph, 1957: 840, fig. 7e-k.

Records. West coast: CP 646C.

Description. A flourishing colony growing on weed. Pedicel generally

smooth, with a single small spherule at distal end, often with regeneration nodes.

Hydrotheca cylindrical, with length 2-3 times height, with 7-9 clearly defined

marginal teeth separated by broad and shallow bays, diaphragm in form of

annular thecal thickening. Marginal teeth sometimes bifurcated at the tip.

Margin often reduplicated.

Perisarc of pedicels very thick, that of hydrothecae variable, sometimes

thick throughout, sometimes thin for the most part though always thickened

at margin and near base.

Gonothecae absent.

Measurements (mm., without reduplications).

Pedicel, length : , : . 0°58-1°87

maximum diameter : ! : : . O*10—-0°17

Hydrotheca, length , : ; : . 0°57—-0:80

diameter at margin . : 4 , : ; : . 0°26—0°33

length/diameter ; : : : , : : . 1°94—2°93

Remarks. The identification of this species must remain doubtful in the

absence of gonothecae, as there are several closely related species with similar

trophosomes. It is close to C. mollis though differing from previous descriptions

of the species in the larger hydrothecae, better defined marginal teeth and

smooth pedicels. A few specimens do, however, show indications of a spiral

twisting at the base of the pedicel. The species has not been recorded from

South Africa before.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 477

Campanularia morganst Millard, 1957

Campanularia morgansi Mallard, 1957: 195, fig. 6. Millard, 1958: 171.

Records. South coast: LAM 30M. MB 47Y. SAMH 202, 221, 230, 2096.

SCD 37W, 84W, 117K, 354F. West coast: WCD 12H.

Genus Clytia Lamouroux, 1812

Remarks on type species. The genus Clytia was established by Lamouroux in

1812 (p. 184) for 3 species, namely:

Sertularia volubilis

Sertularia syringa Linn., 1767

Sertularia verticillata Linn., 1758

Of these, S. syringa has been transferred to Calicella Hincks, 1859, and

S. verticillata is generally considered as a species of Campanularia but has recently

been declared the type species of a new genus Verticillina by Naumov (1960).

In the literature 2 species have been confused under the specific name of

volubilis :

(i) The name Sertularia volubilis was originally established by Linnaeus

in 1758 (p. 811) for Ellis’s ‘small climbing Coralline with bell-shaped

cups’ described and figured by the latter in 1755 (p. 24, pl. 14, fig.

a, A). But Ellis’s description was very inadequate and the material

might be either Campanularia or Clytia. However, the name has been

retained for a well-known species of Campanularia with fixed sporosacs

and a smooth gonotheca which has been declared by Naumov in

1960 to be the type species of this genus (i.e. Campanularia volubilis

(Linn., 1758)).

(ii) In 1786 Ellis and Solander (p. 51, pl. 4, fig. e, f, E, F) described

and figured under the name of Sertularia volubilis material with an

annulated gonotheca which is unmistakably recognisable as the

species later described by Alder in 1856 as Campanularia johnstont.

Ellis and Solander added the corollary that ‘there are different

varieties and sizes of this twining bell-shaped Coralline’. It is to this

description that Lamouroux referred when he created the genus

Clytia.

So far as I can determine no type species has been established for the genus

Clytia and I therefore select Sertularia volubilis Ellis & Solander, 1786, non

Sertularia volubilis Linn., 1758.

However, the specific name volubilis is obviously untenable. The name

Jjohnstoni Alder, 1856, is antedated by the name of the medusa of the same species,

namely Medusa hemisphaerica bestowed by Linnaeus in 1767 on the ‘Medusa

hemisphaerica’ of Gronovius, 1760 (the latter name being part of a Latin descrip-

tion and thus not valid). The same medusa was later placed in the genus

Phialidium Leuckart, 1856, by which name it is now generally known, but which

is antedated by Clytia Lamouroux, 1812.

4.78 ANNALS OF THE SOUTH AFRICAN MUSEUM

To sum up, the name of the type species of the genus Clytia becomes

Clytia hemisphaerica (Linn., 1767).

Synonyms: Medusa hemisphaerica Linn., 1767

Phialidium hemisphaericum (Linn., 1767)

Sertularia volubilis Ellis & Solander, 1786, non Linnaeus, 1758.

