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| |
| ANNALS OF THE

VOLUME 92

SOUTH AFRICAN MUSEUM ~

ANNALE VAN DIE
SUID-AFRIKAANSE MUSEUM

BAND 92

SAATHSONIAR~
NOV - 9 1984

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 92 BAND

PAE TRUSTEES OF THE DIE{ TRUSTEES, VAN: DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD

1983-1984

SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA
BY THE RUSTICA PRESS (PTY) LTD, WYNBERG, CAPE

D828

List OF CONTENTS

GiLes, E. & GOSLINER, T.
Primary type specimens of marine Mollusca (excluding Cephalopoda) in the South
Puinican Museum. (Published April 19835) so. 6 ace aes e eee eae ne ee eens

GOSLINER, T. see GILES, E.
KENNEDY, W. J. see KLINGER, H. C.

KLINGER, H. C. & KENNEDY, W. J.
Cretaceous faunas from Zululand and Natal, South Africa. The ammonite subfamily
Reroniceratinae Elyatt, 1900. (Published)/March 1984))e ass sae eee sone

Prins, A. J.
Morphological and biological notes on some South African arthropods associated
with decaying organic matter. Part 1. Chilopoda, Diplopoda, Arachnida,
@nustaceasand Insecta. (Published(@ctober 19832). 2255.5. 445462 eo

Prins, A. J.

Morphological and biological notes on some South African arthropods associated
with decaying organic matter. Part 2. The predatory families Carabidae,
Hydrophilidae, Histeridae, Staphylinidae and Silphidae (Coleoptera). (Pub-
sheGsRebnularyalOSAo)h tect man uhetee Monit Sea Actas ARON Stak oak da ae Rye Mae ys

Volume 92 is complete in 4 parts.

Page

JUNG)

53)

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CAPE TOWN

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

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

Fiscuer, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634,

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

Tureve, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L. Zoologische
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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 92 #£2Band
April 1983 April
Part 1 Deel

PRIMARY TYPE SPECIMENS OF MARINE
MOLLUSCA (EXCLUDING CEPHALOPODA)
IN THE SOUTH AFRICAN MUSEUM

By

ELIZABETH GILES
&
TERRENCE GOSLINER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

PRIMARY TYPE SPECIMENS OF MARINE MOLLUSCA
(EXCLUDING CEPHALOPODA)
IN THE SOUTH AFRICAN MUSEUM

By
ELIZABETH GILES
&
TERRENCE GOSLINER
South African Museum, Cape Town

[MS accepted 2 November 1982]

ABSTRACT

A list of type specimens of marine Mollusca contained in the collections of the South
African Museum is presented. Three hundred and thirty-seven species are included in this list.

CONTENTS

PAGE
UNtFOGUCHON). © ees vas ce ceri era ie 1
Polyplacophoratinn 5.42 eee 2
Gastropoday o sasesse sania eens 3
Scaphopoday sic avec Mee 4 Aaa eae oes 40
BV al Vila mee iey ea et cretnn at einen eaves 41
FCHEREMCES ree tre in ay ns caer 49

INTRODUCTION

The molluscan collections housed in the South African Museum have
provided the basis for numerous systematic investigations and extensive studies
by well-known malacologists such as G. B. Sowerby, J. R. Tomlin, and W. H.
Turton. In 1958 K. H. Barnard initiated his work on the molluscan fauna,
based largely on material obtained from the dredging operations of the R.S.
Pieter Faure.

As a result of these research activities spanning more than a century, many
primary type specimens have been deposited in the collections of the South
African Museum. In his posthumously published faunal list of South African
molluscs Barnard (1974) noted some of the type material but a complete list has
not been published. ;

The designation of type specimens was not indicated in most of the works
of the above-mentioned authors and their status has been largely determined
on the basis of review of K. H. Barnard’s research notes, which involve
correspondence with most of his predecessors as well as his own work.

Localities and data are given as they appear in the original publications or
have been supplemented by information contained in catalogues of the mollus-
can holdings in the South African Museum.

1

Ann. S. Afr. Mus. 92 (1), 1983: 1-52.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

SPECIES LIST

Class POLYPLACOPHORA

Family Acanthochitonidae

Acanthochiton turtoni Ashby, 1928: 79, pl. 6 (figs 1-4)
= Acanthochitona garnoti (Blainville): Barnard, 1963d: 328.

Holotype: SAM-A32649. Valve of holotype.
Locality: Port Alfred; no depth or date.
Collected by: Col. W. Turton.

Family Lepidopleuridae

- Lepidopleurus chariessa Barnard, 1963d: 332, fig. 29h.

Syntypes: SAM-—A9338. Two shells, disarticulated.
Locality: off Cape St. Blaize; 125 fm.; 1899.
Collected by: R.S. Pieter Faure.

Family Chitonidae

Chiton ashbyi Barnard, 1963d: 343, fig. 29m.

Holotype: SAM—A9337. Middle valve of holotype.
Locality: off Cape Infanta; 46 fm.; 10 July 1900.
Collected by: R.S. Pieter Faure.

Chiton barnardi Ashby, 1931: 46, pl. 6 (figs 74-76), pl. 7 (fig. 77).

Holotype: SAM—A5331. Shell.

Paratype: SAM—A6590. One valve.

Locality: Mozambique Island (?Inhaca); no depth; 1912.
Collected by: K. H. Barnard.

Chiton (Hanleya) sykesi Sowerby, 1903: 225, pl. 5 (fig. 13)
= Lepidopleurus sykesi (Sowerby): Ashby, 1931: 15, pl. 2 (figs 16-19).

Syntypes: SAM-—5342. Two, girdle on slide.
Locality: off Cape Point; 210 fm.; 11 June 1900.
Collected by: R.S. Pieter Faure.

Family Ischnochitonidae

Chaetopleura pomarium Barnard, 1963d: 335, fig. 291.
Holotype: SAM—A93339. Shell with animal.
Locality: western Cape Peninsula; no depth or date.
Collected by: University of Cape Town.

TYPE SPECIMENS OF MARINE MOLLUSCA

Ischnochiton delagoaensis Ashby, 1931: 40, pl. 6 (figs 63-66).

Holotype: SAM-A6589. Valves, disarticulated.
Locality: Delagoa Bay; no depth; 1912.
Collected by: K. H. Barnard.

Ischnochiton hewitti Ashby, 1931: 33, pl. 5 (figs 50-53).

Holotype: SAM-—A6757. Median valve of holotype.
Paratypes: SAM—A33953. Eight valves.

Locality: Mouille Point, Table Bay; intertidal; 1900.
Collected by: R. M. Lightfoot.

Class GASTROPODA

Subclass PROSOBRANCHIA

Family Pleurotomariidae

Pleurotomaria africana Tomlin, 1948: 2, pl. 1.

Holotype: SAM—A29663. Shell.
Locality: 29°48’S 31°23’E; 366 m; 21 July 1931.
Collected by: R.S. Africana IT.

Family Scissurellidae

Scissurella africanae Barnard, 1963b: 13.

Holotype: SAM—A29933. Shell.
Locality: 30°48’S 52°08’E; 400 m; 9 July 1961.
Collected by: R.S. Africana IT.

Scissurella tabulata Barnard, 1964c: 21.

Holotype: SAM-—A9335 (not A9345). Shell with operculum.
Locality: off Cape Natal; 804 m; 4 April 1901.
Collected by: R.S. Pieter Faure.

Family Fissurellidae

Glyphis spreta Smith, 1901: 104, pl. 1 (fig. 18)
= Diodora spreta (Smith): Tomlin, 1932: 160.

Syntypes: SAM-—2459. Three shells.
Locality: Mossel Bay; intertidal; 1898.
Collected by: Miss Powrie.

Emarginula koon Kilburn, 1978: 439, pl. 5.

Paratype: SAM—A339724. Shell.
Locality: Shelly Beach near Port Shepstone; November 1965.
Collected by: R. Cock.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Emarginula natalensis Barnard, 1963d: 297, fig. 23.

Syntypes: SAM-—A9325. Three shells, worn.
Locality: off Cape Morgan; 141 m; 26 July 1901.
Collected by: R.S. Pieter Faure.

Emarginula oppressa Barnard, 1963d: 298, fig. 24a
= Emarginella oppressa (Barnard): Kilburn, 1978: 446, pl. 8 (fig. b).

Holotype: SAM-—A9330. Shell.
Locality: off Cape Natal; 98 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Emarginula pulchreclathrata Tomlin, 1932: 162, fig. 4
= Emarginula agulhasensis Thiele: Kilburn, 1978: 433, pl. 2a—b.

Holotype: SAM-—A3617. Shell.
Locality: Saldanha Bay; 98 m; 13 March 1902.
Collected by: R.S. Pieter Faure.

Emarginula thorektes Kilburn, 1978: 440, pl. 6a—c.

Paratype: SAM-A9323. Shell.
Locality: off Cape Morgan; 141 m; 26 July 1901.
Collected by: R.S. Pieter Faure.

Emarginula vadum Barnard, 1963d: 297, figs 23d—24b.

Holotype: SAM—A9329. Shell.
Locality: off Cape Vidal; 146-183 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Parmaphorella barnardi Tomlin, 1932: 164, fig. 5
= Tugali barnardi (Tomlin): Kilburn, 1978: 452, pl. 11 (figs b-d).

Holotype: SAM—A3623. Shell.
Locality: off Cape Point; 344 m; 27 February 1902.
Collected by: R.S. Pieter Faure.

Puncturella (Vacerrena) christiaensi Kilburn, 1978: 448, pl. 9 (figs c—e).
Paratype: SAM-—A33925. Shell.
Locality: off Sodwana Bay; 50 m; 1975.
Collected by: R.V. Meiring Naude.

Family Trochidae

Angaria lacunosa Barnard, 1963d: 283.

Holotype: SAM-A9281. Shell.
Locality: off Natal; no depth or date.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 5

Bassilissa gelida Barnard, 19636: 440, fig. 8b.
Holotype: SAM-—A9720. Shell, radula, slide.
Locality: 33°26'S 16°33’E; 2 380 m; August 1959.
Collected by: F. H. Talbot on board R.S. Africana IT.

Calliostoma circus Barnard, 1969: 653, fig. 27b.
Holotype: SAM—A30032. Shell with operculum, radula slide.
Locality: off Cape Point; no depth or date.
Collected by: commercial trawler.

Calliostoma glaucophaos Barnard; 19636: 442, fig. 8c.
Syntypes: SAM—A9830. Two shells with opercula, radula slide.
Locality: 33°52’S 16°51’E; 2 520—2 810 m; December 1959.
Collected by: F. H. Talbot on board R.S. Africana II.

Calliostoma (Lischkeia) granoliratum Sowerby, 1903: 222, pl. 5 (fig. 7)
= Calliotropis granolirata (Sowerby): Thiele, 1925: 48.

Holotype: SAM-—A5296. Shell with operculum.

Paratype: SAM—A33690 (previously A5296). Shell.

Locality: off Cape Point; 665 m; 12 June 1900.

Collected by: R.S. Pieter Faure.

Calliostoma iridescens Sowerby, 1903: 223, pl. 5 (fig. 4).
Holotype: SAM-A5294. Shell.
Locality: off Cape Natal; 101 m; 24 December 1900.
Collected by: R.S. Pieter Faure.

Calliostoma perfragile Sowerby, 1903: 222, pl. 5 (fig. 3).
Holotype: SAM-A5275. Shell.
Locality: off Cape Point; 282-304 m; 18 April 1900.
Collected by: R.S. Pieter Faure.

Calliotropis chenoderma Barnard, 1963d: 263, fig. 16c.
Syntype: SAM-A9296. Shell.
Locality: off Cape St. Blaize; 229 m; 21 December 1899.
Syntype: SAM-—A97297. Shell.
Locality: off Cape Morgan; 86 m; 25 July 1901.
Collected by: R.S. Pieter Faure.

Calliotropis persculpta (Sowerby): see Solariella persculpta p. 7.

Calliotropis pompe Barnard, 1963c: 440, fig. 8b.
Syntypes: SAM-—A9883. Two shells, one with operculum.
Locality: off Cape Point, 34°42’S 16°54’E; 1 725-1 789 fm.; December
1959)
Syntype: SAM-A979S. Shell.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Cape Point, 33°50’S 16°30’E; 1 450-1 660 fm.; December
1959.
Collected by: F. H. Talbot on board R.S. Africana II.

Gibbula beckeri Sowerby, 1901: 214, pl. 22 (fig. 22).

Paratype: SAM-A3321. Shell.
Locality: Port Alfred; intertidal; 1915.
Collected by: H. Becker.

Gibbula (Enida) perspectiva Sowerby, 1900: 6, pl. 1 (fig. 17).
Paratype: SAM-—A3320. Shell.
Locality: Pondoland; intertidal; 1915.
Collected by: H. Becker.

Guttula blanda Barnard, 1963d: 265, fig. 17.

Holotype: SAM—A7424. Shell, radula slide.
Locality: off Cape Point; 1 270 m; 20 August 1903.
Collected by: R.S. Pieter Faure.

Minolia adarticulata Barnard, 1963d: 235, fig. 11f.

Syntypes: SAM-—A9277. Two shells.
Locality: off Hood Point, East London; 90 m; 15 July 1901.
Collected by: R.S. Pieter Faure.

Minolia cycloma Barnard, 1964c: 18, fig. 2c—d.

Holotype: SAM-—A9279. Shell.
Locality: off Tugela River, Natal; 67 m; 9 January 1901.
Collected by: R.S. Pieter Faure.

Minolia congener Sowerby, 1903: 223, pl. 5 (fig. 2)
= Solariella laevissima (von Martens): Barnard, 1963d: 239, fig. 10c.

Syntypes: SAM—A5257. Two shells with opercula.
Locality: off Cape St. Blaize; 67 m; 22 October 1900.
Collected by: R.S. Pieter Faure.

Seguenzia fatigans Barnard, 1963b: 13, pl. 2 (figs 10-11).
Syntypes: SAM-—A29930. Two shells.
Locality: 27°48’S 47°19’E; 875 m; 22 June 1961.
Collected by: R.S. Africana IT.

Seguenzia simplex Barnard, 1963d: 265, fig. 16a.
Syntypes: SAM-—A7421. Four shells (one juvenile); radula slide.
Locality: off Cape Point; 1 280 m; 20 August 1903.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA

Solariella beckeri Sowerby, 1901: 214, pl. 22 (fig. 21)
= Solariella laevissima (von Martens): Barnard, 1963d: 239, fig. 10c.

Paratype: SAM-A3323. Shell.
Locality: Kowie (Port Alfred); no depth; 1915.
Collected by: H. Becker.

Solariella dowi Barnard, 1963b: 13, pl. 2 (figs 10-11).

Syntypes: SAM—A29932. Two shells.
Locality: 27°48’'S 47°19’E; 875 m; 22 June 1961.
Collected by: R.S. Africana IT.

Solariella franciscana Barnard, 1963d: 243, figs 10e—11b
= Spectamen franciscana (Barnard): Kilburn, 1977: 178.

Holotype: SAM—A3615. Shell with operculum, radula slide.
Locality: off Cape St. Francis; 137 m; 19 February 1902.
Collected by: R.S. Pieter Faure.

Solariella gilchristi Barnard, 1963d: 242, fig. 11d.

Syntypes: SAM-—A3605. Two shells.
Locality: off Buffalo River; 564 m; 24 April 1901.
Collected by: R.S. Pieter Faure.

Solariella macleari Barnard, 1963d: 241, fig. 11c.

Syntypes: SAM—A7417. Four shells, one apex, two opercula, radula slide.
Locality: ‘off Cape Point N 89°E, 36 miles’; 1 280 m; 20 August 1903.
Collected by: R.S. Pieter Faure.

Solariella palirrous Barnard, 1963a: 12, pl. 2 (figs 7-8).

Holotype: SAM—A29931. Shell broken.
Locality: 36°48’S 52°08’E; 400 m; 9 July 1961.
Collected by: R.S. Africana II.

Solariella persculpta Sowerby, 1903: 223, pl. 5 (fig. 8)

= Calliotropis persculpta (Sowerby): Thiele, 1925: 48.

Syntype: SAM-—A5251. Shell.
Locality: off Cape Natal; 800 m; 4 April 1901.
Collected by: R.S. Pieter Faure.

Solariella turbynei Barnard, 1963d: 243, fig. 11a.

Holotype: SAM-A9276. Shell.
Locality: off Cape Point; 246 m; 17 February 1902.
Collected by: R.S. Pieter Faure.

Turcica helix Barnard, 1964c: 21, fig. e.

Holotype: SAM-A9295. Shell.
Locality: off Cape Vidal; 146-183 m; 27 February 1901.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratype: SAM-A9256. Shell, broken.
Locality: off Umkomaas; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Turcica konos Barnard, 1964c: 20, fig. 3f.
Syntypes: SAM-—A9257. Four shells (two adult, one immature, one
broken).
Locality: off Umkomaas River; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Turcica salpinx Barnard, 1964c: 19, fig. 3a—d.
Syntype: SAM-—A9252. Shell, adult, broken.
Locality: off Cape Morgan; 40 m; 26 July 1901.
Syntypes: SAM-—A9253. Five shells (three broken, two juvenile).
Locality: off Hood Point; 90 m; 15 July 1901.
Collected by: R.S. Pieter Faure.

Family Liotiidae

Brookula coronis Barnard, 1963b: 12, pl. 2 (figs 5-6).
Syntypes: SAM-—A29929. Nine shells.
Locality: 36°48’S 52°08’E; 400 m; 9 July 1961.
Collected by: R.S. Africana IT.

Leptothyra echomenus Barnard, 1963b: 12, pl. 2 (figs 3-4).
Syntypes: SAM-—A29928. Five shells.
Locality: 36°48’S 52°08’E; 400 m; 9 July 1961.
Collected by: R.S. Africana IT.

Leptothyra sola Barnard, 1963d: 231, fig. 9e.
Holotype: SAM-—A9288. Shell, probably juvenile.
Locality: probably Algoa Bay; no depth or date.
Collected by: R.S. Pieter Faure.

Leptothyra syngenes Barnard 1963b: 11, pl. 2, figs 2, 4.
Syntypes: SAM—A29927. Twelve shells, some broken.
Locality: 36°48’S 52°08’E; 400 m; 9 July 1961.
Collected by: R.S. Africana II.

Rufulana sextula Barnard, 1963d: 232, fig. 9f.
Syntypes: SAM—A29634. Three shells.
Locality: probably Algoa Bay; 55 m; 10 November 1898.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA

Family Turbinidae

Astralium (Cyclocantha) gilchristi Sowerby 1903: 221, pl. 5 (fig. 6)
= Astrea gilchristi (Sowerby): Barnard 1963d: 221, fig. 7c.

Syntype: SAM-A5219. Shell.
Locality: off Scottburgh; 168 m; 7 March 1901.
Collected by: R.S. Pieter Faure.

Calcar rhysopoma Barnard, 1964b: 17, fig. 2a—b.

Syntypes: SAM—A9285. Fourteen shells, one operculum, radula slide.
Locality: off Umkomaas River; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Turbo boswellae Barnard, 1969: 652, fig. 27a.

Holotype: SAM—A30031. Shell, operculum, radula slide.
Locality: off Cape Point; 366 m; no date.
Collected by: commercial trawler.

Turbo foveolatus Barnard, 1963d: 216, fig. 6.

Syntypes: SAM-—A9284. Five shells.
Locality: 34°05’S 25°52’E; 95 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

Family Rissoidae

Amphithalamus muiri Barnard, 1963a: 188, fig. 15d.

Syntypes: SAM—A9262. Eleven shells.
Locality: Still Bay; intertidal zone; no date.
Collected by: T. Muir.

Family Vitrinellidae

Cochliopsis catherinae Barnard 1963a: 180, fig. 35a.

Holotype: SAM-A9221. Shell.

Locality: 34°27'S 25°42'E; 473 m; 14 November 1898.
Paratype: SAM—A9341. Shell.

Locality: 33°50'S 25°54'E; 61 m; 11 November 1898.
Collected by: R.S. Pieter Faure.

Cochliopsis tugelae Barnard, 1963a: 180, fig. 35b.

Holotype: SAM-—A9282. Shell.
Locality: off Tugela River; 26 m; 16 January 1901.
Collected by: R.S. Pieter Faure.

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

Vitrinella inclinans Barnard 1963a: 179, fig. 35c.
Syntypes: SAM—A29680. Six shells with animals, one operculum.
Locality: 34°27’S 25°45’E; 463 m; 19 November 1898.
Collected by: R.S. Pieter Faure.

Family Turritellidae

Turritella chrysotoxa Tomlin, 1925: 315, fig. 3a.
Syntype: SAM-—A3561. Shell.
Locality: off Illovo River; 50 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Turritella kowiensis Sowerby, 1900: 6, pl. 1 (fig. 12)

-= Turritella carinifera form kowiensis (Sowerby): Barnard, 1963a: 166.
Syntypes: SAM-—A3315. Two shells.

Locality: Kowie, Port Alfred; no depth; 1915.

Collected by: H. Becker.

Turritella salisburyi Tomlin, 1925: 314, fig. 36

= Turritella sanguinea Reeve: Barnard, 1963a: 169.
Syntype: SAM—A5113. Shell.
Syntype: SAM—A33952. Shell.
Locality: off East London; 95 m; 12 July 1901.
Collected by: R.S. Pieter Faure.

Family Vermetidae

Vermetus (Stoa) corralinaceus Tomlin, 1939: 145, pl. 12 (fig. 4).
Paratypes: SAM-A31494. Two blocks of tubes, radula slide.
Locality: Oudekraal, west Cape Peninsula; no depth or date.
Collected by: University of Cape Town.

Vermetus periscopium Barnard, 1963a: 146, fig. 30a.

Paratypes: SAM-—A29925. Four shells, one operculum, four animals,
radula slide.

Locality: 29°46’S 31°17’E; 110-130 m; 23 April 1958.

Collected by: University of Cape Town.

Vermetus quincunx Barnard, 1963a: 148, figs 28c, 29b-c.
Syntypes: SAM-A29814. Large mass, numerous fragments, protoconchs,
two opercula, radula slide.
Locality: 34°27’S 25°42'E; 470 m; 19 November 1898.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 1

Family Cerithiidae

Alabina telamon Barnard, 1963a: 138, fig. 26d.

Holotype: SAM—A29676. Shell.
Locality: off Tugela River; 118-146 m; 11 January 1901.
Collected by: R.S. Pieter Faure.

Family Cerithiopsidae

Cerithiella natalensis Barnard, 1963a: 126.

Holotype: SAM-—A9045. Shell.
Locality: off Umkomaas River; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Cerithiella nonnitens Barnard, 1963a: 127, fig. 23g.

Holotype: SAM—A9047. Shell.
Locality: off Cove Rock, East London; 146-183 m; 30 July 1900.
Collected by: R.S. Pieter Faure.

Cerithiella (Stilus) subuliapex Barnard, 1963b: 10, pl. 1 (fig. 8).

Holotype: SAM-A29924. Shell.
Locality: 27°48’'S 47°19’E; 875 m; 22 June 1961.
Collected by: R.S. Africana IT.

Cerithiella taylori Barnard, 1963a: 126, fig. 231.

Holotype: SAM-A9711. Shell.
Locality: 33°26’S 16°33’E; 2 260-2 370 m; August 1959.
Collected by: R.S. Africana IT.

Cerithiella vidalensis Barnard, 1963a: 127.

Holotype: SAM-A9046. Shell.
Locality: off Cape Vidal; 146-183 m; 27 February 1901.
Collected by: R.S. Pieter Faure.

Cerithiopsis rota Barnard, 1963a: 124, fig. 23a—c.

Holotype: SAM-—A9043. Shell.
Locality: 34°25’'S 44°33'E; 61 m; 10 November 1898.
Collected by: R.S. Pieter Faure.

Family Finellidae

Cerithidium fragrans Barnard, 1963a: 139, fig. 15c.

Syntypes: SAM-—A29679. Five shells, fossils.
Locality: raised beach, Little Brak River; no date.
Collector unknown.

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Abyssochrysidae

Abyssochrysos tomlini Barnard, 1963a: 143, fig. 27f
= Abyssochrysos melvilli (Schepman): Houbrick, 1979: 10, figs 7d, f, i, 8.
Lectotype: SAM—A9101. Shell.
Paralectotype: SAM-—A33950 (previously A9109). Shell.
Locality: off Cape Natal; 535 m; 4 April 1901.
Collected by: R.S. Pieter Faure.

Abyssochrysos melanoides Tomlin, 1927: 78. figs 1-3.
Syntype: SAM—AS5116. Shell.
Locality: off Cape Point; 1 462-2 724 m; 21 July 1903.
Syntypes: SAM-A33951 (previously A5116). Twelve shells, four
opercula, one radula.
Collected by: R.S. Pieter Faure.

Family Triforidae

Trifora ? axialis Barnard, 19636: 10, pl. 1 (fig. 7) (sic).
Syntypes: SAM-—A29923. Two shells.
Locality: 36°48’S 52°08’E; 400 m; 9 July 1961.
Collected by: R.S. Africana IT.

Trifora bactron Barnard, 1963a: 115, fig. 20c.
Holotype: SAM-—A9077. Shell.
Locality: off Umkomaas River; 73 m; 31 December 1900.
Paratype: SAM-A9076. Shell.
Locality: off Umhloti River; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Trifora baculus Barnard, 1963a: 115, fig. 20b.
Holotype: SAM-—A9078. Shell.
Locality: off Sandy Point north of Cape Morgan; 93 m; 14 August 1901.
Collected by: R.S. Pieter Faure.

Trifora dagama Barnard, 1963a: 112, fig. 19f.
Syntype: SAM—A9073. Shell.
Locality: off Umkomaas River; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Trifora morgani Barnard, 1963a: 111, fig. 19a.
Holotype: SAM-—A9079. Shell, broken.
Locality: off Sandy Point north of Cape Morgan; 93 m; 14 August 1901.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 13

Trifora natalensis Barnard, 1963a: 112, fig. 19c.
Syntype: SAM-—A9071. Shell.
Locality: off Umhloti River; 73 m; 18 December 1900.
Syntype: SAM-—A9072. Shell.
Locality: off Umkomaas River; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Trifora scala Barnard, 1963a: 115, fig. 19d.

Syntype: SAM-A9074. Shell.

Locality: off Umkomaas River; 73 m; 31 December 1900.
Syntype: SAM-—A9075. Shell.

Locality: off Umhloti River; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Family Pyramidellidae

Odostomia tugelae Barnard, 1963a: 93, fig. 15b.

Holotype: SAM-—A9225. Shell.
Locality: off Tugela River; 65-80 fm.; 11 January 1901.
Collected by: R.S. Pieter Faure.

Turbonilla bathybias Barnard, 1963a: 89.

Syntypes: SAM-—A9175. One shell, one apex.
Locality: off East London; 400-450 fm.; 17 April 1901.
Collected by: R.S. Pieter Faure.

Turbonilla? bifilosa Barnard, 1963a: 90, fig. 15a.

Holotype: SAM—A9220. Shell.
Locality: 34°05'S 25°55’E; 122 m; 1 November 1898.
Collected by: R.S. Pieter Faure.

Turbonilla deprofundis Barnard, 1963a: 89, fig. 14b.

Holotype: SAM-A9157. Shell.
Locality: off East London; 400-450 fm.; 17 April 1901.
Collected by: R.S. Pieter Faure.

Turbonilla tugelae Barnard, 1963a: 88, fig. 14f.

Syntypes: SAM—A9153. Shell, apex.
Locality: off Tugela River; 14 fm.; 16 January 1901.
Collected by: R.S. Pieter Faure.

Family Epitoniidae

Acrilla analogica Barnard, 1963a: 106, fig. 18g—h.

Holotype: SAM-—A9026. Protoconch with nine whorls.
Locality: 36°40'S 21°26’E; 366 m; 17 July 1906.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratype: SAM-—A9229. Shell, broken.
Locality: 34°27’S 25°42'E; 467 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

Acrilla natalis Barnard, 1963a: 105, fig. 18e-f.
Syntypes: SAM-—A9028. Two shells, broken.
Locality: off Cape Natal; 99 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Scala blaisei Barnard, 1963a: 102, fig. 18a

= Epitonium blaisei (Barnard): comb. nov.
Holotype: SAM—A9025. Shell.
Locality: off Cape St. Blaize; 228 m; 21 December 1899.
Collected by: R.S. Pieter Faure.

Scala bonaspei Barnard, 1963a: 104, fig. 17b
= Epitonium bonaspei (Barnard): comb. nov.
Syntypes: SAM-—A9761. Two shells with opercula, proboscis, jaws, radula
of larger specimen on slide.
Syntype: SAM-—A9793. Shell.
Locality: off Cape Point; 3 120-3 260 m; December 1959.
Collected by: R.S. Africana IT.

Scala illovoensis Barnard, 1963a: 101, fig. 18b

= Epitonium illovoensis (Barnard): comb. nov.
Holotype: SAM-—A9023. Shell.
Locality: off Illovo River; 50-55 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Scala mathildopsis Barnard, 1963b: 9, pl. 1 (fig. 5)
=Epitonium mathildopsis (Barnard): comb. nov.
Syntypes: SAM-—A29921. Six shells.
Locality: 27°48’S 47°19'E; 875 m; 22 June 1961.
Collected by: R.S. Africana IT.

Scala munda Barnard, 1969: 649, fig. 24b

= Epitonium mundum (Barnard): comb. nov.
Holotype: SAM—A30552. Shell.
Locality: off Cape Point; 1 250 m; 20 August 1903.
Paratypes: SAM-—A7419. Three shells.
Locality: off Cape Point; 1 280 m; 20 August 1903.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 15

Scala tenebrosa Sowerby, 1903: 220, pl. 4 (fig. 6)
= Epitonium tenebrosum (Sowerby): comb. nov.

Syntype: SAM-—A3599. Shell.
Locality: off Cape St. Blaize; 67 m; 22 October 1900.
Collected by: R.S. Pieter Faure.

Family Mathildidae

Mathilda salve Barnard, 1963a: 176, fig. 34.

Holotype: SAM-A9139. Shell.

Locality: off Cape St. Blaize; 229 m; 21 December 1899.

Paratype: SAM-—A9141. Apex of seven whorls, no protoconch.
Locality: off Glendower Beacon, Port Alfred; 183 m; 26 July 1901.
Collected by: R.S. Pieter Faure.

Family Architectonicidae

Fluxina solarium Barnard, 1963b: 11, pl. 2 (fig. 1).

Holotype: SAM—A29926. Shell.
Locality: 27°48’'S 47°19’E; 875 m; 22 June 1961.
Collected by: R.S. Africana IT.

Heliacus numulus Barnard, 1963a: 163, fig. 31e.

Holotype: SAM-A9125. Shell.
Locality: off Cape Morgan; 141 m; 26 July 1901.
Collected by: R.S. Pieter Faure.

Heliacus obolos Barnard, 1963a: 163, fig. 31f-g.

Holotype: SAM-—A9127. Shell, broken.
Locality: off Umhloti River; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Heliacus petasus Tomlin, 1928: 334, pl. 26 (fig. 4).

Holotype: SAM-—A3579. Shell.
Locality: off Scottburgh; 169 m; 7 March 1901.
Collected by: R.S. Pieter Faure.

Family Stiliferidae

Apicalia sowerbyi Barnard, 1963a: 94, fig. 16.
Syntypes: SAM-A9169. Two shells.
Locality: off Hood Point, near East London; 90 m; 15 July 1901.
Collected by: R.S. Pieter Faure.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Fossaridae

Fossarus translucens Barnard, 1969: 647, fig. 23a.
Holotype: SAM—A30052. Shell.
Locality: Gordon’s Bay, False Bay; intertidal; no date.
Collected by: Dr Ackert.

Megalomphalus mosselensis Barnard, 1957: 180, figs b—d.
Holotype: SAM-A29922. Shell, six embryo ex utera, radula slide.
Locality: Mossel Bay; no depth or date.
Collected by: University of Cape Town.

Family Amaltheidae

- Cheila microstriata Barnard, 1963a: 76, fig. 10).

Syntype: SAM-A9004. Shell, broken.

Locality: off Durban; 153 m; 17 December 1900.
Syntype: SAM-—A9005. Shell.

Locality: off Umhloti River; 73 m; 18 December 1900.
Syntype: SAM—A9006. Shell, broken.

Locality: off Durnford Point, Natal; 164 m; 11 July 1900.
Syntype: SAM-A9342. Shell.

Locality: off Cape Vidal; 146-183 m; 27 February 1901.
Collected by: R.S. Pieter Faure.

Family Trichotropidae

Trichotropis zuluensis Barnard, 1963a: 78, figs 12d—13c.
Syntype: SAM—A9011. Shell, radula slide.
Locality: off Durnford Point, Natal; 165 m; 28 February 1901.
Syntype: SAM-A9012. Shell.
Locality: off Cape Vidal; 146-183 m; 27 February 1901.
Collected by: R.S. Pieter Faure.

Family Aporrhaidae

Aporrhais pesgallinae Barnard, 1963a: 67, fig. 8.

Holotype: SAM—A30248. Shell with animal in alcohol, radula slide.
Locality: 26°23’S 14°17’E; 311 m; 28 October 1948.
Collected by: R.S. Africana I.

Family Naticidae

Falsilunatia pseudopsila Barnard, 1963a: 64, fig. 7e; 1969: 646, fig. 22.
Holotype: SAM-A9780. Shell with operculum, radula slide.

TYPE SPECIMENS OF MARINE MOLLUSCA 7

Locality: off Cape Point; 2 920 m; December 1959.
Collected by: R.S. Africana IT.

Polynices cleistopsila Barnard, 1963a: 64, fig. 7d.
Syntypes: SAM-—A9735. One adult shell, two juveniles, three opercula,
radula slide.
Locality: Off Cape Point, 33°36’S 16°15’E; 2 830 m; August 1959.
Collected by: R.S. Africana IT.

Family Cypraeidae

Cypraea fultoni Sowerby, 1903: 218, pl. 4 (fig. 7).
Holotype: SAM—A5053. Shell.

Locality: South Africa; ex pisce; no depth or date.
Collected by: R.S. Pieter Faure.

Family Triviidae

Trivia verhoefi Gosliner & Liltved, 1982: 121, figs 1f, 13-16.
Holotype: SAM—A34387. Shell with animal.
Locality: Llandudno, 34°01'S 18°20’E; 25 m; 22 December 1979.
Collected by: G. Verhoef.
Paratype: SAM—A34388. Shell with animal.
Locality: Llandudno; 20 m; 16 March 1980.
Collected by: W. R. Liltved.
Paratype: SAM—A34389. Shell with animal.
Locality: Llandudno, 25 m; 27 March 1980.
Collected by: W. R. Liltved.

Family Ovulidae

Amphiperas beckeri Sowerby, 1900: 5, pl. 1 (fig. 13)
= Primovula beckeri (Sowerby): Thiele, 1929; Barnard 1963a: 55, figs 5d—6c.

Paratypes: SAM-A8918. Four shells.
Locality: Kowie, Port Alfred; intertidal; 71899.
Collected by: H. Becker.

Family Tonnidae

Eudolium aulacoides Tomlin, 1927: 83, fig. 4a.
Holotype: SAM-—A3529. Shell.
Locality: off Cape Point; 1 020-1 280 m; 17 September 1903.
Collected by: R.S. Pieter Faure.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Cymatiidae

Thalassocyon bonus Barnard, 1960: 440, fig. 3.
Holotype: SAM-A9714. Shell, operculum broken, radula slide A9810.
Locality: off Cape Point; 2 280-2 380 m; August 1959.
Collected by: R.S. Africana IT.

Family Columbariidae

Columbarium angulare Barnard, 1959: 235, fig. 52c.
Syntypes: SAM-—A4608. Three shells, operculum, radula slide.
Locality: off Cape Point; 1 480-1 650 m; 14 July 1903.
Collected by: R.S. Pieter Faure.

_Columbarium formosissimum Tomlin, 1928: 331, pl. 25 (fig. 1).

Holotype: SAM-—A3501. Shell.

Locality: Cape St. Blaize; ‘N by E4E 65 miles’; 90 fm.; 22 December
1899.

Collected by: R.S. Pieter Faure.

Paratype: SAM-—A3499; Shell.

Locality: Cape Seal; ‘N by W 2 W 55 miles’; 87 fm.; 19 February 1902.

Collected by: R.S. Pieter Faure.

Paratype: SAM-—A3450; Shell.

Locality: Cape Seal; ‘N by W 2 W 55 miles’; 87 fm.; 19 February 1902.

Collected by: R. S. Pieter Faure.

Paratype: SAM—A3498; Shell.

Locality: Glendower Beacon; ‘N 2 W 6 miles’; 66 fm.; 10 September 1901.

Collected by: R.S. Pieter Faure.

Columbarium natalense Tomlin, 1928: 331, pl. 25 (fig. 2).

Holotype: SAM-A3497. Shell.
Locality: off Cape Natal; 153 m; 17 December 1900.
Collected by: R.S. Pieter Faure.

Columbarium rotundum Barnard, 1959: 236, fig. 52d.

Syntypes: SAM-—A4592. Two shells, two opercula.
Locality: off Cape Point; 1 400 m; 27 August 1903.
Collected by: R.S. Pieter Faure.

Family Colubrariidae

Epidromus crebrilirata Sowerby, 1903: 220, pl. 4 (fig. 4)

= Nivitriton fictilis (Hinds): Kilburn, 1977: 194.
Holotype: SAM-A4974. Shell.
Locality: off Glendower Beacon, Port Alfred; 183 m; 10 September 1901.
Collected by: R.S. Pieter Faure.

a

TYPE SPECIMENS OF MARINE MOLLUSCA 19

Family Muricidae

Tritonalia juritzi Barnard, 1969: 639, fig. 18d.
Holotype: SAM—A30051. Shell, broken.
Locality: doubtful, probably Cape Peninsula.
Collected by: C. Juritz.

Trophon acceptans Barnard, 1959: 202, figs 40d—43b; 1963c; 435, fig. 7a; 1969:
638.

Holotype: SAM—A3473. Shell with operculum, radula slide.

Locality: off Cape Point, ‘N 64° E 37 miles’; 1 270-1 460 m; 9 September
1903.

Paratype: SAM-—A3449. Three shells.

Locality: off Cape Point, ‘E 7° N 38 miles’; 1 190 m; 9 September 1903.

Paratype: SAM-—A3480. Shell.

Locality: off Cape Point, ‘64° NE 37 miles’; 1 270-1 460 m; 10 September
1903.

Collected by: R.S. Pieter Faure.

Trophon? incertus Barnard, 1959: 203, fig. 32a.
Syntypes: SAM—A8843. Two shells.
Locality: off Cape Natal; 800 m; 4 April 1901.
Collected by: R.S. Pieter Faure.

Trophon johanthielei Barnard, 1959: 206, fig. 44b.
Holotype: SAM-A8844. Shell, broken.
Locality: off East London; 730-820 m; 17 April 1901.
Collected by: R.S. Pieter Faure.

Trophon mioplectos Barnard, 1959: 206, fig. 44a.
Holotype: SAM-A8631. Shell.
Locality: 34°27’S 25°42'E; 466 m; 16 November 1898.
Collected by: R.S. Pieter Faure.

Trophon pistillum Barnard, 1959: 206, fig. 44f.
Holotype: SAM—A8842. Shell, broken; radula slide.
Locality: off Cape Natal; 802 m; 4 April 1901.
Collected by: R.S. Pieter Faure.

Typhus pentaphasios Barnard, 1959: 211, fig. 46h-1
= Siphonochelus pentaphasios (Barnard 1959): Radwin & D’Attilo 1976: 199,
fig. 144.
Syntypes: SAM-—A4949. Two shells.
Locality: off Cape Point; 1 204 m; 10 September 1903.
Collected by: R.S. Pieter Faure.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Coralliophilidae

Coralliophila isosceles Barnard, 1959: 191, fig. 39a.
Syntypes: SAM-—A4951. Two shells.
Locality: off Glendower Beacon, Port Alfred; 121 m; 10 September 1901.
Collected by: R.S. Pieter Faure.

Coralliophila zuluensis Barnard, 1959: 192, fig. 39b.

Holotype: SAM—A8852. Shell.
Locality: off O’Neil Peak, Natal; 163 m; 28 February 1901.
Collected by: R.S. Pieter Faure.

Latiaxis capensis Tomlin, 1928: 332.

Holotype: SAM-A3522. Shell.
Locality: off Sandy Point, Transkei; 173 m; 14 August 1901.
Collected by: R.S. Pieter Faure.

Latiaxis? elstoni Barnard, 1962b: 148, fig. 1.

Holotype: SAM-—A9346. Shell.
Locality: Natal coast; ex pisce; donated 1961.
Collector unknown.

Latiaxis kylix Barnard, 1959: 188, fig. 27
= Latiaxis mawae kylix Barnard: Kilburn, 1973: 565, fig. 9b.

Holotype: SAM—A8850. Shell, juvenile.
Locality: off Cape Natal; 98 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Family Buccinidae

Babylonia pintado Kilburn, 1971: 486, figs 3, 7c, e.
Paratypes: SAM—A4751. Shell.
Locality: off Port Shepstone, “WNW 8 miles’; 27 fm.; 14 March 1901.
Collected by: R.S. Pieter Faure.
Paratype: SAM-—A7706. Shell.
Locality: Natal; ex pisce; no depth or date.
Collected by: Mr Casey.
Paratype: SAM-A8239. Shell.
Locality: Natal; ex pisce; no depth or date.
Collected by: H. W. Bell-Marley.
Paratypes: SAM-—A30956. Six shells.
Locality: Natal; ex pisce; no depth or date.
Collected by: R. Frames.

TYPE SPECIMENS OF MARINE MOLLUSCA 21

Glypteuthria solidissima Tomlin, 1932: 166, fig. 7

= Afrocominella elongata (Dunker): Barnard, 1959: 153, fig. 31f.
Holotype: SAM-A3543. Shell.
Locality: off Cape Point; 82 m; 6 June 1900.
Collected by: R.S. Pieter Faure.

Neptunea bonaespei Barnard, 1960: 440, fig. 3.
Syntypes: SAM-—A9826. Three shells (one broken), operculum, radula
slide.
Locality: off Cape Point, 33°52’S 16°51’E; 2 520-2 820 m; December 1959.
Collected by: R.S. Africana IT.

Nux alabaster Barnard, 1960: 440, fig. 3.
Holotype: SAM-—A9715. Shell with operculum, radula slide.
Locality: off Cape Point, 33°26’S 16°33’E; 2 270-2 380 m; December 1959.
Paratypes: SAM-—A9827. Two shells with opercula.
Locality: off Cape Point; 35°52’S 16°51'E; 2 540-2 820 m; December 1959.
Collected by: R.S. Africana IT.

Prosipho torquatus Barnard, 1963c: 433, fig. 6c—e.

Holotype: SAM-A9884. Shell with operculum, radula slide.
Locality: off Cape Point, 34°37’S 17°03’E; 2 890-2 960 m; December 1959.
Collected by: R.S. Africana IT.

Family Columbellidae

Columbella beckeri Sowerby, 1900: 3, pl. 1 (fig. 7).

Syntype: SAM-A33272. Shell.
Locality: Port Alfred; intertidal; 1915.
Collected by: H. Becker.

Columbella confertilirata Barnard, 1959: 182, fig. 35d.

Syntypes: SAM-—A8879. One adult, two juveniles, fifteen fragments.
Locality: off Umkomaas; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Columbella dibolos Barnard, 1964c: 17

= Mitrolumna stepheni (Melvill & Standen): Kilburn, 1977: 204.

Syntypes: SAM-—A29636. Three worn juveniles, one broken.
Locality: off Umhloti River; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Columbella lightfooti Smith, 1901; 112, pl. 1 (fig. 3).

Paratypes: SAM—A6936. Three shells.
Locality: Kalk Bay, False Bay; no depth or date.
Collected by: R. M. Lightfoot.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

‘Columbella’ polyarosus Barnard, 1959: 182, fig. 35c.
Holotype: SAM-—A8875. Shell, broken.
Locality: off Cape Natal; 99 m; 14 December 1900.
Paratype: SAM-A8876. Apex.
Locality: off O’Neil Peak; 165 m; 28 February 1901.
Paratypes: SAM-A8877. Shell, three fragments.
Locality: off Umhloti River; 73 m; 18 December 1900.
Paratype: SAM-A8878. Apex.
Locality: off Hood Point; 90 m; 15 July 1901.
Collected by: R.S. Pieter Faure.

‘Columbella@’ sigma Barnard, 1959: 185, fig. 36b.
Holotype: SAM-A8890. Shell.
Locality: off Cove Rock, East London; 40 m; 6 August 1901.
Collected by: R.S. Pieter Faure.

‘Columbella’ vitula Barnard, 1959: 184, fig. 36a.
Holotype: SAM-A8887. Shell, juvenile.
Locality: off Cove Rock, East London; 80-130 fm.; 30 July 1901.
Collected by: R.S. Pieter Faure.

Pyrene parhelena Barnard, 1959: 177, fig. 36b.
Syntypes: SAM-—A8870. Two shells (one broken), radula slide.
Locality: off Cape St. Blaize; 229 m; 21 December 1899.
Collected by: R.S. Pieter Faure.

Pyrene salmoneus Barnard, 1963b: 7.

Syntypes: SAM-—A29920. One hundred and fifty-six shells, opercula, many
fragments.

Locality: South of Madagascar, 36°48’S 52°08’E; 400 m; 9 July 1961.

Collected by: R.S. Africana IT.

Family Nassariidae

Nassa muiri Barnard, 1959: 105.
Syntypes: SAM-—A29919. Eleven shells.
Locality: uncertain.
Collected by: University of Cape Town.

Nassa rhysonepia Barnard, 1969: 627, fig. 15a.
Holotype: SAM-A29990. Shell, juvenile.
Paratype: SAM-—A30551. Shell, juvenile.
Locality: off Cove Rock, East London; 40 m; 6 August 1901.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 25

Nassa scopularcus Barnard, 1959: 120, fig. 24a.

Holotype: SAM—A9230. Shell.

Locality: Late Tertiary marine beds, lagoon shore, south of Bogenfels,
South West Africa; 1930.

Collected by: S. H. Haughton, University of Cape Town.

Nassa vidalensis Barnard, 1959: 118, fig. 24b.

Holotype: SAM-A8839. Shell.
Locality: off Cape Vidal; 146-183 m; 27 February 1901.
Collected by: R.S. Pieter Faure.

Family Fasciolariidae

Fasciolaria agulhasensis Tomlin, 1932: 157, fig. 1

= Fasciolaria (Pleuroploca) scholveni Strebel: Kilburn, 1974: 204, fig. 13a.
Holotype: SAM-A6539. Shell.
Locality: Agulhas Bank; 28 fm.; 5 January 1904.
Collected by: R.S. Pieter Faure.

Fasciolaria? holcophorus Barnard, 1959: 98, fig. 21.

Holotype: SAM-A8819. Shell.
Locality: off Cape St. Blaize; 228 m; 21 December 1899.
Collected by: R.S. Pieter Faure.

Fusus africanae Barnard, 1959: 96, figs 19k—20d.

Holotype: SAM-A8610. Shell, juvenile with operculum, radula slide.
Locality: Brown’s Bank; 146-183 m; 17 July 1906.

Paratype: SAM-—A8826. Shell, juvenile, radula slide.

Locality: off western Cape Peninsula; 256-276 m; 28 March 1900.
Collected by: R.S. Pieter Faure.

Fusus bonaspei Barnard, 1959: 95, fig. 20c-f.

Holotype: SAM—A4632. Shell with operculum.

Locality: doubtful, wrong Pieter Faure station number recorded but prob-
ably off Cape Point, depth and date unknown.

Paratypes: SAM—A4629. Two shells with opercula.

Locality: off Cape Point, ‘NE $ N 19 miles’; 266 m; 27 February 1902.

Paratype: SAM—A4630. Six shells, one operculum.

Locality: off Cape Hangklip ‘NE # N 28 miles’; 183 m; 27 August 1903.

Collected by: R.S. Pieter Faure.

Fusus faurei Barnard, 1959: 94, figs 19h—-20e.
Syntypes: SAM-—A4581. Nine shells, six opercula, radula slide.
Locality: off Cape Point, ‘N 41° E 38 miles’; 578-730 m; 16 September
1903.
Syntype: SAM-A4582. Shell.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Cape Point, ‘NE 3 N 39 miles’; 580-1 020 m; 17 September
1903.
Collected by: R.S. Pieter Faure.

Latirus mosselensis Tomlin, 1932: 158, fig. 1.
Syntype: SAM—A3504. Shell.
Locality: off Mossel Bay; 49 m; 14 June 1899.
Collected by: R.S. Pieter Faure.

Family Volutidae

Fulgoraria blaizei Barnard, 1959: 28, fig. 8b; 1969: 620, fig. 11

= Fusivoluta blaizei (Barnard, 1959): Weaver, 1963: 5.
Holotype: SAM-A3433. Shell with operculum.
Locality: off Cape St. Blaize, ‘NE # E 73 miles’; 192 m; 21 February 1902.
Paratypes: SAM-—A3430. Shell, adult, 2 juveniles.
Locality: off Cape St. Blaize, ‘N by E 73 miles’; 230 m; 21 December 1889.
Collected by: R.S. Pieter Faure.

Glypteuthria capensis Tomlin, 1932: 165; 1945: 135
= Fusivoluta sculpturata (Tomlin): Weaver & Dupont, 1970: 185.
Holotype: SAM-—A3446. Shell with operculum.
Locality: off Cape Point, ‘N 41° E 38 miles’; 593-730 m; 16 September
1903.
Paratype: SAM-A3448. Shell.
Locality: off Cape Point, ‘NE by E 4 S 28 miles’; 393-733 m; 28 September
1903.
Collected by: R.S. Pieter Faure.

Fusivoluta decussata Barnard, 1959: 31, fig. 8c.
Holotype: SAM-—A3432. Shell.
Locality: off East London, Buffalo River, ‘N 15 miles’; 566 m; 24 April
1901.
Collected by: R.S. Pieter Faure.

Fusivoluta elegans Barnard, 1959: 32, fig. 8a.
Holotype: SAM-A8803. Shell.
Locality: off East London, Buffalo River; 830 m; 17 April 1901.
Collected by: R.S. Pieter Faure.

Volutocorbis boswellae Rehder, 1969: 202, pl. 40 (fig. 5), pl. 41 (figs 16-19).
Paratype: SAM-—A2010. Shell.
Locality: off Cape St. Blaize, ‘N by E 4 E 65 miles’; 155 m; 28 December
1899.
Paratype: SAM—A2011. Shell.

TYPE SPECIMENS OF MARINE MOLLUSCA 25

Locality: off Cape St. Blaize, ‘N by E 4 E 67 miles’; 164 m; 22 December
1899.

Paratype: SAM—A2009. Shell.

Locality: off Cape Seal, ‘N by E # E 37 miles’; 146 m; 20 February 1902.

Paratypes: SAM—A31976 (previously SAM—A3335). Six shells.

Locality: Brown’s Bank, 36°40'S 21°26’E; 146-183 m; 17 July 1906.

Collected by: R.S. Pieter Faure.

Volutocorbis dispirilis Rehder, 1969: 203, pl. 40 (fig. 4), pl. 41.
Holotype: SAM-—A31977 (previously SAM-—A3335). Shell.
Paratype: SAM—A33687 (previously SAM—A3335). Shell.

Locality: Brown’s Bank, 36°40’S 21°26’E; 146-183 m; 17 July 1906.
Collected by: R.S. Pieter Faure.

Volutilithes gilchristi Sowerby, 1902: 99, pl. 2 (fig. 5)

= Volutocorbis gilchristi (Sowerby): Barnard, 1959: 28.
Paratype: SAM-—A1974. Shell.
Locality: off Cape Natal, ‘W by N #N 11 miles’; 189-366 m; 1 April 1901.
Collected by: R.S. Pieter Faure.

Family Marginellidae

Diliculum inopinatum Barnard, 1962a: 14, fig. 1.
Paratypes: SAM—A29917. Three shells, radula slide.
Locality: Saldanha Bay; 3-28 fm.; 5 May 1960.
Collected by: University of Cape Town.

Persicula alborubida Barnard, 1969: 620, fig. 10d.

Holotype: SAM—A30050. Shell.
Locality: Kommetyjie; intertidal; no date.
Collected by: C. Connolly.

Paratype: SAM-—A30550. Shell.
Locality: Kommetyjie; intertidal; no date.
Collected by: C. Connolly.

Persicula nigrocrocea Barnard, 1969: 619, fig. 10b—c.

Holotype: SAM-—A30049. Shell.
Locality: Simonstown; no depth or date.
Collected by: C. Connolly.

Paratype: SAM-A30549. Shell.
Locality: Simonstown; no depth or date.
Collected by: C. Connolly.

Marginella brocktoni Shackleford, 1914: 98, two figures.
Syntypes: SAM-—A1956. Shell with animal, one shell in pieces.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Cape Point, ‘N 50° E 18 miles’; 329 m; 27 February 1902.
Collected by: R.S. Pieter Faure.

Marginella differens eugenes Barnard, 1959: 8; 1969: 619, fig. 10a.

Holotype: SAM-—A8786. Shell.
Locality: off Umkomaas River; 73 m; 31 December 1900.
Collected by: R.S. Pieter Faure.

Marginella kerochuta Shackleford, 1914: 97, two figures.

Syntypes: SAM-—A1954. Two shells.
Locality: off Cape Point, ‘NE 4 N 18 miles’; 244 m; 27 February 1902.
Collected by: R.S. Pieter Faure.

_Marginella taylori Shackleford, 1916: 194, figs 3-4
= Marginella differens Smith: Barnard, 1959: 8.

Syntypes: SAM-—A7191. Two shells, one broken.
Locality: off Cape St. Blaize, ‘N by E 3 E’; 193 m; no date.
Collected by: R.S. Pieter Faure.

Marginella tomlini Shackleford, 1916: 193, figs 1-2
= Marginella bicatenata Sowerby: Barnard, 1959: 6.
Holotype: SAM—A3704. Shell with animal.

Locality: off Cape St. Blaize, ‘N by E 2 E 68 miles’; 193 m; 21 February

1902.
Collected by: R.S. Pieter Faure.

Marginella walvisiana Tomlin, 1920: 88, fig. 3.

Syntypes: SAM—A30666. Three shells.
Locality: Walvis Bay; no depth or date.
Collected by: P. R. Frames.

Family Olividae

Ancilla errorum Tomlin, 1921: 216, pl. 8, fig. 2.
Holotype: SAM-A1811. Shell.

Locality: Walker Point, ‘NE by N 2 N 7 miles’; 47 fm.; 11 October 1900.

Paratype: SAM-A1816. Shell.
Locality: off Cape Point; 42 fm.; 18 December 1900.
Collected by: R.S. Pieter Faure.

Ancilla rouillardi Kilburn, 1981: 381, figs 6, 50, 87, 88, 99, 104-105.
Paratype: SAM-—A1797. Shell.
Locality: ?Cape St. Blaize; 40 fm.; 13 December 1900.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 27

Family Mitridae

Charitodoron aglaia Tomlin, 1932: 169, fig. 9
= Charitodoron agulhasensis (Thiele): Barnard, 1959: 146.

Holotype: SAM-—A30719 (previously A3440). Shell.
Paratype: SAM-—A3440. Shell.

Locality: uncertain, no Pieter Faure station number recorded.
Collected by: R.S. Pieter Faure.

Charitodoron euphrosyne Tomlin, 1932: 167, fig. 8.

Holotype: SAM—A3441. Shell, broken.
Locality: off Cape Point, N 77° E; 1 220-1 275 m; 28 March 1900.
Collected by: R.S. Pieter Faure.

Charitodoron thalia Tomlin, 1932: 169, fig. 10.

Holotype: SAM-—A8602 (previously A1742). Shell, radula slide.
Locality: off Cape Point; 240 m; 28 March 1900.
Collected by: R.S. Pieter Faure.

Mitra (Dibaphus) bathybius Barnard, 1959: 51, figs 11a—13b.

Holotype: SAM—A8827. Shell, broken, radula slide.
Locality: off Cape Natal; 805 m; 4 April 1901.
Collected by: R.S. Pieter Faure.

Mitra (Mitra) boswellae Cate, 1964: 219, pl. 28 (fig. 1a—b)
= Mitra guttata Swainson: Cernohorsky, 1976: 326.

Holotype: SAM-A29799. Shell.
Locality: uncertain; no date.
Collected by: probably commercial trawler.

Mitra fidis Sowerby, 1916: 491, text figure
= Pusia patula (Reeve): Barnard, 1958: 58.

Syntypes: SAM-A29918. Two shells.
Locality: ‘South Africa’; no date.
Donated by: A. E. Salisbury, 1957.

Family Cancellariidae

Admete decapensis Barnard, 1960: 439, fig. 1a.

Holotype: SAM—A9777. Shell, partly corroded.
Locality: off Cape Point, 34°37’S 17°03’E; 3 070-3 150 m; December 1959.
Collected by: R.S. Africana II.

Cancellaria euetrios Barnard, 1959: 14, fig. 3.
Holotype: SAM-—A8747. Shell.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Cape Recife, 34°26’S 25°42'E; 227 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

Cancellaria euthymei Barnard, 1960: 438, fig. 16.

Holotype: SAM—A9888. Shell.
Locality: off Cape Point, 34°26’S 17°00’E; 2 750-3 260 m; December 1959.
Collected by: R.S. Africana IT.

Cancellaria producta Sowerby, 1903: 220, pl. 4 (fig. 5).
Syntype: SAM-A339. Shell.
Locality: off Umhloti River; 73 m; 21 December 1900.
Collected by: R.S. Pieter Faure.

Family Conidae

Conus eucoronatus Sowerby, 1903: 217, pl. 3 (fig. 9).
Holotype: SAM—A1753. Shell.
Locality: off Cape St. Blaize, ‘N 85° W 4,5 miles’; 27 fm.; 15 June 1899.
Collected by: R.S. Pieter Faure.

Conus gilchristi Sowerby 1903: 217, pl. 3 (fig. 8)
= Conus natalis Sowerby: Walls, 1979: 757.
Holotype: SAM—A1752. Shell.
Locality: Umhlangkulu River mouth, ‘NW by N 7,5 miles’; 50 fm.; 14
March 1901.
Collected by: R.S. Pieter Faure.

Conus patens Sowerby, 1903: 218, pl. 3 (fig. 7).
Holotype: SAM—A1756. Shell.
Locality: Vasco de Gama Peak, ‘N 10° E 13 miles’; 85 fm.; 3 May 1900.
Collected by: R.S. Pieter Faure.

Family Terebridae

Terebra (Abretia) lightfooti Smith, 1899: 247, pl. 6 (fig. 1).
Paratype: SAM-5519. Shell.
Locality: Table Bay; 10 fm.; no date.
Collected by: R. M. Lightfoot.

Terebra planecosta Barnard, 1958: 81, fig. 1la—b.

Holotype: SAM—A8659. Shell.

Locality: off Cove Rock, ‘4,5 miles’; 22 fm.; 6 August 1901.

Collected by: R.S. Pieter Faure.

Paratypes: SAM-—A8658. Two shells.

Locality: off Cape St. Blaize, ‘N by E 73 miles’; 125 fm.; 21 December 1899.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 29

Terebra thielei Barnard, 1958: 80, fig. 1d.
Syntypes: SAM-—A8657. Four shells.
Locality: Algoa Bay; 67 fm.; 10 November 1898.
Collected by: R.S. Pieter Faure.

Family Turridae

‘Acrobella’ acus Barnard, 1958: 149, fig. 21e.
Syntypes: SAM-—A8748. Two shells, one fragment.
Locality: off Cape Recife, 34°27’'S 25°42’E; 468 m; 16 November 1898.
Collected by: R.S. Pieter Faure.

‘Bela’ bella Barnard, 1958: 140, fig. 21c.
Syntypes: SAM-A8639. Three shells.
Locality: off Cape Agulhas; 146-183 m; 17 July 1903.
Collected by: R.S. Pieter Faure.

Clavatula lobatopsis Barnard, 1963c: 420, fig. 2a.
Syntypes: SAM-—A9730. Three shells with opercula, radula slide.
Locality: 33°36’S 16°15’E; 2 780-2 880 m; August 1959.
Collected by: R.S. Africana II.

‘Cythara’ dagama Barnard 1963c: 423, fig. 2g—h.
Syntypes: SAM-—A9731. Four shells, three with animals.
Locality: 33°36’S 16°15’E; 2 790-2 890 m; August 1959.
Collected by: R.S. Africana II.

‘Cythara@’ deliciosa Barnard, 1958: 151, fig. 24

= Eucithera deliciosa (Barnard): Barnard, 1969: 610.
Syntypes: SAM-—A8692. Six shells.
Locality: off Umhloti River; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Cythara? glaucocreas Barnard, 1963c: 422, fig. 2d.

Holotype: SAM-—A9821. Shell, radula slide.

Locality: 33°52’'S 16°51'E; 2 430-2 790 m; December 1959.
Paratype: SAM-—A9753. Shell.

Locality: 33°49’'S 16°30’E; 2 750 m; August 1959.
Collected by: R.S. Africana II.

Daphnella? bitrudis Barnard, 1963c: 428, fig. 1b.
Syntypes: SAM-—A9754. Two shells.
Locality: 33°49’S 16°30’E; 2 750 m; August 1959.
Collected by: R.S. Africana IT.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Daphnella recifensis Barnard, 1958: 161, fig. 30a.
Holotype: SAM—A8757. Shell.
Locality: off Cape Recife, 34°27’S 25°42'E; 468 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

Daphnella? verecunda Barnard, 1963c: 427, fig. 2e.

Syntypes: SAM-—A9697. Three shells, one with animal.
Locality: 33°50’S 17°21’E; 1 100 m; August 1959.
Collected by: R.S. Africana IT.

Drillia armilla Barnard, 1958: 137.

Holotype: SAM—A8714. Shell.
Locality: off Cove Rock; 146-239 m; 30 July 1901.
Collected by: R.S. Pieter Faure.

Drillia bruchia Barnard, 1958: 130, fig. 13b.

Holotype: SAM—A8717. Shell, apex missing.
Locality: off Cape Natal; 440 fm.; 4 April 1901.
Collected by: R.S. Pieter Faure.

Drillia collina Barnard, 1958: 129, fig. 13b.

Syntype: SAM-—A83587. Shell.

Locality: off East London; 32 fm.; 1898.
Syntypes: SAM-—A8710. Shell, two fragments.
Locality: off Hood Point; 49 fm.; 1901.
Collected by: R.S. Pieter Faure.

Drillia diasi Barnard, 1958: 137, fig. 19b.
Syntypes: SAM-—A8566. Two shells.
Locality: 34°26’S 25°42'E; 124 fm.; 1898.
Collected by: R.S. Pieter Faure.

Drillia dovyalis Barnard, 1969: 606, fig. 6a.

Holotype: SAM—A29760. Shell.
Locality: off Sandy Point; 93 m; 14 August 1901.
Collected by: R.S. Pieter Faure.

Drillia erepta Barnard, 1969: 609, fig. b.

Syntype: SAM—A8734. Shell.

Locality: off Cape Morgan; 47 fm.; 25 July 1901.

Collected by: R.S. Pieter Faure.

Syntype: SAM-—A29763. Shell.

Locality: 33°50’S 25°54'E (Algoa Bay); ?30 fm.; 11 November 1898.
Collected by: R.S. Pieter Faure.

Syntypes: SAM—A29764. Two shells.

TYPE SPECIMENS OF MARINE MOLLUSCA

Locality: off Cape St. Blaize; 125 fm.; 21 December 1899.
Collected by: R.S. Pieter Faure.

Drillia falcicosta Barnard, 1958: 132, fig. 15b.

Syntypes: SAM-—A8724. Two shells, one apex.
Locality: off Umhloti River; 40 fm.; 1900.
Collected by: R.S. Pieter Faure.

Drillia laterculoides Barnard, 1958: 129, fig. 12c.

Syntypes: SAM-—A8709. Shell, apex.
Locality: off Hood Point; 90 m; 15 July 1901.
Collected by: R.S. Pieter Faure.

Drillia latisulcus Barnard, 1958: 131, fig. 13c.

Syntypes: SAM-—A8718. Five shells.
Locality: off Hood Point; 90 m; 15 July 1901.
Collected by: R.S. Pieter Faure.

Drillia morgana Barnard, 1958: 133, fig. 16.

Holotype: SAM-A8739. Shell.
Locality: off Cape Morgan; 47 fm.; 25 July 1901.
Collected by: R.S. Pieter Faure.

Drillia omia Barnard, 1958: 123, fig. 11a.
Syntypes: SAM-—A8651. Ten shells.

Locality: ?Port Alfred, probably Table Bay—Saldanha Bay.

Collector unknown.

Drillia oneili Barnard, 1958: 133, fig. 15c.
Holotype: SAM—A8731. Shell, slightly worn.
Locality: off O’Neil Peak; 90 fm.; 1901.
Collected by: R.S. Pieter Faure.

Drillia pecus Barnard, 1969: 607, fig. Sb.
Holotype: SAM—A29761. Shell.
Locality: off East London; 400-450 fm.; 17 April 1901.
Collected by: R.S. Pieter Faure.

Drillia perfluans Barnard, 1958: 132, fig. 15a.
Holotype: SAM-—A8716. Shell.
Locality: off Hood Point; 49 fm.; 15 July 1901.
Collected by: R.S. Pieter Faure.

Drillia pleonastica Barnard, 1958: 133, fig. 17a.
Holotype: SAM-—A8565. Shell.
Locality: 34°26’'S 25°42'E; 228 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

31

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

Drillia pselia Barnard, 1958: 138.
Syntypes: SAM-—A8715. Shell, two broken shells.
Locality: off Cape St. Blaize; 125 fm.; 1899.
Collected by: R.S. Pieter Faure.

Drillia simplicingula Barnard, 1958: 135, fig. 18b.
Holotype: SAM—A8666. Shell.
Locality: off Hood Point; 49 fm.; 15 July 1901.
Paratype: SAM-A3471. Shell.
Locality: off Cape Morgan; 104 m; 6 February 1901.
Collected by: R.S. Pieter Faure.

Drillia spiralis Barnard, 1958: 134, fig. 17b.

Holotype: SAM-A8583. Shell.
Locality: off Cape St. Blaize; 228 m; 21 December 1899.
Collected by: R.S. Pieter Faure.

Drillia tholos Barnard, 1958: 130, fig. 14.
Syntypes: SAM-—A8735. Fourteen shells.
Locality: off Hood Point; 49 fm.; 15 July 1901.
Collected by: R.S. Pieter Faure.

Mangilia (Eucythara) africana Sowerby, 1903: 21, pl. 5 (fig. 9)
= Cythara africana (Sowerby): Barnard, 1958: 115.
Holotype: SAM—A345. Shell.
Paratype: SAM—A33954. Shell, broken, outer lip thin.
Locality: off Umhloti River; 46 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Mangilia extans Barnard, 1958: 156, fig. 27c.

Holotype: SAM—A8756. Shell.
Locality: off Cove Rock; 40 m; 6 August 1901.
Collected by: R.S. Pieter Faure.

Mangilia muiri Barnard, 1958: 153, fig. 26a.

Syntypes: SAM-—A8647. Seven shells, in growth series.
Locality: Still Bay; intertidal; 1921.
Muir Collection.

Mangilia phoxos Barnard, 1958: 155, fig. 27b.
Holotype: SAM-—A8730. Shell.
Locality: off Umhloti River; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

(2
(4)

TYPE SPECIMENS OF MARINE MOLLUSCA

Mangilia sciola Barnard, 1958: 157, fig. 28c.
Syntype: SAM-A8642. Shell.
Locality: off Cape Recife, 34°27'S 25°42'E; 468 m; 14 November 1898.
Syntype: SAM-A8643. Shell.
Locality: near East London, 33°06’S 27°55'E; 78 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

Mangilia serrula Barnard, 1964c: 15, fig. 1a—b.
Syntypes: SAM-—A8728. Five shells.
Locality: off Illovo and Umkomaas; 30-40 fm.; 31 December 1900.
Collected by: R.S. Pieter Faure.

Mangilia tranquilla Barnard, 1958: 158, fig. 28b.
Syntypes: SAM-A8648. Five shells.
Locality: Stull Bay; intertidal; no date.
Collected by: J. Muir.

Mangilia translucens Barnard, 1958: 155, fig. 27a.
Syntypes: SAM-—A8586. Twelve shells.
Locality: off Umhloti River mouth; 40 fm.; 28 December 1898.
Collected by: R.S. Pieter Faure.

Mitramorpha veneris Barnard, 1964c: 16.
Holotype: SAM—A8750. Shell, broken.
Locality: off Cape Vidal; 146-183 m; 27 February 1901.
Collected by: R.S. Pieter Faure.

Monilopsis psilarosis Barnard, 1963c: 421, fig. 4a.
Syntypes: SAM-—A9789. Two shells, radula slide.
Locality: 34°42'S 16°54’E; 3 140-3 270 m; December 1959.
Collected by: R.S. Africana II.

