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96 PART 6 APRIL 1986 ISSN 0303-2515

OF THE SOUTH AFRICAN

MUSEUM

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FiscHER, P. H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 96 Band

April 1986 April

Part 6 Deel

MORPHOLOGY, SYSTEMATICS, AND

VARIABILITY OF THE SOUTHERN

AFRICAN SOFT CORAL ALCYONIUM

VARIABILE (J. STUART THOMSON, 1921)

(OCTOCORALLIA, ALCYONIIDAE)

GARY C. WILLIAMS

Cape Town Kaapstad

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Fig. 1. Living colonies of Alcyonium variabile. A. Red and yellow morphs. B. Peristome from

colony of yellow morph. C. Underwater photograph showing two colonies of the red morph.

D. Underwater photograph showing red and yellow morphs in close proximity. E. Red

morph. F. Yellow morph. G. Red, yellow, and pink variety from the eastern Cape Province.

H. Three polyps from capitulum of red morph. I. Detail of capitulum of yellow morph.

MORPHOLOGY, SYSTEMATICS, AND VARIABILITY OF THE

SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE

(J. STUART THOMSON, 1921) (OCTOCORALLIA, ALCYONIIDAE)

Gary C. WILLIAMS

Marine Biology Department, South African Museum, Cape Town

(With 15 figures and 1 table)

[MS accepted 7 October 1985|

ABSTRACT

The fungiform alcyoniid soft coral, Alcyonium variabile, is shown to be widely distributed in

the colder waters of southern Africa at depths of 13-468 m. New phenotypes from a shallow-

water population found off the Atlantic side of the Cape of Good Hope Peninsula add further

evidence of extreme intraspecific colour variation. Aspects of morphological variation, anatomy,

colour polymorphism, sclerite structure, and geographic distribution are described. The

systematic status of the species and its relations to other species of Alcyonium as well as other

alcyoniid genera are discussed. Alcyonium fungiforme Tixier-Durivault, 1954, A. luteum Tixier-

Durivault, 1954, and Metalcyonium variabile Thomson, 1921, are considered synonyms of

A. variabile.

CONTENTS

PAGE

Ji MEROGUCTNOLND 33:0 als Sig! Goce ety aOR te nee eet ee EA een er 241

Wiaterralie xammniine Cuatiny nrit te ered Mate tn ew taie wees eat hibdede bu Ales 242

DCSCHIPUOME eerie em Weire tt Seah Nt. ae sae ook 243

Morpholoeysandsclentess wanton og re ae nt eee ee eee 243

AENOBMUOLTINT.ed.her 15 cig ck ae ese a See Tol ok eee a eg rea MS

WaniabilttvrandicoloranOnye ie. fo) We bb ec cee De bee et SS)

Distmbutiomandhabitate. 2.120 06.40s 40s fees ce ae ea ce dace U9)

| DIIVEQISSIOID: cesta cod Seaton, cleo sea au ae LAT ca le Ae ssh Gr Raia a ea 259

FAISLOGUCAIISUTVICW ts ee eRe Soe hike ee yes ee ime eae was 29

Sy StemAaliCstatiGne: sete s Sy Ne co cnle. Meek oat «ccs Palas 262

SANCTION 2 ais ououdes achat SG Le cnoes Cc eee Aen an ra? eee 264

Comparison with Alcyonium paessleri May, 1899 ............. 266

ASO E OMANI Me Heer Pratt e Ho ite eee eGo wa os Rie 267

PNCKMOWAC CP EMICM USM emia ene et EE IN ee 268

INCKEN CI CCS remeron LR ON lek sean aig aon 269

INTRODUCTION

Alcyonium variabile (J. Stuart Thomson, 1921) is a highly variable, often

beautifully coloured soft coral from southern Africa. The history of the literature

pertaining to this species, as well as to the genera Alcyonium, Metalcyonium,

Bellonella, Cactogorgia, and Nidalia, has been plagued with considerable

confusion. This is partially due to an often high degree of intraspecific variation,

241

Ann. S. Afr. Mus. 96 (6), 1986: 241-270, 15 figs, 1 table.

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

the dubious nature of the original descriptions of some genera such as

Metalcyonium, the lack of attention by some previous investigators to correct

identification of material or accurate comparison with other known species, and

the lack of clear and consistent morphological distinctions between nominal

genera.

Previously published accounts have been purely taxonomic in nature;

virtually nothing has been recorded on other aspects of the biology of this species.

Previous workers relied on preserved material that was either dredged or trawled

from deep water. Observation of subtidal octocorals in their natural habitats was

technically unfeasible. Study of living colonies in situ was made possible by the

advent of SCUBA, but still very little is known about the ecology, natural history,

and other aspects of the biology of most octocoral species.

Alcyonium variabile has previously been known only from depths exceeding

47 m. A survey of the shallow sublittoral regions of the western side of the Cape

Peninsula by use of SCUBA has revealed a presumably large but previously

unrecorded polymorphic population. This paper presents an examination of

various aspects of the biology of this species from southern Africa, together with

an assessment of its great variability. An examination of the systematics of the

species and a discussion of the status of related genera is also presented.

Alcyonium variabile has been confused in the past with other species of

Alcyonium such as A. paessleri May, 1899, from Patagonia and Antarctica.

These species are here considered separate and a comparison of the two is

presented.

This paper attempts to unify the many known colour varieties of Alcyonium

variabile (many of which were previously assigned specific status) into a single

highly variable species based on consistencies in sclerite distribution and form as

well as other comparative morphological aspects.

The material is deposited in the South African Museum, Cape Town (SAM).

MATERIAL EXAMINED

SAM-—H3165 and H3166: 10 specimens, 23 m depth, Hottentots Huisie,

western side of Cape Peninsula (33°59'S 18°21'E); 15 June 1983; coll. W. R.

Piltved) SCUBAS

SAM-—H3167: 12 specimens; 13-16 m depth, Hottentots Huisie, western

side of Cape Peninsula (33°59’S 18°21’E); 18 August 1983; coll. G. C. Williams,

SCUBA.

SAM-—H3602: 1 specimen; 90 m depth, off East London (33°12’S 28°01'E);

17 July 1984; coll. G. C. Williams (R.V. Meiring Naude, XX 51), dredge.

SAM-—H3271: 1 specimen; 26 m depth, off Llandudno, western side of Cape

Peninsula (34°01’S 18°20’E); 4 February 1984; coll. G. C. Williams, SCUBA.

SAM-—H1042: 2 specimens; 148-159 m depth, 16 km SW of Cape Point

(34°27'S 10°23'E); 28 October 1903; SS Pieter Faure survey, PF 18171, large

trawl.

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 243

SAM-—H895: 5 specimens; 82 m depth; 10 km E of Cape Morgan (32°44'S

28°30'E); 13 August 1901; SS Pieter Faure survey, PF 13388A, dredge.

SAM-—H1040: 8 specimens; 141 m depth; 17 km E of Cape Morgan (32°44’S

28°30'E); 26 July 1901; SS Pieter Faure survey, PF 13174, dredge.

SAM-—H924: 5 specimens; 86 m depth; 36 km SE of Tugela River mouth,

Natal (29°20'S 31°30’E); 29 January 1901; SS Pieter Faure survey, PF 11537,

large dredge.

SAM-—H3245: 2 specimens; 168m depth; 30km S of Cape Hangklip

(34°39'S 18°42’E); 10 February 1948; coll. University of Cape Town Ecological

Survey Collection, Station AFR 882J, dredge.

SAM-—H883: 7 specimens; 95 m depth; 18 km S of Knysna Heads (34°10’S

23°15'E); 2 July 1902; SS Pieter Faure survey, PF 15291, shrimp trawl.

DESCRIPTION

Class ANTHOZOA Ehrenberg, 1834

Subclass OCTOCORALLIA Haeckel, 1866

Order ALCYONACEA Lamouroux, 1816 (emended by Verrill, 1866;

Bayer, 1981)

Family Alcyoniidae Lamouroux, 1812

Genus Alcyonium Linnaeus, 1758

Alcyonium variabile (Thomson, 1921) comb. nov.

Figs 1-15

Alcyonium antarcticum (non Wright & Studer, 1899) Hickson, 1900: 73.

Alcyonium (Metalcyonium) patagonicum (non May, 1899) Kikenthal, 1906: 47 (partim).

Metalcyonium patagonicum (non May, 1899) Thomson, 1910: 562.

Metalcyonium variabile Thomson, 1921: 152; 1924: 47, 69.

Metalcyonium variabile var. molle Thomson, 1921: 162; 1924: 47, 69.

Metalcyonium variabile var. durum Thomson, 1921: 165; 1924: 47, 69.

Alcyonium paessleri (non May, 1899) Molander, 1929a: 50; 19296: 4 (partim).

Alcyonium fungiforme Tixier-Durivault, 1954: 385.

Alcyonium luteum Tixier-Durivault, 1954: 388.

Morphology and sclerites

The colonies examined range in total length from 8 to 70 mm. They are

mushroom-shaped; composed of a spherical capitulum and a conspicuous stalk

(Fig. 1). Both parts are separated by a clear delimitation (Fig. 2A).

The globular capitulum is wider than the stalk. The surface layer is filled with

closely-set sclerites. They vary from stout coarsely tuberculate rods less than

0,22 mm long to capstans less than 0,08 mm long (Figs 3A, 4A—D). Some rods

may be clubbed. Capstans may predominate in some colonies. Some capstans

may be modified and sharply pointed tubercles. The sclerites from the interior of

the capitulum are usually arranged parallel to the gastric cavities. They are

ANNALS OF THE SOUTH AFRICAN MUSEUM

244

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THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 245

AOABRSESE

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Fig. 3. Alcyonium variabile. A. Capitulum sclerites from surface coenenchyme. B. Capitulum

sclerites from inner coenenchyme.

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Alcyonium -variabile. Scanning electron micrographs of capitulum sclerites. A-D. Sclerites of

outer coenenchyme. A. Spindle, 0,17 mm. B. Club, 0,12 mm. C. Detail of middle portion of club

in B, 0,03 mm. D Capstan, 0,07 mm. E-F. Slightly clubbed rods of the inner coenenchyme.

EB) 015 mm: EF) 0)25 mm:

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 247

predominantly rods, sometimes slightly clavate, often with coarse to ornate

tubercles, and vary from approximately 0,07 to 0,27 mm in length (Figs 3B,

4E-F). Modified, sharply pointed capstans may also occur.

The polyps are restricted to the entire surface of the capitulum. When fully

expanded they may reach a length of 12 mm. In expanded condition the polyp

shows two differentiated regions: the basal or proximal region and the distal

anthocodia (Fig. 5A). The proximal portions possess varying concentrations of

capstans or double cones, from 0,04 to 0,12 mm in length. These sclerites are

relatively consistent in form and size (Figs 6C, 7C—F). Partially retracted polyps

can form rounded calyx-like protuberances on the surface of the capitulum;

however, these are not permanent calyces since they are not rigid and are capable

of total retraction into the capitulum (Fig. 1G). The anthocodia consists of three

parts: the neck zone or introvert, the anthocodial wall with crown and points, and

the tentacles (Fig. 5A). Sclerites of the crown, points, and tentacles are rods and

spindles of varying length and coarseness (Fig. 6A). The neck zone is usually

unarmed and translucent—the pharynx and the mesenterial filaments are plainly

visible through it. In one red colony from the western Cape Peninsula, the polyps

possess sparsely scattered capstans in the neck zone. The crown consists of

6-12 tiers of very slender sclerites. These are finely tuberculated needles up to

0,6 mm in length (Fig. 8A, E). Superposing the crown are eight points, each

consisting of numerous spindles of which the undermost are arranged en chevron;

more distally they are longitudinally placed (Fig. 5A). The point sclerites are

mostly very similar to those of the crown, but distally some may be shorter and

more coarsely tuberculated (Fig. 8B—D). The tentacles are narrow and reach a

length of 2,5 mm. On each side they bear one row of approximately 12 pinnules

(Fig. 5B). The backs of the tentacles are armed proximally with the uppermost

point sclerites, and distally with short spindles or coarse rods usually less than

0,13 mm in length (Figs 6A, 8F—G). These sclerites are not arranged in any

distinctive alignment or pattern (Fig. 5A). Rod-like sclerites may also occur in

between the bases of most pinnules (Fig. 5B). Similar short spindles or rods may

occur along the grooves of the peristome in a radial pattern of eight lines

extending from the mouth to the areas between the bases of the tentacles

(Fig. 5C). The wall of the pharynx is strongly impregnated with many short

spindles or rod-like sclerites. These are coarsely tuberculated and usually less

than 0,14 mm in length (Figs 6B, 7A—B).

The stalk is barren, being completely devoid of anthocodiae. The height of

the stalk is variable (Fig. 12). The base may produce finger-like or flattened

projections forming a holdfast. The surface layer of the stalk contains densely set

sclerites that are almost exclusively capstans varying in length from 0,05 to

0,10 mm (Figs 9A, 10A—C). In the interior of the stalk there are mainly large

stout spindles or clubbed forms that are ornately tuberculated. They vary from

0,10 to 0,28 mm in length and up to 0,06 mm in width. Modified, pointed

capstans may occur rarely (Figs 9B, 10D-G).

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Fig. 5. Polyp armature of Alcyonium variabile. A. Expanded polyp showing capstans forming

calyx-like proximal region, short spindles of the pharynx, and long spindles of the anthocodia;

total length of polyp 9 mm. B. Single tentacle showing rod-like sclerites at the base of the

pinnules; total length of tentacle 2,5 mm. C. Peristome showing alignment of rod-like sclerites;

diameter of disc 2 mm.

a—anthocodia, cr—crown, mf—mesenterial filament, nz—neck zone, ph—pharynx, pp—proximal

region of polyp, ps—points.

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 249

Fig. 6. Alcyonium variabile. A. Polyp sclerites from crown, points, and tentacles of anthocodia.

B. Polyp sclerites from wall of pharynx. C. Polyp sclerites from wall of proximal region of polyp.

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

rig. 7. Alcyonium variabile. Scanning electron micrographs of sclerites from pharynx and calyx-like base

of polyp. A-B. Spindles of pharyngeal wall. C-F. Capstans from wall of proximal region of polyp.

\. Scale between squares 30 wm (0,12 mm total length). B. Scale between squares 10 wm (0,11 mm

total length). C—E. 0,07 mm. F. 0,09 mm.

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE

Fig. 8. Alcyonium variabile. Scanning electron micrographs of anthocodial sclerites. A. Needle from crown,

0,60 mm. B-C. Spindles from points. B. 0,21 mm. C. 0,23 mm. D. Detail of central portion of spindle in C;

length of photograph 0,09 mm. E. Needle from crown, 0,37 mm. F-G. Rod-like sclerites from distal portion of

tentacles. F. 0,12 mm. G. 0,12 and 0,09 mm.

By ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Alcyonium variabile. A. Stalk sclerites from surface coenenchyme. B. Stalk sclerites from

interior coenenchyme.

Anatomy

A 35 mm-long red-purple colony collected from the sublittoral of the Cape

Peninsula (SAM—H3166) was sectioned longitudinally. The epidermis is a thin

transparent layer, which separates readily from the underlying mesogloea. The

coenenchyme consists partially of an outer area approximately 0,16 mm thick.

This region is very densely imbedded with primarily red capstans, giving a rough

textured appearance to the colony. Below the outer layer is the inner

coenenchyme, which fills the interior of the colony between the tube-like polyps.

The colour of this region is pink due to the combined effect of white mesogloea

and gastrodermis and light-red sclerites. Numerous spindles and clubbed forms

are contained in the region just below the outer coenenchyme. These sclerites

become less dense toward the interior of the capitulum and stalk regions. The

interior coenenchyme adjacent to the gastric cavities is profusely set with clear to

light-reddish spindles approximately 0,22 mm in length. Many of the gastric

cavities contain spherical ova varying in diameter between 0,06 and 0,12 mm. The

gastrodermal solenial network is readily visible in the inner coenenchyme below

the outer layer of the capitulum. The long thin gastric cavities extend to the base

of the stalk. Anthocodiae are in varying states of contraction and retraction. The

eight-ridged protuberances of the calyces of some polyps are readily visible

(Fig. 11).

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 293

Fig. 10. Alcyonium variabile. Scanning electron micrographs of stalk sclerites. A-—C. Capstans from outer

ceenenchyme. A. 0,06-0,13 mm. B.0,07 mm. C.0,08mm. D-G. Sclerites from inner coenenchyme.

D. Modified capstan, 0,10 mm. E. Clubbed spindle, 0,20 mm. F. Spindle, 0,20 mm. G. Club, 0,17 mm.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Fig. 11. Anatomy of Alcyonium variabile. A. Longitudinal section through capitulum of 35 mm-

long colony from the Cape Peninsula. B. Area of section (shown enlarged from A) from the entire

colony. C. Longitudinal section of a retracted polyp; sclerites of interior coenenchyme omitted;

width of diagram 4 mm. D. Detail of section through coenenchyme of capitulum showing thin outer

layer of capstans and interior of spindles; diameter of diagram 1 mm.

cap—capitulum, cr—crown, e—epidermis, gc—gastric cavity, h—holdfast, ic—interior coenenchyme,

mf—mesenterial filament, nz—neck zone, o-ova, oc—-outer coenenchyme, ph—pharynx, pp—proximal

region of polyp, rp—-retracted polyp, s—solenial tubes, st-stalk.

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 5) 9)

Variability and coloration

The high degree of variability found in many sedentary organisms, such as

octocorals, due to isolation of populations, lack of mobility and ephemeral larval

stages, has been noted by several authors (Thomson 1921: 166; Hickson 1907a:

25-~36).

Alcyonium variabile is remarkable in that the colonies often resemble

young mushrooms of the genus Amanita in shape, appearance, and even colour

in some instances. The most extraordinary aspect of the nature of this species

is its extreme intraspecific variability, particularly in regard to external color-

ation. Stalk development is also variable. Sclerite form and distribution are

the most consistent and least variable characteristics between geographically

isolated populations or individual colonies, and serve to unify the various

widespread populations into one highly variable species. These southern African

populations exemplify a wide range of variability in regard to coloration

(Table 1).

Most southern African material is known to have a well-developed barren

stalk, the length of which may exceed twice the diameter of the capitulum. In

some colonies from the southern Transkei and East London regions the stalk is

reduced in size: the spherical capitulum arises from a very short stalk with a

spreading base (Fig. 12). The development of the stalk is considered a variable

feature in other species as well. In the original description of Metalcyonium

capitatum, Pfeffer (1889: 50) states that the colony has the appearance of a young

mushroom, being short-stemmed with a bulbous head, or the stalk is entirely

missing and the entire colony is of a head-like shape.

It appears that both continuous and polymorphic colour variation are

exhibited in populations of Alcyonium variabile. The recently discovered

localized populations of A. variabile from the western side of the Cape Peninsula

are remarkable in that they exhibit distinctive polymorphism. The individual

colonies of the populations are morphologically very consistent but are sharply

differentiated and discontinuous in colour due to the striking permanent

coloration of calcium carbonate sclerites that are embedded in the outer

coenenchyme.

The populations are obviously dimorphic as two clearly delineated colour

morphs can be recognized (Fig. 1):

Yellow morph. The colonies are a golden-yellow to yellowish-orange. In

some colonies the stalk may have a slightly rose tint due to some sclerites of the

outer coenenchyme being red. The armature of the anthocodial crowns and

tentacles is usually composed of deep orange-yellow sclerites.

Red morph. The colonies are red to mauve to vivid reddish-purple. The

armature of the anthocodiae within a given colony may be composed of either all

red or all yellow sclerites. The pharyngeal sclerites may also be yellowish or light

red; these combined give the anthocodiae as a whole a pinkish-white or yellowish

appearance.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Colour variability in Alcyonium variabile.

mauve-red to

pinkish

W side of Cape Peninsula, | mauve to red mauve to red

Cape Agulhas region

NW side of Cape Peninsula | orange-yellow golden-yellow to orange to pinkish-

SW of Cape Point

S of Cape Hangklip,

Knysna

East London

Cape Morgan

Sandy Point

Port Grosvenor

Durban, Cape Morgan,

East London

Tugela River,

Cape Morgan

Tugela River, Durban,

Port Shepstone, Cape

Morgan

white

rose

yellow

brownish-white

mauve-pink

pinkish-tan

orange

yellow

white to yellow

red

greyish-white

red

yellow

pale pink

red

yellow

orange-red

salmon-pink

bright yellow

yellow

red

yellow

red to red-violet

red

yellow

red

yellow

orange-yellow

white

yellow

brownish-white

mauve-pink

pinkish-tan

red

white

yellow

white to yellow

yellow

greyish-white

pink

orange

pale pink

yellow

red

yellow

salmon-pink

light orange

salmon-orange

yellow

yellowish-white

orange to red

white to yellowish

white to reddish

yellow

white

orange to red

orange

white

brownish-white

white to yellowish-

white

brownish-white

pinkish-tan

yellowish

tan

pink

yellowish

pinkish-white to

reddish

greyish-white

pink

pinkish-white

white

reddish

yellow to white

salmon-pink

pinkish-white

salmon-orange

yellow

pinkish-white

yellowish-white to

rose

white to yellowish

white to pink

yellow

white

white to pink

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 257

Fig. 12. Variability of colony shape and stalk development in Alcyonium variabile. Polyp-bearing

portion (capitulum) is differentiated from stalk or base.

cap-—capitulum, h—holdfast, p—polyp, st—stalk.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

In both morphs, within a given colony, the calyces and outer coenenchyme of

the capitulum and stalk are uniformly coloured. Thus, a colony is entirely red or

yellow. The sclerites in both morphs exhibit tints of red or yellow: sclerites of

other colours have not been observed.

The individual colonies of the yellow or red morphs can occur sympatrically

and may grow as little as 10 mm apart at depths of 13-30 m. Individual colonies

have been observed to be as much as 55 mm in total length. At Hottentots Huisie,

these two morphs are apparently equally represented. At Llandudno and

Kommetjie (34°09’S 18°18’E), however, the red morph is frequently encountered

while the yellow morph has not been observed. The yellow morph seems to be

confined to the Atlantic side of the Cape Peninsula, while the red morph is also

common off the Cape Agulhas region between 24 and 54 m in depth.

A white form, known only from deeper water (150-163 m) off Cape Point,

has not been found in association with either of the other two morphs. It is known

only from five large colonies (52-71 mm in total length) described by Thomson

(1921: 165), collected in 1903 by trawl. Thomson relegated this form to the

complex he named Metalcyonium variabile var. molle. The colonies are a uniform

bright white as sclerites from all parts of the colony are white.

All observed colonies of the dimorphic populations, as well as the white

form, have in common a well-developed and conspicuous stalk. The stalk is at

least as long as the height of the capitulum and is commonly up to twice its height.

One colony of the white form has a stalk length three times the height of the

capitulum. This is the maximum known development of the stalk for the

species.

The numerous colonies that have been collected from Cape Point to Natal

exhibit a more continuous variation. The many populations sampled by the

SS Pieter Faure surveys from 1898 to 1906 and the R.V. Meiring Naude dredge

surveys from 1981 to 1985 exhibit the following differences from the dimorphic

populations and the white form of the western Cape Peninsula: (a) none of the

specimens exceed 40 mm in total length; (b) stalk length is variable, from less

than half the height of the capitulum to approximately twice the height;

rarely monochromatic, usually bicoloured or tricoloured. The stalk, capitulum,

and proximal portions of polyps often exhibit different colours. A wide range of

colour combinations exists (Table 1). The stalk varies from whitish to pale rose to

yellowish, while the surface of the capitulum can be white or greyish-white,

yellow, orange, or red. The calyces of an individual colony are either ail red or all

yellow. Some very striking and beautiful combinations are thus evident.

The colour of all colonies observed is due to the permanent coloration of the

sclerites, for the colonies contain little if any alcohol-soluble pigment. Colour has

been observed to be well preserved in material maintained in ethanol for over

eighty years. Colour patterns are extremely varied and often very striking.

The colour of the individual sclerites ranges from rich orange-yellow, light

lemon-yellow, deep red to maroon or mauve, to white or almost translucent.

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 259

Some spindles are observed to be reddish towards one end and transparent

towards the opposite end.

Capitulum or stalk sclerites of the outer coenenchyme are often quite vividly

coloured, while those of the interior coenenchyme are only slightly tinted with red

or yellow, or are often colourless and transparent.

The high degree of variability in the collected material led Thomson (1921)

to propose the name Metalcyonium variabile for this species and to name two

varieties: (1) M. variabile var. molle for colonies with stalk surface of a ‘leathery

consistency’ and sclerites primarily of capstans; and (2) M. variabile var. durum

with stalk surface of a ‘much harder and somewhat stony consistency’ and

sclerites of capstans, spindles, rods, and clubs. Thomson also recognized a range

of intermediate cases that were not readily identifiable with either subspecies.

Because of the intermediate nature exemplified by some colonies and since the

populations exhibit continuous and extreme variability, the recognition of formal

taxonomic variations or subspecies must be disregarded.

Distribution and habitat

Alcyonium variabile is known only from the following South African

localities: off East London (Hickson 1900, as A. antarcticum); many stations from

Cape Point to central Natal (Thomson 1910, as Metalcyonium patagonicum; 1921,

1924, as M. variabile); Cape Infanta (Molander 1929a, 1929b, as A. paessleri);

Cape Peninsula region (Tixier-Durivault 1954, as A. fungiforme and A. luteum);

western side of Cape of Good Hope Peninsula, southern Natal and Transkei coast

to eastern Cape Province (present study) (Fig. 13).

Alcyonium variabile is a sublittoral benthic organism of shallow to mid-water

and is known from a depth range of 13-468 m. It is recorded by Hickson (1900)

and Thomson (1910, 1921) at depths of 47-468 m. Dimorphic populations of red

and yellow morphs have recently been observed from the western Cape Peninsula

between depths of 13 and 20 m. Thomson (1921: 165) records an entirely white

morph from 146 to 159 m.

Colonies are usually found attached to hard substrata such as shells and rocks

or relatively soft substrata such as encrusting sponges. They are commonly

observed growing on vertical rock walls amongst sponges, gorgonians, bryozoans,

tunicates, and other soft corals.

Colonies have been observed with polyps fully expanded or in varying states

of retraction during daylight hours.

DISCUSSION

HISTORICAL SURVEY

Pfeffer (1889: 49) established the genus Metalcyonium to include two new

species of supposedly clavate and capitate, monomorphic alcyoniids from South

Georgia Island in the southern Atlantic: M. clavatum and M. capitatum.

Unfortunately, as Utinomi (1964: 7) pointed out, Pfeffer did not designate either

ANNALS OF THE SOUTH AFRICAN MUSEUM

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THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 261

of these divergent forms as the type-species of the genus, and gave a vague

diagnosis.

May (1899: 6, 8) described two new species from Patagonia: Alcyonium

paessleri for a globular form and Metalcyonium patagonicum for a clavate species.

He also amended Pfeffer’s original definition of the genus. Tixier-Durivault

(1970: 148) altered the name of M. patagonicum to Bellonella patagonica for

northern Patagonian material.

According to Thomson (1910: 564), Hickson (1900: 73) apparently misiden-

tified two specimens of capitate forms from the East London region of South

Africa as Alcyonium antarcticum Wright & Studer, 1889. Kukenthal (1906: 47)

considered Hickson’s material to possibly be young colonies of M. patagonicum.

Burchardt (1903: 673) described a fungiform-capitate species from the Malay

Archipelago, M. molle. Thomson (1921: 162) later identified material from Natal

as conspecific with this species.

Ktkenthal (1906: 42) revised the genus Alcyonium Linnaeus, 1758, and

relegated Metalcyonium to subgeneric status. However, this was not widely

accepted by workers after that time. Kukenthal considered Alcyonium to be

composed of three subgenera of monomorphic soft corals, differentiated by

colonial growth habit: Alcyonium for all forms with a massive upright stalk and

more or less lobate capitulum; Metalcyonium for unbranched, cylindrical or

conical forms; and Erythropodium for membranous, broadly spreading forms.

This redefinition ignored capitate forms and left their position doubtful.

Broch (1912: 21) changed the subgeneric name Alcyonium to Eualcyonium,

and Kikenthal (1916: 174) proposed the name Parerythropodium for membra-

nous forms since Erythropodium Kolliker, 1865, was previously applied to an

unrelated but presumably superficially similar genus of the scleraxonian family

Anthothelidae (Briareidae of Kiikenthal 1916: 172). Kiikenthal (1906: 48) also

described Metalcyonium novarae, a cylindrical species from the Cape of Good

Hope (False Bay), and Thomson (1921: 167, 172) described two other cylindrical

forms: M. lanceatum and Sinularia unilobata, both from the eastern Cape

Province and Natal.

Thomson (1910, 1921) added to an already chaotic situation by describing the

common and variable South African species under two different names: first

(following Kikenthal 1906: 47) M. patagonicum (1910: 562) and then later as two

varieties of a new species, M. variabile var. molle (1921: 162) and M. variabile

var. durum (1921: 165). Molander (1929a: 50; 1929b: 4) considered Thomson’s

South African material to be identical with the Patagonian and Antarctic species,

Alcyonium paessleri May, 1899. Verseveldt (1967: 10) disagreed with Molander’s

contention and considered A. paessleri and M. variabile as separate species.

Hickson (1907b: 3) identified material from Franklin Island in Antarctica as

A. paessleri. However, Littschwager (1922: 534) considered Hickson’s determi-

nation as a misidentification and gave Hickson’s species (plus apparently similar

material from the Philippines) the new name Alcyonium equisetiforme. Litt-

schwager (1922: 522) and Roxas (1933: 356) provided keys to the 19 worldwide

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

species of Alcyonium (subgenus Eualcyonium) known at that time, and defined

A. paessleri as a globular species without a distinctive stalk.

Yamada (1950: 114, 115) described two new species of globular, capitate soft

corals from northern Japan, A. pacificum and A. muricatum, providing only very

brief descriptions and not designating type material. Uchida (1969: 397) gave a

more detailed description of A. pacificum.

Tixier-Durivault (1954: 385, 388) added two new species of capitate

monomorphic alcyoniids, described as Alcyonium fungiforme and A. luteum,

both from the region of the Cape of Good Hope.

Utinomi (1958: 111; 1964: 7) attempted to arrange some sense of order out

of a chaotic situation by suggesting that in order to validate the status of the genus

Metalcyonium, it should be restricted to markedly capitate forms and that

problematic cylindrical forms such as M. clavatum should be assigned to other

genera. This position is now held to be incorrect (Verseveldt pers. comm.;

present study).

SYSTEMATIC STATUS

The most familiar members of the genus Alcyonium Linnaeus, 1758, are

more or less lobate; the polyp-bearing portions of the colonies are divided into

varying degrees of digitate or lobate processes that arise from a common stalk or

base (Bayer 1981: 916).

However, several other additional species, which are in no way lobate and

show remarkably different growth forms, can be allocated to the genus. For

example, several species that have previously been assigned to the genus

Parerythropodium Kikenthal, 1906, do not have a defined stalk but consist

usually of a thin and pliable form that assumes the shape of the substratum that it

covers, or may be globular to irregular in shape. Groot & Weinberg (1982) have

recently shown that these membranous and encrusting species can be accommo-

dated in Alcyonium since the type-species, P. coralloides, is consistent with

characters of the genus. A number of other species are strongly fungiform in

shape with a single spherical, hemispherical, or flattened disc-shaped capitulum

arising from an unbranched stalk. Several of these have been placed in the genus

Metalcyonium by various authors following the original description of Metal-

cyonium capitatum Pfeffer, 1889 (e.g. Metalcyonium variabile Thomson, 1921;

M. molle Burchardt, 1903).

Species, or individual colonies within a species, are known that grade

between several of these disparate morphological growth forms and thus serve to

amalgamate the various species within the single genus. It appears then, that the

genus Alcyonium is a large variable complex of broad morphological scope, and

that attempts to subdivide the genus on the basis of colony shape must be

considered unjustified or tenuous at best.

Utinomi (1964) maintained that in order to validate the genus Metalcyonium,

M. capitatum should be designated as the type-species of the genus and that

Metalcyonium should be reserved for strongly capitate, mushroom-shaped forms

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 263

with globular or spherical capitulums. This differentiation of genera based solely

on colony shape or growth form is in this case difficult to vindicate, since

‘subcapitate’ forms are also known that tend to be intermediate between

digitiform and capitate forms. Colonies of Metalcyonium patagonicum May, 1899,

for example, are slightly clavate, having a distally swollen or somewhat club-

shaped form that seems to be intermediate between cylindrical and capitate.

Individual colonies of Alcyonium variabile may have thickened stalks that

approach the width of the capitulum, thus making the colony appear more clavate

than capitate. Alcyonium fauri J. S. Thomson, 1910, and A. paessleri May, 1899,

may form globular heads upon a membranous or spreading base. I therefore see

no justification for generic separation of capitate forms and feel that such forms

can be accommodated within the genus Alcyonium. Alcyonium variabile

(Thomson, 1921) should therefore be considered as a new combination of

Metalcyonium variabile Thomson, 1921. I therefore consider the following

worldwide species that possess capitate and unbranched growth forms but lack

permanent calyces, to be members of the genus Alcyonium:

Alcyonium capitatum (Pfeffer, 1889) (South Georgia Island)

(= Metalcyonium capitatum Pffeffer, 1889)

Alcyonium laeve Tixier-Durivault, 1955 (Gabon)

Alcyonium molle (Burchardt, 1903) (Amboina)

(= Metalcyonium molle Burchardt, 1903)

Alcyonium muricatum Yamada, 1950 (Japan)

Alcyonium pacificum Yamada, 1950 (Japan)

Alcyonium planiceps Williams, 1986 (South Africa)

Alcyonium variabile (Thomson, 1921) (South Africa)

(= Metalcyonium variabile Thomson, 1921)

Alcyonium verseveldti (Benayahu, 1982) (Red Sea)

(= Metalcyonium verseveldti Benayahu, 1982)

Alcyonium violaceum Tixier-Durivault, 1955 (West Africa)

However, the status of unbranched, truly digitiform to cylindrical forms with

monomorphic polyps is not completely resolved as yet. These include such species

as Metalcyonium novarae Kikenthal, 1906, M. lanceatum Thomson, 1921,

M. unilobatum (Thomson, 1921) (Fig. 2B), the many species assigned to the

genus Bellonella after the description of (but not including) B. granulata Gray,

1862, and the genera Nidalia Gray, 1834, and Cactogorgia Simpson, 1907. Pfeffer

(1889: 49) originally described Metalcyonium clavatum as a clavate species. It is

actually not clavate, but rather digitiform or cylindrical in shape (Molander

1929a, pl. 4 (fig. 9); Verseveldt pers. comm.).

According to Verseveldt (pers. comm.) most of the species previously

assigned to the genera Bellonella, Cactogorgia, and Metalcyonium can probably

be accommodated in Nidalia by priority—a genus characterized by digitiform or

capitate colony shape and the possession of permanent calyces into which the

anthocodia are retractile. Most other digitiform or capitate species that do not

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

possess permanent calyces can probably be assigned to the morphologically

diverse genus Alcyonium. Both Verseveldt (Zwolle, The Netherlands) and Bayer

(Smithsonian Institution) are currently studying this matter in detail.