Campanularia johnstoni Alder, 1856

Clytia hemisphaerica (Linn., 1767)

Fig. 14 A-F

Laomedea gracilis: M. Sars, 1857: 160, pl. 2 (figs. 1-3, 5).

Campanularia johnstoni Alder, 1856: 359, pl. 13 (fig. 8). Vervoort, 1959: 312.

Clytia raridentata: Vanhoften, 1910: 301, fig. 22. Fraser, 1944: 145, pl. 26 (fig 118). Millard,

1957: 197-

Campanularia raridentata: Stechow, 1919: 58, fig. Q.

Thaumantias raridentata: Stechow, 19234: 107, fig. M. Stechow, 1925): 426.

Clytia gracilis: Stechow, 19256: 431, figs. 9-10. Millard, 1957: 196. Millard, 1958: 172, fig. 3B,

E, G.

Fea Ralph, 1957: 820, 823, figs. 1h-u, 2, 3a—f. Millard, 1958: 172, fig. 3A, D, F.

Laomedea (Phialidium) pelagica: Vervoort, 1959: 313, fig. 558, ¢.

Records. West coast: SB 178C. TB 7, 16. South coast: LIZ 7T, 11K, 40K.

MB 8N, 52K, 60N, 64P, 69B, 81W. SAMH 161, 166, 176, 180, 203, 239, 253,

383. SCD 5L, 37X, 50N, 52U, 56V, 61G, 75H, 79P, 84V, 129E, 179D, 265],

283T, 284E, 330C, 387L, 394G. STJ 31M. TRA 57D, 92W, 150B.

Description. Numerous colonies growing on weeds and other hydroids.

Stems solitary or occasionally sparsely branched, annulated at top and bottom

and occasionally throughout.

Hydrotheca variable in size, with 8-15 marginal teeth. Teeth acute,

sharp or rounded, but always covering a smaller area than the bays between

them, often asymmetrical and leaning towards one side. Diaphragm distinct,

variable in thickness, but always clearly demarcated from the hydrothecal

wall.

Gonotheca generally smooth, with truncated distal end, but sometimes

with 1 or 2 irregular corrugations; none of these specimens with distinct

annulations.

Remarks. Ralph has demonstrated for Clytia johnstoni in New Zealand a

variation in size, shape of marginal teeth and degree of annulation on the

gonotheca which can be correlated with latitude. While specimens from the

southern region correspond to the classical conception of C. johnstoni, with

annulated gonothecae and broad marginal teeth, those from the northern

region have smooth gonothecae and acute marginal teeth and thus cover the

form known as C. gracilis. We thus have no alternative but to combine the two

species. So far as is known there is no difference between the medusae. The

range of variation also covers material from South Africa recorded as

C. raridentata, which is here considered a synonym. See also remarks on p. 477.

Since the latitude on the south coast of Africa is the same as that of the

north end of New Zealand, it might be expected that the material in these

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 479

Fic. 14. Clytia hemisphaerica (Linn.) (A-F), and C. hummelincki (Leloup) (G-L).

A-D. Various hydrothecae showing variation in total size and shape of marginal teeth: A

with triangular teeth, B and D with asymmetrical teeth and C with very slender teeth.

E and F. Gonothecae, corrugated type (a smooth one shown in A).

G. A hydrotheca containing a hydranth, and a gonotheca.

Hand J. Empty hydrothecae.

K and L. Gonothecae containing young medusae.

(A from TB 16, B from TRA 57D, C from Natal, D-F from False Bay, G-L from SCD 328H.)

480 ANNALS OF THE SOUTH AFRICAN MUSEUM

two regions would be similar, and that the variation in South Africa would

continue from where it left off in New Zealand as one moves up the east coast

into warmer waters. This was tested out in the material available and

measurements of hydrothecal size and pedicel length are given below.

Hydrotheca: height (mm.) Pedicel : length (mm.)

Range Mean Range Mean

West coast 0-6-1°0 0:8 0-7-3°4 2°4.

(32-34° S) (16) (16)

South coast 0°4—-0'9 0-6 0°5-5°7 1°9

(34-35. S) (40) (39)

East coast 0:4-0°8 0-6 1-6—3°8 2°5

(32-28° S) (32) (20)

Portuguese

East Africa 0:3-0°6 0°5 0-4-1°6 1:0

(27-23° S) (23) (18)

These figures show that the hydrothecal height from the south coast

corresponds well with that from a similar latitude in New Zealand (Ralph

gives measurements of 0:40-0:81 mm. between 40° S and 34° S) and that there

is a clear decrease in size from the west coast (where the water is colder than

the south coast) round the south and up the east coast. A similar trend in

pedicel length is not so clear, although pedicels from Portuguese East Africa

are certainly shorter than elsewhere. No tendency for a change in the proportions

of the hydrotheca was seen.