Philbertia natalensis Barnard, 1958: 158, fig. 29a.
Syntype: SAM—A8654. Shell.
Locality: off Umhloti River mouth; 40 fm.; 18 December 1900.
Collected by: R.S. Pieter Faure.
Syntype: SAM-—A8758. Shell.
Locality: off Cape Natal, 54 fm.; 14 December 1900.
Collected by: R.S. Pieter Faure.

Pleurotoma (Genotia) belaeformis Sowerby, 1903: 216, pl. 4 (fig. 8)
= *‘Genotia belaeformis (Sowerby): Barnard, 1958: 112, figs 8b—9b.
Paratypes: SAM—A1687. Two shells.
Locality: off Cape Point; 230 fm.; 4 May 1900.
Collected by: R.S. Pieter Faure.

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

‘Pleurotoma’ curricula Barnard, 1969: 612, fig. 8.
Syntypes: SAM-—A9859. Shell, two broken.
Locality: 34°36’S 17°00’E; 2 750-3 200 m; December 1959.
Collected by: R.S. Africana IT.

Pleurotoma (Drillia) fossata Sowerby, 1903: 214, pl. 3 (fig. 5)
= Drillia fossata (Sowerby): Barnard, 1958: 135.
Holotype: SAM—A346. Shell.
Paratype: SAM-A346. Shell.
Locality: off Cape Vidal; 80-100 fm.; 27 February 1901.
Collected by: R.S. Pieter Faure.

Pleurotoma (Clavus) lignaria Sowerby, 1903: 215, pl. 3 (fig. 4)
_= Turris lignaria (Sowerby): Barnard, 1958: 102.

Paratype: SAM-A357. Shell.

Locality: off Table Bay; 240 m; 28 March 1900.
Collected by: R.S. Pieter Faure.

Pleurotoma (Surcula) lobata Sowerby, 1903: 213, pl. 4 (fig. 9)

= Turris lobata (Sowerby): Barnard, 1958: 107, fig. 3i-j.
Syntypes: SAM-—A1673. Five shells, radula slide.
Locality: off Cape Natal; 810 m; 4 April 1901.
Syntypes: SAM-A1674. Three shells, one with operculum.
Locality: off Buffalo River, East London; 560 m; 24 April 1901.
Collected by: R.S. Pieter Faure.

Pleurotoma (Drillia) scitecostata Sowerby, 1903: 214, pl. 4 (fig. 10)
= ‘Drillia’ scitecostata (Sowerby): Barnard, 1958: 119, figs 8f, 20.
Syntypes: SAM-A347. Two shells.
Locality: off Glendower Beacon; 100 fm.; 10 September 1901.
Collected by: R.S. Pieter Faure.

Pleurotoma (Clavatula) turriplana Sowerby, 1903: 215, pl. 3 (fig. 6)
= Clavatula turriplana Sowerby: Barnard, 1958: 144.
Holotype: SAM-A1646. Shell.
Locality: off Cape St. Blaize, ‘N by E 4 E 65 miles’; 155 m; 22 December
1899.
Collected by: R.S. Pieter Faure.

Surcula aditus Barnard, 1969: 604, fig. 4a.
Holotype: SAM—A29759. Shell.
Locality: off Cove Rock; 22 fm.; 6 August 1901.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA 35

Surcula amplisulcus Barnard, 1958: 147, fig. 22a.
Syntypes: SAM-A16839. Five shells.
Locality: west coast of Cape Peninsula; 120 fm.; 4 April 1902.
Collected by: R.S. Pieter Faure.

Surcula faurei Barnard, 1958: 147, fig. 22b.
Holotype: SAM-—A8611. Shell.
Locality: Brown’s Bank, 36,5°S 21°E; 80-100 fm.; 17 July 1906.
Collected by: R.S. Pieter Faure.

Surcula scalaria Barnard, 1958: 146, fig. 22d.

Syntypes: SAM-—A358. Seven shells.
Locality: off Cape Point; 480-800 fm.; 10 September 1903.
Collected by: R.S. Pieter Faure.

Surcula sulcicancellata Barnard, 1958: 145, fig. 22c.

Syntypes: SAM-—A361. Five shells.
Locality: off Cape Point; 130-300 fm.; 28 September 1903.
Collected by: R.S. Pieter Faure.

Turris ambages Barnard, 1958: 148, fig. 23b.

Syntypes: SAM—A8683. Two shells.

Locality: off Cape Natal; 54 fm.; 14 December 1900.
Collected by: R.S. Pieter Faure.

Syntype: SAM-A8684. Shell.

Locality: off Umkomaas River; 40 fm.; 31 December 1900.
Collected by: R.S. Pieter Faure.

Syntype: SAM-A8685. Shell.

Locality: off O’Neil Peak; 90 fm.; 28 February 1901.
Collected by: R.S. Pieter Faure.

Turris saldanhae Barnard, 1958: 109, figs 3f, 7.
Syntypes: SAM—A1738. Seventeen shells.
Locality: Baboon Point, Saldanha Bay; 31 fm.; 18 March 1902.
Collected by: R.S. Pieter Faure.

Typhlomangilia? polythele Barnard, 1963c: 422, fig. 3e-f.
Syntypes: SAM-—A9802. Two shells.
Locality: 33°50’'S 16°30’'E; 1 480-1 660 fm.; December 1950.
Collected by: F. H. Talbot.

Typhlosyrinx chrysopelex Barnard, 1963c: 425, fig. 3g.

Holotype: SAM-—A9857. Shell.
Locality: 34°36'S 17°00’E; 1 500-1 760 fm.; December 1959.
Collected by: F. H. Talbot.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Typhlosyrinx pyrropelex Barnard, 1963c: 423, fig. 2c.

Syntypes: SAM—A1643. Two shells.

Locality: off Cape Point, N 77° E; 650-700 fm.; 29 July 1903.
Collected by: R. S. Pieter Faure.

Syntypes: SAM-—A9858. Three shells, juveniles with animals.
Locality: 34°36’S 17°00’E; 1 500-1 760 fm.; December 1959.
Collected by: F. H. Talbot.

Typhlosyrinx subrosea Barnard, 1963c: 425, fig. 3a—d.
Holotype: SAM—A9822. Shell with animal.
Locality: 33°52’S 16°51’E; 1 380-1 520 fm.; December 1959.
Collected by: F. H. Talbot.
Paratype: SAM-A9822. Shell.
Locality: 33°52’S 16°51'E; 1 380-1 520 fm.; December 1959.
Collected by: F. H. Talbot.

Subclass OPISTHOBRANCHIA

Family Acteonidae

Solidula niecaensis Barnard, 1963d: 317
= Pupa niecaensis (Barnard): comb. nov.

Holotype: SAM—A6553. Shell.
Locality: off Nieca River; 80 m; 7 August 1901.
Collected by: R.S. Pieter Faure.

Family Retusidae

Retusa natalensis Barnard, 1963d: 321, fig. 28a.

Syntypes: SAM—A9409. Two shells.
Locality: off Umkomaas River; 40 fm.; 1900.
Collected by: R.S. Pieter Faure.

Volvula mutabilis Barnard, 1963d: 322, fig. 28b.

Syntypes: SAM-A9410. Two shells.
Locality: off Umhloti River; 40 fm.; 1900.
Collected by: R.S. Pieter Faure.

Family Scaphandridae

Acteocina natalensis Barnard, 1963d: 324, fig. 28c.
Syntypes: SAM—A9388. Two shells.
Locality: off Illovo River; 27-30 fm.; 1900.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA Si

Family Doridoxidae

Doridoxa benthalis Barnard, 1963c: 443, fig. 10.

Holotype: SAM—A9796. Animal, preserved, jaws, radula slide.
Locality: 34°42'S 16°54'E; 1 725-1 780 fm.; December 1959.
Collected by: F. H. Talbot.

Family Chromodorididae

Glossodoris capensis Barnard, 1927: 181, pl. 19 (figs 3-4)
= Hypselodoris capensis (Barnard): comb. nov.

Syntypes: SAM-A6230. Two animals, preserved, two pairs jaws, two
radula slides.

Locality: St. James, False Bay; intertidal; April 1912.

Collected by: K. H. Barnard.

Family Polyceridae

Polycera nigrocrocea Barnard, 1927: 191, pl. 19 (figs 7-8)
=Polycera capensis Quoy & Gaimard: Pruvot-Fol, 1934: 75.

Syntype: SAM-—A6225. Two animals, preserved, radula.
Locality: St. James; intertidal; March 1912.
Collected by: K. H. Barnard.

Plocamopherus apheles Barnard, 1927: 195, fig. 1.
Holotype: SAM—A6232. Animal, preserved, jaws, radula slide.
Locality: Nahoon estuary, East London; no depth; 31 May 1901.
Collected by: J. D. F. Gilchrist.

Family Goniodorididae

Goniodoris ovata Barnard, 1934: 2, 2 text figures.
Holotype: SAM-A6270. Animal, preserved.
Paratype: SAM-—A6270. Radula slide.

Locality: Simonstown; from boat hull; June 1933.
Collected by: K. H. Barnard.

Family Janolidae

Bonisa nakaza Gosliner, 1981: 19, figs 1d-f, 18-25.
Holotype: SAM—A34886. Animal, preserved.
Paratypes: SAM-—A34887. Four animals, preserved.
Locality: off Llandudno, 34°01'S 18°20’E; 20 m; 23 December 1979.
Collected by: W. R. Liltved.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Janolus longidentatus Gosliner, 1981: 9, figs 1c, 10-17.

Holotype: SAM-—A34883. Animal, preserved.

Locality: Miller’s Point, 34°14’S 18°29’E; 3 m; 10 December 1980.
Collected by: T. M. Gosliner.

Paratypes: SAM-—A34884. Two animals, preserved.

Locality: Castle Rocks, 34°18’S 18°29’E; 10 m; 1 October 1980.
Collected by: T. M. Gosliner.

Paratype: SAM-A34885. Animal, preserved.

Locality: Miller’s Point, 34°14’S 18°29’E; 5 m; 11 July 1980.
Collected by: T. M. Gosliner.

Family Flabellinidae

_Flabellina funeka Gosliner & Griffiths, 1981: 106, figs 1a, 2-4.
Holotype: SAM-A34317. Animal, preserved.
Locality: Castle Rocks, 34°14’S 18°29’E; 10 m; 17 January 1980.
Collected by: T. M. Gosliner.
Paratype: SAM-—A34318. Animal, preserved.
Locality: Castle Rocks, 34°14’S 18°29’E; 10 m; 17 January 1980.
Collected by: T. M. Gosliner.
Paratype: SAM—A34319. Animal, preserved.
Locality: Venus Pool, 34°17’S 18°28’E; 10 m; 6 January 1980.
Collected by: T. M. Gosliner.
Paratype: SAM—A34320. Animal, preserved.
Locality: Hermanus, 34°17’S 19°15’E; 17 m; 11 October 1971.
Collected by: R. J. Griffiths.
Paratype: SAM—A34321. Animal, preserved.
Locality: Hermanus, 34°17'S 19°15’E; 17 m; 11 October 1971.
Collected by: R. J. Griffiths.

Family Tergipedidae

Catriona casha Gosliner & Griffiths, 1981: 130, figs 12-14.

Holotype: SAM-—A34871. Animal, preserved.

Paratypes: SAM—A34872. Five animals, preserved.

Locality: Cape Town docks, 33°54’S 18°26'E; 1 m; 26 June 1972.
Collected by: R. J. Griffiths.

Family Facelinidae

Facelina faurei Barnard, 1927: 205, figs. 4-5
= Pruvotfolia faurei (Barnard): Tardy, 1969: 343.
Syntypes: SAM-A6249. Several animals, preserved, jaws, radula slide.

TYPE SPECIMENS OF MARINE MOLLUSCA

Locality: off Saldanha Bay; 33 fm.; 12 March 1902.
Collected by: R.S. Pieter Faure.

Favorinus horridus Macnae, 1954: 19, pl. 1 (fig. 4) figs 11-13
= Phyllodesmium serratum (Baba): Baba, 1955: 56.

Paratypes: SAM-—A29578. Fifteen animals, preserved.
Locality: ‘2 mile S of Kalk Bay Harbour’; intertidal; 1949.
Collected by: W. Macnae.

Hervia quadricolor Barnard, 1927: 203, pl. 20 (figs 9-10)
= Godiva quadricolor (Barnard): Macnae, 1954: 23.

Holotype: SAM—A6248. Animal, preserved.
Locality: St. James (False Bay); intertidal; 20 June 1912.
Collected by: K. H. Barnard.

Family Cratenidae

Cratena capensis Barnard, 1927: 204, pl. 20 (fig. 8).

Syntype: SAM-—A6247. Jaws, radula slide.
Locality: Sea Point; intertidal; 29 November 1913.
Collected by: K. H. Barnard.

Family Aeolidiidae

Aeolidiella saldanhensis Barnard, 1927: 201, figs 2-3
= Aeolidiella indica Bergh: Gosliner & Griffiths, 1981: 119.

Syntype: SAM-—A6257. Jaws, radula slide.
Locality: Saldanha Bay; intertidal; 5 September 1912.
Collected by: R.S. Pieter Faure.

Subclass PULMONATA

Family Siphonariidae

Siphonaria (Patellopsis) anneae Tomlin, 1944: 92, text figure.
Syntypes: SAM-—A2988. Two shells.
Locality: Umhlali; no depth or date.
Collected by: H. C. Burnup.

Siphonaria cyaneomaculata Sowerby, 1906: 37, text figure

= Siphonaria (Patellopsis) deflexa (Helbling): Allanson, 1958: 159.
Syntypes: SAM-—A3330. Two shells.
Locality: Kowie, Port Alfred; intertidal; 1915.
Collected by: H. Becker.

39

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Siphonaria (Patellopsis) dayi Allanson, 1958: 169, pl. 16 (figs 10-15), text
figure.

Holotype: SAM-—A32707. Shell.

Locality: Delagoa Bay; no depth or date.

Collected by: University of Cape Town.

Paratypes: SAM—A33403. Three shells.

Locality: Inhaca Island; no depth; 30 December 1958.
Collected by: University of Cape Town.

Siphonaria (Siphonaria) aspera pallida Allanson, 1958: 171, pl. 4, fig. 17.
Syntypes: SAM-—A33402. Four shells.
Locality: Langebaan Lagoon; intertidal; 25 April 1949.
Collected by: University of Cape Town.

Class SCHAPHOPODA

Family Dentaliidae

Dentalium africanum Sowerby, 1903: 224, pl. 5 (fig. 10).
Paratypes: SAM—A54839. Six shells, radula slide.
Locality: off Umtwalumi River; 25 fm.; 22 March 1901.
Collected by: R.S. Pieter Faure.

Dentalium (Compressidens) capense Tomlin, 1931: 340.
Holotype: SAM—A6191. Shell.
Locality: off Cape Point; 900 fm.; 19 August 1903.
Collected by: R.S. Pieter Faure.

Dentalium engischistum Barnard, 1963d: 352, fig. 30f.
Syntypes: SAM-A5463. Six shells, two with animals, one broken, radula
slide.
Locality: off Cape Natal; 62 fm.; 1900.
Collected by: R.S. Pieter Faure.

Dentalium eualdes Barnard, 1963c: 444.
Syntypes: SAM-—A9736. Two shells, radula slide.
Locality: 33°36’S 16°15’E; 2 780-2 880 m; August 1959.
Collected by: R.S. Africana II.

Dentalium lardum Barnard, 1963c: 445.
Syntypes: SAM-A9768. Three shells.
Locality: 33°39’S 16°30’E; 1 500 fm.; August 1959.
Collected by: R.S. Africana II.

TYPE SPECIMENS OF MARINE MOLLUSCA 41

Dentalium natalense Barnard, 1963d: 350, fig. 30e.
Syntypes: SAM—A9364. Forty-nine shells.
Locality: off Cape Natal; 85 fm.; 17 December 1900.
Collected by: R.S. Pieter Faure.

Dentalium (Fissidentalium) platypleurum Tomlin, 1931: 339, text figure.
Holotype: SAM-—A3631. Shell.
Locality: off Itongazi River; 25 fm.; 14 March 1901.
Collected by: R.S. Pieter Faure.

Dentalium salpinx Tomlin, 1931: 338, text figure.

Holotype: SAM-A5459. Shell.

Paratypes: SAM-—A5459. Nine shells.

Locality: off Cape Point, ‘NE # E, 40 miles’; 700-800 fm.; 15 September 1903.
Collected by: R.S. Pieter Faure.

Dentalium subterlineatum Tomlin, 1931: 337.

Holotype: SAM-A6192. Shell.
Locality: off Cape Point; 900 fm.; 19 August 1903.
Collected by: R.S. Pieter Faure.

Family Cadulidae

Cadulus promontorii Barnard, 1963d: 353, fig. 30h-1.

Syntypes: SAM-—A7460. Four shells, radula slide.
Locality: off Cape Point; 700 fm.; 20 August 1903.
Collected by: R.S. Pieter Faure.

Class BIVALVIA

Family Nuculidae

Nucula aequalitas Barnard, 1964a: 365.

Holotype: SAM—A9472. Two valves.
Locality: off East London; 732-915 m; 17 April 1901.
Collected by: R.S. Pieter Faure.

Nucula fragilis Boshoff, 1968: 95, pl. 8a, text fig. 1.

Syntypes: SAM—A30212. Two valves.
Locality: 29°34’S 31°39’E; 115-118 m; 9 September 1964.
Collected by: R.V. Anton Bruun.

Nucula irregularis Sowerby, 1904: 7, pl. 6 (fig. 12).
Paratypes: SAM-—14843. Two valves.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Struis Point; 28 m; 17 July 1902.
Collected by: R.S. Pieter Faure.

Nucula sculpturata Sowerby, 1904: 7, pl. 6 (fig. 11)
= Nucula pulchra Hinds: Barnard, 1964a: 362.

Paratypes: SAM-—14830. Ten valves, three with dry animals.
Locality: 33°03’S 27°57'E; 62 m; 28 December 1898.
Collected by: R.S. Pieter Faure.

Family Malletiidae

Malletia estheriopsis Barnard, 1963c: 447, fig. 11b.

Syntypes: SAM-A9817. Four valves.
Locality: 33°50’S 16°30’E; 2 700-3 040 m; December 1959.
Collected by: F. H. Talbot on R.S. Africana IT.

Family Nuculanidae

Leda lanceta Boshoff, 1968: 96, pl. 8b—c.

Holotype: SAM—A30213. Two valves.
Locality: 29°21’S 31°58’E; 370 m; no date.
Collected by: R.V. Anton Bruun.

Leda macella Barnard, 1963c: 448, fig. 11d

= Nuculana vestita (Locard): Knudsen, 1970: 28, fig. 11.

Syntypes: SAM-—A9851. Sixty-six complete shells, nine valves.
Locality: 34°05’S 16°58’E; 2 690-2 720 m; December 1959.
Collected by: F. H. Talbot on R.S. Africana IT.

Leda parceplicata Barnard, 1964b: 21

= Portlandia (Portlandia) parceplicata (Barnard): Kilburn, 1973b: 698.
Syntypes: SAM-—A9465. Thirty-nine complete shells, 157 valves.
Locality: off Umhloti River, Natal; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Leda parsimonia Barnard, 1963c: 448.
Holotype: SAM-—A9818. Two valves.
Locality: 33°50’S 16°30’E; 2 700-3 040 m; August 1959.
Collected by: F. H. Talbot on R.S. Africana IT.

Nuculana compta Sowerby, 1904: 6, pl. 6 (fig. 10)
= Leda compta (Sowerby): Barnard, 1964a: 367, fig. 1a.

Paratype: SAM-—14818. Two valves.
Locality: off Cape Natal; 805 m; 4 April 1901.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA

Nuculana gemmulata Sowerby, 1904: 6, pl. 6 (fig. 9)

= Leda gemmulata (Sowerby): Barnard 1964a: 366.
Paratypes: SAM-14785. Four valves.
Locality: off Tugela River; 68 m; 29 January 1901.
Collected by: R.S. Pieter Faure.

Nuculana lamellata Sowerby, 1904: 5, pl. 6 (fig. 8)

= Leda lamellata (Sowerby): Barnard, 1964a: 366.
Paratypes: SAM-14787. Sixteen complete shells, thirteen valves.
Locality: off Cape Natal; 99 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Sarepta natalensis Barnard, 1964c: 22, fig. 4a.
Holotype: SAM-A9474. Valve.
Locality: off Cape Natal; 90 m; 14 December 1900.
Paratypes: SAM—A9470. Seven valves.
Locality: off Tugela River, Natal; 110-146 m; 11 January 1901.
Collected by: R.S. Pieter Faure.

Family Arcidae

Arca (Scapharca) africana Sowerby, 1904: 4, pl. 6 (fig. 4).
Paratypes: SAM-—14831. Complete shell with animal, two valves.
Locality: off Tugela River; 84-100 m; 6 February 1901.
Collected by: R.S. Pieter Faure.

Deltaodon tugelae Barnard, 1962b: 249, figs 2a—e. Kilburn, 1973b: 699.

Lectotype: SAM—A31736. One valve.
Paralectotypes: SAM-A9477. Eighteen valves.
Locality: off Zululand; no depth or date.
Collected by: R.S. Pieter Faure.

Family Limopsidae

Limopsis natalis Barnard, 1964b: 23, figs.

Syntypes: SAM-—A9479. Complete shell, two valves.
Locality: off O’Neil Peak, Natal; 103 m; 28 February 1901.
Collected by: R.S. Pieter Faure.

Family Mytilidae

Crenella striatissima Sowerby, 1904: 3, pl. 6 (fig. 1). Barnard 1964a: 403.
Holotype: SAM-14855. Two valves.
Locality: 37°07’S 25°40’E; 106 m; 14 November 1898.
Paratype: SAM-14856. Valve, broken.

43

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: off Umhloti River, Natal; 183 m; 19 December 1900.
Collected by: R.S. Pieter Faure.

Gregariella simplicifilis Barnard, 1964a: 402.

Syntypes: SAM-6878. Five complete shells, animals in three, five valves.
Locality: Gordon’s Bay, False Bay; intertidal; no date.
Collected by: R. M. Lightfoot.

Musculus (Musculus) virgiliae Barnard, 1964a: 399, fig. 8d.

Syntypes: SAM-—A7776. Six complete shells, two valves, one broken.
Locality: Keurbooms River, eastern Cape; intertidal; 1931.
Collected by: K. H. Barnard.

Septifer bisculpturata Barnard, 1964c: 23, fig. 4c—d.

Holotype: SAM-A9482. Valve.

Locality: off Tugela River, Natal; 86 m; 29 January 1901.
Collected by: R.S. Pieter Faure.

Family Pectinidae

Chlamys fultoni Sowerby, 1904: 2, pl. 6 (fig. 5).
Paratype: SAM-14970. Valve.
Locality: off Amatikulu River, Natal; 48 m; 1 February 1901.
Collected by: R.S. Pieter Faure.

Chlamys gilchristi Sowerby, 1904: 1, pl. 6 (fig. 6). Barnard 1964a: 427.
Holotype: SAM-14853. Shell, complete.
Locality: False Bay; 416 m; 4 May 1900.
Collected by: R.S. Pieter Faure.

Chlamys humilis Sowerby, 1904: 3, pl. 6 (fig. 3)

= Chlamys tinctus (Reeve): Barnard, 1964a: 425.
Paratype: SAM-14858. Shell, complete.
Locality: off Cape St. Blaize, Mossel Bay; 167-183 m; 20 December 1900.
Collected by: R.S. Pieter Faure.

Cyclopecten incubans Barnard, 1964a: 432.

Syntypes: SAM-—A9493. Twelve valves.
Locality: off Cape St. Blaize; 230 m; 21 December 1899.
Collected by: R.S. Pieter Faure.

Cyclopecten vidalensis Barnard, 1964a: 433, fig. 14e-g.
Holotype: SAM-—A9495. Valve.
Locality: off Cape Vidal, Zululand; 146-183 m; 27 February 1901.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA

Family Limidae
Lima abscisa Barnard, 1964a: 441, fig. 16f.
Syntypes: SAM-—A9504. Three valves.
Locality: off Cape Morgan, near East London; 86 m; 25 July 1901.
Collected by: R.S. Pieter Faure.
Lima divericata Barnard, 1964a: 440, fig. 16e.
Syntypes: SAM-A9501. Two valves.
Locality: off O’Neil Peak, Zululand; 165 m; 28 February 1901.
Collected by: R.S. Pieter Faure.
Lima symmetrica Barnard, 1964a: 441.
Holotype: SAM-A9503. Valve.

Locality: off Cape Vidal; 146-183 m; 27 February 1901.
Collected by: R.S. Pieter Faure.

Family Astartidae

Astarte (Digitaria) lunulata Barnard, 1964a: 449, fig. 18b.
Syntypes: SAM-—A9506. Three valves.

Locality: 34°26'S 25°42’'E; 226 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

Family Crassatellidae

Crassatella natalensis Barnard, 1964a: 457, fig. 19b.

Syntypes: SAM—A9509. Four valves, one juvenile.
Locality: off Cape Natal; 98 m; 14 December 1900.

Collected by: R.S. Pieter Faure.
Crassatella pilula Barnard, 1964a: 457, fig. 19c.

Syntypes: SAM-—A3665. Three complete shells, five valves.

Locality: off Cape Morgan, N of East London; 144 m; 26 July 1901.
Collected by: R.S. Pieter Faure.

Family Carditidae

Cardita pulcherrima Sowerby, 1904: 7, pl. 6 (fig. 2). Barnard, 1964a: 459.
Paratypes: SAM-A9512. Nine valves.

Locality: off Cape Natal; 98 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Venericardia fortisculpta Barnard, 1964a: 462.
Syntypes: SAM-—A9519. Four valves.

45

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: Algoa Bay; 95 m; 14 November 1898.
Collected by: R.S. Pieter Faure.

Venericardia nuculoides Barnard, 1964a: 462.

Syntypes: SAM-A29661. Twenty-two complete shells, eight valves.
Locality: 35°05’S 18°17'E; 27 m; 2 July 1961.
Collected by: University of Cape Town.

Family Lucinidae

Phacoides peritaphros Barnard, 1964a: 476, fig. 22.

Syntypes: SAM-—A9524. Five valves.
Locality: off Nieca River, East London; 43 fm.; 7 August 1901.
Collected by: R.S. Pieter Faure.

Phacoides saldanhae Barnard, 1964a: 474, fig. 21c.

Syntypes: SAM-—A4473. Four valves.
Locality: off Baboon Point, Saldanha Bay; 56 m; 18 March 1902.
Collected by: R.S. Pieter Faure.

Phacoides sepes Barnard, 1964c: 25.

Syntypes: SAM-A9529. Two valves.
Locality: off Morewood Cove, Natal; 49 m; 19 December 1900.
Collected by: R.S. Pieter Faure.

Phacoides sudes Barnard, 1964c: 25, fig. 6a—b
= Gonimyrtea sudes (Barnard): Kilburn, 1973b: 701.

Syntypes: SAM—A9531. Six valves.
Locality: off Cape Natal; 98 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Family Laternulidae

Periploma fractura Boshoff, 1968: 97, pl. 8d, text-fig. 2.
Holotype: SAM-—A30211. Shell, complete.
Locality: 29°29’S 30°41’'E; 86 m; 9 September 1969.
Collected by: R.V. Anton Bruun.

Family Thyasiridae

Thyasira unilateralis Barnard, 1964c: 24, fig. 5a.
Syntypes: SAM—A9520. Two valves.
Locality: off O’Neil Peak, Natal; 165 m; 28 February 1901.
Collected by: R.S. Pieter Faure.

TYPE SPECIMENS OF MARINE MOLLUSCA

Family Erycinidae

Tellimya biradialis Barnard, 1964a: 484, fig. 23a.

Syntypes: SAM-—A9535. Two valves.
Locality: off Cape Natal; 155 m; 17 December 1900.
Collected by: R.S. Pieter Faure.

Tellimya trigona Barnard, 1964a: 484, fig. 23b.

Syntypes: SAM-A29741. Five valves.
Locality: Langebaan (Saldanha Bay); no depth; 26 April 1949.
Collected by: University of Cape Town.

Family Montacutidae

Conchentopyx granulosa Barnard, 1964b: 35, fig. 2.

?Holotype: SAM—A33071. Animal, preserved.
Locality: Langebaan (Saldanha Bay); no depth or date.
Collected by: University of Cape Town.

Montacula ornata Barnard, 1964b: 26, fig. 6c—d
= Barrimysia (Callomysia) ornata (Barnard): Kilburn, 1973b: 702.

Syntypes: SAM—A9538. Three valves.
Locality: off Umhloti River; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Montacuta siliqua Barnard, 1964b: 26, fig. 6e
= Nippon mysella (Barnard): Kilburn, 1973b: 702.

Syntypes: SAM—A9539. Three valves.
Locality: off Umhloti River; 73 m; 18 December 1900.
Collected by: R.S. Pieter Faure.

Family Veneridae

Venus (Anaitis) intersculpta Sowerby, 1904: 11, pl. 7 (fig. 2)
= Venus verrucosa Linnaeus: Barnard, 1964a: 496.

Paratype: SAM-14841. Shell, complete.
Locality: Algoa Bay; 18-29 m; 15 March 1899.
Collected by: R.S. Pieter Faure.

Family Mesodesmatidae

Donacilla delagoae Barnard, 1964a: 515, fig. 28b.
Syntypes: SAM-A29742. Two complete shells, juvenile, ten valves.
Locality: Inhaca Island, Delagoa Bay; no depth or date.
Collected by: University of the Witwatersrand.

47

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Tellinidae

Tellina acropisthus Barnard, 1964b: 26. fig. 5b
= Tellina (Tellinella) staurella Lamarck: Boss, 1969: 95, pl. 2 (fig. 2), pl. 3 (figs
1-2), pl. 4 (fig. 2).
Syntype: SAM-A9547. Valve.
Locality: off Cape Natal; 88 m; 17 December 1900.
Collected by: R.S. Pieter Faure.

Tellina europisthus Barnard, 1964b: 27, fig. c
= Tellina (Cadella) semen Hanley: Boss, 1969: 136.

Syntypes: SAM-—A9549. Four valves.
Locality: off Cape Natal; 98 m; 14 December 1900.
Collected by: R.S. Pieter Faure.

Tellina gilchristi Sowerby, 1904: 12, pl. 7 (fig. 3). Barnard, 1964a: 540
= Tellina (Moerella) gilchristi Sowerby: Boss, 1969: 144.

Paratypes: SAM-—14751. Six valves.
Locality: off Cape Point; 92 m; 6 June 1900.
Collected by: R.S. Pieter Faure.

Tellina (Macoma) inclinata Sowerby, 1904: 14, pl. 7 (fig. 9)
= Macoma inclinata (Sowerby): Barnard 1964a: 549, fig. 311.

Syntypes: SAM-—14835. Six valves.
Locality: off Tugela River; 84-92 m; 6 February 1901.
Collected by: R.S. Pieter Faure.

Tellina (Macoma) levior Sowerby, 1904: 13, pl. 7 (fig. 6)
= Macoma levior (Sowerby): Barnard, 1964a: 548.
Paratype: SAM-—14797. Four complete shells, 8 valves.
Locality: Tugela River, ‘N by W 4 miles’; 24 fm.; 21 January 1901.
Collected by: R.S. Pieter Faure.
Paratype: SAM-14798. Six complete shells, 2 valves.
Locality: Amatakulu River, NW by W ? W 12 miles; 26 fm.; 7 February
1901.
Collected by: R.S. Pieter Faure.

Tellina (Macoma) ordinaria Sowerby, 1904: 14, pl. 7 (fig. 7)
= Macoma ordinaria (Sowerby): Barnard, 1964a: 547, fig. 31e.

Paratypes: SAM-14824. Three valves.
Locality: off Saldanha Bay; 18-26 m; 19 March 1902.
Collected by: R.S. Pieter Faure.

Tellina vidalensis Sowerby, 1904: 13, pl. 7 (fig. 6). Barnard 1964a: 541
= Tellina (Moerella) vidalensis Sowerby: Boss, 1969: 141, pl. 16 (figs 2-5).

TYPE SPECIMENS OF MARINE MOLLUSCA 49

Holotype: SAM-14848. Shell, complete.
Locality: off Cape Vidal; 23 m; no date.
Collected by: R.S. Pieter Faure.

Family Pandoridae

Pandora dissimilis Sowerby, 1894: 374; 1897: 21, pl. 6 (fig. 33). Barnard, 1964a:
S12.
Paratypes: SAM-5622. Two shells, complete.
Locality: Table Bay, Green Point; intertidal; no date.
Collected by: E. L. Layard.

Pandora similis Sowerby, 1897: 29. Barnard, 1964a: 572.
Paratypes: SAM-—A29743. Two shells, complete.
Locality: Bluff Channel, Durban; no depth or date.
Donated by: Mr Ponsonby.

Family Cuspidariidae

Cuspidaria nasuta Sowerby, 1904: 18, pl. 7, fig. 14

= Cuspidaria capensis (Smith): Barnard, 1964a: 580.
Holotype: SAM-14819. Shell, complete.
Paratype: SAM-A44372. Shell, complete.
Locality: off Cape Point Lighthouse; 155 m; 3 May 1900.
Collected by: R.S. Pieter Faure.

Cuspidaria optima Sowerby, 1904: 17, pl. 11 (fig. 16). Barnard, 1964a: 580.
Paratype: SAM-—14764. Shell, complete.
Locality: off Umtwalumi River, Natal; 92 m; 11 March 1901.
Collected by: R.S. Pieter Faure.

Family Myochamidae

Myodora rectangulata Barnard, 1964b: 28
= Myodora (Myodora) quadrata E. A. Smith: Kilburn 1973: 709.

Syntypes: SAM-—A9556. Twelve valves.
Locality: off O’Neil Peak, Natal; 92 m; 28 February 1901.
Collected by: R.S. Pieter Faure.

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BARNARD, K. H. 1964b. Two genera of Erycinacea (Bivalvia) from South Africa. Proc. malac.
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BARNARD, K. H. 1964c. The work of the S.S. Pieter Faure in Natal waters, with special
reference to the Crustacea and Mollusca with descriptions of new species of Mollusca from
Natal. Ann. Natal Mus. 16: 9-29.

BARNARD, K. H. 1969. Contributions to the knowledge of South African marine Mollusca. Part
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BARNARD, K. H. 1974. Contributions to the knowledge of the South African marine Mollusca.
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BosuorF, P. 1968. Descriptions of three new species of Pelecypoda dredged off the coast of
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Boss, K. 1969. The subfamily Tellininae in South African waters (Bivalvia, Mollusca). Bull.
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Cate, J. 1964. A new species of Mitra from the western Indian Ocean. Veliger 6: 219-220.

CERNOHORSKY, W. 1976. The Mitridae of the world. Part I. The subfamily Mitrinae. In:
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GosLINER, T. 1981. The South African Janolidae (Mollusca, Nudibranchia) with the description
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TYPE SPECIMENS OF MARINE MOLLUSCA a1

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SZ ANNALS OF THE SOUTH AFRICAN MUSEUM

TOMLIN, J. R. 1931. Reports on the marine Mollusca in the collections of the South African
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ToMLIN, J. R. 1939. A new South African vermetid. J. Conch. Lond. 21: 145.

TOMLIN, J. R. 1944. New South African Siphonaria. J. Conch. Lond. 22: 92-93.

ToMLIN, J. R. 1948. A new species of Pleurotomaria. J. Conch. Lond. 23: 2.

WaLts, J. 1979. Cone shells, a synopsis of the living Conidae. Hong Kong: T. F. H.
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WEAVER, C. 1963. Volute problems. Hawaii. Shell News 11 (5): 5.

WEAVER, C. & Dupont, J. 1970. Living volutes. A monograph of the recent Volutidae of the
world. Delaware Museum of Natural History. Monographic Ser. 1: 1-375.

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———————

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., Syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
“semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid- tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.

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ELIZABETH GILES
&
TERRENCE GOSLINER

PRIMARY TYPE SPECIMENS OF MARINE
MOLLUSCA (EXCLUDING CEPHALOPODA)
IN THE SOUTH AFRICAN MUSEUM

OCTOBER 1983 ISSN 0303-2515

OF THE SOUTH AFRICAN
MUSEUM

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

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

FiscHEeR, P.-H., DuvaL, M. & RAFFry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634. \

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 92 Band
October 1983 Oktober
Part Z Deel

Dr SS
Agu)

MORPHOLOGICAL AND BIOLOGICAL
NOTES ON SOME
SOUTH AFRICAN ARTHROPODS ASSOCIATED
WITH DECAYING ORGANIC MATTER
PART 1
CHILOPODA, DIPLOPODA, ARACHNIDA,
CRUSTACEA, AND INSECTA

By
A. J. PRINS

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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MORPHOLOGICAL AND BIOLOGICAL NOTES ON
SOME SOUTH AFRICAN ARTHROPODS
ASSOCIATED WITH DECAYING ORGANIC MATTER

PART 4

CHILOPODA, DIPLOPODA, ARACHNIDA, CRUSTACEA,
AND INSECTA

By
A. J. PRINS
South African Museum, Cape Town
(With 17 figures and 2 tables)

[MS accepted 5 May 1983]

ABSTRACT

Arthropods found in association with leaf litter and other decaying matter were collected
for forensic purposes in a narrow strip along the south and west coasts. Twenty-one species in
nine orders are discussed in some detail and information is given regarding their morphology,
biology, and ecology. Most of these arthropods play an important role in the breakdown of or-
ganic matter, others are merely predators, which are mainly discussed here.

CONTENTS
PAGE
re HchAMIMIOCMICHOR Sa st 2 kee ese ie Sic asieiae, hse hoi yo Ae ee ee 54
Pealecical and. biolosical notes 22: 2..6.2).- 202s =< fossa e 22s 2 a
ROA ENA OA eh a aed oy rst eee aan EA ce ee 57
Mase cral eNO DIOMOLP Ages ee ere ieee oo ans Sikes Oe wea oe 58
Re retihyp el CMCOPIGAC wrth ers, 2 fone eee pete oe eae ale 58
QUAY CICS (COSIANICH AUC S «ens en Sys nays ne eee 58
ESS plopoddn cena | nee Mae Rae Bice a ole Suet eee ke 60
MR eleteerMLOnta Noe erie toes aes ae as se ese rl pea enci ame Pees 62
| PETTITT Ye (CLC ee eae gk gt A A eee? 62
Cmmatomlussmorelerm (UuCas) 9 oh s i a ea 62
SUE ISSPANE AC NIC Beppe re TEE ie canes en) Be ee Rens Oe eee 65
“UST ANSE aR Ee oe a oe gle eer ea ae 65
ve UE ZANT ETUC ie 2 ae enh i RPO es ae ira a ny CLG ae 70
inte SCORDIONIG dss ata 2 Pes Se erie oe 5 Oats le wits ence 2a 1S
| QYPE Sal VU 1 Cd Ye oh gh Re ee a ee eo yet er eT Oe 8 Cape ee AE is
Wroplectes vanecatus (KOCH), 4. S25525 2 howe ae oer TS
BOE ASA ERIS ACC 509 GE FE ol wy do <a eat cto 245 SONS iN ie tee 78
Meio NIH PRIDOGAT yt ern hese re Sie eh A th arenes 78
NORESIE FEESOOC Ar wet pe eee ets aS ey ad Ae Ge cg ae 80
rently O@lnIS el ACM ets78 8 3 Fey io 2h ab adh SS a, Bil cS a 80
IR ORCCIIO SCORER (athe) ag a5). ne oo coe yee ee 80
Rantbyo Arima gene Aes ate asec Ao 5 at as tle crt ee 81
Armadiidunm vuleare (Uatreille) 2, 2 os. he ae les 81
ROE AS SIU EIS ED Dee ice oer, Sls ES ed ca = ete IE Be ere 82

a3

Ann. S. Afr. Mus. 92 (2), 1983: 53-112, 17 figs, 2 tables.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Order Blattodea yer Ssh tcc a ees er ee wae A 82
Faniily, Blattidac 225 aac eee yr eer Shh eee Ot 82
Pseudoderopeltis foveolata (Walker) .................... 83
lemnopieny x phaleratal(Saussure) hes eee 84

Order’ Mamtodears. ..5 hows steer we te ee ee 86
Bamily Mamntidae sc itt Wie ap eae i a 86
Oxypilusnasuius (Rabies) ieee oe eee ee 86

Order Dermaplerac. kee aoe ee ake ek ee 88
Family Carcinopiondacwr mre ernest res ee ee 89
uboneliatonnulipes: (Ibucas)men sare ee eee 89

Fanly Ronnculiciae. sete merce at) ae ae lal ys i aeh Ses, A Sie 90
Foricula penineuay il (Bn) eee eee 90

Family alcabidinidae, 2 eee tele anion | ase tne sre 92
Labiduramiparion(allas) peek. hee 92
OrdenOrnthopte rai. 5 een a eae ac. a) coy sonar Miata, a 93
Fannily Grylln@ ac nr ee capes aerials. es Che eR ee 93
CophogryllustdelalanduSaussure*.4. 20 ae eeeeee 93
COPHOBIYV USES Deere oho A vic i) hos ee 94
GryllustbunaculaisdeGeen eee eee 95

Order Hemiptera a xge nee teases ry ala oie Gas ae ik cae ates aie 99
Bamuly Anthocoridactm vee eno) ere ee 99
Spain ete. Apacer erase Mee eS. os ane nea ee ee 99
BannilyCorevdaess ohne et te, my icin ne 99
Leptocoris hexophithalma (ihunbere) sa. ee eee 100

Pamnullyel yeacidacesn sane ek her aes... kian lence 100
Melanostethus marginatus (Thunberg) .................. 100
FanullyaBentatomidae nvr. 2.0: Says sc ors ate ieee ree nee 102
Aintestiopsisiorbitalis (Westwood) 99.) ...). 4... ene 102
FamulysPyrnhocondaen a aia ntun eon ee aa ae hee es ee 103
SCONIUS) ONSIEKU AEICIUS) perc ai ae 103

Cenacus carnijed (HADnCIUS) mee ance ee Cee ee 104
FamilyiReduvinidae:! cwisasans oe ae ers Bae Sn coer eee ee 106
ONCOCEPIIGIUS: SPA iat COR HR eras ney Se ee ae 108
ACknOWwledsements *.) 2) Geen tang sro gtens Ocoee oct Ree 110
FRE LEGEMCES oct crepe i a rata, al ete a eee i a 110

GENERAL INTRODUCTION

Intensive studies have been conducted in the past in the field of medical and
veterinary entomology; however, forensic research has been neglected by ento-
mologists and our knowledge on this subject is therefore very limited. The few
facts that are known are usually included as a chapter in occasional major works
on forensic medicine. The most comprehensive data for South African entomol-
ogists are probably various paragraphs in Medical Jurisprudence (Gordon,
Turner & Price 1953) and an article on flies on human faeces and carcasses
(Zumpt & Patterson 1952).

Following requests by the police and State Health Department regarding
cases of murder and cattle theft, the author undertook research to obtain a bet-

ter knowledge of the arthropod successions related to decaying cadavers and |

thereby to provide additional evidence in cases presenting problems in judicial
examinations.

In an investigation of this nature two aspects have to be considered, first, a
survey of the various arthropod populations associated with decaying organic

SOUTH AFRICAN ARTHROPODS a

matter in a certain area, most of which may be indirectly involved, and second,
an assessment of those directly responsible for the destruction of cadavers and
carcasses.

As part of a survey of insects that may be involved in forensic entomology,
a preliminary study was conducted during 1976 to 1979 on the arthropods asso-
ciated with organic matter. Large numbers of arthropods were collected along a
narrow strip of the southern and western coastal regions of South Africa, on the
beach, the dune system, and some 20-30 km inland. A small strip between
Laingsburg and Beaufort West and another between Worcester and Tulbagh
were also included.

As the immature stages of the vast majority of these arthropods are still un-
known, an attempt was made to rear the various species either in their natural
breeding media or in suitable substitutes in order to obtain information on their
life cycles. The condition and colour of the specimens were noted before preser-
vation in 80 per cent alcohol and drawings were made with the aid of a camera
lucida. The average monthly temperature in the laboratory was recorded
throughout the observation periods (Fig. 1A—D). The morphological terms used
in this series of papers in the descriptions of the various instars are those of
Greene (1922), Béving & Craighead (1931), and Hennig (1968), unless other-
wise stated. Identification of the specimens was made by reference to material in
the collection of the South African Museum or as verified by specialists in vari-
ous other institutions.

The first part of this series includes centipedes, spiders, scorpions, ticks and
mites, mantids, earwigs, flower-bugs and assassin-bugs, which are all predacious
except for ticks and certain mites. It also includes millepedes, sow-bugs, beach-
fleas, cockroaches, crickets, red-bugs, chinch-bugs and shield-bugs, which are
mostly phytophagous or are in some or other way associated with decaying or-
ganic matter. These arthropods are really of minor importance to the forensic
entomologist but may play a role either as predators or in breaking down or-
ganic material in the soil. Some species such as the chinch-bugs, red-bugs and
shield-bugs are mainly phytophagous, but they were often found in leaf litter or
accumulations of debris and have therefore been included.

The terrestrial arthropods of a particular region are dependent in most
cases directly or indirectly on the floral elements of that region. The coast, with
its marine, animal and plant detritus, and unique flora of its adjacent dunes in
general, has a very different arthropod fauna from the inland areas.

The beaches along the south and west coasts that were surveyed are usually
separated from the interior by a sand-dune biotope which is characterized by the
presence of a fairly large variety of plants of which the ‘kinkelbossies’, Tetrago-
nium decumbens and T. fruticosa, as well as Arctotheca populifolia, Heteroptilis
suffruticosa, Matricaria sabulosa, Mesembryanthemum crystallinum, and Limo-
neum perigrinum are mostly found on the small fore-dunes. Various other plants
such as Ruppia, Zostera, Phragmitis and Spartina spp. are to be found along the
river-banks and in estuaries. Grasses including Agropyron distichum, Sporo-

ANNALS OF THE SOUTH AFRICAN MUSEUM

56

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LLJbj0)|

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8261

‘040d AON LOO dAS ‘ONV CINE “NOE AVIV ‘YdV “YVW “834 “NVE

9L61

(nn
~)

SOUTH AFRICAN ARTHROPODS

bolus virginicus, and Eragrostis spinosa frequently grow on the beaches, even
reaching the intertidal zone in some areas, and along the Cape south coast Os-
teospermum fruticosa and Polygala mystifolia often reach the shingle-covered
parts of the beach above the highwater mark.

On the beach itself there are mainly brown and green seaweeds such as
Ecklonia maxima and Laminaria pallida, and very often also the dead bodies of
marine animals and birds. In some areas along the south and south-east coast
the large kelps are entirely absent and the beaches are usually completely barren
and devoid of any seaweed, which results in a sparse arthropod fauna. Further
west, however, the number of plants washed up by wave action increases and
provides an excellent habitat for a large variety of arthropods.

Various factors will influence the properties of the wrack arthropod com-
munities and man as an extra-biotopic agent cannot be excluded, particularly in
areas where commercial collection of kelp occurs as some eucoenic species can
exist only in large banks of kelp (Backlund 1945). In certain parts along the west
coast where kelp is collected for commercial purposes, some of the species nor-
mally present were not found during the surveys.

Three types of wrack are generally recognized, wrack strings, wrack flakes,
and wrack banks (Backlund 1945; Egglishaw 1960). The banks are the most im-
portant as they provide the main source of food for the various species of the
shore fauna.

Continuous exposure of the marine rejectamenta to sprays and wind-borne
moisture from the sea affects the composition of the arthropod communities and
there is therefore a marked difference between the amnicolous and thinicolous
associations and the fauna of the greater interior. Although the maritime species
of arthropods exhibit a far greater tolerance to salt than those living further
away from the sea, there is an overlap of the various species along the coast, es-
pecially among certain Coleoptera, Diptera and Lepidoptera. On the other
hand, the vast interior with its variable ecosystems and very large floristic poten-
tial presents a wide range of biotopes supporting a comparatively large inverte-
brate fauna, and a greater number of arthropod species will therefore attack
decaying and dry organic matter.

ECOLOGICAL AND BIOLOGICAL NOTES
CLAss CHILOPODA

Usually active, nocturnal predators with dorsoventrally flattened or de-
pressed bodies. Only one pair of legs on each body segment; first pair modified
into poison claws. Ocelli may be present.

Various centipedes were found to be associated with decaying matter, and
these include the rather small species of the genus Lamyctes with only fifteen
pairs of legs and the very elongate, worm-like species of Eurytion such as E. do-
lichocephalus Attems, which is pale reddish in colour, measures up to 60 mm in
length and has nearly eighty pairs of legs. The latter species was collected in all

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

the areas surveyed and was often found deeply imbedded in semi-dry dung dur-
ing the winter. Cast skins have often been found together with medium-sized
specimens. From collection records it seems to be endemic to the Cape, having
been found along the west and south coasts and inland as far as De Aar and
Hanover. However, further collections may prove otherwise.

Of the Scolopendromorpha, at least two species appeared to be fairly com-
mon under semi-fresh to almost dry cow-pats wherever collections were made.
The one, a bluish-coloured Cormocephalus species (about 50 mm long, and its
larvae), occurred at Hermanus and Stilbaai during the winter; it is apparently
the same species that was collected near Philadelphia during the same period. In
the western and north-western parts, particularly in the coastal areas of Nama-
qualand, a large proportion of the pats contained the bluish-green Arthrorhab-
dus formosus Pocock, most of which were about 45 mm long with a broad,
brownish-red, longitudinal stripe over the dorsum. Specimens from Saldanha,
on the other hand, were more reddish in colour and the mid-dorsal stripe was
greenish. This centipede is very similar to Cormocephalus species, but is easily
distinguished by the spines on the tarsi (absent in Cormocephalus species) and
by the posterior margin of the head plate, which is free (covered by the first
body segment in Cormocephalus species). Prey of both species included spiders,
cockroaches, and silver-fish.

The introduced house centipede, Scutigera coleoptrata (Linnaeus), with its
long slender legs, often appeared at carcasses in various parts of the Cape Penin-
sula during the later post-mortem stage of decay where it mainly devoured some
of the members of the fly population. It is widely distributed in the western,
southern, and eastern Cape and in Natal.

ORDER LITHOBIOMORPHA
Family Henicopidae
Lamyctes castanea Attems

Adult centipede small, brownish red to reddish, 10-12 mm long. Head
somewhat darker in colour with one ocellus on each side. Fifteen pairs of legs,
tibia of twelfth pair not toothed. Posterior angles of all tergites rounded, not
dentate (Fig. 2A-B).

The species seems to be endemic to the Cape Province and is commonly
found under semi-fresh to almost dry cow-pats both inland and on the beach,
often in association with L. africana Porat; it regularly visits carcasses and
corpses to feed on the small soft-bodied insects attracted to the decaying ma-
terial. Both L. castanea and L. africana are more or less of the same length and
colour, but L. africana may be recognized by the toothed tibiae of the twelfth
pair of legs (Fig. 2C). The first-mentioned centipede was found almost through-
out the areas visited, but L. africana occurred mostly in the north-western parts.
A large red species, Lamyctes denticulata Attems, about 13,8 mm long, ap-
peared in fairly fresh dung at Mamre and fed in captivity on soft-bodied insects.

SOUTH AFRICAN ARTHROPODS 59

OV 4

ih
Se Neate
CNS
a

SAI: SO aa w
OO TAS

ey

Fig. 2. A-B. Lamyctes castanea. A. Adult. B. Head of adult showing first and second pairs

of legs, ocellus (oc) and maxillipeds (mxp). (Left lateral view.) C. Lamyctes africanus. Tibia

and tarsus of twelfth pair of legs to show tibial tooth. D. Lamyctes denticulatus. Segments
8-11 to show denticulate posterior border of segments 9 and 11.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

It differs from both L. castanea and L. africana by the toothed posterior angles
of segments 9, 11, and 13 (Fig. 2D).

As these centipedes are long lived, both larvae and adults are usually found
together. According to Cloudsley-Thompson (1957) the Lithobiomorpha (which
includes Lamyctes) hatch with seven pairs of legs, including the poison claws,
but this stage apparently lasts only for a few hours. Almost pure white, anamor-
phic larval forms of L. castanea with seven pairs of legs, excluding the maxil-
lipeds (Fig. 3D) and measuring 1,3—1,5 mm in length, were observed in the soil
around Cape Town during August. They are further characterized by the pres-
ence of an eighth pair of legs visible posteriorly through the integument. Eyes
are apparently absent and the antennae consist of only six segments.

Anamorphic larvae with eight and ten pairs of legs (maxillipeds excluded)
appeared in large numbers during August and September and in both cases two
_ pairs of legs are visible posteriorly through the integument (Fig. 3A—C). The an-
tennae are fourteen-segmented. The specimens with ten pairs of legs are some-
what larger, measuring 2,6-2,8 mm. In both forms the head and antennae are
already pale yellowish red and the tergites, including the intercalary tergites, are
fairly well defined; indications of eyes are present.

The final anamorphic stage larvae with twelve pairs of legs are very similar
to those with ten pairs; however, in specimens collected during October, the an-
tennae consist of sixteen segments and three pairs of posterior legs are visible
through the integument. These larvae measure 2,6—3,5 mm in length.

Various groups of the second or final epimorphic stage individuals were ob-
served during the late spring and almost throughout the summer and autumn.
These specimens, with fifteen pairs of legs, measure 3,8-11 mm in length and
their antennae consist of twenty segments in the 3,8 mm specimens, and thirty-
one segments in the 11 mm specimens.

CLASS DIPLOPODA

Slow-moving herbivorous animals with cylindrical bodies characterized by
presence of diplosegments, each bearing two pairs of legs. Ocelli present or ab-
sent. Defence is effected either by rolling up into a ball or by secretion of pro-
tective chemicals in segmentally arranged glands.

Millepedes are usually present wherever leaf litter or decaying dung is
found, particularly the garden millepede, Ommatoiulus moreleti (Lucas), the
large black Harpagophora nigra Attems, the black and reddish striped Chersas-
tus digrammus (Pocock), a brownish species of the genus Julomorpha, the pale-
brownish Gnomeskelus repandus Attems, and the hothouse millepede, Ortho-
morpha gracilis (Koch).

The last-mentioned millepede is widely distributed in the Republic of South
Africa and, according to Causey (1943), its eggs hatch within 5-10 days (in the
USA). He gives the size of the eggs as 0,35-0,41 mm. It is a polydesmid like
Gnomeskelus repandus, and therefore has a fixed number of instars of which the
eighth, with thirty to thirty-one pairs of legs, is the final or adult form (compare

SOUTH AFRICAN ARTHROPODS 61

Fig. 3. Lamyctes castanea. A. Anamorphic larva with eight pairs of legs (dorsal view).

B. Poison fangs (mxp) and first pair of walking legs of anamorphic larva (left lateral view).

C. Last two pairs of legs of anamorphic larva (left lateral view). D. Anamorphic larva with
seven pairs of legs (dorsal view).

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

with the juliformids, Ommatoiulus moreleti and Chersastus digrammus where a
large number of instars occur).

A tiny unidentified dermestid-like pincushion millepede (Penicillata or Pse-
laphognatha) (Fig. 11H) was found together with Ommatoiulus moreleti under
cow-pats near Jacob’s Bay on the west coast. It is about 2,9 mm long, has twelve
pairs of legs and is brownish red in colour with tufts of dark-brown, thick hairs
on each segment including the head. It was seen to nibble on certain parts of the
dung and its remains were often found in the shelters of reduviids of the genus
Coranus.

Another unidentified species (the specimen collected is about 1,8 mm long
with eight pairs of legs), which is pale yellowish white in colour, is often ob-
served in leaf litter in the Cape Peninsula. It differs from the latter species by
the longer hairs and antennae in relation to body size and by the almost glabrous
dorsum, devoid of any hairs except for some long scale-like setae placed dorso-
laterally. Its habits are unknown. The specimens of both the species collected
are apparently immature, as adult forms usually have at least thirteen pairs of
legs. Lawrence (1981) gives some idea of the life history of these millepedes.

ORDER JULIFORMIA
Family Julidae
Ommatoiulus moreleti (Lucas)

Adults small to medium sized, dark slate-grey to blackish in colour and
20-45 mm long when full grown. Characterized by longitudinal striae on each
body segment and presence of a mid-dorsal posterior extension or tail. After the
adult stage is reached, several further moults occur with an increase in the body
length and the number of legs and ocelli, as is shown in Table 1 for adults col-
lected around Cape Town and Philadelphia during January and February 1979.

This introduced millepede is almost cosmopolitan and is very widely distrib-
uted in the southern and western Cape. It was collected along the coast as far
north as Citrusdal and as far east as Stilbaai. It sometimes occurs in very large
numbers, particularly in soils rich in organic material, and may cause damage to
young plants and even plant bulbs. It is often found in thatched roofs of houses
together with scarabaeid larvae and it is quite possible that their way of feeding
provides the correct stratum for these scarabs. Schubert (1966) gives a good de-
scription of the species and an almost complete distribution in the Cape.

It is commonly found in leaf litter as well as in soil rich in compost and
other organic materials used as manure such as fishmeal, and is also attracted to
decaying carcasses and human cadavers; on the beach it was often seen feeding
on the dry remains of bird and seal carcasses and further inland even appeared
at carcasses during the early part of the dermatophagous stage of decay. It was
the only millepede found at wrack strings in the Cape Peninsula during the sur-
veys. Together with Chersastus digrammus it usually appears in large numbers

SOUTH AFRICAN ARTHROPODS

TABLE 1

Body length of adults of Ommatoiulus moreleti in relation to

Number of leg pairs

number of legs and ocelli.

Number of ocelli
on each side

~]~4J NM ~] ~)

d& GS G2 Wo Wo bo Yo Lo bo bo

hk Od OW ~) UO

TABLE 2

Body length in mm

19.1
18.0
18,0
22,0
19.0

Body length of Ommatoiulus moreleti in relation to number

Number of leg pairs

ONO DN DN Nn hn Nn fe
GO) G2 Gd me Uo ee Rm Uo

ON
On

of legs and ocelli.

Number of ocelli
on each side

10
15
15
15
21

Body length in mm

6.6

12,

—_

~) ON On On fee Go ON Uo &
(=) C=) be) i i i Re ie (i)

SS ee

~~] 00

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

during April and May under semi-fresh cow-pats in sandy areas, feeding on the
wet parts of the dung. It is also common under dry cow-pats.

Copulation between males (19,9 mm long with seventy-three pairs of legs),
and females (32 mm long with eighty-three pairs of legs) of O. moreleti was ob-
served near Philadelphia during May. In most cases this lasted for about 10 min-
utes and after a few days each female laid some 200 eggs in a small hollowed-out
chamber in the topsoil. The eggs are somewhat oblong, slightly shiny and dirty,
yellowish white, measuring from 0,76 X 0,64 mm to 0,80 X 0,60 mm. The incu-
bation period in the laboratory was about 16 days at 22 °C. The first sign of the
so-called preliminary hatching is a rupture in the chorion of the egg, after which
the immobile, legless embrionic pupoid stage (Fig. 4A), which is still enclosed in
an embrionic cuticle and is almost pure white, appears. About 4 days later the
legs become visible through the membrane and anteriorly a tiny eye-spot ap-
pears on each side, and a pair of larger, darkish-orange spots also becomes vis-
ible at.the posterior third of the body. This pupoid stage is about 1,0 mm long.

After about 6 days the embrionic cuticle is shed and the active first instar
with three pairs of legs appears (Fig. 4B). It is about 1,6 mm long, still pure
white and with about eight visible body segments. The darkish orange spot is
then situated on about the fifth segment. After 5—6 days it moults again and then
has seven pairs of legs (Fig. 4C) and one pair of eye-spots and the large lateral
orange spot is then situated near the middle of the body. The first five trunk seg-
ments are more yellowish in colour and the body-length remains about 1,6 mm
long.

In the case of Chersastus digrammus (Trigoniulidae) the second larval instar
is slightly darker and three distinct black ocelli are visible on the head, but the
dark posterior spot is much less conspicuous. Young specimens of the latter
Species were observed along the west coast during December, the smallest found
being 5,3 mm long with a brownish, longitudinal median line on the dorsum, as
in O. moreleti.

After 16-17 days the young larva moults again and is now about 2,8 mm
long with seventeen pairs of legs and three clearly demarcated ocelli on each
side of the head (Fig. 4D). This third instar is characterized by a pair of large,
dark, reddish spots on the fifth body segment as well as five pairs of smaller,
reddish, oval spots posterior to it (these are the openings of the repugnatorial
glands). The posterior extension or tail and the antennae, which are very con-
spicuous at this stage, are already visible from the first instar. The body colour
usually becomes darker and after about a month it moults for the third time,
producing an individual with twenty-nine pairs of legs and six black ocelli on
each side (Fig. 4E). It is then about 4,9 mm long and laterally bears eleven small
reddish spots of which five are very conspicuous, in addition to the large spot.

This fourth stage lasts for about 25 days before it moults again and then
reaches a length of about 6,2 mm. There are thirty-nine pairs of legs, ten ocelli
and laterally eleven small, clearly defined, reddish spots and five inconspicuous
spots in addition to the larger one on the fifth segment (Fig. 4F). The colour of

SOUTH AFRICAN ARTHROPODS 65

the body is pale brownish white, with a fairly broad, longitudinal, brown median
line along the dorsum. This fifth instar lasts for about 32 days and again moults
near the end of September. The sixth instar has about fifty-one pairs of legs and
fifteen ocelli on each side of the head.

The number of legs varies apparently according to the sex of the animal and
therefore fifth-instar specimens with forty-three pairs of legs and ten ocelli were
often collected. As the animal increases in size, the number of legs and ocelli
corresponds with the size of the millepede, as is shown in Table 2 (see also
Table 1).

Baker (1978) described the post-embrionic development and life-history of
this species in Australia and stated that after 1 year O. moreleti was in the se-
venth, eighth or ninth stage. After 2 years the tenth or eleventh stage was
reached and after 3 years the twelfth or thirteenth.

CLASS ARACHNIDA

Arthropods with the prosoma (or cephalothorax) or both the prosoma and
opisthosoma unsegmented; sometimes all the segments fused. Mostly terrestrial
except for a few aquatic mites and spiders. No antennae, compound eyes or
wings present. Respiration by means of tracheae, or book-lungs or both. Gener-
ally carnivorous and cryptozoic.

ORDER ACARINA

Mostly small arthropods, with the prosoma and opisthosoma completely
fused. Larvae with three pairs of legs, nymphs exactly as adults with four pairs
of legs but lacking the genital opening. In certain groups such as the Tetrapodili
adults have only two pairs of legs.

Ticks (Ixodides) have the hypostome barbed and are usually parasitic on ver-
tebrates. Only two species were found to shelter under semi-dry to almost dry
cow-pats, namely the bont-legged tick, Hyalomma truncatum Koch and the red
tick, Rhipicephalus evertsi Neumann. Both males and engorged females of the
two species were recovered during March, H. truncatum in the Philadelphia dis-
trict and near Saldanha and R. evertsi in the Stilbaai area. Males of H. truncatum
measured 4,8 mm from the tip of rostrum to the posterior border of the body and
were dark brown to blackish brown in colour. Engorged females were red and
measured about 13 mm. Females of R. evertsi were somewhat under-developed,
with light-red shields; their bodies were yellowish white and the legs pale reddish.
The females of both species were embedded in the cake of the pats and, in the
case of the red tick, large numbers of eggs were found with them. The eggs are
roundly oval, dark reddish brown to wine coloured and shiny, and measure
0,52 mm in length and 0,38 mm in width. They hatched in about 50 days, pro-
ducing transparent six-legged larvae measuring 0,60 mm shortly after hatching.

Mites (Acari), on the other hand, are usually small to minute with the hy-
postome smooth and without barbs, and are generally found in almost every

ANNALS OF THE SOUTH AFRICAN MUSEUM

66

C. Second stage. D. Third

B. First stage.
F. Fifth stage.

A. Pupoid stage.
E. Fourth stage.

stage.

Fig. 4. Ommatoiulus moreleti.

SOUTH AFRICAN ARTHROPODS 67

habitat available to animal life. Some species are phytophagous, causing severe
losses to crops, others again are parasitic and have developed a close relation-
ship to man, such as the asthma epidermoptid, Dermatophagoides pteronyssinus
(Trouessart), also known as the house-dust mite, which is associated with the
scales of the human skin; it allergizes the bronchi by inhalation (Glass 1971) and
contributes to allergic rhinitis and asthma.

Free-living mites are abundant in debris on the ground or in litter-like accu-
mulations and particularly in soil rich in compost or other decaying organic mat-
ter. The most common species in the Cape Peninsula in soils containing
decaying fish-meal was the Sarcoptiform, Lardoglyphus zacheri Oudemans (Fig.
51), a minute whitish species only 0,2—0,24 mm long. This mite was also found to
infest certain carcasses and when large numbers were present even the larvae of
the skin beetle, Dermestes maculatus de Geer, were covered with numerous
specimens. They apparently did not harm the insects at first, although their
movements were slowed down. Later, however, after prolonged exposure to
large numbers of this mite, growth was retarded and mortality increased.

In the oribatid mites (Fig. 5F) the bodies are usually strongly sclerotized
and darkly coloured and some have large lateral shields or pteromorphs. They
are fairly numerous in rich soils and may play an important role in digesting the
organic matter (see also results by Fujikawa (1979) in Japan). They are found to
be common around carcasses and one species of Baloghobates of the family
Ceratozetidae, which has small pteromorphs, was found to accumulate in large
numbers on the decaying leaves of Arctotheca populifolia along the Strandfon-
tein sand-dune system during the summer and early autumn, in some areas actu-
ally feeding on the growing plants.

The variable Trombidiformes include a large number of phytophagous
species as well as several parasitic mites, particularly of the family Tarsonemidae
of which one species of Tarsonemus (Fig. 5D) was very common on a large variety
of arthropods. It is small, about 0,24 mm long, whitish, and with long setae on the
hind legs. In the Karoo it occurred in exceptionally large numbers on the larvae of
the skin beetle, Dermestes maculatus de Geer, feeding on the droppings of the
Cape grey mongoose, Myonax pulverulentus (Wagner). Heavy infestations of this
species on insects in breeding jars in the laboratory usually resulted in the death of
the hosts. This mite, together with one species of pyemotid (Fig. 5E), was found
to be common around Cape Town in the soil surrounding half-buried carcasses.
The latter is also very small, about 0,5 mm in length, almost globular, and pure
white in colour. Specimens observed during the dermatophagous stage of decay of
various carcasses were found to feed on the eggs of skin-and-hide beetles. The
grain itch-mite, Pyemotes ventricosus (Newport), another member of the:family
Pyemotidae and normally beneficial as it is parasitic on the larvae of grain-
infesting insects, may cause skin irritations or irruptions in humans handling
infested material. (This mite is not to be confused with the human itch-mite,
Sarcoptes scabiei (de Geer) (Sarcoptiformes) which is transmitted by direct con-
tact with infested persons or domestic animals. )

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Prostigmatid mites of the genus Pimeliaphilus, also belonging to the sub-
order Trombidiformes, have been found in association with various reptiles and
arthropods in different parts of the world. Olivier (1977) described four species
from South West Africa of which three were found on insects, viz. P. penrithi
Olivier on cockroaches of the genus Derocalymma, P. buysi Olivier on the tene-
brionid beetle, Stips dohrni (Haag) and P. desertus Olivier on the carabid Anthia
thoracica (Fabricius) (see also discussion on Scorpionida). Most of these species
also seem to be free-living under stones and debris.

In completely dry cow-pats only two undescribed species of prostigmatic
caeculid or rake-legged mites were found. The one species, observed along the
west coast, is about 2,5 mm long, its body trapeziform and broad. Its colour is
brownish to greyish marked with black; the front and sides of the shield are
whitish and the legs are black marked with white. Body-hairs are sparse and
- spatulate. The front legs are strongly developed, with long golden-brown spines
on the inner surface.

The other xerophilous species collected in the Karoo is much smaller, about
1,6 mm long, piceous brown, and covered with longer hairs. It lacks the white
marks of the former species. The colour of these mites blends with that of the
dry dung to bring about a perfect camouflage. In the laboratory they were seen
to stalk other mites, but otherwise their habits are unknown. Lawrence (1939)
maintains that they are probably scavengers, feeding on decaying animal
remains.

A large variety of mostly unidentified mesostigmatid mites were very com-
mon in both fresh and semi-fresh cow-dung. These included various Macrocheles
species such as M. distanti Evans & Hyatt (Fig. 5A), M. peniculatus Berlese,
and Parasitus species (Fig. 5B). An unidentified crimson, prostigmatid spouted
mite of the genus Bdella (Fig. 5G), about 1,6 mm long, appeared regularly but
was not abundant in semi-fresh pats. One of the most numerous species in de-
caying dung was a small, whitish mite about 0,32 mm long and very similar to
Parasitus sp.; it is probably also a member of the Parasitidae. These species are
all predacious, the snouted mite feeding on the Collembola present.

Specimens of M. distanti that were collected measured only 1,1-1,2 mm
long, with the holodorsal shield yellowish brown in colour; specimens of M.
peniculatus, on the other hand, were somewhat bigger, 1,2-1,4 mm long, and
darker in colour. Macrocheles distanti is commonly found on various scarab
beetles such as Onitis, Scarabaeus and Circellium species (Evans & Hyatt 1963).
Parasitus species are more or less oblong mites and, although of the same colour
as the above-mentioned species, they are easily distinguished by the divided dor-
sal plate; their legs are also more slender and more or less of the same size,
whereas in the macrochelids the first pair is elongate and much thinner than the
others. All these mesostigmatid species also occurred in decaying animal car-
casses and human cadavers.