SYNONYMY

Alcyonium fungiforme Tixier-Durivault, 1954, was described from a single

colony collected off Cape Point, South Africa (34°33’S 18°20’E). The colony, as

described by Tixier-Durivault, has a brownish-white stalk and a white capitulum

spotted with rose-red calyces. The obviously fungiform and capitate colony shape

is distinctive. The capstans and thorny spindles figured in the original description

of A. fungiforme and considered by Tixier-Durivault (1954) to be so disparate

from those of material of A. variabile examined by Hickson (1900) and Thomson

(1910, 1921), can undoubtedly be viewed within the context of intraspecific

variation. The spindles illustrated by Tixier-Durivault (1954, fig. 3) from the

capitulum and polyps are very similar to those that have been isolated and

observed from several recently acquired colonies and from Thomson’s SS Pieter

Faure survey material. The stalk sclerites of A. fungiforme are a bit more

pronounced (Fig. 14B). However, this minor discrepancy does not justify the

status of a separate species, since the characteristic sclerite pattern and form are

consistent with A. variabile. The description of A. fungiforme is thus considered

to fall well within the realm of A. variabile; A. fungiforme should therefore be

considered a junior synonym of A. variabile.

Tixier-Durivault (1954: 388) described another new species of capitate

alcyoniid, Alcyonium luteum, from the Cape of Good Hope region (34°39’S

14°42’E). An examination of type material has shown that this species should also

be considered conspecific with A. variabile. The holotype of A. luteum is very

similar to material that Thomson (1921: 164) considered characteristic of

Metalcyonium variabile var. molle (SAM-—H883) from Knysna Heads (34°10’S

23°15'E). These colonies all have lemon-yellow capitulums with yellowish-white

to yellow stalks. The form and distribution of the sclerites are typical of

A. variabile—long spindles forming a transverse band around the base of the

crown of the anthocodia with double rows of spindles arrayed en chevron along

the axis of each tentacle; short capstans in the calyx-like base of the polyp

forming eight ridges; capstans and clubbed forms in the outer coenenchyme with

thick spindles dominating in the inner coenenchyme. The illustration of the

complete colony of A. luteum provided by Tixier-Durivault (1954, fig. 4A) does

not show the obviously capitate, spherical shape of the capitulum. Examination of

the holotype of A. luteum (SAM-—H3245), has shown it to be markedly capitate in

nature (Fig. 14A). The morphological features of these colonies also fall well

within the realm of intraspecific variation of A. variabile. As is the case with

A. fungiforme, I see no justification for considering these species as separate from

the variable A. variabile complex. If A. fungiforme and A. luteum are considered

separate species because of their slight variance from other A. variabile material,

then perhaps a dozen or more other species could also be separated. If one

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 265

0.05 mm

20mm

Fig. 14. Comparative morphology of material synonymous with Alcyonium variabile. A. Coenenchyme

sclerites and two entire colonies of A. Juteum Tixier-Durivault, 1954 (from type material). B. Entire

colony and sclerites of A. fungiforme Tixier-Durivault, 1954 (adapted from Tixier-Durivault 1954).

Stippling represents polyp-bearing capitulum; polyps not shown.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

recognizes the fact of intraspecific variation and the extreme range of variability

possible in sedentary marine organisms such as soft corals, then the coherence of

a single variable complex encompassed by A. variabile is evident.

Molander (19296: 5) maintained that Thomson’s Metalcyonium variabile

complex can be accommodated by Alcyonium paessleri. Molander based his

consideration on the presence of capstans (‘Doppelspindeln’) in the calyx and

outer coenenchyme as characteristic of the species and present in all material

from the three geographically disjunct regions (Patagonia, Antarctica, and

southern Africa). Verseveldt (1967: 10) disagreed with Molander’s contention,

acknowledging the disparate colonial growth forms of the two species in addition

to divergent characteristics of the polyps. I agree with Verseveldt’s assessment

that the Patagonian—Antarctic and southern African forms are indeed two

separate species.

Metalcyonium patagonicum May, 1899, and M. variabile have been confused

by Kikenthal (1906: 47) and Thomson (1910: 562). Kiikenthal listed Hickson’s

(1900) identification of Alcyonium antarcticum as a possible synonym of

A. patagonicum, and Thomson originally identified South African A. variabile

material as M. patagonicum. Metalcyonium patagonicum differs from the variable

A. variabile complex in its clavate growth form and the strongly clubbed nature of

the capitulum sclerites. Tixier-Durivault (1970) placed this species in the genus

Bellonella. Alcyonium patagonicum seems to be the correct binomial designation

for this form but a formal assignment must await a thorough systematic revision of

the cylindrical to clavate alcyoniids.

COMPARISON WITH ALCYONIUM PAESSLERI MAY, 1899

Alcyonium Paessleri May, 1899: 6.

Alcyonium paessleri May, 1899. Molander, 1929a: 50; 1929b: 4 (partim). Verseveldt, 1967: 7.

Alcyonium paessleri May, 1899, has subsequently been described by

Molander (1929a, 1929b) and Verseveldt (1967). May’s type-locality was the

Smyth Channel in south-western Patagonia. Molander (1929a) described material

from off Seymour Island in the Antarctic Peninsula, and Verseveldt described

four colonies from the Straits of Magellan and eastern Patagonia (Fig. 15). The

various specimens are apparently consistent in the lack of a stalk: the polyp-

bearing capitulums are globular swellings that arise directly from the attached

membranous base with no clear differentiation between capitulum and base.

They may often have short knob-like branches. The colonies are recorded as

uniformly whitish to greyish-brown in colour and up to 50 mm in height. Colonies

of Alcyonium variabile usually have well-developed stalks, are unbranched and

upright with well-differentiated capitulum. The colonies are often vividly multi-

coloured.

Although the colony shapes and growth forms of A. paessleri and A. var-

labile are strongly divergent, sclerite form and distribution are remarkably

similar. Both species have essentially the same sclerite patterns: anthocodia with

a crown and points of long, narrow spindles or needles; capstans, double cones,

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 267

Alcyonium variabile

Fig. 15. Map showing known world-wide distributions of Alcyonium paessleri and A. variabile.

or clubbed forms in the polyp bases and outer coenenchyme; and thorny spindles

in the interior coenenchyme. Sclerite form also shows considerable similarity.

The capstans, double cones, needles, clubs, and thorny spindles are similar in

structure and ornamentation in both species.

In A. variabile, the capstan-impregnated polyp bases may completely

withdraw into the capitulum, resulting in eight lines radiating from the centre.

Verseveldt (1967: 8) reported that in colonies of A. paessleri with retracted

polyps, well-defined rounded anthosteles project from the capitular surface, these

not being retractile. The aperture presents a small round hole with no radiating

pattern of lines.

ZOOGEOGRAPHY

As shown above, there is remarkable similarity of sclerite form and

distribution between A. paessleri and A. variabile. This fact led Molander (1929a,

1929b) to consider them synonymous. It can be postulated that this occurrence is

either purely coincidental, or that the two species share a common ancestry.

Powell (1951: 64, fig. A) postulated a possible benthic bridge system for the

distribution of certain prosobranch mollusc genera in the Atlantic Subantarctic

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

region. A continuous bridge between Patagonian South America and the

Antarctic Peninsula is seen as the Scotia Ridge. Similarly, a plausible but

discontinuous zoogeographical link between the Scotia Arc region and southern

Africa is shown by the African—Antarctic Ridge and Agulhas Plateau systems.

These are regions of less than 4 000 m in depth that separate various abyssal

basins of the Atlantic and Southern oceans. This hypothetical model takes into

account periodic declines in sea-level (of at least 140 m) (Dingle & Rogers

1972: 155) during Pleistocene glacial episodes.

According to Powell’s hypothesis, stenothermic species are shown to live in

relatively shallow water in cold-water polar or subpolar regions, and are linked

by deep-water distribution through temperate and tropical latitudes. The west

side of the Cape Peninsula is swept by the Antarctic-influenced Benguela

Current, which contains colder upwelled water. The eastern and southern coasts

of southern Africa are affected by the warmer water of the Agulhas Current,

which comes from tropical latitudes. Alcyonium variabile and A. paessleri are

presumably stenothermic species, which seem only to be tolerant of relatively

cold water. Alcyonium paessleri is known from 150-642 m depths; Alcyonium

variabile is also known only from sublittoral areas; it has not been recorded

intertidally. This alcyonacean is recorded in relatively shallow water (less than

40 m) only from the west side of the Cape Peninsula. It is known from all other

localities in deeper water (up to 468 m). The species is not presently known

from the east side of the Cape Peninsula in False Bay or in shallow-water coastal

regions to the east, which are influenced by the warmer Agulhas Current. The

present incomplete state of knowledge is a consequence of the limited extent of

shallow-water collecting that has taken place by means of SCUBA and the

nature of collecting techniques in the past, i.e. deeper-water trawling and

dredging.

ACKNOWLEDGEMENTS

I thank various staff members of the South African Museum for their kind

assistance: Mr W. R. Liltved for aiding in the collection of live material;

Mrs M. G. van der Merwe and Ms E. J. Hoenson for preparation of photo-

graphic prints and curatorial assistance; Mrs B. Angus for help in locating library

reference material; and Miss L. Wickham for typing the manuscript.

I extend my appreciation to Mr Dane Gerneke and Mr Thomas Smith of the

Electron Microscope Unit, University of Cape Town, for assistance in the

preparation of SEM photographs.

I thank the following for their helpful suggestions and comments: Dr J.

Verseveldt, Zwolle, The Netherlands; Dr F. M. Bayer, Department of Inverte-

brate Zoology, Smithsonian Institution, U.S.A.; Dr P. N. Alderslade of the

Northern Territory Museum, Darwin, Australia; Professor G. M. Branch and

Dr J. A. Day of the Department of Zoology, University of Cape Town.

THE SOUTHERN AFRICAN SOFT CORAL ALCYONIUM VARIABILE 269

REFERENCES

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Anthozoa), with diagnoses of new taxa. Proc. biol. Soc. Wash. 94: 902-947.

Brocu, H. 1912. Die Alcyonarien des Trondhjemsfjordes. I. Alcyonacea. K. norske Vidensk.

Selsk. Skr. 1911: 1-48.

BurcuHarpt, E. 1903. Alcyonaceen von Thursday Island (Torres-Strasse) und von Amboina.

II. Zoologische Forschungsreisen in Australien und dem Malayischen Archipel ausgefihrt

in den Jahren 1891-1893 van Richardsenion. Denkschr. Med.-Naturwiss. Gesellschaft Jena

8: 431-442.

DINGLE, R. V. & Rocers, J. 1972. Pleistocene palaeogeography of the Agulhas Bank. Trans.

R. Soc. S. Afr. 40: 155-165.

Groot, S. & WEINBERG, S. 1982. Biogeography, taxonomical status and ecology of Alcyonium

(Parerythropodium) coralloides (Pallas, 1766). Pubbl. Staz. zool. Napoli I. Mar. Ecol.

3: 293-312.

Hickson, S. J. 1900. The Alcyonaria and Hydrocorallinae of the Cape of Good Hope. Mar.

Invest. S. Afr. 1: 67—96.

Hickson, S. J. 1907a. The differentiation of species of Coelenterata in shallow water seas. Rep.

Trans. Manchr microsc. Soc. 1906: 25-36.

Hickson, S. J. 19076. Coelenterata I. Alcyonaria. Nat. Antarct. Exped. 1901-1904 Nat. Hist.

3: 1-15.

KUKENTHAL, W. 1906. Alcyonacea. Wiss. Ergebn. dt. Tiefsee-Exped. ‘Valdivia’ 1898-1899 13:

1-111.

KUKENTHAL, W. 1916. System und Stammesgeschichte der Scleraxonier und der Ursprung der

Holaxonier. Zool. anz. 47: 170-183.

LoutrscHwaGe_r, J. 1922. Alcyonarien von den Philippinen. I. Die Gattung Alcyonium Linnaeus.

Philipp. J. Sci. 20: 519-542.

May, W. 1899. Alcyonarien. Ergebnisse Hamburger Magalhaensische Sammelreise 4: 1-22.

Hamburg: L. Friederichsen.

Mo.anper, A. R. 1929a. Die Octactiniarien. Further zool. Results Swed. Antarct. Exped.

1901-1903 2 (2): 1-86.

MOLaNnpDeER, A. R. 19296. South and West African Octactiniae in the Gothenburg Natural

History Museum. Géteborgs K. Vetensk.-o. Vitterh Samh. Handl. (B) 1 (7): 3-16.

PFEFFER, G. 1889. Zur Fauna von Sud-Georgien. Jb. hamb. wiss. Anst. 6: 49-55.

PowELL, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecypoda and Gastropoda.

‘Discovery’ Rep. 26: 47-196.

Roxas, H. A. 1933. Philippine Alcyonaria, II. The families Alcyoniidae and Nephthyidae.

Philipp. J. Sci. 50: 345-470.

Tuomson, J. S. 1910. The Alcyonaria of the Cape of Good Hope and Natal. Alcyonacea. Trans.

R. Soc. Edinb. 47: 549-589.

Tuomson, J. S. 1921. South African Alcyonacea. Trans. R. Soc. S. Afr. 9: 149-175.

THomson, J. S. 1924. Charts and comparisons of the distribution of South African Alcyonaria.

With a statement of some of the problems of their dispersal. Trans. R. Soc. S. Afr. 11:

45-84.

TIXIER-DuRIVAULT, A. 1954. Les octocoralliaires d’Afrique du Sud (I. Alcyonacea). Bull. Mus.

natn. Hist. nat., Paris (2) 26: 385-390.

TIxtER-DuRIVAULT, A. 1970. Octocoralliaires. Campagne de la ‘Calypso’ au large des cétes

atlantiques de ’Amerique du Sud (1961-1962). Annls Inst. oceanogr., Monaco 47:

145-169.

Ucuipa, T. 1969. On a northern alcyonarean, Alcyonium pacificum. J. Fac. Sci. Hokkaido

Univ. (6. Zoology) 17: 307-399.

Utinomi, T. 1958. A revision of the genera Nidalia and Bellonella, with an emendation of

nomenclature and taxonomic definitions for the family Nidaliidae (Octocorallia, Al-

cyonacea). Bull. Br. Mus. nat. Hist. 5: 99-121.

UtTinomi, T. 1964. Some octocorals from the Antarctic waters off Prince Harald coast. Scient.

Rep. Jap. Antarct. res. Exped. (E) 23: 1-14.

VERSEVELDT, J. 1967. The Octocorallia collected by R/V ‘Vema’ in the Atlantic Ocean. Am.

Mus. Novit. 2282: 1-19.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

WiLuiams, G. C. 1986. A new species of the octocorallian genus Alcyonium (Anthozoa:

Alcyonacea) from southern Africa, with a revised diagnosis of the genus. J. nat. Hist. 20:

53-63.

WRIGHT, E. P. & STuDER, T. 1889. Report on the Alcyonaria collected by H.M.S. Challenger

during the years 1873-1876. Rep. scient. Results. Voy. ‘Challenger’, Zoology 31: i—lxxu,

1-314.

YAMADA, M. 1950. Descriptions of two Alcyonium from northern Japan. Annotnes zool. jap.

23: 114-116.

6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particu-

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

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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.

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

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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.

GARY C. WILLIAMS

MORPHOLOGY, SYSTEMATICS, AND

VARIABILITY OF THE SOUTHERN

AFRICAN SOFT CORAL ALCYONIUM

VARIABILE (J. STUART THOMSON, 1921)

(OCTOCORALLIA, ALCYONIIDAE)

OF THE SOUTH AFRICAN

CAPE TOWN

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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 la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

FiscHER, P. H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

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

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: ScHuLTZE, L. Zoologische und anthro-

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 96 Band

December 1986 Desember

Part qi Deel

oO"

Alig ce

vio Now

APTIAN AMMONITES FROM THE

ARGENTINIAN AUSTRAL BASIN.

THE SUBFAMILY

HELICANCYLINAE HYATT, 1894

MARIA BEATRIZ AGUIRRE URRETA

Cape Town Kaapstad

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APTIAN AMMONITES FROM THE ARGENTINIAN AUSTRAL BASIN.

THE SUBFAMILY HELICANCYLINAE HYATT, 1894

MARiA BEATRIZ AGUIRRE URRETA

Department of Invertebrate Palaeontology,

South African Museum, Cape Town*

(With 19 figures)

[MS accepted 1 October 1984]

ABSTRACT

Representatives of the subfamily Helicancylinae are locally common in deposits of Aptian

age in the northern central Austral Basin, Patagonia. A stratigraphical synthesis of the Lower

Cretaceous deposits in the area studied is outlined. Schematic sequences of the measured sections

at the principal localities, which also exhibit the various levels containing ammonites, are shown.

The section on systematic palaeontology comprises a discussion of the subfamily Helicancylinae,

and generic and specific descriptions of all taxa represented in the Austral Basin. In addition to

the study of the Patagonian material, bibliographical research reveals the necessity of redefining

the genera Helicancylus and Hamiticeras in order to clarify the systematics of the subfamily. The

following species are identified: Helicancylus patagonicus, Helicancylus bonarellii, Toxo-

ceratoides nagerai, Toxoceratoides cf. biplex, Toxoceratoides? haughtoni, Toxoceratoides? sp.,

and Tonohamites aequicingulatus. The fauna shows some affinities with that of Zululand and

western Europe.

CONTENTS

PAGE

JWR OYGHOVCLSVOVT vas Bitch eB edie Onn ostecneatae put tar et in tial ee ae “Yl

I OCatlOmOlSPECINENS: arian eatin a sate meee wha ie ae: Jaye)

ITIVE MSIOM Seema x eva paca ne becasue ee eMe LE Mj)

Straticraphicsymthesiswi.a. have aaa eo eiev eee ste cela 3 21/3)

FListOnvIolpalacontologicalirescarchy 44-4 4c see 279

Systematic palacontologyann anes aoe See ee nae 280

Concludinesnemarks! iss a ye ee sae oe a See gh le 311

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INCHEROM CC SM rain terse ne ik aaa reine ei AM eR GM & SNE SAE. 8 Sly

INTRODUCTION

The subfamily Helicancylinae comprises a group of small ancyloceratids that

have a nearly worldwide distribution. In the Austral or Magellanes Basin,

Patagonia, Argentina, they are locally common in rocks of Aptian age. Although

small heteromorphs, now referred to this subfamily, were described from this

* Present address: Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires.

This work was carried out under a scholarship sponsored by the Consejo Nacional de

Investigaciones Cientificas y Técnicas, Republica Argentina.

Paid

Ann. S. Afr. Mus 96(7), 1986: 271-314, 19 figs.

DD ANNALS OF THE SOUTH AFRICAN MUSEUM

basin by various authors, those works dealt with general faunal descriptions;

taxonomic and stratigraphic problems concerning this group were still unre-

solved. A description of the representatives of the subfamily Helicancylinae

is necessary in order to advance our knowledge of the Aptian palaeontology and

biostratigraphy of the Austral Basin.

Generic comparisons allow a nearly cosmopolitan correlation. At specific

level, most of the taxa seem to be geographically restricted; some species,

72° 30' \

LOMA PELADA

Tonohamites aequicingulatus

ARGENTINA

PROV. SANTA CRUZ

Helicancylus patagonicus

Toxoceratoides nagerai

Toxoceratoides? haughtoni

Toxoceratoides? sp.

LA MURALLA

Helicancylus patagonicus

oe BAJO COMISION

e@® ‘a Joxoceratoides nageral

oO Toxoceratoides sp. cf. T. biplex @ Ae @.

Ge]

LA HORQUETA

Toxoceratoides? haughtoni

“Tey UPS

own nee

LA FEDERICA

Teller) Us Somtetreli

RIO CARDIEL

Toxoceratoides nagerai

UZ230;

Fig. 1. Index map of the western region of the Province of Santa Cruz,

showing location of the collecting sites.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN ys)

however, show affinities with species described from Zululand and western

Europe.

The localities studied are situated in the north-western province of Santa

Cruz, in the northern central part of the Austral Basin (Fig. 1).

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the source of the material:

CPBA Catedra de Paleontologia, Facultad de Ciencias Exactas y Naturales,

Universidad de Buenos Aires.

MLP Division Paleozoologia de Invertebrados, Museo de Ciencias Natu-

rales de La Plata.

CORD-Pz Catedra de Paleontologia, Universidad Nacional de Cordoba.

DNGM Division Paleontologia, Servicio Geologico Nacional, Argentina.

Casts of some of the figured specimens are deposited at the Department of

Invertebrate Palaeontology of the South African Museum.

Most of the specimens were collected by the author. If not, the name of the

collector is given in the systematic descriptions.

DIMENSIONS

No standard set of abbreviations is in common use for heteromorph

ammonoids. The abbreviations used here are as follows:

JE, = total length

Hy = maximum whorl height

Ho = whorl height opposite to aperture

Ee = minimum whorl height

Wh/Wb = relation between whorl height and whorl breadth.

Dimensions of specimens are given in millimetres.

STRATIGRAPHIC SYNTHESIS

All ammonites studied are (with one exception) from the upper part of the

Rio Mayer Formation. This unit was defined by Hatcher (1897) as ‘a series of

black, very hard, but much fractured slates, with ammonites fairly abundant, but

not sufficiently well-preserved to admit of identification’ on the upper reaches of

the Mayer River. The Rio Mayer Formation outcrops over an extensive belt

parallel to the present main cordillera. The exposures in the area studied,

extending to the north and south of the type locality, have a complex distribution,

controlled by a complicated series of faults and folds (Ramos 1981). Complete

exposures of the Rio Mayer Formation are rare and it is difficult to correlate

partial sections. The selected sections, although not always complete, show at

least definite relations with the under- or overlying formations.

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lee

Green sandstones, ° a

conglomerates and 5 SI @ Tonohamites aequicingulatus

. i

one coquina bed of

= ®

oO --——/

(APR IST RTOS

—————

Black shales with >| G Tropaeum (Tropaeum) sp.

many levels of =

calcareous nodules, SS

interbedded with ———|

fine-grained, green —— 6

calcarenites pak )

Colchidites (C.) vulanensis australis

Heteroceras (H.) elegans

@ Sanmartinoceras africanum

Emericiceras sp.

Rio Mayer Fm.

Fissile black shales

with three levels

of big calcareous

)

Hatchericeras spp.

nodules feesoman)

=| 30m

= 20

= 10

= 0

a Scale

Fig. 2. Stratigraphical section at Loma Pelada, Tucu-Tucu.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 2A

Yellowish-white to M

greenish sandstones, ‘s

tuffs and dark re

shales X

6} Peltocrioceras deeckei

aera

| nee aac

aaa

[i ee ee ea

ss ea

pees

See)

ae)

esa

Ieee eeesssesss Fa FS)

aay

EES sSSey

aes

| ST PEL

= Peltocrioceras deeckei

aa Toxoceratoides cf. biplex

Sassi RN PM inh ie Sanmartinoceras sp.

Australiceras (A.) hallei

Toxoceratoides nagerai

(

{

Australiceras (A.) hallei

G) Sanmartinoceras africanum

Rio Mayer Fm.

)

Fissile black shales + ; ;

wih Galeareous = ‘Aconeceras’ walshense

nodules and thin |

limestones inter- ———

bedded at the base -——}

aes]

aaa

eS)

GES}

—

Ea)

aay

ae)

ras

= 30m

= ee hi

S |x

Stale

Fig. 3. Stratigraphical section at Puesto Bajo Comision, Lake San Martin.

276

ANNALS OF THE SOUTH AFRICAN MUSEUM

Calcareous glauconitic

sandstones, tuffaceous

sandstones and scarce

black shales with

calcareous nodules

Black shales with

Calcareous nodules

interbedded with

dark-grey limestones

and fine-grained

sandstones

Conglomeratic,

quartziferous

sandstones, inter-

bedded with black

shales and coal

horizons

6} Sanmartinoceras patagonicum

Feruglioceras piatnitzkyi

Kachaike Fm.

@ ‘Sanmartinoceras patagonicum’

G' Aioloceras argentinum

Peltocrioceras deeckei

Silesites desmoceratoides

‘Sanmartinoceras patagonicum’

Helicancylus bonarellii

G Toxoceratoides nagerai

Aconeceras sp.

Rio Mayer Fm.

Favrella americana

? Favrella wilckensi

30m

E 20

> 10

Scale

Fig. 4. Stratigraphical section at La Federica, Lake San Martin.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN DET TT

Feruglioceras piatnitskyi

Helicancylus patagonicus

Fine- to medium-

grained greenish. ------— BoE’ G Sanmartinoceras sp. .

yellow sandstones ——s Silesites desmoceratiodes

. ———— G Helicancylus patagonicus

E ae Aconeceras sp.

ie ):| ean

o oe G Peltocrioceras deeckei

—— Eogaudryceras (E.) hertleini

= Toxoceratoides nagerai

iw Toxoceratoides? haughtoni

Toxoceratoides? sp.

Black shales with Helicancylus patagonicus

calcareous nodules

and dark silts

30m

rama

Seana

——

—T

Scale

Greenish-grey

Peet ue Pr 6) Feruglioceras piatnitzkyi

| quinas, | & Silesites desmoceratoides

lenticular limestones 5 8

and yellowish o &

sandstones ae

| @ Sanmartinoceras patagonicum

Phylloceras (Hipophylloceras) sp.

Peltocrioceras deeckei

Black shales with

big calcareous

nodules

Rio Mayer Fen.

Australiceras (A.) cardielensis

G Toxoceratoides nagerai

Fig. 5. Stratigraphical sections. A. Puesto La Senalada, Lake San Martin. B. La Horqueta,

Lake Cardiel.

PET RS) ANNALS OF THE SOUTH AFRICAN MUSEUM

Yellowish-green

sandstones with

green shales

Kachaike Fm.

interbedded at 30m

the base

Black shales with > Peltocrioceras deeckei 10

big calcareous = © Helicancylus patagonicus

nodules °

rd 0

Scale

Fine- tomedium- <&

grained yellowish §&

sandstones, thin Bas

coal levels and S

black shales fe

S Peltocrioceras deeckei

Re Toxoceratoides? haughtoni

Black shales with

big ae = Australiceras (A.) cardielensis

nodules Australiceras (A.) hallei

Tropaeum (T.) inflatum

50m

Scale

Fig. 6. Stratigraphical sections. A. La Muralla, Lake San Martin. B. Rio Cardiel, Lake

Cardiel.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 219

The Rio Mayer Formation consists of a monotonous succession of black

shales, poorly to strongly indurated, sometimes yellowish due to alteration, with

many levels of calcareous nodules. The size of the nodules ranges from a few

centimetres up to more than a metre in diameter. All the fossils are preserved in

these nodules. Different, small-scale facies may be present locally. They are

indicated in Figures 2 to 6, in which the various fossiliferous horizons are also

shown.

The Rio Mayer Formation usually rests conformably on marine and

continental sandstones of the Springhill Formation. In some sections, however,

the unit directly overlies the Jurassic Complejo El Quemado volcanics.

In the northern area the Rio Mayer Formation is conformably overlain by

the sandstones of the Rio Belgrano Formation (Ramos 1979), whilst to the south,

in the Andean region, it is succeeded by the Kachaike Formation. This last unit is

characterized by marine to continental interbedded sandstones and _tuffs

(Riccardi 1971). In the extra-Andean region the Piedra Clavada Formation, a

mainly shallow marine sequence composed of sandstones, shales and tuffs, rests

upon the Rio Mayer Formation (Ramos 1982).

HISTORY OF PALAEONTOLOGICAL RESEARCH

In 1912 Stolley described two specimens of ‘Ancyloceras’ patagonicum

amongst other cephalopods. The material on which Stolley based his study was

collected by Halle (1913), who also studied the palaeoflora and stratigraphy of the

Cretaceous deposits near Bahia de La Lancha, Lake San Martin.

A few years later, Bonarelli & Nagera (1921) published the results of their

geological and palaeontological expedition to Lake San Martin. Amongst other

invertebrate fossils they described and figured two specimens, Leptoceras gr.

silesiacum Uhlig and Leptoceras sp. indet., that were typical of the ‘Level with

Leptoceras’ or ‘Levelc’ in their biostratigraphic subdivision of the Lower

Cretaceous.

A complete regional study dealing with the geology of the western part of the

Province of Santa Cruz was carried out by Piatnitzky (1938). This author also

gave short descriptions and illustrations of the most relevant faunas, including

one specimen of Leptoceras sp. from Arroyo de la Mina and another of

Ancyloceras patagonicum from Rio Cardiel.

After Piatnitzky there were no major palaeontological contributions until

1968 when Riccardi, in an unpublished thesis, described and figured the Lower

Cretaceous invertebrate faunas of Bahia de La Lancha, with a detailed study of

the stratigraphy of this area. Riccardi (1968) described a series of crushed

specimens of ‘Ancyloceras’ patagonicum. He also gave an exhaustive account of

the representatives of this group, not only in the Austral Basin, but also in other

regions of the world.

Finally Leanza (1970), in a monograph dealing with the Cretaceous

ammonite faunas of the Austral Andes, described and figured one specimen of

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

Helicancylus cf. patagonicus (Stolley) and also referred Bonarelli & Nagera’s

Leptoceras to the genus Acrioceras, proposing two new species: A. nagerai and

A. bonarellii.

SYSTEMATIC PALAEONTOLOGY

Class CEPHALOPODA Zittel, 1884

Order AMMONOIDEA Zittel, 1884

Suborder ANCYLOCERATINA Wiedmann, 1966

Superfamily ANCYLOCERATACEAE Gill, 1871

Family Ancyloceratidae Gill, 1871

Subfamily Helicancylinae Hyatt, 1894

Discussion

Casey (1961: 76) grouped in the subfamily Helicancylinae a series of small

ancyloceratids in which the sculpture is simplified on the terminal hook. He

included three Aptian genera, Helicancylus Gabb, Toxoceratoides Spath, and

Tonohamites Spath, while three Barremian genera, Acrioceras Hyatt, Lyto-

crioceras Spath, and Leptoceras Uhlig, were provisionally assigned to the

subfamily.

Casey (1961: 77) also extensively discussed the nomenclatorial problems that

involved the type-genus Helicancylus Gabb, 1869. Gabb (1869) included in

H. aequicostatus a series of fragmentary specimens from which he believed a

complete specimen could be reconstructed. These consisted of an initial helix, a

shaft, and a body chamber; this last fragment was previously referred to

Ptychoceras aequicostatus Gabb, 1864 (pl. 13 (fig. 20)). Anderson (1938: 215)

indicated that the material referred to H. aequicostatus by Gabb (1869) included

at least three species. He restricted the name Helicancylus to the helical part,

redescribing it as H. gabbi (Anderson 1938: 222, pl. 79 (figs 4—5)). He proposed

the new generic name Hamiticeras for the original Ptychoceras aequicostatus

Gabb, 1864 specimen, and for the specimen figured later by Gabb (1869, pl. 25

(figs c-f)). The latter was assigned to Hamiticeras pilsbryi and proposed as the

type-species of the genus. Casey (1961) pointed out that Ptychoceras aequicosta-

tus is the type-species of Helicancylus and that it is congeneric with Hamiticeras

pilsbryi. Thus the latter genus became a synonym of Helicancylus.

The present author disagrees with Casey and partially agrees with Anderson,

in that the original Helicancylus comprises three different genera. The helical

fragment shows no relation to any of the other specimens and is here excluded

from the subfamily. The helical coiling of the specimen recalls Helicancyloceras

Klinger & Kennedy, 1977, and even more Kutatissites Kakabadze, 1970

(= Simionescites Avram, 1976b), being similar in particular to Kutatissites

princeps (Avram, 1976b, pl. 3 (fig. la—c)) and K. rachathasensis Kakabadze,

1981 (pl. 11 (fig. 3a—c)). Although Casey (1961) considered the helical fragment

as ‘irrelevant to the interpretation of Helicancylus’, it can be seen from recent

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 281

literature that this is not so. Its inclusion in the genus and even in the subfamily

only leads to misconceptions. This was the case with Thieuloy (1976), who

referred Kutatissites to the subfamily Helicancylinae when comparing it with ‘the

helical part of Helicancylus’. The large ancyloceratid Kutatissites can hardly be

placed in a subfamily that groups ‘those diminutive ancyloceratids in which the

sculpture is simplified on the terminal hook’ (Casey 1961: 76).

With reference to the shaft and body-chamber fragments assigned to

Helicancylus by Gabb (1869) and to Hamiticeras by Anderson (1938), it is here

believed that Hamiticeras pilsbryi, type-species of Hamiticeras, is not congeneric

with Ptychoceras aequicostatus, type-species of Helicancylus. It is proposed to

consider both as valid genera. Helicancylus will be discussed later; Hamiticeras

can be diagnosed as follows: ‘Small size. Shaft straight and final hook long, almost

parallel to the shaft. Ornament of the phragmocone consists of strong

trituberculate and thin intermediate ribs. Final hook with sharp, high, non-

tuberculate ribbing.’ To Hamiticeras can be referred H. pilsbryi from the Upper

Aptian (Argonauta zone) of California (Anderson 1938) and from the Caucasus

(Drushchits & Kudryavtsev 1960: 295, pl. 11 (figs 6a—b, 7a—b)), and Hamiticeras

sp. (Avram 1976a, pl. 2 (fig. 1 only)) from the Upper Aptian of the Carpathian

mountains (Romania).

After Casey’s (1961) monograph, some authors adopted his classification

(Day 1974; Thomson 1974; Klinger & Kennedy 1977; Martinez 1982), while

others (Murphy 1975; Forster 1975; Etayo Serna 1979; Kakabadze 1981) assigned

the different genera to the family Ancyloceratidae, without any reference to the

subfamilies.

More complicated is the history of the assignation of the Barremian genera

Acrioceras, Leptoceras and Lytocrioceras. Sarkar (1955) described and illustrated

a series of species of Acrioceras and Leptoceras, and reviewed the original

material of Lytocrioceras. He expanded the original conception of Acrioceras

Hyatt, proposing four new subgenera. They were mainly based on the coiling and

on the ornament:

Acrioceras (Acrioceras) s.s.: coiling acrioceratid; ornament with tuberculate ribs.

Acrioceras (Paraspinoceras) (Breistroffer): coiling acrioceratid; non-tuberculate

ribs.

Acrioceras (Aspinoceras) (Anderson): coiling aspinoceratid; non-tuberculate

ribs.

Acrioceras (Protacrioceras) Sarkar: coiling aspinoceratid; tuberculate ribs.

(See Figure 7 for the different types of coiling.)

Unfortunately Sarkar (1955) based his revision on material of D’Orbigny and

from other collections of the nineteenth century, which lack precise stratigraphic

data. According to Sarkar (1955: 26) Acrioceras ranges from the Hauterivian to

the Lower Aptian. He described one species of Leptoceras and redescribed the

type of Lytocrioceras jauberti (Astier, 1851: 25, pl. 9 (fig. 17)) but no photo-

graphic illustration was given.

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

CCG

Fig. 7. Different types of coiling referred to in the text. A. Aspinoceratid. B. Crioceratitid.