The marginal teeth are always acute in South African material as in

northern New Zealand, and on the east coast there is a tendency for the bays

between them to become larger and the teeth narrower (fig. 14).

With one exception the gonothecae are quite smooth or with a few irregular

corrugations, as in those illustrated by Ralph from northern New Zealand.

This is as might be expected, and it is difficult to account for the single sample

where all gonothecae are distinctly annulated (6-8 rings) described from

Portuguese East Africa as C. johnstoni (Millard, 1958, fig. 3D).

The variable nature of the gonotheca explains the discrepancy which

exists in the literature over the nature of the gonotheca of Clytia (Thaumantias)

raridentata (cp. Fraser, 1944, and Stechow, 19232).

The nature of the diaphragm allows for no differentiation between species,

and, although variable in thickness, it is always distinct from the hydrothecal

wall and thus different from the type characteristic of the genus Campanulania.

Clytia hummelincki (Leloup, 1935)

Fig. 14 G-L

Laomedea hummelincki Leloup, 1935: 19, fig. 7.

Records. South coast: SCD 328H.

Description. A rich colony growing on the surface of Lepas sp. taken from a

buoy. Colony stolonic, stem unbranched, long, closely annulated in basal

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 481

region and often for short areas at other irregular intervals, otherwise smooth,

with a single, rather flattened ‘spherule’ at distal end.

Hydrotheca in the shape of an inverted cone and usually with straight

sides, expanding evenly to margin, with height approximately equal to

maximum diameter. Margin untoothed. Diaphragm very delicate, usually

oblique. Hydranth with 15-29 tentacles, completely retractable into

hydrotheca.

Gonotheca arising separately from hydrorhiza on a short pedicel of 2-4

segments, elongated and expanding to distal end which is truncated, containing

I or 2 medusa buds. Oldest medusa deep, with manubrium, 4 radial canals

and 4 tentacle bulbs visible.

Measurements (mm.)

Pedicel, length : A A : 4 ; ; : el G2 > Aas

maximum diameter : ; : : a . 0709-015

Hydrotheca, length : : : ‘ : ; . 0°25-0°38

diameter at margin . 4 : : ; , ‘ . 0°22-0°42

length/diameter : : : : . 0°70-1°36

Gonotheca, length . ; : ; . 0°73-1:26

maximum diameter ; : : : : ; . 0°28-0°45

Remarks. This material agrees entirely with Leloup’s description of

Laomedea hummelincki from the West Indies, except that these pedicels are

somewhat longer and stouter. The nature of the diaphragm and the presence

of medusa buds in the gonotheca (observed here for the first time) shows the

species to be a Clytza. This is only the second record of this rare species, and a

new record for South Africa.

Clytia paulensis (Vanhoffen, 1910)

Fig. 15

Campanularia paulensis Vanhoffen, 1910: 298, fig. 19 a, b.

Clytia paulensis: Stechow, 1919: 45, 155. Stechow, 1923a: 110, fig. N. Stechow, 1925): 428,

fig. 7. Stechow, 1925a: 211.

?Clytia ulvae Stechow, 1919: 47, fig. N. Stechow, 1925): 428.

Records. South coast: LIZ 11M. SAMH 336. SCD 70N, 154G, 258W,

276U, 333D, 387M. TRA 38K.

Description. Several colonies growing on the stems of other hydroids. Stem

unbranched, or giving rise to 1 or 2 secondary pedicels in a sympodial manner,

closely annulated at base, in distal region and at irregular intervals between.

Hydrotheca very thin and fragile, with 7-10 double marginal teeth.

Margin bowed out between teeth giving an undulating outline in end-on view

and sometimes the effect of longitudinal striations in the upper part of the

hydrotheca. Depth of hydrotheca 13 to 3} times diameter at margin.

Gonotheca as described by Stechow: smooth and fragile, with annulated

pedicel and truncated distal end, containing 1-3 medusa buds.

482 ANNALS OF THE SOUTH AFRICAN MUSEUM

el Q

Fic. 15. Clytia paulensis (Vanhoffen).