Some of the Macrocheles species are ectoparasites of flies, as they are often
found attached to their hosts, their mouth-parts deeply embedded in the tissues

SOUTH AFRICAN ARTHROPODS 69

Fig. 5. A. Macrocheles distanti. B. Parasitus sp. C. Cheese skipper, Piophila megastigmata
McAlpine, with two mites, M. distanti attached to abdomen. D. Tarsonemus sp. E. Pyemotid
mite. F. Oribatei mite. G. Bdellasp. H. Aleuroglyphus sp. 1. Lardoglyphus zacheri.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

between the segments (Fig. 5C). Phoresy, therefore, seems to be one of the
main factors influencing the presence and abundance of these mites in cow-pats
and carcasses. According to Axtell (1963) several species of the family Macro-
chelidae are predacious on house-fly eggs and first instar larvae.

The mesostigmatic poultry mite, Dermanyssus gallinae (de Geer), is often
found to shelter in roosting sites of birds in human dwellings, particularly in air-
vents, and may cause minor infestations. In some respects it resembles the larger
Parasitus species, but may be distinguished by the absence of a clearly divided
dorsal plate.

Decaying kelp yielded numerous mites, mainly Mesostigmata and Sarcopti-
formes, especially of the families Laelaptidae and Acaridae. Various species of a
brownish Peletiphis have been seen, while the acarids were represented by a
pale-brown, parasitic Aleuroglyphus sp. (Fig. 5H). Often kelp-flies were so
heavily infested with them, particularly on their legs, that their movements be-
came sluggish. Most of the mites are about 0,68 mm long. A Parasitus sp. (Fig.
5B) also occurred and was found to feed on the eggs of flies laid on the kelp. It
was noticed that when disturbed each mite snatched an egg and disappeared
with it among the Kelp leaves. When fly maggots or beetle grubs were preyed
upon by predacious beetles and their larvae, these mites were seen to approach
the prey and imbibe some of the available body fluids.

ORDER ARANEIDA

In the spiders the abdomen is usually unsegmented except in the Liphistio-
morphae; in this suborder as well as in the Mygalomorphae, which includes the
baboon-spiders, the fangs move up and down. In the rest of the spiders the fangs
move from side to side (or in and out).

Most spiders are poisonous and, although necrotic ulceration or loxoscelism
is caused by the cytotoxic venom of the violin spiders, most of them, such as the
baboon-spiders and the button-spiders, have a neurotoxic poison.

At least four species have been observed in debris in the sand, on dunes,
and on the supratidal zone of the beach. These are mostly burrowing spiders and
they disappear very quickly under the sand when left on the surface. The most
common of these were the armoured spiders (family Zodariidae) (Fig. 6C), and
a small theraphosid. Occasionally a species of Clubiona (family Clubionidae),
found on the dunes, occurred under semi-dry crow-dung both along the south
and west coasts, while one species of the family Ctenidae also frequently ap-
peared under the stranded kelp and debris in the western parts.

These are all small spiders of which the feeding habits are unknown. How-
ever, there is evidence to believe that some of the zodariids prey on the small fly
larvae in the sand, particularly those of the horse-flies belonging to the genus
Limata, which occur in this stratum.

Spiders were common in both semi-fresh and semi-dry cow-pats; the most
widespread appearing to be the wolf-spiders (family Lycosidae). A small Par-
dosa sp., easily recognized by the broad, brown longitudinal band on each side

SOUTH AFRICAN ARTHROPODS WA

of the pale carapace, the arrow-like pale line down the middle of the abdomen,
and the mottled legs, occurred throughout the region surveyed, most often to-
gether with a larger unidentified species. The latter, about 15 mm long, has a
similarly marked carapace, but its pale-brown abdomen is mottled with dark
brown.

Wolf-spiders were also found to be the most common arachnids under
stranded kelp, and a small species of Lycosa (pale reddish-brown with a broad,
brown longitudinal stripe along each side of the body, the middle of the abdo-
men whitish) was very numerous, particularly along the west coast; it was some-
times in association with members of the family Ctenidae, which in some
respects resemble the wolf-spiders. A larger species, about 10,5 mm long and
rather similar to the Lycosa, its abdomen marked with dark brown, seemed to
be just as common along the south coast. Most of them feed on the smaller in-
sects, particularly the flies that are attracted.

The large, brown hairy baboon-spiders, Harpactira spp. and the lesser ba-
boon-spiders, Harpactirella spp. (family Theraphosidae), particularly H. light-
footi Purcell, have all been collected under semi-dry to almost dry cow-pats
along the west coast as far north as Port Nolloth. In some cases the nests were
hollowed out in the pats, while in others they were just below the surface of the
soil, their tunnels leading into the pats. Harpactirella spp. are very similar to
those belonging to Harpactira, but differ from them by the absence of a brush of
short stiff hairs on the external side of each chelicera. The bite of H. lightfooti is
of medical importance (Newlands 1972). Specimens of Harpactira when in a
state of moulting are very susceptible to the attacks by pentatomids of the genus
Halyomorpha. An unidentified dark-grey species, very near H. viridescens
Walker and about 13 mm long, was found in the vicinity of Saldanha feeding
gregariously on fairly large specimens of the brown baboon-spider.

Small (1,9 mm long) blattid-like nymphs, found in the same area, probably
belong to this shield-bug. They are pale greyish yellow on the thorax and con-
nexivum, with greyish-yellow legs and antennae and crimson abdomen.

The family Drassidae were also well represented. In the Saldanha and Veld-
drif areas Amusia cataracta Tucker (Fig. 7B), Zelotes sp., and an unidentified
species very near A. cataracta were mostly present. They are all small, sombre-
coloured, brownish to pale bluish-brown spiders. A similar but much paler
species of Anagraphis was common almost throughout the region, and in the vi-
cinity of the Gouritz River mouth an unidentified Drassodes sp. appeared in pats
containing large numbers of dipterous and scarab larvae. Its colour is pale
brownish red, the abdomen lighter with brownish spots on the posterior half,
and it is easily separated from the other drassids by its long chelicerae carried al-
most horizontally.

The erigonids resemble in some ways small theridiids (button-spiders), but
the males and females are about equal in size. However, in the males there is a
raised area or knob on the front part of the carapace on which their eyes are
situated. Most of them are very small and sombre coloured and five to six

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

species were well represented under semi-dry pats, the most common being a
species of Araeoncus. Other small spiders observed in dry dung were species of
Heriaeus, Clubiona and the lungless spiders, Caponia, which resemble baboon-
spiders.

One of the most beautiful spiders that shelters under semi-dry cow-pats is the
tiny spitting spider, Scytodes sp. (Fig. 6A) (family Sicariidae), which is yellowish
white with dark-brown stripes and spots on the body. It is easily recognized by its
strongly convex and bulbous carapace. Specimens collected near Saldanha
measured only 3—4 mm in length. Prey consisted mostly of small flies, silver-fish,
and other small insects, which are usually immobilized by a sticky saliva.

Nests of the brown or house button-spider, Latrodectus geometricus Koch
(family Theridiidae), were not uncommon under pats and it was observed to
feed on various beetles and other soft-bodied insects such as cockroaches. The
black widow, L. mactans Linnaeus, on the other hand, was much scarcer and
only two nests were observed in the vicinity of Leipoldtville during April, both
under large dry pats, each nest with one or two smooth, oval, white egg cocoons
containing the exuviae of newly hatched spiders. The prey included tenebrionid
beetles, the most abundant victims being the elongate, brownish Oxura setosa
Kirby (16,5-19 mm long) that shelters under dry cow-pats during the summer
and is characterized by the two acute posterior prolongations of the elytra.
Hesse (1942) gives an almost complete list of insect victims of this spider, includ-
ing this tenebrionid. Skeletons of an almost equal number of a black Onymacris
sp. (about 14 mm long), and a few shiny brown Ograbies subdentatus Koch
(7,5-8,3 mm long), were among the remains of the one nest, together with al-
most intact specimens of a small black Psammodes sp. and a Zophosis sp., both
varying in length from 9,9 to 13,0 mm.

It is interesting to note that the sphecid Chalybion spinolae (Lepeletier)
provisions its nest with both species of button-spiders. According to obser-
vations, this wasp accounts for the destruction of a large number of the black
widow in certain areas (J. E. Nel, Durbanville, 1983 pers. comm.).

At least eight spiders were found in the Cape Peninsula in soil surrounding
carcasses, of which a dark crimson species of Dysdera (family Dysderidae) was
the largest (approximately 14 mm). It has a pale-reddish abdomen and excep-
tionally long chelicerae and was particularly abundant during the summer and
autumn. Young individuals collected during the summer varied from 3 to 5 mm
long and were almost pure white. The comb-footed spiders (family Theridiidae)
included a small brownish Enoplognatha sp., its abdomen marked with white
stripes and patches, and an even smaller Anelosimus sp. (Fig. 7A), which is pale
brown with darker brown on its abdomen.

An unidentified Erigone sp. (Fig. 6B) (family Erigonidae), probably the
Same species that was found under cow-pats near Hermanus and Darling, has
the spinners surrounded with black. It occurred together with a related button-
spider-like linyphiid and some larger dictynids (mesh-web spiders) under fairly
decayed carcasses. Hahnia sp. (family Agelenidae), about 4mm long, with

EE

——————————

————E

SOUTH AFRICAN ARTHROPODS

}

SS

Fig. 6. A. Scytodes sp. B. Erigone sp. C. Zodartid spider.
(dorsal and left lateral view).

ts,

mm Ms

D. Gamasomorpha australis

ANNALS OF THE SOUTH AFRICAN MUSEUM

74

FS

‘plopavws nisnuiy ‘g ‘ds snunsojuyp yy ‘Ll ‘34

SOUTH AFRICAN ARTHROPODS TS)

brownish, speckled abdomen, appeared in places where the soil was fairly damp.
Its long spinners are placed in a line along the posterior border of its abdomen;
young individuals collected were almost white.

One of the smallest spiders collected in the soil partly covering carcasses
was the bright brownish-red Gamasomorpha australis Hewitt (family Oonopi-
dae) (Fig. 6D), which is only 1,8 mm long and easily recognized by the two oval
shields covering the abdomen.

Apart from the true spiders, only one brownish harvest spider of the genus
Opiliones (order Phalangida), which is generally found under stones along the
west coast, was collected under dry cow-pats. It feeds on other spiders, flies, and
snails. Near Saldanha Bay a young specimen (37 mm long) of an unidentified
Solpuga sp. (order Solpugida) appeared during midsummer under a large dry
pat and, judging by the remains found near it, this animal must have been feed-
ing on the scorpions Uroplectes variegatus (Koch) and a Parabuthus species.

ORDER SCORPIONIDA

Fairly large nocturnal carnivores with segmented opisthosoma divided into
pre-abdomen or mesosoma, bearing ventrally openings of book lungs and paired
comb-like pectines, and with the tail or metasoma with apical sting. Usually one
pair of dorsal eyes present on cephalothorax and groups of two to five lateral eyes.

These animals, which include burrowing, arboreal, and rupicolous mem-
bers, are usually characteristic of arid environments; however, habitat selection
as in the other arthropods is governed by ecological factors such as vegetation,
geology, and climate.

Their venom apparently consists of two types in which the one produces only
local reaction without systemic effects except in cases of hypersensitivity. This is
usually found in species with a thin metasoma and large flat hands on the pedi-
palpi such as the yellow Cape scorpion, Opisthophthalmus capensis Herbst (fam-
ily Scorpionidae), a species that was periodically observed under large cow-pats.

In the family Buthidae, however, the venom is a neurotoxin, comparable to
that of the colubrine snakes, and may be lethal. In this family the scorpions have
a thick metasoma and the hands of the pedipalpi are ovoid, slender, and rather
small. Only one large species, Parabuthus capensis Hemprich & Ehrenberg,
rarely occurred under dry cow-pats during the surveys and was found to prey on
the smaller and more common Uroplectes variegatus. The latter and Parabuthus
brevimanus (Thorell), the smallest species in the genus, are fairly common
under dry cow-pats all along the west coast where surveys were made.

Family Buthidae
Uroplectes variegatus (Koch)

Full-grown females (32-45 mm long) (Fig. 8G) vary from pale cadmium yel-
low to yellowish brown; some are mottled with brown, with three rather indis-
tinct longitudinal stripes over middle of dorsum; most of them, however,

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

have only a single broken, median line and body segments are marked with light
brown. In all the specimens examined, the largest part of the fifth metasomal
segment is darker in colour than the rest; in some specimens it is piceous brown,
in others very slightly darker. It is rather similar to Parabuthus brevimanus both
in shape and size, but differs from it by the longitudinal body stripes, P. brevi-
manus being of a more uniform coloration.

This species was found inland along the west coast throughout the year,
usually only one specimen per dry cow-pat. Sometimes six out of every ten pats
examined harboured at least one specimen during the summer months; these
scorpions varied from very young (about 12 mm total length) to full-grown fe-
males. In the case of the full-grown specimens, the under surface of the cow-pats
was hollowed out in the form of a chamber in which the animals sheltered.

In the Karoo near Laingsburg, where collections were made during the au-
tumn, U. variegatus was replaced by pale yellowish-brown immature specimens
of U. schlechteri Purcell, which also has a dark fifth metasomal segment; the tail
segments, however, are more slender and longer than in U. variegatus.

Adults of U. variegatus living in association with the pugnacious ant, Ano-
plolepis custodiens (Smith), in the Citrusdal district were found by Eastwood
(1978) to be parasitized by the trombidiform mite Pimeliaphilus isometri Cun-
liffe, which is also found on scorpions in the Philippine Islands (Baker & Whar-
ton 1952). Eastwood regarded this South African scorpion as U. carinatus but,
according to Lamoral (1979), this species occurs only in the northern regions of
the Cape; U. variegatus, on the other hand, is confined to the north-western and
south-western Cape. It is interesting to note that a similar mite Pimeliaphilus
cunliffei Jack, in the U.S.A., uses the cockroach Periplaneta americana (Lin-
naeus) as its natural host (Cunliffe 1952; Jack 1961).

Pregnant females of this species and of Parabuthus brevimanus were ob-
served during September to December, and newborn young were actually found
during the middle of December to the middle of February. Pregnant dissected
specimens of U. variegatus yielded fourteen to twenty embryos of which most
were situated on the sides of the mesosoma. The fully-developed bean-shaped
embryo (Fig. 8C) is enveloped in a transparent membrane and has the meta-
soma folded in under the body. According to observations made, the gestation
period is fairly long, probably lasting a year. Newly emerged young are pure
white with black eyes (Fig. 8B) and an inconspicuous, longitudinal median line
on the dorsum of the mesosoma; the body is devoid of setae except for a few
setae on the pedipalpi (particularly on the fingers) and two or three setae on the
last two apical segments of the legs.

After birth the young immediately climb on to the mother’s back and after
a few days the penultimate metasomal segment becomes pale purplish. Two
days after birth they measure 7,2 mm from the tip of mouth-parts to the tip of
the sting. The developing mouth-parts, claws and sting are still enveloped in the
membranous covering during this stage and are freed only at the first moult,
which occurs 10 days after birth. At this stage the larvae are almost 10 mm long

(=)

Length in mm

SOUTH AFRICAN ARTHROPODS al

Porcellio scaber

30 80 130 180
Number of days

Fig. 8. A-C, G. Uroplectes variegatus. A. Eleven days after birth. B. Two days after birth.

C. Fully developed embryo in adult female. G. Young female. D, F. Porcellio scaber.

D. Adult. F. Increase in body length of the larvae of P. scaber as observed in the laboratory.
E. Armadillidium vulgare (adult).

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Fig. 8A) and, although still very pale, the penultimate metasomal segment is
clearly darker than the rest and the longitudinal median line on the mesosoma is
much more conspicuous. The apical metasomal segment is pure white and the
apical half of the sting brown. The legs are almost white and at this stage some
of the young already begin to leave the mother, although they still cluster to-
gether on the ground. Body hairs are much more abundant, particularly on the
pedipalpi and metasoma; they now clearly resemble the adult, except for the
much narrower mesosoma.

CLASS CRUSTACEA

Mostly aquatic arthropods with calcareous exoskeletons in contrast to the
chitinous covering of insects; with two pairs of antennae and usually five pairs of
legs.

The majority are free-living. They may be vegetarians, predators, or scavy-
engers.

Destruction of stranded kelp by arthropods is mainly brought about by the
large numbers of halophilous crustaceans, both Isopoda and Amphipoda. Their
presence on the beach is indicated by numerous tunnels in the stems of decaying
plants; the crustaceans and large kelp-flies undoubtedly form the most important
part of the wrack fauna that was examined.

ORDER AMPHIPODA

Body usually compressed, thoracic limbs without exopodites and with first
pair modified as maxillipeds.

Apart from Orchestia gammarella Pallas, very few species seem to be eu-
coenic and some of the Talorchestia species observed were even found to breed
in semi-fresh cow-dung in the laboratory. At least four species of the more com-
mon amphipods were collected during the surveys, namely the Atlantic species
T. capensis Dana (Fig. 91), the above-mentioned widespread O. gammarella
(Fig. 9F), which is also found in North America, and the endemic 7. quadrispi-
nosa Barnard, which is known to prey on other beach-hoppers, including T. ca-
pensis (Branch & Branch 1981) and T. australis Barnard (Fig. 9G). All the
species of these light grey to whitish beach-hoppers or sand-fleas are very similar
and mostly scavenge between and just above the intertidal zone, although some
have been found rather far above the supratidal zone on the small dunes. From
the available data it seems as if T. australis and O. gammarella are more restrict-
ed to the southern parts of the Cape, while 7. capensis and T. quadrispinosa are
more widely distributed along the west coast.

The indigenous Talitrus eastwoodae (Methuen) (Fig. 9H), which is very
similar to the beach-fleas but has reddish transverse bands over its body, is a
scavenger confined to forests (Lawrence 1952). It was found in rather large num-
bers around Cape Town in the vicinity of Table Mountain at carcasses lying in
shady surroundings. In the Cape Peninsula it is often found under carpets in
houses during the warmer parts of the year.

SOUTH AFRICAN ARTHROPODS 79

Fig. 9. A. Deto echinata. B. Tylos capensis. C. Marioniscus spatulifrons. D. Ligia dilatata.
E. Bethalus sp. F. Orchestia gammarella._ G. Talorchestia australis. H. Talitrus eastwoodae.
I. Talorchestia capensis.

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

ORDER ISOPODA

A large, variable group with depressed bodies. Thoracic limbs also without
exopodites and first pair always modified as maxillipeds.

The majority are aquatic, but the oniscoid group has adapted to terrestrial
or semi-terrestrial life.

The largest species present under kelp is Ty/os (length up to 50 mm), of
which there are two species, T. capensis Krauss (Fig. 9B), occurring from the
Cape Peninsula eastwards along the coast, and T. granulatus Krauss with a west-
erly distribution. The biggest difference between the two seems to be the texture
of the integument, which is granulate in the case of 7. granulatus and minutely
granulose in 7. capensis (Barnard 1932); their distribution and morphological
similarity, however, seem to indicate that one may rank as a subspecies. Accord-
ing to Kensley (1974), both species are omnivorous but with a bias towards a
herbivorous diet.

Most of the isopods are rather slow-moving, except Ligia species, which are
also characterized by the long antennae and uropods. The common greenish-
brown to olive-grey sea cockroach or shoreslater, L. dilatata Brandt (Fig. 9D), is
widely distributed along the south and west coasts. When collections were made
in wrack during the winter months, large numbers of this isopod were found to
carry small, round, yellowish eggs (0,92—0,76 mm diameter) in their marsupia.

Deto echinata Guerin (Fig. 9A), slate to greyish green with lighter specks, is
also widely distributed from the Namaqualand coast to Cape Agulhas. Large
numbers (some with eggs) were usually present under the kelp during the win-
ter, and very often they were found in seal carcasses in the intertidal zone. They
were often found to prey on other arthropods. The slightly smaller Marioniscus
spatulifrons Barnard (Fig. 9C), of similar coloration, has more or less the same
distribution as Deto echinata, and was usually in association with the latter. Be-
thalus species (Fig. 9E) are darker orange in colour and were mostly found dur-
ing the winter under semi-dry wrack strings along the south coast.

Family Oniscidae
Porcellio scaber (Latreille)

Full-grown females (Fig. 8D) are 15-16 mm long, with a brownish-grey
colour, mottled with pale or creamy white; colour variation, however, is consid-
erable. It cannot roll up into a ball and its uropods are much longer than those
of Armadillidium vulgare (Latreille) (family Armadillidiidae, see below). Males
somewhat more slender with longer uropods.

This species is widely distributed in South Africa and is found in many
places in the world (Barnard 1932).

Adult sow-bugs collected during the first week in September from soil con-
taining decaying carcasses moulted about 10 days later, and after a month pro-
duced about 50 larvae. The latter were kept under observation in the laboratory
and were found to produce larvae 194-200 days after birth. The increase in size

SOUTH AFRICAN ARTHROPODS 81

of the larvae with age is shown graphically in Figure 8F. The average monthly
temperature during the day during the period of observation varied from 21° to
29 °C in September to March and from 18° to 24 °C in April and May.

Young larvae of about 2 mm in length are nearly white with blackish eye-
spots. As their size increases, the colour becomes darker and after about 20 days
they are already pale brownish grey and inconspicuously mottled with dirty
white. After about 80 days they look exactly like their parents and are fairly
dark brownish grey, mottled with creamy white. According to Heeley (1941),
who gives a good account of their biology in England, the development of the
young may be regarded as complete by about the fifth moult.

This eurytopic sow-bug is omnivorous and was found to feed on various ma-
terials, including decaying kelp. It is very common under loose bark (Heeley
1941, Cloudsley-Thompson 1957). It is often found in large numbers under fresh
and semi-fresh cow-pats and plays a vital part in the destruction of the remaining
parts of the dung, especially when the number of fly maggots begins to decrease.
It is sometimes attracted to rotting carcasses and may remain throughout the de-
caying process.

Family Armadillidiidae
Armadillidium vulgare (Latreille)

Adult females (Fig. 8E) about 14,6 mm long and dark slate grey in colour,
mottled with paler spots similar to those of Porcellio scaber, but forming a defi-
nite, inconspicuous, pale longitudinal band over the middle of the dorsum.

This species is very widely distributed in South Africa as well as other parts
of the world (Barnard 1932).

Its biology is very similar to that of P. scaber, but A. vulgare seems to pro-
duce more young than the latter. However, in specimens observed during the
surveys the survival rate of the very young larvae was much lower than in the
case of P. scaber. Young larvae about 1,7 mm long, collected during January
near a turtle carcass, were (as in the latter species) almost pure white with black
eyes. After about a month the body already showed four brownish, longitudinal
lines and the colour gradually darkened until maturity was reached. The growth
of specimens kept under conditions similar to those of P. scaber was, however,
much slower and 200 days after birth some larvae were still only 5 mm long.

This pill bug is mainly omnivorous and is common around houses; it was
abundantly collected in leaf litter and compost heaps all over the Cape Peninsula.
Both Heeley (1941) and Paris (1963) give valuable information on the life cycle of
this species, which is considered by Brereton (1957) to prefer the shade of stones
both in open land and in woodlands. It often occurs in very large numbers
together with P. scaber under semi-fresh to fresh cow-pats, and is also attracted to
decaying carcasses. Albino forms, which are often present among individuals of
this species, point to a calcium deficiency in the soil. They are highly dependent
on humidity and are, therefore, always found in locations that are fairly damp.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

CLASS INSECTA
ORDER BLATTODEA

Family Blattidae

Dorsoventrally compressed exopterygote hexapods with cursorial legs, well-
developed cerci and multi-segmented antennae. Wings usually well developed,
but some are apterous. A few species are sub-social, some are gregarious, and
they may be oviparous, Ovoviviparous, or viviparous; an ootheca of a tough
leathery substance is produced during oviposition.

Although they are usually omnivorous, there is evidence that the Table
Mountain cockroach, Aptera fusca (Thunberg), is a plant-feeding species as it
was observed to feed on the berries of a Cuscuta sp. (Skaife 1979). In the Cape
Peninsula it was sometimes observed under stranded kelp on the intertidal zone,
but repeated efforts to feed the cockroach on this medium were in vain. It often
congregates in familial groups under stones and dry cow-pats as is the case with
other veld species such as Deropeltis erythrocephala (Fabricius) (Fig. 10J).

Of the oviparous species, the American cockroach Periplaneta americana
(Linnaeus) drops the ootheca long before the eggs hatch. This is also the case
with the indigenous Pseudoderopeltis foveolata (Walker), whereas the female
of the German cockroach Blatella germanica (Linnaeus), on the other hand,
carries the ootheca externally until shortly before the eggs hatch. This was also
observed in some specimens of the short-winged mountain cockroach, Tem-
nopteryx phalerata (Saussure), although evidence seems to indicate that this
species behave rather like Periplaneta americana. The Madeira cockroach
Leucophaea maderae (Fabricius), which occurs in many parts of the world and is
widespread in the Subsaharan region (Cornwell 1968), including Natal and
probably also Transvaal, is ovoviviparous. Like D. erythrocephala, it is gre-
garious and may form large colonies outdoors and, according to Scharrer (1951),
in Brazil twenty-five to thirty-two young are produced at one time during the
warm weather. It has been found in sugar-cane on certain North Atlantic
islands.

For many years it has been suspected that cockroaches carry pathogens such
as Mycobacterium tuberculosis (Schroeter) (observed in the faeces of Blatella
germanica), Salmonella typhimurium (Loeffler) (gastro-enteritis), and Chlostri-
dium perfringens (Veillon & Zuber) (gaseous gangrene and one of the causative
organisms of normal decay in carcasses and cadavers), as well as various other
organisms including viruses (Roth & Willis 1957, 1960). Even the organism that
causes bubonic plague, Yersinia pestis (Lehmann & Neumann), has been found
in specimens of the oriental cockroach or black-beetle, Blatta orientalis Lin-
naeus, collected in infected areas (Roth & Willis 1957). The natural vectors of
plague are fleas (Siphonaptera). Cockroaches are also capable of causing allergic
dermatitis (Smith 1973).

Parasitoids include wasps of the genus Tetrastichus, reared from the oothe-
cae and sphecids of the genera Ampulex and Dolichurus, which provision their

SOUTH AFRICAN ARTHROPODS 83

young with cockroaches (Arnold 1928; Roth & Willis 1960). Mites have already
been mentioned (see discussion on Arachnida).

Pseudoderopeltis foveolata (Walker)

Princis (1963) has described the adults (Fig. 10A). Females black and fairly
shiny, with vestigial mesothoracic wings. Depressed sixth and seventh tergites
are characteristic of this genus. Cerci ten- to twelve-segmented in all females
collected and fairly broad and compressed dorsoventrally; penultimate segment
in most cases the longest. Antennae multisegmented (at least sixty to seventy
segments).

This species is widely distributed in South Africa, Zimbabwe and Lesotho
(Princis 1963). It was collected in the interior under semi-dry cow-pats together
with other Pseudoderopeltis spp. (Fig. 11E) and Perisphaeria spp. (Fig. 11F—G),
whereas cow-pats on the beach or on the sand-dune biotope produced mainly
the psammophylic cockroaches Blepharodera discoidalis (Brunner) (Fig. 11D)
and B. ciliata Burmeister (Fig. 10F). B. discoidalis is a blackish species whose
body is surrounded by a pale yellowish white margin; B. ciliata is mottled with
brown; the adult females of the latter measure up to 30 mm in length and males
up to 25 mm with a wing-span of 65 mm; nymphs are mottled with grey and re-
semble the apterous females, except for size. Both species are usually found
under the sand near the stems of plants and both adults and nymphs have been
collected on the beach under the stems of Tetragonia decumbens and Arctotheca
populifolia during October to December. The Perisphaeria spp. vary from pale
light brown, mottled with darker brown, to almost black.

Oothecae of P. foveolata (Fig. 10D), produced from December to January
by adults collected along the west coast, are 7,9-8,8 mm long and are all about
4.9 mm broad. Initially the ootheca is reddish white, but soon darkens to a pale
brownish red and eventually to a red-brown, with the keel somewhat paler. The
dorsal serrations are rounded, not acute. In this species the ootheca is held
upright while the eggs are being deposited, with the keel fitting in the V-shaped
groove in the tenth tergum (Fig. 10B). The ootheca is dropped as soon as all the
eggs have been laid and buried about 5 mm deep in the soil. The incubation
period varies from 47 to 56 days.

The newly hatched young (Fig. 10C) are pale piceous to yellowish brown in
colour, 3,3—3,7 mm long, and smooth and shiny. In all the specimens examined
the cerci are three-segmented as in Periplaneta americana (Guthrie & Tindall
1968) and the antennae twenty-two-segmented. The sixth and seventh tergites
are already depressed in the first instar. The young nymphs remain clustered to-
gether on the ootheca for at least 4-5 days, during which time they turn almost
black.

The first moult was observed 17 days after hatching and by this time they
were about 5,5 mm long. After moulting they are pale whitish brown with al-
most black eyes. In all the second instar nymphs the cerci are six-segmented,

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

with the penultimate segment the longest. The antennae are twenty-five- to
twenty-six-segmented. In the laboratory development was slow and after about a
month the nymphs were only about 6 mm long, the antennae consisted of
twenty-seven to thirty segments and the cerci were still six-segmented. After
about 120 days their length had increased to 10 mm, the cerci were now seven-
to eight-segmented and the antennae thirty-five-segmented. Nymphal growth
over a period of about 130 days is shown graphically in Figure 10E. Some speci-
mens reached maturity only 3-6 months later.

Temnopteryx phalerata (Saussure)

Princis (1963) has given a detailed description of this species. General body
colour of adult (Fig. 10G) light yellowish brown, abdominal segments darker
brown, posterior borders of pronotum and abdominal segments pale yellow.
Legs and antennae yellowish brown, latter multisegmented (more than seventy
segments). Cerci twelve-segmented, measuring about 3,7 mm in adult females.
Total length of body 18-25 mm. A peculiarity of this species is the uneven num-
ber of tarsal segments on the hind legs; out of twenty-five specimens examined,
with only one exception, the hind tarsi on the right side had four segments while
those on the left had five.

According to available collecting data it is endemic to the Cape where it is
widespread along the west and south coasts. This cockroach, which is often ob-
served in leaf litter both on the coastal sand-dune area and in the interior, has
on several occasions been found to feed on certain types of carpets in the Cape
Peninsula; damage reported so far, however, was apparently only very slight.

Whereas in Pseudoderopeltis foveolata the ootheca is always held upright
during oviposition, in this species it is held flat (Fig. 10G) as in the case of the
German cockroach, with the keel in all the specimens examined pointing to the
right side. The oothecae (Fig. 101) are 10,8—15,4 mm long and 3,5-3,7 mm wide
and most of them are dark brown to dark reddish brown in colour, including the
keel of which the serrations are clearly dentate and close together. Oothecae of
this species are very similar to those of Deropeltis erythrocephala (Fig. 10K),
common under cow-pats, but in Deropeltis spp. the serrations resemble those of
Pseudoderopeltis foveolata and are lighter in colour. In Temnopteryx phalerata
the ootheca in most specimens examined was retained by the female for a day
before being buried under 1-2 mm of sand. The incubation period varied from
32 to 39 days during February at a temperature of 30—31°C. In a few specimens
the ootheca was carried for a longer period before being dropped.

Newly hatched nymphs measure 3-3,2 mm in length and are pale yellowish
brown with two white oval spots on the metanotum; the eyes are reddish. The
nymphs soon turn almost black (Fig. 10H) but the posterior borders of the tho-
racic segments. and the first three abdominal segments, as well as the two meta-
notal spots, remain white. A thin white line also runs over the head and thorax.
The legs are piceous with the apical third of the middle and hind femora, as well

SOUTH AFRICAN ARTHROPODS 85

€ 20 40 60 80 100 120
Number of days

mm

Fig. 10. A-E. Pseudoderopeltis foveolata. A. Female (adult). B. Female (left lateral view)

to show oviposition and ootheca (0). C. Newly emerged larva. D. Ootheca. E. Body in-

crease of larvae as observed in the laboratory. F. Blepharodera ciliata (adult). G—I. Tem-

nopteryx phalerata. G. Female with ootheca (o). H. Larva at about 30 days. I. Ootheca.
J—K. Deropeltis erythrocephala. J. Female (adult). K. Ootheca.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

as the tarsi and the apical half of all the coxae, white. The antennae are twenty-
two-segmented, with the second to fourth, the thirteenth and about one-third of
the twelfth and fourteenth segments white in some specimens. In many speci-
mens the thirteenth to the fifteenth segments, as well as the second to fourth,
are also white, the remainder black. The cerci are piceous and three-segmented
as in the other species. In the first instar the tarsi are five-segmented in all the
legs.

Development was very slow and after about 161 days the nymphs were only
about 5 mm long and coloured as shown in Figure 10H. The antennae had
thirty-three to thirty-four segments, with the twenty-first and twenty-second seg-
ments white; the cerci were six-segmented.

Unfortunately, due to an invasion of mites and the excessive use of the in-
secticide Carbaryl to combat ants, most of the specimens died before maturity
was reached.

ORDER MANTODEA
Family Mantidae

Elongate predacious exopterygote hexapods, usually with well-developed
cerci, large eyes and raptorial forelegs. Antennae multi-segmented and prono-
tum long; wings usually well developed, absent in some females. Solitary insects
producing oothecae of a tough material. Subsociality is known to occur in some
species.

Oxypilus nasutus (Fabricius)

Full-grown females (Fig. 11B) 17-20 mm long, apterous and mottled with
dark brown or dark grey and yellowish white. Very striking light coloration on
ventral side of the abdomen and on front femora. As in most mantids, front
femora spined on ventral side, with five spines on exterior margin, of which
basal ones are largest. The three discoidal spines (Fig. 11B) large and agree with
those described by Loxton & Nicholls (1979) for Hierodula membranacea (Bur-
meister), suggesting that this is a generalized predator. Front coxae also have six
to seven spines on anterior margin. Males similar to females, but more slender,
with a wing-span of 45-50 mm.

This species is widely distributed in South Africa and South West Africa,
and is fairly common along the west coast, particularly in Namaqualand. It ap-
peared occasionally in the vicinity of Saldanha Bay under plants growing on the
edge of the fore-dune system adjoining the beach. It is very well camouflaged
against the debris and remains of plant seeds that are found under plants such as
Arctotheca populifolia and is therefore very rarely seen. Prey included small
mantids, blow-flies and other soft-bodied insects.

Oothecae (Fig. 11C) collected during December varied from 8 to 16 mm in
length and were glued to the seeds and to leaf litter under the plants. They are
whitish when freshly produced, but after about a day become reddish brown to

SOUTH AFRICAN ARTHROPODS 87

)

"ee
m
z

Say
7a,
HT ap,

nln y \

Fit jay

10

mm

Fig. 11. A-C. Oxypilus nasutus. A. Newly hatched young. B. Adult showing discoidal
spines (ds). C. Egg packet (lateral view). D. Blepharodera discoidalis. _E. Pseudoderopel-
tis sp. FF. Perisphaeriasp. GG. Perisphaeriasp. H. Penicillata, sp. indet.

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

dark brown with the dorsal edge yellowish white. The incubation period of the
eggs was about 45 days.

The newly hatched mantids (Fig. 11A) are approximately 4,2 mm long,
mottled with brown and pale yellowish white, except the last four or five abdo-
minal segments, which are pale, almost white in colour. The eyes are large and
pale brown. Almost the whole of the pronotum and the tibiae as well as the api-
cal third of the front femora are white. As in the case of the adults, there are
five spines on the exteroventral margin of the front femora, but the anterior
margins of the front coxae are devoid of any spines. Exteroventral margin of
front tibiae has only one strong spine.

In larger nymphs (8,5 mm long) taken during January and which exhibit the
coloration of the adult, the anterior margins of the front coxae possess six to
seven spines and the exteroventral margin of the front tibiae at least two devel-
oped spines and smaller dentations indicating the position of five more spines
present in the adult form.

ORDER DERMAPTERA

Elongate exopterygote hexapods, cerci modified into forceps. Legs cursor-
ial, antennae fairly long with variable number of segments. Forewings reduced
to tegmina, hind wings membraneous, sometimes lacking. Mostly omnivorous;
however, species of the family Hemimeridae are parasitic, such as Hemimerus
talpoides Walker, living in the soft fur of the large African pouched rat, Criceto-
mys gambianus Waterhouse, and feeding on parts of the skin, fungus spores and
other debris (Rehn & Rehn 1935, Walker 1964). Subsociality and viviparity are
known to occur in some earwigs.

Only a few species were collected during the surveys, mainly under cow-
pats, of which Labidura riparia (Pallas) and an unidentified species were the
most abundant. The latter, a brown earwig with pale yellowish stripes and mark-
ings giving it a mottled appearance, was found to be very common in dry pats
near Saldanha Bay. Only wingless specimens, 11-16 mm in length (forceps in-
cluded), with dark-brown heads and usually with two small, oval, lighter yellow
marks on the front, were collected throughout the year. They were found to
shelter in the crust and cake of the pat, their colour blending well with the sur-
roundings, muking them difficult to find. Their legs are pale, marked with
brownish patches and the antennae dark brown with the apical borders of seg-
ments 15 to 16 white.

Young larvae of this earwig measuring 5,2—7,8 mm in length had the same
colour as the adults, but were pale, sometimes pale yellowish and with the pos-
terior part of the body darker as in the adults. The number of antennal segments
varied in the immature stages from eleven (5 mm specimens) to fourteen (7 mm
specimens) and the yellowish patches seemed to be absent on the head. These
nymphs and their exuviae were observed in the pats during both summer and
winter.

SOUTH AFRICAN ARTHROPODS 89

One light-brown specimen of the genus Esphalmenus was collected under
an almost dry cow-pat in open sandveld, near the sand-dune system in the vicin-
ity of Elands Bay during April. It is about 15 mm long (including the forceps)
and differs from EF. peringueyi (Bormans), E. capensis Brindle and E. ecarinatus
Brindle, the only three other species of this genus described from South Africa,
by having a strong tooth on the last tergite, just in front of the base of each
branch of the forceps; the latter also carries a large tooth. Nymphs of the pyr-
rhocorid Scantius forsteri (Linnaeus), sheltered under the same pat.

Family Carcinophoridae
Euborellia annulipes (Lucas)

This earwig is always apterous (Fig. 12A). Females collected are dark red-
dish brown to piceous brown, but the pronotum (and in some specimens also the
meso- and metanotum) is usually paler in colour. Antennae have sixteen to
eighteen segments in most females examined, with fourteenth or thirteenth to
fifteenth white. Forceps more or less straight. Males similar to females, except
that forceps are more curved, particularly on right side (Fig. 12C). Antennae
consist of fourteen to sixteen segments, with twelfth or twelfth to fourteenth
white. Hincks (1947) and Brindle (1978) give good descriptions of males and fe-
males.

This species 1s cosmopolitan and widely distributed in Africa and is the most
widely distributed earwig (Brindle 1978). It is very common in the Cape Penin-
sula and is often attracted to decaying carcasses, feeding on the organic material
and on insects and other arthropods present under such conditions. It is a noc-
turnal species, attracted to houses by lights and, according to observations, it is
thigmotactic as in the case of Labidura riparia. It was never encountered on the
beach or on the dune system during the surveys.

Although this species appeared during the last post-mortem stage of decay
when the surveys were made, it may, depending on various factors such as
weather conditions, location of the carcass and the species of arthropods pres-
ent, be attracted at a very early stage, as is the case with ants, certain beetles,
arachnids, and chilopods.

Males and females were found to copulate during October. When mating
the male and female face away from each other and the male’s abdomen is
twisted so that the genitalia on the ventral side are brought into contact with
those of the female. Copulation in most cases lasted for about 2 minutes and
most females produced a batch of 44-50 eggs about 18 days afterwards. Eggs
were also found during the late summer and early autumn.

The eggs (Fig. 12B) are spherical, shiny white and almost without any
sculpturing. They vary in size from 0,88 X 0,76 mm to 1,06 X 0,90 mm and are
laid in a ground cell excavated by the female, who remains in the cell with the
eggs. When disturbed the female carries the eggs away to a new excavation, or
else devours them. The incubation period is at least 16 days.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

The newly hatched nymph is pure white with brown eyes and about 3,4 mm
long including the forceps, which measure 0,68 mm in the specimens examined.
The antennae at this stage consist of eight segments. In about 3 days’ time they
turn to a pale yellowish brown, the head and posterior three or four abdominal
segments becoming somewhat darker. The legs remain whitish, with a darker
transverse band at about the middle of the femora. The penultimate antennal
segment is also white (Fig. 12D).

At the age of 19 days the number of antennal segments have increased to
eleven (after which stage the first moult apparently occurs) with the penultimate
or, in some cases, the ninth segment remaining white. The general body colour
is yellowish brown, with the head somewhat darker and the legs paler. About 44
days after hatching another moult takes place and the antennae consist of thir-
teen to fourteen segments, with the eleventh segment white. The body length is
then 6,5—7,0 mm. The nymphs were observed to moult again after about 64 days
from the time of hatching and the antennae were then found to consists of fif-
teen to seventeen segments in most cases. In the laboratory the developing
nymphs required about five moults to reach maturity, but some specimens
moulted only four times. The size increase over a period of about 150 days is
shown graphically in Figure 12E. According to Hincks (1947) there are five
instars and the third instar has thirteen-segmented and the fourth instar four-
teen-segmented antennae.

Family Forficulidae
Forficula peringueyi (Burr)

Females (Fig. 12F) measure about 10 mm, forceps included; reddish brown
in colour, with the posterior five segments almost black. Males fairly similar to
females and measure 11—11,6 mm, forceps excluded; latter long, in most cases
measuring about 7,5 mm (Fig. 12G). In both sexes antennae are eleven-seg-
mented. Characteristic of both males and females are the lateral tubercles on
third and fourth abdominal tergites, those on fourth being the largest. Brindle
(1973) gives a description of the sexes.

This species is endemic to the Cape and, according to Brindle (1973), re-
stricted to the western parts where it is common under debris and stones; it has
been found to nest in cow-pats during the winter. The immature stages of this
species look very much like those of F. promontorii (Burr), which was mainly
observed along the south coast, but they seem to be more shiny.

Eggs (Fig. 12K) were found in hollowed-out chambers in fairly dry cow-pats
during July. They are oval and measure 1,2 X 0,72 mm; most of them are shiny,
creamy white to whitish yellow and without any form of sculpture. As in the
case of Euborellia annulipes, the females usually tend the eggs.

Newly emerged nymphs (Fig. 12L) measure about 3,7 mm including the
forceps, and are pure white but soon turn to a pale brown, the head and poster-
ior two to three segments being somewhat darker. The antennae are eight-seg-
mented and of the same colour as the body; the forceps are fairly long and

SOUTH AFRICAN ARTHROPODS 91

Number of days

Fig. 12. A-E. Euborellia annulipes. A. Adult. B. Eggs. C. Forceps of male. D. Nymph

at 3 days. E. Increase in body length of nymphs as observed in the laboratory. F—G, K-L.

Forficula peringueyi. F. Adult female. G. Forceps of male. K. Eggs. L. Newly hatched
nymph. H-J. Labidura riparia. H. Adult. I. Eggs. J. Newly hatched nymph.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

measure 1,1 mm. Observations show that the nymphs require 4-5 months to
reach maturity.

Family Labiduridae
Labidura riparia (Pallas)

This is a large species (Fig. 12H), specimens examined measuring
20-26 mm in length, including the forceps. Usually dark reddish brown, with
central area of abdomen darker, in the shape of two longitudinal, median bands;
pro- and mesonotum also with two darker stripes. Antennae and legs yellowish.
Antennae consist of more than twenty segments, with fourth to about seventh
shorter than the others. Brindle (1973) described both sexes.

This earwig is cosmopolitan and common in Africa, particularly along the
shore (Brindle 1973). It is mainly a predator and generally feeds on the smaller
- staphylinids and scarabs and their larvae as well as on other beetles such as Go-
nocephalum spp. It is attracted to light and is therefore often seen in houses dur-
ing the night. It is sometimes also attracted to semi-fresh cow-dung on the
beach. According to Callan (1964) the pale sand-coloured ecotype is a maritime
form restricted to sandy habitats of the coast. During this survey, however, only
the dark ecotype was found both on the beach and inland. Moderately pale
forms occurred only in the young nymphal stages.

In certain areas along the west coast it was seen to visit dried-out sea-birds
on the high-water mark and was found to feed on other arthropods, particularly
fly larvae, and on bits and pieces of the flesh still left on the skeletons. Near
Elands Bay it was observed in large numbers together with the cicindelid Platy-
chila pallida Fabricius in the top few millimetres of sand under the plant Arcto-
theca populifolia in the supratidal zone; they were associated with the larvae of
the beetle Melyris viridis Fabricius, on which both predators fed. This earwig is
very common under dry cow-pats in sandy areas.

The yellowish-white, shiny eggs (Fig. 121) were collected under semi-fresh
cow-pats during March. They measure 1,1 x 0,96=1,8 x 1,2 mm, vary from
nearly round to oval, and are without any significant sculpture.

The newly hatched nymphs (Fig. 12J) are 3,4 mm long (excluding the for-
ceps) and almost white with pale brownish blotches, particularly on the abdo-
men. As in the two previously mentioned species, a white, longitudinal median
line, representing the posterior part of the ecdysial suture, traverses the thorax.
It is connected anteriorly to the so-called ‘epicranial suture’ of the head. The an-
tennae are brownish and eight-segmented and the eyes brown. Most nymphs
reached a length of 6 mm about 70 days after hatching.

Nymphs 8,7-10 mm long (including the forceps) collected during March,
are darker in colour than newly hatched specimens. The antennae are fifteen-
segmented, with the five apical segments much longer than the others, except
the first and third segments. The eyes are almost black and the forceps measure
about 2,3 mm.

Oe

SOUTH AFRICAN ARTHROPODS 93

In larger nymphs (16,6-17 mm long including the forceps) the colour re-
mains the same, but the stripes on the abdomen are darker. The antennae are
then twenty-segmented, the last ten segments being somewhat longer than the
others, except the first and third segments, which are the longest. Wings are
already partially developed and partly cover the first abdominal segment.

As in the other species, the forceps are used to crush the prey, which is then
lifted over the body and devoured. The nymphs were found to feed on small
scarabaeid larvae and for this purpose they often dug holes in the soil with their
mandibles.

ORDER ORTHOPTERA
Family Gryllidae

Exopterygote hexapods with well-developed cerci, and short antennae with
few segments to long and multi-segmented. Hind legs usually saltatorial, fore-
legs often fossorial and in certain species with tympani. Wings well developed or
absent. Usually omnivorous, but a few species are phytophagous, causing dam-
age to crops and lawns; under certain circumstances they may even become pre-
dacious or cannibalistic. Some sphecids of the genera Tachysphex, Liris,
Chlorion, and Isodontia, and probably also Gasterocericus, prey on crickets (Ar-
nold 1922, 1923, 1928; Bohart & Menke 1976).

Of the somewhat thirty-five species of true crickets (family Gryllidae) that
have been collected in southern Africa, only three were observed in decaying
matter during the surveys, of which the common field cricket, Gryllus bimacula-
tus de Geer, was the most abundant and most widespread. It often shelters
under stranded kelp on the intertidal zone and when reared in the laboratory oc-
casionally accepted pieces of wrack, though it preferred fish-meal and grass cut-
tings. According to Smit (1964) it can be a pest of the vegetable and flower
garden.

Cophogryllus delalandi Saussure

A wingless species (Fig. 13C—D). Ground colour pale creamy white to pale
whitish yellow, with dark-brown marks on dorsal side of body. Cerci, antennae
and ovipositor brownish; the ovipositor being 11-11,6 mm long in mature
females (Fig. 13E). Body length of adult females collected along the west coast
vary from 14,9 to 18,3 mm; males measure about 13,7 mm.

According to Chopard (1955) C. delalandi is widely distributed in the west-
ern and southern Cape Province and has been collected as far east as Mossel
Bay. It was found to be a common prey of the baboon-spider (Harpactira sp.).

Adults were collected almost throughout the year along the west coast
under stones and dry to semi-fresh cow-pats. During the summer and early win-
ter large numbers were often observed, in some cases up to six gravid females
under a single small pat. Eggs (Fig. 13F) were found under the sand under
semi-dry pats during November. These measure from 2,50 x 0,74 mm

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

to 2,7 X 0,91 mm and are fairly dull (only slightly shiny), with a fine superficial
reticulation and varying in colour from creamy white to a dirty brownish white.
Females with eggs were also noticed during April. It was impossible to deter-
mine the incubation period of the eggs, but in a few cases eggs laid during July
hatched only in November. Just before the young cricket emerges the two red-
dish eyes, the mandibles, and the segmentation of the body become visible
through the chorion.

Nymphs that hatched during November (Fig. 13A) measured about 2,5 mm
long and were almost pure white with dark brownish eyes. After about 4 hours
they usually turned to a pale grey colour (Fig. 13B) and after about 11 days the
colour became quite dark. A thin pale longitudinal line over the thorax rep-
resents the ecdysial suture and anteriorly joins the epicranial suture. The legs
are pale except for a dark transverse band near the apices of the femora. As the
nymph increases in size, its body becomes mottled with brown. The size increase
in the laboratory over a period of about 140 days is shown graphically in Figure
13G. Although growth was fairly fast during the first 100 days (the nymphs
reaching a length of about 10 mm in March) some specimens matured only dur-
ing midwinter.

In the females examined, rudiments of the ovipositor appeared about 103
days after hatching and within 50 days had reached a length of nearly 4,6 mm.
At this stage the nymphs are all found in small burrows in the loose sand and in
the laboratory they have to be kept in separate jars to prevent predation. In
field collections made during February large numbers of young females,
11-11,6 mm in length, were found, their ovipositors being only 1,8 mm long; in
specimens collected during June the ovipositors had already reached a length of
5,5-6,1 mm.

Cophogryllus sp.

This species is also wingless and somewhat larger than C. delalandi, females
(Fig. 14A) measuring from 16,6 to 20 mm in length are somewhat more robust.
The body marked with dark-brown stripes instead of the mottled coloration of
C. delalandi and ovipositor much longer, measuring 16,6—25,0 mm. Femora of
hind legs pale yellowish to creamish and not marked with dark coloration.

This species is widespread in the Cape and has been observed even in the
Great Karoo near Beaufort West. In the western Cape adults of this species
were collected under dry cow-pats in sandy areas during the late summer to
early winter, and eggs were collected in the sand under the pats during the latter
part of January.

The eggs (Fig. 14C) are somewhat larger than those of C. delalandi and
vary in size from 2,8 X 0,64 to 3,2 X 0,96 mm. They are yellowish to yellowish
white in colour, shiny and without any form of sculpture. The incubation period
is about 28 days and the newly emerged nymphs (Fig. 14B) are dirty white, al-
most transparent, with the last two abdominal segments and distal half of the
cerci somewhat orange. The head in all the specimens examined is of a pale

SOUTH AFRICAN ARTHROPODS 95

golden yellow, the eyes reddish. Most first-stage nymphs measure from 3 to
3,2 mm. About 4 hours after hatching they become piceous, but have reddish
eyes surrounded by a whitish margin; even the thirty-four-segmented antennae
and the cerci are piceous except for the hair bases, which are white. The borders
of the apical antennal segments also have an orange coloration. As in the pre-
vious species, the ecdysial suture forms a pale, longitudinal median line over the
body and is anteriorly continuous with the epicranial stem. The hind femora are
pale piceous, lacking a dark transverse band.

Characteristic of the nymphs of both this species and those of C. delalandi,
are the long, blackish median hairs on the thorax and posterior borders of the
abdominal segments. Nymphs of this species differ from those of C. delalandi in
that they construct a little burrow soon after hatching, into which they then re-
treat. The burrow was in most cases extended for several millimetres along the
surface, in a similar way to those made by golden moles.

The nymphs are usually very pale after each moult. In the specimens exam-
ined, the first moult occurred 9-10 days after the nymphs had hatched and most
of them at this stage measured about 3,3 mm, with the antennae in almost all
the specimens consisting of about forty-nine segments. After the first moult the
nymph already resembles the adult fairly closely in colour. Further moults were
observed at 44 and 45 days and again at 100 to 102 days after hatching. The first
rudiments of the developing ovipositor appeared after 202 to 207 days. In larger
nymphs (about 14 mm long) collected during November, the ovipositor was only
about 3,3 mm in length. In specimens kept under observation in the laboratory,
the ovipositor reached a length of 4,3 mm about 300 days after hatching. The in-
crease in size of the nymphs over a period of 280 days, as observed in the lab-
oratory, is shown graphically in Figure 13G.

Gryllus bimaculatus de Geer

In the common black field cricket the wings are well developed, with the
hind wings pointed in both sexes and extending beyond apex of abdomen (Fig.
14E). Males are easily recognized by the venation of the front wings (Fig. 14F)
and absence of an ovipositor; front wings pale fulvous in most specimens exam-
ined. The species differs from Cophogryllus spp. by the presence of a distinct
tympanum (Fig. 14J) on both sides of the front tibiae (absent on both sides in
Cophogryllus spp.—Fig. 14D). Wing bases pale yellow and broadly so in males,
but narrower in the case of females. Three distinct ocelli, reddish in colour, are
present.

Very often crickets with reduced hind legs are found. Crickets are aggress-
ive and when nymphs lose a leg in a skirmish, it is replaced at the following
moult by one that is smaller than usual.

_ This species is widely distributed in Asia and Europe and has been collected
all over South Africa and Zimbabwe, including some of the islands along the
west coast such as Bird Island near Lambert’s Bay.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

In the western Cape adults were observed under decaying kelp on the
beach and under semi-fresh to cry cow-pats, both in the winter and summer,
and eggs (Fig. 141) were collected under semi-dry pats throughout the summer
and autumn. In some cases batches of twenty eggs were laid in cracks in
the pats, in others eggs were deposited in the top 4 or 5mm of soil in
batches of fifteen or more; in a few instances, however, eggs were laid singly
in the sand. Those found near Philadelphia measure 2,5 X 0,56-2,8 x 0,76 mm
and are almost banana-shaped. The colour varies from a whitish yellow to
almost golden yellow, the eggs being fairly shiny and without any form of
sculpture.

As the embryo develops the egg becomes slightly darker and after a few
days the body segmentation, the two black eyes, the apices of the mandibles,
and the cerci become visible through the chorion; some eggs develop a dark ring
_ at the anterior pole.

In the laboratory the incubation period was 8-15 days during January. Riv-
nay & Ziv (1963) gave the mean incubation period in Israel as 7,5 days (34 °C)
to 75 days (16 °C).

The newly hatched nymphs are about 2,7 mm long and pale whitish, trans-
lucent, with blackish eyes; three ocelli are indicated by three black marks. They
remain quiescent for 3-4 hours, after which they become piceous (Fig. 14G)
with reddish eyes; they can then easily be distinguished from second instar
nymphs and nymphs of the other two species by the white mesonotum and pale,
almost white posterior half of the pronotum. The pale cerci are white at their
bases, with piceous apices. As in the other species, there is a white, longitudinal
median line over the back and the legs are also piceous. In the few specimens
examined, the antennae consisted of thirty-four segments.

The first moult occurred 7—22 days after hatching in specimens that hatched
during January-February and by this time they were about 4,9 mm long and
black all over (including the pro- and mesonotum), except for the pale, median
dorsal line, the brown eyes, pale brownish cerci, and the small whitish areas on
the sides of the pro- and mesonotum. The antennae consisted of about forty-
nine segments (Fig. 14H). A second moult occurred 40-48 days after hatching,
the nymphs remaining more or less of the same colour as in the second instar.
After each moult the colour is pale brownish white mottled with brownish, but
they assume their normal black colour in 2,5—4 hours.

Development was fairly rapid in the laboratory during the first 100 days or
so and most of the nymphs reached a length of 12,5-14 mm after 94-120 days,
depending on the amount of food available. The size increase during the first
120 days is shown graphically in Figure 13G. The ovipositor appeared 37-52
days after hatching.

As already stated, adults were found throughout the year and large num-
bers of nymphs collected during September at Elands Bay reached maturity dur-
ing that month. According to collection data it seems as if there was only one
generation annually during the surveys with overlapping of the different stages.

oF

SOUTH AFRICAN ARTHROPODS

:
1

ia
My

.

C. delalandei

i

nee

SS

sereseeeeeeeee Cophogryllus sp

ie Onl fen See) )

— « —G. bimaculatus

Length in mm

=>
SS
>

ro)
Now

0 40 80 120 160 200 240 280
Number of days

A. Newly hatched nymph. B. Eleven-day-old

Fig. 13. A-F. Cophogryllus delalandi.
nymph. C. Adult male (left lateral view). D. Adult male (dorsal view). E. Outline of adult

female. F. Eggs. G. Size increase of C. delalandi, Cophogryllus sp., and G. bimaculatus“as
observed in the laboratory over a period of 140 to 300 days.

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

coo
a

he
aN
ff oS
~ ei
Vos =
Sa

a

H
yy
Y
i

i

fi

i

H
ct
H
p

Fig. 14. A-D. Cophogryllus sp. A. Adult female. B. Newly hatched nymph. C. Eggs.

D. Left foreleg (posterior view). E-J. Gryllus bimaculatus. E. Adult female showing

pointed wing apices (wa). F. Adult male showing pointed wing apices. G. Newly hatched

nymph. H. Second instar nymph. I. Eggs. Middle one showing black ring and body segmen-

tation. Right one with developed embryo, showing cerci in lower pole. J. Left foreleg
(posterior view showing tympanum (ty)).

SOUTH AFRICAN ARTHROPODS 99

A peak emergence of adults occurred during the spring and early summer
months with an apparent second peak during the winter.

They were observed to feed on various substances including fish-meal,
grass, and bran, and when they were confined to a small space they devoured
each other; this was also found by Rivnay & Ziv (1963).

ORDER HEMIPTERA
Family Anthocoridae

Small, somewhat flattened, mostly predacious exopterygote hexapods with
three-segmented rostrum and antennae with only a few segments. Cerci absent.
Legs cursorial. Ocelli present.

Sp. indet.

The adults (Fig. 15K) vary from 2,8 mm to 4,2 mm and are dark brown
with a lighter yellowish coloration on the hemelytra. Eyes scarlet and ocelli
clearly visible. Legs with three-segmented tarsi, and antennae pale yellow.

This species was very common in the early dermatophagous stage of decay
of certain animal carcasses around Cape Town and both adults and nymphs were
found to feed on small fly larvae that were present, especially those of the blue-
black carrion-fly Ophyra capensis (Wiedemann). It was also found in the Con-
stantia area in poultry manure in which house-flies, false stable-flies, and the
lesser house-fly were breeding.

Nymphs of all stages were collected during the period January to April.
Those about 1,0 mm long (Fig. 15I) are pale brownish with scarlet eyes, cream-
coloured abdomen and pale, almost translucent legs and antennae. Also three
small, oval, reddish spots present on middle of abdomen, marking the openings
of repugnatorial glands. Characteristic of these nymphs are two long setae on
the apex of the abdomen; these are absent in the 1,4 mm long nymphs. Larger
nymphs, about 2,1 mm long, are very similar to the smaller ones, but the colour
is somewhat darker, especially on the abdomen, and the first two abdominal ter-
gites are whitish in the middle.

In the 3,4 mm specimens (Fig. 15J) there is a clear, whitish-yellow, longi-
tudinal, median line over the head and thorax, and wing rudiments are present.
The six small, semicircular patches dorsally on the abdomen mark the openings
of repugnatorial glands. The colour of these nymphs is yellowish with brownish
patches.

All the immature stages have two-segmented tarsi and ocelli are indicated
by two small scarlet patches in nymphs about 2,0 mm long and larger.

Family Coreidae

Rather similar to the Lygaeidae and also with four-segmented rostrum, but
membrane of front wings with numerous branched veins. Ocelli present. All the
species are phytophagous.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Very few squash bugs were associated with decaying organic matter when
the surveys were made and, apart from the species discussed here, only the dull,
greyish-brown Stenocephalus testaceus Stal, about 9,5 mm long, was found to
shelter under dry cow-pats along the west coast. However, Payne et al. (1968)
found three species in America, belonging to the genera Megalotomus and Aly-
dus, actually feeding on pig carrion.

Leptocoris hexophthalma (Thunberg)

This species is pale crimson to crimson with brownish wings and measures
10-11,5 mm in length (Fig. 15H). It is widely distributed in the Cape, Natal, and
the Transvaal and was also collected in east Africa. It is common in the Karoo
and sometimes congregates in very large numbers on the debris beneath the
bergvygie, Drosanthemum sp., and the skaapbossie, Justicia orchioides. Both
-newly hatched and mature nymphs have been collected on these plants during
March, when they cause extensive damage to the foliage.

The nymphs of this species are quite different from those of the other
Hemiptera inhabiting the same habitat, viz. Melanosthethus marginatus
(Thunberg), Scantius forsteri (Fabricius), and Cenaeus carnifex (Fabricius), as
they are fairly hairy and the repugnatorial gland openings are small and in-
conspicuous. In small (2,0 mm) nymphs (Fig. 15F) the hairs are long, but in
larger nymphs the hairs are short and almost spine-like. All stages of the
nymphs are very similar except for the presence of wing rudiments in the later
instars (Fig. 15G). Ocelli become visible subcutaneously only in nymphs larger
than 7,0 mm; in adult forms the tarsi are three-segmented, as in most other
Heteroptera.

Family Lygaeidae

Similar to Anthocoridae, but with four-segmented rostrum and membrane
of front wings with only a few veins. Ocelli present. Apparently predominantly
seed-feeding; however, a few species are predacious.

Masses of dying and dead stink-bugs, particularly the chinch-bugs Microspi-
lus proximus (Dallas), Geocoris scutellaris Puton, and Nysius binotatus (Ger-
mar) (one of the crop pests, particularly crucifers), and the damsel-bug, Nabis
capsiformis Germar (Nabidae), together with various other beetles such as coc-
cinellids, tenebrionids and carabids, are often found in the intertidal zone along
the west coast. This happens when these insects are blown out to sea by strong
winds and are then washed on the beaches.

Melanostethus marginatus (Thunberg)

This species is 5,8-5,9 mm long, rather dull black, with two large reddish
triangles on wings and clear circular spot on wing membrane (Fig. 16L). Heme-
lytra covered with short silvery hairs, which are absent on disc of each clavus,
thus forming an oval black spot. Pronotum deeply and abundantly pitted.

SOUTH AFRICAN ARTHROPODS 101

Fig. 15. A-D. Antestiopsis orbitalis. A. Egg. B. Egg from which nymph was hatched.

C. Newly hatched nymph. D. Nearly mature nymph. E. Telenomus sp. (egg parasite of A. or-

bitalis). _ F-H. Leptocoris hexophthalma. F. Nymph measuring 2,0 mm. G. Nymph measur-

ing 7,,mm. H. Adult. I-K. Anthocoridae. I. Nymph measuring 1,0mm. J. Nymph
measuring 3,4mm. K. Adult. L. Aderrhis tartareus (adult).

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

It is widely distributed in the Cape Province (Slater 1964) and the southern
Orange Free State, and is a common feeder on Arctotheca populifolia, Senecio
elegans (wild cineraria) and other plants, including Clutia daphnoides. It has also
been found to feed on female Cantharidae. Nymphs are very often found among
debris on the sand-dunes and large numbers of adult bugs have been collected
under dry kelp strings on the beach.

As in the case of all other members of the family Lygaeidae, there are two
ocelli in the adults and the tarsi are three-segmented (Fig. 160). In the small
nymphs (2,8 mm) (Fig. 16M), collected during April, the presence of the ocelli
is indicated only by two dark brownish marks. In this stage the body is scarlet
and the posterior corners and sides of the pronotum as well as the metanotum
are pure white. There is a thin, yellowish, longitudinal line over the middle of
the thorax; the eyes are brown. The two ostioles of the repugnatorial glands are
indicated by two brown patches and the connexivum is marked with white on
each segment. The femora are reddish marked with white on the bases and
apices. The tibiae and tarsi are pale brownish. The first and last antennal seg-
ments are reddish, but the second is pale brownish, and the third whitish.

Larger nymphs (4,0 mm) (Fig. 16N), collected during December, are dark
brownish red on the head and thorax, and the ecdysial suture is yellowish. The
posterior margin of the pronotum is bordered by a white band, which is con-
tinued along the lateral margins; the white coloration of the metanotum is vis-
ible between the two wing rudiments. The antennae and the legs are the same
colour as the body, except for the white apices of the femora and the terminal
antennal segment, which is darker than the others. The two ostioles and extreme
apex of abdomen are dark brown and the connexivum is almost the same colour
as in the 2,8 mm long nymphs.

Family Pentatomidae

Easily recognized by the large scutellar area, rostrum also four-segmented.
Most species are phytophagous such as the bagrada bug, green stink-bug, and
the well-known antestia bug, which is a pest of coffee in the northern parts and a
fruit pest in South Africa, and which will be very briefly discussed here. How-
ever, a few species are predacious, feeding mostly on the caterpillars of Lepi-
doptera.

Antestiopsis orbitalis (Westwood)

Specimens collected on the fore-dunes along the south and west coasts are
pale cream to almost dirty white; some have orangy patches on the head, front
margin of pronotum, scutellum, and along wing embolium. Eyes are brown. In
some specimens there is an orange spot anterior to each eye, as well as two spots
on the anterior part of the scutellum; in others these orange spots are com-
pletely absent. These bugs measure 6,9-7,5 mm in length. Greathead (1966) has
given a good account of the different colour variations and the distribution.

=

SOUTH AFRICAN ARTHROPODS 103

They are widely distributed from the Cape to Kenya but, according to col-
lecting data, they are absent from the Karoo. Eggs, nymphs, and full-grown
specimens of this shield-bug were collected during the summer (December-—
February) on the foliage of Arctotheca populifolia along the coast. Brown
patches on the leaves resulted where large numbers of the feeding stages were
concentrated. They often congregate in large numbers on leaf litter under the
above-mentioned plant.

The eggs of this species (Fig. 15A—B) are about 1,1 mm long and 0,9 mm
wide, creamy white in colour, and the operculum is surrounded by a row of
short micropylar processes; about twenty-seven were counted in most of the
eggs collected. All the eggs found were laid in batches of twelve on the surface
of leaves. After hatching, the black, T-shaped egg-burster can be seen on one
side of the empty eggshell. The newly hatched, chocolate-brown nymphs (Fig.
15C), about 1,2 mm long, with two pale yellowish white patches on the pale ab-
domen, are apparently gregarious, at least up to the first moult, and were often
seen clustered around the eggs during February. As the nymphs increase in size,
they assume the coloration of the adult forms (Fig. 15D).

Skaife (1953) has given a short account of the life-history of this bug and
Greathead (1966) has listed its food plants. According to Annecke & Moran
(1982) it also damages the growth tips of Protea and Leucadendron species. An
egg parasite, Telenomus sp. (Fig. 15E), has been recovered from the Strandfon-
tein area near Muizenberg, Cape Province.

Family Pyrrhocoridae

Similar to Lygaeidae, with membrane of front wings also with only a few
veins, but ocelli absent. Usually brightly coloured with red, orange and black.
Most of them are phytophagous and include the well-known cotton-stainers:
however, a few are predacious.

Scantius forsteri (Fabricius)

The colour of this red bug (Fig. 16A) is dark chocolate-brown, but front of
head, thorax, apex of scutellum, hemelytra, and connexivum are all marked
with crimson. It is very variable in colour and in some specimens the hemelytra
are uniformly chocolate-brown. The antennae and legs are dark chocolate-
brown, almost black in some specimens. Brachyptery is common. Specimens
collected vary from 7,5 to 10 mm in length.

It is widely distributed in Africa and very common along the western parts
of the Cape Province where the surveys were made. Adults and nymphs often
occur in large numbers during the summer and autumn under semi-fresh to dry
cow-pats on the open beach and inland. It is also present on Dassen and Marcus
islands.

The newly emerged, elongate nymphs (Fig. 16B) are pale piceous on the
thorax and the head is pale brownish yellow with posterior border piceous. Eyes

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

scarlet, basal half of abdomen whitish flanked with scarlet and apical half yel-
lowish white behind and scarlet in front; connexivum reddish; position of the re-
pugnatorial glands indicated by three thin, transverse lines dorsally on apical
half of abdomen. Legs pale piceous, first of the two tarsal segments and articula-
tion points white. First two antennal segments pale brownish yellow, third and
fourth pale piceous, with apical third of last segment white. Most of the speci-
mens examined were about 1,6 mm long and in all of them minute denticles or
spinules occur on the ventral side of the apices of the front femora.

As the nymphs increase in size (Fig. 16C), the ecdysial suture lengthens
posteriorly over the first two to three abdominal terga and the wing rudiments
enlarge (Fig. 16D). In the 10 mm newly moulted nymphs, the wings cover the
greatest part of the abdomen and the tarsi are three-segmented. The whole in-
sect is scarlet, except the wings and the largest part of the second antennal seg-
ment, which are almost white. Eyes are piceous. In older adults the colour is
darker and the teeth on the apices of the femora are more strongly developed
(Fig. 16E-G).