C. Acrioceratid. D. Ancyloceratid. E. Toxoceratid. F. Labeceratid.

Wright (1957: L211) maintained Aspinoceras as a valid genus and included

Paraspinoceras as a doubtful synonym of Lytocrioceras. He interpreted Helican-

cylus in the sense of Anderson (1938) and regarded Tonohamites and Toxocera-

toides (the latter with doubt) as synonyms of Hamiticeras. While Thomel (1964)

accepted Sarkar’s (1955) revision of Acrioceras, Wiedmann (1962) dissented and

regarded all four subgenera as synonyms of Acrioceras s.1. Manolov (1962: 531)

proposed the new subfamily Leptoceratinae for a group of small Barremian forms

that he considered to be early representatives of the family Ancyloceratidae. He

also pointed out the close relationship between all these forms. His subfamily

comprised Leptoceras Uhlig, Karsteniceras Royo y Gomez, Veleziceras Wright,

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 283

and Eoleptoceras Manolov. Manolov (1962) furthermore questioned the occur-

rence of Leptoceras in the Berriasian (Nikolov 1960), as that genus is unknown

from the Valanginian and Hauterivian. Thieuloy (1966), however, maintained

that the true Leptoceras occurs in the Berriasian and erected the new genus

Leptoceratoides for the homeomorphic Barremian forms. Dimitrova (1970)

proposed a completely new arrangement of the Cretaceous heteromorphs,

spreading the different genera united in the Helicancylinae by Casey (1961) into a

number of families and subfamilies. Although she based her study on suture lines,

she did not accept Wiedmann’s (1966) suborder Ancyloceratina, which included

all Cretaceous ammonoids with quadrilobate primary suture. Dimitrova’s (1970)

proposal was generally avoided by later authors, except for Avram (1976a), who

followed her classification.

As interpreted here, the subfamily Helicancylinae comprises the Aptian

genera Helicancylus Gabb, Hamiticeras Anderson, Tonohamites Spath, and

Toxoceratoides Spath, and the Barremian Acrioceras Hyatt and ?Lytocrioceras

Spath. The last genus is very enigmatic and as far as can be established it is only

known from the single specimen of the type-species. The Barremian genus

‘Leptoceras’ (= Leptoceratoides) has been referred to a different stock (Manolov

1962; Wiedmann 1973).

In Patagonia, the subfamily Helicancylinae is represented by Helicancylus,

Toxoceratoides, and Tonohamites.

Genus Helicancylus Gabb, 1869

Type-species. Ptychoceras aequicostatus Gabb, 1864, from the Aptian of

California, by original designation (Gabb 1869).

Diagnosis

Coiling variable, usually with a straight or curved shaft and a final hook.

Ornament on the shaft consists of ribs of equal size, each one bearing one to three

rows of tubercles. Final hook with single, non-tuberculate ribs. Suture line with

bifid saddles and asymmetrical, trifid lobes.

Discussion

As interpreted here, Helicancylus differs from Toxoceratoides by the total

lack of intercalatory ribbing and the presence of tubercles on every rib of the shaft

as well as the simple ribbing on the final hook. Toxoceratoides shows strong

trituberculate and fine intercalatory ribs on the shaft and sharp ribs springing in

bundles from umbilical tubercles on the final hook (Casey 1961; Klinger &

Kennedy 1977).

Tonohamites is easily distinguished from Helicancylus by the rounded, non-

tuberculate ribs on the shaft and the strong, rounded or flat ribbing on the final

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

hook. Hamiticeras Anderson has similar ornament on the final hook but shows

strong trituberculate ribs separated by thin intermediaries on the shaft.

According to Casey (1961: 93) the only European record of Helicancylus was

the Upper Aptian ‘Hamites’ sp. figured by Jacob & Tobler (1906, pl. 2 (figs

10—11)). Besides this material, which most probably belongs to the genus, several

species can be assigned to it, although some are included tentatively. They are as

follows:

Ancyloceras elatum von Koenen (1902: 375, pl. 38 (fig. 8a—c), pl. 40 (fig. 2a—b),

pl. 45 (fig. 9), pl. 53 (figs 6-7)).

Toxoceratoides? elatum (von Koenen) (Kemper 1976, pl. 33 (fig. 1)).

Ancyloceras fustiforme von Koenen (1902: 384, pl. 49 (figs 4-5, 7, 9), pl. 53

(figs 8—9)).

Toxoceratoides cf. fustiformis (von Koenen) (Casey 1961: 83, pl. 17 (fig. 4)).

Hamiticeras aequicostatum (Gabb) (Anderson 1938: 216, pl. 37 (figs 2—2a, 3),

al, 79 Cie, ))).

Hamiticeras philadelphium Anderson (1938: 216, pl. 79 (figs 2—3)).

Ancyloceras patagonicum Stolley (1912: 11, pl. 1 (figs 3-3a, ?2—2a)).

Acrioceras bonarellii Leanza (1970: 207, fig. 6 (1)).

Genus uncertain. Group of “Ancyloceras’ patagonicum Thomson (1974: 19, pl. 3

(figs c, g—h)).

‘Ancyloceras’ elatum von Koenen was doubtfully referred to Toxoceratoides

by Kemper (1976), who also figured one fragmentary specimen. This species as

well as “Ancyloceras’ fustiforme von Koenen, assigned to Toxoceratoides by Casey

(1961) are here both included in Helicancylus (see p. 290).

The group of “Ancyloceras’ patagonicum referred to an indeterminate genus

by Thomson (1974) can also be placed in Helicancylus. Thomson compared the

suture line of ‘Hamiticeras’ aequicostatum (illustrated by Anderson 1938, pl. 79

(fig. 6)) with that of the lectotype of ‘Ancyloceras’ patagonicum. He noted that

‘The suture of ““Hamiticeras’’ is more complex and has a narrow external saddle, a

broad trifid first lateral lobe and a slightly smaller second lateral lobe’ (Thomson

1974: 20). He concluded that the ‘A.’ patagonicum-group most probably

represented a new genus.

The suture line of the lectotype of ‘A.’ patagonicum, as illustrated by

Thomson (1974, text-fig. 4b) appears very similar to that of the Patagonian

material studied here and referred to Helicancylus patagonicus. Both are figured

(Fig. 9D-G) for comparison. Recently Thomson (1982) included the Patagonian

material in Helicancylus, but in open nomenclature.

Occurrence

Helicancylus occurs in the Aptian of Antarctica (Thomson 1974), California

(Gabb 1869; Anderson 1938), England (Casey 1961), Germany (Von Koenen

1902; Kemper 1976), Patagonia (Stolley 1912; Riccardi 1968; Leanza 1970), and

Switzerland (Jacob & Tobler 1906) (Fig. 8).

285

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN

‘ovutAOUBOT[OH JO e1oues ueNdy Imoy oy} Jo UONNQINSIP jeorydesZoaSorqoorjeg 8 ‘SI

Selo WweY Vv

SnjAoUedI|o] &

Se}ILWEYOUO] mK

SaPIO}EJIIOXO] + @

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

The report of Helicancylus furcata Kakabadze from the Aptian of the Soviet

Union (Kakabadze 1981) is not accepted here. Kakabadze interpreted the genus

in the sense of Anderson (1938), referring to the initial helix only.

Helicancylus patagonicus (Stolley, 1912)

Figs 9A-G, 10, 11A—C, 12A-F, 19A-B

Ancyloceras patagonicum Stolley, 1912: 11, pl. 1 (figs 3-3a, ?2-2a).

non Ancyloceras patagonicum Stolley: Howarth, 1958: 4, pl. 1 (fig. 4).

non ‘Ancyloceras’ patagonicum Stolley: Riccardi, 1968 (pars), pl. 21 (fig. 1).

non Helicancylus cf. patagonicus Leanza, 1970: 205, fig. 4 (1).

non ‘Ancyloceras’ patagonicum Stolley: Thomson, 1974: 19, pl. 3 (figs c, g—h).

Lectotype

The specimen figured by Stolley (1912, pl. 1 (fig. 3-3a)). Original at

Riksmuseum N Mo. 117877, Stockholm, by subsequent designation Thomson

(1974: 19).

Material

CPBA 11062 from La Muralla, Lake San Martin; CPBA 10898, 10848, 10844

and ?10887 from Puesto La Senalada, Lake San Martin. Rio Mayer Formation.

Upper Aptian.

Description

The most complete specimen, CPBA 11062 (Fig. 11A—C), shows ancylo-

ceratid coiling, with a nearly straight shaft and a recurved crozier. The early stage

of growth is unknown.

The whorl section is initially compressed (Wh/Wb = 1,13—1,20), subovoid,

with rounded dorsum and venter and flat to gently inflated flanks. With increasing

diameter, the whorl section becomes more rounded and on the final hook it is

nearly circular (Fig. 9A—C).

Ornament on the shaft consists of prominent, narrow, obliquely prorsi-

radiate, tuberculate ribs. They are separated by interspaces wider than

themselves. They pass with a slight forward curvature over the dorsum,

sometimes showing duplications, and are distinctly interrupted over the venter.

All the ribs bear at least two rows of tubercles, one siphonal and the other

ventrolateral. In some specimens there is also a third row of small dorsolateral

tubercles.

Towards the end of the shaft the tuberculation gradually disappears and on

the final hook the ornament consists of radial, simple, sharp ribs. They cross the

venter without interruption. On the dorsum they are reduced to striae with a

forward curvature.

The suture line is quite simple, with trifid lateral, umbilical and internal

lobes; the first is broad and slightly asymmetrical (Fig. 9E-G).

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 287

ra VN

Fig. 9. Helicancylus patagonicus (Stolley). A-—C. Whorl section of CPBA 11062. x 2,5.

D. Suture line of lectotype (after Thomson 1974). E-F. Suture line of CPBA 11062. x 4.

G. Suture line of CPBA 11087. x 4.

Dimensions

Specimen lehy, lake Joke

Lectotype* 20,0 14,0 13,0

CPBA 11062 16,0 11,0 9,0

CPBA 10848 14,0 — 11,0

CPBA 10844 20,0 — 16,0

CPBA 10887 10,0 — 4,0

* Deposited at the Riksmuseum N Mo. 117877 (Stockholm). Measurements taken from

Stolley’s (1912, pl. 1 (fig. 3—3a)) original photograph.

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

When Stolley (1912) proposed this species, he described two fragmentary

specimens. He was in doubt whether to refer both to the same species, but

pointed out that the difference in ornament was no greater than that in other

species of ‘Ancyloceras’ known from the Lower Cretaceous of northern

Germany. As far as can be seen from the original illustrations, the main

difference between Stolley’s twe specimens is the degree of curvature of the shaft,

rather than the ornament. The small specimen figured by Stolley (1912, pl. 1

(fig. 2-2a)) is here doubtfully referred to the species. According to Thomson

(1974) both may belong to different genera.

Helicancylus patagonicus (Stolley) differs from H. bonarellii (Leanza) in the

coiling and in the ornament. In the former, the coiling is ancyloceratid, with a

straight shaft and a recurved crozier, and a slow increase in the whorl section. In

the latter, the coiling is open crioceratitid or toxoceratid, with a rapid increase in

the whorl section, especially on the final hook. Besides, in H. bonarellii the ribs

are rounded, closely spaced, and with feeble tubercles, while in H. patagonicus

Fig. 10. Reconstruction of Helicancylus patagonicus (Stolley).

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 289

Fig. 11. Helicancylus patagonicus (Stolley). CPBA 11062 from La Muralla. x 1.

they are high and sharp, with well-developed tuberculation, and they are

separated by wider interspaces.

The Antarctic material figured by Howarth (1958, pl.1 (fig. 4)) as

‘Ancyloceras’ patagonicum or by Thomson (1974, pl. 3 (figs c, g—h)) as group of

‘A.’ patagonicum seems to be more related to H. bonarellii.

Helicancylus aequicostatus Gabb differs from H. patagonicus in the whorl

section, ornament, and suture line. The former species has a subtrapezoid whorl

section with flat dorsum, shaft ornamented with dense rounded ribs that cross the

venter without interruption and bear faint siphonal tubercles only. The suture line

in H. aequicostatus is more incised than in the Patagonian species, with a narrow

ventral saddle (Anderson 1938: 217, pl. 37 (figs 2-3), pl. 79 (fig. 6)).

Helicancylus philadelphium (Anderson) is very closely related to H. aequi-

costatus. Although Anderson (1938) did not compare them, it seems that\a row of

faint ventrolateral tubercles in the former species is the only different feature.

The European species H. fustiformis and H. elatum described by Von

Koenen (1902) are known from fragmentary specimens only. Although Von

Koenen’s descriptions are precise, he described each fragment in detail and it is

difficult to interpret each species as a whole.

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12. Helicancylus patagonicus (Stolley). A-B. CPBA 10844. C-—D. CPBA 10848.

E-F. CPBA 10887. All from Puesto La Senalada. All x 1.

Helicancylus elatum (von Koenen) (1902: 375, pl. 38 (fig. 8a—c), pl. 40

(fig. 2a—b), pl. 45 (fig. 9), pl. 53 (figs 6-7)) has a rapid increase in the whorl

section, and fine dense ribs that are trituberculate on the shaft and bend of the

crozier. Helicancylus fustiformis (von Koenen) (1902: 384, pl. 41 (figs 4-5, 7a—c,

9a—b), pl. 53 (figs 8a—b, 9a—b)) is mostly known by small shaft fragments with

subcircular whorl section and trituberculate ribs. Casey (1961: 83, pl. 17 (fig. 4))

described a fragmentary specimen as Toxoceratoides cf. fustiformis, pointing out

that this species is very similar to T. royerianus but with tubercles in every rib.

In Patagonia, Piatnitzky (1938, pl. 6 (figs 31-32)) figured a fragment of a

shaft of “‘Ancyloceras’ patagonicum. It may belong to this species because of its

well-marked tubercles, although the ribbing is quite dense.

The small specimen referred to Helicancylus cf. patagonicus by Leanza

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 291

(1970) has bi- or trifurcate ribs arising from an umbilical tubercle on the bend of

the crozier. It is here referred to Toxoceratoides nagerai.

It is interesting to discuss the range of this species. Halle (1913) collected one

specimen, illustrated by Stolley (1912, pl. 1), in sandstones referred to his

division 6, in Calafate Stream (fig. 2-2a), and another from the summit of a high

ridge on the south side of a stream (today known as Bajo Comision Stream) also

in his division 6. This informal lithologic unit is at present known as Kachaike

Formation, of Upper Aptian to Albian age.

When describing Halle’s material, Stolley (1912) compared it with European

species of Upper Neocomian to Lower Aptian age, but concluded that

“Ancyloceras’ patagonicum might be of Upper Aptian age. This idea was ratified

in his description of ?Oppelia (Adolphia) sp. in the same publication. Thomson’s

(1974: 20) reference to an Upper Neocomian or Lower Aptian age of this species

seems to be a misreading of Stolley’s work.

All the material described here was collected in the uppermost section of the

Rio Mayer Formation, in a level characterized by the abundance of giant

specimens of Peltocrioceras deeckei (Favre). This fact partially corroborates

Riccardi’s (1968) opinion that ‘A.’ patagonicum was not present in Halle’s

division 6, but only in his division 5 (at present the Rio Mayer Formation).

Unfortunately, this statement was based on a negative fact: that after Halle, no

one has collected specimens of this species in the Kachaike beds, but always in the

Rio Mayer Formation. Another problem is that Riccardi (1968) described as

‘A.’ patagonicum what is here referred to Helicancylus bonarellii and H. pata-

gonicus. The former species is actually restricted to the upper section of the Rio

Mayer Formation in a level below that of Peltocrioceras deeckei, while

H. patagonicus occurs with it.

Further problems arise with the statement by Halle (1913) that ‘A.’ pata-

gonicum occurs above a level with a well-preserved taphoflora in Bajo Comision

Stream. According to Baldoni & Ramos (1981) this apparent position is just

topographic. They pointed out that this species occurs in the middle section of the

Rio Mayer Formation at that locality. The present author had the opportunity to

study the specimen of ‘A.’ patagonicum of Baldoni & Ramos (1981). It does not

belong to Helicancylus patagonicus but to Toxoceratoides nagerai (see

Fig. 17A—B), which actually occurs in the middle section of the Rio Mayer

Formation at the Puesto Bajo Comision locality, well below the level with the

flora.

At the locality La Muralla, which seems to be very close to the place where

Halle collected his second specimen of ‘A.’ patagonicum, this species is associated

with Peltocrioceras deeckei. They were found in the uppermost section of the Rio

Mayer Formation. Although the latter species does not provide a precise age, its

association with Eogaudryceras (Eogaudryceras) hertleinei (Wiedmann) at Puesto

La Sefalada and with Acantohoplites (Nolaniceras) uhligi (Anthula) at Vega

Montes de Oca, together with stratigraphical evidence, points to an Upper

Aptian age for the horizon of Peltocrioceras deeckei (Aguirre Urreta 1985).

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

The author also had the opportunity to study one beautifully preserved

specimen of Helicancylus patagonicus collected by Piatnitzky. The latter stated

(1936) that the specimen was found loose in the Cerro Pelado, in the Rio Cardiel

area. At that locality only the lower member of the Piedra Clavada Formation

(Ramos 1982, fig. 2) is exposed. The lithology of the nodule in which the

specimen is preserved confirms its origin. The Piedra Clavada Formation overlies

the Rio Mayer Formation comformably and has its chronological equivalent in

the Kachaike Formation.

All this indicates that Helicancylus patagonicus is associated with Pelto-

crioceras deeckei and that it appears at some higher horizons. It means that we

cannot rule out its possible presence in the Kachaike Formation, as was already

stated by Halle (1913).

Helicancylus bonarellii (Leanza, 1970)

Fig. 13A-F

Leptoceras gr. silesiacum Uhlig: Bonarelli & Nagera, 1921: 18, fig. 3.

Leptoceras sp. Piatnitzky, 1938: 79, pl. 4 (fig. 20).

?Ancyloceras patagonicum Stolley: Howarth, 1958: 4, pl. 1 (fig. 4).

“Ancyloceras’ patagonicum Stolley: Riccardi, 1968 (pars), pl. 21 (fig. 1).

Acrioceras bonarellii Leanza, 1970: 209, fig. 6 (1).

Paraleptoceras singulare Leanza, 1970: 209, fig. 8 (5).

? “Ancyloceras’ patagonicum Stolley: Thomson, 1974: 19, pl. 3 (figs c, g—h).

Holotype

The specimen figured by Bonarelli & Nagera (1921, fig. 3). Geological

Survey Collection DNGM 9308 from locality Bahia de La Lancha (here referred

to as La Federica), Lake San Martin, Rio Mayer Formation. ?Upper Aptian.

Material

Apart from the holotype, MLP 17094-96 (collected by A. Riccardi),

CORD-Pz 4360 (collected by M. Flores), MLP 16018a—b (collected by

H. Arbe), CBPA 11065—6 from the same level and locality as the holotype. Rio

Mayer Formation. ?Upper Aptian.

Description

All the available specimens are crushed or preserved as impressions. Coiling

is variable. On most of the specimens it is toxoceratid, but in some it is open

crioceratitid with the whorls not touching. The most complete specimen

(Fig. 13A) shows a small open spire followed by a gently curved shaft and a final

recurved crozier. Nothing can be said about the whorl section as the specimens

are extremely crushed.

Ornament consists of fine, simple, rounded ribs, separated by interspaces

narrower than the ribs. On the initial spire rib density is about four per whorl

height, on the shaft or at mid-growth stage (in specimens with crioceratitid

coiling) it is about five, and at the final stage nine ribs are present per whorl

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 293

Fig. 13. Helicancylus bonarellii (Leanza). A. MLP 17094. B. CPBA 11065. C. MLP 16018a.

D. MLP 16018b. E. MLP 17095. F. MLP 17096. All from La Federica. All x 1.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

height. The ribs bear tubercles, but not only their number but also their

appearance and disappearance are extremely variable. When present at the early

stage, the tubercles are only ventral. As size increases, the ribs also show small

ventrolateral and even smaller umbilical tubercles.

On the final hook the tuberculation gradually disappears and the ornament

consists of simple, rounded radial ribs, which cross the venter apparently without

interruption.

The suture line is unknown.

Dimensions

Specimen L Hy Ho HA,

MLP 17094 56,0 10,0 OS) 1,0

MLP 17096 50,0 7/0 9,0 4,0

MLP 16018a 53,0 13,0 8,5 4,0

MLP 17095* 47,0 16,5 — So)

MLP 160185 38,0 7,0 — 3,0

DNGM 9308 - 20,0 SRO 6,0

* Specimens with crioceratitid coiling.

Discussion

Leanza (1970), when proposing this species, indicated that it belonged to the

‘Leptoceras’ silesiacum group. He also pointed out that according to Anderson

(1938) the species had to be referred to the genus Acrioceras Hyatt. Riccardi

(1968) had already stated that Uhlig (1883) never included ‘Crioceras’ silesiacum

in his subgenus Leptoceras. It is irrelevant to discuss here the generic affinities of

Uhlig’s species, but it differs from the Patagonian material in the coiling and in

the ornament of the shaft and final hook (Uhlig 1883: 142, pl. 28 (fig. 4)).

Helicancylus bonarellii differs from H. patagonicus in its smaller size and in

its crioceratitid or toxoceratid instead of the latter’s ancyloceratid coiling. The

ornament of the former species consists of rounded, dense ribs with weak

tubercles.

As stated before, the specimens illustrated by Thomson (1974, pl. 3

(figs c, g-h)) and referred to an indeterminate genus of the group ‘Ancyloceras’

patagonicum, as well as the material figured by Howarth (1958, pl. 1 (fig. 4)) as

‘A.’ patagonicum, show more similarities with H. bonarellii than with H. pata-

gonicus. It is interesting to point out the close morphological resemblance

between some specimens of Antarcticoceras antarcticum Thomson (1974: 20, pl. 3

(figs i-k, m—n)) and those of H. bonarellii with crioceratitid coiling.

Antarcticoceras antarcticum was first referred to an unknown genus of the

subfamily Helicancylinae (Thomson 1971: 158) and afterwards to an uncertain

family (Thomson 1974). Thomson was in doubt whether to assign this taxon to the

Crioceratitidae or to the Ancyloceratidae. He stated that in the morphology of

the shell the genus seems to be allied to the Crioceratitidae, but it also has some

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 295

non-crioceratitid features such as ribbing of similar size, a rather simple suture

line and a high dorsolateral tubercle.

Further research on the possible relationships between the Antarctic

specimens of “Ancyloceras’ patagonicum, Antarcticoceras antarcticum, and the

Patagonian Helicancylus bonarellii might be worthwhile.

Genus Toxoceratoides Spath, 1924

Type-species. Toxoceras royerianum dOrbigny, 1842, from the Lower

Aptian of France, by original designation (Spath 1924: 78).

Diagnosis

Coiling ancyloceratid or toxoceratid. Ornament of initial spire and shaft

consists of trituberculate and intermediate ribs. On the final hook the ornament is

simpler, with single ribs intercalated with others that bi- or trifurcate from an

umbilical tubercle. Suture line with bifid saddles and trifid lobes.

Discussion

Spath (1924: 78) erected this genus without giving any diagnosis. Wright

(1957: L212) doubtfully regarded Toxoceratoides as a synonym of Hamiticeras,

taking into account that the type-species of the former genus was only known by

fragments that do not allow a proper description.

Drushchits & Eristavi’s (1958) first diagnosis of Toxoceratoides referred only

to the coiling, suture line, and ornament of the shaft. Casey (1961: 77) accepted

the validity of the genus and characterized it as: ‘Coiling ancyloceratid or

leptoceratid; may commence with a very small helix. Phragmocone ornamented

as in Ancyloceras, with periodic trituberculate ribs. Final hook with close,

narrow, sharp ribbing which bifurcates or trifurcates irregularly from an umbilical

tubercle. Suture line as in Ancyloceras.’

Although Casey stated the presence of leptoceratid coiling in his diagnosis

none of the species he refers to this genus actually shows it. The term

‘leptoceratid coiling’ is difficult to interpret as Leptoceras includes species with

crioceratitid as well as open coiling. Thus it is preferable to use the term

toxoceratid instead of leptoceratid coiling (Fig. 7E).

None of the species referred to Toxoceratoides shows any trace of an initial

helix, except for a doubtful record of Day (1974: 13). Day identified three

fragments as Toxoceratoides? sp. The present author had the opportunity to see

plaster casts of two of them. The ornament as well as the coiling do not fit in

Toxoceratoides and they may be inner whorls of a big ancyloceratid (e.g. Pelto-

crioceras). The third fragment, as seen in the illustration (Day 1974, pl. 1

(fig. 2)), shows a partially preserved initial helix followed by a shaft ornamented

with narrow, close, non-tuberculate, sharp ribs—a feature not common in

Toxoceratoides. Therefore the presence of an initial helix is ruled out in the

present diagnosis of Toxoceratoides. If this feature proves to be present, it would

be necessary to analyse the taxonomic position of this genus again, as the

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

presence of an initial helix is a typical feature of the nearly contemporaneous

Heteroceratinae (Klinger 1976).

Klinger & Kennedy (1977: 307) partially accepted Casey’s diagnosis; they

pointed out the close relationship between Toxoceratoides and Tonohamites, and

stated that species like Tonohamites decurrens, with strong trituberculation on the

shaft, link both genera.

Later, Etayo Serna (1979: 20) proposed the subgenus Colomboceratoides,

type-species Toxoceratoides (Colomboceratoides) renzoni, with the following

diagnosis: ‘Coiling as in Toxoceratoides but differs from the latter genus by the

development of sculpture characterized by a retarded development of the subdue

lateral trituberculation: early ribs without tubercles, followed by the appearance

on both sides of venter of slender spines, subsequently lateral tubercles appear

and much later the nipple-like peridorsal tubercles show up. The suture line is

much simplified, it has massive subrectangular saddles and subtrifid L.’

According to Kakabadze (1981: 129) this subgenus is superfluous as its main

characteristics are the same as those present in Toxoceratoides. The only apparent

difference is the delayed appearance of the trituberculation, but this is a very

variable character and it does not seem to have any specific value.

Toxoceratoides differs from Tonohamites mainly in the ornament of the final

hook. In the latter genus tuberculation on the shaft is usually reduced but, as

stated by Klinger & Kennedy (1977), species like Tonohamites decurrens, with a

strong trituberculate phragmocone, show the close relationships between these

two genera.

Hamiticeras has a Toxoceratoides-like shaft, but the final hook is long,

parallel to the shaft, and the ribs are sharp, strong, single, and wide-spaced.

These features are sufficient to distinguish the genera.

It is very difficult to find complete specimens of Toxoceratoides, perhaps due

to their small size and open coiling. This has led to the erection of a large number

of species in this genus. Some of these are monotypic or based on fragments,

while others were erected in the nineteenth century and never restudied. There is

no complete agreement between different authors as to which features are of

specific value. The only distinct characteristics for specific separation are the

coiling, whorl section, and the ornament of the body chamber. To a lesser degree,

the ornament of the shaft can be used; little can be said about the initial spire as it

is virtually unknown.

Species referred to or possibly belonging to Toxoceratoides are:

T. biplex (von Koenen) (1902: 381, pl. 41 (figs 3, 10a—b, 11a—b)).

T. biplicatum (von Koenen) (1902: 379, pl. 41 (figs 2a—b, 8a—b)).

I. caucasicus (Kasansky) (1914: pl. 1 (fig. 8a—c)).

I’. corae Murphy (1975: 33, pl. 5 (figs 1, 5)).

I. emericianum (d’Orbigny) (1842: 487, pl. 120 (figs 5—9)).

T.? greeni Murphy (1975: 33, pl. 5 (figs 2-3, 6)).

T.? haughtoni Klinger & Kennedy (1977: 310, figs 59A—-D, 60A-I, 61A-C,

62A—D, 63, 64A-C, 65A-B, 66B, 79A-B).

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 297

. krenkeli Forster (1975: 160, pl. 4 (figs 1-2), text-fig. 33a—b).

. nagerai (Leanza) (1970: 206, fig. 5 (1)).

. obliquatus (Young & Bird) (1828: 278, pl. 18 (fig. 11)).

. proteus (Spath) (1930: 461, pl. 16 (fig. 7)).

(Colomboceratoides) renzoni Etayo Serna (1979: 20, pl. 6 (fig. 19),

text-fig. 30-P).

. rochi Casey (= Ancyloceras royerianum Roch, 1927: 30, pl. 1 (fig. 4)).

. rotundus (Phillips) (1875: 264, pl. 1 (fig. 24)).

. royerianus (d’Orbigny) (1842: 481, pl. 118 (figs 7-11)).

. saulae Murphy (1975: 31, pl. 4 (figs 4, 6)).

seminodosus (Roemer) (1841: 93).

. Sheperdi (Spath) (1924: 173, figs 5-6).

. starrkingi (Anderson) (1938: 207, pl. 59 (fig. 4-4A), pl. 45 (fig. 4A)).

. subproteus Casey (1980: 651, pl. 103 (fig. 3)).

Toxoceratoides sp. 1 Murphy (1975: 35, pl. 6 (figs 1-2, 11)).

Toxoceratoides? sp. 2 Murphy (1975: 35, pl. 3 (fig. 6), pl. 6 (figs 5-6)).

Toxoceratoides sp. nov. Thomson (1974: 16, pl. 3 (figs a, d)).

Ancyloceras (Acrioceras) aff. starrkingi Anderson (Jeletzky 1964: 66, pl. 19

(fig. 2A—C)).

Toxoceratoides sp. 1 Martinez (1982: 140, pl. 24 (fig. 3a—c), text-fig. 21).

Toxoceratoides sp. 2 Martinez (1982: 141, pl. 24 (fig. 4a—d), text-fig. 22).

Toxoceratoides sp. 3 Martinez (1982: 142, pl. 24 (fig. 5a—d)).

Toxoceratoides biplicatum (von Koenen), referred to Toxoceratoides by

Klinger & Kennedy (1977: 307), with bifurcate and ventrally tuberculate ribs on

the shaft, is doubtfully included in this genus. According to Murphy (1975)

Ancyloceras (Acrioceras) aff. starrkingi described by Jeletzky (1964) does not

belong to Anderson’s species and may be new.

SSS Ss

oS oe SS aa

Occurrence

Toxoceratoides occurs in Germany (Von Koenen 1902), Antarctica (Thom-

‘son 1974), Canada (Jeletzky 1964), Colombia (Etayo Serna 1979), Spain

(Martinez 1982), California (Anderson 1938; Murphy 1975), France (D’Orbigny

1842; Roch 1927), England (Spath 1924, 1930; Casey 1961, 1980; Howarth 1962),

Mozambique (Krenkel 1910; Haughton & Boshoff 1956; Wachendorf 1967;

Forster 1975), Romania (Avram 1967a), south-western USSR (Kakabadze 1981),

Zululand (Klinger & Kennedy 1977) and Patagonia (Leanza 1970; present

paper).

Reports of Toxoceratoides from Australia are not accepted here. “Ancylo-

ceras’ taylori Etheridge, referred to Toxoceratoides by Whitehouse (1926), is

believed to be a Tonohamites species, as discussed later in this paper. The

fragments described by Day (1974) as Toxoceratoides? sp. seem to belong to more

than one genus, but not to Toxoceratoides.

According to Casey (1961) Toxoceratoides ranges from the Upper Barremian

to the Lower Aptian (deshayesi zone), and it appears to be replaced by

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tonohamites at the top of the Lower Aptian (bowerbanki zone). However, more

recent publications (Klinger & Kennedy 1977; Etayo Serna 1979) show that

Toxoceratoides ranges up to the Upper Aptian. In Patagonia this genus is present

in deposits of Lower and Upper Aptian age.

Toxoceratoides nagerai (Leanza, 1970)

Figs 14, 15A—-C, 16A-D, 17A-D

Leptoceras sp. indet. Bonarelli & Nagera, 1921: 19, fig. 4.

Helicancylus cf. patagonicus (Stolley): Leanza, 1970: 205, fig. 4 (1).

Acrioceras nagerai-Leanza, 1970: 206, fig. 5 (1).

Holotype

An external mould found in a loose calcareous nodule in the bed of Fésiles

River, Lake San Martin (Bonarelli & Nagera 1921: 19). It seems that the holotype

is lost, but plaster casts are available (Geological Survey Collection DNGM 9297).

Material

Apart from a plaster cast of the holotype, CPBA 10880-81 from Rio

Cardiel; CPBA 10843 from Puesto La Senalada; CPBA 10830, 11061 from

Puesto Bajo Comision; and CORD-—Pz 4368 from La Federica, Lake San Martin

(collection Dr M. Flores). Rio Mayer Formation. Lower—Upper Aptian.

Description

The very early stage of growth is unknown. The coiling is toxoceratid with an

open initial spire followed by a slightly arcuate shaft and a recurved terminal hook

(Fig. 14).

The whorl section is initially subquadrate or suboctagonal if measured over

the trituberculate ribs (Fig. 15 A—B). It is equidimensional or slightly compressed

(Wh/Wb = 1,00-1,07) with a nearly flat dorsum and moderately curved flanks

converging to a rounded venter. With increasing diameter, the whorl section

becomes more rounded. On the body chamber it is nearly circular with inflated

flanks converging to a broad rounded venter (Fig. 15C).

At the smallest diameter (3 mm) ornament consists of single, rounded ribs,

bearing a small ventral tubercle and separated by wider interspaces. At a slightly

larger diameter (6 mm), the ribs become differentiated and the ornament consists

of alternating tuberculate ribs and non-tuberculate intermediaries, both of equal

strength. With increasing diameter the tuberculate ribs become stronger and

ventrolateral tubercles appear. One to five intermediate non-tuberculate ribs are

present at this stage. At a larger diameter (10 mm) the ornament comprises fine

intermediaries and strong trituberculate ribs, with small umbilical tubercles. The

ribs pass straight, or bend forward in a gentle arc, over the dorsum and run

prorsiradiate over the flanks; while the thin intermediaries cross the venter

without interruption, the strong trituberculate ribs end on the ventral tubercle.

Some ribs are duplicated over the dorsum and unite at the umbilical tubercle,

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 299

Fig. 14. Reconstruction of Toxoceratoides nagerai (Leanza).

B C

Fig. 15. Whorl section of Toxoceratoides nagerai (Leanza). A. CPBA 10881.

B. CPBA 10843. C. CPBA 10880. All x 2.

ANNALS OF THE SOUTH AFRICAN MUSEUM

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ay Ul puNoy o1omM suouMIOeds YOg “T8801 WAdO ‘G-O ‘08801 Vado ‘A-V ‘(ezuRe]) msa8vU saplojwss00x0], “OT “B14

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 301

Fig. 17. Toxoceratoides nagerai (Leanza). A-B. CPBA 10830 from Puesto Bajo Comision.

C. Resin cast of CPBA 10843 from Puesto La Senalada after removing the calcite infilling the

phragmocone. D. CPBA 10843. All x 1.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

while others bifurcate on the umbilical wall and cross the flank as two fine

intermediaries.

At the end of the shaft and beginning of the hook, the lateral and ventral

tubercles disappear and the ornament changes to narrow, simple, sharp ribs that

pass radially around the whorl and to ribs arising in groups of two or three from

an umbilical tubercle. On the final hook these tubercles also disappear and the

ribbing is simple.

The suture line is quite simple with trifid internal, umbilical, and lateral

lobes.

Dimensions

Specimen L Ay Ho jabs

DNGM 9297 c. 80,0 ULL 9,0 5d)

CPBA 10880 c. 90,0 16,0 — 8,0

CPBA 10881 c. 88,0 14,0 — SED)

CPBA 10830 OO Gy 13.0) 9,0 3.0

Discussion

Leanza (1970: 206) referred this species to the genus Acrioceras, pointing out

that it belongs to the ‘Ancyloceras’ tabarelli group (Uhlig 1883: 114, pl. 28

(fig. 2)). He also stated that the curved shaft resembles that of ‘Ancyloceras’

silesiacum illustrated by Uhlig (1883: 142, pl. 28 (fig. 4)). Uhlig (1883), however,

did not refer the latter species to ‘Ancyloceras’ but to “Crioceras’. In any case,

although Acrioceras tabarelli shows superficial similarities with Toxoceratoides

nageral, the coiling and ornament of the latter species plead for its attribution to

Toxoceratoides. |

Toxoceratoides royerianus has a depressed whorl section, with a flat dorsum

and a shaft ornamented with alternate strong trituberculate ribs and fine

intermediaries (Casey 1961: 80, pl. 6 (fig. 2a—b), pl. 17 (fig. 3a—b), text-

fig. 30a—h), while 7. nagerai has a compressed to equidimensional whorl section

and more intermediate ribs on the shaft. Besides, the suture line of the first

species is very incised, while in the latter it is quite simple.