A-E and J. Hydrothecae.

F. Gonotheca.

Gand H. MHydrothecae drawn from Stechow’s slide of Clytia ulvae from Marseilles, 1910.

Measurements (mm.)

SAMH TRA SCD SCD SCD —_ Chytia ulvae

336 38K 258W 333D 154G (Marseilles)

Pedicel length : - 0°57-I'14 0°48-1-°82 0:52-0:94 O-QI-I-13 0:85-1:00 0-22—0-90

maximum diameter 0:04-0:06 0:04-0:06 0:04-0:05 0°045-0°05 0°035 0°03—0:035

Hydrotheca, length . 0°35-0°58 0:41-0:72 0:45-0°69 0:53-0:68 0:42-0:50 0°35-0°44

diameter at margin. 0°16-0°33 0:19-0:31 0°15-0:28 0:21-0:25 0°15-0°16 0-12-0-14

length/diameter . 1-76-2:47 1:90-2:68 2:17-3:07 2:20-3:10 2-80-3°13 2°71—3:23

Gonotheca, length : -- 0-715 0:66-1:00 — = =

maximum diameter — 0°385 0-30—0°36 — = ==

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 483

Remarks. Measurements taken from different localities show that the size

and proportions of the hydrotheca are very variable. Vanhdffen’s material

from the Antarctic is well within range, while some of Stechow’s material

from South Africa (19256) and Australia (1925a) is a little smaller, though

the proportions are similar.

It is highly probable that Clytza ulvae Stechow, 1919, is a synonym for

C. paulensis. I have been able to examine a prepared slide of Stechow’s material

of C. ulvae from Marseilles kindly loaned by the Munich Museum and find

that the marginal teeth are in fact double (fig. 15G, H), though the bays

between members of a pair are almost as large as those between pairs. However,

the depth of the former bays is variable within a single colony of C. paulensis

so that the small teeth may be # to 3 the size of the large double teeth. I have

included measurements of Stechow’s material (taken by myself) for comparison.

These show that the hydrothecae are slightly narrower and the proportion

of length/diameter in consequence slightly greater, though the material might

well fit near the end of a series of changing proportions arranged as above.

Obelia dichotoma (Linn., 1758)

Obelia dichotoma: Millard, 1952: 420, 426, 433, fig. 3. Millard, 1957: 198. Millard, 1958: 174.

Millard, 1959): 250.

Obelia dubia: Vanhoffen, 1910: 307, fig. 27. Nutting, 1915: 77, pl. 19 (fig. 1). Stechow, 1925):

435. Fraser, 1937: 86, pl. 17 (fig. 87).

Campanularia obtusidens Jaderholm, 1904: 2, pl. 1 (fig. 1).

Records. West coast: CP 378. LAM 46P. LB 378F. OLF 21C. SAMH 404,

405. TB 8. South coast: KNY 165C (recorded by Day, Millard and Harrison,

1952). LIZ 2G, 11J. MB 37D, 81V. SAMH 147, 160, 223, 338, 342. SCD 112G,

258V, 281D, 312C. TRA 38L, 42W.

Remarks. Obelia dubia is considered to be a synonym for O. dichotoma. The

South African material shows all grades of hydrothecal types from the typical

O. dichotoma form where the marginal teeth are usually not distinct to that of

O. dubia as illustrated by Jaderholm (1904) and Vanh6ffen (1910), where

there are distinct marginal teeth and indications of longitudinal striations. It

is impossible to draw a dividing line between them. The hydrothecal pedicel

is extremely variable in length and the diaphragm may be straight or oblique

within the same colony.

The gonothecae of O. dubia were illustrated by Fraser (1937) and are said

to be ‘almost smooth, or provided with broad, shallow undulations’. In the

South African material the gonothecae are usually smooth, but examples

are also present (with the typical dichotoma hydrothecae) where the gonothecae

are corrugated, approaching very closely the type illustrated by Fraser.

Obelia geniculata (Linn., 1758)

Obelia geniculata: Millard, 1957: 198. Millard, 1959: 250.

Records. West coast: A 383 (reported by Bright, 1938). CP 325. LAM 24H.

LB 127, 314G, 371B. PP 1V. SAMH 357. SB 168G, 235J. TB 6A. TRA 86P.

WCD 81G. South coast: TRA 42V.