Along the south coast a dull, brownish-black brachypterous pyrrhocorid
Aderrhis tartareus (Stal) (Fig. 15L) was fairly generally collected under dry cow-
pats together with Scantius. It is 8-8,5 mm long, with a short, thin, orange-red,
transverse band on the posterior border of the head and an orange-red spot on
the apex of the scutellum. According to Stehlik (1965) it is found only in South
Africa and seems to have predacious habits. A macropterous female, collected
by R. W. Tucker in the Transvaal in December 1913, measured only 7,1 mm in
length.

Cenaeus carnifex (Fabricius)

This species (Fig. 16H) is yellowish red, fairly dull or only slightly shiny;
scutellum, frons and posterior border of head blackish. Antennae, proboscis and
legs blackish, except for basal third of femora, which is reddish. The specimens
examined measure 8,3—10,0 mm in length. Brachyptery is common. The genus is
characterized by the crossed parameres (Stehlik 1965).

It is widely distributed in the Cape and Natal.

Nymphs and adults of C. carnifex were collected under dry cow-pats during
midwinter. Adults are often seen on garden plants during the summer months.
As in Scantius, all the immature stages have two-segmented tarsi, but the
nymphs are quite different in other respects as they are more cimicoid in out-
line, paler in colour and the ostioles of the repugnatorial glands are represented
by three larger blackish patches. In all the immature stages, as well as adults
examined, spines are present only on the apices of the front femora (Fig. 16K).

In the 4,5-5 mm nymphs (Fig. 161), the legs and antennae are piceous, the
articulation points reddish. The head, except the median area, is yellowish and
the thorax brown with yellowish ecdysial suture. The abdomen is pale yellowish
red with a blackish apex.

SOUTH AFRICAN ARTHROPODS

i=

>
5G
* &
ig
ie
2 \

pro

Holga nn)

105

Fig. lo. A—G. Scantius forsteri. A. Adult. B. Newly hatched nymph. C. Nymph measur-
ing 2.3mm. D. Nymph measuring 6.4mm. E. Prothoracic leg of male (posterior view).
F. Mesothoracic leg of male (posterior view). G. Metathoracic leg of male (posterior view).

H-K. Cenaeus carnifex. H. Adult showing crossed parameres (p).

I. Nymph measuring

477mm. J. Nymph measuring 7.8mm. K. front leg (posterior view) showing spine on ante-
rior ventral side. Enlarged to same scale as E-G. L—M. Melanostethus marginatus. L. Adult.

M. Nymph measuring 2,8 mm. N. Nymph measuring 4.2 mm. O. Foreleg (posterior view).

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

Larger nymphs (6,5—8 mm) (Fig. 16J) are similar to the smaller ones, but
rudiments of wings are present and most of the pronotum, except two darker
areas on each side of the middle, is yellowish.

Family Reduviidae

Easily recognized by the strongly developed, curved, three-segmented ros-
trum. Wings usually well developed, though some species are apterous. Many
species have raptorial front legs and many stridulate when handled, the stridula-
tion being prolonged under certain conditions. Predacious, mainly feeding on ar-
thropods. Most species inject a saliva containing a paralysing agent, which also
helps to digest the tissues. In man the bite causes severe pain; the tissues sur-
rounding the puncture become inflamed and harden and the irritation can persist
for several days. The subfamily Triatominae includes species that suck the blood

of vertebrates and transmit disease-causing trypanosomes; however, none of
these are known to occur in South Africa.

Certain assassin-bugs, particularly of the genus Pirates, are attracted to
lights and are often found in houses. An unidentified black species (about
11 mm long) belonging to this genus as well as one shiny, metallic bluish-black,
apterous Glymmatophora sp. (12-15 mm long) have on several occasions been
reported to have bitten people in the western Cape, causing severe pain and
swelling. Some assassin-bugs, including these two widely distributed species that
have both been collected under cow-pats, bite without hesitation when carelessly
handled or when pressed against the skin, and may cause severe symptoms in
hypersensitive people.

Several assassin-bugs were found to shelter under semi-dry cow-pats, feed-
ing on the smaller arthropods present. The most common of these is the hairy,
black Coranus carbonarius (Stal) (Fig. 17H), which is 11-12,5 mm long. Black-
ish nymphs, with blackish-grey abdomen, found in the vicinity, probably be-
longed ‘to this species. Coranus pallescens (Germar), a smaller, greyish bug
(about 8 mm long) was found under similar conditions together with C. papillo-
sus (Thunberg) (Fig. 171), which is known to be a predator of codling moth lar-
vae (Myburgh et al. 1973). Coranus papillosus is about 9,8 mm long, dark
brownish grey in colour and very similar to C. pallescens (which was also found
to be a victim of the black widow Latrodectus mactans (Hesse 1942)). It is well
established on some of the islands along the south coast, e.g. Dyer Island.

A rather small, brownish Didymocephalus braunsi Bergroth (about 7,5 mm
long), covered with fine velvety hairs as in the case of Coranus spp., was fairly nu-
merous near Hermanus during winter, whereas a species of Harpactor, of about
the same length as C. papillosus but almost black, the connexivum coloured black
and yellowish white, was more common along the north-western parts of the
coast. Specimens observed were found to feed on coleopterous larvae.

In the subfamily Holoptilinae the members attract their prey, which con-
sists mainly of ants, by means of a special scent gland or trichome present on the

SOUTH AFRICAN ARTHROPODS 107

Fig. 17. A-G. Oncocephalus sp. A. Female (adult). B. Head of female showing labrum

(labr) and ocelli (oc). C. Egg showing developing eyes of nymph. D. Newly hatched nymph.

E. Head of newly hatched nymph. F. Outline of male. G. Nymph 11,6 mm long (outline of

body), probably belonging to the second species. H. Coranus carbonarius (adult). I. Cora-
nus papillosus (adult).

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

ventral surface of the abdomen. These bugs are sluggish and usually covered
with long hairs. An unidentified species of Holoptilus was discovered in the nest
galleries of the ant Melissotarsus beccarii Emery in the trunks of the
pincushion Leucospermum praemorsum in the vicinity of Clanwilliam (Prins et
al. 1975), the bugs living a ‘double life’ as symphile and predator. Holoptilus
ursus le Peletier & Serville, a small brownish assassin-bug with a brown patch
on the proximal half of the whitish hemelytra, is common on debris in sandy
areas along the Cape south coast where it feeds on the smaller ants, part-
icularly the minor workers of the pugnacious ant, Anoplolepis steingroeveri
(Forel).

Oncocephalus sp.

The full-grown females (Fig. 17A) measure 18 mm in length and are cream
coloured or greyish marked with piceous brown. Largest part of first segment of
~ beak, apical half of second and whole of its third segment is piceous. Body, in-
cluding legs, covered with fine tubercles, those on head and thorax being larger
and tooth-like and each bearing a spatulate seta. Also present are some thick-
ened hairs and fine whitish scale-like hairs that have the look of a pruinescence.
Characteristic of most of the specimens examined are the two black dots on the
median area of most of the abdominal segments and the tiny blackish ostioles of
the repugnatorial glands on the anterior borders of the third and fourth visible
terga.

Head (Fig. 17B) bears two rounded teeth between four-segmented anten-
nae. Second antennal segment very long, about twice as long as first; apical and
praepical segments of about equal length and together slightly more than half
the length of the second. Labrum small and triangular. Pronotum with six teeth,
four on anterior and two on posterior lobe. Also a fairly acute tooth present on
each anteroventral corner of pronotum. Anterior lobe of pronotum fairly convex
transversely and longitudinally.

Front femur swollen, its anteroventral margin bearing about twelve small
teeth; front and middle tibiae clearly marked with two or three dark transverse
bands. Wings atrophied in both females and males; latter similar to females but
easily distinguished by rounded apex of abdomen (Fig. 17F); in females abdomi-
nal apex is pointed.

The specimens studied came from the Saldanha and Elands Bay areas along
the west coast.

This is a rather slow-moving insect that feigns death when disturbed by ex-
tending its front and middle legs forward and its hind legs backward. The speci-
mens were all collected under almost dry cow-pats during autumn and early
winter. The eggs (Fig. 17C) are elongate with a convex operculum and have an
almost matt, dirty yellowish-white colour and a very fine and superficial reticula-
tion; they measure 1,60 xX 0,96 mm to 1,80 x 1,20 mm. Eggs were laid under
the sand under dry pats during July and August and the incubation period was at
least 28 days in the laboratory (at 19 °C).

SOUTH AFRICAN ARTHROPODS 109

Newly hatched nymphs (Fig. 17D) measure about 2,55 mm in length and
are pale brownish white, posterior half of head, pronotum and meso- and meta-
notum being darker. Eyes crimson. First two abdominal segments dark. Abdo-
men pale piceous or greyish and the two ostioles of repugnatorial glands already
visible. A thin, white, longitudinal median line over the thorax represents the
acdysial suture. Tubercles on body pale and each bears a spatulate seta. Legs
pale and front femora already have teeth; hind femora somewhat darker.

These nymphs differ from the mature insects by the absence of both ocelli
and rounded teeth between the antennae. Whole of second segment of beak is
white (Fig. 17E). First and second antennal segments short; first only about half
as long as second, latter about as long as third; apical and penultimate segments
about equal in length. Head and pronotum much larger in relation to body than
in adult, and dorsal teeth absent from thorax but anteroventral teeth fairly long.
As in other Heteroptera the tarsi are two-segmented.

Development was very slow in the laboratory and after about 100 days they
were only 3,6—4 mm long. Larger nymphs (about 5,1 mm long), collected during
November, show the same characteristics as the newly hatched ones, but the
thoracic teeth are more pronounced and each abdominal segment, except the
first visible segment, has two pairs of small, oval, dark spots, the outer ones
smaller; the connexivum also bears six dark spots on each side.

Fairly large nymphs, 11,6 mm long, collected during February, resemble
the adults (including the presence of teeth between the antennae) but still pos-
sess abdominal spots (as in the 5 mm long nymphs).

The young nymphs unfortunately died of starvation in the laboratory, as
they did not accept any prey offered to them.

From the specimens collected this is either a polymorphic species or poss-
ibly two species of similar coloration present in the same area, as some of the
males have much longer wing stumps, the ocelli are absent, the scutellar tooth is
much smaller and the abdomen is more pointed. Nymphs (probably belonging to
males of this second form) measuring 11-12 mm long (Fig. 17G), found during
February, have the connexivum sexdentate on each side and the abdomen more
pointed than in the other form.

Nymphs of another unidentified species collected at Elands Bay during the
late summer, more cimicoid in outline, with slightly darker coloration and meas-
uring about 9,0 mm long, have longer developed wing-covers. In this case the
body is covered with slightly longer hairs, which are simple on the abdomen and
more rod-shaped on the head and thorax. Almost the whole of the rostrum is
dark brown in colour, apical half being almost black; the third segment is much
smaller in relation to the second than in the previously mentioned species. In
younger nymphs (7,3 mm long and also with longer wing-covers) the pilosity,
particularly of the head and thorax, consists of hamate setae. In all the speci-
mens examined, ocelli and the rounded teeth between the antennae so charac-
teristic of the adults and larger nymphs of the first mentioned species, are
absent.

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

I wish to express my gratitude to Dr V. B. Whitehead of the South African
Museum, Cape Town, for valuable suggestions, and to Mr V. Branco, also of
the South African Museum, for some of the drawings.

I should also like to thank the following specialists for identifying material:
Dr B. Kensley, US National Museum (Crustacea); Dr M. K. P. Smit Meyer
(Acari) and Dr A. A. Dippenaar (Araneida), both of the Plant Protection Re-
search Institute, Pretoria; Mrs J. Marshall, British Museum (Natural History),
London (Blattidae); Dr J. B. Walker, Veterinary Research Institute, Onderste-
poort (Ixodidae); and Dr G. Newlands, South African Institute for Medical Re-
search, Johannesburg (Scorpionida).

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ent. Soc. Am. 61: 565—567.

Princis, K. 1963. Blattariae: Revision der Sidafrikanischen Blattarienfauna. S$. Afr. anim. Life
9: 1-318.

Prins, A. J. 1980. The arthropods associated with decaying organic matter in the southern and
western Cape Province. Unpublished Ph.D. thesis (Agriculture), University of Stellen-
bosch.

Prins, A. J., BEN-Dov, Y. & Rust, D. J. 1975. A new observation on the association between
ants (Hym: Formicidae) and armoured scale insects (Homoptera: Diaspididae) J. ent. Soc.
sth. Afr. 38: 211-216.

Rean, J. A. G. & Reun, J. W. H. 1935. A study of the genus Hemimerus (Dermaptera: Hemi-
merina: Hemimeridae) Proc. Acad. nat. Sci. Philad. 87: 457-508.

Rivnay, E. & Ziv, M. 1963. A contribution to the biology of Gryllus bimaculatus De G. in Is-
rael. Bull. ent. Res. 54: 37-43.

112 ANNALS OF THE SOUTH AFRICAN MUSEUM

Rotu, L. M. & WiLuis, E. R. 1957, The medical and veterinary importance of cockroaches.
Smithson. misc. Collns 134(10): 1-147.

Rotu, L. M. & Wixuis, E. R. 1960. The biotic associations of cockroaches. Smithson. misc.
Collns 141: 1-470.

SCHARRER, B. 1951. The Woodroach. Scient. Am. 185: 58-62.

SCHUBART, O. 1966. Diplopoda III. Pselaphognatha, Opisthospermophora, Colobognatha.
S. Afr. anim. Life 12: 19-227.

SKAIFE, S. H. 1953. African insect life. Cape Town: Longmans Green.

SKAIFE, S. H. 1979. African insect life. New Edition. Cape Town, Johannesburg: Struik.

SLATER, J. A. 1964. Hemiptera (Heteroptera): Lygaeidae. S. Afr. anim. Life 10: 15-228.

Smit, B. 1964. Insects in southern Africa; how to control them. Cape Town: Oxford University
Press.

SmiTH, K. G. V. 1973. Insects and other arthropods of medical importance. London: The
Trustees, British Museum (Nat. Hist.).

STEHLIK, J. L. 1965. Mission zoologique de VPI.R.S.A.C. en Afrique orientale (P. Basilewsky.
N. Leloup 1957). Pyrrhocoridae (Het.). Cas. morav. Mus. Brné 50: 211-252.

WALKER, E. P. 1964. Mammals of the world. 2. Baltimore: Johns Hopkins Press.

ZumpT, F. & PATTERSON, P. M. 1952. Flies visiting human faeces and carcasses in Johannes-
burg, Transvaal. S. Afr. J. clin. Sci: 3: 92-106.

Sn,

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6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.

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A. J. PROMS

MORPHOLOGICAL AND BIOLOGICAL
NOTES ON SOME SOUTH AFRICAN
ARTHROPODS ASSOCIATED

WITH DECAYING ORGANIC MATTER
PART 1

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

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

FIsCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull, Bingham oceanogr. Coll. 17 (4): 1-51.

THELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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mNINALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 92 Band
March 1984 Maart
Part 3 Deel

CRETACEOUS FAUNAS FROM
ZULULAND AND NATAL, SOUTH AFRICA
THE AMMONITE SUBFAMILY
PERONICERATINAE HYATT, 1900
By
HERBERT CHRISTIAN KLINGER
&

WILLIAM JAMES KENNEDY

Cape Town Kaapstad

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CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,
SOUTH AFRICA
THE AMMONITE SUBFAMILY PERONICERATINAE HYATT, 1900

By
HERBERT CHRISTIAN KLINGER

South African Museum, Cape Town
&

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

(With 135 figures and 1 table)

[MS accepted 10 May 1983]

ABSTRACT

The systematics of the South African representatives of the subfamily Peroniceratinae are
critically reviewed, with special attention to the almost simultaneously published monographs of
Van Hoepen (27 July 1965) and Matsumoto (20 November 1965) who, although working inde-
pendently, reached a number of similar conclusions and erected a series of genera that are
shown to be synonyms. The genera and subgenera Peroniceras (Peroniceras) de Grossouvre,
1894, P. (Zuluiceras) van Hoepen, 1965, and Gauthiericeras de Grossouvre, 1894, include all
the Zululand peroniceratids.

The present material is the richest assemblage of Peroniceratinae yet described, and it per-
mits clarification of the limits of both inter- and intra-specific variability in the group as well as
their evolution and stratigraphic distribution. Two new species are described: Peroniceras
(Zuluiceras) matsumotoi and Gauthiericeras albiforme.

Subdivision and zonation of the Coniacian stage is critically reviewed as this has bearing on
the systematics of the subfamily.

CONTENTS

PAGE

MN EROGIUC EI OM ete a tate aa Otek Peis aoe esa We lets MR aie ee ER was els 114
OCA OMOLES DE CUNICTIS 6 cas oct aye a ve ais ones ake gots Wes eee 115
FCIONOCAINIC Seen re erate Ue te tee rt oa tel RO Le ere te eee IS)
DIMENSIONS OL SPECIMENS wane eenicie sree 4 oo ances el ae 115
STUUR eRMINO] OL Venn Metra eet cee etr nes oo Plat Aish eat eee 116
Coniacian(straticraphyandicomelation :..-2.+--. 5645 - s25 450s: 116
TT URO CU CLIO Mae ta ete ire Ph onthe: Re here greet de erea Aiea steam 116
Monvavonanithe type re gon qa. sass. ets We a eee 116

she ululanG SUCCESSIOMM.§ 5. e hs ee wel aes Cea eee 120
IIa ae ASC ates latent Se an esa earn Sari ones Ree AE ee ee 122

AAG) ATI srt reyes Ore ahs Ainley SOE Ms wernt ARATE CRT IR, eae lassen ees 124
BINA ee etc ear TO ES, OF Sh Sy tact! ahd ee A a bot 5)
INGRCI SHINES PAIN ete ace a foe wits a ayers re Penske aeaee Ween CHa? 126
ROMINA Ae Se ees ne ors tie ois ce TA ESTE ERR ene 128
GOMCIMSTONMS ee ey tr oe bea 1 WN eh eg ae eae Ba 130

its

Ann. S. Afr. Mus. 92 (3), 1984: 113-294, 135 figs, 1 table.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Systematic palacontolopy ees tae eee ae 130
DISCUSSIONS 5-year 130
GENUS, EKONICETGS FG 0 NR Se eM Eos Oe ok ate ee 135
Subgenus Zeroniceras (Reronicenas) saan ene oh eee 138
Subgenuss2eronicerasn(Zuluiceras nn sae eee ee 180
GENUS GaGUWHhiCriCehass. mn. watts oe ee ; 238
COMCIUSIOMS Hiserch aM ecat eaten Ar eect ea ee acer ae 287
Acknowledgements: Ac. .see eon eee eh eae ee ee eee 289
IRELETE NCES iso eos ees oper foc ats, oS a ees OE esc ee ae 290

INTRODUCTION

Ammonites referable to the subfamily Peroniceratinae Hyatt, 1900, occur
abundantly in two main areas in Zululand, the lower Hluhluwe and the lower
Mzinene rivers, while isolated specimens are known from the environs of Mtu-
batuba (Umkwelane Hill of earlier workers) and recent excavations for the con-
struction of the new north-south highway that transects the region.

Peroniceratids were described by several early workers in the area, includ-
ing Crick (1907), Spath (1921), Besairie (1930), Venzo (1936), and Van Hoepen
(1955). In 1965 there appeared Van Hoepen’s monographical study The Peroni-
ceratinae and allied forms of Zululand. Unfortunately most of Van Hoepen’s
new species were based on single specimens, especially large individuals lacking ~
the inner whorls. This led him to erect 1 new subfamily, 5 new genera, 2 new
subgenera and 26 new species for his material. Only one previously established
species was described, Peroniceras (Peroniceras) tridorsatum (Schliter, 1867).

In the same year as that of Van Hoepen’s monograph, Matsumoto pub-
lished his account of the Peroniceratinae from Hokkaido, Japan, in which he
erected four new genera. Van Hoepen and Matsumoto, working independently,
arrived at very similar conclusions in certain respects. Several of their taxa are
synonyms, as will be shown below. Van Hoepen’s work was published post-
humously on 27 July 1965, four months before Matsumoto’s on 20 November
1965. All these new taxa are reviewed in the present paper in order to stabilize
nomenclature. In the process Van Hoepen’s work is revised in the light of cur-
rent systematic concepts and stratigraphic data. Species not described by Van
Hoepen are also included in this review. The material at the present authors’
disposal sheds some new light on the phylogeny and classification of the subfam-
ily Peroniceratinae.

The following species from Zululand are described:

Peroniceras (Peroniceras) lepeei (Fallot, 1885)
. (P.) tridorsatum (Schliter, 1867)

. (P.) subtricarinatum (d’Orbigny, 1850)

. (P.) westphalicum (von Strombeck, 1859)
. (P.) dravidicum Kossmat, 1895

. (Zuluiceras) zulu van Hoepen, 1965

. (Z.) rarum van Hoepen, 1965

ay a3} ae 2S) oe) Fe

CRETACEOUS FAUNAS FROM SOUTH AFRICA 1S

P. (Z.) henniet van Hoepen, 1965

P. (Z.) undulatocarinatum van Hoepen, 1955
P. (Z.) modestum van Hoepen, 1965

P. (Z.) matsumotoi sp. nov.

P. (Z.) cf. aberlei (Redtenbacher, 1873)
P. (Z.) bajuvaricum (Redtenbacher, 1873)
Gauthiericeras obesum van Hoepen, 1955
G. listeri (van Hoepen, 1965)

G. album (van Hoepen, 1965)

G. albiforme sp. nov.

G. libertae van Hoepen, 1955

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the repositories of the ma-
terial studied:

BMNH British Museum (Natural History), London.

EMP Ecole des Mines, Paris, now in the Université Claude Bernard, Lyon

GG Laboratoire de Géologie, Grenoble

GPIB Geologisches und Palaontologisches Institut der Universitat, Bonn

iL. Oberosterreichisches Landesmuseum, Linz

MNHP Muséum d’Histoire Naturelle, Paris

NMB_ National Museum, Bloemfontein (presently in the collections of the
South African Museum)

OUM Oxford University Museum

SAM South African Museum, Cape Town

SAS South African Geological Survey, Pretoria

SP Collections of the Sorbonne, Paris, now in the Université Pierre et
Marie Curie

BIELD LOCALES

Details of field localities are given by Kennedy & Klinger (1975); fuller de-
scriptions of these localities are deposited in the Department of Palaeontology,
British Museum (National History), London; South African Geological Survey,
Pretoria; and South African Museum, Cape Town.

DIMENSIONS OF SPECIMENS

All dimensions are given in millimetres:
D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical
diameter. R indicates ratio of umbilical to ventrolateral tubercles.

Figures in parentheses are dimensions as a percentage of the total diameter.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916) reviewed and discussed by
Kullman & Wiedmann (1970) is followed here:
I = internal lobe, U = umbilical lobe, L = lateral lobe, E = external lobe.

CONIACIAN STRATIGRAPHY AND CORRELATION

Introduction

The subfamily Peroniceratinae is predominantly Coniacian and members of
the group provide a basis for subdivision and correlation of the stage. Present
subdivision of the stage is in a state of chaos as was made painfully apparent by
Matsumoto’s (1981) recent review of Coniacian biostratigraphy. Some workers
-use only Lower and Upper Coniacian substages, others Lower, Middle and
Upper, with no clear indication as to how these two schemes are related. The
Same species are recorded from the ‘Lower’ Coniacian of one region and the
‘Upper’ Coniacian of another so that the ranges of taxa appear to be anomalous.
Zonal nomenclature is similarly confused.

Kennedy (in press) has reviewed the early history of subdivision of the
Coniacian in France, and proposed a revised zonation and subdivision into
substages as summarized below.

Zonation in the type region

The Coniacian stage was introduced by Henri Coquand in 1857 as the low-
est division of D’Orbigny’s Senonian stage, introduced in 1842-3. It has passed
into the literature as a universally applied division of the Upper Cretaceous. The
standard zonation of the stage is based on ammonites and is the twofold division
of De Grossouvre (1889, 1901) who recognized a lower Barroisiceras [‘Barroi-
sia’| haberfellneri Zone and an upper Paratexanites [‘Mortoniceras’| emscheris
Zone on the evidence of sequences in Aquitaine and Touraine in France. These
zones have been universally accepted as standard zones for the Coniacian and
were adopted by the 1959 Colloque sur le Crétacé supérieur francais (Dalbiez
1960), in the Treatise (Wright 1957), and by Séronie-Vivien (1972) in her defini-
tive revision of the stratotype, among many others. Yet already in 1947 Basse
has pointed out that the Barroisiceras haberfellneri of De Grossouvre was based
on a misinterpretation of Von Hauer’s species, and that the French material all
belonged to a distinctive form which she named Reesideoceras gallicum Basse,
1947 (a synonym of Ammonites petrocoriensis Coquand, 1859), although never-
theless referring to a ‘haberfellneri’ Zone in some of her later works. True
Barroisiceras does not occur in France, indeed, there is no good evidence that
the species B. haberfellneri actually occurs in the Coniacian. Similarly with the
Paratexanites emscheris Zone, the index species does not occur in France.

Confusion is not limited to nomenclature. Collignon (e.g. 1960) used the
haberfellneri-emscheris standard in his early works, and developed a threefold

CRETACEOUS FAUNAS FROM SOUTH AFRICA i oF

zonation in Madagascar, with zones of Peroniceras dravidicum (oldest), Koss-
maticeras theobaldianum and Barroisiceras onilahyense, and Prionocycloceras
guyabanum and Gauthiericeras margae (youngest). In 1979 Collignon et al. re-
cognized a sequence of three zones in the Coniacian of the Beausset Basin (Var,
France): Peroniceras tricarinatum (oldest), Barroisiceras haberfellneri, and Gau-
thiericeras margae (youngest), although the middle zone was not proven by am-
monites. Robaszynski et al. (1980) recognized a Peroniceras tridorsatum Zone at
the base of the Coniacian in the Boulonnais, France (with, however, only eight
specimens of the genus Peroniceras from the area), and French textbooks now
cite a standard sequence of tricarinatum—haberfellneri-margae Zones despite the
fact that haberfellneri does not occur in France.

In those areas where a faunal sequence can be determined, Forresteria
(Harleites) petrocoriensis (Coquand, 1859)—the Barroisiceras haberfellneri of
authors—always precedes Peroniceras species and marks the base of the Conia-
cian in the type area (Kennedy, Wright & Hancock 1983; Kennedy in press).
Yet more recently, Matsumoto (in Matsumoto et al. 1981) proposed as standard
zones for the European Coniacian a sequence of Peroniceras tricarinatum Zone
(below) and Protexanites bourgeoisi Zone (above) on the basis of what could be
gleaned from the literature.

Ammonites are generally very rare in the Contacian of France (and indeed
most of north-western Europe), and only five areas, the Aquitaine Basin, a
small area in Touraine in the southern Paris Basin, a few square kilometres of
outcrop around Dieulefit (Drédme), the Beausset Basin (Var), and the environs
of Rennes-les-Bains in the Corbiéres (Aude), have yielded ammonites in any
numbers. In their revision of the Craie de Villedieu, Jarvis et al. (1983) do not
mention new records of Coniacian ammonites around Villedieu-le-Chateau, the
most famous Coniacian ammonite locality in France. In spite of these difficul-
ties, the French succession provided Kennedy with the basis for the following
subdivision of the Contacian:

Lower Santonian Texanites gallicus Zone (provisional)
Paratexanites serratomarginatus Zone

Upper Coniacian 5 ae
PP Gauthiericeras margae Zone

Middle Coniacian Peroniceras (Peroniceras) tridorsatum Zone
Lower Coniacian Forresteria (Harleites) petrocoriensis Zone
Upper Turonian Subprionocyclus neptuni Zone

Details are as follows:

1. Forresteria (Harleites) petrocoriensis Zone

The base of the zone and the Coniacian stage are marked by the appear-
ance of the index species, specimens of which occur only centimetres above the
regional hardground or discontinuity surface that separates basal Coniacian
sands and marls and underlying Turonian rudistid limestones in Aquitaine.
Other ammonites are rare, with only single specimens of Scaphites (Scaphites)
cf. meslei de Grossouvre, 1894, Metatissotia desmoulinsi (de Grossouvre, 1894),

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

and M. nanclasi (de Grossouvre, 1894). This is the fauna of Arnaud’s (1877) As-
size K of the Aquitaine sequence.

2. Peroniceras (Peroniceras) tridorsatum Zone

The base of the zone is marked by the appearance of diverse Peroniceras
(Peroniceras) species. The fauna of this zone is widely recorded in Arnaud’s As-
size L! in Aquitaine and is marked by P. (P.) tridorsatum (Schliter, 1867) and
other Peroniceras (Peroniceras) species with rarer P. (Zuluiceras) bajuvaricum
(Redtenbacher, 1873). Other ammonites are Onitshoceras? ponsianum (de
Grossouvre, 1894), Tongoboryceras sp. nov., Placenticeras fritschi de Gross-
ouvre, 1894, Forresteria (Harleites) nicklesi (de Grossouvre, 1894), Gauthieri-
ceras? boreaui (de Grossouvre, 1894), Tissotia redtenbacheri (de Grossouvre,
1894), Metatissotia slizewiczi (de Grossouvre, 1894), Metatissotia ewaldi (von
Buch, 1847), Metatissotia nodosa (Hyatt, 1904), Tissotioides haplophyllus (Red-
tenbacher, 1873), Scaphites meslei (de Grossouvre, 1894), Otoscaphites arnaudi
(de Grossouvre, 1894), and nodose Baculites.

Sufficient of these ammonites occur together at single localities to indicate
that a single fauna is present, but the evidence from the old collections is not
unequivocal. More recent collecting showed Peroniceras (Zuluiceras) bajuvari-
cum, Metatissotia ewaldi, and Scaphites (Scaphites) meslei occurring together at
Antignac (Charente-Maritime), but suitable outcrops are now rare in Aquitaine.

Further evidence for the composition of the P. (P.) tridorsatum Zone fauna
in France comes from Dieulefit in Drome, where the ammonite fauna includes
many forms known only from L! in Aquitaine. Species in common are P. (P.)
tridorsatum, P. (Zuluiceras) bajuvaricum, Metatissotia robini, Tissotioides haplo-
phyllus, and S. (Scaphites) meslei, with as additional elements Peroniceras (P.)
lepeei (Fallot, 1885), P. (P.) westphalicum (von Strombeck, 1859), P. (Zulu-
iceras) isamberti (Fallot, 1885), and Scalarites sp.

The association from the Marnes de Ceyreste in the Beausset Basin, Var,
also suggests that the assemblage represents a single zonal fauna with the co-
occurrence of Peroniceras (P.) subtricarinatum, Parapuzosia (Parapuzosia) sp.,
Placenticeras sp., Forresteria (Forresteria) alluaudi (Boule, Lemoine and
Thévenin, 1907) Scaphites (Scaphites) meslei and Otoscaphites arnaudi.

3. Gauthiericeras margae Zone

The base of the zone is marked by the appearance of Gauthiericeras margae
(Schliiter, 1867) and G. nouelianum (d’Orbigny, 1850). Also present are
G. ?boreaui (de Grossouvre, 1894), which ranges from below, Tridenticeras sp.,
and nodose Baculites.

The position of the zone is unequivocally demonstrated by specimens of
G. margae from Arnaud’s (1877) Assize L? in Aquitaine, and the record from
the middle part of the Grés de Baguier of the Beausset Basin (Collignon et al.
IOWD).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 119

4. Paratexanites serratomarginatus Zone

The base of the zone is marked by the appearance of the index species.
Other elements of the assemblage that are restricted to the zone in France are
Protexanites (Protexanites) bontanti (de Grossouvre, 1894), Placenticeras semior-
natum (d’Orbigny, 1850), Phlycticrioceras trinodosus (Geinitz, 1850), and no-
dose Baculites. Protexanites (P.) bourgeoisi (d’Orbigny, 1850) ranges from below.

This is the fauna of Bed 4 of the Craie de Villedieu in Touraine (Kennedy
in press). Its position is indicated by its occurrence above the condensed Cal-
caires Durs, which contains Gauthiericeras margae there, by the index species
and Protexanites (Protexanites) bourgeoisi occurring very high in the Coniacian
of northern Aquitaine, and by the index occurring above G. margae in the
Beausset Basin in Var.

5. The Coniacian—Santonian boundary in France

At Javrezac (Charente), a locality mentioned by Coquand as a type section
for both Coniacian and Santonian stages, the boundary is easily drawn at a hard-
ground between glauconitic limestones of the Coniacian below and marls of the
Santonian above. Defining this boundary by using ammonites is much more
difficult. The precise position of ammonites within the lower part of the San-
tonian—Arnaud’s assize M'!—is poorly documented, and ammonites are very
rare indeed. De Grossouvre (1894) recorded Paratexanites |‘Mortoniceras’| ser-
ratomarginatus (Redtenbacher, 1873) from both the Coniacian and Lower San-
tonian; in 1901 he decided that the Coniacian specimens all belonged to
P. [‘Mortoniceras’| emscheris (Schliter, 1876) and that P. serratomarginatus was
exclusively Santonian. He never illustrated Santonian serratomarginatus, and
none survive in museum collections that can be attributed to Arnaud’s M!. In-
deed, later work has shown that P. serratomarginatus and P. emscheris are not
synonyms and are both from high in the Coniacian (Wiedmann 1979; Kennedy,
Klinger & Summesberger 1981). De Grossouvre (1901) also indicated in a table
that Protexanites [‘Mortoniceras’| bourgeoisi (d’Orbigny, 1850) also extended
into the lower part of the Santonian, but made no mention of this in the text and
no specimens from M! survive.

Far more widely recorded is Texanites texanus (ROmer, 1852), and this was
taken as the index of the lower zone of the Santonian by De Grossouvre. Collig-
non (1948) correctly interpreted R6mer’s species (excluding all European
records), which he referred to two distinct varieties, regarded by subsequent
workers as either subspecies or separate species. Material from M! in Aquitaine
(De Grossouvre 1894: 80, pl. 16 (figs 2, 4)) was referred to the variety gallica
Collignon, 1948; the two figured syntypes are actually from M? at Niel-le-Virouil
(Charente-Maritime). The only other specimens seen from Aquitaine are two
specimens labelled Ammonites coniaciensis Coquand, 1857, in the Ecole des
Mines Collection ex Boucheron Collection (now in the collections of the Univer-
sité Claude-Bernard, Lyons) from La Valette (Charente), a locality referred to

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Assize L? by Arnaud, and a specimen in the Sorbonne Collections ex Toucas
Collection (now in the Université Pierre et Marie Curie, Paris) from M!, at
Rentes, Cognac. This specimen is a crushed Texanites (Texanites) sp.

Apart from texanitids, there are poorly preserved Placenticeras from L! in
the Arnaud Collection and records of Baculites incurvatus (Dujardin, 1837) (e.g.
De Grossouvre 1901).

The conclusion is that it is not possible to define the base of the Santonian
in ammonite terms in Aquitaine with any great confidence. Only from occur-
rences elsewhere in France e.g. Provence (Fabre-Taxy 1963) and the Corbiéres
(Collignon & Bilotte 1983) is it possible to suggest that the appearance of Texa-
nites sensu Stricto is the best marker for the base of the stage, and even then the
name of the zone is uncertain—the fauna may be that of De Grossouvre’s
T. texanus Zone, but the correct name for the European ‘texanus’ is unresolved.

~The Zululand succession

Kennedy & Klinger (1975) presented a series of working subdivisions of the
Zululand Coniacian. These were not intended to be taken as zonal divisions but
as a working framework only until such time as the taxonomic revisions needed
to formulate a proper zonation were complete. The position has still not been
achieved, but fuller faunal lists can now be given (see below). These lists in-
cluded only those species known to be definitely from particular divisions. In the
Upper Coniacian in particular, there are many specimens (especially in Van
Hoepen’s collection) collected loose from fields and hillslopes along the
Hluhluwe and Mzinene rivers that may be from Coniacian IV or V (previously
recorded as ‘Coniacian IV-V’, e.g. by Klinger & Kennedy (1980), which
misleadingly suggested that the species ranged through these divisions).

Coniacian I

Gaudryceras cf. varagurense Kossmat, 1895, Pachydesmoceras denisoni-
anum (Stoliczka, 1865), Pachydesmoceras sp., Kossmaticeras (Kossmaticeras)
theobaldianum theobaldianum (Stoliczka, 1865), K. (K.) aff. theobaldianum
crassicostatum Collignon, 1954, Placenticeras kaffrarium Etheridge, 1904, and
Didymoceras (Didymoceras) indicum (Stoliczka, 1865). Of this assemblage, only
Kossmaticeras (K.) theobaldianum provides indisputable ammonite evidence for
a Coniacian date.

Coniacian IT

Gaudryceras varicostatum van Hoepen, 1921, Parapuzosia(?) donlisteri (van
Hoepen, 1968), Kossmaticeras (Kossmaticeras) theobaldianum crassicostatum
Collignon, 1954, K. (K.) jonesi Collignon, 1965, Placenticeras kaffrarium Ether-
idge, 1904, Peroniceras (Peroniceras) tridorsatum (Schliter, 1867), P. (P.) lepeei
(Fallot, 1885), P. (P.) westphalicum (von Strombeck, 1859), P. (P.) dravidicum
Kossmat, 1895, P. (Zuluiceras) cf. aberlei (Redtenbacher, 1875), P. (Z.) baju-
varicum (Redtenbacher, 1873), Forresteria (Forresteria) alluaudi (Boule, Le-
moine & Thévenin, 1907), F. (F.) cf. hobsoni (Reeside, 1932), Fo) (=)
madagascariensis (Collignon, 1965), Yabeiceras orientale Tokunaga & Shimizu,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 121

1926, Y. cf. orientale, Y. transiens Kennedy, Wright & Klinger, 1983, Y. anki-
natsyense Collignon, 1965, Y. costatum, Collignon, 1965, Y. manasoaense Collig-
non, 1965, Y. aff. manasoaense, Y. crassiornatum Kennedy, Wright & Klinger,
1983, nodose Baculites, Scaphites (Scaphites) meslei de Grossouvre, 1894, and
Otoscaphites arnaudi (de Grossouvre, 1894).

Coniacian III

Placenticeras kaffrarium Etheridge, 1904, Peroniceras (Peroniceras) lepeei
(Fallot, 1885), P. (P.) tridorsatum (Schliter, 1867), P. (P.) dravidicum Kossmat,
1895, P. (Zuluiceras) cf. aberlei, P. (Z.) bajuvaricum, nodose Baculites, Sca-
phites (Scaphites) meslei de Grossouvre, 1894, and Otoscaphites arnaudi (de
Grossouvre, 1894).

Coniacian IV

Peroniceras (Zuluiceras) zulu van Hoepen, 1965, P. (Z.) rarum van
Hoepen, 1965, P. (Z.) modestum van Hoepen, 1965, P. (Z.) matsumotoi sp.
nov., P. (Z.) henniei van Hoepen, 1965, P. (Z.) undulatocarinatum (van
Hoepen, 1955), Gauthiericeras obesum van Hoepen, 1955, G. listeri (van
Hoepen, 1965), G. albiforme sp. nov., G. planulatum van Hoepen, 1955, G. li-
bertae van Hoepen, 1955, G. album (van Hoepen, 1955), Protexanites (Protexa-
nites) cynci (van Hoepen, 1968), Paratexanites australis Klinger & Kennedy,
1980, P. aff. P. emscheris (Schliter, 1876) (Protexanites and Paratexanites are
very rare in situ in Coniacian IV), and nodose Baculites.

Coniacian V

Protexanites (Protexanites) cynci (van Hoepen, 1968), Paratexanites australis
Klinger & Kennedy, 1980 (many loose Protexanites and Paratexanites are prob-
ably from Coniacian V, outcrops of which are poor), Pseudoschloenbachia aff.
primitiva Collignon, 1965, and feebly nodate and non-nodate Baculites.

Santonian I

The base of the Santonian is taken at the appearance of diverse Texanites
species.

Correlation of the Zululand succession with that recognized by Kennedy in
France is rendered difficult by the low diversity of the ammonite faunas of the
European Lower Coniacian Forresteria (Harleites) petrocoriensis Zone and
Coniacian I in Zululand, plus the difficulty of finding in situ faunas in Coniacian
V.

Coniacian II and III correlate with the Middle Coniacian Peroniceras (Pero-
niceras) tridorsatum Zone, having many elements in common. Coniacian I may
be equivalent to the Lower Coniacian F. (H.) petrocoriensis Zone. It is equally
possible (at least on the ammonite evidence) that this is absent, with early Coni-
acian time represented in the sub-Coniacian unconformity. In the latter case the
sparse fauna of this division could simply be an impoverished Middle Coniacian
assemblage. Only evidence from other groups can resolve this problem.

The base of Coniacian IV in Zululand is to be correlated with the base of
the European Upper Coniacian Gauthiericeras margae Zone on the basis of the
close similarities between the Gauthiericeras in the two regions. The presence of

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

several (albeit rare) Protexanites and Paratexanites species may indicate that the
top of Coniacian IV extends into the base of the succeeding Paratexanites
serratomarginatus Zone, but P. zeilleri (de Grossouvre, 1894) is a pre-P. serrato-
marginatus Zone species (the holotype is from the condensed Calcaires Durs de
la Ribochére (petrocoriensis to margae Zones)), and there are records of rare
Protexanites below the P. serratomarginatus Zone.

Coniacian V in Zululand is probably to be correlated with the European
Upper Coniacian P. serratomarginatus Zone, but the paucity of ammonite
species and poor exposure do not permit certainty that it does not extend into
the lowest Santonian. While the appearance of Texanites in numbers marks the
Lower Santonian in both areas, it is still not clear if this is, indeed, the base of
the stage.

Madagascar

Coniacian faunas from Madagascar are the richest known, and are de-
scribed in works by Boule, Lemoine & Thévenin (1906-7), Collignon (1931,
1948, 1954, 1965), Besairie (1936), and Basse (1927, 1931, 1947, etc.).

Well illustrated as the faunas are, their stratigraphic distribution presents
many problems and detailed successions are generally lacking.

In 1960 Collignon provided a threefold zonation maintained in his later
works:

Lower Santonian Texanites oliveti Zone

Upper Coniacian Prionocycloceras guyabanum and Gauthiericeras margae
Zone

Middle Coniacian Kossmaticeras theobaldi and Barroisiceras onilahyense Zone

Lower Coniacian Peroniceras dravidicum and P. subtricarinatum Zone

Upper Turonian Romaniceras deveriai and Coilopoceras requieni Zone

In understanding these successions, the faunas from individual localities are
more revealing than those of the synthetic accounts.

In the Diego Basin Besairie & Collignon (1972: 49) note that the Senonian
consists of sandstones and sandy marls with several limestone beds, and that
there are no vertical successions. Faunas recorded indicate both Middle and
Upper Coniacian (in the sense used here) to be present.

In the Majunga Basin the same authors record only a fauna with Barroisi-
ceras from Antsira. The assemblage consists of Barroisiceras onilahyense Basse,
1947, Lewesiceras beantalyense Collignon, 1952, and Peroniceras sp.

In the Morondava Basin Besairie & Collignon (1972: 249) recognize Coni-
acian with Peroniceras rousseauxi de Grossouvre, 1894, and Lewesiceras tongo-
boryense Collignon, 1952, above post-Lower Turonian basalts. The same
relationship is observed between Demoka and Manambolo, with ‘Lewesiceras’
tongoboryense Collignon, 1952, Peroniceras rousseauxi De Grossouvre, 1894,
and ‘Hyporbulites’ masiaposensis Collignon, 1956, east of Bevitsika, and Peroni-
ceras sp. Gaudryceras analabense Collignon, 1956, and Tetragonites wieli Collig-
non, 1956, north-west of Bemonta. At Ankotrofotsy, a 7-8 m sequence of marls

CRETACEOUS FAUNAS FROM SOUTH AFRICA 123

and reefal limestones resting on basalt yielded (Hourcq 1950) Forresteria (For-
resteria) alluaudi (Boule, Lemoine & Thévenin, 1907), Kossmaticeras (Kossma-
ticeras) recurrens Kossmat, 1897, Oxybeloceras, Phylloceras (Hypophylloceras),
Gaudryceras, Puzosia, and Pachydiscus. The first of these faunas was regarded
as Lower Coniacian, but the presence of Peroniceras at the first two localities
and Forresteria (F.) alluaudi at the third all indicate the Middle Coniacian Pero-
niceras (P.) tridorsatum Zone as defined here.

In the centre of the Morondava Basin the Turonian forms a continuous strip
from Manambolo to Andranomena (Besairie & Collignon 1972) with Lower
Turonian containing Fagesia, and an Upper Turonian containing Romaniceras.
The facies is mixed, with coarse red sandstones, sometimes with silicified wood.

At Masiaposa, 3 km west of Antsarona, there is a rich fauna with (among
other stratigraphically unimportant taxa) Romaniceras (Romaniceras) deveri-
anum (d’Orbigny, 1841), Coilopoceras requienianum altesselata Collignon, 1965,
Kossmaticeras (Kossmaticeras) recurrens Kossmat, 1897, Subprionocyclus nep-
tuni (Geinitz, 1850), Subprionocyclus casterasi Collignon, 1965 (? = S. normalis
Anderson, 1958), Masiaposites spp., and Hourcquia spp.

The Turonian and Coniacian are separated by a basaltic interlude, suc-
ceeded by ‘Coniacien Inférieur’; between Manambolo and Tsiribihina, there are
- approximately 20 m of yellow-grey marls with sandy limestones, yielding many
ammonites referred to the Peroniceras dravidicum Zone. Important taxa are
P. (Peroniceras) dravidicum Kossmat, 1895, P. (P.) subtricarinatum (d’Orbigny,
1850), P. (P.) cf. tridorsatum (Schliiter, 1867), P. (P.) westphalicum (von
Strombeck, 1859), P. (P.) moureti de Grossouvre, 1894, P. (Zuluiceras) aff.
czoernigi (Redtenbacher, 1873), P. (P.) besairiei van Hoepen, 1965, P. (P.)
tenue van Hoepen, 1965, P. (P.) guerini Collignon, 1965, ‘Lewesiceras’ spp.,
Yabeiceras menabense Collignon, 1965, and Y. ankinatsyense Collignon, 1965
(see full list, by locality, in Besairie & Collignon 1972: 283).

Above, the ‘Coniacien Moyen’ and ‘Supérieur’ are represented by 30 m of
yellow marls, with the Zone a Kossmaticeras theobaldi et Barroisiceras onila-
hyense yielding: Barroisiceras onilahyense Basse, 1947, Subbarroisiceras mahafa-
lense Basse, 1946, Forresteria (Forresteria) alluaudi (Boule, Lemoine &
Thévenin, 1907), F. (F.) madagascariensis (Collignon, 1965), Yabeiceras bituber-
culatum Collignon, 1965, Y. costatum Collignon, 1965, Y. ankinatsyense Collig-
non, 1965, Y. magnumbilicatum (Basse, 1946), Kossmaticeras (Kossmaticeras)
theobaldianum (Stoliczka, 1865) and other species, plus numerous other forms
listed by Besairie & Collignon (1972: 283). The ‘Coniacien Supérieur’ Zone a
Prionocycloceras guyabanum et Gauthiericeras margae yields only the index
species and long-ranging taxa (Besairie & Collignon 1972: 284).

The base of the Santonian Texanites oliveti Zone is marked by the appear-
ance of the index species and many other taxa.

These observations indicate that there is no ammonite evidence for the
Lower Coniacian F. (H.) petrocoriensis Zone of the European standard, and
that the ‘Coniacien Moyen’ Peroniceras (P.) dravidicum Zone is equivalent to

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

part of the European P. (P.) tridorsatum Zone. The paucity of common ele-
ments between the succeeding B. onilahyense and K. (K.) theobaldi Zone and
European faunas makes comparison difficult, but Forresteria (F.) alluaudi is a
common species, suggesting that the zone is also Middle Coniacian and equiva-
lent to part of the P. (P.) tridorsatum Zone.

Comparisons with Zululand are close but pose problems. Whereas the
peroniceratids of the Madagascan P. (P.) tridorsatum Zone indicate correlation
with Coniacian II and III, the absence of Forresteria (Forresteria) in the P. (P.)
tridorsatum Zone in Madagascar but its presence in the succeeding Barroisiceras
onilahyense—Kossmaticeras theobaldi Zone is the reverse of the order seen in
Zululand where this genus occurs with Peroniceras in Coniacian II but is absent
in Coniacian III, with Peroniceras persisting.

There is insufficient evidence to resolve this apparent inconsistency.

The fauna of the Prionocycloceras guyabanum—Gauthiericeras margae Zone
has littlke in common with European faunas; the one fragment of G. margae
figured by Collignon (1965, pl. 435 (fig. 1798)) is so poor as to be indeterminate
from the figure. Correlation with the G. margae Zone is suggested. Correlation
with Zululand is easier: there are Prionocycloceras species common to this zone
and Coniacian IV.

Japan

Matsumoto (1981) has presented the most recent review of the sequence of
ammonite faunas in the Japanese Coniacian, the chief sections being in Hok-
kaido, Kyushu, and Shikoku. Interpretation of the sequence is often difficult
due to complex tectonics and the general rarity of ammonites.

A twofold division based on inoceramids is recognized, with a zone of Jno-
ceramus uwajimensis below and I. mihoensis above. These species are strati-
graphically separated in some areas (e.g. the Ikushumbets, Futaba, and
Orogawa areas), but elsewhere (e.g. the Obira area of Hokkaido) they overlap,
with I. mihoensis being the later species to appear.

Potentially important species for long-term correlation from these divisions
are as follows:

Inoceramus uwajimensis Zone

Nowakites mikasaensis Matsumoto, 1979, Prionocycloceras wrighti Matsu-
moto, 1971, P. sigmoidale Matsumoto, 1965, Prionocycloceras sp. aff. P. lenti
(Gerhardt, 1897), Ishikariceras binodatum Matsumoto, 1965, Paratexanites
serratomarginatus (Redtenbacher, 1873), Barroisiceras onilahyense Basse, 1947,
Pseudobarroisiceras nagaoi Shimizu, 1932, Barroisiceras (Basseoceras) inorna-
tum Matsumoto, 1970, F. (Muramotoa) yezoensis Matsumoto, 1970, F. (M.)
muramotoi Matsumoto, 1970, Yabeiceras orientale Tokunaga & Shimizu, 1926,
‘Harleites cf. harlei (de Grossouvre, 1894)’ (apparently a feebly ornamented
Forresteria (F.) alluaudi), and Hourcquia hataii Hashimoto, 1973. Recorded
from either this zone or that succeeding it are Peroniceras (Peroniceras) yuba-
rense Matsumoto & Muramoto, 1981 (said to be a close ally of P. (P.) besairiei

CRETACEOUS FAUNAS FROM SOUTH AFRICA 125

van Hoepen, 1965) and P. (P.) aff. platycostatum van Hoepen, 1955 (P. (P.)
dravidicum Kossmat, 1895, according to the present revision).

Elements undoubtedly from this zone listed above show that it is certainly
in part Middle Coniacian, equivalent to the European tridorsatum Zone and to
Coniacian II of the Zululand sequence. There is no ammonite evidence for
Lower Coniacian. The only anomalous record is of Paratexanites serratomargina-
tus, a species known only from the Upper Coniacian P. serratomarginatus Zone
in Europe and only from the Upper Coniacian elsewhere.

The basis of the record is given in Matsumoto (1970: 260). One specimen
was from a fallen nodule near the Coniacian—Santonian boundary, and a second
was also collected loose in Hokkaido. The only well-localized specimen is the
fragment figured by Matsumoto (1970, pl. 36 (fig. 2)) from his ‘Locality IK2112,
Go-no-sawa, Pombets, a tributary of the Ikushumbets’, zone of Jnoceramus
uwajimensis K5 (Coniacian).

Inoceramus mihoensis Zone

Peroniceras (Peroniceras) latum Matsumoto & Muramoto, 1981 (probably
P. (P.) westphalicum), Sornayceras omorii Matsumoto, 1965 (= Peroniceras
(Zuluiceras) bajuvarieum (Redtenbacher, 1873)), P. (Z.) proteus Matsumoto,
1965, Paratexanites orientalis (Yabe, 1925), and P. mikasaensis Matsumoto,
1970.

This assemblage appears to be in part equivalent to the European Peroni-
ceras (P.) tridorsatum Zone and includes species allied to those of the Gauthieri-
ceras margae and Paratexanites serratomarginatus zones, indicating it to be in
part Middle and in part Upper Coniacian. When compared with the Zululand
succession it has common (or closely comparable) species to those of Coniacian
III and IV.

Texas

The Coniacian ammonite faunas of this vast region have been described by
Young (1963), Wolleben (1967), and Young & Powell (1978).

The Coniacian is represented in the Austin Chalk and its correlatives. Over
much of Texas east of the Pecos there is a regional break between the Austin
and the underlying Eagle Ford. The base of the Austin is Coniacian (but not
lowest Coniacian) in the Dallas area, but becomes progressively older when
traced southwards, until it is Upper Turonian along the Rio Grande. The stan-
dard zonation proposed by Young is: Prionocycloceras gabrielense Zone (young-
est), Peroniceras westphalicum Zone, and Peroniceras haasi Zone (oldest).
Faunas of these zones are rather poor in number and species.

Peroniceras (P.) haasi Young, 1963 (less than twenty specimens) and Coilo-
poceras austinense Adkins, 1931 (Adkins 1931: 50, explanation of pl. 4), one speci-
men, the holotype, are from Bear Creek ‘about 3 mile west of Marchaca’, which is
said to be in both Travis and Hays counties. The only precisely localized P. (P.) haasi
specimens are fragments from ‘30 feet above the base of the Austin’ from Williamson
County, a specimen from the basal ‘15 feet of the Austin Chalk’ on

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Boudin Creek, Travis County, and the basal ‘20 feet of the Austin’ at Watters
Park, Travis County (Young 1963: 72). Inoceramus species collected from 2 m
above the base of the Austin at Cedar Hill Scarp, Johnson County, are Cremno-
ceramus deformis (Meek, 1871) (an early form of Kauffman (OUM
KT5386-9)).

According to Kauffman, Cobban & Eicher (1978), Cremnoceramus defor-
mis (early form) characterizes the upper part of the lower half of the Scaphites
preventricosus Zone of the U.S. Western Interior.

The Peroniceras (P.) westphalicum Zone yielded P. (P.) westphalicum
(seven specimens) and P. (P.) moureti (interpreted here as P. (P.) tridorsatum)
(two specimens). Prionocycloceras adkinsae Young, 1957, was inferred to be
from this zone on phylogenetic grounds alone.

The Prionocycloceras gabrielense Zone yielded P. gabrielense Young, 1957
(five specimens), P. hazzardi Young, 1957 (the holotype plus two doubtful ju-
veniles), Parabevahites sellardsi Young, 1957 (three specimens), and Protexanites
planatus (Lasswitz, 1904) (eleven specimens). Wolleben (1967), working in
Trans-Pecos Texas, added Placenticeras colquitti Wolleben, 1967 (two speci-
mens), and Prionocycloceras adkinsae Young, 1963 (one specimen) to the
P. gabrielense Zone assemblage.

It is concluded that the ammonites of the Peroniceras haasi and P. west-
phalicum Zones indicate the Middle Coniacian, and are equivalent to the Euro-
pean P. (P.) tridorsatum Zone. There is no ammonite evidence for the Lower
Coniacian.

Northern Spain

A series of papers by Wiedmann (1960, 1964, 1979, 1980) and Wiedmann &
Kauffman (1978) have documented the sequence of Upper Cretaceous ammon-
ite faunas in northern Spain, building on early work by Karrenberg (1935), Ciry
(1940, 1951), and others.

It is important to note that the highest Turonian ammonites, which include
Romaniceras (Romaniceras) deverianum (d’Orbigny), are generally separated
from the first Coniacian ammonites by a rudistid limestone complex that is 25 m
thick at Picofrentes in the Celtiberic Ranges and 50-80 m thick in the Nela area
of the Vascogotic Ranges. The succeeding Coniacian consists of sandy and glau-
conitic marls up to 200 m thick in the Vascogotic Ranges. Two important sec-
tions are given in detail as a basis for discussion.

Ollogoyen, Navarra (1°32'15"E 42°41'15’N)

The section consists of 110 m of grey clays and silty marls. Details are as
follows (after Wiedmann 1960: 718; 1979: 177, with original nomenclature):

1. Upper part of Middle Turonian (Zone VII): more than 10 m with Roma-
niceras inerme (de Grossouvre, 1889), and Proromaniceras pseudodeverianum
(Jimbo, 1894).

2. Upper Turonian (Zone VII): about 50 m with Romaniceras ornatissi-
mum (Stoliczka, 1865), R. (R.) cf. deverianumy (d’Orbigny, 1841), Collignoni-

CRETACEOUS FAUNAS FROM SOUTH AFRICA 127

ceras? sp., Pachydesmoceras denisonianum (Stoliczka, 1865), and Puzosia cf.
P. planulata (J. de C. Sowerby, 1827).

3. Lower Coniacian (I and ?II): about 45 m with Tissotia (Metatissotia) cf.
robini (Thiolliére, 1848), Reesideoceras cf. R. camerounens Basse, 1947, and
Proplacenticeras sp.

4. Middle—Upper Coniacian (Zones ?III and IV): about 50 m of massive
limestones.

5. Upper Coniacian (Zone V): sandy marls and marly limestone of the Col
d’Ollogoyen with Paratexanites cf. P. emscheris (Schliter, 1876), Gauthiericeras
aberlei (Redtenbacher, 1873), Gaudryceras vascogoticum Wiedmann, 1962, and
Scaphites compressus d’Orbigny, 1842.

Terradillos de Sedano, Burgos (0°8'05”E 42°39'10’N)

1. Middle—Upper Turonian (Zones VII and VIII) massive, well-bedded
limestones with Vaccinites.

2. Lower Coniacian (Zone I): 32 m marls and marly limestones, glauconitic
sandstones and marls with Tissotioides haplophyllus (Redtenbacher, 1873),
T. crassus Ciry, 1940, Metatissotia robini (Thiolliére, 1848), M. ewaldi (von
Buch, 1847), Parapuzosia cf. P. corbarica (de Grossouvre, 1894), to which can
be added Tongoboryceras sp. (OUM KZ 14001) and Scaphites meslei (de Gross-
ouvre, 1894) (OUM KZ 14006).

3. Lower Coniacian (Zone II): 4 m calcarenites and limestones with
Reymentoceras hispanicum (Wiedmann, 1960) and Protexanites bourgeoisi
(d’Orbigny, 1850).

4. Middle Coniacian (Zone III): 16 m grey marls with Gauthiericeras vallei
Ciry, 1940, G. margae (Schliiter, 1867), G. gordum (Karrenberg, 1935), Barroi-
siceras cf. B. sequens (de Grossouvre, 1894), Texasia iberiense (Basse, 1947),
Reesideoceras cf. camerounense Basse, 1947, R. cf. R. nicklesi (de Grossouvre,
1894), Paratexanites zeilleri (de Grossouvre, 1894), and Scaphites ex aff. arnaudi
de Grossouvre, 1894.

5. Upper Coniacian (Zone IV): 12 m of brown marls with Hemitissotia turzoi
Karrenberg, 1935, Texasia cf. dartoni Reeside, 1932, and Protexanites sp. nov.

6. Upper Coniacian (Zone V): 5 m of marls similar to those of Zone IV
with Hemitissotia lenticeratiformis Wiedmann, 1978.

7. Lower Santonian (Zone I): 7,5 m of marls and 8 m of marly limestones
with Texanites hispanicus Collignon, 1948, and Lenticeras cf. lissoni Knetchel,
1947.

8. Upper Santonian: 20 m red limestones with larger benthonic Foramini-
ferida.

At first sight this succession appears wholly incompatible with the sequence
developed by Kennedy (in press) for the French succession. De Grossouvre
(1901) had long ago recognized that the acme of Tissotidae in France occurred
at least at the level of appearance of Peroniceras. Subsequent work has
confirmed this and the following species are of the Tissotioides haplophyllus
Zone: T. haplophyllus (of which T. crassus is a synonym (Kennedy in press)),

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

Metatissotia robini and M. ewaldi, which are restricted to the P. (P.) tridorsatum
Zone in France, whereas Tongoboryceras and Scaphites (Scaphites) meslei are
commonest in this zone. It is concluded that the Tissotioides_haplophyllus Zone
of Wiedmann is equivalent to the P. (P.) tridorsatum Zone.

The only common element of the Texanites hispanicum Zone and the
French sequence is Protexanites bourgeoisi. In France this is generally upper-
most Coniacian, but Protexanites does occur low in the Upper Coniacian in
Zululand.

The fauna of the Gauthiericeras vallei Zone presents most apparent anoma-
lies. It does include G. margae, but the other recorded species of *‘Gauthiericeras’
(G. turzoi, G. gordum, G. vallei) are all Prionocycloceras. So far as can be judged
from large new collections, the Prionocycloceras at this level includes com-
pressed, flat-sided forms as well as the depressed strongly ribbed types already
_ noted, and Barroisiceras iberiense Basse (1947: 118; lectotype herein designated,
the original of Basse’s pl. 7 (fig. 2)) is one such compressed form; Barroisiceras cf.
sequens of Wiedmann & Kauffman (1978, pl. 11 (fig. 3)) and Wiedmann (1980,
pl. 11 (fig. 3)) is probably another, as may be the Reesideoceras cf. came-
rounense recorded from the zone. (Unpublished observations by C. W. Wright
on Colombian Prionocycloceras show a comparable range of variation from
depressed and coarsely ribbed to compressed and feebly ribbed specimens. )

Of other species noted by Wiedmann, Paratexanites zeilleri can be dated
only as pre-Paratexanites serratomarginatus Zone in France, and Otoscaphites
[‘Scaphites’| ex. aff. arnaudi is impossible to assess without illustrations. O. ar-
naudi itself appears in the P. (P.) tridorsatum Zone. The Gauthiericeras vallei
Zone is thus directly equivalent to the G. margae Zone of the French sequence.

The Gauthiericeras turzoi and Hemitissotia lenticeratiformis zones cannot be
correlated with the sequences in France (or indeed elsewhere), having no de-
scribed elements in common.

Romania

Szasz (1981) presented the most recent review of the Coniacian succession
in Romania. Two areas were investigated — the Babadag Basin (north Dobro-
gea) and the Cheia Valley—Vinturarita (south Carpathians). In the Babadag
Basin two successive zones were recognized, a zone of Barroisiceras haberfell-
neri below and one of Peroniceras moureti above. In the Vinturarita region, only
one zone of Forresteria (Reesideoceras) petrocoriensis was recognized, and equa-
ted with the Barroisiceras haberfellneri Zone of the Babadag Basin.

Details of the zonal assemblages are as follows:

Zone of Barroisiceras haberfellneri

Barroisiceras haberfellneri haberfellneri (von Hauer, 1866), Harleites bentori
Parnes, 1964, Yabeiceras sp. (aff. Yabeiceras orientale Tokunaga & Shimizu,
1926), Pseudokossmaticeras sp. (sp. nov?), Nowakites? macoveii Szasz, 1981,
Scaphites ex gr. compressus d’Orbigny, 1842, Gaudryceras aff. varagurense
Kossmat, 1891, Gaudryceras aff. mite (von Hauer, 1866), Damesites spp.,

CRETACEOUS FAUNAS FROM SOUTH AFRICA 129

Austiniceras cf. mobergi (de Grossouvre, 1894), Neocrioceras (Schlueterella)
kossmati (Simionescu, 1899), and Neocrioceras sp. Also recorded from this level
are Harleites harlei (de Grossouvre, 1894), Gaudryceras denseplicatum (Jimbo,
1894), and Pseudokossmaticeras ?sp. nov.

Zone of Peroniceras moureti

Peroniceras moureti de Grossouvre, 1894, Peroniceras tridorsatum
(Schliiter, 1867), Peroniceras sp., Nowakites carezi (de Grossouvre, 1894), and
Puzosia ex. gr. orientalis Matsumoto, 1954. Previous records from this level in-
clude Peroniceras czoernigi (Redtenbacher, 1873), Peroniceras rousseauxi de
Grossouvre, 1894, Eupachydiscus sayni (de Grossouvre, 1894), and ‘Anisoceras’
sertum Miller & Wolleman, 1904.

Zone of Forresteria (Reesideoceras) petrocoriensis

Forresteria (Reesideoceras) petrocoriensis (Coquand, 1865), Sornayceras aff.
omori Matsumoto, 1965, Nowakites tallavignesi (d’Orbigny, 1850), Nowakites
lemarchandi (de Grossouvre, 1894), Pseudokossmaticeras brandti (Redten-
bacher, 1873), Neocrioceras (Schlueterella) kossmati (Simionescu, 1899), Hypor-
bulites cf. bizonatus (Fritsch, 1872), Gaudryceras aff. varagurense Kossmat,
1895, Gaudryceras aff. mite (von Hauer, 1866), Scaphites ex. gr. compressus
dOrbigny, 1842, Tetragonites (Epigoniceras) sp. nov. (aff. T. (E.) epigonum)
Kossmat, 1895, Tetragonites? sp., and Puzosia sp.

The P. moureti Zone is clearly equivalent to the Middle Coniacian P. tri-
dorsatum Zone as used here. The F. (R.) petrocoriensis Zone is also part at least
equivalent to the tridorsatum Zone (some of Szasz’s F. (R.) petrocoriensis are
actually Middle Coniacian F. (H.) nicklesi (de Grossouvre) (e.g. Szasz 1981, pl.
11 (fig. 1)). The B. haberfellneri Zone also presents difficulties; the only com-
mon element with faunas we have studied is the Yabeiceras, which suggests this

zone to be Middle Coniacian.
TABLE 1

Coniacian correlation
SUBSTAGE FRANCE ZULULAND MADAGASCAR SPAIN

LOWER
SANTONIAN ‘gallicus’ I oliveti

hispanicus
?—?—?—?
lenticeratiformis
turzol
vallei

UPPER
CONIACIAN

guyabanum
Cul
margae

serratomarginatus

?—?—?—?

theobaldi
MIDDLE et :
CONIACIAN tridorsatum onilahyense hispanicum
- +
dravidicum haplophyllus

et
subtricarinatum

LOWER
CONIACIAN petrocoriensis

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Conclusions

The preceding observations lead to the correlation shown in Table 1. In the
following sections the terms Lower, Middle and Upper Coniacian are as used in
the present discussion, and not as used by previous authors.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA
Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order AMMONITIDA Hyatt, 1889
Superfamily ACANTHOCERATACEAE de Grossouvre, 1894
Family Collignoniceratidae Wright & Wright, 1951
Subfamily Peroniceratinae Hyatt, 1900

Discussion

Wright (1957: L428—429) included four genera in the subfamily Peronicera-
tinae: Prionocycloceras Spath, 1926, Gauthiericeras de Grossouvre, 1894, Pero-
niceras de Grossouvre, 1894, and Yabeiceras Tokunaga & Shimizu, 1926. The
authors follow Matsumoto (1964, 1965) in transferring Yabeiceras to the subfamily
Barroisiceratinae and Prionocycloceras to the subfamily Collignoniceratinae.

Gauthiericeras, with type species Ammonites margae Schliter, 1867, was
erected by De Grossouvre (1894: 87) for a series of species ‘dont les tours a sec-
tion subquadrangulaire présentent sur leur bord externe une quille médiane tres
saillante: en dehors de ce caractére extérieur, les cloisons offrent aussi certaines
particularités spéciales qui les placent comme formes de transition entre celles
des Mortoniceras et celles du genre suivant, Peroniceras’.

Peroniceras, with type species P. moureti de Grossouvre, 1894, was diag-
nosed (De Grossouvre 1894: 93) as including a group of species presenting ‘d’as-
sez grandes affinités avec les Mortoniceras et les Gauthiericeras, mais s’en
distinguant par leur bord externe orné de trois quilles et par une ligne suturale
beaucoup plus découpée et plus élancée qui rapelle celle des Stoliczkaia: le pre-
mier lobe latéral est, comme dans ce dernier genre, assez étroit et terminé par
une fourche bien nette. La troisi¢éme selle latérale a aussi son extrémité
supérieure bien au-dessus de celles des deux premieres.’

Concerning the suture-line of Peroniceras [‘Schloenbachia’| dravidicum,
Kossmat (1895: 190) noted that ‘der erste Lateralsattel von beiden Seiten her
stark eingeschniirt und von sehr zierlicher Gestalt, der zweite Lateralsattel ist
nicht mehr selbstandig, sondern dem ersten Lateralsattel als grosser schiefer Ast
angehanet...’. Kossmat also maintained that all Peroniceras species possessed a
similar suture-line, though not always as pronouncedly developed. In this re-
spect he mentioned Peroniceras czoernigi (Redtenbacher), P. tridorsatum
(Schliiter) and P. moureti de Grossouvre.