Toxoceratoides rochi Casey is very close to T. royerianus, but differs from

that species and from 7. nagerai by the presence of very strong ribs on the final

hook with very weak umbilical tubercles (Roch 1927: 30, pl. 1 (fig. 4)).

Toxoceratoides saulae Murphy, 1975, differs from T. nagerai mainly in

having an ovoid, depressed whorl section and coarser trituberculate ribs on the

end of the shaft.

Toxoceratoides krenkeli Forster (1975, pl. 4 (figs 1-2)) has a more depressed

whorl section, mainly trituberculate ribs on the shaft with few intermediaries and

coarse ribbing on the final hook.

Toxoceratoides proteus (Spath) shows lateral tubercles on the final hook from

which the bifurcate ribs arise (Casey 1961: 82, pl. 10 (fig. 2a—c)). This feature

allows an easy separation from all the other known species of the genus.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 303

The shaft of 7. starrkingi (Anderson) bears some resemblance to that of

T. nagerai, but the final hook has not only umbilical but also lateral and ventral

tubercles (Anderson 1938: 207, pl. 59 (fig. 4-4A), pl. 65 (fig. 4A)).

Tonohamites decurrens has a Toxoceratoides-like shaft, which resembles that

of Toxoceratoides nagerai, especially in the number of fine intermediate ribs.

However, both species are easily distinguished by the ornament of the final hook.

In the former, there are radial, broad and rounded single ribs (Casey 1961: 80,

pl. 5 (fig. 3a—b), pl. 21 (fig. 2)), while in 7. nagerai the ribbing on the final hook

is sharp, narrow and with some umbilical tubercles.

Leanza (1970, fig. 4 (1)) described a fragmentary specimen of Helicancylus

cf. patagonicus. He stated the close similarities with Stolley’s (1912, fig. 3-3a)

“Ancyloceras’ patagonicum. Although Leanza’s specimen is poorly preserved, it

shows umbilical tubercles on the bend of the hook, as well as bi- and trifurcate

ribs. These features are not present in Helicancylus patagonicus and Leanza’s

specimen is therefore referred to Toxoceratoides nagerai.

Toxoceratoides cf. biplex (von Koenen, 1902)

Figs 18C—D, 19C

Compare:

Ancyloceras? biplex von Koenen, 1902: 381, pl. 49 (figs 10a—b, 11a—b).

Toxoceratoides cf. biplex (von Koenen): Casey, 1961: 83, pl. 20 (fig. 6).

Material

CPBA 10910 from Puesto Bajo Comisi6n, Lake San Martin. Rio Mayer

Formation. Upper Aptian.

Description

One specimen is available, consisting of the end of the shaft and final hook,

and preserved as an internal cast partially covered with the original shell.

The whorl section on the earliest preserved part, which coincides with the

end of the phragmocone, is depressed (Wh/Wb = 0,83), ovoid, with a flat dorsum,

a broadly rounded umbilical edge, strongly inflated flanks, and a broadly rounded

venter. The maximum width is at the dorsal third of the flanks. On the final hook

the whorl section is more rounded, slightly depressed (Wh/Wb = 0,93) with a

narrower dorsum, moderately inflated flanks converging to a rounded venter

(Fig. 18C—D).

Ornament consists of fine, sharp, dense ribs. They run prorsiradiate over the

flanks. On the bend of the crozier there are frequent low-angle bifurcations near

the umbilical margin. On some ribs, slight tubercle-like elevations are present at

the umbilical edge. Over the dorsum the ribs are reduced to striae with a forward

flexure. The upper half of the flanks as well as the venter are heavily abraded, but

it seems that the ribs cross the venter without interruption, and at the distal end

they are simple, dense and radial.

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. A. Whorl section of Toxoceratoides? haughtoni Klinger & Kennedy,

CPBA 10901. x 2. B. Whorl section of Toxoceratoides? sp., CPBA 11049. x 2,5.

C-D. Whorl sections of Toxoceratoides cf. biplex, CPBA 10910. x 2,5.

Discussion

The single specimen compares well with that of Von Koenen (1902, pl. 49

(figs 10b—11b)) in ornament and whorl section, although the latter is more

depressed and with the maximum width on the dorsal half of the flank. Von

Koenen (1902: 381) indicated the presence of lateral tubercles, but did not

illustrate these. In the Patagonian specimen faint elevations can be seen

occasionally on some ribs where the original shell is still preserved.

Drushchits & Kudryavtsev (1960: 295, pl. 39 (fig. 3a—c)) described and

figured two fragmentary specimens of Leptoceras biplex von Koenen. The

presence of strong bituberculate ribs on the shaft and bifurcate ribs on the bend of

the crozier casts doubt on the specific assignation of these specimens. According

to Forster (1975: 162) the Russian material may belong to Toxoceratoides

fustiformis. As the latter species does not have umbilical tubercles on the bend of

the crozier and the shaft is ornamented with trituberculate ribs, it is doubtful that

the Russian material can be assigned to T. fustiformis or to T. biplex.

Casey (1961: 83, pl. 20 (fig. 6)) referred a small fragmentary specimen to

IT. cf. biplex, which resembles the one here described. According to him, the

most characteristic feature of this species is the presence of ‘fine, sharp, wiry

ribbing’.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 305

Toxoceratoides? haughtoni Klinger & Kennedy, 1977

Figs 18A, 19H-J

Toxoceratoides? haughtoni Klinger & Kennedy, 1977: 310, figs 5YA—D, 60A—I, 61A—C, 62A-C,

64A-C, 66B, 79A-B.

? Toxoceratoides (Colomboceratoides) renzoni Etayo Serna, 1979: 20, pl. 6 (fig. 19), text-fig. 3O.

Holotype

SAS 64/T; from locality 168, Mfongozi Creek, northern Zululand, Ap-

tian III-IV (Upper Aptian). South African Geological Survey Collection,

Pretoria. Collected by H. Klinger, 1970.

Material

CPBA 10901 from La Horqueta, Cardiel River (collection Lic. G. Marin),

and CPBA 10849 from Puesto La Senalada, Lake San Martin. Rio Mayer

Formation. Upper Aptian.

Description

Both specimens are fragments of curved shafts. The whorl section is initially

subcircular, slightly compressed (Wh/Wb = 1,07) with a feeble convex dorsum

and rounded flanks converging to a flattened venter. As size increases the whorl

section becomes more laterally compressed (Wh/Wb = 1,25) (Fig. 18A).

At the smallest diameter (Wh=7,5 mm) ornament consists of single

rounded ribs, slightly prorsiradiate and with rounded ventral tubercles, which are

marginal to a siphonal depression. At this stage small ventrolateral tubercles can

also be seen. At a diameter of 10 mm there are two kinds of ribs; some are fine,

non-tuberculate, and the others are strong, high with ventrolateral and ventral

tubercles. Both types alternate regularly. While the first type crosses the venter

without interruption, the second one ends on both sides of a siphonal depression.

Both cross the dorsum straight or slightly curved and run prorsiradiate over the

flanks.

Some of the strong ribs duplicate from the ventrolateral tubercle and on the

lower flank and dorsum they form two fine ribs, while between the ventrolateral

and ventral tubercle there is only a single flat and broad rib. At large diameters,

the intercalatory ribs disappear.

The suture line cannot be traced on the present material.

Discussion

The shaft fragments correspond well with those of Klinger & Kennedy

(1977). These authors assigned this species to Toxoceratoides with doubt because

of the peculiar ornament of the early whorls as well as the absence of simple

ribbing on the recurved crozier. According to Klinger & Kennedy (1977) those

atypical features serve to distinguish 7. ? haughtoni from other species assigned to

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 19. A-B. Helicancylus patagonicus (Stolley), CPBA 10898 from Puesto La Sefalada.

C. Toxoceratoides cf. biplex, CPBA 10910 from Puesto Bajo Comisién. D-F. Tonohamites

aequicingulatus (von Koenen), CPBA 11897 from Loma Pelada. G. Toxoceratoides? sp.,

CPBA 11049 from Puesto La Sefalada. H—-J. Toxoceratoides? haughtoni Klinger & Kennedy.

H-I. CPBA 10849 from Puesto La Sefialada. J. CPBA 10901 from La Horqueta. All x 1.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 307

this genus. Unfortunately neither the early whorls nor the final hook are

preserved in the Patagonian material.

Toxoceratoides krenkeli is to some extent a comparable species, as the

ornament on the shaft is similar to that of T.? haughtoni. However, the former

species has a depressed whorl section and the tuberculation appears at a very

small diameter (Forster 1975; Klinger & Kennedy 1977).

Etayo Serna (1979) proposed a new subgenus and new species of

Toxoceratoides: T. (Colomboceratoides) renzoni. The ornament and whorl

section of the single fragment of this species shows close similarities to

T.? haughtoni, so that T. renzoni may be a junior synonym.

Occurrence

Toxoceratoides? haughtoni occurs in the Upper Aptian of Zululand and

Patagonia.

Toxoceratoides? sp.

Figs 18B, 19G

Material

CPBA 11049 from Puesto La Senalada, Lake San Martin. Rio Mayer

Formation. Upper Aptian.

Description

The single fragment is 80 mm long and comprises the upper part of the shaft

(55 mm) and the beginning of the final hook. The ventral region is heavily

abraded.

At the smallest diameter (Wh = 14 mm) the whorl section is ovoid, laterally

compressed (Wh/Wb = 1,27) with a feeble convex dorsum, slightly curved flanks

and rounded venter. The whorl section, measured over a tuberculate rib, is

subhexagonal (Fig. 18B). As size increases, the whorl section becomes more

rounded.

The shaft is ornamented with strong tuberculate ribs and thin non-

tuberculate intermediaries, which are arranged in an irregular pattern. Both types

cross the dorsum with a forward curvature, are prorsiradiate over the flank and

straight over the venter, where the strong ribs are interrupted. The ribs show two

rows of tubercles: one ventral and the other ventrolateral. On the upper part of

the shaft some bear a third row of small tubercles near the umbilical edge.

As in Toxoceratoides? haughtoni, some of the strong ribs are duplicated at

the ventrolateral tubercle and cross the dorsum as two fine single ribs; between

the tubercles they are broad and flat. Only the dorsolateral part of the end of the

shaft and the beginning of the crozier is preserved. The ornament consists of

single, fine, narrow, dense, non-tuberculate ribs. There is no indication of

umbilical tuberculation.

The suture line, partially exposed, shows relatively high elements and it is

quite incised, with trifid lateral, umbilical, and internal lobes.

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Generic allocation of the specimen is difficult as it shares characteristics of

Toxoceratoides and Tonohamites.

The absence of umbilical tubercles on the bend of the crozier and the

ornament of single ribs are features of Tonohamites rather than of Toxo-

ceratoides. However, in the former genus the ribs are usually broad and rounded,

not thin and sharp as in the Patagonian specimen. The shaft ornament is more

Toxoceratoides-like, but not typical if compared with species like Toxoceratoides

royerianus (Casey 1961: 78, pl. 6 (fig. 2)) or T. krenkeli (Forster 1975: 160, pl. 4

(figs 1-2)). Besides, Tonohamites decurrens has the shaft ornamented as in

Toxoceratoides, which shows the close relationship between both genera, as

stated by Klinger & Kennedy (1977: 319).

The only feature that allows a comparison with another species is the

longitudinal duplication of the strong ribs. This character is also present in

Toxoceratoides? haughtoni. Both species differ markedly in the ornament of the

bend of the crozier.

Finally, Tonohamites and Toxoceratoides show a typical ancyloceratid

pattern in the suture line, with bifid saddles and trifid lobes, but the former

usually has low, simple elements while in the latter the suture line may be more

incised with relatively higher elements.

Based on the suture line and to a lesser extent on the ornament of the shaft,

the present fragment is referred with doubt to Toxoceratoides, aware that the

ornament of the final hook is atypical.

Genus Tonohamites Spath, 1924

Type-species. Tonohamites decurrens Spath, 1924, from the Lower Aptian of

Germany, by original designation.

Diagnosis

Coiling toxoceratid or labeceratid. Ribbing usually rounded, tuberculation

may be present, but the tubercles are weak and mostly confined to the venter. On

the body chamber the ribs are simple, strong, rounded or flat and non-

tuberculate. Suture line simple, with bifid saddles and trifid lobes.

Discussion

The type-species of this genus is difficult to interpret, and nomenclatural

problems are involved. Casey (1961: 84) extensively discussed this point. Wright

(1957: L212) regarded Tonohamites as a synonym of Hamiticeras, while Casey

(1961: 84) maintained the genus and gave the first diagnosis.

The type-species of Tonohamites does not show the main features present in

other species assigned to this genus. In fact, the ornament of the shaft, with strong

trituberculate ribs, closely resembles that of Toxoceratoides and isolated

fragments can hardly be distinguished. However, the ornament of the body

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 309

chamber of Tonohamites, with broad, rounded, or flat ribs, allows an easy

separation from Toxoceratoides, which shows fine, sharp, single ribs intercalated

with bi- or trifurcate ones arising from umbilical tubercles on the final hook.

Hamiticeras Anderson, as interpreted here, differs from Tonohamites in its

coarse trituberculate and intermediate ribs on the shaft and its long, parallel final

hook with radial, sharp, high ribbing.

Helicancylus Gabb, redefined here, is easily distinguished from Tonohamites

by the complete lack of minor ribbing, and the presence of tubercles on every rib

on the shaft and on the recurved crozier.

According to Klinger & Kennedy (1977), Casey (1961) referred the following

species to Tonohamites:

Tonohamites decurrens Spath (1924: 85). (Lectotype is the specimen illustrated by

Von Koenen 1902, pl. 33 (fig. 2, and the lower part of fig. 3a).)

Tonohamites aequicingulatus (von Koenen) (1902: 394, pl. 37 (figs 5a—c, 6a—e)).

Tonohamites? hunstantoniensis Casey (1961: 90, pl. 21 (fig. 1a—d)).

Tonohamites koeneni Casey (1961: 89). (Holotype is the specimen illustrated by

Von Koenen 1902, pl. 33 (fig. 3a, upper part only).)

Tonohamites limbatus Casey (1961: 89, pl. 21 (fig. 3a—b), pl. 22 (figs 3a—c, 4)).

Tonohamites? eichwaldi (Jasykow) (in Sinzow 1872: 36, pl. 6 (figs 7—9)).

Tonohamites? undosus (von Koenen) (1902: 393, pl. 35 (fig. 13a-f)).

It is interesting to mention that Casey (1961) noted similarities between some

fragments of Tonohamites aequicingulatus and Hamites? undosus von Koenen.

He also indicated that the latter species is only known by a small fragment of the

shaft and that it is really difficult to decide whether it is a separate species. Finally,

Casey decided.to join all the fragments as belonging to a single, variable species

and placed 7.? undosus as a possible synonym of T. aequicingulatus.

To the list given above must be added Tonohamites? caseyi Klinger &

Kennedy (1977: 324, figs 46, 49) and perhaps Tonohamites? taylori (Etheridge).

Etheridge (in Jack & Etheridge 1892: 498, pl. 42 (fig. 13)) described and figured

one specimen of ‘Ancyloceras’ taylori showing a tightly coiled initial spire

followed by a straight shaft, both ornamented with simple annular ribs. Later the

same author (Etheridge 1909: 162, pl. 49 (figs 3-6)) included that fragment with

other specimens and described them all as ‘Crioceras’ taylori.

The type specimen of ‘Ancyloceras’ taylori was placed in Toxoceratoides by

Whitehouse (1926), who figured a small additional fragment, while the specimens

of ‘Crioceras’ taylori were included in the Albian genus Labeceras Spath.

Finally Day (1974: 14) noted that ‘Ancyloceras’ taylori might be placed more

suitably in Tonohamites than in Toxoceratoides. This view is supported by the

Tonohamites-like ornament of the small fragment illustrated by Whitehouse

(1926: pl. 36 (fig. 5)).

Kakabadze (1981) recorded Tonohamites picteti (Ooster) from the Lower

Aptian of southern USSR. As he did not describe nor figure the specimens, it is

not possible to decide if they belong to this genus or not. Ooster’s (1857, pl. 50

(figs 1-6)) type specimen of ‘Ancyloceras’ picteti does not resemble Tonohamites.

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

Specific differentiation within the genus Tonohamites is rather difficult,

especially when dealing with fragments. It is based mainly on the coiling,

ornament, and whorl section.

Occurrence

Tonohamites occurs in the Lower Aptian of Germany (Von Koenen 1902),

Spain (Martinez 1982), England (Casey 1961), and ?southern USSR (Kakabadze

1981). It also occurs in the Upper Aptian of Madagascar (Collignon 1962) and

Zululand (Klinger & Kennedy 1977). This is the first record from Patagonia

(sce Fig] 8):

Tonohamites aequicingulatus (von Koenen, 1902)

Fig. 19D-F

Hamites aequicingulatus von Koenen, 1902: 394, pl. 37 (figs Sa—c, 6a—e).

Tonohamites aequicingulatus (von Koenen): Casey, 1961: 87, pl. 9 (figs 2a—b, 3a—b, 4), text-

fig. 32. Klinger & Kennedy, 1977: 322, figs 38C, 68A-—E, 88D.

Tonohamites sp. aff. aequicingulatus (von Koenen): Collignon, 1962: 14, pl. 221 (fig. 960).

Lectotype

The original of Von Koenen (1902, pl. 37 (fig. Sa—c)) from the Lower Aptian

of northern Germany, by subsequent designation of Casey (1961: 87).

Material

CPBA 11897 from Loma Pelada, Tucu-Tucu. Rio Belgrano Formation.

Lower Aptian.

Description

The small part of a shaft, 35 mm long and preserved as an internal cast, is

partially covered with the original shell.

The whorl section is ovoid, depressed (Wh/Wb = 0,75-0,77), with flat

dorsum, slightly convex flanks, and broadly rounded venter. Ornament consists

of annular, rounded ribs. They are nearly radial on the dorsum and prorsiradiate

on the flanks. There are four ribs within a distance equal to the whorl diameter. In

the early stage some ribs bear siphonal, lateral and umbilical tubercles; these are

very small and rounded and disappear with increase in size.

The partially exposed suture line is simple with asymmetrical lateral lobe.

Discussion

As Casey (1961: 88) noted, this species is only known from fragments. The

Patagonian specimen resembles both the European (Von Koenen 1902; Casey

1961) and Zululand (Klinger & Kennedy 1977) material. The only apparent

difference between those specimens and the present fragment is that in the latter

some early ribs bear three rows of tubercles, whilst the figured specimens show

none or a ventral row only (Von Koenen 1902, pl. 37 (fig. 5); Casey 1961, pl. 9

(fig. 2b), text-fig. 32b).

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN Silat

The Madagascan specimen figured by Collignon (1962, pl. 221 (fig. 960))

shows a more rounded whorl section and the ribs are stronger. It has tentatively

been referred to this species.

Tonohamites decurrens Spath has strong trituberculate ribs separated by

intermediaries on the shaft (Casey 1961: 86, pl. 21 (fig. 2), pl. 5 (fig. 3a)).

Tonohamites limbatus Casey has a slightly compressed subrectangular whorl

section and strongly prorsiradiate, narrow ribs (Casey 1961: 89, pl. 20

(figs 3a—c, 4)).

According to Klinger & Kennedy (1977: 322) the body chamber of

T. koeneni Casey resembles that of 7. aequicingulatus, but the latter species

shows ventral tubercles on the shaft.

Martinez’s (1982: 142, pl. 24 (fig. 6a—b), text-fig. 23) small shaft fragment

referred to Tonohamites sp. has comparable ornamentation and whorl section. It

differs, however, in its more simple suture line at the same diameter.

Occurrence

Tonohamites aequicingulatus is known from the Aptian of Germany (Von

Koenen 1902) and Lower Aptian (bowerbanki zone) in England (Casey 1961).

The Madagascan and Zululand specimens are from the Upper Aptian (Collig-

non’s (1962) Aconeceras nisus and Melchiorites melchioris zone, and Kennedy &

Klinger’s 1975 Aptian HI-IV—see Klinger & Kennedy (1977)).

CONCLUDING REMARKS

The most important results of this study are:

— The redefinitions of Helicancylus (type-genus of the subfamily Helicancylinae)

and of Hamiticeras clarify the systematics of the subfamily Helicancylinae.

— Helicancylus, as here interpreted, accommodates some species (one of which

is Helicancylus patagonicus) of hitherto uncertain affinities.

— The systematic study allows the recognition of three genera of this subfamily

in the Austral Basin: Helicancylus, Toxoceratoides and Tonohamites. They are

recorded for the first time in this basin. Seven species have been identified.

— The representatives of this subfamily are locally common in several horizons

of the Rio Mayer Formation and in one level of the Rio Belgrano; all are of

Aptian age. Their recognition means an important increase in our knowledge of

the Aptian biostratigraphy of the northern part of the Austral Basin.

— The identified fauna facilitates the correlation with previously known

assemblages, especially from western Europe and south-eastern Africa.

ACKNOWLEDGEMENTS

The Consejo Nacional de Investigaciones Cientificas y Técnicas, Argentina,

supported this study through a research scholarship at the South African

Museum. The South African Museum helped to make this publication possible.

312 ANNALS OF THE SOUTH AFRICAN MUSEUM

The Servicio Geologico Nacional, Argentina, provided the means for fieldwork.

To these institutions I am particularly indebted.

I am most grateful to Dr H. Klinger (South African Museum) for his advice

and stimulating discussions during my stay in Cape Town; to Dr V. Ramos

(Servicio Geoldégico Nacional) for valuable help in the field and criticism on the

stratigraphy; to Dr A. Riccardi (Museo de La Plata) and Dr M. R. A. Thomson

(British Antarctic Survey) for useful discussions; and to Dr R. Levi (Servicio

Geoldgico Nacional), Dr M. Hiinicken (Universidad Nacional de Cordoba), and

Dr A. Riccardi (Museo de La Plata) who kindly lent original material.

Special thanks are also due to Lic. G. Marin and Lic. M. Palma (Servicio

Geoldgico Nacional) for their help in the field; to Miss S. Dove (South African

Museum) for taking the photographs, and to Miss J. Blaeske and Mr V. Branco

(South African Museum) for preparing the illustrations.

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Drusucuits, V. V. & Kupryavtsev, M. P. 1960. [Atlas of the Lower Cretaceous fauna of the

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ETHERIDGE, R. 1909. Lower Cretaceous fossils from the sources of the Barcoo, Ward and Nive

rivers. Part 2. Cephalopoda. Rec. Aust. Mus. 7: 135-188.

APTIAN HELICANCYLINAE FROM ARGENTINIAN BASIN 313

Forster, R. 1975. Die geologische Entwicklung von Sid-Mozambique seit der Unterkreide und

die Ammoniten-Fauna van Unterkreide und Cenoman. Geol. Jber 12: 3-324.

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1: i-xx, 1-243.

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Hatcuer, J. B. 1897. On the geology of southern Patagonia. Am. J. Sci. (4) 4: 327-354.

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oriental Portuguesa). Bolm Servs Ind. Minas Geol. Lourengo Marques 17: 3-24.

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Graham Land. Scient. Rep. Falkld Isl. Depend. Surv. 21: 1-16.

HowartH, M. K. 1962. The Yorkshire type ammonites and nautiloids of Young and Bird,

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Argentina. Actas II° Congr. Geol. Chileno 1 (B): 1-32.

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

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France. Mém. Soc. géol. Fr. (N.S.) 43 (101): 1-78.

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Tuomson, M. R. A. 1982. A comparison of the ammonite faunas of the Antarctic Peninsula and

Magellanes Basin. J. geol. Soc. 139: 763-770.

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Wien. 46: 127-290.

WACHENDORF, H. 1967. Zur Unterkreide-stratigraphie von Sud-Mozambique. Neues Jb. Geol.

Paldont. Abh. 129: 272-303.

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195-242.

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Aptychi. Abh. math.-naturm. Kl. Wiss. Mainz 1962: 1-148.

WIEDMANN, J. 1966. Stammesgeschichte und System der postriadischen Ammonoideen. Ein

Uberblick. Neues Jb. Geol. Paldont. Abh. 125: 49-79; 127: 13-81.

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HALLAM, A. ed. Atlas of palaeobiogeography: 309-316. Amsterdam: Elsevier.

Wricut, C. W. 1957. Mesozoic Ammonoidea. In: Moore, R. ed. Treatise of invertebrate

palaeontology. Part L, Mollusca 4: L80-—L465. Kansas: Geological Society of America &

University of Kansas Press.

Younc, G. & Birp, J. 1828. A geological survey of the Yorkshire coast. Whitby.

Manuscript received February 1984.

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

FIscHER, P. H. 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

FiscHEeR, P. H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

expérimentale et générale 74 (33): 627-634.

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

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische und anthro-

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 96 Band

December 1986 Desember

Part 8 Deel

ay LR

SSS

SB: 9.9.9.9

2 4

UPPER BARREMIAN HETEROCERATINAE

(CEPHALOPODA, AMMONOIDEA)

FROM PATAGONIA AND ZULULAND,

WITH COMMENTS ON THE

SYSTEMATICS OF THE SUBFAMILY

MARIA BEATRIZ AGUIRRE URRETA

HERBERT CHRISTIAN KLINGER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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UPPER BARREMIAN HETEROCERATINAE

(CEPHALOPODA, AMMONOIDEA)

FROM PATAGONIA AND ZULULAND, WITH COMMENTS ON THE

SYSTEMATICS OF THE SUBFAMILY

MARIA BEATRIZ AGUIRRE URRETA

Departamento de Geologia, Facultad de Ciencias Exactas y Naturales,

Universidad de Buenos Aires, Argentina

HERBERT CHRISTIAN KLINGER

Department of Invertebrate Palaeontology, South African Museum, Cape Town

(With 26 figures)

[MS accepted 17 September 1985]

ABSTRACT

Representatives of the ammonite subfamily Heteroceratinae from the province of Santa

Cruz, Patagonia, Argentina, include Heteroceras (H.) elegans Rouchadzé and Colchidites

vulanensis Egojan australis Klinger, Kakabadze & Kennedy. Examination of this material and

that of Zululand casts doubt on the current systematic concepts applied within the subfamily,

shows distinct size-related dimorphism in Colchidites, and further illustrates the scope of

intraspecific variation.

CONTENTS

PAGE

JIVE ETECONGNEUCTONOVIN Pas cease, Gee ona keenest air Oe ene eae 316

EE OCANOMOMSPECIMNENS Meme te ae fate an Ge ea a ae 316

Field localities

| PENTENOVTUIENSS 5 2. cr pPOMn ee ean Cnn Ree ne ONT RO ra shed, ner mag ar Venti rer 316

ATO NU ENING |. 6s GaSe AE oP Re ae ee Gat 316

SUCUREMCHIMIMNOLOCV erat ixe te alee ee ees rr ieee waht ag 6 3, 4 39)

Systematic palaeontology

Subfamilyseleteroceratinacy- «a. aee nee eee see oo: 319

GeMUSHHCICKOCENAST A oe oats sens anh Pee es 322

GemMUSEOlCHIGICS We wee nbs vet ai ee a oe 325

Discussion

Validity of genera and species groups ............. 343

Sexualidimorphisiitwes. nner cee oe eee a 350

IntKaspecwiCvanlablon omens | see cee ron 350

SUUIPOUTEN AT soe StS aNa oA or Gaia tee alia he ta miei Oa ee Hana Papa re oe PR 350

List of species of Heteroceras and Colchidites ............... Soil

PNCKMOWICAEEMICMIES, Minas nee cues yh Woe © Sale es ee os we 35

INGIETEM CE SIs ery ree eI Met uh a. Sed oS rate eed 355

SMS)

Ann. S. Afr. Mus. 96(8), 1986: 315-358

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

The Upper Barremian ammonite subfamily Heteroceratinae Spath, 1922, is

best known from the south-western part of the U.S.S.R. Recent description of

abundant representatives of this heteromorph group from Zululand, South Africa

(Klinger 1976; Klinger et al. 1984) has shown distinct faunal similarities with the

Caucasus; a trend that continues well into the Aptian, as indicated by other

heteromorph ammonite groups (Forster 1975a, 1975b; Klinger & Kennedy 1977).

Blasco et al. (1980) recorded the first occurrence of the heteroceratine genus

Colchidites from Tucu Tucu in the province of Santa Cruz, Patagonia, Argentina.

Since then, more material was collected by Aguirre Urreta in Patagonia for

comparison with the Zululand and Caucasian material. Also, additional material

from Zululand became available.

This material casts doubt on the current systematic concepts applied in the

subfamily and displays the extensive range of intraspecific variation within the

group. In addition, size-related dimorphism is demonstrated in the group for the

first time.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the repository of material:

SAM = South African Museum, Cape Town

BMNH = British Museum (Natural History), London

GIAS = Geological Institute, Academy of Science, Georgian S.S.R., Tbilisi

CPBA = Catedra de Paleontologia, Facultad de Ciencias Exactas y Naturales,

Universidad de Buenos Aires

UP = Geology Department, University of Pretoria (Boshoff collection)

SAS = Geological Survey, Pretoria, South Africa

FIELD LOCALITIES

Patagonia

Most of the Patagonian material was collected at two localities: Chorrillo del

Medio and Loma Pelada, situated in the central-western province of Santa Cruz,

approximately 48°25’S 72°00’ W (Fig. 1). These were described by Aguirre Urreta

(1983) and details of the sections are shown in Figure 2. Only one specimen was

collected at locality Cerro Cornillos, situated south of the former, 10 km north of

Lake San Martin, approximately 48°47'S 72°23'W.

Zululand

All the Zululand material was collected at locality 170 of Kennedy & Klinger

(1975: 302, fig. 11), cliff and gully sections 2 km north-west of Mlambongwenya

Trading Store on the north side of the Mlambongwenya Stream, 27°10'10"S

32°10'13"E. Details of the section are shown in Figure 3, which is a composite for

this locality and extends over several hundred metres. It corresponds to

Haughton’s (1936: 293) localities L7-L,3.

UPPER BARREMIAN HETEROCERATINAE S19)

ARGENTINA

Laguna Moyano

o Sterea

PROV.

SANTA CRUZ

RIO

GALLEGOS

LEGEND

(| Quaternary Dep.

: : . Kachaike Fm.

Ea Tucu-Tucu Rio Belgrano Fm.

48°30' A Rio Mayer Fm.

LOWER

CRETACEOUS

+ Fossiliferous locality

Fig. 1. Locality map of Patagonian exposures.

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

LEGEND

CHORRILLO

DEL MEDIO FOSSILS

Ammonites

LOMA O Hatchericeras spp.

PELADA ri Colchidites (C.) vulanensis

australis

Sanmartinoceras africanum

Kachaike Fm.

Emericiceras sp.

Kachaike Fm.

Heteroceras (H.) elegans

Tropaeum (T.) sp.

Tonohamites aequicingulatus

Heteroceratinae indet.

Belgrano

Bivalves

4 Apiotrigonia sp.

Crustaceans

# Hoploparia longimana

* Enoploclytia sp.

wx Protocallianassa patagonica

Rio Mayer Fm.

LITHOLOGY

== Black shales

Rio Mayer Fm.

Black shales with calcareous nodules

| Sandstones

©. Sandstones with concretions

“NM"| Cross-bedded sandstones

0 Cee) 9.

0°56 oo | Conglomerates

TT T| tufts

Potty Limestones

Fig. 2. Stratigraphic section at Loma Pelada and Chorrillo del Medio, Patagonia.

At least 10m o

buff concretion}

Ancyloceras, Tr

Prominent concn

Tropaeum and ot|

1*6m poorly ex

2m poorly expo

+| Concretions wit

ancyloceratid f|

1e5m silts wit

The APTIAN/BARR

mo] Concretions wit

Silts with scat

molluscs, inclu(

Concretions crov

Silts

Concretions crov

occasional aconé

Glauconite-rich

r= ee Concretions with

. -:|Colchidites and

Grey-buff, burrdq

Grey-buff, burrd

Concretions with

Heteroceras and

Poorly exposed g

Concretions with

and trigoniids.

Fig. 3. Stratigraphic

UPPER BARREMIAN HETEROCERATINAE

At least 10m of poorly exposed silts with brittle pale

buff concretions yielding Cheloniceras spp., Valdedorsella?,

Ancyloceras, Tropaeum, Australiceras and other forms

ed ee TOF OR IGULLY.

Prominent concretion layer with Procheloniceras?

Tropaeum and other ancyloceratids

“f 1*6m poorly exposed silts

Concretions with abundant Procheloniceras?

2m poorly exposed weathered silts

Concretions with abundant Procheloniceras? and

ancyloceratid fragments

(ea a pe a RRS BE

1*5m_ silts with scattered concretions.

The APTIAN/BARREMIAN BOUNDARY is at this level

Concretions with trigoniids and rare Colchidites

Silts with scattered concretions yielding diverse

molluscs, including Colchidites

£)Concretions crowded with Colchidites

Silts

Concretions crowded with Colchidites, Panopea and

occasional aconeceratids

4] Glauconite-rich silts with molluscs

wo [oo eo

)Concretions with molluscs, including Phylloceras,

Colchidites and Sanmartinoceras?

Grey-buff, burrowed silts with bivalves and wood debris

VEFES eS

Grey-buff, burrowed silts with bivalves and wood debris

sCey,\ Concretions with molluscs, notably trigoniids,

Heteroceras and Colchidites

Poorly exposed silts

AlConcretions with drifted molluscs, e.g. Gervillella

pnd trigoniids. Logs common

5 Buff silts with molluscs and large Teredo-bored Togs

26 }*:

Concretions with 'Crioceratites'

Grey-buff, burrowed silts with wood debris and

drifted molluscs

Teredo-bored logs occur

Concretions with wood and drifted molluscs,

including ‘Crioceratites' and aconeceratids

so 8 [ee ea

Caw w

Poorly exposed silts with huge doggers, crowded

With molluscs, including abundant ammonites:

‘Emericiceras', 'Acrioceras', Eulytoceras,

PhyTloceras serum, Sanmartinoceras?, Guvenile

aconeceratids and hemihoplitids. Large oyster—

encrusted and Teredo-bored logs common

TOP OF RIVER CLIFF

Calcareous concretions with many molluscs,

eg. Megatrigonia, Panopea

Buff, burrowed silts

Shelly concretions with abundant bivalves (trigoniids,

Gervillella). 'Crioceratites' sp.

eal steal

Wa es

Poorly exposed, grey-buff burrowed and cross-bedded

silts with scattered concretions

Drifted molluscs common

Concretions crowded with drifted molluscs and

& large logs. Megatrigonia, Gervillella

Sg) SS

Buff, burrowed silts r

Shelly concretions

Buff, burrowed silts a

Concretions with Phylloceras, 'Crioceratites' and :

Colchidites

Concretions with drifted molluscs and logs

Buff silts with drifted molluscs

3m grey-buff bioturbated silts with traces of

cross-bedding. Layers of drifted molluscs

common, yielding heterodonts, trigoniids,

Gervillella and oysters.