484 ANNALS OF THE SOUTH AFRICAN MUSEUM

SUMMARY

A total of 43 species of hydroids is recorded, including 25 Gymnoblasts

and 18 Calyptoblasts. Of these 5 are new species, namely Monocoryne minor,

Myriothela tentaculata, Bicorona elegans, Aequorea africana and Campanularia

laminacarpa, and 8 are new records for the Republic of South Africa.

The new genus Bicorona is established for a Corynid species with 2 whorls

of tentacles and fixed sporosacs.

The status of the Campanularian genus Clyiza is discussed and a type species

selected.

ACKNOWLEDGEMENTS

The Trustees of the South African Museum acknowledge gratefully the

receipt of grants from the University of Cape Town and from the Council for

Scientific and Industrial Research towards the cost of publication. The author

wishes to acknowledge the receipt of a grant from the Fourcade Bequest of the

University of Cape Town for the purchase of a microscope.

REFERENCES

ALDER, J. 1856. A notice of some new genera and species of British hydroid zoophytes. Ann.

Mag. nat. Hist. (2) 18: 353-362.

ALLMAN, G. J. 1864. On the construction and limitation of genera among the Hydroida. Ann.

Mag. nat. Hist. (3) 13: 345-380. ;

ALLMAN, G. J. 1871-2. A monograph of the Gymnoblastic or Tubularian hydroids. London: Ray

Society.

ALLMAN, G. J. 1876. Diagnoses of new genera and species of Hydroida. 7. Linn. Soc. (Xool.) 12:

251-284.

ALLMAN, G. J. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years

1873-76. Part Il1.—The Tubularinae, Corymorphinae, Campanularinae, Sertularinae and

Thalamophora. Rep. Voy. Challenger 1873-76 23 (70): 1-90.

ANNANDALE, N. 1915. Fauna of the Chilka Lake. The Coelenterates of the lake, with an account

of the Actiniaria of brackish water in the Gangetic Delta. Mem. Indian Mus. 5: 65-114.

Bepot, M. 1901-25. Matériaux pour servir 4 histoire des Hydroides. I-VII. Rev. suisse Zool.

9, 13, 18, 20, 24, 26, 32.

BILLARD, A. 1906. Hydroides. Expéd. sci. ‘Travailleur’ et du ‘Talisman’ 8: 153-244.

BILLARD, A. 1907. Hydroides de Madagascar et du sud-est de l’Afrique. Arch. Zool. exp. gén.

(4) 7: 335-396.

Bitiarp, A. 1910. Revision d’une partie de la collection des Hydroides du British Museum.

Ann. Sci. nat. zool. (9) 11: 1-67.

Brazier, J. 1887. Notes on the distribution of Ceratella fusca Gray. Proc. Linn. Soc. N. S. W. (2)

1: 575-576.

Briccs, E. A. 1929. Studies on Australian athecate hydroids. No. 2. Development of the

gonophores and formation of the egg in Mpriothela australis Briggs. Rec. Aust. Mus., 17:

244-264.

Briccs, E. A. 1931. Notes on Australian athecate hydroids. Rec. Aust. Mus. 18: 279-282.

Bricut, K. M. F. 1938. The South African intertidal zone and its relation to ocean currents. II.

An area on the southern part of the west coast. Trans. roy. Soc. S. Afr. 26: 49-65.

Brocu, H. 1916. Hydroida I. Dan. Ingolf-Exped. 5 (6): 1-66.

Brocu, H. 1918. Hydroida II. Dan. Ingolf-Exped. 5 (7): 1-205.

Browne, E. T. and Kramp, P. L. 1939. Hydromedusae from the Falkland Islands. Discovery

Rep. 18: 265-322.

HYDROZOA OF THE SOUTH AND WEST COASTS OF SOUTH AFRICA 485

Carter, H. J. 1873. Transformation of an entire shell into chitinous structure by the polype

Hydractinia, with short descriptions of the polypidoms of five other species. Ann. Mag. nat.

Hist. (4) 14: I-15.

Day, J. H. 1959. The biology of Langebaan Lagoon: a study of the effect of shelter from wave

action. Trans. roy. Soc. S. Afr. 35: 475-547-

Day, J. H., Mittarp, N. A. H. and Harrison, A. D. 1952. The ecology of South African

estuaries. Part II]. Knysna: a clear open estuary. Trans. roy. Soc. S. Afr. 33: 367-413.