CRETACEOUS FAUNAS FROM SOUTH AFRICA ‘0

The diagnoses of Peroniceras and Gauthiericeras as given by De Grossouvre
and slightly elaborated on by Kossmat remained virtually unchanged during the
ensuing 60 years. In 1957 the diagnoses of Peroniceras and Gauthiericeras were
given in the Treatise (Wright 1957: L429) as:

Gauthiericeras. ‘Rather evolute; whorl section trapezoidal to subquadrate; with
entire or feebly crenulate high keel bearing a distinct groove on either side;
strong ribs spring at first in pairs, later usually singly, from single or double um-
bilical tubercles and end in strong ventrolateral tubercles.’

Peroniceras. ‘Very evolute, with oval, trapezoidal or square whorl section;
venter with 3 entire keels, middle one as high as others or higher; regular
rounded or conical umbilical tubercles are joined to slightly larger number of
similar ventrolateral ones by short straight rounded ribs; also, lateral tubercles
may occur.’

Reyment (1957: 65) added a new subgenus to Peroniceras, P. (Reginaites),
with type species P. (Reginaites) quadrituberculatum Reyment, 1957. This is an
evolute form with tricarinate venter but with four rows of tubercles. As the
authors have previously shown (Klinger & Kennedy 1980), Reginaites is more
appropriately referred to the subfamily Texanitinae Collignon, 1948, and is
closely allied to and often homoeomorphic with Plesiotexanites. In addition, it
seems to be restricted to the Upper Santonian and possibly the lowermost
Campanian (in Zululand), rather than Coniacian as initially presumed by
Reyment.

Wiedmann (1960) described a new subgenus of Gauthiericeras, G. (Ciryella),
type (and only) species G. (Ciryella) vascogoticum Wiedmann, 1960, from the
‘Lower’ Coniacian of Spain. As shown above, the type species is no older than
Middle Coniacian. This is a widely umbilicate form with a single keel, fastigiate
venter, two rows of tubercles, and a rather simple suture, in several respects in-
termediate between Peroniceras and Gauthiericeras, according to Wiedmann.

In 1965 this relatively simple classification, comprising four genera (or sub-
genera) only, was extended by the addition of a new subfamily, nine new genera
and two subgenera by the independent studies of Van Hoepen and Matsumoto.

Van Hoepen’s publication received a rather narrow circulation, and his
diagnoses are given here in full; Matsumoto’s diagnoses are summarized where
appropriate.

Reiterating Kossmat’s (1895: 190) views, Van Hoepen (1965: 4) defined
Peroniceras as ‘ammonites with three keels at a full-grown stage and with a su-
-ture-line which is deeply incised and of which the third lateral saddle is rooted in
the inner lower corner of the second lateral saddle and which is separated from
the auxiliary saddles by a more or less horizontal, sometimes deeply penetrating
lobe’.

The new subgenera of Peroniceras, P. (Zuluiceras), type species P. Zulu-
iceras zulu van Hoepen, 1965, and P. (Zuluites), type species P. (Zuluites)
modestum van Hoepen, 1965, were defined as follows:

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

P. (Zuluiceras). ‘Ammonites with umbilicus narrower than in Peroniceras sensu
stricto and whorl section more or less square in the early stages, but increasing
more rapidly in height than in Peroniceras sensu stricto and tending to become
inflated and rounded in later stages; the ribs are stronger relative to the tuber-
cles than in Peroniceras sensu stricto’ (Van Hoepen 1965: 9);

P. (Zuluites). ‘A further development of the trend exhibited by Zuluiceras; dis-
tinguished from that subgenus by the still greater whorl-height and oval whorl
section. Ornamentation gradually becomes weaker and eventually disappears
completely.’ (Van Hoepen 1965: 23).

In the new subfamily Gauthiericeratinae (ex Gauthiericeratidae van Hoe-
pen (1955), Van Hoepen (1965) included ‘ammonites with three keels at a full-
grown stage of which the suture-line is little incised; the axes of the three lateral
saddles and the two lateral lobes are more or less parallel; the third lateral
Saddle is short and high up the suture’.

Apart from the nominate genus Gauthiericeras, Van Hoepen included five
new genera in the subfamily Gauthiericeratinae: Andersonites, type species
A. listeri van Hoepen, 1965; Fluminites, type species F. albus van Hoepen, 1965;
Hluhluweoceras, type species H. fugitivum van Hoepen, 1965; Falsebayites, type
species F. peregrinus van Hoepen, 1965; and Fraudatoroceras, type species F.
besairiei van Hoepen, 1965. With the exception of Falsebayites, the type species
of which was based on two specimens, all the other taxa were based on single
specimens. In order of pagination they were defined as follows:

Andersonites. ‘Ammonites with three keels, wide umbilicus, more or less square
section, very strong, straight ribs each with two tubercles, a very large shoulder
tubercle and only slightly smaller umbilical tubercle’ (Van Hoepen 1965: 29).
Fluminites. ‘Ammonites with three keels, wide umbilicus and a section which is
at first higher than broad. The ribs are strong and slightly concave forwards.
They are at first close together, but later far apart. External saddle very low
down, reaching nearly as far back as the first lateral lobe. First lateral saddle and
first lateral lobe broad; second lateral saddle narrow; second lateral lobe very
short. Axes of these elements more or less parallel’ (Van Hoepen 1965: 31).
Hluhluweoceras. ‘Ammonites with suture-line consisting of a square, broad first
lateral saddle, a narrower second lateral saddle and a broad but very short third
lateral saddle. The external saddle is short and fairly high up. The first lateral
lobe is deep and narrower than the second lateral saddle. The axes of all these
elements are parallel to the keel, with the exception of the axis of the short but
robust second lateral lobe which extends obliquely, more or less in the direction
of the hind end of the first lateral lobe. There is only one keel which is slightly
crenulated; the venter is smooth, the section square’ (Van Hoepen 1965: 33).
Falsebayites. ‘Ammonites with a suture-line consisting of a square, broad, first
lateral saddle, a narrower but still broad first lateral lobe the sides of which are
slightly convergent backwards, a short second lateral saddle and a short second
lateral lobe (both these elements being narrower than the first lateral lobe); then
follows a very short third lateral saddle which is as broad as high, and on the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 133

umbilical surface are another three small lobes and saddles. The axes of all these
elements are parallel to the keel. The external saddle is short and fairly low down.
The whole suture is a more or less regularly meandering line. The venter is
rounded; along its middle there is a broad zone which is ever so slightly raised and
might be regarded as representing a keel. The ribs form a broad, very low band
passing across the venter in a slightly forward direction’ (Van Hoepen, 1965: 34).
Fraudatoroceras. ‘Ammonites with three keels, very wide umbilicus, square sec-
tion, fairly large, clavate shoulder tubercles and very long, narrow umbilical tu-
bercles reaching up to the middle of the flanks. External saddle is half as high as
the first lateral saddle which is as broad as high. First lateral lobe is very narrow.
Second lateral saddle is not as broad as the first. Second lateral lobe is narrow
and short. Third lateral saddle broad and short, but not as broad as the second
and not reaching as far forward nor as far backwards. Third lateral lobe narrow
and short not reaching as far forward or backward as the second. The umbilical
seam passes through its middle. The dorsal saddle is high and narrow, higher
than the first lateral saddle. There is a very small, oblique, internal lateral lobe
and a small, first dorsal lateral lobe, just touching the umbilical seam. The axes
of all these elements, except that of the internal lateral lobe, are parallel’ (Van
E¥eepen 1965: 35).

According to Matsumoto’s interpretation of Peroniceras, (Matsumoto 1965:
210) the genus is characterized by *... evolute whorls, distinctly tricarinate
venter, bi-tuberculate ribs and deeply incised, particularly specialized sutures’,
whereas Gauthiericeras (Matsumoto 1965: 210) ‘... typically has less evolute
whorls, an entire or feebly crenate ventral keel with a groove on either side,
stronger ventrolateral tubercles, often double umbilical tubercles and simpler,
normal collignoniceratid sutures’.

Matsumoto (1965: 210) furthermore believed that ‘Peroniceras is clearly
separated from Gauthiericeras (s. 1.) by the different pattern of sutures and dis-
tinctly tricarinate-bisulcate venter’.

Apart from Peroniceras and Gauthiericeras, Matsumoto referred Reginaites
and Ciryella to the subfamily Peroniceratinae and added four new genera: Cob-
banoceras, type species C. tanakai Matsumoto, 1965; Sornayceras, type species
S. proteus Matsumoto, 1965; Ishikariceras, type species [. binodosum Matsu-
moto, 1965; and Reymentites, type species R. hataii Matsumoto, 1965. Contrary
to the opinions of both Matsumoto and Wiedmann (1978: 670), the authors pre-
fer to place Reymentites, a tri- or quadrituberculate form with tricarinate venter,
in the subfamily Texanitinae Collignon, 1948, rather than in the subfamily
Peroniceratinae. Moreover, Reymentites is regarded as possibly a junior syno-
nym of Reginaites, which the authors also referred to the subfamily Texanitinae
(Klinger & Kennedy 1980).

Cobbanoceras is evolutely coiled with a wide umbilicus, subquadrate whorls
and three entire keels on the venter, of which the middle one is higher than the
lateral ones. Ribs are mostly single, and each has an umbilical and ventrolateral
tubercle. The suture is rather simple.

134 ANNALS OF THE SOUTH AFRICA.V MUSEUM

Sornayceras has a moderate to large shell, consisting of more or less evolute
whorls with wide or moderate umbilicus. The whorl section varies from
subquadrate to high subrectangular with a roughly fastigiate venter. There is a
distinct median keel bordered on either side by a shallow groove without distinct
lateral keels. The median keel is entire or feebly crenate. The flanks are orna-
mented by radial ribs which are intercalated or branched, bearing umbilical and
ventrolateral tubercles. The suture is deeply and finely incised and much compli-
cated.

Ishikariceras has a shell of moderate size, evolute with wide umbilicus. The
whorl section is subquadrate in early stages, but semi-elliptical in the adult. The
venter is unicarinate with two shallow lateral grooves. Lateral keels, if present,
are virtually indiscernible or very weak. The suture is rather simple with massive
and subquadrate elements.

Etayo-Serna (1979: 97) introduced the genus Gloriaceras, type species
~G. correai Etayo-Serna, 1979, which has peroniceratine characteristics. The
original diagnosis is as follows:

‘Conchs small to moderately large. The earliest whorls observed are discoidal
with fastigate, unicarinate median ventral region; the ornament consists of faint
falcoid or biconcave ribs that cross the venter in an acute adoral bend; the ribs
may bifurcate on the umbilical third of the flank, at this stage no tubercles are
developed.

Shortly after, the ribs weaken on the median ventral line and develop a rounded
somewhat spirally elongated external lateroventral node; the admedian ventral
end of the ribs elongates longitudinally and which soon interconnect forming al-
most continuous keels on each side paralleling the median keel but the ribs can
still be observed crossing the venter. The ribs sporadically develop weak peri-
umbilical swellings. With growth the ribs tend to straighten on flanks, become
club-like and connect the ventrolateral carinae. The periventral tubercles are
much stronger than the peri-umbilical ones which sometimes are missing. The
suture line has E/L saddle large, rectangular and bifid with the external branch
foliate and narrow at its stem; L lobe is wide, deep, bifid (pseudotrifid), L/U2
saddle is slender, taller than E/L saddle, bifid, phylloid; U2 lobe is digitated,
short. U2/U3 and U3/U1 are subequal. U3 lobe is digitated, glove-like; U1 lobe
is bifid, narrower but as deep as U2; U1/I saddle is single, with a unique phylloid
head.’

Another possible candidate for the subfamily Peroniceratinae is Neo-
gauthiericeras Collignon, 1969, type species N. zafimahovahi Collignon, 1969,
from the basal Campanian of Madagascar, erected for evolute forms with fasti-
giate venter and bituberculate ribs and suture similar to that of Gauthiericeras,
or very close to it. Collignon did not indicate whether the genus was to be
placed in the subfamily Peroniceratinae, but the name seems to imply this. The
authors (Klinger & Kennedy 1980: 111) previously suggested that Neogauthiert-
ceras may possibly be derived from Reginaites at the Santonian—Campanian
stage boundary and should thus also be referred to the subfamily Texanitinae.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 135

Patrulius & Szasz (1975: 158) described a pentacarinate form of Peroniceras
from the Coniacian of Romania. The material is fragmentary but, if more com-
plete, would possibly be eligible for separate subgeneric or generic rank.

All these taxa, with the exception of those previously referred to the sub-
family Texanitinae by Klinger & Kennedy (1980) and the enigmatic specimen
from Romania are summarized in Figure 1. From the diagnoses it appears that
the main characteristics for distinguishing between the various taxa are, in un-
weighted order:

(i) Unicarinate versus tricarinate venter,
(ii) Complex versus simple suture-line,

(111) Evolute versus involute coiling.

The dividing lines become vague in the cases of Fluminites, Falsebayites,
and Sornayceras. The validity of these various taxa and their extent is reviewed
below in their proper context, as exemplified by the various species.

Genus Peroniceras de Grossouvre, 1894

Type species
_ Peroniceras moureti de Grossouvre, 1894: 100, by original designation =
Ammonites tridorsatus Schliter, 1867: 26.

Diagnosis

Medium-sized to large, moderately to very evolute, whorl section com-
pressed trapezoidal to depressed quadrate. Venter with three subequal entire
Keels, or with the siphonal keel strongest. Ribs may be simple, prorsiradiate,
straight or concave throughout, with or without umbilical bullae that migrate out
to an inner flank position at maturity, or may arise in pairs or intercalate on
inner whorls. All ribs develop ventrolateral tubercles. Ribs may persist to ma-
turity, or weaken and decline on the adult body chamber.

Suture variable, from simple with short elements, as in ancestral Collignoni-
ceratinae, or complex, with long, deeply incised elements; L more or less bifid;
U2 small and inserted below L/U.

Discussion

The diagnoses given above (pp. 130-5) by various authors indicate con-
fusion in generic distinctions, yet Peroniceras is one of the most easily identified
taxa if the degree of complexity of the suture is ignored. A detailed description
of the type species is given below; it shares with many other species very evolute
coiling, a tricarinate venter and ornament of mainly simple ribs each with an
umbilical and a ventrolateral tubercle.

Its suture is relatively simple with rather broad, blocky, little-indented
saddles E/L and L/U and a narrow L; U2 and inner elements are all narrow and
small. Other species, such as P. dravidicum Kossmat, 1895, universally referred
to the genus, have much more complex, long, narrow, finely indented saddles

ANNALS OF THE SOUTH AFRICAN MUSEUM

136

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CRETACEOUS FAUNAS FROM SOUTH AFRICA 137

Fig. 2. External suture-line of typical Peroniceras, here referred to as

P. (Peroniceras) dravidicum-type, with highly incised elements and narrow-

stemmed saddles and lobes. Note the thin, stemmed saddle U3/U2 attached
obliquely to the base of saddle L/U2. x 1. (After Van Hoepen 1965, fig. 2a.)

with lobe U2 distinctly inserted below saddle L/U (Fig. 2). There is great varia-
tion between species in these characters and it seems absurd to pick out particu-
lar degrees in a gradational series to typify subgenera. The simple form of suture
closely resembles that of evolute species of Subprionocyclus Shimizu, 1926 (Col-
lignoniceratinae) and Protexanites Matsumoto, 1955, and other Texanitinae ex-
cept that the inner elements are narrower to accommodate the more evolute
whorl. The evolutionary trend was thus from the collignoniceratine type to the
dravidicum type.

Fraudatoroceras van Hoepen, 1965, of which Cobbanoceras Matsumoto,
1965, is a junior synonym, have type species with sutures that are only very
slightly less developed along this line than those of Peroniceras tridorsatum
(= P. moureti) and the species assigned to them, which differ in no other signifi-
cant characters, should be referred to the group of Peroniceras centred round
the type species. Indeed, Fraudatoroceras besairiei, the type species of Van Hoe-
pen’s genus Fraudatoroceras, resembles P. tridorsatum (= P. moureti) much
more than do other species always referred to Peroniceras, both in the early
stage at which tricarination sets in and in the shape of the pinched latero-umbili-
cal bullae.

Occurrence

Middle Coniacian, world-wide. Upper Coniacian of Japan and Zululand,
?Santonian of Japan.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

Subgenus Peroniceras (Peroniceras) de Grossouvre. 1894

(= Fraudatoroceras van Hoepen, 1965: 35; Cobbanoceras Matsumoto, 1965:
219: Gloriaceras Etayo-Serna, 1979: 96)

Diagnosis

Very evolute, keels strong and subequal, ribs and tubercles persist to the
end of the adult body chamber.

Discussion

Observations under the discussion of the genus Peroniceras (p. 137) explain
the basis for regarding Fraudatoroceras and Cobbanoceras as synonyms of Pero-
niceras (Peroniceras).

The genus Gloriaceras Etayo-Serna, 1979 (type species G. correaui Etayo-
Serna, 1979: 97, pl. 13 (fig. 18). pl. 14 (fig. 10). text-fig 9C-D. H) was intro-
duced for two Colombian species that were separated from Peroniceras on the
basis that they had an early stage with flexuous bifurcating ribs that cross the
venter forming chevrons. a character retained in some adults, and also differed
in suture-line. This was described as having a large, rectangular and bifid E/L
saddle with the external branch foliate and narrow at its stem, L lobe wide, deep
and bifid (pseudotrifid): saddle L/U2 slender. taller than saddle E/L. bifid and
phylloid: U2 digitated and short; Ux/U: and Us/U: subequal: U: digitated and
glove-like: U; bifid. narrower but as deep as U2; and Uy/I single with a unique
phylloid head. This was contrasted with the suture of P. moureti on the basis of
de Grossouvre’s figure, which he described as having a wide E/L. with narrow.
highly incised digitated L. UUs accessory to L/U2, and a similar interdepen-
dency of L/U—U2/U: was noted in P. dravidicum on the basis of Kossmat’s
figure (1895, pl. 23 (fig. 3)).

Juvenile Peroniceras (Peroniceras) of several species have, at 10-15 mm di-
ameter, a strong, entire or undulose siphonal keel flanked by shallow grooves
and weak lateral keels. which may also be undulose, strengthening at the ter-
mination of the ribs. This stage is rapidly replaced by a distinctly tricarinate one
in which undulations are either lost or are irregular and correspond to the nbs.
Gloriaceras correaui differs only in the more obvious development of this fea-
ture and the retention of rather more marked undulations of the lateral keels, m____
these respects retaining the phylogenetically primitive features of the ancestral
Collignoniceratinae a little longer than some other specimens. Etayo-Serna also
noted that the ribs extend to the siphonal keel in Gloriaceras. This is no more so
than in some specimens of the type species of Peroniceras (Peroniceras): the
juvenile SAS Z734, illustrated as Figure 16D-E (the original of Van Hoepen
1965, pl. 2), in fact shows all the supposedly diagnostic features of Gloriaceras,
which is here treated as a strict synonym.

Zuluiceras van Hoepen, 1965, of which Zuluites van Hoepen, 1965, and
Sornayceras Matsumoto, 1965, are regarded as synonyms, is discussed fully

'

CRETACEOUS FAUNAS FROM SOUTH AFRICA 139

below (p. 180). They represent a series of progressively more involute forms.
the earliest of which. Zuluiceras aberlet (Redtenbacher. 1873) and Z. bajuvari-
cum (Redtenbacher. 1873) occur with Peroniceras (Peroniceras). These early
forms (Zuluiceras s.s.) are inked by a contmuum to much more mvolute later
forms (“Zuluites’). the series showing progressive declme m ornament. The
earliest species are close to Peroniceras (Peroniceras) and Zuluites is m conse-
quence afforded subgeneric status only. The two differ m the greater mvolution
of Zuluiceras. the compressed whorls that tend to round; lateral keels that are
weaker than the siphonal one. and adults that retam only weak omament or
none.

Peroniceras (Peroniceras) is easily distinguished from Gauthiericeras de
Grossouvre, 1894. which has a strong. sometimes serrated siphonal keel flanked
by grooves and only moeipient lateral keels. and strong primary nbs that may
branch at umbilical or lateral tubercles.

Ishikariceras Matsumoto. 1965, has mner whorls with a subquadrate sec-
tion. coarse bituberculate mbs and a strong siphonal keel but no. or only very
weak. side Keels. the body chamber being oval and virtually smooth with only
constrictions and a single Keel.

Reginaites Reyment. 1957. of the Texanitmae, of which Reymentites Matsu-
moto. 1965. is a synonym (Klinger & Kennedy 1980). is ticarmate with umbili-
cal and ventrolateral tubercles only on nuclei, the ventrolateral splitting m two
at a later stage when feeble lateral tubercles may appear.

Occurrence

Middle Coniacian. world-wide. P. (Peroniceras) |'Cobbanoceras’| tanakai
Matsumoto. 1965. is questionably dated as approximately Santoman (Matsu-
moto 1965: 220).

Peroniceras (Peroniceras) tridorsatum (Schiiter. 1867)

Figs 3-15. 16D-E

Ammonites tridorsatus Schiiter. 1867: 26. pl. 5 (fig. 1)-

Ammonites ci. tridorsatus Schiiiter. 1876: 138. pl. 41 (figs 35)-

? Ammonites (Schloenbachia) cz6rnigi Redi- Fallot. 1885- 229. pl. 1 (fig. 1)-

Peroniceras subiricarinatum wat. tridorsaian Schiater- de Grossouvre. 1894: 96. pi. 10 (figs
2-3). pl 11 (fig. 1).

Peroniceras mouret: de Grossouvre, 1894: 100. pl. 11 (figs 34). text-figs 37. 39. Pruvost. 1910-
367_ Siolley. 1916- 89. Diener, 1925: 150. Venzo_ 1936: 101. pl. 9 (fis 1). Young. 1963: 73.
prada pl 27 (fig. 1). text-fig. 13(a). ?Collignon 1965- 60. pl. 440 (fiz. 1807). Szasz

& Lacainsu. 1974- 209. pl. 6 (fig. 3). pl. 7 (fig. 3). Wiedmann. 1978: 669. pl. 2 (Ags +5).
Amedro & Robaszynski. 1978: 37. pl. 1 (fig. 2). Colhgnon e al. 1979: 392. pl. 2 (fig. 3)-
Szasz_ 1981- 106. pl. 9 (figs 34).

Peroniceras (Peroniceras) besairiei van Hoepen. 1965: 4. pl. 1, text-fig_ la.

Fraudatoroceras besairiei van Hoepen. 1965: 36, pl. 27. text-fig. 92D.

Peromceras besairiei van Hoepen- Collignon. 1965: 34. pl. 437 (fig. 1805). pl. 438 (fic. 1805).

Peroniceras rousseauxi de Grossouvre. 1894: 102. pl 11 (fs. 5). Diener. 1925- 151-

Peroniceras ci_ moureti de Grossouvre: Maury. 1907: 87. Diener. 1925: 151-

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

Peroniceras tridorsatum (Schliter): Stolley, 1916: 89. Diener, 1925: 152. Van Hoepen, 1965: 7,
pl. 2. Amedro & Robaszynski, 1978: 36, pl. 2 (fig. 1), pl. 3 (figs 1-2). Szasz, 1981: 107, pl.
2) ie, 2),

? Peroniceras tricarinatum @Orbigny: Scupin, 1913: 93, pl. 3 (fig. 5).

Peroniceras lepeei (Fallot): Mirauta & Mirauta 1964: 336, pl. 7 (fig. 2), pl. 11. ?Joja & Chiriac,
1964: 46, pl. 1 (fig. 5).

? Peroniceras cf. tridorsatum (Schliter): Collignon, 1965: 58, pl. 439 (fig. 1805, non 1806).

? Peroniceras aff. tridorsatum (Schliter, 1867): Amedro & Robaszynski, 1978: 37, pl. 2 (fig. 2).

Peroniceras sp. (aff. Peroniceras rousseauxi Grossouvre): Szasz & Lacatusu, 1974: pl. 2 (fig. 4),

pl. 7 (fig. 4).
Peroniceras (Peroniceras) tridorsatum (Schliter, 1867): Kennedy, in press.

Type specimens

The holotype of the species, from the Coniacian of Westphalia, is lost.
The holotype, by original designation, of Peroniceras moureti de Grossouvre,
~ 1894, is the original of De Grossouvre’s pl. 11 (fig. 4) from the ‘Calcaires
tendres gréseaux de la base de la Craie de Villedieu. Courtiras, pres Venddme
(Loir-et-Cher)’. It is in the collections of the Ecole des Mines, now in
the Université Claude Bernard, Lyon. Paratypes include the specimen figured
by De Grossouvre as his pl. 11 (fig. 3), an unregistered specimen in the
collections of the Sorbonne (now in the Université Pierre et Marie Curie, Paris)
from Assize L' of Arnaud at La Boulenerie, near Jonzac (Charente-Maritime)
and a specimen (ML 7) from the Coniacian of Lezennes (Nord) in the Musée
Gosselet, Lille.

The holotype, by monotypy, of Peroniceras rousseauxi de Grossouvre,
1894, is an unregistered specimen in the collections of the Sorbonne from Assize
L! of Arnaud at St. Simon-de-Jonzac (Charente-Maritime).

The holotype, by original designation, of Peroniceras (Peroniceras) besairiei
van Hoepen, 1965, is SAS 2248.

The holotype, by original designation, of Fraudautoroceras besairiei van
Hoepen, 1965, is SAS Z756.

Material

SAS Z1117, SAS A63, and NMB D1145a-e, all from locality 72, degraded
river cliff on the north side of the Mzinene River downstream from the junction
with the Munywana River, Zululand, St. Lucia Formation, Coniacian II and ILI.
SAS Z1590 and Z1522 from locality 91, degraded river cliff and artificial cut
near the Hluhluwe River estuary, east-south-east of Hluhluwe, Zululand,
St. Lucia Formation, undifferentiated Coniacian debris. SAS Z1623a—b, Z1607,
and SAM-—PCZ5942 from locality 92, bulldozer scrapings and adjacent hillslopes
on the farm Panplaas east-south-east of Hluhluwe, Zululand, St. Lucia
Formation, Coniacian II and HI. SAS Z248, Z730 and Z1120 from locality 73,
degraded river cliffs on the north bank of the Mzinene River downstream of
the Munywana River junction, Zululand, St. Lucia Formation, Coniacian I
and III.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 141

Fig. 3. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). A. SAS A63.
B. NMB-D1145a. C-E. SAS Z1117. All x 1.

142

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). NMB D1145e.
Specimen with distinct latero-umbilical tubercles on inner whorls, but indistinct
on outer whorls, as in P. (P.) lepeei (Fallot, 1885). x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 143

| Fig. 5. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). NMB D1145C.
| Specimen with narrower umbilical width than usual. x 1

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

5 mm

Dimensions
D Wb Wh Wb/Wh U R
D1145a 31,0 9,0(29,0) 9,0(29,0) 1,0 16,0(51,6) 32:34
Z1623b ADT eV ASPZGOE9) U2 S(29,2)) LOS 20,5(48,0) 28:28
Z1623a 52,4 17,0(32,4) 12,5(23,8) 1.3 27,0(52,0) 26:26
aa) 5930, 17402858) 17,0(28,8) 1,0 SRO (5225) 38:38
Z1607 60,0 17,0(28,0) 1772955) 1,04 29,1(48,5) 30:30
Z1120 63,3 19,0(30,0) 19,6(30,9) 0,96 30,5(48,0) Diem
Z1590 S258) 2456(2987) 22,0(26,5) OL 48 ,0(57,9) 28:28
Z730 ILO ZS (29.11) D303 (2279) 1,09 60,0(59,0) 30:30
21522 IZOD 30F0 Cor) 26,0(23,2) oS 65,0(58,0) 24:24
D1145c 119,0 — 40,0(33,6) — 55,0(46,2) —
Z756 12050" 33,7(@8,0) Zell 2251) 123) 71,0(59,0) 35°33
D1145e 124,0 36,0(29,0) 34,0(27,4) 1,05 66,0(53,2) 38:38
Z248 ISZ0 S30(Z50) 33,0(25,0) LO 73,0380) 29:29

Fig. 6. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). External suture of SAS A63.

Description

Coiling is very evolute, with an umbilical width of the order of 48 to 59 per
cent of the diameter, and with the whorls overlapping only slightly.

The whorl section is rounded to subrectangular, varying from compressed
to depressed, with the greatest costal width at the umbilical tubercle.

There is an initial smooth stage, which may persist to a diameter of 9 mm;
thereafter umbilical tubercles connecting to ventrolateral tubercles via weak ribs
appear. With increasing diameter all ornament becomes stronger, with promi-
nent umbilical bullae situated well away from the umbilical edge on the flanks
and connected to rounded or slightly clavate ventrolateral tubercles by promi-
nent ribs. The ribs are generally single but occasional bifurcations occur, es-

CRETACEOUS FAUNAS FROM SOUTH AFRICA 145

pecially on the inner whorls. All ribs are slightly prorsiradiate and curved.
Strong spiral ridges are commonly developed.

The venter is prominently tricarinate, with the central keel strongest. The
suture is relatively simple but variable with blocky elements (Figs 12-13).

Discussion

Kennedy (in press) has discussed the difficulties in interpreting Schliter’s
lithograph of the lost holotype, and concluded that the variable, closely and sim-
ply ribbed Peroniceras (P.) with persistent umbilical-umbilicolateral bullae that

Fig. 7. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). SAS Z248, the holotype of
P. (Peroniceras) besairiei van Hoepen, 1965. X 1.

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B

Fig. 8. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). SAS Z248 the
holotype of P. (Peroniceras) besairiei van Hoepen, 1965. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 147

occur in France and Germany (P. tridorsatum, moureti, and rousseauxi) rep-
resent but a single species, a view followed here.

P. (P.) lepeei (Fallot, 1885) differs from the present species in having deli-
cate dense ribs that lack umbilical bullae; it is discussed further below. There
are, however, transition forms such as NMB D1145e (Fig. 4) that have a more
angular whorl section and weakening ornament on the outer whorl.

P. (P.) besairiei van Hoepen, 1965, is based on a depressed variant of this
species that is close to the holotype of De Grossouvre’s P. (P.) rousseauxi, also
regarded as a synonym. Van Hoepen was misled in his interpretation of P. (P.)
besairiei by the illustrations of Besairie (1930, pl. 64 (figs 4-6)) and Venzo (1936,

Fig. 9. Peroniceras (Peroniceras) tridorsatum (Schliiter, 1867). SAS Z786, the holotype of
Fraudatoroceras besairiei van Hoepen, 1965. X 1.

148

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Peroniceras (Peroniceras) tridorsatum (Schliiter, 1867). SAS Z786,
the holotype of Fraudatoroceras besairiei van Hoepen, 1965. x 1,2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 149

Fig. 11. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867).
SAS Z1522, with stronger lateral ornament than the other specimens, and
distinct spiral ornament. x 1.

O 20mm

[a

Fig. 12. Peroniceras (Peroniceras) tridorsatum (Schliiter, 1867).
Complete suture-line of SAS Z786.

150

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Peroniceras (Peroniceras) tridorsatum (Schliiter, 1867). Suture-

lines of different specimens. A. SAS Z248, part of the suture-line of the

holotype of Van Hoepen’s Peroniceras (Peroniceras) besairiei, alleged to

be of the P. (P.) dravidicum type. B-C. SAS Z1623. D. SAM-—PCZ5942.
E. SAS Z730.

CRETACEOUS FAUNAS FROM SOUTH AFRICA ifoyl

pl. 11 (fig. 9)). According to Van Hoepen (1965: 36), P. (P.) besairiei and his
Fraudatoroceras besairiei are ‘practically the same in shape and ornamentation.
... The only difference is the suture-line’. He believed that F. besairiei had a
simple block-type suture-line, whereas P. (P.) besairiei (interpreted according to
Besairie’s figures) was alleged to have a complex Peroniceras (dravidicum)-like
suture-line. Examination of the holotype of P. (P.) besairiei shows that the
suture-line (which according to Van Hoepen could not be developed) is of the
same type as that of F. besairiei (Figs 12-13). They are synonyms, homonyms
when in the same genus, and both synonyms of P. (P.) tridorsatum. Van
Hoepen also erred in regarding his P. (P.) besairiei as the same as Besairie’s
(1930: 634, pl. 46 (figs 4-6)) Peroniceras subtricarinatum and Venzo’s (1936: 99,
pl. 10 (fig. 1) pl. 11 (fig. 9)) Peroniceras dravidicum, and he assumed that the
suture-lines would be identical. The specimens described by Besairie may partially
include P. (P.) tridorsatum as some specimens were said to have single ribbing,
but the figured specimens definitely do not belong to P. (P.) tridorsatum as the
complex suture-line and bifurcating ribbing shows. The specimen figured by
Venzo probably belongs to P. (P.) dravidicum. Collignon (1965: 54, pl. 437 (fig.
1803), pl. 438 (fig. 1803)) described and figured a specimen under the name of
Peroniceras besairie:, but unfortunately did not describe the suture-line; neither
is it visible in the figures.

Fig. 14. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). A-C. SAS Z1120.x 1.

Sz ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. Peroniceras (Peroniceras) tridorsatum (Schliiter, 1867). SAS Z756, the outer
phragmocone whorl of the holotype of Fraudatoroceras besairiei van Hoepen, 1965. xX 1.
(See also Figs 9-10.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 153

Specimens the authors refer to Peroniceras (P.) subtricarinatum may resem-
ble P. (P.) tridorsatum superficially, but differ in having occasionally bifurcating
ribs and a more complex, incised suture-line.

P. (P.) ninakawai (Matsumoto 1965: 223, pl. 37 (fig. 2), text-figs 7-8) from
the ‘Santonian’ of Hokkaido has a narrower umbilicus (41-42 %) than many
P. (P.) tridorsatum specimens (48-49 %) and develops strong, widely spaced
auricular ribs on the outer whorls.

Occurrence

Middle Coniacian P. (P.) tridorsatum Zone of western, central and
southern Europe, Gulf coast of the United States, Coniacian II and III of Zulu-
land.

Peroniceras (Peroniceras) lepeei (Fallot, 1885)
Figs 16A-—C, 17-18

Ammonites (Schloenbachia) L’Epeei Fallot, 1885: 231, pl. 1 (fig. 2.).
Peroniceras L’Epeei (Fallot): Desio, 1920: 208, pl. 12, (figs 6-7), pl. 14 (fig. 6). Diener, 1925:
150. Kennedy, in press.

Type specimen

The holotype, by monotypy, is the original of Fallot (1885 pl. 1 (fig. 2)),
from the Grés Verts de Dieulefit of Dieulefit, Dr6me, south-eastern France,
Middle Coniacian, Peroniceras (P.) tridorsatum Zone. The specimen is pre-
served in the collections of the Laboratoire de Géologie of the Faculté des
Sciences, Grenoble.

Material

SAS Z1568 from locality 92, bulldozer scrapings and adjacent hillslopes on
the farm Panplaas, east-south-east of Hluhluwe, Zululand, St. Lucia Formation,
Coniacian II or III. SAS A1989 from locality 72, degraded river cliff on the
north side of the Mzinene River downstream from the junction with the Mu-
nywana River, Zululand, St. Lucia Formation, Coniacian II or II.

Dimensions

D Wb Wh Wb/Wh U
Z1568 52,0 15,5(29,8) 14,2(27,3) 1,09 POS.)
Description

Coiling is polygyral, very evolute with an umbilical diameter of 53 per cent.
The whorl section is variable, both between specimens and at different stages of
ontogeny (Fig. 18A). It is initially ovoid and compressed, but thereafter
rectangular with flattened flanks.

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 16. A-—C Peroniceras (Peroniceras) lepeei (Fallot, 1885). SAS Z1568.
D-E. Peroniceras (Peroniceras) tridorsatum (Schliter, 1867). SAS Z734.
All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA £55

Fig. 17. Peroniceras (Peroniceras) lepeei (Fallot, 1885). Large specimen showing the
relatively simple suture-line and faint spiral ornament on the body chamber.
SAS A1989.x 1.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

L
U2
G
@) 10 mm
nen aneeememmeermerrs

Fig. 18. Peroniceras (Peroniceras) lepeei (Fallot, 1885). A. Whorl section of SAS Z1568.
x 1,5. B. External suture of SAS Z1568. C. External suture of SAS A1989.

It remains so in SAS Z1568 (Fig. 16A—C), but reverts to being rounded in
SAS A1989 over the greater part of the phragmocone and body chamber, with
no clearly defined umbilical edge (Fig. 17).

On the innermost whorls of SAS Z1568 the flanks are completely smooth
up to a diameter of 6 mm, and the venter is fastigiate with a feeble siphonal
ridge. Beyond that diameter tubercles appear in the umbilical seam of the
succeeding whorl, and at a diameter of 18 mm adult ornament is fully devel-
oped. This consists of delicate, concave, prorsiradiate ribs each bearing a clavate
tubercle. There are 50 ribs per whorl in SAS Z1568 (Fig. 16 A—C). Ribbing is
generally simple but occasional bifurcations occur in SAS A1989 (Fig. 17). Parts
of the body chamber of both specimens show delicate spiral ornament. The
venter is tricarinate, with the median keel strongest. The suture is rather simple,
with massive saddles and lobes (Fig. 18 B—C).

CRETACEOUS FAUNAS FROM SOUTH AFRICA 157

Discussion

This is a very poorly known species, thus far described only from the
Middle Coniacian of France and Italy. The species is characterized by the deli-
cate, concave prorsiradiate ribs, absence of umbilical tubercles, and flattened
flanks.

It is very close to P. (P.) tridorsatum (= P. (P.) moureti), and the two occur
together. De Grossouvre (1894: 101) had already pointed to the similarities be-
tween the species, and Desio (1920: 208) even included P. (P.) moureti in the
synonomy of P. (P.) lepeei. The only difference is the absence of umbilical tu-
bercles in P. (P.) lepeei. The specimen figured by Schliiter (1876, pl. 4 (figs
3-5)) as Ammonites cf. tridorsatus, and included in the synonomy of P. (P.)
moureti by De Grossouvre, has umbilical tubercles on the inner whorls but these
become progressively weaker on the outer whorl. Similar though more coarsely
ornamented specimens of P. (P.) tridorsatum also occur in Zululand and this
suggests a close relationship between the two species.

The suture-lines of the two species are similar, having relatively simple
blocky elements but lack the narrow-stemmed elements typical of the P. (P.)
dravidicum-type suture.

Occurrence

Middle Coniacian, P. (P.) tridorsatum Zone of Dieulefit (Drome), Conia-
cian (precise horizon unknown) of Touraine (condensed Calcaires durs de la
Ribocheére) in France, Italy, and Coniacian II or II of Zululand.

Peroniceras (Peroniceras) subtricarinatum (d’Orbigny, 1850)
Figs 19A-B, D-E, 20-23

Ammonites tricarinatus d’Orbigny, 1841: 307, pl. 91 (figs 1-2). Schliter, 1872: 44, pl. 13 (figs
1-4). Fritsch 1872: 26, pl. 1 (figs 1, 3, non 2).

Ammonites subtricarinatus dOrbigny, 1850: 212. ? Ooster, 1858: 151, pl. 26 (figs 10-11).
Drescher, 1863: 331, pl. 8 (figs 2, 4). Non Stoliczka, 1865: 54, pl. 31 (fig. 3).

Ammonites (Schloenbachia) subtricarinatus d’Orbigny: Fritsch, 1889: 70; 1893: 74; 1897: 36.

Peroniceras subtricarinatum d’Orbigny: de Grossouvre, 1894: 94 (pars), pl. 10 (fig. 1, non 2-3),
non pl. 11 (figs 1-2). Non Sturm, 1900: 60, pl. 3 (fig. 5). Pervinquiere, 1907: 249. Pruvost,
1910: 365. Stolley, 1916: 89. Burckhardt, 1921: 112, pl. 29 (figs 5-6 only). Diener, 1925:
151. Besairie, 1930: 634, pl. 66 (figs 4-6), text-fig. 2.1. Andert, 1934: 406. Collignon, 1965:
58, pl. 439 (fig. 1804). Thomel. 1969: 119, pl. F (figs 1-2). Amedro & Robaszynski, 1978:
Sapplemiaues 3) Collicnonrer al. 19792 390; ple 1 (fis. 1):

Schloenbachia (Peroniceras) subtricarinatum d’Orbigny: Boule, Lemoine & Thévenin, 1907: 22.

Peroniceras aff. tricarinatum d’Orbigny: Bose, 1928: 268, pl. 11 (figs 2-4).

Peroniceras tenuis van Hoepen, 1965: 7, pl. 3, text-fig. 1b-e.

Peroniceras tenue van Hoepen: Collignon, 1965: 66, pl. 443 (fig. 1814).

Peroniceras (Peroniceras) subtricarinatum (d’Orbigny, 1850): Kennedy, in press.

Type specimens

D’Orbigny (1841: 308) referred two specimens to his Ammonites tricarina-
tus, which he subsequently (1850: 212) renamed Ammonites subtricarinatus be-

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 19. A-B, D-E. Peroniceras (Peroniceras) subtricarinatum (d’Orbigny, 1850). The lectotype
from the Lower Coniacian near Rennes-les-Bains, France. X 1. CC. Gauthiericeras margae
(Schliiter, 1867), holotype. x 0,6. (See Figs 94-95 for details.)

SAR ee n,m

159

CRETACEOUS FAUNAS FROM SOUTH AFRICA

L9

‘oO x

.

COOLZ SVS

(OS8T ‘AUsIQIO,.P) wnNoUlIDI14NgQNS (SD4dIIUOAAG) SDAIDIUOAA ‘(YZ

“SI

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160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 21. Peroniceras (Peroniceras) subtricarinatum (d’Orbigny, 1850). SAS Z511, the holotype of
Van Hoepen’s Peroniceras (Peroniceras) tenuis (1965: 7, pl. 3, text-figs 1b, e); a compressed speci-
men representing the inner whorls of P. (P.) subtricarinatum. X 1. (See also Fig. 22.)

cause of prior usage of the combination Ammonites tricarinata by Poitiez &
Michaud (1838). The syntypes survive in the D’Orbigny Collection (now in the
Muséum Nationale d’Histoire Naturelle in Paris) under the catalogue number
7183. The larger of these, reillustrated here as Fig. 19A—B, D-E is herein desig-
nated lectotype of the species. It is from the environs of Sougraines, near
Rennes-les-Bains (Aude).

Material

SAS Z1002, Z511, SAM-—PCZ5943 from locality 93, hillslopes on the
boundary of lots H101 and H102, east-south-east of Hluhluwe, Zululand, St.
Lucia Formation, Coniacian II, SAS Z1521 from locality 92, bulldozer scrapings

161

CRETACEOUS FAUNAS FROM SOUTH AFRICA

C17 “SIq 998) “90 x “SO6T ‘UodooH{ UA sinual (‘d) “d
O} Je[IWIIS spIOyM JOUUT YM UoWUTOOdS ‘EP6SZOd-WVS ‘(OS8I ‘AUSIQIO,.P) MéniDUlADILIGNS (SDAdIIUOAIg) SDAINIUOAI “ZZ “BIA

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

and adjacent hillslopes on the farm Panplaas, east-south-east of Hluhluwe,
Zululand, St. Lucia Formation, Coniacian II and III.

Dimensions
D Wb Wh Wb/Wh U R
7éS)\\Il 13770 37,0(28,0) 34,0(35,8) 1,08 TxA 0(SD53)) 29:42
Z1002 152.00 33,0C1em) 31,5(20,7) 1,05 92,0(60,2) 26 : 36
PCZ5943. 195,0 —40,0(20,5) 41.0(21.0) 0,97 113,0(57,9) 12x2:18x2
A 100 — XO@DS5) 27,0(22,5)’ 1,0 70,0(58,3) 14x2:19x2
Description

Coiling is polygyral and evolute with an umbilical diameter of 55 to 60 per
_ cent of the total diameter. The adult whorl section is depressed, fastigiate with a
prominent central keel and subordinate lateral keels and more or less parallel
flanks in intercostal section. The dorsal zone of impression is very shallow and
whorl overlap minimal. In the juvenile stage the whorl section is more rounded
and the venter appears unicarinate. In costal section maximum width is across
the dorsal third of the flanks.

Ornament consists of prominent rounded to bullate umbilical tubercles
and smaller spirally elongated ventrolateral tubercles connected by weak ribs.
Bifurcations are present both on the inner and outer whorls and in some cases
one of the ribs of a bifurcating pair is weaker than the other, thus creating
the appearance of intercalated ventrolateral tubercles. On the outer phrag-
mocone whorls the umbilical tubercles may enlarge considerably and migrate
away from the umbilical edge. Strength of ornament in the various specimens
studied varies considerably as the figures show. In SAS Z511 (Fig. 21) faint
spiral ornament is visible in the area between the ventral tubercles and the lat-
eral keel.

The suture is finely subdivided. Details may vary considerably in response
to position relative to ornamentation which affects mainly width and incision of
the elements (Fig. 23).

Discussion

This species has been treated in various ways since its original description.
De Grossouvre (1894: 94 et seq.) included Schliter’s Ammonites tridorsatus in
the synonomy of P. (P.) subtricarinatum. According to De Grossouvre’s inter-
pretation they could, however, be separated at varietal rank. The typical P. (P.)
subtricarinatum has a nearly equal number of single and bifurcating ribs,
whereas P. (P.) tridorsatum has all the ribs simple. The lectotype of P. (P.) sub-
tricarinatum, here refigured as Figure 19A—B, D-E shows D’Orbigny’s figures to
be considerably restored, especially as far as the inner whorls are concerned.
The specimen does, however, show the typical common and distinctive bifurca-
tions on the outer whorls.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 163

¢) 10 mm
ey

Fig. 23. Peroniceras (Peroniceras) subtricarina-

tum (d’Orbigny, 1850). Two successive sutures

of SAS Z1002 to illustrate influence of orna-

ment on details of suture, in this case especially
the width of the stem of saddle L/U2.

Desio’s (1920: 206, pl. 25 (fig. 3)) P. subtricarinatum var. etruscum has
weak ornament on the inner whorls, and lacks umbilical tuberculation on the
outer whorls. Furthermore, the whorl section is much higher than wide. As
Desio (1920: 207) pointed out, the specimen is very close to P. (P.) dravidicum.

P. cf. subtricarinatum var. tridorsatum of Desio (1920: 205, pl. 12 (fig. 10))
is poorly preserved, but because of the single ribbing falls beyond the limits of
this species.

P. (P.) tenuis van Hoepen (1965: 7, pl. 3, text-figs 1b-e) merely represents
the inner whorls of a weakly ornamented form of P. (P.) subtricarinatum as Fig-
ures 21—22 show, and is thus included in the synonomy.

P. haasi Young (1963: 72, pl. 34 (figs 3-4), pl. 35 (figs 1-2)) from the Coni-
acian of Texas has the general appearance of P. (P.) subtricarinatum but weaker
ornament. Given more material it might be possible to include it in the strict
synonomy of P. (P.) subtricarinatum, as was done with P. (P.) tenuis van
Hoepen.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence

Middle Coniacian P. (P.) tridorsatum Zone of western and central Europe,
north Africa, Madagascar, and Mexico. Coniacian II and II of Zululand.

Peroniceras (Peroniceras) westphalicum (von Strombeck, 1859)
Figs 24-28 |

Ammonites westphalicus von Strombeck, 1859: 56. Schliiter, 1867: 30, pl. 6 (fig. 2); 1872: 45, pl.
13 (figs 5-6).

Peroniceras westphalicum (Schliter): de Grossouvre, 1894: 98, pl. 12 (figs 1, 4), text-fig. 38.
Pruvost, 1910: 366. Non Scupin, 1913: 94, pl. 4 (fig. 8). Desio, 1920: 214, pl. 12 (fig. 2), pl.
13 (fig. 3). Diener, 1925: 152. Adkins, 1933: 407, 453. ? Andert, 1934: 407. Young, 1963:
74, pl. 28 (figs 2-4), pl. 29 (figs 1-2), text-fig. 15d. Collignon, 1965: 60, pl. 440 (fig. 1807).

Peroniceras westphalicum (Schliter) var. australis Venzo, 1936: 100, pl. 9 (fig. 4), pl. 11 (fig. 8).

- Peroniceras stephaninii Venzo, 1936: 102, pl. 9 (fig. 3), pl. 11 (fig. 10).

Peroniceras guerini Collignon, 1965: 64, pl. 442 (fig. 1811).

Peroniceras cf. westphalicum (Schliter), 1867: Amedro & Robaszynski, 1978: 37, pl. 1 (fig. 1).

? Peroniceras latum Matsumoto & Muramoto, 1981: 52, pl. 6 (fig. 1), pl. 7 (fig. 1), text-fig. 1.

Type specimens

Von Strombeck’s originals, from the Coniacian of Westphalia, have not
been traced. They were presumably in Berlin.

Material

NMB D1146, D1145, D1060, SAS A2941, A1972, all from locality 72, de-
graded river cliffs on the northern banks of the Mzinene River downstream of
the junction with the Munywana River, Zululand, St. Lucia Formation, Coni-
acian II and III. SAS Z1616 from locality 93, bulldozer scrapings and adjacent
hillslopes around the pumping station on the farm Panplaas, east-south-east of
Hluhluwe, Zululand, St. Lucia Formation, Coniacian II and III.

Dimensions

D Wb Wh Wb/Wh U R
Z696 144.0 41 ,0(28,4) 43,0(29,8) (U5 85,0(59,0)- 1se2 aleve
A1972 155,0 35,0(22-5) 40,0(25,8) 0,87 89,0(57,4) —
Z1616 195,0 — 50,0(25,6) — 100,0(51,2) 1ex2 ier
D1146 195).0) 49 ,0(25,2) 56,0(28,0) 0,87 97,0(49,7) PAVED)
Description

Coiling is evolute with an umbilical diameter of about 50 to 60 per cent of
the total diameter. The whorl section is compressed throughout, higher than
wide with little-inflated flanks, and has a very low umbilical wall and broadly
rounded venter.

Ornament on the inner whorls is more or less of the P. (P.) dravidicum type
with frequently bifurcating ribs, though not as strongly developed as in the latter
species. With increasing diameter the umbilical tubercles become progressively

;

CRETACEOUS FAUNAS FROM SOUTH AFRICA 165

weaker, and eventually lateral ornament consists of broad, low ribs that end or
weaken considerably before reaching the clavate ventrolateral tubercles. At the
same time there is a tendency towards a decrease in the number of bifurcations.
On the outer whorls of the specimens available, virtually all ribs are single. De-
tail of the ornament varies considerably, from densely costate as in SAS Z1616
(Fig. 27) to sparsely costate in NMB D1146 (Fig. 24).

The species may grow to a very large size. The suture is highly incised
(Fig. 28).

Discussion

In the absence of type specimens, interpretation of this species presents
difficulties. It is here interpreted in terms of the specimen from near Altenessen

Fig. 24. Peroniceras (Peroniceras) westphalicum (von Strombeck, 1859). NMB D1146,
specimen with typical ornament consisting of low, broad ribs. x 0,6.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 25. Peroniceras (Peroniceras) westphalicum (von Strombeck,
1859). NMB D1060. Evolute specimen with typical ornament.
x 0,69.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 167

Fig. 26. Peroniceras (Peroniceras) westphalicum (von Strombeck, 1859). SAS Z696, specimen
with ornament approaching that of P. (P.) dravidicum Kossmat, 1895. x 0,77.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 27. Peroniceras (Peroniceras) westphalicum (von Strombeck, 1859). SAS Z1616, specimen
with ornament transitional to P. (P.) dravidicum Kossmat, 1895. x 0,70.

figured by Schliiter (1872, pl. 13 (figs 5—6)). This clearly shows the low, rounded
and broad bullae, which clearly distinguish the species from the majority of
other peroniceratids.

One of the specimens figured by De Grossouvre (1894, pl. 12 (fig. 1)), from
the Craie de Villedieu of Touraine, lacks the typical broad lateral ornament and
may possibly be a transition towards P. (P.) subtricarinatum.

P. westphalicum var. italicum Desio (1920: 216, pl. 16 (fig. 4)) lacks the bul-
lae of typical P. (P.) westphalicum and has, instead, sharp-crested, narrow,
widely spaced lateral ribs. In this respect it bears resemblance to P. stefaninii

CRETACEOUS FAUNAS FROM SOUTH AFRICA 169

Sa mae ¢ \
a “ee i] \
5 ; |
\ CA |
\
oe
J at =~ 4
| cs Coy ere
| Looks ala ana cote
— = jae La oe
lf :
Ss
@) : 10mm

Fig. 28. Peroniceras (Peroniceras) westphalicum (von Strombeck, 1859).
Partial suture-line of SAS Z996.

Venzo (1936: 102, pl. 9 (fig. 3)) from the Coniacian of Zululand. Specimen
NMB D1146 (Fig. 24) also shows a tendency towards narrowing of the ribs as in
these forms. They may thus safely be included in the strict synonomy of P. (P.)
westphalicum, in contrast to the conclusions of Matsumoto & Muramoto (1981:
55):

P. westphalicum var. australis Venzo (1936: 100, pl. 5 (fig. 4)), also from
the Coniacian of Zululand, resembles the specimen figured by De Grossouvre
(mentioned above) in having a rather prominent umbilical tubercle and a more
quadrate whorl section than typical P. (P.) westphalicum. It is morphologically
transitional to P. (P.) subtricarinatum.

P. guerini Collignon (1965: 64, pl. 442 (fig. 1811)) has typical P. (P.) west-
phalicum ornament, and differs only in possessing a prominent depression separ-
ating the lateral ribs from the ventrolateral tubercles. The authors doubt if this
feature merits specific separation, and include the species in the strict synonomy
of P. (P.) westphalicum.

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

P. latum Matsumoto & Muramoto (1981: 52, pl. 6 (fig. 1), pl. 7 (fig. 1),
text-fig. 1) was considered to be a close ally of P. stefaninii Venzo, 1936, differ-
ing only in a wider umbilicus (U = 63 %) and numerous (twenty-eight) ribs on
the outer whorl. It is also a probable synonym of the present species.

Occurrence

Middle Coniacian of western Europe, Madagascar, Zululand, and the Gulf
Coast of North America.

Peroniceras (Peroniceras) drayidicum Kossmat, 1895
Figs 29-42

_ Ammonites subtricarinatus d’Orbigny: Stoliczka, 1865: 54, pl. 31 (fig. 3)

Schloenbachia (Peroniceras) dravidica Kossmat, 1895: 190, pl. 22 (fig. 3).

Peroniceras dravidicum Kossmat: Solger, 1904: 181, figs 71-72. Lombard, 1930: 294, pl. 30
(fig.1). Venzo, 1936: 99, pl. 10 (fig. 1), pl. 11 (fig. 9). Non Ciry, 1940: 212. Reyment, 1958:
65. Collignon, 1965: 52, pl. 436 (fig. 1802), p. 62, pl. 441 (fig. 1809); 1978: 63, pl. 28
(Gays. 11).

Peroniceras cfr. subtricarinatum d’Orbigny var. dravidicum Kossmat: Desio, 1920: 207, pl. 12
(ie, QD).

Peroniceras cf. dravidicum Kossmat: Spath, 1921: 295, pl. 23 (fig. 1).

Peroniceras sp. aff. dravidicum Kossmat: Collignon, 1967: 51, pl. 30 (fig. 6).

Peroniceras platycostatum van Hoepen, 1955: 367, figs 15-17.

Peroniceras spathi van Hoepen, 1955: 368.

Peroniceras (Peroniceras) multinodatum van Hoepen, 1965: 8, pl. 4; text-fig. 1f.

Peroniceras (Zuluiceras) obliquenodatum van Hoepen, 1965: 14, pl. 8; text-fig. 3f.

Peroniceras (Zuluiceras) bessiae van Hoepen, 1965: 18, pl. 13.

Type specimen

The lectotype is the original of Kossmat (1895, pl. 23 (fig. 3a-d)) by the
subsequent designation of Matsumoto (1965: 213).

Material

SAS A536, Z253, Z753, Z1017, SAM—PCZ5944, PCZ5495, K5494, K4950,
NMB D2145b, D1148 from locality 72, degraded river cliff and northern banks
of the Mzinene River downstream from the junction with the Munywana River,
Zululand, St. Lucia Formation, Coniacian II and III. SAS Z663, Z811, Z1607,
ZN521, Zis2ic, Z1616, 21632, Zi1650, Z1662, Zi1672, Zi674 trom@locality so?
bulldozer scrapings and adjacent hillslopes around the pumping station on the
farm Panplaas, east-south-east of Hluhluwe, Zululand, St. Lucia Formation,
Coniacian II and III. SAS Z1 and Z892 from locality 93, hillslopes on either
side of the boundary fence separating lots H101 and H102, east-south-east of
Hluhluwe, Zululand, St. Lucia Formation, Coniacian II and III. SAS A609
from locality 24, caisson excavations for new Nyalazi River bridge, Zululand,
St. Lucia Formation, inferred to be Coniacian II and III. Numerous fragments
and uncatalogued material from the above localities.

CRETACEOUS FAUNAS FROM SOUTH AFRICA Al

Fig. 29. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z1017 showing details of ontogeny up
to the body chamber. Note especially the unicarinate venter on the inner whorls. x 0,9.

72

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 30. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS PCZ5945 showing details of ontogeny.

horls. A-B X 1, C X 3.

inate inner w

, unicarina

Note especially the smooth

CRETACEOUS FAUNAS FROM SOUTH AFRICA 17/3

Fig. 31. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z1633 partial suture-line
showing effect of ornamentation on details of elements, in this case saddle U2/U3.

Dimensions

D Wb Wh Wb/Wh iO R
D1148 44,0 11,0(25,0) 14,0(31,8) 0,79 19,0(43,2) 19:30
Z1607 46,0 13,0(28,3) 14,0(30,4) 0:97 21,0(45,6) 19:24
LAGT2 46,0 12,0(26,1) i351(28-3) 0,92 ZIAVSO0K, 027-38
L253 6530) —«18,0(@27,7) ZAVO(G2-3) 0,86 29,0(44,6) 20:32
L392 74,0 20,0(27,0) 21,0(28,4) 0,95 37,0(50,0) i
Z811 g5.0. » 24,0(32,0) 24,0(32,0) 1,0 34,0(45,3) 26:29
K5494 fs-0- 720,005-6) 24,0(30,8) 0,83 37,0(47,4) 20:34
NG SZ BOLOr = 2050(25,0) 23-0267) 0,87 40,0(50,0) = 23:27
Z1662 8020) 19.0(23:7) 23,0(28,7) 0,83 39,0(48,8) 20:31
Z663 S00" — 18:0(22,2) 29,0(35,8) 0,62 39,0(48,1) 15:34
TA SDA 82,0 20,0(24,4) 25,0(30,5) 0,80 eA le iy DAS
A536 Oi Om 220(25-3) 27,0(31,0) 0,81 39,0(44,8) 20:36
ZASZ 1c 67.0), 22.0(25,3) DAO CSi0) 0,82 40,0(45,9) 18:24
PCZ5944 9720) 2650(26.8) ZSLU(Z5-8) 1,04 OE O\nin 25250
D1145b 104,0 29,0(27,9) 29,0(27,9) 0 54,0(51,9) 20:26
D1145a LIGOH" 33200855) 34,0(29,3) 0,97 STAO(4I DD) «26:32
A912 1222062 3120054) 32,0262) 0597 66,0(54,1) 34:34

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

D Wb Wh Wb/Wh U R
Z753 124,0 31,0(25,0) 41,0(33,1) 0,75 55-0443) =
Z1616 141,0 38,0(26,9) 45,0(31,9) 0,84 ~—62,0(43,9) 20:35
Z1674 151,0 37,0(24,5) 49,0625) 075. (72,0477) mueoes
PZ5495 150,0 35,0(23,3) 37,0(24,7) 0,95 — 83,0(55,3) 28:35
Z1017 183,0 =e 45,0(24,6) — 104,0(56,8) 37:37
Z1650 203,0 = 55,0(27,1) .— (114.0660) estees

Description

This is the largest known collection of P. (P.) dravidicum, and exhibits all
stages of ontogeny, as well as the extensive intraspecific variation.

Coiling is generally evolute, the umbilicus comprising up to 56 per cent of
- the diameter in large individuals, but it may be as low as 43 per cent in smaller
specimens and what appear to be stratigraphically younger representatives of the
species.

The whorl section is generally rectangular, higher than wide throughout,
but may be quadrate or even slightly wider than high in some individuals.

Details of the early ontogeny can be seen in SAS Z1017 (Fig. 29),
NMB D1148 and SAM-—PCZ5945 (Fig. 30). The early whorls are smooth, with
parallel flanks converging to an acute, unicarinate venter. The smooth stage lasts
up to about 8 mm, whereafter rounded tubercles start appearing at the umbilical
edge and clavate ventrolateral ones in the umbilical suture of the succeeding
whorl. They are connected by weak, inconspicuous bifurcating and single ribs.
This ornament remains basically the same thoughout, except on the later part of
the phragmocone and on the body chamber of the larger specimens, where the
ribs become more club-shaped and predominantly single, and the umbilical
tubercles become indistinct. This latter ornament is very reminiscent of P. (P.)
westphalicum.

The central Keel is strongest throughout ontogeny.

The suture-line is complex, generally with highly incised elements. Details
differ, however, according to the relative position of the sutural elements to or-
nament (Fig. 31). If saddles coincide with tubercles, they tend to be rather
broad, whereas if situated in the interspaces, they are narrow-stemmed as in
typical Peroniceras. The attachment of the U2/U3 saddle to the base of the saddle
L/U2 is prominent in the species.

Discussion

Specimens referred to this species form the bulk of the Peroniceras material
before us. Variants show gradations in some characters, both in contemporary
and successive populations, towards other species. It is easily recognized, how-
ever, by the numerous bifurcating ribs on the inner whorls, compressed
rectangular whorl section, evolute coiling and weak lateral keels.

175

CRETACEOUS FAUNAS FROM SOUTH AFRICA

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ANNALS OF THE SOUTH AFRICAN MUSEUM

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ow
So
eae
N-2
Na
<C
es
vale
oo 2B
Sl (Sy
§ |S
SO
g&°
ZS
ma
me
Sa
Osn
ee
S ae
3S
Qu
Sw
i> GN
Os
See
S 6
a6
=e
oi
Se
=
Ss
eS
oS
“Ss
ans)
a8
obs
= ™
Ha

Fig. 34. Peroniceras (Peroniceras) dravidicum Kossmat, 1895.

Suture-line of SAS Z1119.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 77

aoe,
eee

@ 10 mm
ae a

Fig. 35. Peroniceras (P.) dravidicum Kossmat, 1895. Suture-lines of juvenile specimens.
A. SAS Z1633. B. SAS Z1615. C. Uncatalogued fragment.

A number of species described by Van Hoepen (1955, 1965) are here re-
garded as synonyms of this species. Peroniceras spathi van Hoepen (1955:
368 = Peroniceras cf. dravidicum Stoliczka of Spath, 1921: 295, pl. 23 (fig.
la—d)) (Fig. 32B—C herein) is clearly a synonym of P. (P.) dravidicum. P. platy-
costatum van Hoepen (1955: 367, figs 15-17) (Fig. 33 herein) was said to differ
from P. (P.) dravidicum in having slightly converging flanks and somewhat dif-
ferently disposed umbilical and ventrolateral tubercles. Van Hoepen also consid-
ered his species to differ from the specimen of P. (P.) dravidicum figured by
Venzo (1936, pl. 10 (fig. 1), pl. 11 (fig. 2)) on the basis of the denser ribbing and
higher whorl section. Given the variation shown by the material at the authors
disposal, these differences are obviously within the limits of intraspecific varia-
tion, and P. platycostatum is included in the strict synonymy of P. (P.) dravidi-
cum.

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B C

Fig. 36. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z663, the holotype of
Van Hoepen’s P. (P.) multinodatum. X 1.

P. (P.) multinodatum van Hoepen (1965: 8, pl. 4, text-fig. 1f) (Fig. 36
herein) differs from the commoner forms of P. (P.) dravidicum in having an un-
usually high ratio of ventral to umbilical tubercles (34:15), but has all the other
characteristics of the species.

P. (Zuluiceras) obliquenodatum van Hoepen (1965: 14, pl. 8, text-fig. 3f)
(Fig. 37 herein) is nothing more than a coarsely ornamented variant of P. (P.)
dravidicum, although the narrow umbilical width (45 %) suggests that it is in this
respect transitional to P. (Zuluiceras). Another form transitional to P. (Zulu-
iceras), but which still falls within the limits of P. (P.) dravidicum, is P. (Zulu-
iceras) bessiae van Hoepen (1965: 18, pl. 13) (Fig. 38 herein).

Some of the very large forms of P. (P.) dravidicum here figured (e.g. Figs
39-40, with very evolute whorls and club-like, simple ribs on the outer whorls,
are reminiscent of P. (P.) westphalicum. In typical P. (P.) westphalicum, how-
ever, the ribs are broad and widely spaced. The inner whorls of both species are
remarkably similar, suggesting a close relationship.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 179

Fig. 37. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z753, the holotype of
P. (Zuluiceras) obliquenodatum van Hoepen, 1965. This specimen is transitional to subgenus
P. (Zuluiceras). X 1.

Specimens of P. (P.) dravidicum with more quadrate whorl sections and
stronger ornament are in these respects transitional to P. (P.) subtricarinatum,
e.g. SAS A609 and SAM-PCZ5946 (Fig. 32A). Typically, however, P. (P.) sub-
tricarinatum has a quadrate to depressed whorl section with very strong umbili-
cal bullae. In addition, the venter is generally more rounded and the coiling
more evolute.

P. (P.) dravidicum is the ancestor of P. (Zuluiceras). The transition in-
volved the development of tighter coiling, retention of a single keel to greater
diameters, and more prominent development of rounded ribbing on the outer
whorls (Fig. 41). In fact, one of the paratypes of P. (Z.) zulu, SAS Z14 (Fig.
45), is still very close to P. (P.) dravidicum.

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 38. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z255, the holotype of P. (Zuluiceras)
bessiae van Hoepen, 1965, another specimen transitional to subgenus P. (Zuluiceras). X 1.

In addition to the above changes, there appears to have been a general in-
crease in overall size.

Occurrence

Coniacian of Italy, north, west and central Africa, Madagascar and India,
Coniacian II and III of Zululand.
Subgenus Peroniceras (Zuluiceras) van Hoepen, 1965
(= Zuluites van Hoepen, 1965: 23; Sornayceras Matsumoto, 1965: 226)
Type species

Peroniceras (Zuluiceras) zulu van Hoepen, 1965: 23, by original desig-
nation.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 181

Fig. 39. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z1650, a large form with
club-like simple ribs on outer whorls. x 0,6.

Diagnosis

Progressively more involute than Peroniceras (Peroniceras) with adult
whorls tending to become rounded and smooth. Lateral keels are weak to
absent on outer whorls and siphonal keel may be entire or serrated. Suture
complex, of P. (P.) dravidicum type.

Discussion

Van Hoepen separated P. (Zuluiceras) from P. (Zuluites) on the basis of
the greater involution and loss of ornament in adults of the latter. Stratigra-
phic work shows that there is a progression from early, relatively evolute
Middle Coniacian species with persistent ornament that are close to Peronice-
as (Peroniceras) (e.g. P. (Zuluiceras) bajuvaricum (Redtenbacher, 1873),

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 40. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z1663 a large form with
club-like ribs on outer whorls. x 0,84.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 183

Fig. 41. Peroniceras (Peroniceras) dravidicum Kossmat, 1895. SAS Z1674, specimen with
narrower umbilical width transitional to subgenus P. (Zuluiceras). X 0,75.

P. (Z.) aberlei (Redtenbacher, 1873), P. (Z.) isamberti (Fallot, 1885)) to Upper
Coniacian species that are much more involute and lose ornament when adult.
There is no morphological break between the members of the series, and the
separation proposed by Van Hoepen is considered unnecessary.

Sornayceras Matsumoto, 1965, is a further, junior synonym of P.
(Zuluiceras). The type species, Sornayceras proteus Matsumoto, 1965, is
evolute and coarsely ribbed, the ribs persisting, suggesting that it is an
early member of the lineage. Matsumoto (1965: 227) also mentioned that
P. undulatocarinatum van Hoepen, 1955, here regarded as a good example
of P. (Zuluiceras), was a Sornayceras, which supports the present conclusions.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B C

Fig. 42. Peroniceras (Peroniceras) dravidicum Kossmat, 1895.
WXo Syexs) Z4NSVA 10AC, SYANS) ANSE, JENIN <I,

P. (Zuluiceras) most closely resembles P. (Peroniceras) amongst the Peroni-
ceratinae. The more involute coiling, compression, lateral keels subsidiary to the
siphonal, and loss or weakening of ornament at maturity distinguishes them.

Occurrence

Middle Coniacian of Touraine, Aquitaine, Dieulefit (Dr6me), the Cor-
biéres (Aude), Var, and Provence in France, Germany, Czechoslovakia, Aus-
tria, Italy, Romania, Armenia, north Africa, Zululand, Madagascar, and Japan.
Upper Coniacian of Zululand and, possibly, Japan.

Peroniceras (Zuluiceras) zulu van Hoepen, 1965
Figs 43-52

Peroniceras sp. a Crick, 1907: 226, text-fig. to p. 226.

Peroniceras sp. 8 Crick, 1907: 227.