Wood debris frequent

Calcareous concretions with juvenile aconeceratids

3m Grey-buff, burrowed silts with traces of cross-

beddings Abundant drifted molluscs and wood debris,

as below

Calcareous concretions. A belemnite fragment

Grey-buff bioturbated silts with traces of cross-

bedding. Large logs common, together with oyster

bioherms. Orifted molluscs - oysters, Grvillella,

Megacucul laea, Exogyra, trigoniids and heterodonts

Concretions with abundant molluscs including

juvenile aconeceratids

Grey-buff burrowed silts with traces of cross-bedding.

Orifted molluscs include Phylloceras serum

Calcareous concretions with Megatrigonia and

Megacucullaea

3m Grey-buff, burrowed silts with traces of cross-

bedding. Abundant drifted molluscs, including

Steinmanella henningi. Large Megacucullaea in

Tfe position. lyster bioherms up to 2m diameter,

Dri ftwood

Concretions with abundant molluscs, including

Eulytoceras, Crioceratites, juvenile aconeceratids

an gatrigonia

RIVER BED

Fig. 3. Stratigraphic section at locality 170, Mlambongwenya Creek, Zululand. Reproduced with permission of British Museum (Natural History),

London.

UPPER BARREMIAN HETEROCERATINAE 319

Note should be taken of the different lithologies. In Zululand the fauna

occurs in a near-shore facies of silts and sandstones with abundant plant material,

including well-preserved impressions of Zamites recta and Cladophlebis dundro-

diensis. In Patagonia the dominant lithologies are anoxic black shales, with

several levels of calcareous nodules in which the fossils are preserved.

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916; see Kullmann & Wiedmann

1970 for a recent review) is followed in the present work.

I = internal lobe; U = umbilical lobe; L = lateral lobe; E = external lobe.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA Cuvier, 1797

Class CEPHALOPODA Zittel, 1884

Order AMMONOIDEA Zittel, 1884

Suborder ANCYLOCERATINA Wiedmann, 1966

Superfamily ANCYLOCERATACEAE Gill, 1871

Family Heteroceratidae Spath, 1922

Subfamily Heteroceratinae Spath, 1922

The subfamily Heteroceratinae Spath, 1922, has a virtually world-wide

distribution (see e.g. pp. 322, 325) but is best known from the south-western part

of the U.S.S.R. (Georgia, Caucasus and Turkmenia) through the monographical

studies of Djanélidzé (1926), Rouchadzé (1933), Eristavi (1955), Egojan (1965),

Kotetishvili (1970) and Kakabadze (1967, 1971a, 1975); especially the latter. The

systematics of the subfamily are based mainly on the work of Rouchadzé (1933),

and subsequently elaborated by Kakabadze (1967, 1971a, 1975). According to

Kakabadze (1967, 1971a, 1971b, 1975) the group is elevated to familial level, and

consists of two subfamilies that are further subdivided as follows:

Heteroceratinae Spath, 1922 Colchiditinae Kakabadze, 1967

Heteroceras (Heteroceras) d’Orbigny, 1850 Imerites Rouchadzé, 1933

Heteroceras (Argvethites) Rouchadzé, 1933 Fristavia Kakabadze, 1967

Hemibaculites Hyatt, 1900 Colchidites Djanélidzé, 1926

Paraimerites Kakabadze, 1967

With the exception of Hemibaculites, which is an enigmatic genus and of

uncertain affinities, all the above genera are closely related, both morphologically

and phylogenetically.

Division at subfamilial level is based on coiling. In Heteroceratinae an initial

helix is immediately followed by an uncoiled section, which may either be a

straight shaft or a broadly curved section, ending in a recurved hook of which the

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

shafts may be parallel or divergent. In the Colchiditinae an initial helix is followed

by a planispirally coiled section of variable duration, which embraces the initial

helix, finally uncoiling into a shaft and recurved hook. In some forms an uncoiled

section has not yet been recorded, and it is suspected that they may reach

maturity in the planispirally coiled stage.

As discussed earlier (Klinger 1976: 8) division into separate subfamilies as

advocated by Kakabadze is considered superfluous.

Differentiation at generic or subgeneric level is based mainly on the presence

or absence of tuberculation. Heteroceras s.s. is non-tuberculate throughout,

whereas H. (Argvethites) has a pair of ventral tubercles on the shaft. Colchidites is

non-tuberculate throughout; Paraimerites has a pair of variably developed ventral

tubercles on part of the planispiral section and Evistavia and Imerites are both

quadri-tuberculate on part of the planispiral whorls. Ervistavia differs from

Imerites in having intercalatory and bifurcating ribs; the latter originate at the

ventrolateral tubercles and continue over the dorsum.

Within the genera Heteroceras and Colchidites, several species groups have

been recognized. These were already identified by Rouchadzé (1933) and

subsequently either retained or elevated in rank by Kakabadze (1967, 1971a,

1975). Within Heteroceras s.s. Rouchadzé (1933) recognized two species groups:

Heteroceras astieri d’Orbigny and Heteroceras helicoceroides Karsten (misspelled

heliceroides). The group of H. astieri includes large forms with a high helix, and

axis of coiling oblique to the dorso-ventral symmetry of the shell. The shaft is long

and curved, with a recurved crozier. The group of H. helicoceroides includes

small forms with the axis of coiling of the helix oblique to the plane of symmetry

of the shell, with a slender, straight shaft and relatively long recurved crozier.

Heteroceras (Argvethites) is similar in size to the latter species group, but differs

by the possession of a row of ventral tubercles on the shaft, and often a siphonal

depression on the crozier.

In Colchidites, Rouchadzé (1933) identified three species groups:

(1) Colchidites gr. ex intermedius Djanélidzé, with a well-developed helical stage

consisting of 5—8 whorls, a poorly developed planispiral part not exceeding

one whorl, and a well-developed shaft and hook.

(2) Colchidites gr. ex colchicus Djanélidzé, with 4-7 whorls in the helical stage,

one or two planispiral whorls, and a relatively well-developed uncoiled part.

(3) Colchidites gr. ex shaoriensis Djanélidzé with a low helix consisting of

2-3 whorls and a well-developed planispiral section with two or more

whorls. An uncoiled part has as yet not been found in this species group.

According to Kakabadze (1975) these three species groups of Colchidites

form a phylogenetic sequence, demonstrating yet another trend towards recoiling

amongst heteromorphs (cf. Wiedmann 1969)—starting with Heteroceras at the

one end, and ending at the other with a near-ammonitic coiled, streamlined form,

C. gr. ex shaoriensis, which in turn may have given rise to the ‘normally’ coiled

Deshayesitidae. It is unknown whether similar species groups and trends are

UPPER BARREMIAN HETEROCERATINAE 321

present in Paraimerites, as only one, monotypical uncoiled species, P. breviheli-

coides Kakabadze, is known. No uncoiled forms are known in either J/merites or

Eristavia. Both genera appear stratigraphically before Paraimerites and Colchi-

dites from a common ancestor with Heteroceras, but range into the uppermost

Barremian with Paraimerites and Colchidites.

The validity of this classification will be discussed below on the basis of the

Patagonian and Zululand material.

Fig. 4. Lumachelle with Colchidites from locality 170, Mlambongwenya Creek, Zululand. Note

the co-occurrence of fine and coarsely ribbed forms. SAM—PCZ6393. X 0,65.

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Heteroceras s.s. d Orbigny, 1850

Type-species. Turrilites emerici d Orbigny, 1842, from the Upper Barremian

of south-western France.

Diagnosis

Open-coiled heteromorphs with an initial helix followed by a straight or

curved shaft and recurved crozier. Coiling in helix dextral or sinistral, with whorls

tightly coiled or scarcely in contact. Ornament on helix consists of sinusoidal,

single ribs. On the shaft and on the bend of the crozier, there are single,

bifurcating and intercalatory ribs. These cross the dorsum with a slight forward

curvature, sometimes duplicating there, and straight over the venter. In some, a

slight siphonal depression may interrupt ribbing over the venter on part of the

shaft and/or crozier. Suture line rather simple with low elements.

Occurrence

Heteroceras s.s. occurs in south-western U.S.S.R. (Georgia, Caucasus,

Turkmenia) (Rouchadzé 1933; Eristavi 1955; Egojan 1965; Kotetishvili 1970;

Kakabadze 1975), Bulgaria (Nikolov 1964; Dimitrova 1967), Hungary (Fulop

1964), Czechoslovakia (Silesia) (Uhlig 1883), France (d’Orbigny 1842, 1850,

1851; Kilian 1888a, 1888b), California (Murphy 1975), Colombia (Karsten 1858,

1886; Royo y Gomez 1945), Canada (Jeletzky 1970), Japan (Obata & Ogawa

1976; Obata et al. 1976), Zululand (Klinger 1976; Klinger et al. 1984; herein),

Patagonia (herein).

It has also been listed in Algeria (Blayac 1905; Simionescu 1905), Morocco

(Roch 1930), Tunisia (Burollet et al. 1983), Tanzania (Spath 1930), Spain

(Simionescu 1905) and Romania (Simionescu 1905). Reference to this genus in

Peru seems to be due to geographic misinterpretation of Karsten’s (1856, 1886)

works.

Many unsubstantiated references to Heteroceras are probably misidentifica-

tions of Upper Cretaceous nostoceratids.

Heteroceras (Heteroceras) elegans Rouchadzé, 1933

Figs 5, 10E-I

Heteroceras elegans Rouchadzé, 1933: 232, pl. 13 (fig. 4), text-fig. 36. Eristavi, 1955: 118.

Kakabadze, 1975: 89, pl. 3 (figs 1, 3-5), pl. 8 (fig. 5).

Heteroceras astierianum d’Orb.: Drushchits & Eristavi, 1958, pl. 49 (fig. 3).

Heteroceras (H.) sp. aff. H. (H.) astieranum d’Orbigny: Klinger, 1976: 11, pl. 1 (figs 1-2), text-

figs Sa—c, 6a.

Heteroceras (Heteroceras) elegans Rouchadzé: Klinger, Kakabadze & Kennedy, 1984: 44,

figs 2A-F, 3A-G, 4A-C.

Type

Lectotype by subsequent designation (as holotype) by Kakabadze (1975) is

the specimen figured by Rouchadzé (1933, pl. 13 (fig. 4)) from western Georgia,

Goresha, Upper Barremian zone of Colchidites securiformis, housed in the

UPPER BARREMIAN HETEROCERATINAE 323

collections of the Geological Institute of the Academy of Science, Georgian

S.S.R., cat. no. GIAS 350/1016.

Material

CPBA 11898-11900 from Loma Pelada, Tucu Tucu, CPBA 11121 from

Cerro Cornillos, Lake San Martin, Patagonia, Rio Mayer Formation, Upper

Barremian, and SAS H54/29hi, BMNH C80025a—c, 80026a—b, 80028-9 from

locality 170, Mlambongwenya Creek, northern Zululand, Makatini Formation,

Upper Barremian.

Description

The helix consists of 4-5 contiguous whorls, coiled either dextrally or

sinistrally. The last helical whorl passes directly into the straight to slightly curved

shaft. The recurved crozier is relatively long, and nearly parallel to the shaft.

The whorl section in the helix is subcircular; on the shaft it varies from

slightly compressed to depressed, and on the crozier it becomes equidimensional

to subquadrate.

Ornament on the helix consists of fine, single, sinusoidal ribs. Ribbing on the

shaft is usually single and slightly prorsiradiate, but at the end of the shaft and in

the bend of the crozier bifurcating ribs are common. On the final part of the

crozier the ribs are single, slightly curved and separated by interspaces wider than

themselves.

A siphonal depression is present in some, but not all of the Zululand and

Patagonian specimens (cf. Fig. 5A). It may already occur on the shaft, or only on

the crozier, or on both. Ribs may be slightly thickened here, but no distinct

tubercles develop.

Comparison

The Patagonian and Zululand material compares favourably with the

specimens of H. (H.) elegans figured from western Georgia (Rouchadzé 1933;

Kakabadze 1975; Klinger et al. 1984). According to Kakabadze (1975: 66) a

siphonal depression occurs in the majority of tuberculate heteroceratids, 1.e.,

H. (Argvethites). Our material shows that this feature also occurs at random in

non-tuberculate forms, and seems to be of very low taxonomic value.

The specimens here described also resemble the single helix and shaft

fragment referred to Heteroceras aff. astieri by Obata & Ogawa (1976, pl. 4

(ina)

Heteroceras (H.) elegans differs from H. (H.) helicoceroides (Karsten) by the

presence of chevron-like ornament on the venter of the latter—a rather

Hemibaculites-like feature.

In H. (H.) eristavii the helix does not pass directly into the shaft; the latter is

shorter and the recurved crozier forms a very tight bend (Kakabadze 1975: 90,

pl. 4 (figs 1, 3), pl. 5 (fig. 5), pl. 8 (fig. 7)).

324

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Heteroceras (Heteroceras) elegans Rouchadzé, 1933. A—C.CPBA 11898.

D-F. CPBA 11121. G—H. SAS H54/29hi. All x 1.

UPPER BARREMIAN HETEROCERATINAE 325

Heteroceras (H.) vermiforme Rouchadzé is only known from fragments. It is

a very small species, with fine ribbing and an open, recurved crozier (Rouchadzé

1933, pl. 13 (fig. 5); Kakabadze 1975, pl. 3 (figs 6-8), pl. 4 (fig. 2)).

According to Kakabadze (1975: 92), H. (H.) isocostata Kakabadze is related

to H. (H.) elegans, but differs in its greater size, stronger ornament on the helical

part and the presence of bifurcating ribbing at the beginning of the shaft.

Occurrence

Upper Barremian of northern Caucasus, western Georgia, Zululand and

Patagonia.

Genus Colchidites Djanélidzé, 1926

Type-species. Colchidites colchicus Djanélidzé, 1926, from the Upper

Barremian of Georgia.

Diagnosis

Three distinct coiling modes occur during ontogeny—an initial helical spire

consisting of between two and eight whorls, followed by a planispiral, discoidal

section coiled more or less at right angles to the spire, which in turn may or may

not end in a straight shaft with recurved crozier. Ornament consists of non-

tuberculate, single, dichotomizing or intercalatory ribs.

Occurrence

Colchidites occurs in south-western U.S.S.R. (Turkmenia, Caucasus, Geor-

gia) (Djanélidzé 1926; Rouchadzé 1933; Eristavi 1955; Drushchits & Kudryavtsev

1960; Kotetishvili 1970; Kakabadze 1971a), France (Kilian 1888a, 1888)),

Turkey (Pelin & Thieuloy 1975), Colombia (Royo y Gémez 1945; Etayo Serna

1964), Patagonia (Blasco et al. 1980; herein); Zululand (Klinger 1976; Klinger et

al. 1984). Myczynski’s (1977) record of Colchidites sp. aff. C. colchicus from

Cuba is accepted with reservation. The illustrated specimen is a poorly preserved,

very small open-coiled heteromorph with no initial helix.

Colchidites vulanensis Egojan australis Klinger, Kakabadze & Kennedy, 1984

Figs 4, 6-9; 10A—D, J; 11-25, 26A—D

Colchidites sp. A Klinger, 1976: 16, pl. 1 (figs 6-7), pl. 3 (fig. 1), text-figs Sg, 6f—g.

Colchidites sp. B Klinger, 1976: 17, pl. 3 (fig. 2), text-fig. 5h.

Colchidites aff. C. colchicus Djanélidzé: Blasco, Nullo & Ploszkiewicz, 1980: 45, pl. 1 (figs 1-8).

Colchidites vulanensis Egojan australis Klinger, Kakabadze & Kennedy, 1984: 45, figs 6A-F,

7A-F, 8A-L, 9A-N.

Type

Holotype is SAS 3304/L54 from locality 170, Mlambongwenya Creek,

northern Zululand, Makatini Formation, Upper Barremian.

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

CPBA 11784, 11786, 11789, 11791, 11797-11803, 11805-6, 11809-11,

11814-18, 11821-23, 11829-30, 11839-47, 11849, 11851, 11856-61, 11863,

11865—-67, 11874, 11876, 11878, 11884—5, 11888, 11901-2, all from Loma Pelada,

Tucu Tucu (Bald Mountain of Hatcher 1903: 139) and CPBA 11828, 11838,

11848, 11852, 11894—6, all from Chorrillo del Medio, Tucu Tucu, Province of

Santa Cruz, Patagonia, Rio Mayer Formation, Upper Barremian; SAS L54/36h,

L54/3288, BMNH C80013, 80015—20, 80022-—25, SAM—PCZ6388—92, and more

than 50 uncatalogued specimens, including blocks of colchiditid lumachelle, all

from locality 170, Mlambongwenya Creek, northern Zululand, Makatini Forma-

tion, Upper Barremian.

Description

Dimorphic colchiditid with an initial helix consisting of at least five whorls,

surrounded by one or two or more planispiral whorls. No uncoiled section is

present.

The whorls of the helix are tightly coiled and impressed; they are either

dextral or sinistral. Of 57 specimens, 30 are dextral and 27 sinistral. The most

complete specimen (Fig. 6E) shows five whorls in the helix, thus the actual

number may be as high as six or seven. The apical angle of the helix varies

between 42 and 64 degrees. The axis of coiling of the helix is oblique to the plane

of coiling of the planispiral section, forming an acute angle of between 31 and

42 degrees. The apex of the helix rests on the umbilical edge of the planispiral

section. The whorl section in the helix is slightly depressed, subquadrate with

rounded edges.

Ornament on the helix consists of sharp, narrow, sinusoidal ribs, which

coalesce in twos or threes at the umbilical edge. Density varies between 24 and

32 per whorl.

Coiling in the first half of the planispiral whorl is irregular, with lateral

deviations, but then becomes regular. In most specimens the planispiral whorl

embraces the helix tightly, leaving no openings. The first planispiral whorl is

higher than wide (Wb:Wh = 0,97-0,83), but the whorl section varies consider-

ably from subrectangular to subcircular in different specimens. The ornament

also shows a wide range of variation. In some specimens the ribs are high and

sharp, slightly sinusoidal, usually simple, separated by wide interspaces, with rib

density between 20 and 22. In others, the ornament consists of dense, rounded,

sinusoidal ribs, which are simple or bifurcate from midflank. Bifurcations are

asymmetrical on either side of the flanks, and are not arranged in a regular

pattern. Rib density varies between 32 and 36. However, between these extremes

there is a complete series of intermediate forms, so that separation on these

grounds alone is impossible.

Some specimens are adult after one planispiral whorl at a diameter of about

50 mm. There are distinct apertural modifications consisting of strong, widely

spaced prominent ribs. These are here interpreted as microconchs. Other

327

UPPER BARREMIAN HETEROCERATINAE

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UPPER BARREMIAN HETEROCERATINAE 331

specimens have more than two planispiral, still septate whorls. It is estimated that

fully grown specimens must have had at least two and a half to three planispiral

whorls at a diameter of 300 mm. No signs of uncoiling or uncoiled fragments are

known.

Ornament on the planispiral section is very variable, but generally tends to

weaken on the outer whorls, becoming nearly smooth in some specimens.

Dimensions

Specimen D Wi Wi W,/Wh U

Holotype 44,2 18,0 (0,40) 17,0 (0,38) 1,06 15,5 (0,35)

L54/36h 91,0 38,0 (0,42) 31,0 (0,34) | 22 31,0 (0,34)

C80022 37,0 16,5 (0,44) | 15,0 (0,40) 1,10 1230; (0,32)

C80016 SIL 21,0 (0,41) 21,0 (0,41) 1,00 19,0 (0,37)

C80018 54,0 2510) (O40) 9 Ailes (O40) AG 20,0 (0,37)

C80015 50,0 23,0 (0,46) 23,0 (0,46) 1,00 18,0 (0,36)

C80017 Son0) 22,0 (0,41) 21,0 (0,40) FOS) 8,07(0534))

PCZ6389 IAW 46,0 (0,41) 34,0 (0,31) 1.339 —_— —

CPBA 11848 3 0 150 (ay ss (O.32)) 1,20 W220 (32)

CPBA 11828 S510) 16,5 (0,46) 16,0 (0,44) 1,03 20 (0.23)

CPBA 11821 5655 AES (O40) =) 13555102377) LOW ID20S(0F33))

CPBA 11817 41,0 17,0 (0,41) 16,5 (0,40) 1,03 12,0 (0,29)

CPBA 11809 45,5 IS.5 (O41) 16,0 (O35) LS — —

CPBA 11815 45,0 18,0 (0,40) 16,0 (0,36) 1,13 14,0 (0,31)

CPBA 11851 47,5 IDS (OA) 16.0 (O37) il 2a 15,0 (0,32)

CPBA 11839 40,0 15,0 (0,38) 15,0 (0,38) 1,00 IDE Sm(O731))

CPBA 11800 39,0 16,5 (O40 (Osis) 1,10 130 (533)

CPBA 11791 32), ZA) (O39) — W9)0) (Oso) Pel 22,0 (0,41)

CPBA 11830 54,0 ZOFS(O338) i) 20507 (0537) 1,03 21,0 (0,39)

CPBA 11867 48,5 20,0 (0,41) 20,0 (0,41) 1,00 14,5 (0,30)

CPBA 11894 60,0 D0) (O62) AASV (Oss7)) 1,14 21,0 (0,35)

CPBA 11797 160,0 65,0 (0,41) 48,0 (0,30) 11.3)5 50,0 (0,31)

Comparison

Most of our material compares favourably with Egojan’s (1965) original

figured specimens, especially as far as the large size of the macroconchs is

concerned. The Zululand material was separated from the Caucasian material at

subspecific level by Klinger et al. (1984) on account of the broader and flatter

venter of the planispiral section, but on the basis of the present material we doubt

if even this is necessary.

Egojan (1965: 120) compared C. vulanensis to C. djanelidzei Rouchadzé but

the two species differ on account of the disparate number of whorls in the helical

section; C. djanelidzei only has 2-3 as compared to 6-7 in C. vulanensis australis.

Occurrence

Colchidites vulanensis s.1. occurs in the Upper Barremian of the Caucasus,

Zululand and Patagonia.

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. A-D. Colchidites vulanensis australis. Coarsely ribbed microconch. CPBA 11821.

E-G. Heteroceras (H.) elegans Rouchadzé, 1933. CPBA 11900. H-I. Heteroceras (H.) elegans

Rouchadzé, 1933. CPBA 11899. J. Colchidites vulanensis australis Klinger, Kakabadze &

Kennedy, 1984. Immature macroconch? CPBA 11784. All x 1.

UPPER BARREMIAN HETEROCERATINAE 333

Fig. 11. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984. A-—C. Holotype.

SAS 3044/L54. D-G. Immature macroconch? CPBA 11838. All x 1.

334

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984. Macroconch. SAS L54/36h. X 1.

335

UPPER BARREMIAN HETEROCERATINAE

TX “68E9ZOd-WVS ‘Youosoine! “pR6l “Apouusy 2 ozpeqeyey ‘s198uTPy syv.usnv SISUBUDINA SappIyZOD “¢{ B14

ANNALS OF THE SOUTH AFRICAN MUSEUM

336

< 0:65.

Kakabadze & Kennedy, 1984. Macroconch.

PCZ6388.

is Klinger,

SAM

ls austra

Colchidites vulanens

. 14

Fig

Sei)

UPPER BARREMIAN HETEROCERATINAE

Macroconch.

565

Kakabadze & Kennedy, 1984

x 0

)

inger

SAM-PCZ6388

Fig. 15. Colchidites vulanensis australis

ANNALS OF THE SOUTH AFRICAN MUSEUM

338

565

Kakabadze & Kennedy, 1984. Macroconch.

x 0

is Klinger

—PCZ6388

Colchidites vulanensis austral

SAM

Fig.

S39

UPPER BARREMIAN HETEROCERATINAE

1984. Macroconch.

rile

Kakabadze & Kennedy,

inger

lanensis australis K\

CPBA 11896

ites vu

Rigel a Colehid

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. Colchidites vulanensis australis Klinger,

Kakabadze & Kennedy, 1984. Macroconch.

CPBA 11896, x 1

341

UPPER BARREMIAN HETEROCERATINAE

arly

Kakabadze & Kennedy, 1984. Macroconch.

is Kl

CPBA 11797

inger

is austral

Fig. 19. Colchidites vulanens

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 20. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984. Macroconch.

CRBAVI/STE x 1k

UPPER BARREMIAN HETEROCERATINAE 343

Fig. 21. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984. CPBA 11799.

xan

DISCUSSION

Validity of genera and species groups

Description of the Patagonian and Zululand material raises a number of

questions that cast doubt on the current systematic concepts employed within the

Heteroceratinae, and these merit discussion.

The first concerns the tripartite division of Colchidites into species groups

C. intermedius, C. colchicus and C. shaoriensis. This was originally conceived by

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

Rouchadzé (1933) and retained by Kakabadze (1971a). Egojan (1965), however,

had already cast doubts on the correct allocation of C. vulanensis. The well-

developed planispiral part places it in the group of C. shaoriensis, whereas the

well-developed helix places it more suitably in C. colchicus. These same doubts

were raised by Klinger (1976) and Klinger et al. (1984) and two alternatives were

proposed: either an uncoiled section was never developed; or, the Zululand

assemblage was an ontogenetically segregated population, with the representa-

tives of the discoidal stage inhabiting the shallow, near-shore water, and the

uncoiled ancyloceratid forms inhabiting the deeper parts of the basin. The

discovery of microconchs with complete apertures, however, clearly rules out the

Fig. 22. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984. CPBA 11799.

mile:

UPPER BARREMIAN HETEROCERATINAE 345

Fig. 23. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984. Macroconch.

CPBA 11829. x 1.

presence of an uncoiled section in C. vulanensis australis. Thus, as far as the

number of whorls in the helix and planispiral sections are concerned, C. vu-

lanensis australis fits none of the three recognized species groups. Instead, it more

closely resembles representatives of Paraimerites or Imerites. With the exception

of Paraimerites brevihelicoides Kakabadze, a monotypical species, none of the

ANNALS OF THE SOUTH AFRICAN MUSEUM

346

‘WOTDNS [IOUM Jeuosiijqns YUM YOUOOOIeY, “p86

‘TX “06¢9Z0d-WVS

‘Apouuoy 2 ozpeqeyey

‘JOBUI[Y] SYVAISND siIsuaUBjNA sapzip1yoj0D

‘y7 “SI

347

UPPER BARREMIAN HETEROCERATINAE

“UONSIS JIOYM [BUOSINQNS YIM YOUCdOIDR]

v86l

‘TX “06€9ZOd-WVS

‘Apouusy 2 ozpeqeyey

JOSUITY syosnv sisuaunjna sayipiyajo)

Sé

sty

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 26. A-C. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984.

SAS L54/3288. D. Colchidites vulanensis australis Klinger, Kakabadze & Kennedy, 1984.

CPBA 11805. E-G. Paraimerites haughtoni Klinger, Kakabadze & Kennedy, 1984.

SAS L54/D5. All x 1.

UPPER BARREMIAN HETEROCERATINAE 349

other species referable to either Paraimerites or Imerites has an uncoiled section.

Affinities of C. vulanensis australis with Paraimerites or even Imerites become

more obvious in specimens with distinct quadrate to rectangular whorl section.

Here the ribs are slightly flared near the venter, and the ventrolateral edges of

these may easily be interpreted as tubercles. Also, there is a slight thickening of

the ribs on the ventrolateral flanks that coincides with a slight forward flexure

and/or bifurcation of the ribs. This could be interpreted as incipient Imerites

ornament.

Having established that Colchidites vulanensis australis does not fit the

traditionally accepted tripartite division of genus Colchidites, but instead shows

more affinities toward Paraimerites and Imerites, division at generic level may be

scrutinized.

Differentiation within the subfamily Heteroceratinae at generic or sub-

generic level is based mainly on the presence or absence of tubercles, as outlined

above (p. 320). The presence or absence of tubercles has been used extensively

in the classification of other heteromorph taxa. In most cases this is an objective,

unambiguous criterion and easy to apply, and forms the mainstay of hetero-

morph taxonomy. Recent work on various heteromorph groups, especially the

Hamites—Protanisoceras—Anisoceras plexus (Casey 1961; Wiedmann 1962;

Wiedmann & Dieni 1968; Scholz 1979), the Turrilitinae (Klinger & Kennedy

1978; Scholz 1979), Baculitidae (Cooper & Kennedy 1977; Scholz 1979), etc., has

made it painfully obvious how divided workers are in assessing the importance of

tuberculation. Thus the tuberculate forms of the baculitid genus Lechites are

accorded subgeneric rank by Cooper & Kennedy (1977) while the same are

merely regarded as subspecific variants of the non-tuberculate species by Scholz

(1979). Indeed, Scholz (1979) has shown that in some heteromorph groups

(Lechites), tubercles may appear iteratively, and that the proportion or absence

of tuberculate to non-tuberculate forms may differ in different geographic

regions. In addition, transitions occur between tuberculate and non-tuberculate

forms, e.g. Hamites praegibbosus Spath and Protanisoceras gr. ex blancheti (cf.

Casey 1961: 94) where allocation to one genus or another becomes subjective

and a matter of opinion.

Bearing these data in mind, we provisionally accept the criterion of presence

or absence of tubercles in the systematics of the subfamily Heteroceratinae. Thus

those colchiditid specimens without distinct tubercles are referred to genus

Colchidites, and those with a pair of siphonal rows, to Paraimerites, even though

the coiling strategies do not fit the diagnoses as applied to the material from

south-western U.S.S.R.

These same criteria are applied to differentiating between Heteroceras s.s.

and H. (Argvethites). According to Kakabadze (1975: 66) a siphonal furrow on

the shaft occurs mainly in tuberculate forms, i.e., H. (Argvethites). However, a

siphonal furrow occurs in two of the non-tuberculate specimens of H. (GEL)

elegans from Argentina. In addition, siphonal furrows are known from other

unrelated heteromorph groups, e.g. Toxoceratoides (Aguirre Urreta 1986),

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hamites (Stomohamites) virgulatus (see Wiedmann & Dieni 1968, pl. 5 (fig. 2),

text-figs 21-22) and thus seem to be of no or little taxonomic significance.

Sexual dimorphism

Dimorphism has not been discussed previously in Heteroceratinae. In other

groups of ammonites it has been well documented over the past two decades (see

e.g. Makowski 1963; Westermann 1969; Kennedy & Cobban 1976; Callomon

1981; Donovan et al. 1981; Wright 1981 for discussions), but evidence has come

mainly from Jurassic forms—the notable Cretaceous exception being some

scaphitids (see Cobban 1969). Callomon (1981) mentioned that sexual dimorph-

ism in the Cretaceous remains largely to be explored, especially amongst the -

Ancyloceratina. Unfortunately, apart from the above-mentioned scaphitids, data

here are still very scant. Dimorphism has been demonstrated in the baculitid

genus Sciponoceras (see Kennedy & Juignet 1983: 17 for summary) and Lechites

(Cooper & Kennedy 1977) and suggested in Hamites by Cooper (unpublished

data, see Kennedy & Juignet 1983: 12).

Dimorphism in Heteroceratinae was only found in the austral representa-

tives of Colchidites vulanensis, but we suspect that it may also be present in

Heteroceras. The two species groups recognized within Heteroceras, H. gr. ex.

helicoceroides and H. gr. ex. astieri may possibly be dimorphic pairs.

Intraspecific variation

Finally, the Patagonian and Zululand material illustrates the extent of

intraspecific variation magnificently. Admittedly, coarsely and finely ornamented

forms can be identified, as can forms with quadrate whorl sections approaching

Paraimerites ornament; but these are all connected by transitions, so that it

would be futile to separate these at any formal level. This variation also casts

doubt on the validity of the great number of species referred to the genera

Heteroceras and Colchidites, as listed below. Egojan (1965) had already

commented on the disparate number of species to specimens in the Georgian

collections but, without direct access to this material, we have to refrain from

definite comments on the synonymies.

SUMMARY

The ammonite subfamily Heteroceratinae is represented in Patagonia by

Colchidites vulanensis australis Klinger, Kakabadze & Kennedy and Heteroceras

(H.) elegans Rouchadzé, both species also known from Zululand and the

Caucasus. The representatives of Colchidites vulanensis australis demonstrate

dimorphism in the genus: microconchs mature at diameters of about 50 mm and

one planispiral whorl; and macroconchs with more than two planispiral whorls

still septate at diameters exceeding 150 mm. The microconchs show apertural

modifications in the planispiral section, clearly ruling out the possibility of an

uncoiled section. The ratio of helical to planispiral coils, plus the absence of an

uncoiled section in C. vulanensis australis fits none of the three species groups

UPPER BARREMIAN HETEROCERATINAE 351

traditionally accepted within the genus Colchidites, but instead shows greater

affinity with Paraimerites or Imerites. Comparison with other heteromorph

groups shows that the presence or absence of tubercles may be of very little

taxonomic significance, and in cases, be ambiguous. Siphonal depressions occur

in various unrelated heteromorph groups, and seem to be of no taxonomic

significance. The wide range of intraspecific variation as seen in the Patagonian

and Zululand material casts severe doubt on the validity of the multitude of

species erected for the material from the south-western U.S.S.R., and suggests

that systematics can be greatly simplified.

LIST OF SPECIES OF HETEROCERAS (HETEROCERAS) AND

COLCHIDITES

Species described in open nomenclature are omitted. Most of the references

listed below provide descriptions and/or figures of the species concerned.

Genus and subgenus Heteroceras d’Orbigny, 1850

(= Lindigia Karsten, 1858)

Type-species. Turrilites emericianus d’Orbigny, 1842, by subsequent desig-

nation Meek (1876: 477).

Heteroceras emericianum (d’Orbigny) (1842: 580, pl. 141 (figs 3-6); 1851: 220,

pl. 3 (fig. 1); Kakabadze 1971la: 48, pl. 3 (fig. 2)). Upper Barremian of

France.

Heteroceras emericianum (d’Orbigny) var. costata Rouchadzé (1933: 242, pl. 15

(fig. 3); Kakabadze 1971a: 49, pl. 3 (fig. 1)). Upper Barremian of western

Georgia and northern Caucasus.

Heteroceras astierianum d’Orbigny (1851: 219, pl. 4 (fig. 1); Kilian 1888a: 430,

pl. 3 (fig. 2); 18885: 687, pl. 21 (fig. la—b); Dimitrova 1967: 65, pl. 37

(fig. 2); Kakabadze 1975: 86, pl. 1 (fig. la—b), pl. 2 (fig. 1), pl. 8 (fig. 2)).

[= Heteroceras imericum Rouchadzé (1933: 230, pl. 13 (figs 1-2),

text-fig. 34).] Upper Barremian of Bulgaria, Caucasus, France and western

Georgia.

Heteroceras bifurcatum d’Orbigny (1851: 221, pl. 3 (figs 2-3); Kilian 1888a: 432,

pies (tiges)bNikoloy. 19642 124 plh 4° (figs 1-2)- pls (fig. 5a—b);

Dimitrova 1967: 64, pl. 39 (fig. 2)). Upper Barremian of Bulgaria and

France.

Heteroceras bifurcatum d’Orbigny var. trifurcata Kilian (1888a: 432). Upper

Barremian of France.

Heteroceras devii Rouchadzé (1933: 231, pl. 13 (fig. 3), text-fig. 35; Kakabadze

1975: 87, pl. 1 (fig. 2), pl. 2 (fig. 3), pl. 8 (fig. 1)). Upper Barremian of

western Georgia.