EHRENBERG, C. G. 1834. Beitrage zur physiologischen Kenntnis der Corallenthiere im allge-

meinen, und besonders des Rothen Meeres, nebst einem Versuche zur physiologischen

Systematik derselben. Abh. disch. Akad. Wiss. Berl. 1832: 225-380.

Es, J. 1755. An essay towards a natural history of the corallines, and other marine productions of the

like kind, commonly found on the coasts of Great Britain and Ireland. London.

Exus, J. and SoLANDER, D. 1786. The natural history of many curious and uncommon zoophytes,

collected from various parts of the globe by the late Fohn Ellis, esq., F.R.S. London.

Ewer, D. W. 1953. On a new Tubularian hydroid from Natal. Ann. Natal Mus. 12: 351-357.

Fraser, C. McL. 1937. Hydroids of the Pacific coast of Canada and the United States. Toronto:

University Press.

Fraser, C. McL. 1944. Hydroids of the Atlantic coast of North America. Toronto: University Press.

Gitcurist, J. D. F. 1919. On a species of the crawling medusa Eleutheria, from the Cape of

Good Hope (Cnidonema capensis, g. et sp. n.) and the southern Eleutheriae. Quart. F. micr. Scz.

63: 509-529.

Gray, J. E. 1868. Notes on the Ceratelladae, a family of keratose sponges. Proc. zool. Soc. Lond.

1868: 575-579:

HAECKEL, E. 1879. Das System der Medusen. Jena.

Hamonp, R. 1957. Notes on the Hydrozoa of the Norfolk coast. 7. Linn. Soc. (Zool). 43: 294-324.

Hartavs, C. 1897. Die Hydromedusen Helgolands. Wiss. Meeresuntersuch. (N.F.), Bd. 2, Abt.

Helgoland, Heft 1, 10: 449-512.

Hincks, T. 1868. A history of the British hydroid zoophytes. London.

Huve#, P. 1952. Revision des polypes Campanulinides mediterranéens, 1 Partie. Rec. Trav. Sta.

mar. Endoume 4: 34-47.

Hyatt, A. 1877. Revision of the North American Poriferae; with remarks upon foreign species,

Part 2. Mem. Boston Soc. nat. Hist. 2: 481-554.

JADERHOLM, E. 1904. Hydroiden aus den Kiisten von Chile. Ark. Zool. 2: 1-7.

JADERHOLM, E. 1909. Northern and Arctic invertebrates in the collection of the Swedish State

Museum (Riksmuseum). IV. Hydroiden. K. svenska Vetensk Akad. Handl. 45: 1-124. ,

JADERHOLM, E. 1917. Hydroids from the south seas. Norrképings Hégre Allmanna Léroverk, Lasaret,

1916-1917: I-23.

JADERHOLM, E. 1923. Hydroids from West and South Africa. Medd. Géteborg Mus. Zool. 26: 1-7.

Kramp, P. L. 1959. The hydromedusae of the Atlantic Ocean and adjacent waters. Dana Rep.

46: 1-283.

Kramp, P. L. 1961. Synopsis of the medusae of the world. 7. Mar. biol. Ass. U. K. 40: 7-469.

Lamouroux, M. 1812. Extrait d’un mémoire sur la classification des polypiers coralligénes non

entierement pierreux. Nouv. Bull. Sci. Soc. philom. 3: 181-188.

Lamouroux, J. V. F. 1816. Histoire des polypiers coralligénes flexibles, vulgairement nommés zoophytes.

Caen.

LeLoup, E. 1935. Hydraires Calyptoblastiques des Indes Occidentales. Mem. Mus. Hist. nat.

Belg. (2), 2: 1-73.

Letoup, E. 1937. Hydropolypes et Scyphopolypes recueillis par C. Dawydoff sur les cétes de

V’Indochine Frangaise. Mem. Mus. Hist. nat. Belg. (2) 12: 1-73.

LeLoup, E. 1952. Faune de Belgique. Coelentérés. Bruxelles.

Linn£, C. 1758. Systema Naturae. Ed. 10. Holmiae.

LinnE, C, 1767. Systema Naturae. Ed. 12. Holmiae.

Manton, S. M. 1940. On two new species of the hydroid Myriothela. Sci. Rep. Brit. Grahamld

Exped. 1 (4): 255-293.

MarsHALL, W. 1892. Spongiologische Beitrage. Festsch. Rudolf Leuckart. Leipzig.