Peroniceras (Zuluiceras) zulu van Hoepen, 1965: 9, pl. 5, text-figs 1 g—i, 2b.
Peroniceras (Zuluiceras) charliei van Hoepen, 1965: 11, pl. 6, text-fig. 1).
Peroniceras (Zuluiceras) regularis van Hoepen, 1965: 14, pl. 9, text-fig. 3d-e.
Peroniceras (Zuluiceras) multilineatum van Hoepen, 1965: 23, pl. 18, text-fig. Sd-e.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 185

Fig. 43. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. SAS Z746, the holotype. x 0,46.

Type specimen

The holotype, by original designation, is SAS Z746 (Figs 43-44), from
locality 91, degraded river cliffs and artificial cut west of the boundary of the St.
Lucia Game Reserve and Lot H103, east-south-east of Hluhluwe, Zululand.
This area is strewn with rubble derived from the St. Lucia Formation, Coniacian
IV and V, and the precise age of the holotype is thus unknown.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

ae

Fig. 44. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. SAS Z746, the holotype. x 0,62.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 187

Fig. 45. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. SAS Z14, one of
Van Hoepen’s paratypes, still close to P. (P.) dravidicum. X 0,8.

The type specimens of Van Hoepen’s P. (Z.) charliei, P. (Z.) regularis, and
P. (Z.) multilineatum, SAS Z735, Z736, and Z758 respectively, are all from the
same locality as the holotype of P. (Z.) zulu. Their precise age is thus uncertain.

Dimensions

D Wb Wh Wb/Wh U R
ASS H10:0- 4.39,0G5,5) 38 ,0(34,5) 1,03 53,0(48,2) —
2136 141,0 55,0(39,0) 44 .0(31,2) 122) 68,0(48,2) 22:23
Z758 15850 52,0(2,9) 47,0(29,7) 110 74,0(46,8) 23:23

Z746 296,0 87,0(29,4) 92,0(31,1) 0,94 146,0(49,3) 25:25

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Coiling is polygyral, evolute with an umbilical diameter of 47 to 50 per cent
of the total diameter. Successive whorls overlap to the ventrolateral row of
tubercles.

The whorl section varies from slightly compressed to slightly depressed
with a rounded to overhanging umbilical wall, slightly converging flanks
and a broadly fastigiate venter with a distinct median keel (which may be
serrated) as well as weaker lateral keels. The latter may disappear completely
in adults.

Se oo

Fig. 46. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. Suture line and whorl section of
holotype. x 1. (After Van Hoepen 1965, figs 1g, 2b.)

Ornament consists of rounded to radially elongated umbilical tubercles con-
nected by ribs of varying strength to clavate ventrolateral tubercles. On the
inner whorls the ribs may bifurcate or intercalate occasionally. On the outer
whorls they are generally single, stronger and more distantly spaced than on nu-
clei. The ventral area between the row of ventral tubercles and the median keel
is often ornamented by faint spiral striae. Where preservation is good, spiral or-
nament may also be observed on the flanks.

The suture is highly incised and of the P. (P.) dravidicum type (Fig. 46).
The maximum body chamber length observed is half a whorl.

Discussion

This species appears to include the two specimens described by Crick
(1907: 226-228) as Peroniceras spp. a and 8, here refigured as Figures 51—52.
Van Hoepen’s choice of the holotype is unfortunate as this is a large, septate

CRETACEOUS FAUNAS FROM SOUTH AFRICA 189

Fig. 47. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. SAS Z735, the holotype of
Van Hoepen’s P. (Z.) charliei. X 1.

specimen lacking the inner whorls (Figs 43-44) and thus difficult to compare
with the other material. However, as interpreted here, P. (Z.) zulu is an evolute
form with an umbilical diameter between 45 and 50 per cent of the total di-
ameter with whorls more or less as wide as high.

One of Van Hoepen’s syntypes, SAS Z14 (Fig. 45) differs from the typical
form of the species in having a more rectangular whorl section and more distinct
lateral keels, and probably being in those respects transitional to Peroniceras
(Peroniceras) of the dravidicum group.

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

i <>
Pi ie & : : y
mag
~~ e > ee ‘s Lae
aa vat : sere haa
‘ :

i

Fig. 48. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. SAS Z736, the holotype of
Van Hoepen’s P. (Z.) regularis. x 0,92.

P. (Z.) zulu includes P. (Z.) charliei (Fig. 47), P. (Z.) regularis (Fig. 48),
and P. (Z.) multilineatum (Figs 49-50), all of which are from the same locality as
the holotype and have comparable proportions and ornament: the differences
cited by Van Hoepen are trivial and within the range of intraspecific variation
shown by other peroniceratids. .

P. (Z.) zulu differs from P. (Z.) henniei in having a wider umbilicus and
more angular whorl section, but it is clear that the latter is derived from P. (Z.)
zulu through progressive increase in involution and rounding of the whorl
section. In addition, there is a reduction of the lateral keels and progressive
serration of the median keel.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 191

Fig. 48. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. SAS Z736, the holotype of
Van Hoepen’s P. (Z.) regularis. x 0,92.

In P. (P.) dravidicum the whorl section is more rectangular and the coiling
more evolute.

Occurrence

The horizon of the type material is difficult to establish as the specimens
were collected loose. Material collected in situ is restricted to Coniacian IV,
which is presumed to be the level of the type specimens. The species is restricted
to Zululand.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Wy

) zulu van Hoepen, 1965. SAS Z758, the holotype of

Van Hoepen’s P. (Z.) multilineatum. X 0,8.

iceras

(Zulu

. 50. Peroniceras

Fig

193

CRETACEOUS FAUNAS FROM SOUTH AFRICA

1965.
5

s (1907) Peroniceras sp. a. X 0,8

:

an Hoepen

(Zuluiceras) zulu v

g. 51. Peroniceras

o

i

The original of Crick

15

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 52. Peroniceras (Zuluiceras) zulu van Hoepen, 1965. The original of Crick’s (1907) Peroniceras sp. 8. X 1.

|

CRETACEOUS FAUNAS FROM SOUTH AFRICA 195

Peroniceras (Zuluiceras) rarum van Hoepen, 1965
Fig. 53

Peroniceras (Zuluiceras) rarum van Hoepen, 1965: 16, pl. 12, text-fig. 4(c).

Type specimen

Holotype, by monotypy, is SAS Z742 from locality 91, degraded river cliffs
and artificial cut west of the boundary of St. Lucia Game Reserve and Lot
H103, east-south-east of Hluhluwe, Zululand, St. Lucia Formation, Coniacian
IV or V, probably the former.

Material

BMNH C83865, from locality 83, foreshore exposures around Mason’s
Camp, western shores of False Bay, east-north-east of Hluhluwe, Zululand, St.
Lucia Formation, Coniacian IV.

Dimensions

D Wb Wh Wb/Wh U R
L142 KOOO — 3250(32,0) 36,0(36,0) 0,89 38,0(38,0) —
Description

The holotype consists of part of a septate, crushed whorl. Coiling is rather
involute with an umbilical diameter of 38 per cent. The whorl section through-
out is compressed with little inflated, parallel flanks and a gently rounded, fasti-
giate venter bearing a distinct crenulate central keel and poorly defined lateral
keels.

Lateral ornament consists of weak umbilical bullae, narrow, rectiradiate,
occasionally bifurcating ribs, and rounded to clavate ventrolateral tubercles.
Faint traces of spiral ornament are visible in the area between the ventrolateral
tubercles and the faint lateral keels.

The suture is only partially exposed, but appears to be of the general
P. (P.) dravidicum type.

Discussion

It is regrettable that Van Hoepen erected a new species on such incomplete
material, which renders interpretation difficuit. As far as the whorl section is
concerned, the species is allied to the group of P. (P.) dravidicum, but the cre-
nulate keel and more involute coiling places it closer to the group of P. (Zulu-
iceras). It differs from P. (Z.) zulu by the tighter coiling. The more tightly coiled
group of P. (Z.) henniei differs from P. (Z.) rarum in having a more inflated
whorl section at comparable diameters.

Lacking more material for additional observations, the species is here main-
tained with doubt.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

Occurrence

The type locality includes rubble derived from Kennedy & Klinger’s (1975)
divisions Coniacian IV and V, so that the precise age is unknown. The second
specimen is firmly dated as Coniacian IV. The species is known only from Zulu-

land.

Fig. 53. Peroniceras (Zuluiceras) rarum van Hoepen, 1965.
SAS 2742, the holotype. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 197

Peroniceras (Zuluiceras) henniei van Hoepen, 1965

Figs 54-64

Peroniceras (Zuluiceras) henniei van Hoepen, 1965: 12, pl. 7, text-fig. 1k.

Peroniceras (Zuluiceras) alfredmeintjesi van Hoepen, 1965: 15, pl. 10, text-fig. 3b—c.
Peroniceras (Zuluiceras) simplicicostatum van Hoepen, 1965: 16, pl. 11, text-fig. 3a, g.
Peroniceras (Zuluiceras) undulatocostatum van Hoepen, 1965: 19, pl. 14, text-fig. 4a—b.
? Peroniceras (Zuluiceras) similis van Hoepen, 1965: 19, pl. 15, text-figs 2a, 4d.
Peroniceras (Zuluiceras) proximum van Hoepen, 1965: 20, pl. 16 text-fig. Sa—b.
Peroniceras (Zuluiceras) ultimum van Hoepen, 1965: 22, pl. 17, text-fig. 5c.

Type specimens

The holotype, by original designation, is SAS Z638 from locality 91, de-
graded river cliffs and artificial cut west of the boundary of St. Lucia Game Re-
serve and Lot H103, east-south-east of Hluhluwe, Zululand, St. Lucia
Formation, Coniacian IV or V. The type specimens of P. (Z.) alfredmeintjesi,
SAS 2738, P. (Z.) simplicicostatum, SAS Z740, P. (Z.) proximum, SAS Z747,
P. (Z.) ultimum, SAS Z752, P. (Z.) undulatocostatum, SAS Z744, P. (Z.) sim-
ilis, SAS Z745 and SAS 104, are all from the same locality as the holotype, and
are equally imprecisely dated.

Dimensions

D Wb Wh Wb/Wh U R
Z638 115,0 41,0(35,6) 42 ,0(35,6) 0,98 46,0(40,0) 12x12:12x2
Lils2 130,0 46,0(35,4) 44 .0(33,8) 1,04 56,0(43,1) 21:34
Z738 150,0 54,0(36,0) 53;,0G5;3) 1,02 64,0(42,7) 26. Sil
Z745 178,0 61,0(34,3) 60,0(33,7) 1,02 79,0(44,4) --
Z7A47 262,0 81,0(30,9) — 87,0(33,2) —-0,93 115,0(43,9) =
Z740 283,0 88,0(31,1) 97,0(34,3) 0,91 115,0(40,6) 23525
Z744 B4510 139,0(40.3) 12510663) 1,11 140,0(40,6) =
Description

The species has an umbilical diameter varying between 40 and 45 per cent
of the total diameter. The whorls are slowly expanding, overlapping up to or be-
yond the ventrolateral tubercle. The whorl changes through ontogeny; it is in-
itially quadrate with flattened flanks, but in later stages the umbilical wall
becomes rounded and the flanks inflated. The venter is broadly rounded with a
serrated central keel and weak lateral keels.

The adult whorl section varies among specimens as discussed below.

Ornament on the inner whorls is of P. (P.) dravidicum type, with umbilical
bullae giving rise to one or two ribs that link ventrolateral clavi. On the outer
whorls ribbing becomes bolder, and bifurcations rather rare. The ventral area
between the ventrolateral tubercles and the central keel is ornamented by fine,
spiral, interrupted striae. The undulations on the keel are far more numerous
than the ventrolateral clavi.

The suture-line is of the P. (P.) dravidicum type (Fig. 61).

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 54. Peroniceras (Zuluiceras) henniei van Hoepen, 1965.
SAS Z638, the holotype. x 1.

Discussion

The authors’ interpretation of the species is somewhat different from that of
Van Hoepen (1965) and includes all forms with an umbilical diameter of be-
tween 40 and 45 per cent, more or less equal whorl breadth and height, serrated
central keel, and very weak or no lateral keels. This species continues the mor-
phological trend set by P. (Z.) zulu towards tighter coiling and loss of ornament
on the outer whorls.

The description given above covers a number of species described by Van
Hoepen, all collected from the same locality. These species (listed in the syn-
onomy) take the name P. (Z.) henniei.

———$—$—$<———

CRETACEOUS FAUNAS FROM SOUTH AFRICA 199

Fig. 55. Peroniceras (Zuluiceras) henniei van Hoepen. This specimen and that in Figure 57 show
the features of the species to the best advantage. x 0,8.

The holotype of P. (Z.) henniei (Fig. 54) consists of parts of three consec-
utive whorls, of which the last quarter of the outermost one is body chamber.
The impression of an earlier whorl shows that the venter is already unicarinate
and serrated at a diameter of 12,5 mm. As far as the angular whorl section is
concerned, this specimen is perhaps closest to P. (Z.) zulu.

The holotype of P. (Z.) alfredmeintjesi (Figs 55—56A) is the best preserved
of all the specimens and shows the main features of the species. The inner
whorls are typically quadrate, whereas the outer whorls are more rounded. A

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 56. A. Peroniceras (Zuluiceras) henniei van Hoepen, 1965. SAS Z738, the holotype of
Van Hoepen’s P. (Z.) alfredmeintjesi. X 0,9. B. Peroniceras (Zuluiceras) zulu van Hoepen. :
x 0,88.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 201

Fig. 57. Peroniceras (Zuluiceras) henniei van Hoepen, 1965. SAS A374. x 1.

nodate Baculites, presumably of the group B. capensis Woods, is present in the
body chamber of this specimen.

The holotype of P. (Z.) simplicicostatum (Figs 58-59) is an enormous indi-
vidual, still septate at a diameter of 306 mm. Unfortunately the inner whorls up
to a diameter of 90 mm are not preserved. The whorls are much more massive
and increase in size is more rapid than in the preceding specimens. Also, whorl
height exceeds whorl breadth considerably. In this respect the specimen is closer
to P. (Z.) undulatocarinatum described below, but has a slightly greater umbili-
cal width than the latter species. The specimen figured by De Grossouvre (1894,

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 58. Peroniceras (Zuluiceras) henniei van Hoepen, 1965. SAS Z740, the holotype of
P. (Z.) simplicicostatum van Hoepen. X 0,47.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 203

Fig. 59. Peroniceras (Zuluiceras) henniei van Hoepen,
1965. SAS Z740, the holotype of P. (Z.) simplicicosta-
tum van Hoepen, 1965. x 0,53.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 60. Peroniceras (Zuluiceras) henniei van Hoepen, 1965. SAS Z744, inner whorls
of the holotype of P. (Z.) undulatocostatum van Hoepen, 1965. X 1.

Fig. 61. Peroniceras (Zuluiceras) henniei van Hoepen, 1965. Suture line and whorl section of
SAS Z744. x 1. (After Van Hoepen 1965, fig. 4a—b.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 205

Fig. 62. Peroniceras (Zuluiceras) henniei van Hoepen, 1965. SAS Z747, the holotype of
P. (Z.) proximum van Hoepen, 1965. x 0,6.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 63. Peroniceras (Zuluiceras) henniei van Hoepen, 1965. SAS Z752, the holotype of
P. (Z.) ultimum van Hoepen, 1965. x 1.

pl. 9 (fig. 1)) as an adult of Gauthiericeras bajuvaricum is a comparable but
poorly preserved European analogue.

The holotype of P. (Z.) undulatocostatum is another gigantic specimen with
a maximum measured diameter of 533 mm. Unfortunately only part of the inner
whorls survive, but Van Hoepen’s figure of the holotype is adequate also to
show it as a transitional form between P. (Z.) henniei and P. (Z.) undulatocari-
natum. Part of the inner whorl is preserved with some of the shell still intact.
This shows the finely serrated central keel and finely striate radial ornament in
the ventral area between the keel and the ventrolateral tubercles (Fig. 60).

207

CRETACEOUS FAUNAS FROM SOUTH AFRICA

imen

a loosely coiled spec

, 1965. SAS 104,
SG0L/3-

) zulu

Move (Z

transitiona

Fig. 64. Peroniceras (Zuluiceras) henniei van Hoepen

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

The holotype of P. (Z.) similis is not available for study, but from the de-
scriptions and figures it seems to be a more evolute form connecting P. (Z.)
henniei with P. (Z.) zulu. The holotype of P. (Z.) proximum (Fig. 62) is also a
large specimen consisting of part of two successive whorls of which only a small
part is body chamber. Again the looser coiling suggests that it connects morpho-
logically with P. (Z.) zulu.

The holotype of P. (Z.) ultimum (Fig. 63) is a septate disc 130 mm in di-
ameter. Apart from details of preservation, the outer whorls of this specimen
are indistinguishable from the inner whorls of the holotype of P. (Z.) undulato-
costatum, and is thus a further clear synonym of P. (Z.) henniei.

The species is here interpreted more widely than by Van Hoepen, but the
authors doubt if the differences between the various ‘species’ given by that
author are of specific significance. All the species are based on single specimens
that occur at essentially the same stratigraphic level, which further causes one to
doubt their validity. In consequence they are all here included in P. (Z.)
hennieéi.

The next species to be discussed, P. (Z.) undulatocarinatum, continues the
trend towards closer coiling, but here a progressive loss of ornament on the
outer whorls spans the boundary between Van Hoepen’s ‘subgenera’ P. (Zulu-
iceras) and P. (Zuluites).

Occurrence

The type locality is strewn with rubble from Coniacian IV and V; the
species is probably from the former. It may possibly also occur in France.

Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955
Figs 65-75

Peroniceras undulatocarinatum van Hoepen, 1955: 369, figs 18-21.

Peroniceras (Zuluites) georgemeintjesi van Hoepen, 1965: 25, pl. 20, text-fig. 6c—e.
Peroniceras (Zuluites) lettiae van Hoepen, 1965: 27, pl. 21, text-fig. 7a—d.

? Peroniceras (Zuluites) dubium van Hoepen, 1965: 28, pl. 22, text-fig. 7b—c.

Type specimens

The holotype, by original designation, is SAS Z15 (Fig. 73) from locality
91, degraded river cliffs and artificial cut west of the boundary of St. Lucia
Game Reserve and Lot H103, east-south-east of Hluhluwe, Zululand. This
locality is strewn with rubble from the St. Lucia Formation, Coniacian IV and
V, and the specimen cannot be precisely dated, although it is probably from
Coniacian IV. The holotype of P. (Zuluites) georgemeintjesi is SAS Z733, the
holotype of P. (Z.) lettiae is SAS Z185; the holotype of P. (Z.) dubium is SAS
Z750. Other specimens are SAS Z19, Z631, Z742, and Z868, all from the same
locality. Some specimens are associated with nodose Baculites, suggesting that
they are from Coniacian IV.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

209

Fig. 65. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955. SAS Z19, inner whorls
of paratype of Van Hoepen (1955). x 0,95.

Dimensions
D

Z19 142,0
Z750 148.0
PENS 159,0
A535 ‘at 154,0

at 218,0
Z185 174,0

Wb

49,0(34,5)
42,0(28,3)
54,0(34,0)
48,0(31,2)
63,0(28,9)
62,0(35,6)

Wh
55,0(38,7)
54,0(36,5)
59,0(37,1)
60,0(39,0)
87,0(39,9)
68,0(39,1)

Wb/Wh

0,89
0,78
0:95
0,80
0,72
0,91

U
55,0(38,7)
58,0(39,2)
64,0(40,2)
55,0(35,7)
84,0(38,5)
72,0(41,4)

R
132-30
i x<A13s x2
14x2:14x2

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 66. A-B. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955 SAS Z19,
inner whorls of paratype of Van Hoepen (1955). x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 67. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1965. SAS Z19,
outer whorl of paratype of Van Hoepen (1955). x 0,75.

211

DN ANNALS OF THE SOUTH AFRICAN MUSEUM

™

, c
N

Fig. 68. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955. Suture-line and whorl
section of the holotype, SAS Z15. x 1. (After Van Hoepen 1955, figs 19-20.)

L
E Us

é

Cc
Ww

—_—_

Fig. 69. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955. Suture-line of paratype
SAS Z19. x 1. (After Van Hoepen 1955, fig. 22.)

Coiling is involute for Peroniceras, with an umbilical diameter of between
35 and 40 per cent of the total diameter. Successive whorls overlap up to and be-
yond the ventrolateral tubercles.

The whorl section is distinctly compressed, with a vertical to overhanging
rounded umbilical wall, little-inflated, rounded flanks converging to a broadly
rounded-fastigiate venter (Fig. 68).

Ornament on the inner whorls consists of prominent bullate to auricular
umbilical tubercles, weak prorsiradiate ribs and rounded to clavate ventrolateral
tubercles. The latter are situated near the ventral third of the flanks. In the

CRETACEOUS FAUNAS FROM SOUTH AFRICA PENS

Fig. 70. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955. SAS Z733,
the holotype of Van Hoepen’s P. (Zuluites) georgemeintjesi. x 0,66.

214 ANNALS OF THE SOUTH.AFRICAN MUSEUM

oe

Fig. 71. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955. SAS Z185,
the holotype of P. (Zuluites) lettiae van Hoepen, 1965. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 2\9

fastigiate area between the ventrolateral tubercles and the central keel, numer-
ous spiral striae occur. In some specimens, e.g. SAS Z15 (Fig. 73), these striae
also cover the ventrolateral tubercles. The central keel undulates, but the undu-
lations do not coincide with ventrolateral clavi. The central keel is bounded on
either side by a shallow depression producing slight spiral ridges, but these can
hardly be termed lateral keels.

On the outer whorls of the specimens available (nearly all of them are still
septate), all ornament weakens and the whorls become nearly smooth.

Discussion

P. (Zuluiceras) undulatocarinatum continues the trend towards increased
involution and compression, forming a link between P. (Z.) henniei and P. (Z.)
modestum. In addition, loss of ornament on the outer whorls is obvious. This is
comparable to that found in the transition from Texanites to Submortoniceras
(Klinger & Kennedy 1980: 172).

Differences between undulatocarinatum, henniei and modestum are grada-
tional in some respects. There is no clear-cut boundary between them in respect
of umbilical diameter: P. (Z.) henniei has an umbilical width of 40 to 45 per
cent, P. (Z.) undulatocarinatum between 35 and 40 per cent, whereas P. (Z.)
modestum has an umbilical width below 30 per cent. These reductions in umbili-
cal width are, however, accompanied by changes in whorl section and loss of or-
nament.

P. (Zuluiceras) wundulatocarinatum, as here interpreted, includes
P. (Zuluites) georgemeintjesi van Hoepen and P. (Z.) lettiae van Hoepen. Differ-
ences between these latter ‘species’ and P. (Zuluiceras) undulatocarinatum are
minor and restricted to details of the inner whorls.

me Full

Fig. 72. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955. Suture-line and whorl
section of SAS Z185, the holotype of P. (Zuluites) lettiae van Hoepen, 1965. x 1. (After Van
Hoepen 1965, figs 7d, a.)

ANNALS OF THE SOUTH AFRICAN MUSEUM

216

tum van Hoepen, 1955.

iceras) undulatocarina

iceras (Zulu
SAS Z15

Peroni

ISH

Fig

x OWA.

the holotype

b)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 217

B

Fig. 74. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955. SAS Z733,
the inner whorls of P. (Zuluites) georgemeintjesi van Hoepen, 1965. X 1.

P. (Zuluites) dubium (Fig. 75) is more compressed than the holotype of
P. (Zuluiceras) undulatocarinatum and has finer ornament. It bears great simi-
larity to the inner whorls of the holotype of P. (Zuluites) georgemeintjesi
(Fig. 74), The outer whorls of the latter are, however, virtually identical with
those of P. (Zuluiceras) undulatocarinatum. The holotype of P. (Zuluites) lettiae
(Figs 71-72) has stouter, more quadrate inner whorls than the holotype of
P. (Zuluiceras) undulatocarinatum and is in these respects transitional to P. (Z.)
henniei and P. (Z.) zulu.

In summary, inner whorls are rather variable as far as dimensions and orna-
ment are concerned; this is analogous to the situation in the similarly com-
pressed Submortoniceras condamyi Collignon, 1948, and S. woodsi (Spath,
1921), from Zululand and Pondoland (Klinger & Kennedy 1980).

Occurrence

The type locality is strewn with rubble from divisions Coniacian IV and V,
the species probably coming from the former level.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 75. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1955.
SAS Z750, the holotype of P. (Zuluites) dubium van Hoepen, 1965. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 219

Peroniceras (Zuluiceras) modestum van Hoepen, 1965
Figs 76-85

Peroniceras (Zuluites) modestum van Hoepen, 1965: 24, pl. 19, text-fig. 6a—b.

Type specimen

The holotype, by original designation, is SAS Z739 (Fig. 76, 77A) from
locality 91, degraded river cliffs and artificial cut west of the boundary of St.
Lucia Game Reserve and Lot H103, east-south-east of Hluhluwe, Zululand, St.
Lucia Formation, probably Coniacian IV.

Material

SAS 221, Z104, Z379, Z631, and Z868 from the same locality and inferred
horizon as the holotype, SAS 106 from locality 89, St. Lucia Formation, Coni-
acian IV. BMNH C83863 and C83864 from the same horizon at locality 82.

Dimensions

D Wb Wh Wb/Wh U R
Z868 145,0 49,0(33,8) 59,9(40,7) 0,83 ADNVG3.8) 2835
SAS106 220-5 10(33-6) 67,0(44,1) 0,76 42,0(27,6) —
ZS 6050 57,0(5.6) IZ 04:0) O79 46,0(28,7) —
Jas, t6520* 56,0G3,9) 74,0(44,8) 0,76 50206053) —
SAS104 229,0 64,0(27,9) 102,0(44,5) 0,63 59,0(25,8) —
Description

Coiling is very involute, the umbilical diameter varying between 25 and 33
per cent, and up to 50 per cent of the previous whorl being covered.

The whorl section is compressed ovoid throughout ontogeny, with a vertical
to distinctly overhanging umbilical wall, slightly inflated flanks converging to a
broadly rounded venter, with no distinct demarcation between the latter and the
flanks. The venter bears one narrow, serrated keel. Slight depressions occur on
either side of the keel, especially on the inner whorls, but distinct lateral keels
are not developed. On the outer whorls of the phragmocone these depressions
virtually disappear and the venter is unicarinate without sulci.

Ornament on the inner whorls is as in P. (Z.) undulatocarinatum, but rap-
idly disappears on the outer whorls, leaving the shell completely smooth but for
slight undulations representing the umbilical tubercles. Spiral striae occur on the
ventral regions where the shell is well preserved.

All the specimens are still septate, indicating that the species grew to a very
large size. The suture is of the P. (P.) dravidicum type.

ANNALS OF THE SOUTH AFRICAN MUSEUM

220

Fig. 76. Peroniceras (Zuluiceras) modestum van Hoepen, 1965. SAS Z739, the holotype. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA ess |

Fig. 77. A. Peroniceras (Zuluiceras) modestum van Hoepen, 1965. SAS Z739.
B. Peroniceras (Zuluiceras) undulatocarinatum van Hoepen, 1965. SAS Z750, the
holotype of P. (Zuluites) dubium van Hoepen, 1965. Both x 0,92.

pas ANNALS OF THE SOUTH AFRICAN MUSEUM

E L

Fig. 78. Peroniceras (Zuluiceras) modestum van Hoepen, 1965.
Suture-line of the holotype. x 1.
(After Van Hoepen 1965, fig. 6a.)

Discussion

This species is easily distinguished from the preceding Peroniceras (Zulu-
— iceras) species by the high degree of involution and virtual lack of ornament on
the later whorls of the phragmocone.

This appears to be the last species of the P. (P.) dravidicum stock, a line
that evolved rapidly through the Middle and lower Upper Coniacian in Zulu-
land, becoming progressively larger and showing increasing involution and whorl
compression, losing lateral ornament and the pair of lateral keels. Although the
end member is very different from the ancestral P. (P.) dravidicum, the suture-
line remained remarkably similar throughout, especially the U2/Us lateral sad-
dle, which remained attached obliquely by a thin stem to the root of the lateral
L/U2 saddle.

Occurrence
Coniacian IV of Zululand.

Peroniceras (Zuluiceras) matsumotoi sp. nov.
Figs 86-88

Type specimen

The holotype is NMB D1140 from locality 73, degraded river cliffs on the
north bank of the Mzinene River downstream of the junction with the Muny-
wana River, Zululand, St. Lucia Formation, probably Coniacian IV.

Etymology

The species is named for Emeritus Professor Tatsuro Matsumoto of Kyushu
University.

Dimensions
D Wb Wh Wb/Wh U R
D1140 168,0 40,0(23,8) 52,0(30,9) Ona 76,0(45,2) —

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 79. Peroniceras (Zuluiceras) modestum van Hoepen, 1965. SAS Z743.
Note the similarity to P. (Z.) matsumotoi sp. nov. (Figs 86-88) apart from the
more inflated whorl section and narrower umbilical width. x 0,68.

223

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 80. Peroniceras (Zuluiceras) modestum van Hoepen, 1965. SAS Z743. Note the serrated,
unicarinate venter already present on the inner whorls. x 0,71.

225

CRETACEOUS FAUNAS FROM SOUTH AFRICA

SAS 104. x 0,67.

1965.

>)

Fig. 81. Peroniceras (Zuluiceras) modestum van Hoepen

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

pt OBOE

Fig. 82. Peroniceras (Zuluiceras) modestum van Hoepen, 1965.
A. SAS Z868. x 1 B. SAS 104. x 0,69.

CRETACEOUS FAUNAS FROM SOUTH AFRICA D2)

Fig. 83. Peroniceras (Zuluiceras) modestum van Hoepen, 1965. SAS Z868. xX 1.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 84. Peroniceras (Zuluiceras) modestum van Hoepen, 1965. SAS 106. X 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 229

Fig. 85. Peroniceras (Zuluiceras) modestum van Hoepen, 1965. SAS 106. x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

230

Holotype NMB D1140. x 1.

i sp. nov.

Fig. 86. Peroniceras (Zuluiceras) matsumoto

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘
,

seein sca a

pital

2h bhai

ate

Fig. 87. Peroniceras (Zuluiceras) matsumotoi sp. nov.
Holotype NMB D1140. x 1.

31

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

0 10

ee
mm

Fig. 88. Peroniceras (Zuluiceras) matsumotoi sp. nov.
Suture-line of holotype NMB D1140.

SSS oesssoqqnssss

Fig. 89. Ishikariceras binodosum Matsumoto, 1965. Whorl section and partical suture. Xx 1.
(After Matsumoto 1965, text-figs 17-18.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 233

Description

Coiling is very evolute, with only 25 to 33 per cent of the preceding whorl
covered, the degree of overlap increasing through development. The umbilical
diameter is 45 per cent of the total diameter.

The whorl section is compressed throughout, with subparallel, slightly in-
flated flanks and a broadly rounded unicarinate venter. The venter is unicarinate
at the smallest diameter observed but in later growth it is bound by shallow de-
pressions, although no distinct lateral keels develop. The keel is serrated on the
inner whorls but entire on the internal mould of the outer whorls.

The ornament of inner whorls consists of umbilical bullae and rounded ven-
trolateral tubercles. These are linked by radial ribs, most of which are single,
with only occasional bifurcations at or near the umbilical tubercle. Ornament
weakens progressively in later growth and, at the greatest diameter preserved,
the flanks bear only feeble undulations. The suture is incompletely exposed
(Fig. 88) but is of the P. (P.) dravidicum type. The body chamber is unknown.

Discussion

P. (Z.) matsumotoi differs from all the other Zululand species in being both
compressed and very evolute.

The precise relationship of this species to other P. (Zuluiceras) species is
not clear, as transitional or superficially similar forms do not occur.

The most similar form occurs in Madagascar: ‘Schloenbachia’ schneeblii
Boule, Lemoine & Thévenin (1907: 36, pl. 5 (figs 2—2a), text-fig. 20). It resem-
bles P. (Z.) matsumotoi in having coarse ornament on the inner whorls but vir-
tually smooth, unicarinate outer phragmocone whorls. At comparable
diameters, however, the suture-lines are totally different. That of ‘Schloen-
bachia’ schneeblii is very simple, with broad parallel saddles and lobes, markedly
different from the highly incised, narrow saddles and lobes of P. (Z.) matsumo-
tot. Moreover, the siphonal keel of ‘Schloenbachia’ schneeblii is both broader
and more prominent than that of P. (Z.) matsumotoi.

Another comparable form is_ Ishikariceras binodosum Matsumoto
(1965: 236, pl. 43 (fig. 1), text-figs 17-18) from the Coniacian of Hokkaido,
Japan. The inner whorls have a more quadrate section but the Japanese species
show the same loss of ornament on the outer whorls. The suture of the Japanese
species is strikingly different from that of P. (Z.) matsumotoi and is simple, with
little-incised, broad elements (Fig. 89). This is a situation comparable to that be-
tween the extremes of Peroniceras s.s. Here, however, no intermediate forms
linking Ishikariceras to Peroniceras are known. Although it seems probable that
such forms will turn up, retention of /shikariceras as a separate genus seems wise
at present.

Other comparable forms are two specimens described by Matsumoto
(1965: 233) as Sornayceras sp. cf. propoetidum from the ‘Upper’ Coniacian of
Hokkaido, Japan. These specimens have suture-lines and whorl sections like

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

those of the present species, but are more narrowly umbilicate (U = only 30 per
cent of the total diameter as against 45 per cent in P. (Z.) matsumotoi)

Occurrence
Coniacian IV of Zululand only.

Peroniceras (Zuluiceras) cf. aberlei (Redtenbacher, 1873)
Figs 90-91, 92A

Compare

Ammonites aberlei Redtenbacher, 1873: 111, pl. 25 (fig. 4a—e).
De Grossouvre, 1894: 89.

Type specimen

Redtenbacher failed to designate a holotype for this species. A lectotype
designation will be undertaken in a pending version of some of the Gosau faunas
(Kennedy & Summesberger in preparation).

Material

NMB D1147 from locality 72, degraded river cliff and stream sections on
the north side of the Mzinene River downstream of the junction of the Muny-
wana River, Zululand, St. Lucia Formation, Coniacian II or III.

Description

A single specimen consisting of parts of two successive septate whorls is re-
ferred to Redtenbacher’s species. The section of both whorls is distinctly com-
pressed, with a fastigiate venter bearing a single keel. The keel appears entire
and is bounded on either side by distinct grooves, creating the appearance of
weak lateral keels. Ornament on the inner whorls consists of fine, frequently bi-

Fig. 90. Peroniceras (Zuluiceras)
cf. aberlei (Redtenbacher, 1873).
NMB D1147. x 1.

i)
oe)
Nn

CRETACEOUS FAUNAS FROM SOUTH AFRICA

@) 10mm

a ee |
Fig. 91. Peroniceras (Zuluiceras) cf. aberlei (Redtenbacher, 1873). Suture-line of NMB D1147.

furcating, prorsiradiate ribs. On the outer whorl, ornament is coarser and the
umbilical tubercles are very prominently rounded to slightly auricular.
The suture is highly incised and of the P. (P.) dravidicum type (Fig. 91).

Discussion

Redtenbacher based this species on a series of specimens; a cast of one of
these is here illustrated as Figure 92A. A full discussion of the species is to be
given in a forthcoming revision of the Gosau Fauna (Kennedy & Summesberger
in preparation). P. (Z.) aberlei can be distinguished from the similarly com-
pressed P. (Z.) propoetidum (Redtenbacher, 1873) and P. (Z.) bajuvaricum
(Retenbacher, 1873) by virtue of its fewer, coarser ribs and tubercles, more of
which arise in pairs from the umbilical bullae. P. (Z.) proteus (Matsumoto,
1965) is much more coarsely ornamented than P. (Z.) aberlei.

Occurrence
Coniacian of western and southern Europe, and Zululand.

Peroniceras (Zuluiceras) bajuvaricum (Redtenbacher, 1873)
Figs 92B-C, 93

Ammonites bajuvaricus Redtenbacher, 1873: 107, pl. 24 (fig. 2a—c).

Gauthiericeras bajuvaricum (Redtenbacher): de Grossouvre, 1894: 88, pl. 9 (fig. 1), pl. 12 (figs
2a—b, 3a—b), text-fig. 35.

Schloenbachia (Gauthiericeras) bajuvarica (Redtenbacher): Boule, Lemoine & Thévenin,
1907: 22, pl. 12 (fig. 1—1a), text-fig. 22.

Gauthiericeras bajuvaricum (Redtenbacher): Diener, 1925: 152 (with synonomy).

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Gauthiericeras bajuvaricum (Redtenbacher) var. skoenbergensis Venzo, 1936: 105, pl. 9 (fig. 5),
pl. 11 (fig. 7).

Sornayceras bajuvaricum (Redtenbacher): Matsumoto, 1965: 226 et seq.

Sornayceras omorii Matsumoto, 1965: 230, pl. 42 (figs 1-2), text-figs 13-15.

Sornayceras aff. bajuvaricum (Redtenbacher): Szasz, 1976: 208, pl. 5 (fig. 2).

Sornayceras cf. bajuvaricum (Redtenbacher): Atabekjan & Akopjan, 1972: 10, pl. 3 (figs 4-5).

Type specimen

The lectotype is Redtenbacher’s (1873 pl. 24 (fig. 2a—c)) original figured
specimen from the Gosau Beds of Austria, designated by Reyment (1958: 43).

Material

NMB D1059 and SAS A343 from locality 72, degraded river cliff and north-
ern banks of the Mzinene River downstream of the Munywana River, Zululand,
St. Lucia Formation, Coniacian II or HI. SAS Z1632a—b from locality 92, bull-
_ dozer scrapings and adjacent hillslopes around pumping station on the farm Pan-
plaas, east-south-east of Hluhluwe, Zululand, St. Lucia Formation, Coniacian II
or III.

Dimensions

D Wb Wh Wb/Wh U R
Zio32a ~~ 55,0) 16,0951) 19508455) 0584 22,0(40,0) 11x2:18x2
ZENO (COO LV OC3.3)) ZN O@S.0) O85 23,0(38,3) ZAG
A343 WO IVC) AoVs5s) ~ O78 30,0(39,0) —
D1059 oi) A204) S20Ge53) On7il 35 ,0(40,2) os
Description

Coiling is involute with an umbilical diameter of about 38 to 40 per cent of
the total diameter, each whorl covering the ventral third of the preceding one.
The whorl section is compressed throughout, with flat parallel flanks and a fas-
tigiate venter. The venter bears a strong central keel bounded on either side by
a slight depression. There are weak lateral ridges that cannot be termed true lat-
eral keels. The umbilical wall is vertical to overhanging.

Ornament consists of weak umbilical bullae connected to clavate to
rounded ventrolateral tubercles by numerous single and bifurcating ribs.

The suture-line is complex with narrow-stemmed saddles of the P. (P.) dra-
vidicum type (Fig. 93).

Discussion

In Zululand the species is easily identified by the narrow umbilicus, high,
compressed whorl section, unicarinate, fastigiate venter, and ornament consist-
ing of single and bifurcating ribs.

Venzo (1936: 105-106) commented on the simplicity of the suture-line, but
sutures that he described appear to be those of juveniles. Suture-lines of speci-
mens of P. (P.) dravidicum taken at similar diameters (Fig. 35) appear identical

CRETACEOUS FAUNAS FROM SOUTH AFRICA DAT

and there can be no doubt that this is, indeed, a representative of P. (Zulu-
iceras).

Pending the revision of the Gosau material, a discussion of the affinities of
the species is omitted, except that the possibility exists that a number of super-
ficially similar, compressed species such as P. (Z.) omorii (Matsumoto)
(1965: 230, pl. 42 (figs la—b, 2a—b)) and P. (Z.) isamberti (Fallot) (1885: 232, pl.
2 (fig. 1)) may be mere variants of the species or subspecies only. P. (Z.) aberlei
is more sparsely ribbed on the outer whorls, as is P. (Z.) propoetidum (Redten-
bacher, 1873).

Occurrence

Middle Coniacian of western and southern Europe, Armenia and possibly
Japan; Coniacian II of Zululand.

Fig. 92. A. Peroniceras (Zuluiceras) aberlei (Redtenbacher, 1873). Cast of one of

Redtenbacher’s original specimens from the Oberésterreichisches Landesmuseum, Linz.

B-C. Peroniceras (Zuluiceras) bajuvaricum (Redtenbacher, 1873). B. SAS Z1632a
C. SAS A343. All x 1.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

—i— =
=

' : 0
mm

Fig. 93. Peroniceras (Zuluiceras) bajuvaricum (Redtenbacher, 1873).
Suture-lines of SAS Z1632a.

Genus Gauthiericeras de Grossouvre, 1894

(= Ciryella Wiedmann, 1960: 763; Andersonites van Hoepen, 1965: 29; Flumi-
nites van Hoepen, 1965: 31; Hluhluweoceras van Hoepen, 1965: 33; Falsebayites
van Hoepen, 1965: 34)

Type species
Ammonites margae Schliiter, 1867: 29, pl. 5 (fig. 2) by original designation
of de Grossouvre, 1894: 87.

Diagnosis

Medium sized to large, moderately evolute to involute, whorl section de-
pressed to compressed, trapezoidal to quadrate with strong, entire, undulating
or serrated siphonal keel flanked by a groove on either side, the outer edge of

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 94. Gauthiericeras margae (Schliiter, 1867). Holotype, the specimen figured by
Schliter (1867, pl. 5 (fig. 2)). x 0,6. (See also fig. 19c.)

239

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 95. Gauthiericeras margae (Schliter, 1867). Holotype, the specimen figured by
Schliiter (1867, pl. 5 (fig.2)). x 0,6.

which may be strengthened into a low ridge, but not a lateral keel. Strong ribs
arise from umbilical bullae in pairs or singly and may branch again from inner to
mid-lateral bullae, if present. Shorter intercalated ribs sometimes present. All
ribs terminate in strong ventrolateral clavi.

Suture rather simple, with little-incised elements; saddles square.

Discussion

Gauthiericeras is easily distinguished from other peroniceratids by its strong
siphonal keel, lack of true lateral keels, strong ribs and the presence, in the type

CRETACEOUS FAUNAS FROM SOUTH AFRICA 241

species and some others, of a lateral tubercle. Persistent ornament and paired or
branching ribs distinguished it from /shikariceras.

Wiedmann (1960: 763, pl. 8 (figs 1-4), text-fig. 16) introduced the subgenus
Gauthiericeras (Ciryella), type species G. (C.) vascogoticum. The original diag-
nosis is as follows: ‘Coquille a enroulement extrémement évolute qui laisse voir
les tubercles marginaux. Section de tours rectangulaire. Les cOtes restent sim-
ples et serrées chez l’adulte et sont munies d’un tubercle ombilical allongé dans
le sens de la cote et d’un tubercle marginal en massue. Au stade jeune, les cotes
qui couvrent seulement les flancs sont bifurquées sur le tubercle ombilical ou
simples. Région siphonale tectiforme et avec une caréne siphonale entiére. Cloi-
sons, comme Gauthiericeras et Peroniceras.’

In Wiedmann’s discussion, Ciryella was suggested as an intermediate form
between Gauthiericeras and Peroniceras, with the general coiling and evolute
umbilicus of the latter (U = 40 % at D = 42 mm, 51 % at D = 73 mm, and
52 % at D = 125 mm) and similar ribbing. The features recalling Gauthiericeras
were said to be the single, entire siphonal keel.

Wiedmann gave the age of his species as Lower Coniacian, and it is based
on a specimen recorded by Ciry (1940: 217). As discussed previously (p. 128), the
earliest Coniacian ammonite zone recognized by Wiedmann in his 1960 paper
and subsequently, that of Tissotioides haplophyllus, would appear to be Middle
Coniacian, while Wiedmann’s Zone III of Gauthiericeras vallei, also placed in
the Lower Coniacian in 1960, is Upper Coniacian. It is suggested here that Ci-
ryella is actually an Upper Coniacian form. It differs from Gauthiericeras only in
being very evolute, yet specimens of Gauthiericeras margae from France have U
= up to 42 %. G. nouelianum (d’Orbigny, 1850) has U = as little as 20 % and
G. obesum van Hoepen, 1955, has U = up to 46,5 %. Ciryella thus stands in the
same relationship to Gauthiericeras as does Peroniceras (Peroniceras) to the
more involute P. (Zuluiceras), but without the distinctive differences in orna-
ment shown by these two subgenera. At most it is a subgenus of Gauthiericeras,
and even this seems unnecessary.

Van Hoepen (1965) was led astray by the wide extent of intraspecific varia-
tion shown by the Zululand Gauthiericeras, the numbers of which by far exceed
that of all other described Gauthiericeras collections. This led to his erection of a
series of genera: Andersonites, Hluhluweoceras, Falsebayites, and Fluminites. In
1970 and 1971 low water-levels in False Bay permitted the authors to collect a
range of material in situ, on the basis of which it is possible to demonstrate a
series from very evolute individuals close to G. roquei (Peron, 1897) via more
narrowly umbilicate forms referred to G. obesum van Hoepen, 1955, to nar-
rowly umbilicate forms with rapidly expanding whorls that include Van
Hoepen’s Andersonites, Falsebayites, and Hluhluweoceras, the type species of
which are all regarded as synonyms, the name Gauthiericeras |‘Andersonites'’ | lis-
teri van Hoepen, 1965, having priority. The type species of Fluminites, F. albus
van Hoepen, 1965, is retained as a further species of Gauthiericeras, allied to a
very involute species G. albiforme sp. nov. Gauthiericeras libertae van Hoepen,

ANNALS OF THE SOUTH AFRICAN MUSEUM

242

‘1 x ‘((0-8] 8) Sz ‘Id ‘EZ8T) Je4yoequelpoy Aq poinsy uouniseds ay], “(2981 ‘1oINIYOS) avsuvwu spsaq14alyINVy 96 “314

CRETACEOUS FAUNAS FROM SOUTH AFRICA 243

1955, of which G. planatum van Hoepen, 1955 is a synonym, is also maintained
as a separate species. Gauthiericeras shows a number of similarities to Prionocy-
cloceras Spath, 1926, and, indeed, Spath (1926: 80) considered it as transitional
between Collignoniceras and Gauthiericeras. The type species differ in the
strong umbilical and lateral tubercles of Gauthiericeras margae, and the absence-
of the huge spines of adult Prionocycloceras guyabanum. Both share a simple su-
ture and strongly serrated Keel flanked by shallow grooves but, whereas the ribs
extend from the ventrolateral tubercles to the siphonal keel in Prionocycloceras,
they decline in Gauthiericeras. Many P. guyabanum specimens have fine inter-
calated flank ribs.

Some species show intermediate features, but the two genera are distinct.

Prionocycloceras may well be the link between Upper Turonian Priono-
cyclus and Upper Coniacian Gauthiericeras. Of species referred to Gauthieri-
ceras by recent workers, G. margae peruanum Briggen, 1910, is a Forresteria
(Forresteria), as 1s Gauthiericeras hoepeni Collignon, 1965. Of Spanish species,
Gauthiericeras margae turzoi Karrenberg, 1935, G. margae gorda Karrenberg,
1935, G. vallei Ciry, 1940, and G. vallei gorda Ciry, 1940, are Prionocycloceras.

Occurrence

Upper Coniacian of France, Spain, central Europe, north, east and west
Africa, Madagascar, New Caledonia, Mexico, Colombia, and Peru. Middle
Coniacian records are doubtful.

Gauthiericeras obesum van Hoepen, 1955
Figs 97-114

Gauthiericeras margae (Schliter): Besairie, 1936: 203, pl. 24 (figs 17-18). Venzo, 1936: 104, pl.
10 (fig. 2). ? Collignon, 1965: 49, pl. 435 (fig. 1798).
Gauthiericeras obesum van Hoepen, 1955: 374, figs 25-28.

Type specimen

The holotype, by monotypy, is SAS Z12, (Figs 107-108), collected loose
from the mouth of the Hluhluwe River, Zululand.

Material

SAS 85, 92/2, H200/33, H200/37, H200/76, H200/94, H200/109, SAS Z200,
ZS) 2517, Z201I5, Z2031, 22035, BMNH €83760—C83790, and
SAM-PCZ5947-50, from locality 83 locally known as Mason’s Camp, Zululand,
St. Lucia Formation, Coniacian IV. SAS H203/8, SAS Z12, SAM-—PCZ5951
from boulder- and concretion-strewn slopes west of the St. Lucia Game Park
fence, east-south-east of Hluhluwe, Zululand, St. Lucia Formation, Coniacian
IV. SAS Z1055 and Z1057 from locality 73, degraded river cliffs and stream sec-
tions on north side of the Mzinene River downstream of the Munywana River,

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Zululand, St. Lucia Formation, Coniacian IV. SAS A1510 and BMNH
C83801—C83806 from locality 22, on the north side of the Nyalazi River near
Nyalazi River Trading Store, Zululand, St. Lucia Formation, Coniacian IV.

Dimensions
D Wb Wh Wb/Wh U R

A1754 98,0 350G5,7) B30S750) 0,95 38,0(38,8) —:17
H200/76 81,0 WI (YU CS53)}) — SNOGS,3) 0,87 31,0(38,3) —:—
PCZ-5948 85,0 27,0(31,8) 30,0(35,3) 0,90 —35,0(41,2) 18:18
H200/34 108,0 36,0(33,3) 40,0(37,0) 0,90 40,0(37,0) —:—
PCZ5497 108,0 37,0(34,2) 42,0(38,9) 0,88 41,0(40,0) —:17
PCZ5949 108,0 HUGS) “SOO G,10) 0,97 43 ,0(39,8) 10x20
Z200 123,0 45,0(36,6) 50,0(40,6) 0,90 48 ,0(39,0) —:20
SAS-85 124,0 = 47,0(37,9) =» 485068,9) AiG
Z1055 I7R0 40,0(31,5) 46,0(36,2) 0,87 53,0(41,7) 20:20
ZA73 129,0 44,0(34,1) 47,0(36,4) 0,94 53,0(41,1) 17;
H203/8 130,0 41,0(31,5) 47,0(36,1) 0,87 57,0(43,8) 18:18
H200/33 134,0 45,0(33,6) 48,0(35,8) 0,94 53,0(39,6) —:—
Z1057 134,0 39,0(29,1) 45,0(33,6) 0,87 —-52,0(38,8) 21:21
Lisi 140,0 48,0(35,0) 52,0(37,1) 0,92 20710) 19:19
Z191 150,0 4,0(29,3) 51,0(34,0) 0,86 +~—«64,0(42,7) 16:—
Z12 150,0 52,0(34,7) 57,0(38,0) 0,91 59,0(39,3) 18:18
H200/94 164,0 us 61,0(37,2) — Gil.) 17:18
Z2021 TAO) 5480283) S810G 457), 0193) oon GiES) i
DTD 167,0 55,0(32,9) 59,0(35,3) 0,93 ~—-67,0(40,1) 23:23
PCZ5950 185,0 59,0(31,9) 66,0(5,7) 0,82 76,0(41,1) 20:20
Z2035 190,0 58,0(30,5) 67,0(35,3) 0,87 82,0(43,2) 19:19
Z874 204,0 60,0(29,4) 71,0(34,8) 0,84 91,0(44,6) PEED
92/2 221,0 = 71 ,0(32,1) SOLO CS a) 2s
Z2030 225,0 65,0(28,9) 74,0(32,9) 0,88 98,0(43,6) 18:18
92 235,0 70,0(29,8) 80,0(34,0) 0,87 ~—:103,0(43,8) DieO
H200/109 MBTAD 73,0(30,8) 85,0(35,9) 0,86 100,0(42,2) Vee
TLAAVIS) 245,0 72,0(29,4) 80,0(32,5) 0,90 106,0(43,3) IGS
Z877 285 ,0 75,0(26,3) 90,0(31,6) 0,83 136,0(43,7) 19:19
PCZ5951 308,0 83,0(26,9) 101,0(32,8) 0,82 141,0(45,8) 18:18
ZLB TS 311,0 83,0(26,7) 98,0(31,5) 0,85 144,0(46,3) tee
Description

The umbilical width of the species varies between 37 and 46 per cent of the
total diameter, but the majority of specimens have umbilical widths of 39 to 44
per cent (Fig. 97). Coiling is very evolute and the whorls are only slightly over-
lapping, so that the dorsal zone of impression is very shallow. It appears that the
more widely umbilicate specimens grew to a larger size than the more narrowly
umbilicate ones, but this may possibly be only an artefact of preservation rather
than an indication of dimorphism. The whorl section is compressed throughout,
Wb/Wh ratios ranging from 0,82 to 0,97.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 245

No. of specimens

Si aaI0 839 AO 41” 142) 45) 44 AS 4

Umbilical @

Fig. 97. Histogram illustrating distribution of umbilical width in Zululand population of
Gauthiericeras obesum van Hoepen, 1955.

Density of ribbing in the population is reasonably constant, varying be-
tween 16 and 23 per whorl, but the majority of specimens have a rib density of
17 to 18 per whorl. Strength of ornament is variable, but this depends to some
extent on preservation, and is often accentuated through abnormal thickening of
the shell as a result of diagenetic growth of calcite. Specimen SAS Z473 (Fig.
98) has much of the inner whorls preserved. Here ornament consists of radial to
slightly rursiradiate, very strong ribs, apparently all single, each bearing two
rows of tubercles. The umbilical ones are pinched bullate, protruding far beyond
the level of the ribs, especially in places where the shell has been thickened dia-
genetically. The ventrolateral tubercles are clavate and pointed, and lean
against and impress into the umbilical wall of the succeeding whorl. On the
outer whorls, the ventrolateral tubercles become weaker and rounded clavate,
whereas the umbilical tubercles remain prominently bullate and merge into the
ribs.

Differences in strength of ornament in the adult stage become obvious
when comparing specimens SAS Z869 (Fig. 99) and SAS Z2030 (Fig. 100), the
latter with strong ribbing and tuberculation, and the former with virtually
smooth flanks.

Some specimens retain the body chamber (e.g. SAM-—PCZ5951) (Figs
101-102), consisting of about three-quarters of a whorl. Here ribbing becomes
bolder and wider spaced. The umbilical bullae may disappear completely, fusing
with the rib. The ribs may also curve gently forward or become slightly sinuous.

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 98. Gauthiericeras obesum van Hoepen, 1955. SAS Z473. Specimen with part of inner
whorls preserved, showing early strong ornament. X 1.

247

CRETACEOUS FAUNAS FROM SOUTH AFRICA

th weak ornament

i

Specimen w

SAS Z869
100. x 0,67

.

1955
igure

b)

besum van Hoepen
compared to F

lericeras O

Fig. 99. Gauth

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

A very large body chamber fragment, NMB D1060 (Fig. 103), with two strong
ribs but distinct umbilical and ventrolateral tubercles, may belong here or per-
haps to G. listeri described below.

The suture lines are very simple as shown in Figures 99-100, and 105.

Discussion

Gauthiericeras obesum most closely resembles the type species, G. margae
(Schliiter, 1867). Schliiter’s figure is highly idealized, and a cast of the basis of
Schliter’s figure is shown in Figures 20C and 94-95. The outer whorl is crushed
almost flat and the inner whorls survive only as an external mould. A topotype
specimen, also in Schliter’s collection, the specimen figured by Redtenbacher
(1873: 109, pl. 25 (fig. la—c)) from the Gosau Beds of Austria, De Gross-
‘ouvre’s material (1894: 90, pl. 15 (figs 1-2)) (see Fig. 96 herein), plus new ma-
terial from Touraine show that G. margae differs from G. obesum in several
respects. The ribs commonly arise from bullae in pairs in juveniles, a feature not
seen in G. obesum; there is a lateral tubercle from which the ribs also branch,
and the siphonal keel is strongly serrated. Adults are more closely comparable
but, whereas the umbilical bullae perch on the umbilical shoulder in G. obesum,
they migrate outward to an inner flank position in G. margae.

G. nouelianum (d’Orbigny, 1850) has been illustrated by Sornay (1955); it is
much more involute than G. obesum (U = 20-27,5 %; see Kennedy in press for
details). It co-occurs with G. margae and, like that species, has branching ribs,
lateral tubercles, and a serrated keel, all of which render it easily separable from
G. obesum. G? boreaui (de Grossouvre) (1894: 111, pl. 7 (fig. 3)) is a diminu-
tive species, involute, feebly and flexuously ribbed, and lacks ventrolateral
tubercles.

Gauthiericeras roquei Peron, 1897: 52, pl. 8 (fig. 1), pl. 9 (figs 1-2), pl. 17
(fig. 6)) (Fig. 115 herein) is a very evolute species (U = 43-44 %). Ornament is
very distant, with only ten ribs on the outer whorl compared with nearly double
that number in similarly evolute G. obesum.

Gauthiericeras listeri (van Hoepen, 1965), of which Hluhluweoceras fugiti-
vum van Hoepen, 1965, Falsebayites peregrinus van Hoepen, 1965, and
Gauthiericeras listeriforme Collignon, 1965, are synonyms, is a contemporary of
G. obesum. It differs in being more involute (U = 32-36 %) with a compressed
whorl section in most specimens. It stands in the same relationship to G. obesum
as does G? nouelianum to G. margae. These involute and evolute forms may be
dimorphs, but this cannot be proven.

Occurrence

Coniacian IV of Zululand. Upper Coniacian Gauthiericeras margae and
Prionocycloceras guyabanum Zone of Madagascar.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 249

Fig. 100. Gauthiericeras obesum van Hoepen, 1955. SAS Z2030.
Specimen with strong ornament compared to Figure 99. x 0,63

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 101. Gauthiericeras obesum van Hoepen, 1955. SAM-—PCZ5951. Large, evolute specimen :
with body chamber up to three-quarters of a whorl in length. x 0,48. |

eal

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 102. Gauthiericeras obesum van Hoepen, 1955. SAM-—PCZ5951,

ions of the body chamber. X 0,6

dorsal views of two sect

DD) ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 103. ?Gauthiericeras obesum van Hoepen, 1955. NMB D1060, large body chamber
fragment. x 0,72.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 253

Fig. 104. Gauthiericeras obesum van Hoepen, 1955. SAS A1510. Individual differing from most
Zululand specimens in showing numerous rib bifurcations. x 0,82.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 105. Gauthiericeras obesum van Hoepen, 1955. SAS Z877,
large specimen with wide umbilicus. x 0,5.

255

CRETACEOUS FAUNAS FROM SOUTH AFRICA

5 28 OLoile

1cus

th wide umbil

imen WI

ceras obesum van Hoepen, 1955. SAS Z877,

large spec

teri

Fig. 106. Gauth

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

SEN

Fig. 107. Gauthiericeras obesum van Hoepen, 1955. SAS Z12, the holotype. x 0,8.

|

2571

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 108. Gauthiericeras obesum van Hoepen, 1955. SAS Z12, the holotype. x 1.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 109. Gauthiericeras obesum van Hoepen, 1955. SAS Z872. X 0,88.

Z59

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 110. Gauthiericeras obesum van Hoepen, 1955. SAS Z872. x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

260

. SAS Z2015. x 0,6.

besum van Hoepen, 1955

lericeras O

. 111. Gauthi

1g

F

261

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘TX “LOPSZOd-WVS ‘SS6] ‘Uodo0p{ URA WNsago sp4saold

AIYINDE) “ZI

|

262

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 113. Gauthiericeras obesum van Hoepen, 1955. SAM-PCZ5948. x 1.

263

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘TX ‘0b6SZOd

WVS SSol

‘usdooH URA WiNSagO SpAdIAAIYINDD “pL “Bl

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 115. Gauthiericeras roquei Peron. (After Peron, 1896, pl. 8 (fig. 1). Reduced.)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 265

Gauthiericeras listeri (van Hoepen, 1965)

Figs 116-123 -

Andersonites listeri van Hoepen, 1965: 31, pl. 23, text-fig. 8b, d.
Hluhluweoceras fugitivum van Hoepen, 1965: 33, pl. 25, text-fig. 8a, c.
Falsebayites peregrinus van Hoepen, 1965: 34, pl 26. text-fig. 9f-g.
Gauthiericeras listeriforme Collignon, 1965: 49, pl. 435 (fig. 1797).

Type specimens

The holotype, by original designation, is SAS Z754 (Figs 116-117B), from
locality 91, degraded river cliffs and artificial cut west of the boundary fence of
St. Lucia Game Reserve, east-south-east of Hluhluwe, Zululand. This area is
strewn with rubble derived from the St. Lucia Formation, Coniacian IV and V,
and the precise horizon cannot be stated. The holotype of Hluhluweoceras fugiti-
vum is SAS Z755, from the same horizon as the holotype of Gauthiericeras lis-
teri. The holotype of Falsebayites peregrinus is SAS Z341, the paratype is SAS
Z339, both from a little to the north of the previous site. Their precise horizon is
similarly uncertain.

Material

SAS H206/2, a topotype, was also collected loose; SAS Z342 is from ‘close
by’. BMNH C83791 is from locality 83, where it is firmly dated as Coniacian IV.

Dimensions

D Wb Wh Wb/Wh U R
H206/2 119.0 42 ,0(35,3) 48 ,0(40,3) 0,87 43 ,0(36,1) 18:19
TSS) 180,0 65,0(36, 1) 71,0(39,4) 0,91 64,0(35,6) —
Z341 c.190,0 c.68,0(35,8) c.72,0(37,9) c.0,94 c¢.65,0(34,2) —
Z754 195,0 74,0(37,9) 73,0(37,4) 1,01 70,0(35,9) —

G. listeriforme (after Collignon)
102,0 47,0(46,0) 42,0(41,2) 2 = san 0G453) 9:10

Description

The whorls expand rapidly and the umbilicus is narrow, comprising 32 to 36
per cent of the total diameter. The whorl section is compressed in the majority
of specimens but may become slightly depressed on the later septate whorls. The
umbilical wall is generally rounded, the flanks are very little inflated, and the
venter is broadly rounded.

Only one specimen has the inner whorls preserved (SAS H206/2 Fig. 122).
The ventrolateral tubercles are concealed in the umbilical seam and the umbili-
cal tubercles are displaced somewhat up the flanks, as in G. margae. They are
conical to radially elongated, but very irregularly developed. Intercalatory ribs,
which do not bear umbilical tubercles, sometimes occur.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

econ tieia

Fig. 116. Gauthiericeras listeri (van Hoepen, 1965). Holotype SAS Z754. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 267

A B

Fig. 117. Gauthiericeras listeri (van Hoepen, 1965). A. SAS Z342, inner whorls of paratype of
Falsebayites peregrinus van Hoepen. X 1. B. SAS Z754, inner whorls of holotype of G. listeri.
xX 2.

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 118. Gauthiericeras listeri (van Hoepen, 1965). SAS Z342, paratype of Van Hoepen’s
Falsebayites peregrinus. X 0,64. .

CRETACEOUS FAUNAS FROM SOUTH AFRICA 269

Fig. 119. Gauthiericeras listeri (van Hoepen, 1965). SAS

Z342, paratype of Van Hoepen’s Falsebayites peregrinus;

section on body chamber showing fastigiate venter and
slight constriction. X 1.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 120. Gauthiericeras listeri (van Hoepen, 1965). SAS Z341, holotype of Van Hoepen’s
Falsebayites peregrinus. X 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA pay

Fig. 121. Gauthiericeras listeri (van Hoepen, 1965). SAS Z755. Holotype of Van Hoepen’s
Hluhluweoceras fugitivum. x 0,71.

LD ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 122. Gauthiericeras listeri (van Hoepen, 1965). SAS H206/2, specimen showing early
whorls. Xx 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 27S

= eae eee 0

ee)

Fig. 123. Gauthiericeras listeri (van Hoepen, 1965). Suture-lines of
Z755, Z754, Z341. x 1. (After Van Hoepen 1965, figs 8a—b, 9g.)

The venter is distinctly unicarinate, with a single, strong central keel
bounded on either side by a depression of variable strength. The ribs often con-
tinue beyond the ventrolateral tubercle to the area of these depressions, thus
creating the impression of wavy lateral keels, but these are not true keels.

Adult ornament is very variable as the figures show, but differences are all
within bounds of specific variation. It ranges from weak (SAS Z755) (Fig. 121)
through fairly well developed (as in SAS Z341) (Fig. 120) to very strong (SAS
SA aGeig. 116).

A large part of the body chamber is preserved in SAS Z342, occupying
about three-quarters of a whorl. At the apertural end of this specimen the ribs
become increasingly prorsiradiate, the central keel less prominent, and prorsi-
radiate striae cross the venter (Fig. 119).

Suture-lines are shown in Figure 123A-C.

Discussion
The species as here interpreted comprises virtually the whole of Van

Hoepen’s subfamily Gauthiericeratinae and includes the type species of his ‘gen-
era’ Andersonites, Hluhluweoceras, and Falsebayites. This species is probably

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

derived from G. obesum through reduction of the umbilical width and increase
in rate of whorl expansion.

‘Andersonites’ listeri is the most strongly ornamented morphological variant
of the species and the inner whorls are somewhat reminiscent of Peroniceras
(P.) subtricarinatum as far as lateral ornament is concerned. The venter, how-
ever, 1S distinctly unicarinate, and not tricarinate as stated by Van Hoepen
(1965: 29) (Fig. 117B). There is only one central keel with a spiral furrow on
either side. A slight extension of the lateral ribs to these depressions creates the
appearance of wavy lateral keels. This together with imperfect shell preserva-
tion, probably led Van Hoepen to believe that the specimen was tricarinate.

‘“Hluhluweoceras’ fugitivum (Fig. 121) has massive quadrate inner whorls
with weak lateral ornament, but can be connected to G. listeri by transitional
forms, so that separation is artificial.

‘Falsebayites’ peregrinus (Figs 117A, 118-119) the only species in Van
Hoepen’s subfamily Gauthiericeratinae based on more than one specimen,
shows a very rapid increase in whorl height and an umbilical diameter of only 32
per cent. The general pattern of ornament, however, is that of G. Jisteri.

Collignon (1965: 49, pl. 435 (fig. 1797)) reported a species from the Middle
Coniacian of Madagascar G. listeriforme, that is very much like G. listeri. The
whorl section is much wider than high, the whorl breadth to height ratio being
1,12. It, too, is a synonym of G. listeri.

Occurrence

In Zululand well-dated specimens are restricted to Coniacian IV. The
Madagascan specimen referred to the species is said to be from the ‘Coniacian
Moyen’.

Gauthiericeras album (van Hoepen, 1965)
Figs 124-126

Fluminites albus van Hoepen, 1965: 31, pl. 24, text-fig. 9c—e.

Type specimen
The holotype is SAS Z757 (Fig. 124) from localities 91 or 92 near the estu-
ary of the Hluhluwe River, Zululand, St. Lucia Formation, Coniacian IV or V.

Material

BMNH C83792 is from locality 83 and is precisely localized as Coniacian
IV. A more complete specimen, SAS Z871, is from the same locality and hori-
zon as the holotype.

Dimensions

D Wb Wh Wb/Wh U R
Z871 243,0 69,0(28,4) 86,0(35,4) 0,80 86,0(35,4) T7ei@
TENS — 31,0 40,0 Oa

CRETACEOUS FAUNAS FROM SOUTH AFRICA DAS

Fig. 124. Gauthiericeras album (van Hoepen, 1965). Holotype SAS Z757. x 1.

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 125. Gauthiericeras album (van Hoepen, 1965). SAS LST erxe 0563

PAT

CRETACEOUS FAUNAS FROM SOUTH AFRICA

0565:

(van Hoepen, 1965). SAS Z871.

Fig. 126. Gauthiericeras album

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Coiling is moderately involute, with an umbilical diameter of 35 per cent in
the only complete specimen available. The whorl section is ovoid throughout
ontogeny, compressed, with little-inflated flanks (the maximum width at mid-
flank), and a broadly rounded venter. The umbilical wall is gently rounded.

Ornament on the inner whorls of the holotype is very weak, consisting of
umbilical bullae, prorsiradiate ribs and ventrolateral clavi. The venter appears
tricarinate, but this is a result of the strong development of depressions on either
side of the keel. On the outer whorls ornament strengthens considerably. The
ribs become sharp-crested and widely spaced, whereas the umbilical tubercles
become progressively radially elongated and eventually merge with the ribs. The
ventrolateral tubercles similarly merge with the ribs, giving the latter a club-like
appearance. Part of the body chamber is preserved in SAS Z871 (Fig. 125) and
_ here adult ornament is fully developed.

Discussion

Unfortunately this species is poorly represented in numbers, so that the
range of intraspecific variation cannot be established. It is easily distinguished
from G. obesum by the ovoid, rather than quadrate to rectangular whorl section
of the latter. The outer whorls of G. listeri are remarkably similar to those of
G. album, but, again, the whorl section of the former is more quadrate.

Occurrence
Coniacian IV of Zululand.

Gauthiericeras albiforme sp. nov
Figs 127-129

Type specimens

The holotype is SAS 92/4 (Figs 127-128) from between localities 83 and 84,
near Picnic Point, False Bay, Zululand, St. Lucia Formation, Coniacian V.
Paratypes are SAS 92/2-3, from the same locality and horizon as the holotype.