Heteroceras elegans Rouchadzé (1933: 232, pl. 13 (fig. 4), text-fig. 36; Kaka-

badze 1975: 89, pl. 3 (figs 1a—b, 3-5), pl. 8 (fig. 5); Klinger er al. 1984: 44,

figs 2A-F, 3A—G, 4A-C). Upper Barremian of western Georgia, northern

Caucasus, Zululand and Patagonia.

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

Heteroceras eristavii Kakabadze (1975: 90, pl. 4 (figs 1a—b, 3), pl. 5 (fig. 5), pl. 8

(fig. 7)). Upper Barremian of western Georgia.

Heteroceras haugi Karakasch (1907: 144, pl. 26 (fig. 2)). Upper Barremian of

Crimea.

Heteroceras helicoceroides (Karsten) (1858: 103, pl. 1 (fig. Sa—c); 1886: 27, pl. 1

(fig. 5a—c); Royo y Gdmez 1945: 466, text-fig. 3). Upper Barremian of

Colombia.

Heteroceras isocostata Kakabadze (1975: 91, pl. 4 (fig. 5a—b)). Upper Barremian

of western Georgia.

Heteroceras jeletzkyi Murphy (1975: 36, pl. 7 (fig. 4)). Upper Barremian of

California.

Heteroceras kotetishviliae Kakabadze (1975: 88, pl. 1 (fig. 4)). Upper Barremian

of western Georgia.

Heteroceras tardieui Kilian (1888a: 433, pl. 4). Upper Barremian of France.

Heteroceras tskaltuboensis Kakabadze (1975: 87, pl. 2 (fig. 2), pl. 8 (fig. 3)).

Upper Barremian of western Georgia.

Heteroceras vermiforme Rouchadzé (1933: 233, pl. 13 (fig. 5); Kakabadze 1975:

90, pl. 3 (figs 6-8), pl. 4 (fig. 2a—b). Upper Barremian of western Georgia.

Genus Colchidites Djanélidzé, 1926

(= Heteroceras (Santandericeras) Royo y Gémez, 1945)

Type-species. Colchidites colchicus Djanélidzé, 1926.

Colchidites colchicus Djanélidzé (1926: 256, pl. 1 (fig. 1); Kakabadze 1971a: 54,

pl. 6 (figs 1-2)). Upper Barremian of western Georgia.

Colchidites colchicus Djanélidzé var. phasiensis Rouchadzé (1938: 169, pl. 5

(figs 5-6), text-fig. 21; Kakabadze 1971la: 56, pl. 8 (fig. 1)). Upper Barre-

mian of western Georgia.

Colchidites colchicus Djanélidzé var. renngarteni Rouchadzé (1938: 169, pl. 5

(fig. 4), text-fig. 20; Kakabadze 1971a: 55, pl. 7 (fig. 1), text-fig. 23). Upper

Barremian of western Georgia.

Colchidites apolinarii (Royo y G6mez) (1945: 468, pl. 74 (fig. la—b), pl. 75

(fig. 1), text-figs 4-5; Etayo Serna 1968: 59, pl. 1 (figs 4, 6), pl. 2 (figs 1-2),

text-figs 3B, 4,4; Kakabadze 1971a: 59, pl. 9 (fig. 2a—c)). Upper Barremian

of Colombia.

Colchidites atsharensis Rouchadzé (1933: 246, pl. 17 (fig. 2), text-fig. 16;

Kakabadze 1971a: 76, pl. 9 (fig. 4), pl. 18 (fig. 2a—b), text-fig. 46). Upper

Barremian of western Georgia and western Turkmenia.

Colchidites belaiaensis Kakabadze (1971a: 66, pl. 15 (fig. 2a—c)). Upper Barre-

mian of northern Caucasus.

Colchidites bethleviensis Kakabadze (1971la: 56, pl. 5 (fig. 4a—c), text-

figs 24-25). Upper Barremian of western Georgia.

Colchidites colleti Rouchadzé (1933: 249, pl. 18 (fig. 2), text-fig. 48; Kakabadze

1971a: 69, pl. 14 (fig. 1), text-fig. 39). Upper Barremian of western Georgia.

UPPER BARREMIAN HETEROCERATINAE 353

Colchidites colleti Rouchadzé var. shaoriensis Rouchadzé (1938: 170, pl. 6

(fig. 3)). Upper Barremian of western Georgia.

Colchidites cuneicostatus Kakabadze (1971a: 57, pl. 8 (fig. 2a—c), text-figs 26-27).

Upper Barremian of western Georgia.

Colchidites djanelidzei Rouchadzé (1933: 247, pl. 17 (fig. 1), text-fig. 47;

Kakabadze 1971la: 77, pl. 18 (fig. la—b), text-fig. 47). Upper Barremian of

western Georgia.

Colchidites elissoae Kakabadze (1971a: 60, pl. 5 (fig. 3), text-figs 28-29). Upper

Barremian of western Georgia.

Colchidites ellipticus Rouchadzé (1933: 252, pl. 20 (fig. 1); Drushchits &

Kudryavtsev 1960: 296, pl. 40 (fig. Sa—b); Kakabadze 197la: 64, pl. 11

(fig. 2a—b), pl. 12 (fig. la—b), text-fig. 35). Upper Barremian of western

Georgia and northern Caucasus.

Colchidites ellipticus kvadaurensis Rouchadzé (1933: 253, pl. 20 (figs 33-34);

Kakabadze 197la: 65, pl. 5 (fig. 2), text-fig. 36). Upper Barremian of

western Georgia. |

Colchidites eristavii Kotetishvili (1970: 80, pl. 12 (fig. 3a—c); Kakabadze 1971a:

59, pl. 9 (fig. 2a—c)). Upper Barremian of western Georgia.

Colchidites gamkrelidzei Rouchadzé (1933: 251, pl. 19 (fig. 2), text-fig. 50;

Kakabadze 1971a: 63, pl. 10 (fig. 2a—b), text-figs 33-34). Upper Barremian

of western Georgia.

Colchidites intermedius Djanélidzé (1926, fig. 14; Kakabadze 1971a: 51, pl. 4

(fig. 3)). (Non Colchidites intermedius in Rouchadzé 1933: 239, pl. 14

(fig. 6), text-fig. 40.) Upper Barremian of western Georgia.

Colchidites kakabadzei Kotetishvili (1970: 81, pl. 13 (fig. 3); Kakabadze 1971a:

75, pl. 17 (fig. la—c)). Upper Barremian of western Georgia.

Colchidites kakhadzei Rouchadzé (1938: 170, pl. 6 (fig. 4), text-fig. 23; Kaka-

badze 197la: 61, pl. 10 (fig. la—b), text-fig. 30). Upper Barremian of

western Georgia and south-eastern France.

Colchidites kutatissiensis Kakabadze (1971a: 53, pl. 5 (fig. 1), text-figs 21-22).

Upper Barremian of western Georgia.

Colchidites latecostatus Rouchadzé (1933: 246, pl. 16 (fig. 3), text-fig. 45);

Kotetishvili 1970: 83, pl. 14 (fig. la—b); Kakabadze 1971la: 75, pl. 16

(fig. la—b)). Upper Barremian of western Georgia.

Colchidites leenhardti (Kilian) (18885: 688, pl. 20 (fig. 3), pl. 21 (fig. 2);

Kakabadze 1971la: 52, pl. 3. (fig. 3)). Upper Barremian of south-eastern

France and western Georgia.

Colchidites longicostatus Kakabadze (1971la: 71, pl. 19 (fig. 6), text-fig. 42).

Upper Barremian of western Georgia.

Colchidites longus Rouchadzé (1933: 240, pl. 14 (fig. 7), pl. 15 (fig. 1),

text-fig. 41; Kakabadze 197la: 50, pl. 4 (fig. 4)). Upper Barremian of

western Georgia.

Colchidites multicostatus Kakabadze (1971a: 67, pl. 12 (fig. 3a—c), text-fig. 38).

Upper Barremian of western Georgia.

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colchidites ratshensis Rouchadzé (1933: 254, pl. 20 (fig. 6); 1938: 170, pl. 4

(fig. 6); Kakabadze 1971a: 72, pl. 12 (fig. 5a—b), pl. 14 (figs 2, 4-5), text-

figs 43-44). [= Colchidites nicortsmindensis Rouchadzé (1933: 255, pl. 20

(fig. 7); Tovbina 1963: 110, pl. 3 (fig. 5)).] Upper Barremian of western

Georgia and Turkmenia.

Colchidites ratshensis Rouchadzé var. kopetdaghensis Tovbina (1963: 111, pl. 3

(fic. 6); Kakabadze 197la: 73, pl.l2 (ig: 2)). Upper Barenianmon

Turkmenia.

Colchidites rionensis (Simonovich, Batsevich & Sorokin) (1875: 173, pl. 6

(fig. la—b); Rouchadzé 1933: 243, pl. 15 (figs 4-5), text-fig. 43; Kakabadze

1971la: 74, pl. 13 (fig. 1), text-fig. 45). Upper Barremian of western

Georgia.

Colchidites rotundus Rouchadzé (1933: 241, pl. 15 (fig. 2), text-fig. 42; Drush-

chits & Kudryavtsev 1960: 296, pl. 37 (fig. 2a—b); Kakabadze 1971a: 49,

pl. 3 (fig. 4), text-fig. 19). Upper Barremian of western Georgia and

northern Caucasus.

Colchidites rouchadzei Eristavi (1955: 121; Kakabadze 1971a: 51, pl. 4 (fig. 1)).

Upper Barremian of western Georgia.

Colchidites sarasini Rouchadzé (1933: 250, pl. 18 (fig. 3), pl. 19 (fig. 1), text-

fig. 49; Kakabadze 197la: 62, pl. 11 (fig. la—c), text-figs 31-32). Upper

Barremian of western Georgia and Lower Aptian? of north-western

Caucasus.

Colchidites securiformis (Simonovich, Batsevich & Sorokin) (1875: 166, pl. 4

(fig. 3a—b); Rouchadzé 1938: 168, pl. 4 (fig. 5); Kotetishvili 1970: 84, pl. 10

(fig. 4); Kakabadze 1971a: 81, pl. 172g. 4), ple 19 (ies) text -tean os) p

Upper Barremian of western Georgia.

Colchidites shaoriensis Djanélidzé (1926, pl. 1 (fig. 2); Kakabadze 1971la: 79, —

pl. 14 (fig. 3), pl. 19 (fig. 3), text-fig. 50). Upper Barremian of western

Georgia and western Turkmenia.

Colchidites tenuicostatus Kakabadze (1971a: 82, pl. 17 (fig. 2), pl. 19 (fig. 4),

text-fig. 54). Upper Barremian of western Georgia.

Colchidites tinae Eristavi (1955: 121, pl. 4 (fig. 11); Kakabadze 1971a: 52, pl. 4

(fig. 2)). Upper Barremian of western Georgia.

Colchidites tovbinae Kakabadze (1971a: 80, pl. 17 (fig. 3), text-fig. 51). Upper

Barremian of western Georgia.

Colchidites trifurcatus Kakabadze (197la: 66, pl. 12 (fig. 4a—b), text-fig. 37).

Upper Barremian of northern Caucasus.

Colchidites tzotnei Rouchadzé (1933: 254, pl. 20 (fig. 5), text-fig. 51; Kakabadze

1971a: 68, pl. 13 (fig. 2a—c)). (Non Colchidites tzétnei in Dimitrova 1967:

65, pl. 26 (fig. 2)). Upper Barremian of western Georgia.

Colchidites veleurensis Kakabadze (1971a: 83, pl. 19 (fig. 5), text-fig. 55). Upper

Barremian of western Georgia.

Colchidites vulanensis Egojan (1965: 119, pl. 1 (figs la—b, 3), pl. 2 (figs 1-2);

Kakabadze 1971la: 70, pl. 15 (fig. 1), text-fig. 41). Upper Barremian of

north-western Caucasus.

UPPER BARREMIAN HETEROCERATINAE 355

Colchidites vulanensis Egojan australis Klinger, Kakabadze & Kennedy (1984: 45,

figs 6A-F, 7A-F, 8A-L, 9A-N). Upper Barremian of Zululand and

Patagonia.

ACKNOWLEDGEMENTS

The Consejo Nacional de Investigaciones Cientificas y Técnicas, Argentina,

supported this study through a research scholarship to Aguirre Urreta at the

South African Museum. The South African Museum helped to make this

publication possible. The Servicio Geoldgico Nacional of Argentina provided the

means for fieldwork in Patagonia. Aguirre Urreta is particularly indebted to

these institutions.

Special thanks are due to J. V. Ploszkiewicz, E. Perea and V. Ramos

(Servicio Geologico Nacional, Argentina) for their valuable help in the fieldwork

carried out in Patagonia by one of us (MBAU).

Assistance with the photography by S. Dove (South African Museum) is

gratefully acknowledged. J. Blaeske (South African Museum) kindly assisted

with some of the illustrations.

REFERENCES

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358 ANNALS OF THE SOUTH AFRICAN MUSEUM

WIEDMANN, J. 1962. Unterkreide-Ammoniten von Mallorca. 1. Lieferung: Lytoceratina,

Aptychi. Abh. math.-naturw. Kl. Akad. Wiss. Mainz 1962 (1): 1-148.

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Ammonoidea. Systematics Assoc. Spec. Vol. 18: 157-174.

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Family Nuculanidae 4

Nuculana (Lembulus) bicuspidata (Gould, 1845)

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MARIA BEATRIZ AGUIRRE URRETA

HERBERT CHRISTIAN KLINGER

UPPER BARREMIAN HETEROCERATINAE

(CEPHALOPODA, AMMONOIDEA)

FROM PATAGONIA AND ZULULAND,

WITH COMMENTS ON THE

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Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

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THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: ScHULTZE, L. Zoologische und anthro-

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——EEE

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EVIDENCE FOR THE LOW-ALTITUDE ORIGIN

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DERIVED FROM THE SYSTEMATIC REVISION

OF THE GENUS COLOPHON GRAY

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S. ENDRODY-YOUNGA

Cape Town Kaapstad

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EVIDENCE FOR THE LOW-ALTITUDE ORIGIN OF THE

CAPE MOUNTAIN BIOME DERIVED FROM THE SYSTEMATIC

REVISION OF THE GENUS COLOPHON GRAY

(COLEOPTERA, LUCANIDAE)

S. ENDRODY- YOUNGA

Transvaal Museum, Pretoria

(With 31 figures)

[MS accepted 31 March 1986]

ABSTRACT

The Cape high-mountain stag beetles have been studied during the past five years. They are

recognized as representatives of a biome restricted today to the high-altitude ranges of the

mountains. This biome is often referred to as a mountain relict. An evaluation of the kinship

relations of the Colophon species presented a distribution pattern where closely related species

are isolated on mountain ranges separated by wide stretches of low-altitude plainland. These

plains are now uninhabitable for Colophon species. It was concluded that the present high-

mountain biome evolved on the low-lying plainland during a period when the environmental

conditions there were similar to those of the high altitudes today. Thus the high-mountain biome

is not an autochtonous relict, but is in refuge in an area to which it has retreated after its area of

origin became uninhabitable due to aridification and temperature increase.

The systematic revision and cladistic analysis of the genus Colophon includes the

descriptions of two new species, C. barnardi and C. moniisatris.

The discussions are introduced by sections on speciation, environment-—organism relation-

ships and interspecific competition.

CONTENTS

PAGE

HME RO CNC ULOM err aer trie site LR an de. ce ibecrrnn tions sib Wilseat Seeces. 4 360

Past cool temperate and humid climate in the Cape, and lowland

onsimothe hieh-mountaim biome asso. sagas see see] oe 361

Climate and vegetation of the past lowland habitats ............... 363

hhemaspectolabsolute time, 224.24 0ccl. es Ava ee WHS de sede yt ees 365

The geographic extent of past humid and cool temperate climate .... 365

Obsetvationsjand backeround!studics .25--...-.-.028:+s- snes 2: 366

Speciationes 4545 4.005 NESS ERO LOT OA ESRRER soc ok e ER 366

Cladisigmethodomtracingekinship) (eee 4q5- ses. 4s s esses 366

The organism—environment relationship..................... 367

Competitions. ween tae tees Ren es. Fes Seaudie ds & 368

When is a biome a relict, and when is it in refuge? ............. 370

Obscrnvationsomlive specimens 2.624. .405 546s see oe sae Sil

@ladisticanalysisiof the’\Colophon species.....5.---4.55+---52+-2-: 374

The present-day distribution of Colophon species compared with

(OSI? HONNTLOVEN S46 oles cyanide Sates NG Gee ae pee ae S/T)

Systematic revisionotthelsenus Colophon .......:...5-+.+-+-+:-- 380

PBTCMMIStORVOMSUUGIES Perry ee an cae eae ec ene oe et oe 380

The position of genus Colophon in Lucanidae ................ 381

359)

Ann. S. Afr. Mus. 96 (9), 1988: 359-424, 31 figs.

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Material’ éxamitied (24+ 320225573 ee CO ee 382

Key toiColophonimalles).. 2: Seneccs oa hee eee 383

Descriptions of the’specics, os. 2132 as ee oe ee eee 385

ihe: plesiomorphierouplolkispecicsssse see see eee 385

he aponrerphigroupiokspecies Baaoeee ae Cee eee 396

Acknowledgements). .0¢ )490:.552..8. Vea poe eee ee 422

References ss. occu hie ta dy Be Ree ee eee 423

INTRODUCTION

The Coleoptera Department of the Transvaal Museum, Pretoria, has

adopted historic biogeography as a central theme of research. During the past ten

years geographic areas that are of significance in the biohistory of the

subcontinent have been selected for fieldwork.

While tracing the origin of the Namib Desert biome, the west-coast area was

investigated from the Namib Desert to the winter-rainfall area in the south.

Simultaneously the faunal contacts of isolated indigenous forest areas in the

Drakensberg and southern Cape mountain ranges were studied. All observations

so far lead towards the conclusion that the roots of our primordial fauna are

southern in their origin. The Gondwana origin and post-Gondwana radiation of

some of our oldest faunal elements from south to north have been discussed

previously (Endrédy-Younga 1978). An historic contact between the Cape and

south-western biogeographic zones (Endrédy- Younga 1978) also became evident,

but was this by derivation or by simultaneous evolution under distinct zonation of

environmental conditions? It was clear that the next logical step was to learn more

about the oldest stratum of the Cape biome, the Gondwana relicts.

As a starting point, Barnard’s (1929, 1932a, 1932b) studies on the lucanid

genus Colophon were selected. Barnard and his fellow members of the Mountain

Club discovered a number of species of this genus living exclusively in high-

altitude mountain habitats of the southern Cape. He compared this distribution

pattern to that of the isopod crustacean genus Phreatoicus (Barnard 1929). In

doing so, he was the first to base on evidence from the invertebrate fauna the

suggestion that all the high-mountain organisms—and not only the

vegetation—are members of a distinct biome of common origin, and that this

habitat has historic significance.

Barnard’s hypothesis (1929: 180) attributed geological ages to the inhabiting

of mountains, and explained the fragmentation of populations by fragmentation

of the mountains. While a time scale thus visualized is exceedingly long, it would

in any case solve only a few of the problems. Nevertheless, the relict character of

the high-mountain biome is still widely accepted today.

Intensive collections of invertebrates have been made at and around known

habitats of Colophon species, in a search for phylogenetic and geographic

connections between the faunal elements. This fieldwork and its evaluation are

still in progress.

REVISION OF THE GENUS COLOPHON GRAY 361

Fig. 1. Colophon moniisatris sp. nov. Male in its natural environment in the Swartberg Range.

It was not my intention to revise the ‘marker’ genus Colophon until one of

the species collected in 1978 proved to be undescribed (Fig. 1). As the revision

progressed, the problems relating to the faunal history grew alarmingly, but the

solution presented itself when the results of the cladistic analysis of the species

were compared with their present-day distributions.

The derivation of the conclusions is lengthy and the time for reading is

limited for everyone. The paper is therefore presented in the reductive style.

Thus I start with my statements in medias res, with ample references to the

succeeding sections, where the reasoning can be followed and checked.

PAST COOL TEMPERATE AND HUMID CLIMATE IN THE CAPE,

AND LOWLAND ORIGIN OF THE HIGH-MOUNTAIN BIOME

Many components, plants and animals, live today exclusively in the alpine

regions of the Cape mountains. As all organisms are persistent in environmental

conditions to which they have adapted during their speciation (see p. 367), the

present high-mountain conditions must be largely the same as they were when the

component taxa evolved. There are two possible reasons for this situation: either

the biota live where they have evolved (autochtonous) and the conditions in the

area have not changed markedly, or the biota evolved somewhere else where the

conditions were similar to those where they live now and the biota changed

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

geographic position due to a shift of climatic zonation. In the first case the whole

biome represents a relict, and in the second it is in refuge (see p. 370).

The conclusion that the high-mountain biome has been translocated from its

area of origin, which was the country of low elevation around and between the

mountains, has been reached by the following sequence of hypotheses and

observations.

(a) If the biome is autochtonous, a clinal type of character transformation would

be expected along the Swartberg and Langeberg ranges. Dispersion and

subsequent speciations could occur only along the gradients of the ranges. Under

these circumstances distant isolated conspecific populations cannot occur.

(b) Conspecific populations split between mountains ranges are not known in

Colophon, although some populations are separated by deep passes (C. primosi),

or by hilly country, far below their present altitude (C. stokoei). Different species

of the monophyletic genus are, however, separated from each other by extensive

plains, of which the most striking example is C. westwoodi on Table Mountain.

Distantly isolated conspecific populations are well known among plants,

e.g. Protea pruinosa occurs on Blesberg (in the middle of the Swartberg) and in

the Cedarberg. The same can also be expected in other invertebrates, as Barnard

pointed out in the isopod genus Phreatoicus (1929: 182).

closest kinship relation might occur today in different mountain ranges. Colophon

whitei and C. montisatris sp. nov. live at the eastern distribution limits of the

genus in the heights of the Swartberg. Species from which they cannot be derived

(C. primosi and C. neli) block their west-to-east route of mountain dispersion.

Their closest kins, both in the apomorphic (C. izardi) and the plesiomorphic

condition (C. thunbergi and C. barnardi sp. nov.), inhabit sections of the

Langeberg, on the other side of the Little Karoo.

(d) The only hypothesis that resolves the problem is that the high-mountain

biome evolved around the mountain ranges, where most of the evolutionary

lineages have also been separated (in some cases down to the extant species).

Considering the persistence of species in their native climatic conditions (see

p. 367), the climate of the Little Karoo and a part of the Great Karoo (see p. 363)

is expected to have been approximately as humid and as cool in the past as the

alpine regions of the Cape mountains today. In order to attain these conditions in

the surrounding Karoo, the regional temperature would have to drop to a level

that would result in a high-altitude snow cover throughout most of the year.

Due to gradual aridification and temperature increase the low-lying country

became uninhabitable, but at the same rate the climate of the mountains became

hospitable. Populations of Colophon species and their ecological allies gradually

moved to higher altitudes in the mountains. Those species that had no access to

high mountains became extinct when their refuges became uninhabitable.

The ancestral Colophon population had ample space for dispersion,

particularly when they gained access to the Little and Great Karoo. The

REVISION OF THE GENUS COLOPHON GRAY 363

speciating populations could form a mosaic-type distribution on the open flat

country. The dispersion-cum-speciation process was not necessarily along a

gradient, as it would have been if it had occurred on a mountain range.

Phylogenetically distant species would then be able to find refuge on the same

range, e.g. from either side (see kinship of species on the Swartberg). Similarly,

the distribution of conspecific populations of closely related species could split

and shift towards different ranges (Protea pruinosa on Cedarberg and Swartberg,

or Colophon whitei and C. montisatris sp. nov. on Swartberg and C. thunbergi

and C. izardi on Langeberg).

(ce) The role of competition as a driving force for species to occupy alien biotopes,

at least in the animal kingdom, is rejected (see p. 368). Interspecific ‘competition’

might have a certain role in the food chain; thus the appearance of hitherto absent

predators might accelerate the extinction of their prey.

CLIMATE AND VEGETATION OF THE PAST LOWLAND

HABITATS

The concept of persistence of organisms in their native environmental

conditions (see p. 367) suggests that their original requirements had to be similar

to those prevailing at present in the alpine regions, i.e. considerably more humid

and considerably cooler than they are now in the southern parts of the Great

Karoo. The similarity of native conditions of different species is expressed clearly

by the altitudes at which they were taken in the different ranges. In the Swartberg

range the species occupy only the highest altitudes (approximately 2 000 m), and

are absent from any lower intervening sections. In the Langeberg, however, they

are less restricted to the heights of the peaks within their distribution areas (from

1 300 m up). Of these two ranges the northern (Swartberg) is the drier, with less

frequent cloud cover, higher summer temperatures, and a long dry spell in

summer. The relevant altitudes in both ranges (and even lower altitudes) are

regularly snow covered during two to three months in winter. The lowest zone on

Table Mountain that is occupied by a Colophon species is below 1 000 m.

Field observations (see p. 372) revealed that adults are active in any period

of the summer months but appear only under optimal weather conditions. All

specimens were sighted (in both Swartberg and Langeberg ranges) in the morning

or afternoon hours, mostly in thick fog, and on one occasion (Langeberg) in light

rain (Fig. 2). Specimens were seen in the late afternoon hours in the spreading

shade of peaks in clear weather, but not during sunny noon hours as Barnard

mentioned (1929: 178).

Their high humidity requirement is quite evident from the habitat data and

field observations. The low temperature as a factor of native conditions is more

difficult to test directly. There is, however, some circumstantial evidence to

suggest that low temperature was also part of the native climate.

If high humidity was the sole requirement, Colophon species could inhabit

the forests that are, in places, within a few kilometres of their present habitats.

364 ANNALS OF THE SOUTH AFRICAN MUSEUM

sae ea a

Fig. 2. A-B. Colophon izardi Barnard, female. The first live Colophon specimen

photographed. A. Dorsal view. B. Lateral view.

The forest habitat, however, is alien to Colophon and apparently much more so

than the low temperatures of humid alpine altitudes and the regular snow cover.

Thus to accept low temperature, Colophon required little or no adaptation,

i.e. deviation from its native condition.

The Northern Hemisphere glacial faunal fluctuation, as observed by Coope

(1979) in the British Isles, suggests a rather strong temperature adherence of the

coleopterous fauna (see p. 368).

REVISION OF THE GENUS COLOPHON GRAY 365

THE ASPECT OF ABSOLUTE TIME

The study of Colophon does not offer any direct evidence (see p. 377) of the

time when the habitat shift from the lowland to the alpine habitats occurred.

However, the Colophon species are strictly associated with the fynbos vegetation

type, and are restricted to its stratum of alpine altitudes with the implied climatic

conditions. Dated changes in the extent of the fynbos vegetation could provide

such evidence.

The first evidence (to my knowledge) of lowland fynbos far outside the

present distribution area was identified recently by Scholtz (1985), who found

Protea pollen deposits at 30°22’S 18°26'E.

The climatic change accompanied by the habitat shift to the mountain heights

seems to be a Plio—Pleistocene event coloured by several waves of lesser climatic

fluctuations (see Brain 1981). The close kinship of isolated Colophon species and

many known fragmented conspecific populations in other groups of organisms

(see p. 362) strongly support this hypothesis. Australian researchers have come to

similar conclusions with regard to their local conditions.

THE GEOGRAPHIC EXTENT OF PAST HUMID AND COOL

TEMPERATE CLIMATE

Global climatic zonation was not of the same intensity throughout the

Earth’s history; it is conceivable, however, that it always had an effect on the

climate of the subcontinent. It can thus be assumed that the cool temperate

conditions became reduced northwards in the middle of the area.

Present distribution records of Colophon demarcate the area between the

Great Karoo and the southern coastal flats as the area where the evolution of the

genus took place. The unrelated western species (C. primosi, C. neli) of the

Swartberg range, blocking the dispersion range of the two apomorphic species

(C. whitei and C. montisatris sp. nov.) (see p. 377 and Fig. 1), suggest that the

former two species might have reached their present positions from the Great

Karoo. We do not know, however, how far to the north of the Great Karoo was

originally inhabited by Colophon. Recent visits to some heights of the Witteberg

and Cedarberg ranges yielded no trace of Colophon, although these ranges have

well-developed fynbos vegetation. The ancestral stock of Colophon was

associated with the extremely cool and humid type of flatland fynbos in the

south-western corner of the Cape, and did not necessarily disperse with it into its

ecologically marginal northern types. Nevertheless, it is possible that some of its

populations shifted northwards when aridification first commenced in the central

Karoo.

Past occupation of the southern coastal plains is uncertain. The ancestral

species of the genus were certainly already apterous, non-wood-boring (in the

larval stage), and likely to be associated with open (low and scattered canopy)

fynbos-type vegetation. Areas that were taken over by dense forest vegetation in

more humid periods have thus been excluded from the past distribution of

Colophon.

366 ANNALS OF THE SOUTH AFRICAN MUSEUM

OBSERVATIONS AND BACKGROUND STUDIES

In the present paper a monophyletic group of stag beetles is studied. The

relationships between the 14 recognized species were established by cladistic

analysis. The cladogram thus obtained is compared with the present distribution

of the species and conclusions are drawn as to what environmental conditions

might have facilitated the dispersion of populations to reach their present loci.

Before entering the discussion the premises of this study need to be outlined.

Some of them are widely accepted and sufficiently described; others, however,

have been neglected or even sharply disputed as valid forces of evolution.

SPECIATION

The genetic process of evolution will not be discussed, but rather the status

and fate of populations that can or cannot lead to a speciation event. Phyletic

gradualism, which proposes sympatric speciation and an accumulation of

imperceptible changes as a means of speciation, is rejected (Mayr 1942; critique

in Hennig 1966; Paterson 1978). The accepted alternative pattern is that of

punctuated equilibria (Hennig 1966; Eldredge & Gould 1972; Paterson 1978).

According to this pattern, speciation is an event and not a continuous process; the

species are real entities and not subjective fabrications of the observer when he

slices convenient sections of the phenetic continua. The species retain their

genetic identities for a long period of time relative to the short period of the

speciation event leading to them. They do not transform in themselves, but cease

to exist when the last population representing them becomes extinct in time.

Their genetic continuity is manifested in their daughter species (if formed), or

their lineages become extinct with the last surviving species. The physical

precondition of any speciation event is the accidental formation of population

fragments. Such fragments might speciate (should they survive) if their

populations are sufficiently small, the environmental conditions of their new

habitats are sufficiently different, and they remain in isolation from other

populations for a sufficiently long period of time. It is evident that these rather

independent conditions would seldom coincide. Thus the frequency of speciation

events will depend both on the genetic constitution of the species involved and on

the lability of climatic conditions, aggravated or eliminated by the other

components of the environment. The same environmental changes would affect

populations of different species (with respect to ‘isolation’ and speciation) quite

differently. This observation emphasizes that ‘isolation’ hardly depends on the

magnitude of the environmental differences, but primarily on the genetically

determined reaction of the organism (see also p. 367).

CLADISTIC METHOD OF TRACING KINSHIP

It is claimed that phenetic similarity, if evaluated correctly, reflects the

degree of kinship. The theory formulated by Hennig (1966) has an ever-growing

literature where the methodology of cladistic analysis is adequately described.

REVISION OF THE GENUS COLOPHON GRAY 367

Here only some of the basic premises of the cladistic theorem will be recalled.

The analysis is based on the occurrence of the same derived characters

(synapomorphies) in two or more species. It means that those species that show

the highest numbers of synapomorphies, i.e. the latest acquired stages of the

transformation of the phenotypic characters, have the closest kinship. This also

implies that the weighting of characters is not a part of the evaluation. Actually,

the common occurrence of a normally infraspecific character in the phenetic

spectrum of two species would indicate the closest, sister-species relationship

(Endrédy-Younga 1981).

THE ORGANISM—ENVIRONMENT RELATIONSHIP

At every single speciation event a set of adaptive characters becomes fixed

within the genetic spectrum of a newly established species. The ultimate ‘reason’

for speciation is to provide the forthcoming generations with a set of inherited

characters that enables the individuals to cope with the environmental conditions

into which they are born, without the necessity of ever repeating adaptation.

For any organism to survive it is absolutely essential to possess a basic set of

adaptations as an inherited genetic asset to cope with the ‘normal’ or average

fluctuations of its native habitat. Thus its apparently limited adaptive capacity is

left open to cope with changes within its environment. If this were not the case,

any new-born land organism, even today, would first have to adapt to conditions

on land, leaving little chance ever to enter complex environmental situations.

Terrestrial life would still be struggling for survival along the coastlines.

If speciation is a complex genetic response to a new set of environmental

conditions into which the new population fragment was forced, it is easy to

understand that the speciated population will be conservative in its further habitat

selection; in other words it will stick to its native habitat.

Thus no population will enter a habitat alien to its native type, a habitat for

which it is not equipped in its adaptation complex. A population might, however,

be stranded in such a habitat by accident; thus it is much rather a ‘shipwreck’ than

a ‘colonizer’ population. For example, parts of populations of a forest-dwelling

species will not endeavour to enter and colonize the surrounding savannah

habitats, but may be ‘shipwrecked’ on them if the forest, which the species was

inhabiting, first became fragmented, then gradually disappeared due to climatic

changes. The classical island ‘colonizers’ (e.g. of Hawaii) are in this sense no less

‘shipwrecks’. They all reach the islands accidentally and have to encounter the

new situation with their various capacities for adaptation.

It is evident that the balance between the magnitude of difference under the

native and the new environmental conditions on the one hand, and the adaptive

capacity of the population involved on the other, will determine whether the

‘shipwreck’ population will be able to adapt and survive, or will become extinct.

The magnitude of adaptive capacity is a specific character, expressed in habitat

utilization with specialists and generalists as the two extremes.

368 ANNALS OF THE SOUTH AFRICAN MUSEUM

The ever-changing environment is a permanent stress on the organic world

that might be countered in two different ways. Either the tolerance to such

changes increases infinitely in the organism, resulting in a few infinitely flexible

forms, or parts of the organic world specialize to one or other type of condition,

resulting in increasing diversity. Apparently an infinite flexibility could not be

attained and the mechanism to adapt to particular conditions has developed. The

species with its genetically fixed adaptation complex is a foothold (and apparently

the only one) to a further change that the environment might provoke.

The inherited adaptation asset is the tool of survival for the individual in its

native environment, and the fixation of new adaptations through speciation is the

mechanism of the biota to survive the stress of the ever-changing environment.

The key statement to the present theory is specific habitat conservatism (see

above). Clear evidence for this statement was provided by Coope (1979), who

studied the coleopterous fauna of peat accumulations deposited through a

number of successive glacial and interglacial periods. It was found that practically

identical assemblages of species disappeared and reappeared with the fluctuations

of the glacial periods. At present, in the middle of an interglacial, most of the

species known from the glacial peat deposits in England are known to be extant in

their cool-climate refuges as far away as in central Asia. There is no evidence of

phenetic transformation according to the phyletic model of speciation, and it is

reasonable to expect that the same species will reappear again if glaciation were

to recur.

It can be concluded that in speciation a new, adjusted habitat preference will

be attained along with other simultaneously fixed characters. The species will

stick to its acquired habitat, as its whole set of adaptations refers to that particular

habitat and not to any other. As a result the species will readily extend its

distribution area if its habitat is expanding spatially (due to climatic change), but

its distribution will again contract if the climatic trend is reversed.

COMPETITION

Interspecific competition as a driving force of evolution in the animal

kingdom is emphatically rejected.