Mitrarp, N. 1952. Observations and experiments on fouling organisms in Table Bay eben

South Africa. Trans. roy. Soc. S. Afr. 33: 415-445-

Mitrarp, N. A. H. 1955. New species of Hydrozoa from South Africa. Ann. S. Afr. Mus. 41:

215-222,

486 ANNALS OF THE SOUTH AFRICAN MUSEUM

Mitiarp, N. A. H. 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus. 43:

173-243-

Mitrarp, N. A. H. 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa. Part I.

Calyptoblastea. Ann. S. Afr. Mus., 44: 165-226.

Mitrarp, N. A. H. 19594. Hydrozoa from the coasts of Natal and Portuguese East Africa. Part IT.

Gymnoblastea. Ann. S. Afr. Mus. 44: 297-313.

Mitxiarp, N. A. H. 19596. Hydrozoa from ships’ hulls and experimental plates in Cape Town

docks. Ann. S. Afr. Mus. 45: 239-256.

Muinuarp, N. A. H. 1962. The Hydrozoa of the south and west coasts of South Africa. Part I.

The Plumulariidae. Ann. S. Afr. Mus. 46: 261-319.

Mitrarp, N. A. H. 1964. The Hydrozoa of the south and west coasts of South Africa. Part II.

The Lafoeidae, Syntheciidae and Sertulariidae. Ann. S. Afr. Mus. 48: 1-56.

Naumov, D. V. 1960. Hydroids and Hydromedusae of the marine, brackish and freshwater

basins of the U.S.S.R. (In Russian). Opred. Faune SSSR 70: 1-585.

Nurttine, C. C. 1915. American hydroids. Part III. The Campanularidae and the Bonneviellidae.

Spec. Bull. U.S. nat. Mus. 4 (3): 1-126.

Pennycurk, P. R. 1959. Faunistic records from Queensland. Part V.— Marine and brackish

water hydroids. Pap. Dep. Zool. Univ. Qd. 1: 141-210.

PicarpD, J. 1951. Note sur les hydraires littoraux de Banyuls-sur-Mer. Vie et Milieu 2: 338-349.

PicarpD, J. 1955. Hydraires des environs de Castiglione (Algérie). Bull. Soc. Aquic. Péche (N.S.)

7: 181-199.

Raupy, P. M. 1947. The hydroid and medusa of Cnidonema vallentini (Anthomedusae) from

Wellington, New Zealand. Trans. roy. Soc. N.Z. 76: 414-420.

Ravpu, P. M. 1957. New Zealand thecate hydroids. Part I.—Campanulariidae and Campanu-

linidae. Trans. roy. Soc. N.Z. 84: 811-854.

Ra.pu, P. M. 1958. New Zealand thecate hydroids. Part II.— Families Lafoeidae, Lineolariidae,

Haleciidae and Syntheciidae. Trans. roy. Soc. N.Z. 85: 301-356.

Rees, W. J. 1938. Observations on British and Norwegian hydroids and their medusae.

J. Mar: biol. Ass. U.K. 23: 1-42.

Rees, W. J. 1939. A revision of the genus Campanulina van Beneden, 1847. Ann. Mag. nat. Hist.

(II) 3: 433-447.

Rees, W. J. 1956. On the hydroid Merona cornucopiae (Norman). 7. Mar. biol. Ass. U.K. 35:

499-506.

Ritcute, J. 1909. Supplementary report on the hydroids of the Scottish National Antarctic

Expedition. Trans. roy. Soc. Edinb. 477: 65-101.

RussE.L, F. S. 1938. On the nematocycsts of Hydromedusae. 7. Mar. biol. Ass. U.K. 23: 145-165.

RussELL, F. S. 1953. The medusae of the British Isles. Cambridge: University Press.

Sars, M. 1857. Bidrag til kundskaben om Middelhavets Littoral-fauna, reisebemaerkninger fra

Italien. Nytt Mag. Naturv. 9: 110-164.

SPENCER, W. B. 1892. On the structure of Ceratella fusca (Gray). Trans. roy. Soc. Vict. 2: 8-24.

StecHow, E. 1909. Beitrage zur Naturgeschichte Ostasiens: Hydroidpolypen der japanische

Ostktiste. Abh. bayer. Akad. Wiss. math.-nat. 1 (6): 1-111.

StEcHow, E. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer

Gebiete. Zool. Fb. System. 42: 1-172.