Dimensions

D Wb Wh Wb/Wh U R
92/3 113505 ~ 36,0159) 48 ,0(42,5) OS 30,0(26,6) —
92/2 160,0 48,0(30,0) 58,0(36,2) 0,83 55,0(34,4) —
92/4 131205 Pole 0G3:7) 72,0(39,8) 0,85 62,0(4,2) 1Se20
Description

The umbilicus is narrow in the early growth stages, comprising only 27 per
cent of the diameter, but increases to about 34 per cent in larger individuals. In
adults the whorl section is distinctly subtrigonal, with an overhanging umbilical
wall and little-inflated flanks converging to a narrow, unicarinate venter.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 279

Fig. 127. Gauthiericeras albiforme sp. nov. Holotype SAS 92/4. x 0,86.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 128. Gauthiericeras albiforme sp. nov. Holotype SAS 92/4. x 0,96.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 281

B

Fig. 129. Gauthiericeras albiforme sp. nov. Whorl section and suture-line. x 0,5.

Ornament throughout consists of distinctly pinched, auricular umbilical bul-
lae that merge into strong, more or less radial or slightly prorsiradiate concave
ribs. These terminate in indistinct ventrolateral tubercles. On the outer whorls
ribbing is generally single, but on the inner ones bifurcations may occur at the
umbilical tubercle. Rib density is 19 to 20 on the outer whorls.

The venter is distinctly unicarinate, with only shallow spiral depressions on
either side of the keel.

Discussion

Distinguishing features of this species are the narrow umbilicus and the sub-
trigonal whorl section with an overhanging umbilical wall. As far as ornament is
concerned, there is great similarity to G. album, described above. Although the

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

material is limited, it may be possible to derive G. albiforme from G. album
through slight decrease in umbilical diameter and increasing rate of whorl ex-
pansion. Given enough material connecting the two species, separation at sub-
specific level would probably be sufficient.

The subtriangular whorl section serves to distinguish the species from
G. obesum and G. listeri.

Occurrence

Upper Coniacian of Zululand. This appears to be the last surviving species
of Gauthiericeras.

Gauthiericeras libertae van Hoepen, 1955
Figs 130-134

Gauthiericeras libertae van Hoepen, 1955: 372, figs 23-25.
Gauthiericeras planulatum van Hoepen, 1955: 374, figs 29-31.

Type specimens

The holotype of Gauthiericeras libertae is SAS Z4 (Figs 132-134), it was
found by Van Hoepen in a pile of rubble probably at locality 91 or 92 of
Kennedy & Klinger (1975) near the Hluhluwe estuary, Zululand, St. Lucia For-
mation, probably Coniacian IV. The holotype of G. planulatum is SAS Z3, also
from near the Hluhluwe estuary close to locality 89 or 90 of Kennedy & Klinger,
St. Lucia Formation, probably Coniacian IV.

Material

SAS Z1136, from locality 73, degraded river cliffs and stream exposures on
the Mzinene River downstream from the junction with the Munywana, Zulu-
land, St. Lucia Formation, Coniacian IV or V. SAS H205/22 is from locality 90,
SAS H205/1 from locality 90 or 91, both near the Hluhluwe estuary, Zululand,
Coniacian IV or V. SAS Z2018 and BMNH C83797—C83799 from locality 83,
foreshore exposures at Mason’s Camp, Zululand, St. Lucia Formation, Coni-
acian IV.

Dimensions

D Wb Wh Wb/Wh U R
Z2018 131,0 41,0(31,3) 52,0(39,7) 0,79 48,0(36,6) 22:22
Z3 134,0 37,0(27,6) 51,0(38,1) O72 46,0(34,3) 18:21
H205/22 13500 37-0 C72) 49 ,0(36,0) OFS 53,0(38,9) —
Z1136 137,0 43,0(31,4) 50,0(36,5) 0,86 50,0(36,5) —
H205/1 148,0 41,0(27,7) 53,0(35,8) Od 54,0(36,5) 16:16
ZA 150,0 40,0(26,7) 54,0(36,0) 0,74 55,0(6,7) ~ ies0

CRETACEOUS FAUNAS FROM SOUTH AFRICA 283

Fig. 130. Gauthiericeras libertae van Hoepen, 1955. SAS Z3 holotype of
G. planulatum van Hoepen, 1955. x 0.96.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

——.~—
_—
oS

Fig. 131. Gauthiericeras libertae van Hoepen, 1955.

Whorl section and suture-line of the holotype of

G. planulatum. x 1. (After Van Hoepen 1955, figs
30-31.)

Description

Coiling is evolute, the umbilical diameter varying between 34 and 38 per
cent of the total diameter, with successive whorls overlapping to the ventral
tubercle or a little beyond, so that about 25 per cent of the previous whorl is
covered. The whorl section is compressed throughout, with whorl breadth to
height ratios that vary between 0,72 and 0,86. The inner flanks are gently
rounded, the outer flanks flattened and subparallel or slightly convergent, the
venter narrow and fastigiate.

There are 16 to 22 weak to prominent umbilical bullae per whorl. These
give rise to either single ribs or pairs of ribs that vary from weak to strong be-
tween individuals but are straight and prorsiradiate to mid-flank and curve
slightly forward to terminate in weak ventral clavi. The ribs are prolonged for-
ward to the edge of a shallow groove that flanks a blunt undulating siphonal
keel. The outer edges of the grooves are also undulose, giving an appearance of
incipient lateral keels.

Suture simple, as in other members of the genus (Figs 131, 134).

Discussion

Van Hoepen separated Gauthiericeras libertae and G. planulatum on the
basis of the predominance of single ribs in /ibertae and the predominance of
paired ribs in planulatum, as well as sutural details. New material shows that
there is variation in this respect, and the common features of overall shell form
and the distinctive ventral ornament with crenulate siphonal keel and crenulate
edges to the flanking grooves suggest that only a single species is present.

Ribbing style, evolute coiling combined with compressed whorls and the
distinctive venter distinguish this species from all others.

Occurrence
Coniacian IV of Zululand.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 285

Fig. 132. Gauthiericeras libertae van Hoepen, 1955. Holotype SAS Z4. x 0,85.

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 133. Gauthiericeras libertae van Hoepen, 1955. Holotype SAS Z4. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 287

Fig. 134. Gauthiericeras libertae van Hoepen, 1955.
Whorl section and suture-line of holotype. x 1.
(After Van Hoepen 1955, figs 24-25.)

CONCLUSIONS

A general reappraisal of Coniacian stratigraphy and revision of the very rich
peroniceratid faunas found in Zululand provide an outline of the evolution of
the group that is of some significance for interregional correlation and elucida-
tion of evolutionary patterns among late Cretaceous ammonites (Fig. 135).

The earliest genus of the Peroniceratinae is Peroniceras itself. The appear-
ance of P. (Peroniceras) (of which Fraudatoroceras van Hoepen, 1965, Cobba-
noceras Matsumoto, 1965, and Gloriaceras Etayo-Serna, 1979, are synonyms)
marks the base of the Middle Coniacian as that term is used here. There is a hi-
atus in the record between the first appearance of Peroniceras and the
Collignoniceratinae, which are believed to be the ancestral stock spanning the
Lower Coniacian. Matsumoto (1965: 213) believed that the origin of Peroniceras
lay in Prionocyclus, and the Prionocyclus lineage certainly survived to give rise
to Prionocycloceras, which extends to the Upper Coniacian. Matsumoto sug-
gested that the lateral keels of Peroniceras arose by the elevation of the outer
edges of the grooves flanking the keel of ancestral Prionocycloceras. Indepen-
dent observations by Etayo-Serna (1979) and the present authors shows that the
Keels arise by elongation and eventual absorption of outer ventrolateral clavi
into a keel, and that both lateral and siphonal keels of several Peroniceras (Per-
oniceras) species show undulations at some stage in ontogeny.

The suture of the type species of Peroniceras (Peroniceras), P. (P.) moureti
de Grossouvre, 1894 (a synonym of P. (P.) tridorsatum (Schliiter, 1867)) is rela-
tively simple, like that of ancestral Collignoniceratinae. Other species, such as
P. (P.) dravidicum Kossmat, 1895, have a much more complex suture, with
long, narrow and finely indented saddles. Evolution must have been from the
simple P. tridorsatum type to complex P. dravidicum type, although there is no

ANNALS OF THE SOUTH AFRICAN MUSEUM

288

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detailed stratigraphy to support this. In Zululand, P. (Peroniceras) of both types
are confined to the Middle Coniacian. Only in Japan is there a suggestion that
P. (Peroniceras) extends above this and into the Santonian; however, the sup-
porting evidence is not wholly convincing.

Peroniceras (Zuluiceras) van Hoepen, 1965 (of which Zuluites van Hoepen,
1965, and Sornayceras Matsumoto, 1965, are synonyms) also appears in the
Middle Coniacian, with some slight evidence (in Zululand at least) that it
slightly post-dates the nominate subgenus. It has a relatively complex suture of
P. dravidicum type. The earliest species are relatively evolute, but through the
higher Middle and lower Upper Coniacian there is a trend towards increasing in-
volution, compression of the whorls, and loss of ornament.

Gauthiericeras (of which Ciryella Wiedmann, 1960, Fluminites van Hoepen,
1965, Andersonites van Hoepen, 1965, Hluhluweoceras van Hoepen, 1965, and
Falsebayites van Hoepen, 1965, are synonyms) appears at the base of the Upper
Coniacian in Zululand. Only the cryptic diminutive ‘Schloenbachia’ boreaui de
Grossouvre, 1894, which may appear in the Middle Contacian in France, points
to a possible slightly earlier appearance. The genus shows much stronger simI-
larities to Prionocycloceras than to other, earlier Peroniceratinae, and this is be-
lieved to be its origin. Both evolute and involute species occur in the fourth
division of the Zululand Coniacian, but there is insufficient evidence to allow in-
terpretation of this in terms of an evolutionary trend as has been demonstrated
in the Peroniceras — Zuluiceras series through the Middle and Upper Conia-
cian. Ishikariceras Matsumoto, 1965, with its evolute strongly ornamented inner
whorls with strong umbilical and ventral tubercles, a strong siphonal keel
flanked by weak grooves, and an almost smooth body chamber, appears to be a
distinct genus. Its precise horizon within the Coniacian is unknown. If, as
appears likely, the two known specimens are adult, it is a micromorph.

Neogauthiericeras Collignon, 1969, described from the Santonian—Cam-
panian boundary of Madagascar, is, by virtue of its stratigraphic isolation from
other Peroniceratinae, more likely to be a Gauthiericeras homoeomorph of texa-
nitine affinities analogous to Reginaites, a texanitine homoeomorph of Peroni-
ceras.

ACKNOWLEDGEMENTS

We thank Dr H. W. Ball, Dr M. K. Howarth and Mr D. Phillips of the
British Museum (Natural History), Dr C. W. Wright (Oxford), Prof. Dr
J. Wiedmann (Tubingen), the late General M. Collignon (Moirans) and Emeri-
tus Prof. T. Matsumoto (Kyushu) for advice and criticism. The assistance of the
technical staff of the South African Museum, Cape Town, and the Geological
Collections, University Museum, Oxford, is gratefully acknowledged. Mr
W. Wetzel (Tubingen) photographed Van Hoepen’s type specimens.

Financial aid from the South African Council for Scientific and Industrial
Research to Klinger, and the Trustees of the Sir Henry Strakosh Bequest and

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

the Royal Society and the Natural Environment Research Council to Kennedy is
gratefully acknowledged. Part of the research was undertaken by Klinger in
Tubingen during the tenure of a research Fellowship of the Alexander von
Humboldt Foundation.

Access to the late Dr E. C. N. van Hoepen’s material was provided through
the courtesy of the Director of the Geological Survey of South Africa, Pretoria.

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KAUFFMAN, E. G., CoBBAN, W. A. & Ercuer, D. L. 1978. Albian through Lower Coniacian
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“KENNEDY, W. J., WRIGHT, C. W. & KLINGER, H. C. 1983. Cretaceous faunas from Zululand
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Africa. The ammonite subfamily Texanitinae Collignon, 1948. Ann. S. Afr. Mus. 80:
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J. Fac. Sci. Tokyo Univ. 1 (6): 181-212

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

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36: 59-133.

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ZiTTEL, K. A. von. 1884. Handbuch der Paldontologie. 1 Abt., II Band, Lief. III. Cephalopoda.
Munich & Leipzig: Oldenbourg.

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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) eae Gould, 1845: 37.
Leda plicifera A. Adams, 50!
Laeda bicuspidata Hanley, igs): 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861:
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HERBERT CHRISTIAN KLINGER
&
WILLIAM JAMES KENNEDY

CRETACEOUS FAUNAS FROM
ZULULAND AND NATAL, SOUTH AFRICA
THE AMMONITE SUBFAMILY
PERONICERATINAE HYATT, 1900

AE 92 PART 4 - FEBRUARY 1984
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

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

FISCHER, P.-H., DuvAL, M. & RarFy, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51. :
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.

Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. :
(continued inside back cover)

PNINALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 92 Band
February 1984 Februarie
Part] 4 Deel

MORPHOLOGICAL AND BIOLOGICAL NOTES
ON SOME SOUTH AFRICAN ARTHROPODS
ASSOCIATED WITH DECAYING
ORGANIC MATTER

PART 2
THE PREDATORY FAMILIES CARABIDAE,
HYDROPHILIDAE, HISTERIDAE,
STAPHYLINIDAE AND
SILPHIDAE (COLEOPTERA)
By
A. J. PRINS

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

MORPHOLOGICAL AND BIOLOGICAL NOTES ON
SOME SOUTH AFRICAN ARTHROPODS
ASSOCIATED WITH DECAYING ORGANIC MATTER

PART 2

THE PREDATORY FAMILIES CARABIDAE,
HYDROPHILIDAE, HISTERIDAE, STAPHYLINIDAE AND
SILPHIDAE (COLEOPTERA)

By
A. J. PRINS
South African Museum, Cape Town

(With 16 figures)

[MS accepted 16 June 1983]

ABSTRACT
The morphology of the immature stages of fourteen species of beetles belonging to the
families Carabidae, Hydrophilidae, Histeridae, Staphylinidae, and Silphidae, as collected along
a narrow strip between Mossel Bay and Elands Bay and in a small area around Laingsburg and
Tulbagh, is illustrated. Available data on their biology and ecology are given and their associa-
tion with decaying organic matter is noted. These insects play an important part in the preda-
tion of the arthropods attracted to carcasses and cadavers and other decaying organic matter.

CONTENTS

PAGE

ATE GNU CULO eet eke Seer eae alah oh a eee a Le ee ee ee 296
Morphologicalland biologicalnotes=- >... 242. - +4... 4.2400 eee ee 298
amMilWAG@akaDiGae A ti cexc ec ciel eink ons. 4 One uta ake COR RC 298
IbaermOStenuUs Gormplanatus Deleanias | ee ee ae eee 300
mi lWalycrophilGacy i. <j. te. .le en gd ees Po Se se Mens sere 304
CErGy ONNNanitiMmusIKIMSCh. =.) «2. eee a ee 305
Sphacnadwum cajfrum Laporte & Castelnau .4)-- 7255-45-45. 4- 308
FMI BEMNS GeTAC ACs yen a 2 Oe, Gon Gia oe nel Bae areata ie teria 310
Nap uinussbicolor(HaWneciUS) eb yaad cea a eee S13
Carcinops minutia Ranracus)| =.) seers eeae oe eee eee 316
ATM RS CAP IMNVHIMIGAC. aes. cin ae Pal Sess os ad Ges cls eee 318
VM COCIAUGSS Diet atone Waster Ae cat Heo eye a aide erly he ep 321
Omaliunvoarenarium: (Bemmhauen) eee ene eee ee 324
Oxvyiclus.cafier: (ETICHSOM) aos. see a fly are eee eee Se B27,
iloniwus natalensis Boheman wane) - sense eae soe oe oe 330
PAO MIAUS: SPD eMnetee ley. & koe ee Meee eae See tne Oho eee 334
SEGDIMVLUTLUS SSD ra iy erate: adhe loe jee! een Ie Cees ens ee ost lane tae 387
KOMI TLS S OC S oe SOL A ee HT ee ee ee et aoe 341
AEM BOUIN AS ew es yee ee pede es le Meet wih eae hoe MacNee. 345
Sil PNG NICANS PA hICMUShs ow. tetas: See eet ae ceo et a eee ee 345

Si phaipunciilata Olivicte.s = ame teste bes cee ee 350
PNCKNOWICASCIMEM (Sacer in. ave ose he eee ad 8 thd nev ct ven saesbtuo Ste atne senaye 3S;
IRCHETEDICES: 5 5, is anette Mae estate aac mele A ERR ai bn Losi inte 2 Ae 553
PND TC VIAL ONS Mia een cine Se CIT LS 5 Neu it den tc aaa egies er oe 354

Ug)

Ann. S. Afr. Mus. 92 (4), 1984: 295-356, 16 figs.

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

In the first part of this series of papers some specimens belonging to the
myriapods, arachnids, crustaceans, and hemimetabolous hexapods are dealt
with; in this second part the immature stages of the first group of the holometa-
bolous families are considered.

This group comprises one species of the adephagous Coleoptera and thir-
teen species of the polyphagous series Staphyliniformia, which includes the
superfamilies Hydrophiloidea, Histeroidea, and Staphylinoidea.

The surveys covered the beach, the dune system and the adjacent inland
area along a narrow strip of the southern and western coastal regions of South
Africa; a small strip between Laingsburg and Beaufort West, as well as between
Worcester and Tulbagh, was also included.

Nearly half of all known insects are beetles and, although they are of such
diverse form and habits and include some of the largest and also some of the
smallest of insects, only a very few of them have developed a tendency to social-
ity. Of the some 150 known families only about nine show some signs of sub-
social behaviour and these include the Staphylinidae, Silphidae, Hydrophilidae,
and also the Scarabaeidae. However, none of the South African species or, at
least, those that have been collected during the surveys, can be regarded as sub-
social, except perhaps hydrophilids belonging to the genus Spercheus; some may
well be described as gregarious as in the case of Harpalus capicola Dejean and
H. agilis Péringuey (Carabidae); Gonocephalum arenarium (Fabricius), G. sim-
plex (Fabricius), Zophosis boei Solier, and Stenocara longipes (Olivier) (Tene-
brionidae), and various other species often collected in very large numbers
under semi-dry to dry cow-pats, such as the vegetable weevil, Listroderes costi-
rostris Schonherr (Curculionidae).

All the species discussed in this paper are geophiles and all are predacious
both in the adult and larval stage, except the hydrophilids, the adults of which
are coprophages. All are attracted to decaying organic matter in which other
arthropods are breeding, and therefore play an important part in reducing
the number of coprophagous species. Nevertheless, they will feed on the
colloids oozing from the decaying matter if circumstances necessitate this,
particularly the hydrophilids and silphids and, to a certain extent, the staphy-
linids.

The larvae of most species are campodeiform, and in the case of the Sphae-
ridiinae the legs are reduced and they resemble the scarabaeoid forms.

As the amount of food available plays an important part in the develop-
ment of these insects, and as this food source is often limited, dwarfed individ-
uals are common in nature, sometimes differing markedly from the normal
forms. This is clearly demonstrated by some specimens of an unidentified species
of Philonthus (Fig. 1B) collected in various localities during the survey and in-
cluded here in the discussion on the Staphylinidae. To the naked eye the adults
appeared to be two different species; however, there were no morphological

SOUTH AFRICAN ARTHROPODS 297

Fig. 1. Families Carabidae, Staphylinidae.
A. Amblystomus capenis (adult). B. Philonthus sp. (adult). C. Oxytelus caffer (adult).
D. Liamegalonychus sp. (adult).

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

differences when the larvae were examined under the microscope and it is there-
fore assumed that they are variants of the same species.

The morphological terms used in this paper in the descriptions of the imma-
ture stages are those of Richmond (1920), Béving & Craighead (1931), Hafez
(1939a—d), Dorsey (1940), Van Emden (1942), and Kasule (1968).

MORPHOLOGICAL AND BIOLOGICAL NOTES
Family Carabidae

Flat to oval, usually dark-coloured beetles with the thorax generally
narrower than the elytra, except in certain groups such as the tribe Scaritini.
Head narrower than pronotum and tarsi five-segmented. Antennae eleven-
segmented. Wings well developed or absent.

The ground beetles can be divided into species that live near or in water, on
the ground and in trees; of these groups the geophiles are the most abundant,
the vast majority of which belongs to the subfamilies Carabinae and Harpalinae.
The eggs of the different species are laid either in the soil or in little mud pack-
ets and there are three larval stages. Pupation usually occurs in the soil in a
pupal cell.

Certain species are commonly found under kelp strings and flakes just
above the highwater mark, such as the black Acanthoscelis ruficornis (Fabricius)
(Fig. 2B), which was on various occasions excavated from the burrows of the
isopod Tylos capensis Krauss and was also observed to feed on beach-fleas (Or-
chestia and Talorchestia spp.), fly maggots, and even on the ladybird Lioadalia
flavomaculata de Geer, which is often found on the kelp. It was now and then
collected from dead sea-birds. When feeding, the prey is held between the fore-
legs, and in the case of maggots almost everything is consumed except the
mouth-hooks and a few pieces of the skin. Its food included the smaller, pale,
straw-coloured cicindelid, Platychila pallida (Fabricius) (Fig. 2G), which oc-
curred under similar conditions. This tiger beetle seems to favour the more arid
parts of the north-western Cape Province.

Scarites rugosus Wiedemann (Fig. 2A) closely resembles A. ruficornis, but
is much larger (30 mm) and has a wider distribution in South Africa, both along
the coast and inland. Péringuey’s (1896) note that it is generally found on the
sea-shore, or at no great distance inland, is therefore not entirely correct. It was
often found in the sand under shore plants; otherwise its habits are unknown.

Various other carabids have been collected in decaying and dry kelp, most-
ly of the flake and string types, of which Harpalodes xanthorhaphus (Wiede-
mann), Harpalus fuscoaeneus Dejean, Tetragonoderus immaculatus LaFerté,
Agonum rufipes (Dejean), and Liamegalonychus spp. (Fig. 1D) were the most
common. The first two species are widely distributed in South Africa and, to-
gether with Liamegalonychus (which also feeds on the flesh-fly Sarcophaga mari-
tima Engel) have often been collected under cow-pats. Hystrichopus vigilans
(Sturm), which resembles Liamegalonychus spp. and sometimes occurs in ma-
nure in the western Cape Province, has been found by the author to cause mild

SOUTH AFRICAN ARTHROPODS

299

G2asIa

e
a

Fig. 2. Families Carabidae, Cicindelidae, Silphidae.

A. Scarites rugosus (adult). B. Acanthoscelis ruficornis (adult). C. Geobaenus lateralis

(adult). D. Harpalus capicola (adult). E. Tetragonoderus thunbergi (adult). F. Silpha mutila-
ta (adult). G. Platychila pallida (adult). H. Thachys capicola (adult).

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

irritation of the skin in hypersensitive people when carelessly handled. All these
species are black, except the first named, which is brown. Agonum rufipes has
light-reddish legs.

Tetragonoderus immaculatus is rather similar to T. thunbergi (Crotch)
(Fig. 2E), which occurred together with Harpalus agilis Péringuey and
H. fuscoaeneus in soils rich in organic matter such as fish-meal and compost, but
it is somewhat bigger and lacks the two pale-yellowish patches on the apical
parts of the elytra.

At least twelve carabids were collected under dry cow-pats, the largest be-
ing the previously mentioned Liamegalonychus sp. (14-15 mm). Harpalomor-
phus capicola Péringuey, Harpalus capicola Dejean (Fig. 2D), H. agilis Périn-
guey, H. fulvicornis Thunberg, H. fuscoaeneus Dejean, and Cratognathus
mandibularis Dejean are all medium-sized species (7-12 mm) and often oc-
- curred in large numbers under pats along the west coast, particularly during the
winter. Harpalomorphus capicola, which was not as abundant, was collected
only in the north-western parts. Harpalodes xanthorhaphus, common under
kelp, and Geobaenus lateralis Dejean (Fig. 2C) are both widely distributed and
occurred under dry pats in fair numbers in almost all the areas surveyed.

Some carabids are generally attracted to fresh dung, particularly Amblysto-
mus capensis (Motschulsky) (Fig. 1A), but only very small numbers were ob-
served. On the other hand the shiny, reddish-black Tachys capicola Péringuey
(Fig. 2H), which is about the same size as A. capensis, is a common species in
leaf litter in the Cape Peninsula and also occurs in soil rich in compost.

A small, oval, pale straw-coloured carabid, Omophron capicola Chaudoir
(about 6,9 mm long), is often encountered in sandy areas along the western
parts of the southern coastal region. It resembles the tenebrionid, Pachyphaleria
capensis (Castelnau), but can easily be distinguished by the metallic green
patches on the head, median area of the prothorax and elytra, and by the sickle-
shaped mandibles, the large eyes and longer antennae. It was observed to stalk
Machilinus sp. (Order Microcoryphia) common on the debris in these regions,
and probably feeds on this machilid.

Laemostenus complanatus Dejean

DESCRIPTION
Adult (Fig. 3J).

Previously described by Péringuey (1896) and Hinton (1945). Black to dark
brownish black in colour, head and thorax fairly shiny, elytra duller, with some-
what silky sheen. Antennae, palpi and tarsi reddish brown; first three antennal
segments devoid of pubescence, segments 4-11 pubescent. Elytral striae fine but
clearly indicated.

This is a cosmopolitan species and, according to Hinton (1945), apparently
a native of Europe or north Africa. According to collection data found only
around Cape Town, on the Cape Flats, Robben Island, and near Stellenbosch.

SOUTH AFRICAN ARTHROPODS 301

In Britain it appears in granaries and in glue and chemical works (Hinton 1945)
and, according to Pescott & Miller (1937), the adults were observed to prey on
the larvae of the codling-moth, Cydia pomonella (Linnaeus) in Australia
Around Cape Town it is commonly attracted to decaying carcasses and compost
heaps.

Larva (Fig. 3A-B)

A general description of carabid larvae is given by Van Emden (1942).
Elongate, subcylindrical and 16,6-17 mm long when full grown.

Body creamy white, with tergites and sternites pale piceous brown. Head
directed slightly upwards. Pronotal tergite largest and also more strongly sclero-
tized, all tergites divided by a thin, pale, longitudinal median line. Mesothoracic
spiracle largest and oval; abdominal spiracles (of which one on first segment is
larger than rest) annular (Fig. 3P). Two lateral abdominal sclerites present, of
which upper one bears five and lower one six setae, except in eighth segment
where both have five setae in all specimens examined. Ninth segment with only
one lateral sclerite bearing only two setae, and also with two long cerci, each
with nine long setae. Tenth segment tubular and functioning as a proleg. Sternal
region (Fig. 3L) of abdominal segments 1-7 each with single large anterior ven-
trite bearing eight to ten setae and four posterior postventrites, of which inner
pair bears two and outer pair four setae. One pair of minute praeventrites also
present. Sterna 8 and 9 with sclerites fused into a single plate, which is divided
posteriorly on eighth segment. Sternal plate on eighth segment with thirteen to
fourteen setae, that on ninth segment with ten to twelve. Each tergum with
nineteen to twenty setae, except that of eighth segment, which has ten to twelve
only, and ninth segment, which has four, one of which is situated on base of
each cercus.

Legs (Fig. 3D) with some scattered setae on coxae, each trochanter with
two posterior and about four anterior spine-like setae, as well as two long ven-
tral setae. Femur with a double ventral row of four to five spine-like setae and
single short seta dorsally near apex. Tibia with about six spine-like setae around
apex. Tarsus slightly longer than tibia, bearing two spine-like apical setae dorsal-
ly and two simple, acute claws, which are only slightly shorter than tarsus. Com-
pare with Staphylinidae where legs are four-segmented (Figs 9E, 10F, 11E, 13F,
14B).

Young larvae collected during September resemble mature specimens in
form, shape, and number of setae present on head and thorax, but middle tooth-
like process in mandibles is much more acute, and (in the few specimens exam-
ined) third seta on cercus is branched (Fig. 30). Abdominal sterna are not fully
differentiated into various sclerites present in later instars.

Head capsule (Fig. 3C, F—G)
Dark reddish brown, finely reticulate, somewhat shiny. Frontal and epi-
cranial sutures distinct; neck absent. Cervical keels well indicated. Nasale

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Family Carabidae.
Laemostenus complanatus. A. Larva (dorsal view). B. Larva (left lateral view). C. Head
(dorsal view). D. Right mesothoracic leg (posterior view). EE. Mandibles (dorsal view).
F. Head (left lateral view). G. Head (ventral view). H. Labium (dorsal view). I. Right max-
illa (dorsal view). J. Adult. K. Last abdominal segment of pupa (dorsal view) to show depres-
sion. L. Second abdominal sternite. M. Pupa (ventral view) with beetle already fully devel-
oped. N. Egg. O. Third seta on cercus to show branch (highly magnified). P. First
abdominal segment to show spiracle.

SOUTH AFRICAN ARTHROPODS 303

somewhat prolonged, shallowly emarginate in the middle and also very slightly
so on each side; adnasalia slightly concave; two parietalia dorsally and laterally
with about eight setae each. Chaetotaxy as illustrated. Six ocelli present on each
side. Antennae four-segmented, fourth segment small, slightly shorter than
third; latter with sensorial appendages on exterior side; penultimate segment
with one short dorsal seta and two lateral ones; apical segment (fourth) also with
one dorsal seta and two lateral ones, and also bearing short setae at extreme
apex. Antennal base not very clearly demarcated and bearing single lateral seta.
Hind margin of head deeply excavated medially on dorsal side, ventrally only
slightly so. Gular suture distinct and with V-shaped notch nearly half-way
between anterior and posterior margins.

Mandibles (Fig. 3E)

Falcate, length slightly more than twice the width at base. Apex fairly
acute, colour dark reddish brown, almost black in old specimens. Retinaculum
present; subbasal penicillus represented by one or few setae in most specimens
examined. Exterior lateral side with depression, bearing single long seta.

Maxillae (Fig. 31)

With large almost rectangular stipes. Ventrally smooth and with about six
setae, of which one is very long, on external margin; dorsally beset with numer-
ous setae on mesal side. Outer lobe of maxilla palpiform and two-segmented,
with single ventral seta on first segment. Inner lobe present as small conical pro-
cess, with single long style-like seta at apex. Cardo ventrally with a single seta.
Maxillary palpus three-segmented, with apical segment small, about half the
length of first. Palpifer segment-shaped and with single ventral seta. Palpi de-
void of setae.

Labium (Fig. 3H)

Trapezoidal, with base narrower than apex. Ventrally divided by shallow
longitudinal furrow into two lobes, each bearing single seta apically. Ligula pro-
longed in front and bisetose. Lateral margin of labium sclerotized and bearing
numerous setae. Palpiger with single seta. Labium basally covered with long,
fine hairs. Labial palp two-segmented, with apical segment small, about half the
length of first.

Pupa (Fig. 3M)

Pupa about 16,6 mm long. Almost pure white soon after pupation, but
mandibles gradually become red; 3—4 days before beetle emerges, black eyes be-
gin to show through integument. Head and thorax with some scattered brown
setae dorsally. Abdominal terga with about nine long brownish-black setae on
each side; laterally also with two setae, except the last two or three terga, which
bear only few setae. Developing genital capsule represented by two conical tu-
bercles. Terminal abdominal tergum with almost square depression, the lateral
sides of which bear four to six setae (Fig. 3K).

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

BIOLOGY

Both adults and larvae are predacious and feed on various insects in the
soil; those collected from the carcasses fed in the laboratory mainly on fly larvae
and the grubs of skin-and-hide beetles. The maggots and pupae of the cheese
skipper, Piophila megastigmata McAlpine, also seemed to be a favoured source
of nourishment, as well as the caterpillars of tineids.

Beetles with fully developed eggs were collected under carcasses during
April and May. These eggs (Fig. 3N) measure 1,6 mm X 0,88 mm and are oval
shaped, almost pure white, with the surface microscopically wrinkled or sha-
greened and somewhat dull. Some beetles were found hibernating under the
bark of trees and under stones; those collected during September survived for at
least 6 months and bred larvae during the early winter. Full-grown larvae ap-
peared from September to October and pupae were found from October to
- November. There was only one generation a year during the survey period;
however, overlapping occurs and young larvae measuring 4,5 mm were observed
during spring. The fully mature larvae construct clay cells in which they remain
dormant for a short period (3-20 days) after which they pupate. Pupal stages
varied from 11 to 13 days in the laboratory during the late spring and early sum-
mer (21-23 °C).

Newly emerged beetles are pale whitish brown, but after about a day the
head and thorax become blackish; some 3 days later they usually assume their
normal black colour.

Family Hydrophilidae

Small to medium-sized oval beetles, the dorsum smooth and convex, and
the head prominent. Maxillary palpi fairly long and antennae seven to ten-
segmented. Tarsal formula 5—5—5 or 5—4—4. Wings well developed.

Some seventy species have been described from southern Africa, most of
which are aquatic or at least semi-aquatic; those of the subfamily Sphaeridiinae
are terrestrial, restricted to wet or damp places and breed in dung and other de-
caying matter; larval breathing in this case is pseudometapneustic. Very little is
known about the habits of one genus, Coelostoma; according to Boving & Hen-
riksen (1938) the presence of suckers on the underside of the abdomen of the
larvae may indicate that they are adapted for moving over rough or hard sur-
faces such as rotten leaves, etc. Adults of Coelostoma punctulatum (Klug) were
collected on the green algae growing in stagnant pools in the vicinity of Laings-
burg.

At least six species of water scavenger-beetles belonging to the above-
mentioned subfamily are common in fresh cow-pats in the areas surveyed. When
the dung starts to dry out, however, some of these beetles, together with some
of the histerids and staphylinids, which are attracted, generally leave the pats,
except those that prefer the drier media. The mandibles of the adult beetles are
much better sclerotized than in the coprophagous Aphodius species (Scarabaei-
dae) and they are thus adapted to feed on much drier and harder dung particles,

SOUTH AFRICAN ARTHROPODS 305

and predation may not be entirely excluded. Most of these beetles therefore re-
main in the dung much longer than the smaller scarabs and are often found in
fairly dry pats.

Cercyon maritimus Knisch was the most common species found in dung. In
Mamre (south-western Cape) quite a large number of the rather small Palaearc-
tic and Nearctic hydrophilid Cercyon pygmaeus (Illiger) occurred in association
with C. maritimus during the early spring, but its larvae could not be traced. It
was fairly abundant along the south coast during the late summer and autumn in
both semi-fresh and almost dry cow-dung.

Sphaeridium caffrum Laporte & Castelnau appeared in large numbers dur-
ing the winter and spring in the western parts of the Cape and was very numer-
ous during the late summer when larvae and pupae were present in fresh dung in
almost all areas. The related S. guinquemaculatum Fabricius (length about
3 mm), which is similar to, but smaller than, S. caffrum occurred during the
summer in Montagu. It is widespread, not only in the Cape Province but also in
other parts of the world such as Sri Lanka, southern Asia, China, and Taiwan
(Knisch 19246). Sphaeridium caffrum, however, is known only from the Sub-
saharan and Malagasy regions.

Pachysternum capense (Mulsant), on the other hand, which is also small
(2,3-2,5 mm long), occurred in fair numbers together with Sphaeridium quin-
quemaculatum in Montagu and can be distinguished from the latter by having in-
conspicuous longitudinal striae on the elytra. It was common in the Karoo and
along the west coast, often in association with S. caffrum. In the Sandveld an
unidentified species (2,3-3,1 mm long), with blackish head and pronotum and
pale straw-coloured elytra, appeared with P. capense in the same cow-pats.

Cercyon maritimus and C. gigas d’Orchymont usually also breed in decay-
ing kelp and may even be found in the decaying carcasses of marine animals and
birds. According to collection data, the latter species is endemic to the Cape
Province, and is mainly dark brown with narrow, longitudinal blackish stripes
over its elytra.

A third, peculiar, small hydrophilid Spercheus cerisyi Guerin-Méneville
(about 4 mm long and pale brownish grey in colour) appeared to be common on
kelp during the winter and spring in estuaries around the Cape Peninsula. It was
very often associated with the chrysomelid Monolepta bioculata (Fabricius) (of
about the same size), which has four large, oval white spots surrounded by black
margins on the dorsum of its orange body. Both beetles were found to feed on
the fluids oozing from the decaying kelp. This hydrophilid (subfamily Sperchei-
nae) is apparently a water-living species.

Cercyon maritimus Knisch
DESCRIPTION
Adult (Fig. 4A)

Previously described by Knisch (1924a). Small, rather variable beetles,
varying in colour from brown to almost pitch black; in some cases elytra brownish,

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

marked with black. Each elytron with about nine shallow punctured striae and
densely pitted all over, the pits small and as wide apart as their diameter.
Beetles vary from 3,7 to 4,6 mm in length.

Widely distributed along the Cape coasts.

Larva (Fig. 4B)

Resembles larva of histerids, but easily distinguished by the last abdominal
segment or stigmatic atrium (Fig. 4M), which consists of the tergal plate of the
eighth segment, the median and lateral lobes of the ninth segment (lateral lobes
also known as acrocerci), the true cerci or mesocerci, each with a long filament
at apex, and the procerci, which are actually processes of the eighth pleurites.
All spiracles including those of mesothorax of equal size and of biforous type;
seven pairs laterally on abdomen, eighth pair opening in atrium. Pro-, meso- and
~ metathoracic shields well developed, though small on last two thoracic seg-
ments. Full-grown larvae measuring 10-11 mm in length and dirty to creamy
white in colour, except head, thoracic shields and eighth abdominal plates,
which are reddish brown and somewhat shiny. Body integument leathery, tough
and covered with microscopic spinules (Fig. 4N). Legs present, with femur and
tibia distinguishable although minute (contradicts Richmond’s (1920) statement
that legs are entirely absent in Cercyon).

Head capsule (Fig. 4C—D)

Oval, somewhat upwardly directed in most specimens seen, with small ocel-
lus on each side. Frontal sutures not visible. Antennae three-segmented, with
first segment about twice as long as the second; latter with small, apparently
two-segmented appendage; third segment smaller than the second and with
some sensory pegs apically. Postgenae ventrally separated by gular suture, which
is Y-shaped and delimits a pregular area anteriorly between its arms. Arrow-
shaped tentorial pit posteriorly, almost in centre. Chaetotaxy as illustrated.

Mandibles (Fig. 4H, L)

About half as long as cranium and nearly one-third longer than wide.
Broad basally, cutting edge of left mandible without teeth, that of right man-
dible with single short tooth (ti) about half-way between apex and molar area.
Teeth very similar to those of C. quisquilius (Linnaeus) (Hafez 1939d). Molar
area thin and almost trenchant in both mandibles, forming a ridge rather than
teeth; that of left mandible somewhat broader than that on right. Penicilli not
present in the specimens examined. Ventral side of left mandible with longitudi-
nal, curved median groove visible through dorsal integument for reception of
right mandible.

Maxillae (Fig. 41I-J)

With broad cardo and stipes, nearly twice longer than wide and with more
or less shiny, oval area dorsolaterally furnished with fine setae; mesal margin
bearing eight to nine short setae. Fairly long seta also present apically on lateral

SOUTH AFRICAN ARTHROPODS 307

SS (jo EN
= a

See a

Fig. 4. Family Hydrophilidae.
A-N. Cercyon maritimus. A. Adult. B. Larva (dorsal view). C. Head (dorsal view).
D. Head (ventral view). E. Pupa (left lateral view). F. Pupa (ventral view). G. First three
segments of larva (left lateral view). H. Left mandible (dorsal view). I. Left maxilla (dorsal
view). J. Left maxilla (ventral view). K. Labium (dorsal view). L. Right mandible (dorsal
view). M. Atrium, straight from behind. N. Integument showing spinules (highly magnified).
O-X. Sphaeridium caffrum. O. Atrium, straight from behind. P. First three segments of larva
(left lateral view). Q. Atrium (dorsal view). R. Adult. S. Labium (dorsal view). T. Left
mandible (dorsal view). U. Left maxilla (dorsal view). V. Left maxilla (ventral view).
W. Labium (ventral view). X. Right mandible (dorsal view).

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

side. Ventrally without setae. Maxillary palp three-segmented, with segments
tapering towards the apex and of about equal length, the second segment bear-
ing long seta. Apical segment with sensory pegs at apex. Palpifer segment-like
and bearing two long setae ventrally; also two to three shorter setae near apical
border on mesal side, probably remnant of galea. Short spine-like seta also pres-
ent basally on mesal margin.

Labium (Fig. 4K)

Small, represented by prementum and two-segmented palpi. Mentum dor-
sally asymmetrically developed in all specimens seen, forming long, almost tri-
angular projection on left side, which bears some short setae on anterior part.
When viewed from below a small part of this projection is visible on mesal side
of left cardo-stipes.

_ Pupa (Fig. 4E-F)

Almost pure white after pupation, but soon changes to straw-yellow with
blackish eyes. Head with four tubercles, each bearing long, thick seta or stylus;
pronotum with about ten stylus-bearing tubercles on anterior and posterior mar-
gin as well as two near middle. Meso- and metathorax with stylus-bearing tuber-
cle on each side of median line. Abdomen on each side with three rows of sty-
lus-bearing tubercles, of which one row is situated laterally. Wings folded down
over ventral side of body. Developing genital capsule present as two small,
elongate cones situated close together. Fleshy cerci divergent in most specimens
studied, each with long filament attached to larval skin. Five pairs of spiracles
clearly visible, sixth pair small and inconspicuous. Body length of specimens
examined 3,7—-5,5 mm.

BIOLOGY

Adults and larvae were observed almost throughout the year, mostly in
kelp, but sometimes also in fresh cow-dung. As already stated, the larvae are
mainly predacious, but will feed on the colloids oozing from decaying kelp. Lar-
vae collected in this medium along the west coast during spring had a life-span
of at least 40 days before pupation occurred. Most pupae were found in the top
few centimetres of soil in oval earthen cells constructed by the larvae. Pupal
stages varied from 9 to 12 days during August and September, the newly
emerged beetles being yellowish white with some orange coloration on prono-
tum. Those observed assumed their normal dark colour after a few days to
about a week.

Sphaeridium caffrum Laporte & Castelnau
DESCRIPTION
Adult (Fig. 4R)

Length 4,3-5,5 mm. Oval, black, only slightly shiny, with fulvous or paler
spot near base of each elytron as well as fulvous or paler area on apical third of

SOUTH AFRICAN ARTHROPODS 309

elytra. Latter without longitudinal striae but densely punctate, the punctures be-
ing small and shallow. External border of elytra also pale fulvous. Legs brown-
ish, furnished with spines.

Widely distributed in South and east Africa and Madagascar (Knish 19245).
Specimens studied here were collected mostly in the western parts of the Cape
Province.

Larva

Very similar to Cercyon maritimus, but easily distinguished by absence of
extra tooth on right mandible and by stigmatic atrium, which bears two fleshy
projections on each side (the procerci and acrocerci) (Fig. 4Q). Eighth pair of
spiracles also opens into atrium. Mesocerci much shorter than in C. maritimus
and also furnished apically with filament or seta; median lobe broader than in
latter species (Fig. 4O), its hind margin almost quadridentate as in Sphaeridium
scarabaeoides (Linnaeus) (Hafez 1939b). Seven pairs of biforous spiracles pres-
ent laterally on abdomen. Legs much larger than in C. maritimus and, apart
from femur and tibia (with circle of about nine short setae around apex), third
or tarsal segment with one or two setae at tip also present (Fig. 4P). Integument
leathery and covered with minute spinules similar to those of C. maritimus; also
some short setae present. Full-grown larvae measure 8—9,5 mm and are of same
colour as those of the latter species.

Head capsule

Very similar to that of C. maritimus, including structure of gular suture;
also without frontal suture. With two to three ocelli on each side. Integument
fairly shiny and brown and cranium wider behind than in front. The antennae
three-segmented as in C. maritimus, but projection on second segment lacking;
second segment also about half the length of first; third segment smallest and
with short setae or sensory pegs at tip.

Mandibles (Fig. 4T, X)

Similar to those of C. maritimus, including the ventral groove on left man-
dible, but lacking tooth between apex and molar area on right mandible. Molar
areas trenchant. Two small setae usually present on exterior sides, one situated
near base.

Maxillae (Fig. 4U-V)

Very similar to those of C. maritimus, with cardo-stipes somewhat extended
on exterior side and with long seta and also with about four shorter setae ven-
trally. Mesal margin with numerous very fine setae, otherwise exactly as in
C. maritimus, including setae on palpifers. Second palpal segment with short
seta on both mesal and lateral margin; apical segment with short spine-like seta
near base on mesal side, and with sensory pegs at apex.

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

Labium (Fig. 4S, W)

Very similar to that of C. maritimus, but differs from it by longer labial pal-
pi, longer and more pointed projection on left side, which is furnished with fine
setae on anterior margin and by presence of a short ligula, which is about three-
quarters the length of first segment of palpi. Palpiger about as long as first seg-
ment of palpus.

Pupa

Very similar to that of C. maritimus, including apical cerci and styli. Num-
ber of styli on body and particularly those on head and pronotum also similar to
those of latter species, although much longer and thinner in all specimens exam-
ined. Cerci somewhat shorter than in C. maritimus. Most pupae about 5,4 mm
long. Colour of pupae at first more or less creamy white, but eyes become
brown after a few days. Whole pupa changes to darker colour just before beetle
emerges.

BIOLOGY

Large numbers of beetles were collected in fresh and semi-fresh cow-pats in
the western parts of the Cape during the summer and autumn and larvae were
observed during the early part of the summer, producing pupae during Decem-
ber and January. The larvae pupate in small cells constructed in the drier parts
of the dung and, to judge from their intestinal contents, most of the larvae col-
lected in the pats must have been feeding mostly on the decaying material, par-
ticularly the liquids oozing from the wet particles. When crowding occurs they
become cannibalistic.

Family Histeridae

Very small to medium-sized, oval hard-bodied beetles, head deeply sunk
into prothorax, short legs and eleven-segmented antennae capable of being re-
tracted. Tarsal formula 5—5—5 or 5—5—4. Wings well developed; elytra usually not
covering last two abdominal segments.

Some 150 species of histerids have already been described from southern
Africa, most of which are predacious on coleopterous and dipterous larvae and
other smaller arthropods. According to Nuorteva (1970), who experimented
with histerids in Finland, there is a very high negative correlation between the
occurrence of blow-flies and histerids in fish carcasses. Bornemissza (1968) re-
ported up to 50 per cent kill of fly maggots in cow-pats by Pachylister chinensis
(Quensel) in Fiji. Some species, however, live in the burrows of small mammals
or in the nests of birds (Arnett 1963), particularly species of Saprinus, Gnathon-
cus, and Hister; others again have developed a myrmecophilous or termitophi-
lous life, such as species of Monoplius as well as Hister. Species of Teretrius,
which are cylindrical in form, live in the burrows of wood-boring beetles, where-
as the flat species of the genera Hololepta, Pachycraerus, and Paromalus are

SOUTH AFRICAN ARTHROPODS 311

Fig. 5. Family Histeridae.
A. Hister nomas. B. Pachylopus dispar. C. Saprinus cupreus. D. Rhypochares saprinoides.
E. Saprinus elegans. F. Saprinus nitidus. G. Gnathoncus rotundatus. H. Hypocacc..lus
specillum. 1. Hypocacculus harmonicus. J. Carcinops minuta.

S12 ANNALS OF THE SOUTH AFRICAN MUSEUM

found under the bark of plants. At least four species of Pachycraerus have been
collected in Natal under the bark of Euphorbia species.

One of the largest histerids found in South Africa is the black, almost
smooth Macrolister maximus (Olivier) about 25 mm long, widely distributed in
the Republic of South Africa, Botswana, Zimbabwe, and west Africa but it is
not common. During the recent surveys at least fifteen species have been found
in decaying matter, of which Saprinus bicolor (Fabricius), S$. cupreus Erichson,
and S. nitidus (Wiedemann) were the most abundant. Saprinus bicolor occurred
mostly in carcasses, both on the beach and inland, whereas S. cupreus visited
both carcasses and cow-dung. Saprinus nitidus was the most common species in
fresh dung and was followed by Hister nomas Erichson, H. contemptus Marseul,
Atholus conformis (Erichson), the small Hypocacculus harmonicus (Marseul),
and Abraeus setulosus Fahraeus.

Saprinus nitidus (Fig. 5F) is a rather small beetle (2,8-4,2 mm long), shiny
black with a fairly large, somewhat diffuse reddish spot as well as two short
oblique striae on each elytron. Saprinus cupreus (about 3,3 mm long) (Fig. 5C),
is a shiny black beetle about the same size as S. nitidus (Fig. 5F), but lacks the
reddish spots on the elytra. Hypocacculus harmonicus (Fig. 51) is also shiny
black but smaller than Saprinus nitidus and is only 1,2-1,4 mm long. Abraeus
setulosus (1,2-1,8 mm long) is almost dull, densely punctate, and characterized
by the presence of short club-shaped hairs. These beetles are widely distributed
in South Africa and even occur in semi-fresh and almost dry dung.

The shiny black Hister nomas (about 8 mm long) (Fig. 5A), with about
seven striae on each elytron, was collected mostly in fresh dung in Heidelberg,
Cape Province, particularly during the spring; it was accompanied by the smaller
and similar Atholus conformis. Hister contemptus is about the same size as
H. nomas but has only about five striae on each elytron. None of these beetles
seemed to breed in the dung. Species belonging to the genera Hister and Atholus
mentioned here are easily recognized by the stria that runs parallel to the lateral
and anterior margins of the pronotum (Fig. 5A).

Three other species, Rhypochares saprinoides (Erichson) (Fig. 5D), Hypo-
cacculus specillum (Marseul) (Fig. 5H), and Tribalus capensis (Paykull) were
found to be prevalent in semi-dry and dry cow-pats and were never collected in
fresh dung. All these species are black. Rhypochares saprinoides (about 4,5 mm
long), with indistinct striae on the elytra, and H. specillum (2,3 mm long), dull
black with five shiny patches on its body, are endemic to the Cape Province
according to collection data, and are common, but not numerous, in the western
parts that were surveyed. The widespread Tribalus capensis (about 2,9 mm long)
is slightly shiny and oval, the striae on the elytra being almost obsolete. Most of
these beetles were observed to feed on the small scarab larvae and on the larvae
of any smaller flies that happened to be present.

Soils rich in decaying organic matter, such as compost and fish-meal, at-
tracted the shiny greenish-black Saprinus elegans (Paykull) (Fig. 5E), whereas
fowl manure, particularly in fowl-pens of the closed type in which the larvae of

SOUTH AFRICAN ARTHROPODS B16

the lesser meal-worm, Alphitobius diaperinus (Panzer), and those of the skin-
and-hide beetle, Dermestes maculatus de Geer, were present, produced large
numbers of Carcinops minuta (Fahraeus) (Fig. 5J) and Guathoncus rotundatus
(Kugelann) (Fig. SG). These two species are common in the Cape Peninsula in
decaying carcasses when fly larvae develop, together with the previously men-
tioned Saprinus bicolor and S. cupreus. Both Carcinops minuta and Gnathoncus
rotundatus are rather small beetles, the latter more quadrate, with coarser punc-
turation than the first, the striae covering only a portion of the elytral surface.

In decaying kelp usually two species only of histerids were observed, and
these fed mainly on the immature stages of the flies that were present. Pachy-
lopus dispar Erichson (Fig. 5B), the larger of the two species (3,1-3,5 mm), is
dull black in colour without any sculpture on its body and was often found feed-
ing on small maggots in decaying red bait (Pyura stolonifera (Heller)). The other
species, Acritus lightfooti Lewis, is very small (1,4 mm), red-brown in colour and
its body is finely rugosopunctate. There is evidence, however, that this species
may also feed on Collembola, particularly the purplish Anurida maritima
(Guerin).

Saprinus bicolor (Fabricius)
DESCRIPTION
Adult (Fig. 6L)

Oval, shiny, 4-6 mm long. Pronotum coppery black. Elytra blackish green
to reddish black and finely rugosopunctate, with about five striae on each side
near base; central disc of elytra smooth and only shallowly punctate. Pronotum
punctate, with punctures larger on sides. Exposed tergites shiny and densely
punctate. Legs shiny, brownish black, with all leg segments flattened and tibiae
spined on dorsal side. All femora with grooves for receiving tibiae; grooves on
front femora situated on anterior side, those on other legs on posterior side.

Widespread in South Africa and Zimbabwe (Thérond 1956).

Larva (Fig. 6B)

Full-grown larva 12-12,5 mm long and dirty creamy white, with head and
pronotal shield yellowish brown; subcylindrical. First eight abdominal segments
each with four annulets dorsally and lateroventrally with fleshy swellings which,
in conjunction with small tenth segment, function as pseudopods. Cerci small,
bisegmented, each bearing two terminal setae. Body sparsely covered with golden
hairs; integument covered with conical spines (Fig. 6C) visible only under
high magnification. All spiracles almost of same size, oval and of biforous type;
twin opening clearly visible on ventral side. Mesothoracic spiracle situated ven-
trally above leg and tilted lengthways, so that long axis is in longitudinal posi-
tion, with opening pointing forward. Legs (Fig. 6D) small but well developed,
tarsungulus long, slightly shorter than tibia, which is nearly twice as long as fe-
mur. Both trochanter and femur with ventral seta and tarsungulus with small
seta ventrally near base.

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Family Histeridae.
Saprinus bicolor. A. Head (dorsal view). B. Larva (left lateral view). C. Part of integument
highly magnified to show spiracle and spines. D. Right leg (posterior view). E. Head (ventral
view). F. Left mandible (dorsal view). G. Left mandible (exterior lateral side). H. Right
mandible (ventral view). I. Left maxilla (dorsal view). J. Labium (dorsal view). K. Left
mandible (ventral view). L. Adult. M. Pupa (dorsal view). N. Pupa (ventral view).

SOUTH AFRICAN ARTHROPODS Sil)

Head capsule (Fig. 6A, E)

Slightly upwardly inclined, about as long as wide and about as wide behind
as in front; posterior margin deeply and angularly emarginate on dorsal side,
ventrally only slightly so; dorsally somewhat shiny, microscopically reticulate
and light reddish brown. No ocelli visible. Chaetotaxy as illustrated. Nasale ex-
tended and emarginate in middle, the two small teeth thus formed asymmetrical
in specimens seen; right tooth larger than the left one; sinuate on each side of
median extension and forming broad, rounded tooth on each lateral angle,
cephalad of the antennae. Frontal and coronal sutures obsolete, but represented
by some pale lines as shown in Figure 6A. Small pregula present ventrally be-
tween arms of Y-shaped gular suture, of which short stem ends in small poste-
rior tentorial pit at centre of head; postgenae fused posteriorly and gular suture
visible only as very shallow longitudinal impression. Antennae about as long as
the mandibles; three-segmented, with third segment small, and penultimate one
bearing sensory cone; all segments devoid of setae.

Mandibles (Fig. 6F-H, K)

Almost triangular, about half the length of cranium and about a third longer
than wide at base. Cutting edge of both mandibles entire, apex rounded and
with a small tooth (t; and t2) near molar area, which distally bears a single
rounded tooth (m; and m2); proximally with a penicillus. Lateral margin rounded
dorsoventrally, without dorsal carina and with small seta near base. Basal
portion cf mandible raised above the dorsal surface (Fig. 6G) for preartis to join
condyle.

Maxillae (Fig. 61)

Cardo and stipes united; latter dorsally with longitudinal excavation and
with three setae on exterior margin and eight to ten setae on mesal; ventrally
with only one seta. Apical border with six to eight dorsal setae. Palpifer seg-
ment-like, bearing one lateroventral and about four dorsal setae as well as small
conical galea on mesal side. Palpus three-segmented, about as long as cardo and
stipes and without setae. Galea with two small, palus-like setae.

Labium (Fig. 6J)

Labial stipes (prementum) long, with sides almost parallel and only two
ventral setae near apex. Dorsally with row of about ten fairly long setae on each
side of basal half and with two setae near apex. Labial palpi two-segmented,
with apical segment about twice as long as basal one; both without setae.

Pupa (Fig. 6M-N)

Robust, measuring nearly 6 mm; white in colour with reddish head and
mandibles. Body dorsally sparsely covered with short setae. Terminal abdominal
segment with a crescent-shaped appendage at apex; dorsum of this segment al-
most triangular and with longitudinal carina; developing genitalia visible as two
swellings on ventral side.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

BIOLOGY

Eggs of this species were found in the soil near decaying organic material
containing developing fly maggots. The larvae feed mainly on small insects and
when mature, construct a soft, silk-lined earthen cell in the soil in which they
pupate. Diameter of most of the cells examined was 9,8 mm. The pupal stage
lasted for about 13 days during April. The newly emerged beetle is dark brown
on the head, with brownish-yellow pronotum. About two-thirds of the pronotal
disc in most specimens examined is of a dark brown, the elytra and abdomen
pale yellow, with brownish-yellow legs. After a day or two they assume their
normal coloration.

Carcinops minuta (Fahraeus)
_ DESCRIPTION
Adult (Fig. 5J)

Rather small, oval beetle, about 2,5 mm long, reddish black in colour, head
black. Fairly shiny all over. Head and pronotum densely and minutely punc-
tured; elytra with seven striae each, striae almost reaching hind margin; spaces
between striae also have some fine punctures.

Collection records come from Cape Town and surrounding suburbs, and
from Paarl. Also present in Natal, east Africa and Zaire (Thérond 1956).

Larva

Similar to that of Saprinus bicolor but smaller, about 6,3 mm long when full
grown, and more elongate. Cerci bisegmented and much longer than in latter
species, apical segment with two long terminal setae (Fig. 7B). Legs much longer
in relation to body than in S. bicolor (Fig. 7A). Integument of specimens
examined without conical spines, but with rounded tubercles in pseudopodal
areas (Fig. 71).

Head capsule (Fig. 7C—D)

Slightly longer than wide, measuring 0,44 mm wide and 0,50 mm long in
most specimens seen, about as wide behind as in front, the sides almost parallel,
only slightly convex and somewhat sinuate behind antennae. Almost smooth and
shiny, reddish brown to dark brown depending on degree of sclerotization; dor-
sally with some indistinct sulci as in drawing. Posterior margin dorsally deeply
and angularly emarginate, ventrally widely and shallowly so. No ocelli observed.
Chaetotaxy as in Figure 7C-D. Pregula very narrow. Nasale (Fig. 7E) in all
specimens seen with two asymmetrical lobes in middle, that on left bidentate,
right one stronger developed and tridendate. Characteristic of this species is the
oblique carina dorsally on posterior one-third of cranium, which is absent in
S. bicolor. Antennae slightly longer than mandibles, three-segmented; apical
segment short, slightly more than half as long as the second, with terminal sen-
sory pegs. First segment longest, about one-quarter longer than penultimate

SOUTH AFRICAN ARTHROPODS S07

f.2

i - ffn= =- o-

Fig. 7. Family Histeridae.
Carcinops minuta. A. Head and thoracic segments (left lateral view). B. Last two or three
abdominal segments (left lateral view). C. Head (dorsal view). D. Head (ventral view).
E. Nasale (highly magnified). F. Left mandible (dorsal view). G. Left maxilla (dorsal view).
H. Labium (dorsal view) showing lobe-like structures with brush of hairs. I. Part of integu-
ment highly magnified to show tubercles.

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

one, which bears sensory cone and sensory pegs. All segments devoid of setae in
the few specimens examined.

Mandibles (Fig. 7F)

Similar to those of S. bicolor, but more sickle-shaped, molar area much
smaller, tooth t2 (also ti-Fig. 6H) large and acute and situated in about the mid-
dle. Mandibles less than half the length of cranium, penicillus fairly long.

Maxillae (Fig. 7G)

Similar to those of S. bicolor, but united cardo and stipes much longer, dor-
sally with single row of about twelve long, fine hairs as well as two setae near
apex, inner one the longest. Ventrally with a long seta; also two short setae on
external side. Palpifer as in S. bicolor, with a ventral seta and conical galea,
-which is much more developed in this species and bears a long terminal seta.
Maxillary palpi three-segmented, apical segment about as long as first and sec-
ond taken together.

Labium (Fig. 7H)

Labial stipes very short, with two ventral and two dorsal setae, mentum
produced on each side in form of a blunt tooth with a terminal spine-like seta. In
this species there is dorsally a small lobe-like structure on each side of mentum,
bearing a brush of thick hairs. Labial palpi two-segmented, with apical segment
nearly twice as long as basal one, both without setae.

Family Staphylinidae

A very large family of elongate, sombre-coloured rove-beetles, with short
elytra exposing nearly always about half of the abdomen. They vary from very
small to fairly large (about 30 mm or more); antennae usually eleven-segmented
(sometimes nine- to ten-segmented), ocelli present in certain forms; hind wings
usually well-developed. Tarsal formula 4-5-5, 4—4-—4 or even 3-3-3 or 2-2-2.

The beetles abound wherever decaying organic matter accumulates, both
along the shore and inland. Many species (subfamily Aleocharinae) live as ter-
mitophiles and myrmecophiles, while some species of the genus Aleochara are
ectoparasitic on fly puparia, such as A. salsipotens Bernhauer (Fig. 8A), which
were observed to destroy the pupae of the kelp-flies Fucellia capensis (Schiner)
and Coelopa africana Malloch; however, this predation accounts for the destruc-
tion of only a very small percentage of the kelp-fly populations.

Some species, particularly those belonging to the subfamily Paederinae,
when crushed against the skin, produce a blistering fluid that may cause severe
irritation.

The larvae of staphylinids are campodeiform, active, predacious and
usually somewhat flattened or depressed. Legs fairly long, four-segmented with
a tarsungulus. Ocelli present or absent. Abdomen with ten segments.

SOUTH AFRICAN ARTHROPODS 319

Various rove-beetles have been observed in decaying kelp on the beaches,
particularly Cafius xantholoma (Gravenhorst), which is widespread in Europe
(Bernhauer & Schubert 1914) and regarded by Backlund (1945) as eucoenic.
Morley (1907) suggests that it may even be omnivorous and will feed on mam-
mal carcasses if necessary. During the present study it was found on carcasses of
marine mammals on a few occasions, but appeared to feed on dipterous maggots
under these circumstances.

Species of Aleochara, including the previously mentioned A. salsipotens
(Fig. 8A), are some of the most numerous of all the Staphylinidae associated
with kelp, together with species of Omalium, Xantholinus and Bledius, particu-
larly B. alutellus Bernhauer. Most of them were found all along the coast as well
as further inland in the survey area. A tiny, brown, Omalium-like Phyllodrepa
hessei Bernhauer, (about 1,9 mm long) occurred only along the western part of
the Cape Peninsula in kelp where the larger species were absent. The large
black Staphylinus hottentotus Nordman (Fig. 8C), which is distributed through-
out South Africa (Scheerpeltz 1973), was as numerous in kelp banks and flakes
as in semi-fresh cow-dung and other decaying organic matter such as carcasses of
marine and land animals.

The immature stages of Aleochara spp., Bledius spp., and Staphylinus hot-
tentotus were never found in kelp during the surveys and their breeding habits
are therefore unknown. However, the larvae and pupae of an unidentified Aleo-

Fig. 8. Family Staphylinidae.
A. Aleochara salsipotens. B. Leucoparyphus variipes. C. Staphylinus hottentotus.

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

chara sp. (Fig. 9) were observed in semi-fresh cow-dung near Elands Bay during
the winter, and those of Omalium arenarium (Bernhauer) (Fig. 10) in wrack
near Kommetijie.

Fresh cow-dung on the beach and in the interior attracted mainly rove-
beetles of the genera Aleochara, Philonthus, Xantholinus, and Leptacinus, all of
them generally feeding on the flies and their larvae present in the dung, particu-
larly the young larvae of the green dung-fly, Orthellia peronii (Robineau-Desvoi-
dy), and those of the scarabs. Oxytelus caffer (Erichson) (Fig. 1C) and an un-
identified species of both Aleochara and Philonthus (Fig. 1B), were the most
numerous. All are black, except the one Philonthus sp., which has brownish ely-
tra, and normally they all breed in the dung.

The Philonthus sp. (7,5-9 mm long) is fairly shiny, the Aleochara sp. (only
4,5-5 mm), on the other hand, is duller due to the presence of body hairs. All
_ the species occurred almost throughout the year and are widely distributed
throughout the whole area that was surveyed.

Aleochara bipustulata (Linnaeus) (3,2-3,7 mm long), another widespread
species in the Cape Province as well as in the Palaearctic and Nearctic regions,
occurred in cow-pats together with various other Aleochara and Leptacinus spp.
and seems to have the same parasitic habits as A. salsipotens.

An unidentified Philonthus sp., very similar to the black species previously
mentioned but more polished, particularly on the elytra, was found mainly in
decaying birds in the vicinity of Cape Town. Occasionally the medium-sized,
shiny, black Philonthus natalensis Boheman (Fig. 11J), occurred in fresh cow-
pats near Heidelberg. Its large, oval, white eggs together with immature and
mature larvae, which are characterized by the very long urogomphi, were col-
lected occasionally during the late spring.

A black Xantholinus sp., (6,6-7,9 mm long), was fairly common in fresh
dung throughout the region surveyed and also appeared in decaying kelp along
the west coast. It is probably the same species that was found to visit decaying
carcasses.

The peculiar Leucoparyphus variipes Fauvel (Fig. 8B), a small, shiny, black
staphylinid (about 3,7 mm long), is common all along the south coast and was
mostly attracted to cow-pats in which large numbers of the smaller flies belong-
ing to the families Mycetophilidae, Sciaridae, and Sepsidae were breeding. It is
very agile and distinctly resembles members of the order Thysanura.

When cow-pats have dried out to a certain extent, some of the species disap-
pear to be replaced by others such as Aleochara maura Sachse, Staphylinus hot-
tentotus, Philonthus caffer Boheman, and an elongate, yellowish-brown Astenus
sp. Oxytelus caffer, Aleochara bipustulata, and some of the Xantholinus spp.
were also observed to remain in fairly dry dung for long periods and were even
found to continue to breed in pats as long as there was some moisture available.

A fairly large, agile and aggressive, unidentified Staphylinus sp. (Fig. 13M),
chequered with golden yellow and brown, was found to breed in fairly dry to
semi-fresh pats, both along the south and west coasts. When a cow-pat is picked

SOUTH AFRICAN ARTHROPODS 321

up, the large brownish larvae of this species are often found with open jaws and
trying to escape into the soil. In the laboratory these larvae fed on various in-
sects, but did not accept isopods.

The staphylinids that visited carcasses during the surveys included Aleo-
chara salsipotens (mostly on the beach), A. lateripennis Bernhauer, Oxytelus caf-
fer, O. grandis (Eppelsheim), Staphylinus hottentotus and at least one undeter-
mined species each of Aleochara, Oxytelus, Philonthus, and Xantholinus (Fig.
14A). Oxytelus grandis is a shiny brown species that is widely distributed, occur-
ring in Natal, Zimbabwe as well as in west Africa; all the other rove-beetles are
black, except Aleochara lateripennis and the Xantholinus sp., which have light-
reddish and brownish elytra respectively.

According to Kaufmann (1937) Oxytelus spp. and genera of the tribe Oma-
lini are absent during the warm weather and prefer the cold. However, in this
study Oxytelus caffer was found throughout the year in fairly large numbers,
both in cow-dung and carrion, and immature stages of this species and those of
O. grandis were collected in bird carcasses along the west coast throughout the
summer and autumn.

Aleochara sp.
DESCRIPTION
Adult (Fig. 9A)

Black and shiny, abundantly covered with medium-long, brownish, ad-
pressed setae, particularly on head and thorax; abdomen more sparsely covered
and therefore more shiny. Hairs grow from small pits, which are oval and larger
on abdomen. Legs and antennae dark brownish. First three and apical anten-
nal segments longer than wide, others as wide as or wider than long. Length
4.8-5 mm.

Widely distributed, collected in cow-dung almost throughout the areas sur-
veyed.

Larva (Fig. 9B)

More robust than in Philonthus and Staphylinus spp., and more depressed
dorsoventrally. Colour bright yellow to cadmium yellow, with oval brownish
patch on each side of all body segments except pronotum. Mature specimens
4.6—-5 mm long; sparsely covered with medium-long setae, of which four charac-
teristically arranged medially near posterior border of the pro-, meso- and meta-
notum. Urogomphi (Fig. 9C) very small, apparently one-segmented, each with
one long apical seta and one or two preapical setae. Spiracles circular and those
on abdomen of same size and situated laterally; mesothoracic spiracles (Fig. 9K)
ventral in position, somewhat larger and without setae (there is, however, a seta
anterior to it). Legs (Fig. 9E) similar to those of other species, but with reduced
number of setae, lacking especially rows of spines on femur; tarsungulus with
one ventral and one dorsal spine. Integument dorsally covered with very fine, al-
most ultramicroscopic spicules; very few triangular denticles present and body
hairs mostly simple.