The term competition was introduced in biology for situations where

individuals exert pressure on one another in order to obtain more (enough) of

their environmental necessities. These necessities range from food and breeding

ground to the most general term of ‘Lebensraum’. It is claimed that by this action

entire populations might be driven into other, less advantageous habitats. It is

also often implied that such enforced habitat changes, leading to speciation,

represent an important factor of evolution. Without trying to comment on rare

and isolated cases (such as sessile benthonic animals) of habitat change for this

reason and possible speciation resulting from it, I believe that the course of

evolution rests on the balance of the organism—environment relation, as discussed

before. In other words, evolution does not ‘benefit’ from competition, and least

of all from that within biota of the same trophic level.

REVISION OF THE GENUS COLOPHON GRAY 369

What does interspecific competition imply? It is evident that those

individuals whose demands in the ‘Lebensraum’ are the most similar would be in

the fiercest competition against each other. This is undoubtedly the case between

conspecific individuals, which share characters including that of identical habitat

requirement. Thus it is only logical that competition should act most strongly

within the species rather than between species.

Undoubtedly, there is interspecific competition in the plant kingdom, e.g. if

a dense-canopy tree grows in the midst of a carpet of sun-loving plants, it will

successfully compete with them because neither can change position in order to

move out of the disadvantageous sphere of influence of the other. The outcome

will be that some of the badly affected individuals would die in the shaded area.

But assume that in time the entire deep-soil, well-irrigated flat area is taken over

by the forest; would the herbaceous plants of this habitat gradually move onto the

shallow soil of the over-drained hillside?

It is certain that one of the advantages achieved in the animal kingdom by the

‘invention’ of locomotion was that animals capable of it can ‘step aside’ if another

individual enters their sphere of interest.

The concept of interspecific competition in zoology was certainly borrowed

from botany. All sorts of distribution patterns, unclarified situations of balance in

ecosystems, and behavioural differences in cohabiting species were simply

explained as results of competition. To illustrate the above, the Namib Desert

biome may be cited as an example.

It is found that congeneric species have slightly different habitat preferences

on the dunes; it could be claimed that interspecific competition keeps them apart.

I believe that the native microhabitat of the different species is reflected in their

present habitat selection. The aliopatric speciation areas of many species involved

is proven (Endrédy- Younga 1982). The same applies to the high-mountain biome

under discussion.

In the desert biome several tenebrionid species might live together in the

same hummock or on the same dune; some of them are diurnal, others nocturnal.

It has been claimed that the activity pattern of the different species has shifted

apart due to competition. The time pattern is, however, a generic or more often a

tribal character (Eurychorini and Caenocrypticini are nocturnal, Zophosini and

Adesmiini diurnal, in their sometimes continent-wide distribution area). Thus the

activity pattern is a much older genetic character of the evolutionary lineages than

the co-habitation of a few of their extant representatives that might be thought to

compete.

It is claimed in this paper that competition is not a driving force of evolution

or dispersion of biota. Unusual pressure on an ecosystem by the importation of

new species as a result of various causes, ranging from human importation to conti-

nental collision, might be expected. This might drastically alter the composition

of the fauna (e.g. Marsupialia in South America), but mainly in the direction of

extinctions. We have little evidence that rapid speciation has ever started in an

indigenous fauna owing to the arrival of successfully competing biota.

370 ANNALS OF THE SOUTH AFRICAN MUSEUM

Competition in the food chain could be another possibility. It can be claimed

that the prey species move out into an alien habitat to avoid the pressure of

predator(s). I have not heard, however, of antelopes vacating the savannah and

moving into the forest due to the activity of lions. The competition between

predator species for a limited quantity of prey could still be a possibility. In this

case, however, conspecific individuals have identical habitat and prey selection,

but members of other predator species do not. It would be more likely that the

excess population of the predator that upset the balance would be eliminated than

that another prey species would be badly affected. Thus neither the difference in

habits nor habitat selection between predator species is due to competition.

Cheetahs have similar habits and habitats throughout their distribution areas and

these are different from those of lions, whether or not they occur together.

Probably the best support for this is provided by examples of biological control;

pests and predators live together in an oscillating equilibrium without either

becoming extinct. Even human influence can tip the balance only if new predators

or parasites are introduced, but not by the propagation of the indigenous ones.

WHEN IS A BIOME A RELICT, AND WHEN IS IT IN REFUGE?

The term relict is used equally for biota and for habitats if they are isolated

remnants of earlier periods of larger distributions and habitat continua. Thus it

has a clear implication in time. Does the term relict, however, mean

unequivocally that biota were left behind in situ where they evolved and that the

relict habitat of today formed a part of the earlier continuum? In other words, has

the term relict a similarly clear spatial implication?

It has been seen (p. 367) that species persist in environmental conditions that

were associated with their speciation; in other words, species are conservative in

their habitat selection. In an ever-changing environment the biota are forced to

follow the spatial shifting of climatic zonation. Parts of populations might be left

behind in situ where conditions are persistent: e.g. gallery forests remaining from

a forest belt during an arid period. Such a gallery forest is, in time and space, a

relict habitat and its biota might consist largely of relict populations, i.e. relicts in

the sense of time and space.

Climate zones might, however, together with their persistent biota, sweep

through areas that were previously under the regime of different climatic zones.

Under particular conditions such a moving biome might be stranded in a

geographic position distant from its original position, where it no longer exists.

Such a habitat and its biota are relicts in the temporal sense, but not in the spatial

sense. They were left behind from an earlier epoch, but not in situ. In other

words, these biota are not autochtonous.

The term for such translocated relict habitats is refuge habitat or refugium,

and applies equally to the biota characterizing them. There is terminological

difficulty in describing in one word the biota or species in a refuge habitat, and

this might have been the reason why they are simply called ‘relicts’ without

distinction.

REVISION OF THE GENUS COLOPHON GRAY a7

It will be shown in the following section that the distinction between relict

and refuge habitats and their biome is crucial to the understanding of the history

of an area where relicts, in the broad sense, occur. It will also be shown that the

biota of the Cape high mountains are in refuge, and do not represent a relict in

the spatial sense.

OBSERVATIONS ON LIVE SPECIMENS

Very little is known of the biology of the Colophon species, and observations

are restricted to a few live specimens in nature (Fig. 3) and in captivity.

In October 1977 several high-mountain areas were visited, both in the

Swartberg and Langeberg ranges. Although on that occasion no live specimens

were seen, it was possible to mark two sites for further studies. The indications of

the prospective sites (one in each range) were fragments of dismembered

specimens of Colophon beetles found in debris and under stones, Large numbers

of ground-traps with preservative were set in both areas but they yielded no

specimens of Colophon, although later several specimens were sighted within a

few metres of operating traps. It was concluded that none of the baits used,

including fermenting banana, had any attraction for these species. During three

subsequent years the two selected sites were visited five times. On three occasions

live specimens were seen and observed; each time some of the specimens were

taken live to Pretoria, where they were observed for periods of up to 84 days. Our

scanty knowledge comes from these field and laboratory observations of live

specimens.

Fig. 3. Colophon izardi Barnard, male, in its natural environment in the

Langeberg Range.

372 ANNALS OF THE SOUTH AFRICAN MUSEUM

The native habitat in terms of habitat diversity is markedly different for the

two species observed. The habitat of the Swartberg species, Colophon moniisatris

sp. nov., was demarcated in a tiny area of about 400 m horizontally and 20 m

vertically. Outside this area not even fragments of specimens could be found,

despite the fact that no obvious change in any of the environmental conditions

could be seen. However, the Langeberg species, C. izardi, was found at sites

8 km apart and differing in altitude by about 400 m. The same species is known

from five additional localities at increasing distances from the observation area.

Although a detailed evaluation of environmental differences between the altitude

extremes is not available, the differences in other parameters, such as soils,

vegetation, exposure, etc., seemed to be considerable between the high open

crest and the broad valley between two crests.

The striking difference between the Colophon habitats on the two mountain

ranges is obviously due to their general climatic situation. The Swartberg

population appears to occupy the last remnant of a habitat still acceptable to it,

and it is likely that it is on the verge of natural extinction unless the regional

climate soon turns more humid and cooler, thus extending spatially its native

habitat. This is a classical ‘bottle-neck’ situation, providing selective pressure for

a trapped population to speciate. Indeed, the climate of the Swartberg is drier and

warmer compared to the Langeberg; rain and mist are less regular in the summer

months. The same altitudes at which Colophon lives in the Langeberg (down to

1 400-1 600 m) are very dry after December in the Swartberg, where the peak of

activity of beetles in general has already passed. In the Langeberg the mist zone is

lower, and although dry spells might occur there too, the climate is more humid

throughout the summer. Colophon specimens were seen live in both localities in

February too, but with the described difference in their altitude range.

I am of the opinion that Colophon species in general have a much less-

restricted seasonality of adult activity than Barnard (1929: 178) suggested.

Barnard recorded, for example, C. izardi in October and November, and I have

seen them in December and February; thus the adult activity period covers the

whole summer at that altitude. The habitats might often be covered in snow until

August or September, persisting from March or April. The surface activity of

adults is likely to be triggered by optimal conditions at any time during the

summer and is not restricted to any particular period of it. The pattern of

scattered days of surface activity of an individual is likely to last through several

years. This opinion might be supported by specimens, particularly females, that

have the anterior tibiae strongly eroded. This degree of wear also occurs in

Tenebrionidae, but only in species that could be kept in captivity for many years.

According to observations Colophon species are decidedly diurnal in

activity. On all occasions when live specimens were sighted ‘observation shifts’

were organized, in order to scan the area with powerful lamps throughout the

night. However, not a single Colophon was sighted, although hundreds of other

Coleoptera were collected. Observation on live captive specimens confirms the

diurnal activity.

REVISION OF THE GENUS COLOPHON GRAY 373

The optimal weather conditions for surface activity can also be defined. Five

specimens of Colophon monitisatris sp. nov. were sighted within an hour, in thick

fog after sunset (Swartberg, February 1979). One specimen of C. izardi was seen

(Langeberg, December 1977—the very first live specimen) at about 16h00 in

drizzling rain and biting cold wind. All other specimens were seen in late

afternoon hours, often after sunset but before dark. Barnard (1929: 178)

mentions 06h00-—08h00 as the collecting hours of most of his specimens, and

records as exceptional two specimens that were collected ‘at midday in hot sun,

though amongst low vegetation’. Nevertheless, it is clear that these beetles prefer

the cool and humid periods of summer days, when their refuge habitat is not

under snow. Again this is an indication of the climatic conditions of their lowland

native habitat.

Fig. 4. Colophon montisatris sp. nov., male. This specimen’s activity was

observed for over an hour.

Circumstantial evidence suggests that Colophon species have a subterranean

life, where their larvae feed on the roots of plants. On two occasions males of two

species (C. izardi and C. montisatris sp. nov.) were observed undisturbed for

over an hour. I had hoped that they would lead me to a female (that of

C. montisatris sp. nov. was not yet known at that time), or would chew on

something, dig in, or withdraw to their own burrow when it became dark; they did

absolutely nothing. Their slow movement seemed to be entirely undirected and

aimless and revealed nothing at all of their habits (Fig. 4). Specimens in captivity

did not accept any food provided for them; this included a range of plants from

their habitat, obtained from the Botanical Gardens, Pretoria. None of the captive

specimens were observed to feed (seven specimens, over periods ranging from

15 to 84 days) unless on the vegetation debris that was richly supplied in the soils

brought with them.

374 ANNALS OF THE SOUTH AFRICAN MUSEUM

CLADISTIC ANALYSIS OF THE COLOPHON SPECIES

As will be explained (p. 381), the higher classification of this group can only

be clarified in conjunction with a comprehensive revision of the whole family. At

this stage we have to be satisfied with the recognition of the genus as a

monophyletic group. The general structure displayed by all 14 species suggests

that this group is not only monophyletic but forms one single genus.

The 14 species form two distinct groups indicated by the symmetry of the

aedeagus and the single correlated character found, i.e. the level of the clypeal

margin (see p. 385). However, separate genera could not be allocated to these

species groups, as one of them—comprising the plesiomorph species—

apparently does not possess any apomorphic character states among the

numerous characters studied.

The two species groups are interpreted as two distinct evolutionary lineages

within the genus. The first group, comprising the plesiomorph species, is a result

C B

Fig. 5. Lineage characters in the genus Colophon. A. Symmetrical aedeagus in the plesiomor-

phic lineage, as in C. haughtoni Barnard. B. Asymmetrical aedeagus in the apomorphic lineage,

as in C. primosi Barnard. C. Apical portion of penis with the non-rectractable inner sack as in

C. primosi Barnard. D. Asymmetrical anal sternite in C. montisatris sp. nov., typical for the

males of the apomorphic lineage.

REVISION OF THE GENUS COLOPHON GRAY 375

of the first cleavage and possesses as apomorphies only those characters that

ensure the generic position of Colophon itself. The generic apomorphies can only

be recognized through out-group comparison, but at this stage we do not know

what should be considered as an out-group for Colophon. However, the existence

of true generic-level apomorphies (applicable to all species included) is con-

fidently postulated. The phenetic distinctness of the Cape stag beetles proves

their integrity at least at generic level. The problem is indeed to find a relative to

them at all.

The two evolutionary lineages represented by the two species groups can be

characterized as follows. Plesiomorphic lineage— Aedeagus of symmetrical type,

parameres very nearly identical or symmetrical; anal sternite evenly rounded

posteriorly. Penis feebly sclerotized with distinct transverse undulation on the

shaft (Fig. 5A). Clypeus meets labrum on an even level. Apomorphic

lineage—Aedeagus of asymmetrical type, right paramere strongly dilated

towards apex and forms a sharp hook at its inner margin, left paramere slender

and without inner hook (Fig. 5B). Anal sternite correspondingly asymmetrical at

its posterior margin, emarginate to the right of its median axis so as to provide

space for the enlarged right paramere at ejection (Fig. 5D). Penis well sclerotized

with very distinct and asymmetrical apical portion (Fig. SC). Anterior margin of

clypeus raised above the level of labrum at their suture.

The symmetrical structure of the penis is a Key and irrefutable character to

denote the plesiomorphic position in the lineage of species that share this

character.

The above group characters are not used in the following analysis, which is

based on the transformation series given below. The analysis of each lineage is

carried out separately. Often the same characters are used, and are sometimes

represented in both transformation series in the manner: absence to presence

(plesiomorph group), presence to distortion (apomorph group).

Transformation series in the plesiomorphic lineage

1. Apico-ventral process of anterior tibia small to large.

2. A longitudinal crest at the ventral side of anterior tibia more or less uniform

and moderate over whole length to sharply raised in its basal section.

3. Apico-ventral process of anterior tibia in central position to shifted to inner

margin.

4. Dorsal process of mandible absent to present.

5. Ventral process of mandible concealed to free standing and prominent.

6. Lateral margin of pronotum evenly curved along whole length to deeply

emarginate before reaching posterior lateral angle.

7. Anterior margin of pronotum immarginate at its median section to con-

tinuously marginate.

Gula flat along whole length to sharply concave (lateral view).

9. Prosternal apophysis simple to forming a sharp tubercle. at its caudal

(intercoxal) end.

376 ANNALS OF THE SOUTH AFRICAN MUSEUM

stokoei

eastmani

haughtoni

cameroni

1 2 3 4 5 6 7 8 9

Fig. 6. Tabulated record of apomorphies in the plesiomorphic lineage of the genus Colophon.

Figures below the histograms refer to characters evaluated in the text.

cassoni

berrisfordi

primosi

westwoodi

barnardi

thunbergi

whitei

montisatris tf iy PH Su

izardi

1 2 3 4 5 6 7 8 9 10

Fig. 7. Tabulated record of apomorphies in the apomorphic lineage of the genus Colophon.

Figures below the histograms refer to the characters evaluated in the text.

Transformation series in the apomorphic lineage

1. Apico-ventral process of anterior tibia present, reducing to absent.

2. Inner angle of anterior tibia rounded to forming a prominent process.

3. Anterior tibia straight to bent inward.

4. Longitudinal ventral elevation of anterior tibia flat and rounded to sharply

crested.

5. Lateral margin of anterior tibia with several spaced teeth, through trifid to

bifid apical portion.

6. Anterior tibia evenly curved at external margin to a deep U-shaped

emargination in pre-apical position.

REVISION OF THE GENUS COLOPHON GRAY B77)

7. Dorsal process of mandible present to absent.

Ventral process of mandible in basal, to median or apical position.

9. Lateral margin of pronotum moderately to sharply and deeply emarginate

posteriorly.

10. Gena evenly rounded behind eye to a distinct postgenal protuberance.

Fig. 8. Simple female mandibular structure in Colophon

westwoodi Gray. The mandibles are very similar in

females of all Colophon species.

Where applicable, the very uniform display of character states in females was

used for out-group comparison in deciding on the polarity of the transformation

series. Females generally, but notably in Lucanidae, are more conservative in

phenetic changes and thus can be considered in most characters to be close to a

hypothetical ancestor (Figs 8, 14A). In some other characters, stages uniformly

displayed by the symmetrical-aedeagus lineage, were considered as plesiomorphic

states in the genus, e.g. the presence of an apico-ventral process of the anterior

tibia and the presence of a dorsal process of the mandible.

The synapomorphies, as displayed in the plesiomorphic and in the apo-

morphic lineages of species respectively, are shown in Figures 6 and 7. From

these data a cladogram was constructed (Fig. 9). On the branches the numbers of

synapomorphies are marked in squares and the numbers of autapomorphies in

circles, indicating the position of terminal species.

THE PRESENT-DAY DISTRIBUTION OF COLOPHON SPECIES

COMPARED WITH THEIR EVOLUTION

It is assumed that Colophon, as a monophyletic genus, had a single ancestral

species. From this ancestral population, by the repeated speciation of marginal

population fragments, an increasing number of species has evolved. In a slow-

moving, wingless and terrestrial type of organism such as Colophon, the process

of dispersion must have been slow and the chances of population fragmentation,

378 ANNALS OF THE SOUTH AFRICAN MUSEUM

! ‘i Hl IV 5 6 7 8 9 10 1 12 13 14

aga n

sid

os eg - — S&S

= i= ° no) = D p

= fo) ca —- c qe - {o) mo) = o 2

S p oO i) S n n {S) iS o — 1) ma)

a fo) nay 2 (7) RQ, - i= » (S = ~ »

oO = WY fo) n c - — %) ‘= S Ste i o

CG c oO wn oO a = Qa = a p = = ie

QO OH DOOOOGE EQ) OS

\ \ : \ BE

[e) EB \ \

Fig. 9. Cladogram of Colophon species. Figures in squares show the numbers of synapomor-

phies found; figures in circles show the numbers of autapomorphies found in the individual

species.

affecting spatially advanced marginal populations, rather good. If the character

transformations in species of such a group of apterous organisms are studied

along a geographical gradient, a clinal pattern will usually be found. This is only

to be expected as the phenetic reflection of a chain of subsequently forming

daughter species. Similar observations have also been made on apterous

tenebrionids (Endrédy-Younga 1982).

The assumption that Colophon and its entire high-mountain biome is an in

situ relict of past climatic epochs should be dramatically proved by the phenetic

cline of Colophon species along the mountain ranges where they live. It would be

expected that consecutive east-west sections of the Swartberg and Langeberg

ranges should be occupied by species of closest kinship in the sequence of their

specific character transformations.

However, the actual situation is very different from that expected. The

plesiomorph group of species occupies the mountain ranges in the south-western

corner of the area (Fig. 10). Most of the apomorphic species occupy the east—

west ranging Swartberg and Langeberg mountains, but one (C. westwoodi) lives

ai)

REVISION OF THE GENUS COLOPHON GRAY

‘(SOWILU IO} 6 “BI 90S) advouT] s1ydiowode sy} Jo satoeds ay) soinsy o1qeie ‘adeouTy

o1ydioworsayd ay) Jo sorsads oy) yuasoidar sandy uewoy ‘sotoods woydojoy say) Jo uoNNaqinsip Aep-juasoid pur uorssodsip feonomiodAH “Oy “314

ce Lc (0X4 6L

380 ANNALS OF THE SOUTH AFRICAN MUSEUM

on Table Mountain, where it is isolated at the western extremity of the mountain

groups occupied exclusively by the plesiomorph species.

Similarly, the three terminal apomorphic species of the apomorphic species

group are separated at the east end of the major mountain ranges: C. whitei and

C. montisatris sp. nov. on the Swartberg and C. izardi on the Langeberg, with the

arid Little Karoo between them.

Thus the dispersion and speciation of the Colophon stock could not have

occurred on and along the mountain ranges but only on the lowland situated

around the mountains. At the end of the humid and cool temperate climatic

period, when the presently inhabited altitudes of the mountains were under snow

most of the year, an intricate mosaic pattern of Colophon species inhabited the

low-altitude country. Because of the gradual changes towards a dry and hot

summer, the populations withdrew to the nearest mountains where they could

persist under their native conditions. It is likely that several species retreated into

areas that could provide only temporary shelter. Populations that became

stranded in hilly country of insufficiently high altitudes could not survive during

continuing aridification.

The phylogenetic position of Colophon primosi and C. neli raises further

problems. They belong undoubtedly to the apomorphic species group of the

genus; however, they cannot be derived from the same ancestral stock as all other

species of this group. There is a strong indication that the dispersion and

speciation of the C. whitei—C. izardi lineage proceeded eastwards between the

main mountain ranges. It is unlikely, however, that the ancestry of C. primosi

and also of C. neli followed the same route. It is conceivable that the ancestral

territories of those latter lineages occupied sections of the southern Karoo, north

of the Swartberg range. The same geographical situation can also be the reason

why only those two species have survived wheras their closest relatives, having no

access to nearby mountains for retreat, became extinct.

SYSTEMATIC REVISION OF THE GENUS COLOPHON

THE HISTORY OF STUDIES

The first Colophon specimens must have been collected around the turn of

the eighteenth and nineteenth centuries by unknown travellers. The genus, with

its first known species, C. westwoodi, was described by Gray in 1832; locality,

date or collector of the type-specimen (in the British Museum (Natural History),

London) are not known. A second, similarly old specimen in the Hope Museum,

examined by Parry in 1870, bears no information regarding its origin, but was

already labelled as a ‘type’ in Parry’s time. Gray (1832) knew, however, only of

one male.

The second species, C. thunbergi, described by Westwood (1855), is

dedicated to the notable traveller and naturalist C. P. Thunberg, who visited the

western Cape in the 1790s. There is no evidence that he actually collected the

specimen in question, but a passage in his diary (1793) coincides with the locality

REVISION OF THE GENUS COLOPHON GRAY 381

of a specimen of C. thunbergi in Oberthur’s collection (Muséum national

d’Histoire naturelle, Paris), referring to the Swellendam section of the Langeberg

range. These hints led Barnard to the discovery of a population of this

controversial species at the Zuurbrak Peak in 1932 (Barnard 1932b).

Dr K. H. Barnard revived interest in these curious beetles. As a curator, and

after Péringuey, the director of the South African Museum, Cape Town, he was

not only a Keen student of aquatic zoology but also a devoted mountaineer. It is

only to be expected that the curious high-mountain dwellers would not long evade

his interest. As far as can be ascertained, the first specimens of Colophon

collected by Barnard were those found in 1916 in the Hottentots Holland

Mountains. They proved to be different from the two known species and were

described as C. stokoei Barnard, 1929. The second record came from Dr

S. Haughton, who collected a single male of yet another unknown species in the

Matroosberge in 1917—C. haughtoni Barnard, 1929, dedicated to its collector.

Thus encouraged, Barnard’s mountaineering colleagues paid attention to these

curious beetles during their excursions, and their efforts are reflected in Barnard’s

dedications of new species collected. All nine of Barnard’s species are named

after members of the Mountain Club of South Africa. (I have completed the

eminent list by naming one of the new species after the former President of the

Club, Dr K. H. Barnard.)

In his first paper on Colophon, Barnard (1929) described five new species:

C. stokoei, C. haughtoni, C. cameroni, C. izardi and C. primosi, and made the

first character evaluation in the genus, pointing out the main specific characters of

the male mandibles, anterior tibiae, and aedeagi. Studying the aedeagi, he first

discovered that the taxonomic position of the genus under the subfamily

Lampriminae is, at least, doubtful.

A plate with photographs of all species of Colophon, including those to be

published in a forthcoming scientific paper, was published in a popular article

(Barnard 1932a). Although the names to be used accompanied the photographs,

this article cannot be considered as the place of publication (see International Code

of Zoological Nomenclature 1964: Article 12). The proposed names became avail-

able in a subsequently published scientific paper (Barnard 1932b), where descrip-

tions of C. neli, -C. whitei, C. cassoni, C. berrisfordi and C. eastmani were given.

THE POSITION OF THE GENUS COLOPHON IN LUCANIDAE

The genus Colophon was until very recently associated with Southern

Hemisphere subfamilies of the Lucanidae. McLeay (1819) recognized the

subfamilies Chiasognathinae and Lampriminae as separate subfamilies. Follow-

ing this classification, Didier & Séguy (1953) placed Colophon under Lamprimi-

nae. The same classification was maintained by Landin (1955) and Brinck (1956).

For a long period of time these two subfamilies (both Southern Hemisphere) were

combined under the name Chiasognathinae, and Colophon was considered a

member of it (Parry 1864; Handlirsch 1908; Van Roon 1910; Jeannel 1942). Thus

the actual affiliation of the genus has not changed.

382 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lacordaire (1856), who recognized Chiasognathinae and Lampriminae,

placed Colophon under the subfamily Lucaninae, or ‘lucanides vrais’. All

classifications mentioned so far were based on differently evaluated external

characters, in particular those of males that are strongly subjected to secondary

sexual character transformations.

The first author to examine the male genitalia of Colophon species was

Barnard (1929), who was by no means a specialist of Lucanidae. Nevertheless he,

referring to Sharp & Muir (1912), observed that these genitalia are distinctly

more like the Lucaninae than the Lampriminae-type, and shed the first doubt on

the association of this South African genus with the Lampriminae. His

noteworthy discovery was disregarded by several subsequent authors.

In 1960 Holloway published her study on a wide range of species of the

family Lucanidae comparing, among other characters, the structures of male

genitalia. She categorically excluded Colophon from Lampriminae, and placed

the genus under Lucaninae as Lacordaire (1856) and Barnard (1929) had done,

although on a more intuitive basis.

Being concerned with the relationship and origin of the genus Colophon,

I have repeated Holloway’s study on less extensive, but representative, material

from South America, Africa and Australia, including species classified under the

subfamilies Chiasognathinae, Lampriminae and Lucaninae. I agree that accord-

ing to Holloway’s (1960) classification Colophon does not belong to either of the

Southern Hemisphere subfamilies, but to the cosmopolitan Lucaninae. Neverthe-

less, the problems of relationship and origin remain unresolved. It became

evident that there is no evolutionary link between Colophon and the other, old-

world stock genera of African Lucanidae. Thus, although Colophon belongs to

the subfamily Lucaninae, it seems not to be derived from its African old-world

lineages, nor could the latter be derived from the stock of Colophon. Although no

conclusive studies were made in this direction, some of the Southern Hemisphere

genera of the subfamily Lucaninae (such as Pholidotus McLeay in Brazil, and

Rhyssonotus in Australia) seem to be more closely related to Colophon than any

of the African genera.

Thus the Gondwana origin of Colophon can be maintained under its

classification in Lucaninae, but the problems in the evolution of the subfamily are

compounded. Holloway (1960) envisaged the necessity of further subdivision of

the subfamily to resolve diversity problems. Such a study might also clarify the

problems surrounding origin, evolution and dispersion within the group.

MATERIAL EXAMINED

In the course of the present study nearly all Colophon specimens known to

be deposited in collections were re-examined. In all 136 complete specimens and

numerous fragments were seen, some of the latter being listed below under

primary and secondary types. The material included four primary and 38

secondary type specimens, 46 further specimens and numerous fragments from

the South African Museum, Cape Town; eight primary and 18 secondary type

REVISION OF THE GENUS COLOPHON GRAY 383

specimens and eight further specimens from the British Museum (Natural

History), London; the type specimen of C. thunbergi from Oxford; and one

primary and 10 secondary type specimens and 11 further specimens and fragments

in the collection of the Transvaal Museum, Pretoria. By kind courtesy of the

South African Museum, some of the duplicates could be deposited in the

Transvaal Museum, Pretoria.

The value of fragments, which are more frequently found than intact or live

specimens, was fortunately already realized by Barnard. Some fragments, such as

male heads and anterior tibiae, are perfectly characteristic of the species they

represent. Barnard had already made use of such fragments in his character

evaluations and listed them in his species descriptions. Indeed, in several

instances the single, or few, intact specimens alone would not permit conclusive

recognition of specific characters and individual variations. In this paper more

importance than usual is given to fragments; where appropriate they are

designated as paralectotypes and have been marked accordingly.

In listing material examined the locality data have been taken from the actual

specimens labels, hence various inconsistencies in spelling, etc. Any additional

information added is placed within square brackets.

KEY TO COLOPHON MALES

1(8) Edge of clypeus in flush with labrum. Aedeagus of symmetrical type, left

and right parameres closely similar. Penis less sclerotized with distinct

transverse undulation, apical portion symmetrical (Fig. 5A). Apical

sternite symmetrical at apical margin.

2 (3) Mentum with sharply raised anterior portion forming one or two

(bilateral) rounded elevations or a deep aciculate anterior surface;

sickle-shaped mandibles with ventral process at base only (Fig. 17A).

Anterior tibia emarginate before apico-lateral dilation (Fig. 20C).

21-27 mm. Western Cape mountains ............. C. stokoei Barnard

3 (2) Mentum flat or moderately and evenly convex.

4(5) Apex of mandible and pre-median dorsal process connected by a

flattened shiny surface (Fig. 11B). Ventral process of mandible broadly

obtuse-angled. Prosternal apophysis without any process on posterior

portion. (Anterior margin of pronotum not furcate-marginate.) Anterior

tibia slightly and evenly curved and dilated (Fig. 14D); apico-ventral

process extremely large, cylindrical and perpendicular to tibia.

20-23 mm. Western Cape mountains .......... C. haughtoni Barnard

5 (4) Dorsal process of mandibles more or less cylindrical, not forming a flat

inner surface; position more or less postmedian.

6(7) Ventral process of mandible well developed and situated in basal

position. It is flat, acute-angled, without forward-pointing cylindrical

process. Dorsal process in median position, apex symmetrically truncate

(Fig. 11A). Anterior tibia evenly but somewhat more dilated (Fig. 14B).

16—26 mm. Western Cape mountains ........... C. cameroni Barnard

384 ANNALS OF THE SOUTH AFRICAN MUSEUM

7 (6) Ventral process of mandible indistinct, situated in a basal position not

surpassing mentum. Dorsal process about median in position, narrow,

cylindrical, perpendicular to main arm (Fig. 11C). Anterior tibia dis-

tinctly arcuate at apical portion (Fig. 14E), ventral crest evenly and

sharply elevated from base, abruptly contracted postmedially, forming a

right angle (Fig. 14F). Apico-ventral process entirely in inner marginal

position, also visible in dorsal view (Fig. 14E). 17—22,5 mm. Western

Capesmountainss: 2.2 2.2.0 en an ee ee C. eastmani Barnard

8 (1) Anterior margin of clypeus sharply elevated above the level of labrum.

Aedeagus of asymmetrical type, left and right parameres strongly

dissimilar, the left one angularly dilated (Fig. 5B). Penis strongly

sclerotized, surface smooth with longitudinal lateral margination, apex

asymmetrical (Fig. 5C).

9 (10) Mandibles long, beak-like and prognathous with matching inner margin

(Fig. 21B), together with legs orange-brown in contrast to black body;

small ventral processes in basal position. Anterior tibia straight, evenly

dilated with four sharp lateral teeth (Fig. 20A). 28-35 mm. Swartberg

TATOO eo elec e So Cesc ae ae Pe ae C. primosi Barnard

10 (9) Mandibles arcuate, black.

11 (12) Anterior tibia narrow, rectangularly bent inward in apical third

(Fig. 27F). Ventral process of mandibles in apical position; apex of main

arm in closed position of mandibles forms a circle with ventral process

(Fig. 29C). Pronotum about as long as broad, normally with a pair of red

dots on disc (Fig. 31). 21,8—25,3 mm. Langeberg range. ...............

ee EE eo Or noe doe ccs duds aso 0: C. izardi Barnard

12 (11) Anterior tibia straight or bent at varying angles, but not rectangularly

bent or filiform.

13 (14) Anterior tibia flat and oblong with parallel margins, about three times

longer than broad (Fig. 27A); ventral crest sharp at basal half of length,

ending here abruptly, forming a rectangle. Apico-ventral process large.

Mandible with ventral process only (Fig. 21A). 15-19 mm. Swartberg

E10 SA aE RE Na US NC) MUR IE era ein alia at atrstaen ba C. neli Barnard

14 (13) Tibia not regularly oblong.

15 (16) Frons behind clypeal declivity with a rounded elevation. Main arm of

mandibles broadly truncate and slightly bifid at apex, with rounded

dorso-median elevation. Dorsal process reduced to a tiny basal knob.

Basally situated ventral process small, hardly surpassing labrum

(Fig. 24A). Anterior tibia moderately arcuate and dilated with four or

five often irregular lateral processes (Fig. 20B). 21-25 mm. Table

Mountaitts, 9.5) au opm le ia bel eel ai na ce C. westwoodi Gray

16 (15) Frons flat in middle or finely concave at clypeal declivity.

17 (20) Mandible without dorsal process.

REVISION OF THE GENUS COLOPHON GRAY 385

18 (19) Anterior tibia very strongly dilated from base to apical four-fifths, then

abruptly reduced into a U-shaped emargination before rising into bifid

apico-lateral process (Fig. 27B). Pronotum deeply emarginate pos-

teriorly, posterior angle rectangular. 23—27,8 mm. Swartbergrange. ....

oe a ag AO Ee STM na teeter ae ge ae Le te a C. montisatris sp. nov.

19 (18) Basal three-fifths of anterior tibia almost straight and parallel in dorsal

view, apical dilation trifid, not contracted before dilation (Fig. 20E).

Posterior third of pronotum contracted, but lateral margin here not

concave, posterior angle of pronotum obtuse-angled, ampliate lateral

margin of disc rounded. 21 mm. Langeberg range. C. barnardi sp. nov.

20 (17) Mandibles with dorsal and ventral processes.

21 (22) Anterior tibia very strongly dilated from base to deep pre-apical

emargination of lateral margin; apico-lateral process narrow and bifid

(Fig. 27D). Dorsal process of mandibles in median position forming an

oblong surface with pointed angle at its inner apical corner; the whole

process is sunk below the dorsal level of the main arm. Ventral process

with its forward-pointing apex is positioned between the apex of main

arm and dorsal process (Fig. 29A). 26-31 mm. Swartbergrange. .......

5 Sie Gn ee Un Oar HIVES Rene cc eae parr ea a eee C. whitei Barnard

22 (21) Anterior tibia not strongly dilated laterally, tibia semi-parallel, expanding

evenly to apical process. Dorsal process of mandibles is not an oblong

structure.

23 (24) Ventral process of mandibles is produced sharply forward; it is narrow

and cylindrical. According to the shape and position of the dorsal

processes this species is known in two sympatric morphs (Fig. 17B, C).