STEcHOow, E. 1923a. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und

anderer Gebiete. II. Zool. Fb. System. 47: 29-270.

STEcHow, E. 1923b. Ueber Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen

uber einige andre Formen. Zool. Anz. 56: 97-119.

StEcHow, E. 19252. Hydroiden von West- und Siidwestaustralien nach den Sammlungen von

Prof. Dr. Michaelsen und Prof. Dr. Hartmeyer. Zool. 7b. System. 50: 191-269.

STEcHOW, E. 19255. Hydroiden der Deutschen Tiefsee-Expedition. Wiss. Ergebn. ‘Valdivia’ 17:

383-546.

VANHOFFEN, E. 1910. Die Hydroiden der Deutschen Siidpolar-Expedition 1901-1903. Dtsch.

Stidpol. Exped. 11 (Zool. IIT): 269-340.

Vannuccl, M. 1954. Hidrozoa e Scyphozoa existentes no Instituto Oceanografico. II. Bol.

Inst. oceanogr., S. Paulo 5: 95-138.

Vannuccr, M. and Regs, W. J. 1961. A revision of the genus Bougainvillia (Anthomedusae).

Bol. Inst. oceanogr., S. Paulo 114: 57-100.

HYDROZOA OF THE SOUTH AND WEST CQASTS OF SOUTH AFRICA 487

VeERVvooRT, W. 1946a. Exotic hydroids in the collections of the Rijksmuseum van Natuurlijke

Historie and the Zoological Museum at Amsterdam. Zool. Meded. 26: 287-351.

VERVOORT, W. 1946). Fauna van Nederland. Aflevering XIV. Hydrozoa (C1). A. Hydropolypen.

Leiden.

VeERVooRT, W. 1959. The Hydroida of the tropical west coast of Africa. Atlantide Rep. 5: 211-325.

VERVooRT, W. 1962. A redescription of Solanderia gracilis Duchassaing & Michelin, 1846, and

general notes on the family Solanderiidae (Coelenterata: Hydrozoa). Bull. Mar. Sci. Gulf

Caribb. 12: 508-542.

Warren, E. 1906. On Tubularia solitaria sp. n., a hydroid from the Natal coast. Ann. Natal Mus.

1: 83-06.

Warren, E. 1907. On Parawrightia robusta gen. et sp. nov., a hydroid from the Natal coast;

and also an account of a supposed schizophyte occurring in the gonophores. Ann. Natal

Mus. 1: 187-208.

Warren, E. 1908. On a collection of hydroids, mostly from the Natal coast. Ann. Natal Mus. 1:

269-355.

WEILL, R. 1934. Contribution a l’étude des Cnidaires et de leurs nématocystes. I. Recherches

sur les nématocystes. II. Valeur taxonomique du cnidome. Trav. Sta. zool. Wimereux 10:

1-347; IX: 349-701.

Ann. S. Afr. Mus., Vol. XLVIII Plate 1

Solanderia procumbens (Carter).

A. Whole colony of form A on left (WCD 158F) and form B on right (WCD 158E).

B. Details of the skeleton in form A (WCD 158F). An old stem shown on left with reticulate

meshwork, and a younger stem on right with more marked longitudinal trabeculae and

many hydrophores.

(Photos: Dr. G. 7. Broekhuysen)

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,

including TABLE OF CONTENTs and Summary. Position of text-figures and tables must be

indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. x 7 in. (73 in.

including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in

alphabetical order of authors’ names (Harvard system). References at end of paper must be

given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,

not ‘and’. Year of publication; several papers by the same author in one year designated by

suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper

names (except in German). Title of journal, abbreviated according to World list of scientific

periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).

Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).

Part number, only if separate parts of one volume are independently numbered. Page numbers)

first and last, preceded by a colon (without prefix ‘p’). Thus:

SmirH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;

date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,

with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of

author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of

author of book; his initials; title of book, underlined; edition, if any; volume number, if any,

in arabic numerals, with wavy underlining; pagination of paper; place of publication; name

of publisher. Thus:

SmitH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. and ed. 3: 63-95.

London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species

has been previously designated (subsequent to 1758) are listed in chronological order, with

abbreviated bibliographic references to descriptions or citations following in chronological

order after each name. Full references must be given at the end of the paper. Articles and

recommendations of the International code of zoological nomenclature adopted by the XV International

congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.

When misidentified as another species:

Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).