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

Head capsule (Fig. 9D, F-G)

Smooth and shiny, pale golden yellow in colour; slightly wider behind than
in front. Posterior margin slightly or widely emarginate, anterior margin straight
and without teeth. Frontal sutures very indistinct and in some specimens indi-
cated only by Y-shaped depression, but epicranial stem usually clearly demar-
cated. Setal pattern as illustrated. Neck absent. One ocellus present on each
side, in some specimens not pigmented and may easily be overlooked. Hind
margin deeply emarginate ventrally. Gula small, but clearly visible. Antennae
three-segmented; apical segment small with three long subapical setae of which
one is ventral in position, and with about four sensory pegs at apex; first and
second segments of about equal length, second segment with two small ventral
spines near apex and with sensory cone surrounded by about three long setae.
Labrum free, trapezoidal, anterior margin slightly emarginate and bearing four
long and two short setae; also transverse row of six setae medially, two of which
are situated on oblique lateral face.

Mandibles (Fig. 9L, N)

Broad at base and tapering towards apex, thus appearing falcate; about half
the length of cranium. Cutting edge of both mandibles bidentate, subapical
tooth smaller than other. Molar area absent. Lateral margin fairly rounded
dorsoventrally and with two setae placed far ventrally in specimens examined.

Maxillae (Fig. 9H, J)

Definite division between galea and lacinia (visible only on ventral side)
and long seta on lateral margin. Lacinia with five spines and long seta on mesal
margin; maxilla dorsally without setae but ventrally with three long setae on
stipes and single long seta on cardo. Latter with Y-shaped suture. Maxillary pal-
pi appear four-segmented in specimens examined, first segment short and bear-
ing single seta on ventral side; second and apical segments of about equal
length; penultimate segment slightly longer and bearing a single lateral and ven-
tral seta.

Labium (Fig. 91, M)

Labial stipes (prementum) apically with two-segmented labial palpi; short,
broad, almost conical ligula present and two long subapical setae on ventral
side. Mentum fairly large and almost conical and bearing four setae on ventral
side. Boundary between mentum and prementum more strongly sclerotized and
ventrally bearing short seta on each side. Submentum as broad as mentum and
also bearing four setae ventrolaterally.

Pupa (Fig. 9O-P)

Pale lemon to cadmium yellow in colour, very pale soon after pupation;
length 3,9-4 mm. Head with about eighteen short, brownish setae; pronotum
with some short setae on front and hind margins and a parabolic transverse row

SOUTH AFRICAN ARTHROPODS 323

Fig. 9. Family Staphylinidae.
Aleochara sp. A. Adult. B. Larva (dorsal view). C. Abdominal segments 8-10 of mature
larva (dorsal view). D. Head (ventral view). E. Right mesothoracic leg. F. Head (dorsal
view). G. Head (left lateral view). H. Left maxilla (dorsal view). I. Labium (ventral view).
J. Left maxilla (ventral view). K. Left mesothoracic spiracle. L. Left mandible (dorsal
view). M. Labium (dorsal view). N. Right mandible (dorsal view). O. Abdominal apex of
pupa (ventral view). P. Pupa (ventral view). Q. Pupal cocoon.

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

of about eight over its middle. Each abdominal tergum with transverse row of
about nine short setae near posterior margin. Meso- and metanotum also with
few short setae. Abdomen with few short setae ventrally as illustrated; first two
sternites medially almost devoid of any setae. Developing genitalia (Fig. 9O)
visible as small cone-like structure with two setae flanked on each side by some-
what raised area with centrally placed seta.

BIOLOGY

Larvae of this species were found in association with larvae of Oxytelus caf-
fer in the western parts of the Cape during June; they were in semi-fresh cow-
pats, which also contained larvae of the scarab, Aphodius laetus Wiedemann.
Mature larvae construct pale-brownish oval cocoons in the dung (Fig. 9Q), using
their saliva and mixing it with dung particles. Pupation occurred after a short
_ prepupal stage; the pupae observed appeared in July and the beetles emerged in
August. Although the beetles were attracted to carcasses around Cape Town
they did not reproduce under these circumstances.

Omalium arenarium (Bernhauer)

DESCRIPTION

Adult (Fig. 100)

Light brownish in colour; abdomen somewhat darker, in some specimens
piceous; head and scutellum black. Sparsely punctate; elytra also rugulose,
shiny. Length 3,3-4,8 mm.

According to collection data it is endemic to the Cape, and during the sur-
veys it was found along the west coast from Paternoster to Kommetjie.

Larva (Fig. 10P)

About 3,8 mm long when full grown and in dorsal view very similar to that
of Oxytelus caffer, but cerci somewhat longer and, although also dark, they are
not piceous; tenth segment also tubular and longer in relation to body. Sparsely
covered with setae most of which are of ‘frayed’ type (Fig. 11L), particularly
those forming transverse row in middle of each abdominal tergite (except last
two tergites), which are stronger developed than others. As in O. caffer a thin,
longitudinal, median line traverses thoracic segments and joins epicranial suture.
Abdominal shields entire. Legs also similar to those of O. caffer and sparsely
covered with spines; the tarsungulus long, acute and simple and with short pos-
terodorsal and longer ventral spine some distance from base. Spiracles similar to
those of O. caffer and similarly situated. Cerci (Fig. 10U) undivided and bearing
two ‘frayed’ type setae on basal half, one on external margin and one dorsal, as
well as two long ventrally situated setae, of which one is situated near apex; also
two small spine-like setae, one just below exterior ‘frayed’ seta and one on inter-
nal margin, between the two long setae. Apex of cercus conical with single short

SOUTH AFRICAN ARTHROPODS 325

seta at base of cone. Integument of dorsal shields lacking triangular denticles
found in O. caffer.

Head capsule (Fig. 10Q, T, V)

Sides slightly convex and frontal and epicranial sutures fairly distinct; neck
absent. Five ocelli present on each side as illustrated. Hind margin almost
straight or slightly emarginate with clypeofrontal suture obsolete in specimens
examined. Labrum free, its anterior margin widely convex and devoid of any
setae dorsally in most specimens seen, except two short submedian setae near
anterior border. Extreme apical portion somewhat raised and with two longer
setae and some spicules ventrally. Chaetotaxy as illustrated; some hairs of
‘frayed’ type. Antennae three-segmented, with first segment short, second about
twice as long as first and bearing three long subapical setae and fairly long
finger-shaped sensory cone, which is about as long as third antennal segment.
Latter about as long as the first segment, with three long subapical setae and
three to five short spines or sensory pegs apically. Postgenae ventrally separated
by short gula.

Mandibles (Fig. 10W, Y)

Almost sickle-shaped and about twice as long as width at base; about half
as long as cranium. Cutting edge bidentate (with one long apical and one smaller
preapical tooth). Molar area without teeth but trenchant and bearing prostheca
distally. Dorsal carinae rounded and indication of scrobis present, bearing long
seta distally and shorter one some distance from base.

Maxillae (Fig. 10R)

Mala long and tapering with about six spicules or cones ventrally on mesal
side near apex; lateral margin with short seta some distance from base and one
on mesal side near base. Another short mesal seta situated about half-way be-
tween base and apex. Palpifer short and segment-like in some specimens and
ventrally bearing short seta. Palpus three-segmented; first two segments of equal
length, third longer; second segment with two lateral setae. Stipes ventrally with
two setae. Sclerite between cardo and mala, bearing two short setae near base,
which is raised and fairly convex dorsoventrally.

Labium (Fig. 10S)

Mentum small, membraneous and bearing four small setae on basal sclero-
tized area. Prementum bearing two segmented palpus and two setae on ventral
side close to finger-shaped ligula, which is nearly as long as palpi. Prementum
minutely setose near base of ligula, otherwise devoid of setae.

Pupa (Fig. 10Z)
Yellowish white in colour, becoming dark shortly before beetle emerges.
Length 2,5-2,8 mm. Characterized by long setae: two on front, two on hind

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Family Staphylinidae.
A-N. Oxytelus caffer. A. Larva (dorsal view). B. Larva (left lateral view). C. Head (ventral
view). D. Head (dorsal view). E. Left urogomphus. F. Right mesothoracic leg. G. Eggs.
H. Pupa (ventral view). I. Left maxilla (dorsal view). J. Labium (dorsal view). K. Left man-
dible (dorsal view). L. Apex of left mandible (lateral view). M. Right mandible (dorsal
view). N. Head (left lateral view).
O-Z Omalium arenarium. O. Adult. P. Larva (left lateral view). OQ. Head (dorsal view).
R. Left maxilla (dorsal view). S. Labium (dorsal view). T. Head (ventral view). U. Left uro-
gomphus. V. Head (left lateral view). W. Left mandible (dorsal view). X. Apex of pupa
(ventral view). Y. Right mandible (dorsal view). Z. Pupa (ventral view).

SOUTH AFRICAN ARTHROPODS 327)

margin of pronotum and seven laterally, two submedially on abdominal terga
1-7. Also two long setae on front margin of head and some smaller ones on both
head and pronotum and on abdominal sterna. First three abdominal spiracles
circular and clearly visible, the others inconspicuous. Abdominal apex with two
fleshy projections, each with a long seta; developing genital capsule either pres-
ent as a rounded swelling or as two projections, each with short seta apically
(Fig. 10X).

BIOLOGY

Larvae of this staphylinid were observed in decaying kelp along the west
coast during the winter months. The larval period seems rather short and most
larvae collected in June pupated during the same month. Most of the young lar-
vae used for rearing of the adults were kept in decaying kelp from which all
other arthropods were removed. These larvae developed to maturity and pro-
duced normal adults.

Oxytelus caffer (Erichson)

DESCRIPTION
Adult (Fig. 1C)

Black, matt or slightly shiny, particularly abdomen; strongly punctate and
also rugulose, with rugulae more or less longitudinally arranged; areas between
rugulae and punctures also minutely reticulate. Legs reddish brown. Easily re-
cognized by presence of four longitudinal, rounded carinae over the middle of
the pronotum. Length 3,2-4,2 mm.

One of the most numerous and widely spread staphylinids; according to
Scheerpeltz (1973) it is distributed over a large area including South West Afri-
ca, South Africa and east Africa.

Larva (Fig. 10A-B)

Whitish yellow to lemon yellow, in some specimens even cadmium yellow
in colour. About 7,0 mm long when full grown, subcylindrical and rather similar
to that of Aleochara sp. Thoracic tergites divided medially by distinct line, but
those of abdomen undivided. Anterior borders of all dorsal abdominal shields
clearly demarcated by a thin, dark line. Thorax and abdomen sparsely covered
with brownish hairs placed more or less in transverse rows; some of these hairs
long, particularly those over middle of segments.

Ninth abdominal segment with two short piceous to blackish urogomphi
(Fig. 10E), which appear undivided; each urogomphus with long apical seta,
three long subapical setae, smaller seta about half-way between base and apex
and small seta near base. Tenth segment forming short tube, with anal opening a
short Y-shaped slit; stem of the Y pointing downwards and ending in pointed
lobe. Abdominal spiracles circular and all more or less of the same size. Meso-

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

thoracic spiracles also circular, slightly larger than those on abdomen and also
situated on side of body between pro- and mesothorax. Dorsal shield without
any fine denticles; hairs simple.

Legs (Fig. 10F) as in Aleochara sp. with reduced number of spines. Tibia
also with three spines around apex, one being dorsal; claws simple, acute and
with two small spines placed ventrally near base.

Head capsule (Fig. 10C-D, N)

Pale golden brown with front one-third darker in older specimens; fairly
smooth and shiny. Front transversely and finely rugulose; frontal and epicranial
sutures rather indistinct in some specimens. Chaetotaxy as illustrated. Sides
somewhat convex, hind margin widely and shallowly emarginate on dorsal side.
One fairly large ocellus present on each side. Antennae appearing three-
segmented; first segment about as long as second; latter with three long sub-
apical setae and a large sensory cone on mesal side; third segment small, about
as long as sensory cone, with three long preapical setae and about three short
setae at apex. Neck absent. Labrum free, trapezoidal, and with rounded lobe on
each side at base; also with almost triangular sclerite in middle at junction with
clypeus. Frontoclypeal suture only partly developed on each side. Postgenae
ventrally separated by small gula. Hind margin of cranium ventrally deeply
emarginate.

Mandibles (Fig. 10K—M)

Narrow; broader basally and about twice longer than width at base. Fairly
rounded dorsoventrally on external side, without dorsal carina. Long seta pres-
ent near base (rather dorsally placed) and smaller seta (on external side) near
base of apical teeth. Each mandible with three apical teeth although, when
viewed from above, mandibles appear bidentate. Molar areas absent, but ven-
tral side with oval, flat area bearing dorsal trenchant ridge which forms broad
tooth (ts) on mesal side.

Maxillae (Fig. 101)

Rather similar to those of Aleochara sp., but galea and lacinia not separ-
ated or delineated by suture. Mala oblique, almost trenchant, and furnished
with eight to nine strong, palus-like teeth; basal teeth largest and extreme basal
one in some specimens accompanied by one or two smaller ones. Basal half of
mala ventrally also with three to four smaller teeth and apical part with five to
six. Maxilla dorsally without setae, except for one long seta on lateral margin
below palpus. Mesal margin of mala with few very small spines (or sensory pegs)
near junction with maxillary articulating area. Maxilla ventrally with about four
setae of which three are on apical half and one on basal half. Maxillary articula-
ting area ventrally present as large convex lobe on mesal side of stipes. Cardo
with one seta only on ventral side and with Y-shaped suture. Palpifer small and

SOUTH AFRICAN ARTHROPODS 329

bearing single seta ventrally. Palpus three-segmented, with first and apical seg-
ments of about equal length; first is shaped like wide S with a row of very small
spine-like setae round its apex; second segment short, about half the length of
first and with two ventral setae.

Labium (Fig. 10J)

Rather similar to that of Aleochara sp.; submentum fairly large and bearing
two setae. Mentum wider than submentum and only partly sclerotized in young
larvae; it bears four setae. Prementum with broad, lobe-like ligula and two-
segmented palpi. As in maxillary palpi, first segment with row of tiny, spine-like
setae or spicules on ventral side of its apex; apical segment with some sensory
cones; also few tiny spines at base of first segment on palpiger. Stipes ventrally
with two long setae as well as two short ones close to base. Labium dorsally be-
set with fine setae or spicules and two tiny, spine-like setae at apex of ligula.
Four small circular sensory areas, two on ligula and two on stipes, clearly visible
dorsally.

Pupa (Fig. 10H)

Oblong and light yellow in colour, but turns almost black just before emer-
gence of beetle. Length 4,9-6,8 mm. Abdominal apex rather similar to that of
Xantholinus sp. (Fig. 14L), with the developing genital capsule in most speci-
mens visible as small conical tubercle just above two fleshy conical projections.
First three abdominal spiracles circular, small and clearly visible; others incon-
spicuous. First abdominal tergum with two long submedian setae; second to
seventh abdominal segments each with lateral setae. Head with three long setae
on each side and two on occiput. Pronotum with two long setae on anterior mar-
gin and some long setae (about ten) along posterior and lateral margins. Other-
wise devoid of setae.

BIOLOGY

Large numbers of the beetles were observed in cow-pats almost throughout
the year and occurred in the wet as well as in the drier parts of the dung. Larvae
and pupae were collected during autumn and winter and both larvae and beetles
were found to feed on dung particles and on colloids oozing from the dung. Eggs
(Fig. 10G) of this species are oval, smooth, and shiny, and are whitish yellow to
bright lemon yellow in colour; those collected during June along the south coast
measure 0,68 X 0,36 mm to 0,76 X 0,46 mm.

When full-grown, the larvae construct small oval cells in the soil or in the
drier parts of the dung, in which the prepupae and pupae are formed. Pupal
stages lasted 10-14 days in the laboratory during the winter months. Prepupae
are somewhat wrinkled and pale yellow in colour. Newly emerged beetles are
black with white wings, which darken after a few hours.

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

Philonthus natalensis Boheman
DESCRIPTION
Adult (Fig. 11J)

Shiny black beetle, 11-12,5 mm long. Elytra short and covered with
medium-long, golden-whitish hairs. Body microscopically striatoreticulate and
with silky sheen; abdomen more shiny. Eyes large, oval, slightly less than twice
as long as wide. Head smooth, except for row of large punctures around eyes
and few postocular punctures, each bearing single long brown seta. Pronotum
laterally with some long setae, particularly the transverse row parallel with ante-
rior border on each shoulder; also about eight punctures on each side of the pro-
notal disc, each with long seta. Antennae brown, legs piceous with spines and
hairs on tibiae and tarsi.

Widespread in east, south-east and South Africa (Scheerpeltz 1973).

Larva (Fig. 11A-B)

When mature, about 15,6 mm long. Head and pronotum dark reddish
brown, meso- and metathorax yellowish brown with dark spot on each side;
mesothorax darker in middle. Underside of thorax and abdomen creamy white;
each tergum with two brownish sclerotized plates. Tenth abdominal segment
long, tube-like and functioning as pseudopod. Cerci very long, inarticulate and
about as long as first seven abdominal segments; basal ninth of each cercus beset
with small spines, remainder of cercus with fine hairs; spine-like seta present
dorsally near base of each cercus. Head slightly downwardly directed and about
as wide as thorax. Pronotum narrower in front than behind and with shallow,
oblique, more weakly sclerotized depression on each side near anterior angles.
Sclerotized areas fairly smooth and shiny.

Body sparsely covered with long golden brown setae, some of which of
‘frayed’ type (Fig. 11L) with branched tips. Body integument dorsally with
microscopic triangular denticles, visible only under high magnification. Ecdysial
suture visible as pale reddish-yellow longitudinal line over the middle of thoracic
terga, anteriorly continuous with epicranial stem. Sterna similar to those of Sta-
phylinus sp., but most setae of ‘frayed’ type. Mesonotal spiracle large, oval, and
with single seta anteriorly. Abdominal spiracles almost circular, those of first ab-
dominal segment larger. |

Only coxae and femora with some short, fine hairs (Fig. 11E), otherwise
legs devoid of hairs, but with strong spines or spine-like setae; those on femora
arranged in double ventral row; space between rows bare. Each trochanter with
six to seven ventral spines, and long slender apical seta. Tibiae with spines dis-
tributed over whole surface. Claws as in Staphylinus sp. (Fig. 13F), but each one
with three short spines, one of which is ventral, the other two situated more or
less on posterior side.

The larvae are characteristic of the genus Philonthus and easily recognized
by their very long cerci.

SOUTH AFRICAN ARTHROPODS 331

Fig. 11. Family Staphylinidae.
Philonthus natalensis. A. Larva (dorsal view). B. Larva (left lateral view). C. Head (dorsal
view). D. Head (left lateral view). E. Right leg (posterior view). F. Head (ventral view).
G. Left mandible (dorsal view). H. Right mandible (dorsal view). I. Labium (dorsal view).
J. Aduit. K. Left maxilla (dorsal view). L. Modified seta and triangular denticles on integu-
ment (highly magnified). . Pupa (ventral view). N. Pupa (left lateral view). O. Ventral
view of last two abdominal segments of pupa. P. Egg.

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

Head capsule (Fig. 11C-D, F)

Dark yellowish brown, fairly smooth and shiny. Sides almost parallel, pos-
terior corners rounded. Long epicranial stem and shorter frontal sutures visible
as thin pale-reddish lines. Neck short. Chaetotaxy as illustrated. Four ocelli
present. Cheeks rounded dorsoventrally and without carinae. Frons small, with
two setae on each side. Each half of epicranium bearing only about eight dorsal
setae and only faint indication of paler longitudinal lines. Nasale shaped as in
Staphylinus sp., septemdentate and with same number of setae.

Ventral aspect of cranium very similar to that of Staphylinus sp.; pregula,
however, much longer and more acute posteriorly and with same number of
setae. Posterior tentorial pits visible as short, elongate black marks laterad to
arms of gular suture, not adjacent to stem as in Staphylinus sp. Antennae
four-segmented, with one sensory cone on apex of fourth segment and one
-mesally near apex of penultimate segment; latter and apical segment with pre-
apical setae.

Mandibles (Fig. 11G—H)

Almost falcate, slightly less than half the length of cranium. Retinaculum
absent; each mandible with dorsal carina on cutting edge extending over apical
two-thirds of its length. Apices of both mandibles rounded, not pointed as in
Staphylinus sp. Lateral side of mandible rounded, without carina, and with
single fairly long seta near base. Molar areas and penicilli absent.

Maxillae (Fig. 11K)

Similar to those of Staphylinus sp. Stipes fairly long, both ventral and dor-
sal faces devoid of setae, except for single long seta ventrally near base and in-
conspicuous seta on base of palp; also two setae on lateral margin, one of which
is fairly long, and about four setae on mesal side (mala). Inner lobes absent,
outer lobes or galea conical and about as long as penultimate segment of labial
palp and bearing two setae at apex as well as single short seta near base. Maxil-
lary palps appear three-segmented, with apical and penultimate segments of
about equal length, latter with single lateral and mesal seta; terminal segment
with apical third narrowed. Palpifer with single seta as in Staphylinus sp. Cardo
long, about three-quarters length of stipes and bearing single ventral seta.

Labium (Fig. 111)

Ligula conical as in Staphylinus sp. and nearly as long as penultimate seg-
ment of palp. Latter three-segmented, with first segment about twice as long as
penultimate one, apical segment less than half the length of latter. As in Staphy-
linus sp., prementum can be retracted into mentum and bears ventral seta near
each palpal base. Labial stipes, including palpigers, dorsally covered with fine
silky hairs as in Staphylinus sp. Mentum on each side with tooth, as in latter
species.

SOUTH AFRICAN ARTHROPODS 333

Pupa (Fig. 11M-—O)

Semi-obtect, almost smooth and shiny and very similar to that of Staphyli-
nus sp. Light reddish yellow in colour. Length 7,9-8,7 mm. Anterior pronotal
margin with sixteen to twenty long barbed setae and lateral margins of abdomi-
nal segments 7 and 8 also with fairly long setae. Spiracles as in Staphylinus sp.
All legs with six to nine pointed tubercles. It differs from pupa of Staphylinus in
possessing two conical projections at posterior end, each with an elongate seta-
like filament. Developing genital capsule (Fig. 110) also represented by two
spine-like projections apically on ventral side.

BIOLOGY

The oval, almost pure-white eggs (Fig. 11P) are fairly large, about
2,2 mm long and about 1,4 mm wide, matt and with indistinct striae and
longitudinal tuberculate ridges alternately arranged, as illustrated. They were
laid singly in semi-fresh dung during October and hatched in 4 or 5 days. On
hatching the chorion splits from the top over one or both sides and just
before hatching the mandibles of the larva can be seen in the middle of the
egg on one side.

The newly hatched larva looks very much like the mature larva, but is pure
white with golden-brown mandibles and four brownish eyes on each side of the
head. It measures about 6,1 mm in length shortly after hatching and its long cer-
ci are about as long as the almost transparent body. The spiracles are very con-
spicuous at this stage and in most of the specimens examined there are only a
few setae on the body and most of these, even some of the spines on the legs
(except a few on the head, thorax and ventral side of the 8th abdominal seg-
ment), are of the ‘frayed’ type. The claws are conspicuous and bear only two
ventral spines. The tenth segment is very large in relation to the rest of the body
and is about as long as the head. In the latter the coronal and frontal sutures are
obsolete and the two small setae on the middle of the frontal disc, as well as the
two posterior setae near the posterior border of the head, are absent. The man-
dibles are fairly pointed, and the small apical part of the third segment of the
maxillary palp is already differentiated. After a few hours the larva becomes
greyish in colour, but the first two body segments as well as the head are golden
brown.

The lifespan of the larva is short and those reared in the laboratory were
mature in about 14 days at 25°-27°C after which they pupated in their larval
skins in small clay cells either in the soil or in the remains of the dung. Hafez
(1939a) gives the duration of the larval stage of the cosmopolitan Philonthus
quisquiliarius (Gyllenhal) in Cairo as 6 to 8 days at 30°C. Pupae were recovered
during November and the adult beetles emerged after about 9 days. The newly
emerged beetles are black with almost pure-white wings and elytra.

This species was very abundant in fresh cow-dung along the south coast.
Both larvae and adults are predacious on other insects and also feed on the
juices oozing from the dung.

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

Philonthus spp.
(Fig. 1B)

Two forms of this beetle were collected in cow-dung. The one (sp. A) is
somewhat larger than the other (sp. B) and was collected mostly along the south
coast; sp. B was found throughout the area surveyed. As a proper identification
of the two forms was impossible, they are here treated as two separate species;
however, there are no appreciable morphological differences between the
larvae.

Philonthus sp. A (large)
DESCRIPTION
Adult

Similar to P. natalensis but smaller, only 7,5—-7,9 mm long; colour black,
head and pronotum very shiny. Head narrower than in latter species and more
oval in shape.

This species was found in association with P. natalensis and is apparently
widely spread along the south coast; it was also observed at Montagu.

Larva (Fig. 12A)

Rather similar to that of P. natalensis but smaller, measuring only 10-12
mm in length when mature, and cerci (Fig. 12L) much shorter, consisting of two
segments. Second segment of cercus slightly less than half the length of first and
with long apical seta, as well as small subapical, spine-like seta; first segment
with about six long setae as well as some shorter ones. Tenth abdominal seg-
ment tube-like, but shorter than in P. natalensis. Whole body sparsely covered
with medium-long brownish setae, some of which are of ‘frayed’ type (Fig. 12B).
Otherwise exactly as in P. natalensis. The dorsal integument covered with denti-
cles similar to those of P. natalensis, but somewhat larger. Spiracles similar to
those of P. natalensis, including anteriorly situated seta on mesothoracic pair.
Legs also similar to those of P. natalensis, including tarsal spines and long seta
on apical portion of trochanter.

Head capsule (Fig. 12C—D)

Light brown to yellowish brown in colour; very finely reticulate and with
some rugae; smooth and shiny, about as long as wide, with sides parallel and
hind margin almost straight. Neck as in P. natalensis, posterior angles rounded.
Nasale, ocelli antennae and ecdysial sutures as in P. natalensis. Ventral aspect of
cranium also very similar. Chaetotaxy as in Figure 12C—D.

Mandibles

Similar to those of P. natalensis and much shorter than cranium; also bear-
ing seta close to base.

SOUTH AFRICAN ARTHROPODS 335

Fig. 12. Family Staphylinidae.
A-J. Philonthus sp. A. (large). A. Larva (left lateral view). B. Integument of dorsal shield,
showing hair and denticles (highly magnified). C. Head (dorsal view). D. Head (left lateral
view). E. Left maxilla (dorsal view). F. Labium (dorsal view). G. Left maxilla (ventral
view). H. Egg. I. Pupa (ventral view). J. Abdominal apex of pupa (ventral view).
K-T. Philonthus sp. B (small) K. Larva (left lateral view). L. Urogomphus. M. Integument
of dorsal shield (highly magnified). N. Head (dorsal view). O. Left maxilla (dorsal view).
P. Labium (ventral view). Q. Labium (dorsal view). R. Left maxilla (ventral view).
S. Abdominal apex of pupa (ventral view). T. Head (left lateral view).

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

Maxillae (Fig. 12E, G)

Very similar to those of P. natalensis. However, maxillary palpi clearly
four-segmented, tapering towards apex, with apical and first segments of
about equal size; second segment slightly less than twice as long as first, and
third segment slightly longer than apical one; only second segment with two
setae. Inner lobes absent, outer lobes present as finger-like structure, about as
long as first palpal segment and bearing two to three setae at apex. Otherwise
as in P. natalensis. Stipes with only one spine-like seta mesally and two on
lateral side; ventrally with three setae and dorsally either with one near mesal
margin or without any setae. Cardo as in P. natalensis. Palpifer also with
single seta.

Labium (Fig. 12F)

Ligula finger-shaped and conical, almost as in P. natalensis, and about as
long as penultimate segment of three-segmented palp. First segment of palpus
long, about twice as long as the second. Sides of ligular stipes (prementum) scle-
rotized, median area beset with fine hairs dorsally, as in P. natalensis; palpigers
not as convex as in latter. Labium ventrally with two submedian setae on poste-
rior boundary of palpigers (Fig. 12P), exactly as in P. natalensis; also with tooth
on each side of mentum.

Pupa (Fig. 121I—J)

Very similar to that of P. natalensis but smaller, only 4,8—5 mm in length.
Colour pale whitish yellow, but becoming piceous just before beetle emerges.
Differs from pupa of P. natalensis in that setae on anterior margin of pronotum
are much longer in relation to body; there are furthermore only five to six setae
on each side, compared with eight to eleven in P. natalensis. In most specimens
examined, developing genital capsule (Fig. 12J) appeared as a flat lobe flush
with body wall and, except in few cases, the two spine-like projections were ab-
sent. Every abdominal segment with medium-long seta laterally, whereas in
P. natalensis only segments 7 and 8 bear lateral setae.

BIOLOGY

The eggs (Fig. 12H) of this species are very similar to those of P. natalensis
but are smaller, measuring about 1,5 mm by 0,96 mm and the tubercles or den-
tations on the ridges are larger; the colour of most of the eggs was dirty white.
They were laid singly in fresh dung at the same time as those of P. natalensis.
The larval lifespan is more or less the same as in P. natalensis and pupae ap-
peared during November. As in the latter species, the larvae construct clay cells
in the soil in which they pupate. Most of the larvae were reared in fresh dung
that did not contain any arthropods, except for mites and a few sciarid larvae;
when crowded they became cannibalistic.

SOUTH AFRICAN ARTHROPODS 337

Philonthus sp. B (small)
DESCRIPTION
Adult

Black, head and pronotum smooth and very shiny. Elytra brownish black
and densely setose. Abdomen shiny and densely covered with setae. Similar to
sp. A except for size: most specimens collected measure only 3,7-5,3 mm.

This is one of the most common rove-beetles in cow-dung and was observed
in all the areas surveyed.

Larva (Fig. 12K)

Very similar to sp. A in general form, setal pattern and colour, but some-
what smaller, mature larvae measuring 9-10 mm. Spiracles and legs similar to
those of sp. A, including long seta on apical portion of trochanters and anterior-
ly situated seta on mesothoracic spiracles. Dorsal shields with triangular teeth as
in sp. A; some of the teeth broad and tri- or multidentate as in sp. A (Fig. 12M)
and to some extent also in P. natalensis. Body setae of ‘frayed’ type as in other
species.

Head capsule (Fig. 12N, T)

Very similar to that of sp. A; however, nasale somewhat more produced in
front. Colour golden brown. Almost smooth and shiny, very finely reticulate and
with some rugae, particularly near the frons. About as long as wide or slightly
wider than long in some specimens. Chaetotaxy as in Figure 12N, T. Ventral
aspect similar to that of sp. A. Antennae as in latter and also four-segmented.

Mandibles, maxillae (Fig. 120, R), and labium (Fig. 12P, Q)

Exactly as in sp. A.
Pupa

Similar to sp. A but smaller, varying from 3,8 to 4,5 mm in length and most
specimens collected have the two spine-like projections present (Fig. 12S).

BIOLOGY

Beetles and larvae were found breeding in semi-fresh cow-dung throughout
the summer and autumn and the larval lifespan seems to be the same as for sp.
A. In both cases the larvae construct loose cells in the soil in which they remain
dormant for a short while; the pupal stage in sp. B lasts about 6 days in January.

Staphylinus sp.
DESCRIPTION
Adult (Fig. 13M)

Large, beautiful, and aggressive species, 15,8-17 mm long, densely covered
with fine, silky, golden-brown hairs, which give it a chequered appearance. True

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

colour of body dark brown, except elytra, borders of prothorax and apical mar-
gins of segments, which are brownish red. Densely punctate and dull all over,
except for a narrow longitudinal, smooth and shiny line over middle of prono-
tum. Coloration of anterior border of cranium, including a V-shaped area in
middle of anterior border and basal part of labrum, reddish yellow. A narrow
area around the eyes, extending nearly to the posterior border of the head as
well as the clypeus, also reddish yellow. Labrum deeply cleft in middle. Anten-
nae and legs pale yellowish brown.

Distributed throughout the interior of the southern and south-western parts
of South Africa where the surveys were made.

Larva (Fig. 13A-B)

Length 23-24 mm when mature. In general shape very similar to the
larva of Laemostenus complanatus, but head is carried horizontally, with
mandibles directed slightly downward. Head and pronotal shield dark reddish
brown, head almost piceous in old larvae; meso-, and metathoracic and first
abdominal tergum light brown. Thoracic terga with thin, white median line
(ecdysial suture), which is anteriorly continuous with epicranial stem. Head as
wide as, or slightly wider than, pronotum and in mature specimens examined
it measured 4 mm long, including closed mandibles. Legs and antennae pale
brownish yellow.

Abdomen creamy white and tergal plates only slightly demarcated on seg-
ments 3-9. Second abdominal segment with tergum almost as strongly sclero-
tized as first. Each abdominal sternum with two sclerotized plates bearing setae,
of which at least some are of ‘frayed’ type; plates rather obscure, except on first
abdominal segment, which has two smaller, clearly demarcated plates with
about five setae each. Body integument dorsally with microscopic, triangular
denticles similar to those of Philonthus natalensis. Cerci same colour as abdo-
men and fairly long. Tenth segment tubular, almost as long as eighth and func-
tions as proleg. Spiracles oval, with mesothoracic pair (Fig. 13D) largest and
with strong seta anteriorly; those on first abdominal segment (Fig. 13E) larger
than other abdominal spiracles. Inner surface of each spiracle lined with fine,
microscopic hairs. Some setae on abdomen with branched tips. Each cercus
(Fig. 13H) with six long setae, of which one is situated apically; also six to eight
short ‘frayed’ setae at base of each cercus. Latter appears two-segmented, with
apical segment thin, nearly half the length of first, and situated on short seg-
ment-like extension of first.

Legs (Fig. 13F) yellowish brown. Tarsungulus simple, acute and about half
the length of tibia, bearing two small setae on ventral side, about half-way be-
tween its base and apex. All segments except the coxae with strong, almost
palus-like setae or spines. Femur on each side with lateroventral row of spines,
which meet at apex; ventral area between rows with short basal row of three
spines; laterally also with row of finer setae above the spines and with few small
scattered dorsal setae; otherwise its dorsum devoid of hairs. Spines on tibia

SOUTH AFRICAN ARTHROPODS 339

Fig. 13. Family Staphylinidae.
Staphylinus sp. A. Larva (dorsal view). B. Larva (left lateral view). C. Head (dorsal view).
D. Right mesothoracic spiracle. E. Right abdominal spiracle. F. Right mesothoracic leg.
G. Head (left lateral view). H. Urogomphus. I. Head (ventral view). J. Left mandible (dor-
sal view). K. Right mandible (dorsal view). L. Labium (dorsal view). M. Adult. N. Left
maxilla (dorsal view). O. Pupa (ventral view). P. Pupa (dorsal view). Q. Pupa (left lateral
view).

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

similarly arranged, but with only two spines on ventral area between rows. Tro-
chanters with about twelve strong setae, situated ventrally and laterally; apical
one very long.

Head capsule (Fig. 13C, G, I)

About as long as wide, or slightly wider than long. Sutures distinct; epicra-
nial stem long, frontal sutures enclosing small frons. Dark reddish brown in
colour. Distal half of cranium and largest part of frons microscopically reticulate;
basal half more or less smooth, very superficially reticulate and also with some
rugulosity. Whole head somewhat shiny, particularly front half. Chaetotaxy as in
illustration. Epicranium on each side with narrow, smooth longitudinal area
divided by fine, branched stria; each area with about four setae. Nasale septem-
dentate, median and lateral teeth small. Intervening spaces between teeth each
with short seta; long seta present on each large lateral tooth, and shorter one
posteromesad of latter.

Four-segmented antennae situated on each side of nasale. Second antennal
segment largest, more than twice as long as first; fourth segment small and about
half the length of third; latter with sensory cone on the mesal side; fourth seg-
ment with sensory cone at apex. Both penultimate and apical segments with
three setae, two on mesal side in penultimate segment and one on mesal side in
apical segment.

Four ocelli on each side. Gular suture distinct and Y-shaped, enclosing a
narrow pregula; tentorial pits situated just behind latter. Collum short, but dis-
tinct. Cranium on each side with weak carina, which ends posteriorly in tooth-
like tubercle.

Mandibles (Fig. 13J—K)

Falcate, about half the length of cranium and almost symmetrical. Cutting
edge with uninterrupted dorsal and ventral carina_and with indication of dorsal
tooth or retinaculum half-way between apex and base. Molar areas and penicilli
absent. Single, medium-long seta present near base on external margin and
minute one opposite dorsal tooth.

Maxillae (Fig. 13N)

Stipes long and devoid of setae ventrally and dorsally, except for two short
setae, one on inconspicuous base of three-segmented palp and one on apex of
stipes. Mesal and lateral margins both with four setae. Cardo with a single long
seta on lateral margin. Inner lobes of maxillae absent; outer lobes one-
segmented, conical and slightly less than twice as long as apical segment of palp,
and with two apical setae; also with small seta on apical half of lateral margin.
First two segments of palp about equal in length, apical one about half the
length of penultimate one; latter with single lateral seta on each side. Palpifer
segment-like and small, less than half the length of first palpal segment.

SOUTH AFRICAN ARTHROPODS 341

Labium (Fig. 13L)

Ligula conical, elongate and one-segmented. Palpi two-segmented; apical
segment shorter than ligula and nearly half the length of basal segment. Labial
stipes (prementum, with ventral seta near palpal bases) can be retracted into
mentum. Pointed tooth present on hypopharyngeal bracon on each side of men-
tum at base of which is short ventral seta. Mentum basally also with two ventral
setae. Labium dorsally with fine, silky hairs on its base and on stipes, extending
forward to base of ligula and also covering bases of palpi.

Pupa (Fig. 130-Q)

Semi-obtect, almost smooth and shiny, heavily sclerotized, orange-brown,
and about 12,5 mm long. Body devoid of setae, except for row of about twenty-
one barbed setae on anterior margin of pronotum and single barbed seta on lat-
eral sides of segments 7 and 8. Each mesothoracic leg with about sixteen small
tubercles and each metathoracic leg with about five. Median area of mesonotum
immediately anterior to M-shaped suture and upper boundaries of legs swollen
and somewhat conical. First four spiracles on each side conical and conspicuous,
last four small and inconspicuous. Labrum narrowly but acutely excised in mid-
dle, with short suture extending from emargination. Developing genital capsule
indicated only by two almost flat areas (each with tiny depression) on ventral
side of apical segment. Pupa darkens after a few days and four small black spots
(eyes) become visible on each side of head.

BIOLOGY

Young larvae were collected under semi-fresh cow-pats from April to Au-
gust and pupation took place from July to September. The larval stages lasted
for 100-140 days and the pupal stages at least 35 days. Most of the pupae were
found in small clay cells in the soil, but a few were found in cells hollowed out in
cow-pats lying on fairly dry and hard soil. The larvae collected fed mostly on lar-
vae of the dung fly, Orthellia peronii (Robineau-Desvoidy).

Large, oval, creamy-white eggs measuring about 2,9 mm by 2,5 mm, with
the surface densely covered with microscopic spinules or tubercles, and found in
dung containing its larvae, apparently belong to this species. The eggs very
closely resemble those of some of the larger scarabs.

Xantholinus sp.
DESCRIPTION
Adult (Fig. 14A)

Dark blackish brown to almost black in colour, with apical part of elytra
brownish. Very finely reticulate-rugulose, fairly shiny and sparsely punctured
with fairly large oblong punctures on head, thorax, and elytra. Covered with
medium-long golden hairs. Eyes large, situated near anterior margin of head.
Mandibles, legs, and antennal flagellum reddish brown. Length 6,3—7,5 mm.

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

Widely distributed in the southern Cape Province, having been observed in
carcasses around Cape Town and in semi-fresh to fresh cow-pats along the south
coast.

Larva (Fig. 14B)

Mature larvae measure about 7,8 mm and are creamy white, except for
head and pronotum, which are light reddish brown or brownish red, and meso-
and metanotum and legs, which are pale yellowish red. Sparsely covered with
medium-long brownish hairs more or less arranged in two transverse rows on
tergites. Urogomphi (Fig. 14D) shorter than the tubular tenth segment and two-
segmented; the terminal segment thin and with long apical seta. Spiracles oval;
meso-thoracic one transverse (Fig. 14F) and with seta anterior to it; those on ab-
dominal segments obliquely arranged; first abdominal and mesothoracic spiracle
of about equal size and larger than rest. Legs (Fig. 14E) similar to those of Phil-
onthus spp., but spines somewhat longer; trochanter also with a long seta anter-
iorly; claws acute and with only two spines more or less on ventral side. Median
longitudinal line (ecdysial suture) over thorax distinct, anteriorly continuous
with epicranial stem. Tergites on abdominal segments divided medially by fairly
broad longitudinal line. Dorsal integument without triangular denticles, but in
some places with long rows of fine spicules (Fig. 14G). Setae and spines simple.

Head capsule (Fig. 14C, H-I)

About as wide as pronotum, sides almost parallel and somewhat sinuate;
posterior corners rounded and posterior margin almost straight. Golden brown
to light brownish red in colour, almost smooth and shiny; anterior half rugose in
middle and also finely reticulate between rugae; posterior half faintly rugulose.
Nasale prolonged in front and consisting of median lobe with three teeth, of
which middle one is smallest, and two lateral lobes, each with four teeth of
about equal size. Chaetotaxy as illustrated. Frontal and epicranial sutures as in
Philonthus spp. Neck fairly wide. Antennae appear four-segmented, with basal
segment small, much shorter than second; the latter and third antennal segments
of about equal size; apical segment with three long subapical setae and about
three smaller setae at apex; slightly shorter than third segment, the latter with
three long subapical setae and a long sensory cone on the ventral side close to
apical border. Single ocellus present on each side. Gular suture Y-shaped and
anteriorly enclosing fairly wide pregula, beset with numerous small, rounded
tubercles.

Mandibles (Fig. 14N)

Falcate and much shorter than cranium; lateral side rounded, without dor-
sal carina and with three lateral setae, of which basal one is smallest. Molar area
absent but small denticle, reminiscent of prostheca, is present. Cutting edge
pointed apically and with dorsal trenchant ridge running from apex to about half
the length of mandible.

SOUTH AFRICAN ARTHROPODS 343

Fig. 14. Family Staphylinidae.
Xantholinus sp. A. Adult. B. Larva (left lateral view). C. Head (dorsal view). D. Left uro-
gomphus. E. Right mesothoracic leg. F. Light mesothoracic spiracle. G. Integument of dor-
sal shield of abdomen (highly magnified). H. Head (left lateral view). I. Head (ventral
view). J. Left maxilla (dorsal view). K. Left maxilla (ventral view). L. Last two abdominal
segments of pupa (ventral view). M. Pupa (ventral view). N. Left and right mandible (dorsal
view). O. Labium (ventral view). P. Labium (dorsal view). Q. Pupa (left lateral view).

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

Maxillae (Fig. 14J—K)

Similar to that of Philonthus natalensis, but maxillary palp clearly four-
segmented, with segments tapering towards apex; second segment with two lat-
eral setae of which one is mesal. Palpifer segment-like and bearing two setae
ventrally. Stipes long, ventrolaterally with about six setae and with another seta
almost in middle of stipes. Cardo ventrally also with one long seta situated more
to mesal side. Stipes and cardo dorsally without setae, except for short one situ-
ated laterally just below outer lobe and in some specimens second one just be-
low first. Outer lobe finger-like, nearly as long as second palpal segment and
bearing about three apical setae.

Labium (Fig. 140-P)

Very similar to that of P. natalensis and with three-segmented palpi. Ligula
conical and beset with fine hairs dorsally (including also the palpigers and medi-
an area). Palps tapering towards apex; first segment longest and apical one
shortest; tiny spine present laterally just below palpiger.

Pupa (Fig. 14L-M, Q)

Semi-obtect. Newly formed pupae are white but soon turn to a whitish yel-
low with the posterior projections brownish red and mandibles red-brown.
Length 3,9-4,8 mm. Very finely reticulate-punctate, shiny and without any
setae, except two diverging, seta-like filaments borne on two conical caudal pro-
jections.

Pupae rather similar to those of Philonthus natalensis except for absence of
hairs. Developing genital capsule (Fig. 14L) appears as a raised or somewhat
conical area ventrally on abdominal apex. First four abdominal spiracles fairly
large and clearly demarcated, diminishing in size posteriorly; remainder very in-
distinct.

BIOLOGY

Larvae of this species were found in carcasses in Cape Town from Septem-
ber to October; they were feeding mainly on fly larvae, especially those of the
cheese-skipper, Piophila megastigmata McAlpine. The duration of the larval
period was approximately 19 days. Mature larvae construct oval clay cells, mea-
suring 7—7,5 mm in length, in the soil for pupation. In most cases there was a pre-
pupal stage of 3-4 days, during which time the larvae became almost pure white,
except for the reddish head. The pupal stage lasted 11-14 days in November.
Larvae collected in decaying cow-dung from June to August had a somewhat
longer lifespan, the pupal period being 20-22 days.

Newly emerged beetles are blackish with pure white wings, which usually
become blackish within 4-6 days. Some of the beetles were found to feed on
cheese-skipper maggots, but small grubs of skin-and-hide beetles were also at-
tacked; in the laboratory even some of the large white collembolans were de-
voured.

SOUTH AFRICAN ARTHROPODS 345

Family Silphidae

Small to moderately large (about 40 mm) flattened beetles, usually sombre
coloured, often with metallic blue sheen; some with red or orange markings.
Apical abdominal segments often exposed. Antennae ten- to eleven-segmented.
Tarsal formula 5—5—5. Most of them are carrion feeders, both in the adult and
larval form; some feed on snails and lepidopterous larvae and a few are phyto-
phagous (not present in South Africa). Species of the genus Nicrophorus (not
occurring in southern Africa) have the habit of burying the carrion to ensure an
adequate food-supply for the larvae; they are therefore known as burying
beetles.

Only about ten species have been described from the Subsaharan region, of
which the endemic Silpha mutilata Laporte & Castelnau and the more wide-
spread S. micans Fabricius have been introduced into Europe. The latter, to-
gether with S. caeruleoviridans Dohrn (also widely distributed in the Cape Prov-
ince, South West Africa and Zimbabwe), were described under the subgenus
Chalcosilpha; however, according to Arrow (1909) these two species are conspe-
cific. The largest species found in the Cape Province are S. capicola Péringuey,
S. peringueyi Portevin, and S. punctulata Olivier (which is common in fynbos
along the western parts of the southern Cape), all measuring 17 to 20 mm in
length. 3

Both S. micans and S. mutilata (which differs from the first-mentioned
species by the small elevated area or ridge in the middle of each elytron) (Fig.
2F) were found to be common on carcasses of mammals and birds both on the
coast and inland, and were also attracted to soil containing decomposing fish-
meal. However, only the one species, S. micans, was observed to breed in this
medium and in the carcasses examined.

Silpha micans Fabricius
DESCRIPTION
Adult (Fig. 15A)

Elytra blackish to blackish brown with rest of body, including legs and an-
tennae, blackish blue, in some cases even iridescent blue; legs perhaps more
bluish brown; eyes black. Integument leathery, elytra not covering three to four
terminal abdominal segments. Each elytron with three almost indistinct longi-
tudinal rugae, of which the exterior one is the most prominent and reaches only
to about the posterior one-third of the wing cover. Tibiae with numerous spines.
Wings well developed. Antennae with three-segmented club. All tarsi five-
segmented. Length 12,6-13,8 mm.

Widely spread in the Subsaharan region (Jeannel & Hatch 1928).

Larva (Fig. 15C)

General description of the larva of a Silpha sp. is given by Dorsey (1940).
Depressed dorsoventrally and wider in front than behind. Thoracic terga rounded
laterally. Abdominal terga, except ninth and tenth, with anterolateral angles

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

rounded, posterolateral angles acute, and lateral and posterior borders furnished
with setae, some of which (particularly lateral ones) are more strongly devel-
oped. Head narrower in front than behind; nearly cordiform. Nine pairs of spi-
racles situated ventrally, with mesothoracic one largest; first abdominal pair
somewhat larger than the other abdominal spiracles; all oval or annular; meso-
thoracic pair (Fig. 15P) furnished with two setae each, situated on internal side.
Colour dark brown to blackish brown, particularly posterior third of each seg-
ment. Integument leathery and covered with small tubercles and medium-long
setae (Fig. 15S) most of which are cleft at tip, or even cleft two or three times.
Sculpture on thoracic terga present as wide, irregular reticulation, particularly
on the anterior part of the pronotum. Ecdysial suture distinct in most segments,
except perhaps last two or three. Tenth abdominal segment almost tubular,
about as long as ninth and tapering towards apex; ninth fringed with fine setae.
Legs (Fig. 15H) well developed, slightly increasing in size from hind legs to
forelegs; coxae large, with longitudinal groove on exterior face distinct; strong
spine-like setae present, both ventrally and dorsally on tibiae, ventrally only on
femora; dorsal setae on latter short but slender; tarsungulus acute and with one
ventral and one posterior spine about half-way between base and apex. Uro-
gomphi borne on ninth abdominal segment; fairly long, appearing two-
segmented, with first segment much longer than terminal one and furnished with
spine-like setae; latter segment subdivided into a smaller proximal and larger
distal part in some specimens; with one or two pre-apical setae and with a
medium-long seta apically. Mature larvae measure about 14 mm in length.

Head capsule (Fig. 15E-G)

Slightly less than twice as wide as long. Sides very convex, epistomal and
frontal sutures clearly indicated. Distal ends of dorsal tentorial arms, although
contiguous with frontal sutures, are not in line with antennal sockets as stated by
Dorsey (1940) but further back, more or less in line with dorsal ocelli. Latter
four in number and situated on lateral margin, about half-way between base and
apex. Ventral ocelli two in number, with anterior one situated just below anten-
nal socket. Frontoclypeal suture indicated only laterally, obsolete over middle
portion. Posterior margin of head widely and shallowly emarginate dorsally. An-
tennae three-segmented, with first and apical segments of about equal length,
second one somewhat longer and bearing projection on mesal part of its apex.
Last two segments with setae. Chaetotaxy of head in most cases as illustrated.
Sculpture consisting of small tubercles arranged in form of a wide reticulation,
areas between striae minutely reticulate and shiny. Frons more rugulose, with
reduced number of tubercles. Head divided ventrally into two halves by deep,
emarginate hind border; gula short.

Labrum and clypeus

Labrum and clypeus united, almost triangular; boundary between them in-
dicated by more weakly sclerotized cuticle. Labrum more or less smooth, with

SOUTH AFRICAN ARTHROPODS 347

Fig. 15. Family Silphidae.
Silpha micans. A. Adult. B. Newly hatched larva (dorsal view). C. Adult larva (dorsal
view). D. Egg. E. Head (left lateral view). F. Head (dorsal view). G. Head (ventral
view). H. Right mesothoracic leg. I. Left mandible (dorsal view). J. Right mandible (dorsal
view). K.Epipharynx. L. Labium (ventral view). M. Left maxilla (dorsal view). N. Labium
(dorsal view). O. Left maxilla (ventral view). P. Left mesothoracic spiracle. Q. Pupa (ven-
tral view). R. Pupa (left lateral view). S. Dorsal integument showing tubercles and hair
(highly magnified).

348 — ANNALS OF THE SOUTH AFRICAN MUSEUM

few superficial rugae and minute reticulation between rugae; anteriorly bilobed
due to presence of short medial cleft; lateral margins sinuate just behind cleft.
Labrum with four strong setae on each side of which posterior one is situated on
small, lateral, triangular sclerotized area in front of clypeal border; tiny seta also
present on each side of labral cleft. Clypeus more or less rugoso-reticulate,
about as long as, or slightly longer than, labrum; trapezoidal and bearing trans-
verse row of about six setae behind anterior margin.

Epipharynx (Fig. 15K). Bilobed in front, anterior and lateral margins scle-
rotized, each lobe in front with rounded denticles, as well as triangular tooth
(Dorsey’s (1940) rectangular tooth), lateral margins each with short spine situ-
ated in line with second porous area. Median area concave, forming more or less
a lobe on each side, bearing an oval area beset with spiculi; the two quinquepor-
ous areas clearly indicated in most specimens with anterior parabolic row of fine
pores. First porous area visible as two small circles just behind triangular teeth.
Anterior to each spiculum-bearing area and just behind anterior lobes in line
with first porous area is a small transverse area with fine setae.

Mandibles (Fig. 15I-J)

Slightly shorter than cranium, fairly narrow, with broad subtriangular base
and nearly twice as long as width at base. Each mandible with an oblique, trans-
verse dorsal depression separating cutting edge from base. Molar areas absent,
cutting edge on each side comprised of two rounded teeth of which apical one is
largest. Dorsal carinae absent; mandibles rounded on lateral face, each with
single seta.

Maxillae (Fig. 15M, O)

Elongate, mala and stipes fused, lacinia and galea free only at extreme
apex; latter with tuft of very fine hairs. Lacinia with eight to nine broad palus-
like spines giving it a comb-like oblique anterior margin; mesal area of lacinia
with some minute denticles at base. Maxilla dorsally without setae, ventrally
with about five, of which one is situated half-way between galea and palpifer and
one just below palpifer. Cardo somewhat oval, with Y-shaped suture (arms of Y
pointing posteriorly) and with only one seta laterally on apex. Maxillary palpus
appearing four-segmented, with apical segment the longest; third segment shorter
than the second; first segment very small and with single seta ventrally; both
apical and penultimate segments with setae. In his study of six American
species, Dorsey (1940) gives the number of palpal segments as three.

Labium (Fig. 15L, N)

Different sclerites fairly easily recognizable; postmentum large and triangular
or trapezoidal, usually bearing four setae near anterior margin. Mentum small
and less sclerotized. Prementum large, consisting of basal sclerite, ventrally with
four setae of which two are situated on its anterior, less sclerotized half. Apical
sclerite with two-segmented palpi; this sclerite ventrally also with four setae.

SOUTH AFRICAN ARTHROPODS 349

Hypopharynx (Fig. 15N). Glossa deeply cleft in the middle, the two lobes
densely covered with fine hairs; hypopharyngeal scleromes (= paraglossae,
Boving & Craighead 1931) distinct; hypopharyngeal bracon well developed and
symmetrical. Superlinguae united as one broad structure.

Pupa (Fig. 15Q-R)

Exarate, clearly showing broad pronotum of the adult. Anterior margin of
pronotum with four long setae and some smaller hairs on disc; small hairs also
present on posterior part of head. Abdominal segments each with long lateral
setae; apex of abdomen with two strong setae or styli. Developing genital cap-
sule visible as broad lobe between caudal styli. Anterior border of metathorax
ventrally with two fleshy spines. Only first four pairs of abdominal spiracles large
and clearly visible. Length 11-12 mm.

Newly formed pupae are almost pure white with golden brown hairs; as
they develop they become darker, and just before emergence of beetles they are
piceous.

BIOLOGY

Beetles were collected during midsummer and midwinter and usually ap-
pear wherever carcasses and skins are found. They feed on the latter or on other
arthropods present, particularly on fly and blow-fly larvae. They are attracted to
fish-meal in large numbers and eggs were often found during January and Feb-
ruary in soil containing this medium.

Most of the eggs (Fig. 15D) collected in Claremont, Cape Town, vary from
2,20 by 1,4 mm to 2,4 by 1,6 mm and are oval, yellowish white and almost
smooth and shiny. Most eggs were laid singly in the upper few millimetres of soil
that contained fish-meal and hatched after 2-3 days during January. Shortly be-
fore hatching, two narrow dark lines separated by a thin pale line are visible
near the anterior pole of the egg.

Newly hatched larvae (Fig. 15B) are similar to the mature larvae except in
colour and size, and are almost pure white with reddish eyes, golden-coloured
setae, and two dark or piceous parallel lines on the pronotum (already visible
through the chorion in the embryo). These two black lines are visible only in the
newly hatched specimens; after the first moult the lines disappear. The newly
hatched larvae measure 44,2 mm in length and within 2-3 hours they turn to al-
most pitch black, with reddish eyes. In the laboratory the first instar lasted for
about a day. After each moult the larvae are nearly white; second and third
instar larvae usually assume their black colour after 2—4 hours, as in the first
instar.

The second moult occurs 2—5 days after the first and the larvae then mea-
sure 9-10 mm. Within 3-4 days after the second moult they are 13-14 mm long
and usually enter the soil, where they construct thin-walled clay cells in which
they remain dormant for another 3-4 days before pupation occurs. The total
lifespan of the third instar larvae varies from 6 to almost 8 days and the pupal

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

stages also last for 6-7 days, the adult beetles emerging 17—24 days after oviposi-
tion.

Newly emerged beetles are almost white with yellowish-brown pro- and
mesonotum; the head is darker and the eyes dark brown; the last abdominal seg-
ment, legs, and antennae are pale yellowish brown. After a day or so they as-
sume their normal dark coloration.

Silpha punctulata Olivier
DESCRIPTION
Adult

Dull, brownish black, about 19 mm long, somewhat more oval in outline
than S. micans, the elytra covering the abdomen. Head and prothorax fairly
densely punctate, the punctures coarser on the lateral areas of pronotum than
on its disc. The six elytral rugae very prominent, areas between them punctate,
the punctures coarser than on pronotum. Legs microscopically reticulate-
punctate or rugulose, tibiae spined as in S. micans. Antennal club not very
prominent, the three segments clearly demarcated by presence of fine, short
hairs causing pruinescence, rest of antennae slightly shiny as in the other
species. It differs from S. micans by absence of hairs on dorsal surface of prono-
tum and elytra.

This species is widespread in the Subsaharan region (Jeannel & Hatch
1928); collection records also come from the western Cape Province.

Larva (Fig. 16A)

When mature about 25 mm long and brownish black in colour. Very similar
to that of S$. micans, but somewhat more elongate; abdominal terga lobe-like on
each side, these lobes acuminate or pointed posteriorly. Meso- and metathoracic
segments with posterior angles less acute than those of abdomen; prothoracic
angles rounded; all posterior angles of body segments each with a short spine-
like seta; anterior angles, except those of pronotum, each with three to four
short setae. Ecdysial suture fairly distinct in most segments except probably last
three or four. Urogomphi shorter in relation to body length than in S. micans,
appearing two-segmented, with second segment also subdivided in most speci
mens examined and the setation similar to that of the latter species.

Legs similar to those of S. micans with somewhat more spine-like setae
present on both femur and tibia. Body integument (Fig. 16C) dorsally covered
with short scale-like setae forming a single row along posterior border of seg-
ments, except that of abdominal segment nine, which is without such a row. Se-
tae on posterior border of tenth segment longer and spine-like. The integument
differs from that of S. micans by the distinct reticulation and by the smaller and
less obvious tubercles that are present; apices of scale-like setae entire, not cleft.
Spiracles similar to those of the latter species, but lacking the two setae on inter-
nal side of the mesothoracic pair.

SOUTH AFRICAN ARTHROPODS 351

Fig. 16. Family Silphidae.
Silpha punctulata. A. Adult larva (dorsal view). B. Epipharynx. C. Dorsal integument
showing tubercles and hair (highly magnified). D. Head (dorsal view). E. Left mandible
(dorsal view). F. Left maxilla (dorsal view). G. Labium (ventral view). H. Labium (dorsal
view). I. Left maxilla (ventral view). J. Right mandible (dorsal view).

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

In small larvae measuring about 10,5 mm long that were collected during
January, all the antennal segments are about of equal length. In most of the
specimens examined there are only about six long setae on the sternal plate just
behind the head (about twenty-four in the mature specimens) and there is an
extra-long seta on each lateral margin of the central disc of the labrum. There
are also only two long setae ventrally on apical segment of labium, the other se-
tae present are short and almost spine-like. Otherwise exactly as in the mature
larva.

Head capsule (Fig. 16D)

Dark brown to blackish brown, about as long as wide or only slightly wider
than long; minutely, or rather microscopically, reticulate-punctate and some-
_what shagreened: cranial chaetotaxy as illustrated, similar to that of S. micans
but differing from it by presence of short scale-like setae and absence of long
setae on dorsal surface of epicranium, except two long setae in front of the four
dorsal ocelli just behind antennal base, and one short spine-like seta dorsolater-
ad of median ocellus. Antenna three-segmented, first segment slightly longer
than third (of about equal length in S. micans); second segment about as long as
apical one (as in the case of the latter species) and without a conical projection.
All segments with setae (in S. micans the first segment is without hairs).

Ventral side of head similar to that of S. micans, including two ventral ocel-
li, but postmentum with more setae.

Labrum and clypeus (Fig. 16D)

Labrum and clypeus united as in S. micans and similar in shape; labrum
fairly deeply emarginate on anterior border, lateral margins sinuate behind cleft;
dorsal surface with ten long setae, of which two are situated on each side on the
sclerotized triangle, just in front of clypeal border. There is no demarcation line
between clypeus and frons, except the sulcus on each side as in other species.

Epipharynx (Fig. 16B). Rather similar to that of S. micans; however, tri-
angular teeth at apex are absent. Oval area with fine hairs present just behind
anterior margin on each side of median concavity. Lateral margins with three
short almost sickle-shaped setae (absent in S. micans) and denticles present on
anterior margin of the latter species absent in this case.

Mandibles (Fig. 16E, J)

Slightly shorter than cranium, fairly narrow and more elongate than in
S. micans due to narrow bases. As in latter species, a fairly prominent oblique
depression present on each mandible separating cutting edge from base. Lateral
faces fairly rounded, each mandible with two setae of which proximal one is
longest. Two teeth of cutting edge distinct, apical one usually bilobed and larger
than the other.

SOUTH AFRICAN ARTHROPODS 353

Maxillae (Fig. 16F, I)

Similar to those of S. micans, but in this species the lacinia and galea en-
tirely fused, the demarcation line still visible, particularly on dorsal side. Galea
forming a lobe with fine golden setae as in latter species; lacinia with twelve to
thirteen palus-like spines, of which distal five are shorter with rounded apices,
the other eight or nine longer. Mesal area of lacinia also with minute spines at
base. Maxilla dorsally without setae, ventrally with about six setae as indicated
in Figure 161; exterior margin below palp with five short setae. Cardo as in
S. micans, but with one long and one short seta near apex.

Maxillary palpi four-segmented as in S. micans, first segment small with one
ventral seta, third segment longer than second (shorter than second in S. mi-
cans), apical one the longest. Both apical and penultimate segments with setae,
second with only a single dorsolateral seta.

Labium (Fig. 16G—H)

Also similar to that of S. micans, postmentum triangular with about four-
teen setae. Mentum smaller, less sclerotized and without setae. Prementum
large and with about four setae ventrally on each side of its median area. Apical
sclerite ventrally with about ten setae and with two-segmented palpi of which
basal segment is slightly longer than apical one and bearing about four setae on
exterior margin; apical segment with only a single short seta on its mesal margin
in most specimens examined. Glossa deeply cleft in middle, the two lobes dense-
ly covered dorsally with fine hairs; hypopharyngeal sclerites clearly demarcated.

BIOLOGY

Nothing is known about the life history of these beetles. Larvae were ob-
served wandering about on the ground covered with fynbos.

ACKNOWLEDGEMENTS

I am grateful to Dr S. S. Walters (Dept. of Agriculture, Elsenburg) and Dr
V. B. Whitehead (South African Museum) for reading the manuscript and for
helpful suggestions, and to Mr V. Branco (South African Museum) for prepar-
ing some of the illustrations.

REFERENCES

ArneTI, R. H. 1963. The beetles of the United States. Washington: Catholic University of
America Press.

Arrow, G. J. 1909. Systematic notes on Coleoptera of the clavicorn families. Ann. Mag. nat.
Hist. (8) 4: 190-196.

BACKLUND, H. O. 1945. Wrack fauna of Sweden and Finland. Ecology and chorology. Opusc.
ent. 5: 1-236.

BERNHAUER, M. & SCHUBERT, K. 1914. Staphylinidae IV. Coleoptm Cat. 57: 289-408.

BoRNEMISSZA, G. F. 1968. Studies on the histerid beetle Pachylister chinensis in Fiji, and its
possible value in control of buffalo-fly in Australia. Aust. J. Zool. 16: 673-688.

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

Bovine, A. G. & CRAIGHEAD, F. C. 1931. An illustrated synopsis of the principal larval forms
of the order Coleoptera. Entomologica am. 11: 1-351.

Bovine, A. G. & HENRIKSEN, K. L. 1938. The developmental stages of the Danish Hydrophili-
dae. Vidensk. Meddr. dansk naturh. Foren. 102: 27-162.

Dorsey, C. K. 1940. A comparative study of the larvae of six subspecies of Siljpha (Coleoptera:
Silphidae). Ann. ent. Soc. Am. 33: 120-139.

EmDEN, F. I. Van. 1942. A key to the genera of larval Carabidae (Col.). Trans. R. ent. Soc.
Lond. 92: 1-99.

Harez, M. 1939a. The life history of Philonthus quisquiliarius Gyllh. Bull. Soc. ent. Egypt 23:
302-311.

Harez, M. 1939b. The life history of Sphaeridium scarabaeoides L. Bull. Soc. ent. Egypt 23:
312-318.

HAFEZ, M. 1939c. The external morphology of the full-grown larva of Oxytelus latiusculus Kr.
Bull Soc. ent. Egypt 23: 333-338.

Harez, M. 1939d. The external morphology of the full-grown larva of Cercyon quisquilius L.
Bull. Soc. ent. Egypt 23: 339-343.

Hinton, H. E. 1945. A monograph of the beetles associated with stored products 1. London: The
Trustees, British Museum (Nat. Hist.); Norwich: Jarrold & Sons.

JEANNEL, R. & Hatcn, M. H. 1928. Silphidae. Coleoptm Cat. 95: 63-244.

KasuLe, F. K. 1968. The larval characters of some subfamilies of British Staphylinidae (Col-
eoptera) with keys to known genera. Trans. R. ent. Soc. Lond. 120: 115-138.

KAUFMANN, R. R. U. 1937. Investigations on beetles associated with carrion in pannal ash near
Harrogate III. Entomologist’s mon Mag. 73: 268-272.

KniscH, A. 1924a. Neue Afrikanische Palpicornier. Wien. ent. Zig 41: 17-24.

KniscH, A. 19246. Hydrophilidae. Coleoptm Cat. 79: 1-306.

Mor ey, C. 1907. Ten years’ work among vertebrate carrion. Entomologist’s mon. Mag. 43:
45-51.

NuorteEva, P. 1970. Histerid beetles as predators of blowflies (Diptera, Calliphoridae) in Fin-
land. Annls zool. fenn. 7: 195-198.

PERINGUEY, L. 1896. Descriptive catalogue of the Coleoptera of South Africa. Trans. S. Afr.
phil. Soc. 7: 99-623.

Prescott, R. T. M. & MILLER, L. W. 1937. Codlingmoth and William pears. Control methods
investigated. J. Dep. Agric. Vict. 36: 479.

RICHMOND, E. A. 1920. Article 1. Studies on the biology of the aquatic Hydrophilidae. Bull.
Am. Mus. nat. Hist. 42: 1-94.

SCHEERPEL7Z, O. 1973. Coleoptera: Staphylinidae. S$. Afr. anim. Life 15: 43-394.

THEROND, J. 1956. Histeridae. S. Afr. anim. Life 3: 295-303.

ABBREVIATIONS
acc acrocercl
adn adnasale
ant antenna
bt teeth on base of labium
car cardo
carc carina (see also ck)
ce cercus (see also ur)
ck cervical keel (see also carc)
cly clypeus
col collum
c & st cardo and stipes
doc dorsal ocelli
dta dorsal spot of distal tentorial arms
ep epicranium

epc epipharyngeal cleft

SOUTH AFRICAN ARTHROPODS 355

f frons

fa finger-shaped air-tube of biforous spiracle
fe femur

g developing genital capsule
ga galea

gl glossa

gs gular suture

gu gula

hb hypopharyngeal bracon
hps hypopharyngeal sclerome
hy hypostomal margin

il inner lobe

jx juxta

] ligula

fab) - labium

labr labrum

lac lacinia

Ict ligular cleft

Im prostheca

Ip labial palp

ls labial stipes

m mentum

mi, M2 molar area

ma mala

mcc mesocercus

mel mesothoracic leg

ml median lobe

mila superlinguae/maxillulae
mp maxillary palp

mx maxilla

mxs maxillary articulating area
n neck

na nasale

oc ocellus

of occipital foramen

ol outer lobe

par parietalia

pec procercus

pen penicillus

pf palpifer

pfg palpiger

pg pregula

pm (1 pm, 2 pm) prementum

1 ppa first porous area

2 ppa second porous area
ppm postmentum

pre preartis

prp parabolic row of pores
prvt praeventrite

pvt postventrite

qppa quinque porous area
ret retinaculum

rs triangular tooth

S] sensory appendix

sm submentum

spir spiracle

st maxillary stipes

t tarsungulus

356

tits
ta
tb
tpp
tr
ur
VOC
vt

ANNALS OF THE SOUTH AFRICAN MUSEUM

scissorial teeth

tarsus and claw

tibia

tentorial pit

trochanter

urogomphus (see also ce)
ventral ocellus

ventrite

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

‘An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name ‘(and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text

’ ‘

eo. -... the Figure depicting C. namacolus ...’; *. . . in'C. namacolus (Fig. 10) ..

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

A. J. PRINS

MORPHOLOGICAL AND BIOLOGICAL

NOTES ON SOME SOUTH AFRICAN
ARTHROPODS ASSOCIATED

WITH DECAYING ORGANIC MATTER

PART 2

THE PREDATORY FAMILIES CARABIDAE,
HYDROPHILIDAE, HISTERIDAE, STAPHYLINIDAE
AND SILPHIDAE (COLEOPTERA)

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