More than 18 mm. Western Cape mountains. ...... C. cassoni Barnard

24 (23) Ventral process of mandibles forms a forward-upward slanting ridge from

base, its corners are rounded obtuse-angled (Fig. 19). Dorsal process

pointed triangular. Length about 21 mm. Swartbergrange. ............

is AION AN RMB ON) thai) SUES Re i bois 3 C. berrisfordi Barnard

DESCRIPTIONS OF THE SPECIES

THE PLESIOMORPH GROUP OF SPECIES

The common characters of the species forming this group are the

symmetrical type of aedeagus (Fig. 5A) and evenly arcuate posterior margin of

anal sternite in males. A correlated character is the structure of the forehead,

where the margin of the clypeus is level with the labrum (in males). Phenetic

characters do not show the split into the two main lineages in females.

The parameres in this species group are nearly similar, symmetrically arcuate

at their apex, with the left paramere also simple (enlarged in the species of the

apomorphic group). Penis less sclerotized than in the apomorphic species, with

characteristic undulation on the shaft. Apex evenly and symmetrically rounded,

ending in the long flagellum of the ductus ejaculatorius. Minor differences were

386 ANNALS OF THE SOUTH AFRICAN MUSEUM

seen between species in minute details of the aedeagus; however, these will not be

described, as no particular importance was attributed to them in the evaluation.

External characters of males were found to be sufficiently constant and much

more evident than those of the aedeagi. In C. stokoei, the only species where

considerable variability of characters could not be evaluated conclusively

(individual or subspecific), there are not enough males known to provide the

support of genital characters.

Colophon cameroni Barnard, 1929

Figs 11A, 12, 14B

Colophon cameroni Barnard, 1929: 172, fig. 7; 1932a: 23, figs. Didier & Séguy, 1953: 76. Brinck,

1956: 309, 320. Holloway, 1960: 335.

Diagnosis

Black, medium size; mandible with large dorsal and small ventral processes.

Anterior tibia almost straight, ventral crest sharply elevated basally, flat apically

with apico-ventral process. Aedeagus of symmetrical type.

A B Cc

Fig. 11. Male mandibular structure in Colophon species. A. C. cameroni Barnard.

B. C. haughtoni Barnard. C. C. eastmani Barnard.

Description

Male. Head broad and short, slightly tapering in front of eyes; rounded

ocular canthi high, vertex flatly depressed between. Frons steeply deflected

towards clypeus, which is level with flat and broad labrum. Punctation finer and

denser near ocular canthi than at vertex. Mandibles prognathous, finely and

evenly arcuate, pointed apex not bent inward. Dorsal process large, slightly

raised above the level of main arm; slightly forward-pointing, truncate apex

slightly dilated. Ventral process short and broad at base of mandible, rounded

apex rectangular or obtuse-angled (Fig. 11A). Mentum more than twice as broad

as long, corners narrowly rounded, anterior margin arcuately truncate. Gular

sulcus shallow.

REVISION OF THE GENUS COLOPHON GRAY 387

Pronotum distinctly broader than elytra, lateral margins semi-parallel

(hardly converging forward) in median third, more convergent in anterior third,

posteriorly not emarginate. Lateral and basal margins margined, anterior margin

smooth. Punctation very fine and scattered.

Elytra smooth, densely shagreened and very finely punctate, wanting distinct

subhumeral carina.

Anal sternite only slightly asymmetrical.

Anterior tibia straight, evenly dilated from base to apex; bifid apico-lateral

process short and broad, not marked off basally by deep insinuation of lateral

margin (Fig. 14B). Ventral crest sharply elevated in basal two-thirds, gradually

flattened anteriorly; apico-ventral process small.

Aedeagus of symmetrical type. Penis feebly sclerotized, apex symmetrical

and smoothly rounded, shaft sharply and deeply wrinkled transversely. Parameres

Fig. 12. Colophon cameroni Barnard, male;

paralectotype; length 24,5 mm.

388 ANNALS OF THE SOUTH AFRICAN MUSEUM

relatively symmetrical, right paramere broader, apex more pointedly turned

inward; left paramere narrower, inward-turning apex short and rectangular.

Female. No female is known from the type locality.

Length

Male: 25 mm (range, estimated from heads, 16-26 mm).

Distribution

Western Cape, Hex River Mountains.

Material examined

Males: 3 complete specimens, 1 anterior part of body, 24 heads and

10 anterior legs. Females: 3 complete specimens, 2 bodies and 1 head. Lecto-

type ¢, paralectotype 2 (complete); 1d and 12 (complete), 4 ¢ heads

(paralectotypes) in British Museum (Natural History), London. All other

specimens in South African Museum, Cape Town.

Lectotype. 3: Waai Hoek Mts., 5 000 ft. [= 1524 m], 1.1.1925, K. H.

Barnard, in British Museum, London.

Paralectotypes. 1 complete 6, 7 6 heads and 5 ¢ anterior legs: Waai Hoek

Mts., 5 600 ft.[= 1 707 m], [April] 1928, K.H. Barnard in South African

Museum, Cape Town; 1 complete 2, Waai Hoek Mts., 5 600 ft. [= 1 707 m], in

British Museum, London. 17 ¢ heads (13 in South African Museum, 4 in British

Museum): Fonteintjieberg, Worcester distr., 6 500 ft. [= 1 982 m], [March]

1929, K. H. Barnard.

Additional material. 1 3, 1 2, Milner Ridge Peak, Hex River Mountains,

6 000 ft.[= 1 829 m], January 1934, K. H. Barnard, in British Museum, London.

1 5,2 2 (specimens found dead) and 2 6 anterior legs: Great Winterhoek top,

6 800 ft. [= 2 073 m], August 1929, K. H. Barnard, in South African Museum.

3 6 anterior legs: Witzenberg range, Tulbagh, 1932, F. Berrisford, in South

African Museum.

Variability

There is a considerable size variation in the 21 ¢ heads measured. They

range between 4 and 6 mm, measured between transverse sulcus of vertex and

apex of mandibles. No allometric variation was found. A limited individual

variation could be observed in the angle of the basal process of mandibles, and in

the apical curvature of mandibles, as well as in the length of the ventral elevation

of anterior tibia. Nevertheless, the Waai Hoek and Fonteintjiesberg specimens

are certainly conspecific.

A pair from Milner Ridge Peak, Hex River Mountains (see above) represent

the smallest specimens [d 16 mm, 2? 17 mm). Head length of male (measured as

above) is only 3,7 mm. In this male the mandibular structure is similar but the

dorsal processes are narrower and are not dilated at their apex; ventral process

flat, obtuse-angled. Elytra in both specimens with faint and short subhumeral

REVISION OF THE GENUS COLOPHON GRAY 389

elevation. Anterior tibia similar, but elevation of ventral crest less than half of

tibial length.

Colophon haughtoni Barnard, 1929

Figs 11B, 13, 14D

Colophon haughtoni Barnard, 1929: 171, fig. 6; 1932a: 19, figs. Didier & Séguy, 1953: 77. Brinck,

1956: 308, 320.

Diagnosis

Black; pronotum and elytra more shiny than in the other species. Medium

size. Pronotum immarginate anteriorly. Anal sternite moderately asymmetrical,

aedeagus of symmetrical type.

Description

Male. Head broadly oblong. Genal margin in front of eye distinctly

emarginate. Ocular canthus high but rounded, vertex evenly concave in between,

steeply deflected in middle to broad and finely emarginate clypeus, which is on

the same level as the labrum. Punctation of vertex distinct, somewhat denser

laterally. Mentum broad, semicircular. Transverse gular sulcus broad and rather

shallow. Mandibles moderately long, evenly arcuate from base to apex. Apex

appears as split, inner dorsal portion being curved backwards to form a broad

concave surface between the two apices. Dorsal process absent. Ventral process

small and obtuse-angled at base of mandibles (Fig. 11B).

Pronotum broadest at about the basal third of its length, evenly arcuate

anteriorly in one, roundly arcuate to apical angle in another specimen. Prebasal

section of lateral margins contracted but not emarginate. Posterior angle obtuse.

Anterior margin of pronotum immarginate. Disc very finely and sparsely

punctate, punctures somewhat larger laterally. Base of prosternal apophysis

without a trace of callous elevation.

Elytra short and broad, not much narrower than pronotum. A short humeral

crest at base distinct and, particularly in one of the two males, a faint longitudinal

striation discernible.

Anal sternite moderately asymmetrical.

Anterior tibia arcuate, moderately dilated towards apex with an apical and a

pre-apical triangular process, which appears in some other species as the bifid

apico-lateral process (Fig. 14D). Ventral crest sharp and elevated in basal two-

thirds, flattened pre-apically, forming a very large conical process perpendicular

to tibia.

Aedeagus of symmetrical type. Penis feebly sclerotized together with apical

portion. Parameres (right) unarmed pre-apically, apical portion of parameres

nearly symmetrical.

Female. Shape of pronotum and elytra, and sculptural features of latter,

similar to that of males.

390 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Colophon haughtoni Barnard, male;

length 23 mm.

A B Cc D E F

Fig. 14. Anteror tibiae in Colophon species. A. C. primosi Barnard, female. B. C. cameroni

Barnard, male. C. C. cassoni Barnard, male. D. C. haughtoni Barnard, male. E. C. eastmani

Barnard, male. F. C. eastmani Barnard, male in semi-ventral view.

REVISION OF THE GENUS COLOPHON GRAY 391

Length

Males 20—23 mm, female 18 mm.

Distribution

Hex River Mountains.

Material examined

Holotype. 6: Matroosberg, Hex River Mountains; 1917, S. H. Haughton; in

British Museum (Natural History), London.

Additional material. 2 3, 1 2°: Matroosberg, Hex River Mountains; January

1941, K. H. Barnard, in South African Museum, Cape Town. 2 2 (found dead):

Groot Hoek Peak, Hex River Mountains; September 1933; K. H. Barnard, in

British Museum, London.

Colophon eastmani Barnard, 1932

Figs 11C, 14E-F, 15

Colophon eastmani Barnard, 1932b: 174, fig. 6. Didier & Séguy, 1953: 76. Brinck, 1956: 309, 320.

Diagnosis

Medium-sized black species. Mandible with dorsal process in middle and

ventral process basally. Clypeus is flush with labrum, pronotum immarginate

anteriorly. Anterior tibia in male moderately arcuate with bifid apico-lateral

process. Anal sternite moderately asymmetrical, aedeagus of symmetrical type.

Description

Male. Head transversely oblong. Genal margins in front of eye emarginate,

not converging. Ocular canthi large, broad and rounded, vertex evenly concave in

between, anteriorly straight or shallowly sinuate. Vertex steeply deflected to

clypeus. Clypeus broadly truncate or finely emarginate; flush with broad and flat

triangular labrum. Disc finely and rather sparsely punctate. Mandible evenly

arcuate, sickle-shaped apex pointed, perpendicular dorsal process behind middle

with large rounded apex. Ventral process very short, pointed, obtuse-angled; in

closed position of mandibles concealed by labrum (Fig. 11C). Mentum flat, twice

as broad as long, laterally evenly arcuate. Transverse gular sulcus deep, anteriorly

vertical.

Pronotum moderately broader than elytra, broadest between middle and

posterior third of length; lateral margins almost evenly arcuate between lateral

angles, posteriorly flattened but not emarginate, posterior angle rounded obtuse-

angled. Pronotum anteriorly immarginate. Punctation hardly visible on disc,

more distinct laterally.

Anal sternite forms a small callus at margin, which is almost in middle;

hardly asymmetrical.

392 ANNALS OF THE SOUTH AFRICAN MUSEUM

ce ~ ong eS

Fig. 15. Colophon eastmani Barnard, male;

paralectotype; length 22 mm.

Anterior tibia moderately arcuate inward, slightly dilated, apico-lateral

process produced, bifid (Fig. 14E, F). Ventral crest sharp, gradually elevating to

abrupt end in front of middle; apico-ventral process small, triangular, forward-

pointing, visible in dorsal view (Fig. 14E, F).

Aedeagus of symmetrical type. Penis feebly sclerotized. Parameres not quite

symmetrical.

Female. Unknown.

Length

Male 17—22,5 mm (lectotype 22,5 mm).

Distribution

Western Cape mountains.

Material examined

6 d (dead), 2 5 heads, 10 ¢ anterior legs.

Lectotype. 3: Keeromberg (Worcester), Sept. 1930; K. H. Barnard, in

British Museum, London.

REVISION OF THE GENUS COLOPHON GRAY 393

Paralectotypes. 1 6: Keeromberg (Worcester), Sept. 1930; K. H. Barnard,

in British Museum, London. 3 6 (partially fragmented), 2 d heads, 10 6

anterior legs; Keeromberg (Worcester), Sept. 1930; K. H. Barnard, in South

African Museum, Cape Town.

Additional material. 1 ¢ (fragmented): Dassieberg, Robertson, 5 550 ft.

[= 1 692 m]; January 1936, K. H. Barnard; South African Museum.

Colophon stokoei Barnard, 1929

Figs 16, 17A, 20C—D

Colophon stokoei Barnard, 1929: 168, figs 3—5. Didier & Séguy, 1953: 77. Brinck, 1956: 308, 320.

Diagnosis

A species with symmetrical type of aedeagus and anal sternite; clypeus flush

with labrum. Mandibles without dorsal process, anterior tibiae arcuate, apically

bifid in front of lateral emargination. Mentum prominently thickened.

Remarks

A range of variations was described as ‘local variation’ by Barnard, who

marked the different types as ‘f. typ. [forma typica], var. A, B, C, and D’.

Considering the large and fragmented distribution area of C. stokoei, the marked

differences could indicate subspecifically distinct populations. The locality

records, however, seem to contradict such a situation. One of Barnard’s reasons

for not attaching specific or subspecific value to any of these variations could have

been the inconclusive distribution patterns drawn from the available records.

Some of the locality data, often provided by laymen, are considered as doubtful

(e.g. the Blesberg record of C. whitei, see p. 417), and no further records have

become known since Barnard’s (1929) description. Though I doubt that some of

the considerable differences are due only to individual variations, I refrain from

drawing any conclusions due to the inadequacy of the material. The differences

will be briefly discussed and the varieties are denoted below by the same letters as

used by Barnard.

Description

Male. Head transversely oblong, anterior lateral angle almost pointed, but

more obtuse-angled than in C. westwoodi. Internal subocular crests rounded but

distinct, with tubercle at the anterior inner end of each crest. Frons evenly

depressed between tubercles and arcuately slanting to clypeus. Barnard’s

‘anterior margin of head’ refers to the angle of declivity of frons towards clypeus

and not to a difference in levels between clypeus and labrum. The shape of frons

and its form of declivity show a high degree of variation. It is sharp-edged with an

overhanging clypeus in C. stokoei var. B. Clypeus meets triangular labrum on the

same level.

Mandibles comparatively short and simple, each nearly semicircular, dorsal

surface of apical portion furcate. Dorsal process lacking. Ventral process

394 ANNALS OF THE SOUTH AFRICAN MUSEUM

prominent, rectangular (C. stokoei var. C) or forward-pointing acute-angled

(Fig. 17A).

Mentum very thick with a high degree of variation (simple in all other

species). The actual form of mentum is the basis of Barnard’s division of the

species into varieties. In all varieties mentum evenly elevating from base in lateral

view, and forming a vertical line or surface anteriorly. Anterior margin

transverse, sometimes more or less tuberculate at either side (C. stokoei forma

typica and C. stokoei var. B), anterior surface broad and large, deeply excavated

(C. stokoei var. A), or simply roundly pointed (C. stokoei var. C and D).

Pronotum evenly arcuate at lateral margins, hardly and only very slightly

emarginate near posterior angle. In general appearance moderately enlarged.

Elytra short and broad with distinct variation in proportions. Smooth in most

varieties but in the ‘holotype’ of C. stokoei var. C, elytron with two distinct

lateral elevations and a rounded inner longitudinal elevation (costae).

Anterior tibia more or less arcuate and dilated towards apex. Apex laterally

bifid (C. stokoei forma typica, var. A, B and D) with deep U-shaped emargina-

tion pre-apically (Fig. 20C) (C. stokoei forma typica, var. A and B, less deep in

C. stokoei var. D); or apico-lateral armature tridentate with indistinct pre-apical

emargination (C. stokoei var. C) (Fig. 20D). Ventral longitudinal crest distinct,

but not sharply raised basally, almost continuous to apex in C. stokoei var. C.

Apico-ventral process long and narrow in all varieties.

Aedeagus of symmetrical type and similar in all varieties where known

(C. stokoei forma typica, var. A and C) and all seven dissected specimens. As

specific differences in aedeagi can be found between all recognized species of the

genus, the similarity of aedeagi in C. stokei varieties might indicate their

infraspecific status. Dorsal line of left paramere in lateral view emarginate in

apical third, and forms an angle where it turns into the finely convex basal two-

thirds; in C. haughtoni apical portion almost straight, angle flat and indistinct.

Right paramere less pointed than in C. haughtoni.

Female. Apparently identical between varieties where known (i.e. from

localities of C. stokoei forma typica, var. B, C and D).

Length

Male 24-27 mm, female 18-22 mm (Barnard (1929)—20-—22 mm in males,

measured without mandibles).

Distribution

Western Cape mountains from Hottentots Holland to the Stellenbosch and

Wellington mountains.

Material examined

10 d (two dismembered and seven dissected), 4 2. Of these 2 ¢ and2 @ are

in British Museum (Natural History), London; all other specimens in South

African Museum, Cape Town.

REVISION OF THE GENUS COLOPHON GRAY 395

Type material. The designation of a primary type should actually only be

applied to the ‘forma typica’ described by Barnard as C. stokoei; specimens listed

under his lettered varieties should not be designated as paralectotypes of the same

species. However, I intend to designate primary types for the ‘varieties’ as well,

so as to avoid increasing the existing confusion, not only in the evaluation of the

variability of the species but probably also in the given locality records. The label

data are quoted exactly from the specimen labels and additional information,

partly from Barnard (1929), is given in brackets.

C. stokoei forma typica

Lectotype. 1 6: Hott.-Holl. [Hottentots Holland] Mts., 4 000 ft.

[4 000-5 000] ft. [= 1 220-1 524 m], Caledon C.C. [Cape Colony], Barnard,

[January] 1916; in British Museum, London.

Fig. 16. Colophon stokoei Barnard, male; paralectotype

of Barnard’s forma typica; length 24 mm.

396 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paralectotype. 1 3: data as for lectotype; in South African Museum, Cape

Town.

A third male with identical data to the above specimens, but not mentioned

in Barnard’s (1929) description, is in the South African Museum, Cape Town. It

is identical to the ‘lectotype’ of C. stokoei var. A (see below).

C. stokoei var. A.

‘Lectotype’, d6: Kleinmond Mt., C.P. [Cape Province]; 1.8.1927, T. P.

Stokoe, in the South African Museum, Cape Town.

The identity of this specimen, the second listed by Barnard (1929), is not

quite certain, though the area, date and collector agree. The first specimen listed

by Barnard as C. stokoei var. A could not be found, unless it is the one listed

above but bearing different locality. Though both specimens agree with the

description of C. stokoei var. A, they prove how unreliable the locality data are.

C. stokoei var. B.

Syntypes consisting of 1d head, 1 2 head, 4 d anterior legs (rather

dissimilar in details): Plattberg, 3 000 ft. [915 m], Palmiet Riv. Mts., K. H.

Barnard, Febr. 1927 [South African Museum]. Syntype, 1 2 (dismembered):

locality as for other syntypes; in British Museum (Natural History), London.

C. stokoei var. C.

Holotype, ¢: Drakenstein Peak [Stellenbosch District], 4 000 ft. [1 220 ml],

2.1.1920; K. H. Barnard [and T. P. Stokoe]; in South African Museum, Cape

Town.

C. stokoei var. D.

Syntypes consisting of 1 6 head and 3 6 anterior tibiae: Up. [Upper]

Snoukop, Wellington 4 500 [—5 000] ft. [= 1 372-1 524 m], K. H. Barnard [and

R. Primos], [January].

Additional specimens. 1 3 (forma typica): Sneeukop, Hottentots-Holland

Mts., Somerset West (name of collector illegible), 20.xii,1945. 1 ¢ (forma

typica): Bushmans Castle, Berg River Range, Franshoek, Dec. 1943. 1 3 (closest

to C. stokoei var. C, but more slender and straight anterior tibia): Sneeukop,

Hottentot-Holland* Mts., 4 500 ft. [1 372 m], 15.xii.1935, K. H. Barnard. 2 ¢

and 2 (morphologically between C. stokoei forma typica and C. stokoei

var. C): Berg River Range, Emerald Dome, Apr. 1944, K. H. Barnard. 1 @ (see

Barnard 1929, C. stokoei var. C): Helderberg, 5.xii.1926, K. H. Barnard.

THE APOMORPH GROUP OF SPECIES

The common characters of this species group are the strongly asymmetrical

shape of the aedeagus and the correspondingly asymmetrical posterior margin of

the anal sternite in males. A correlated character expressed in the same form in

all the species of this group is the anterior margin of the clypeus being raised

above the level of the labrum. As the asymmetry of the aedeagus and the position

of the labrum are not functionally related, these correlated apomorphic

characters indicate convincingly the monophyletic origin of this well-defined

evolutionary lineage.

REVISION OF THE GENUS COLOPHON GRAY 397

The parameres in these species are asymmetrically expanded, with the left-

hand one in particular forming a large apical or pre-apical triangular process

pointing towards the penis. Thus the two parameres form a closed sheath or

funnel around the penis in a closed position, probably in order to protect the

penis and the long flagellum at penetration at the start of copulation. (Note that

the flagellum is not retractile into the penis as it is in the Lampriminae.) The penis

is strongly sclerotized, its apical portion asymmetrical. Its integument is firmly

sclerotized with longitudinal furrows (Fig. 5B) and is not transversely undulating

as in the plesiomorphic species group (Fig. 5A). The posterior margin of the anal

sternite is emarginated above the enlarged left paramere and bears a more-or-less

(according to species) distinct, rounded projection above the right paramere

(Fig. 5D). More or less distinct differences in the structure of the parameres and

the penis were observed between species. Although these differences are likely to

be constant and specific, a detailed evaluation was not possible owing to the

scarcity of the material. While male genital characters might improve the

phylogenetic evaluation of the species, their importance in species identification 1s

limited, as male external characters are ample and appear to be specifically

constant.

Colophon cassoni Barnard, 1932

Figs 14C, 17B-C, 18

Colophon cassoni Barnard, 1932b: 172, fig. 4. Didier & Séguy, 1953: 76. Brinck, 1956: 311, 320.

Diagnosis

Small to medium-sized black species. Male mandibles with dorsal and

prominent ventral processes. Aedeagus of asymmetrical type.

Description

Male. Head broadly oblong. Genal margin in front of eye straight, parallel or

slightly converging. Anterior margin deeply insinuated and deflected to vertically

positioned clypeus. Clypeus narrow, pointed in small specimens, raised above

Fig. 17. Male mandibular structure in Colophon species. A. C. stokoei Barnard (forma

typica). B.C. cassoni Barnard (forma typica). C. C. cassoni m. recurva morpha nov.

398 ANNALS OF THE SOUTH AFRICAN MUSEUM

level of small triangular labrum. Mandibles rather short, main arms curved

inwards at varying angles. Apices roundly pointed or truncate, sometimes slightly

emarginate. Ventral process sharply produced anteriorly, joining the main arm,

slightly sinuate at its inner margin; apex reaches basal third to half of length of

mandible. Dorsal process thick, almost cylindrical, rising smoothly from surface

of main arm, pointing slightly forward (Fig. 17B). Mandibular armature is subject

to allometric variation. Mentum slightly convex, roundly triangular, coarsely

punctate. Gular sulcus flat.

Pronotum hardly broader than elytra in specimens seen (all small). Margins

semi-parallel in median third, evenly arcuate, converging anteriorly, finely

emarginate behind. Pronotum with margination only interrupted in middle of

anterior margin. Punctation extremely fine on disc, more distinct at posterior

angle.

Elytra smooth, finely punctate, sutural margins smoothly and rather

indistinctly elevated.

Anal sternite asymmetrical, without distinct off-centre protuberance of

margin (small specimen).

Anterior tibia similar to that of C. cameroni; almost straight at inner margin,

arcuately dilated towards apex. Bifid apico-lateral process more produced,

sometimes with a third rudimentary proximal process (Fig. 14C). Ventral crest

sharply elevated at basal two-thirds (some variation in length), arcuately

interrupted pre-apically; apico-ventral process rather large; in an inner-side view

the process looks as if split from body of tibia.

Aedeagus: right paramere forms a large process at its inner margin; margin

between process and apex straight, angularly emarginate between process and

base. Left paramere unarmed, wedge-shaped with roundly pointed apex.

Fig. 18. Colophon cassoni Barnard,

male; paralectotype, fragment; length

14 mm. An estimated total length of the

specimen could be about 23 mm.

REVISION OF THE GENUS COLOPHON GRAY 399

Female. Distinguished by the antericr margination (interrupted in middle) of

pronotum, emargination of posterior converging portions of pronotal margins,

smooth and rather broad fine elevation of sutural margins. Pronotal punctation

more distinct than in male.

Length

Male 18mm (range estimated from size of heads 16-24 mm); female

16-17 mm.

Distribution

Swartberg, the northern range of Cape mountains.

Variability

The mandibular armature of males shows two distinct forms, without any

intermediates. Barnard (19325: 172) referred to intermediates but these differed

only in thickness and shape of the apex of the dorsal processes (as is often seen in

the genus) and not in the position and direction of the processes. No similar

dimorphism has been seen in any other species of the genus. However, as no

correlated characters could be found and only one complete male is known of

each form, the actual value of this unusual difference cannot be evaluated. All

specimens and fragments referred to this species were collected on two occasions

(1930 and 1932) and both samples include both mandibular forms. It can thus be

assumed that the two forms occur sympatrically. For the time being the two

varieties will be considered as morphs of the same species (see below).

Colophon cassoni Barnard, 1932 (forma typica)

Fig. 17B

Description

Dorsal process of mandibles in median position, pointing diagonally forward.

With mandibles in closed position, the small space bounded laterally by the main

arms and apically by the dorsal processes is of an inverted heart-shape (Fig. 17B).

Material examined

Lectotype. 3: Meiringspoort Berg, Zwartberg Range, K. H. Barnard, Febr.

1932; in British Museum (Natural History), London.

Paralectotypes. 1 @ (identity doubtful); Meiringspoort Berg, Zwartberg

range, K. H. Barnard, Febr. 1932; in British Museum, London. 21 ¢ heads,

26 6 anterior legs; data as for lectotype; in South African Museum, Cape Town.

2 ¢ (identity doubtful), 3 d heads, Meiringspoort Berg, Zwartberg Range,

F. Berrisford, 1930; in South African Museum.

400 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colophon cassoni morpha recurva morph. nov.

Fig. 17C

Description

Dorsal process of mandibles in basal position pointing diagonally backward.

Space between the apex of mandibles and dorsal processes ovate and much larger

(Fig. 17C).

Material examined

Holotype. 3: Meiringspoort Berg, Zwartberg Range, Febr. 1932, K. H.

Barnard; in British Museum (natural History), London.

Paratypes. 2 3 heads (data as for holotype); 1 ¢ front body, 2 d heads (data

as for holotype, but coll. F. Berrisford, 1930); in South African Museum, Cape

Town. 6 head (data as for holotype) in Transvaal Museum, Pretoria.

Remarks

Barnard’s (1932b) description gave the data for specimens collected by

F. Berrisford as ‘Blaauw Punt near Meiring’s Poort, Zwartberg Range, 6 500 ft.

(F. Berrisford and P. Casson Scott-Hayward), March 1930’. As is often the case,

the data published by Barnard are not consistent with those of the labels; there is

no doubt, however, that they refer to the same specimens.

The identity of Meiringspoort Berg could not be established among the peaks

west of the poort.

Colophon berrisfordi Barnard, 1932

Fig. 19

Colophon berrisfordi Barnard, 1932b: 173, fig. 5; 1932a: 19, figs. Didier & Séguy, 1953: 76.

Brinck, 1956: 311, 320.

Remarks

The holotype (the only specimen known) is a fragment, generally in a fresh

condition, consisting of the front part of the body of a male (head with both

antennae and pronotum with both anterior legs, one complete with tarsi). The

aedeagus and anal sternite, which bear the most important species-group

characters, are wanting. Sharp-edged clypeus, elevated above labrum, refers the

species to the apomorphic group, as this character is correlated with the

asymmetrical type of aedeagus in all known species. Structure of anterior tibia is

practically identical with that of C. cassoni, and rather similar to that of

C. cameroni.

Description

Male. Head very similar to that of C. whitei, with the sharp and narrow

transverse dorsal impression level with eyes; genal canthus in front of eye less

dilated. Moderately arcuate mandibles with well-developed premedian dorsal

REVISION OF THE GENUS COLOPHON GRAY 401

process, and with small tubercle between this process and apex (more distinct on

left mandible). Ventro-basal process of mandible forms a forward- and upward-

slanting ridge with rounded corners (Fig. 19).

Pronotum sharply margined all round, marginal fossa only flattened at a

point in middle of anterior margin. Shape similar to C. whitei, surface densely

shagreened, punctation extremely fine in middle of disc, gradually more distinct

towards lateral margins.

Anterior tibia slightly arcuate, semi-parallel to apical dilation, not contracted

pre-apically. Apical dilation armed with three teeth. Ventral crest strongly

elevated from base to in front of middle, here almost forming a right angle,

arcuately flattened to the base of the small rectangular apico-ventral process.

Ventral crest and apico-ventral process are comparable to those of C. neli and

C. cassoni. Inner apical corner of tibia forms a sharp, slightly acute-angled

process, not rounded as in C. neli.

Female. Unknown.

Length

From apex of mandibles to base of pronotum 13 mm, breadth of pronotum

10,0 mm, suggesting a smaller specimen than the smallest known specimen of

C. whitei; total length probably in the range of 20—22 mm.

Distribution

Central part of Swartberg Range, west side of Meirings Poort.

Fig. 19. Colophon berrisfordi Barnard, male;

holotype, fragment; length 13 mm. An esti-

mated total length of the specimen could be

about 21 mm.

402 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B (S D E

Fig. 20. Male anterior tibiae in Colophon species. A. C. primosi Barnard. B.C. westwoodi

Gray. C. C. stokoei Barnard forma typica from Bushmans’ Castle. D.C. stokoei Barnard,

holotype of Barnard’s var. C. E. C. barnardi sp. nov.

Material examined

Holotype. 3 (anterior part of specimen): Meirings Poort Berg, Zwartberg

range, F. Berrisford, 1930; in British Museum (Natural History), London. (Name

label in Barnard’s hand.)

Colophon neli Barnard, 1932

Figs 21A, 27A

Colophon neli Barnard, 1932b: 170, fig. 2; 1932a: 19, figs. Didier & Séguy, 1953: 77. Brinck,

1956: 310, 320.

Diagnosis

Colophon neli is the smallest known species of the genus. Black. Mandible

lacking dorsal process. Anterior tibia broad, straight and parallel. Clypeus raised

above labrum. Anal sternite and aedeagus of asymmetrical type.

Description

Male. Head broadly oblong. Ocular canthus and a pair of anterior elevations

at the base of the mandibles rounded. Vertex deflected in a V-shape to clypeus,

which is narrow with rounded or emarginate apex and is raised high above narrow

triangular labrum. Surface coarsely punctate, particularly in depressions.

Mandible simple sickle-shaped, with only prominently pointed ventral process at

the base (Fig. 21A). Gular sulcus very flat and shallow.

Pronotum broadest at about basal third of length; lateral margins evenly

arcuate, converging anteriorly, finely emarginate towards obtuse-angled posterior

angle. Margination only interrupted in middle of anterior margin. A fine

punctation of surface visible only near the posterior angle. Base of prosternal

apophysis not calloused as in most species.

REVISION OF THE GENUS COLOPHON GRAY 403

Elytra conspicuously small in relation to large pronotum, almost circular.

Broad sutural margins smooth and raised. No subhumeral callus or other

conspicuous longitudinal sculptural elements. The very fine shagreen produces an

iridescent sheen on the elytra of fresh specimens (lectotype).

Anal sternite asymmetrical, but does not form an off-centre callus.

Anterior tibia broad from the base, straight and parallel, it forms a short and

broad bifid apico-lateral process (Fig. 27A). Ventral crest sharp and well raised

basally, then terminates abruptly between half and two-thirds of tibial length,

forming a rectangle; after a short stretch of interruption forms a small triangular

process near apex.

Aedeagus asymmetrical. Penis strongly sclerotized, apical portion asymmet-

rically dilated. Right paramere forms a broad triangular process at its inner

margin, well removed from apex of paramere; margin between apex and inner

process slightly emarginate. Left paramere contracts towards apex, apex

rounded.

Female. It resembles C. cassoni in raised suture line, which is less distinct

than in male. Anterior tibia, however, has an apico-ventral process, which is

absent in C. cassoni.

Length

Male 15-19 mm, female 15-17 mm.

Distribution

Swartberg Mountains, the northern range of the Cape mountains.

Material examined

74,3 2,16 front body, 14 d heads, 3 2 heads, 12 ¢ anterior legs.

Lectotype. 6: Zwartberg Pass, K. H. Barnard, Nov. 1929; in South African

Museum, Cape Town.

Paralectotypes. 3 3, 2 2, 8 3 heads, 9 ¢ anterior legs: Zwartberg Pass,

K. H. Barnard, Nov. 1929; in South African Museum, Cape Town. 3 6, 1 2:

Zwartberg Pass, K. H. Barnard, Nov. 1929; in British Museum (Natural

History), London.

Additional material. 1 6 front body, 6 d and 3 @ heads, 3 o anterior legs:

Swartberg Pass, Dec. 1968, R. H. Jones and J. H. Potgieter; Transvaal Museum,

Pretoria.

Remarks

The altitude data on the labels of some of the type specimens range between

5 500 and 6 000 ft. [1 677 m and 1 829 ml], and not 5 500 and 6 500 ft. [1 677 m

and 1 982 m] as stated in Barnard’s (1932b) description.

404 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colophon primosi Barnard, 1929

Figs 14A, 20A, 21B, 22A—C

Colophon primosi Barnard, 1929: 175, fig. 9; 1932a: 19, figs. Didier & Séguy, 1953: 76. Brinck,

1956: 314, 320.

Description

Male. Head broadly oblong, rather similar to that of C. izardi, but ocular

corner not sharply raised. Vertex only depressed, frons level transversely and

only slightly deflected to produced clypeus, which is very highly elevated above

the level of flat triangular labrum. Mandibles straight, prognathous, longer and

narrower in lectotype from Seven Weeks Poort Berg than in the Blue Berg

specimens. Apex uniformly emarginate at inner margin, with sharp inward-

turning points (Fig. 21B, 22A, C). Ventral process clearly marked in Blue Berg

specimens, more rounded in lectotype. Surface of head distinctly punctate,

punctatation of mandibles hardly visible. Labrum slightly transverse, oblong;

gular sulcus sharp but rather shallow.

Pronotum with some variations in details; similar but somewhat broader than

in C. izardi. Punctation, if discernible at all, visible only at lateral and anterior

margins.

Fig. 21. Male mandibular structure in Colophon species. A. C. neli Barnard. B. C. primosi

Barnard.

REVISION OF THE GENUS COLOPHON GRAY 405

Elytra more evenly tapered towards apex, less semi-parallel anteriorly than

in C. izardi; subhumeral carina rounded but very distinct; submarginal lateral

depression coarsely corrugated transversely.

Prosternal apophysis hump-like between coxae. Anal sternite asymmetrical,

off-centre apex produced in a transversely oblong and flat process.

Anterior tibia very simple, straight, with four lateral teeth and an inner

apical tooth (Fig. 20A). Ventral crest serrate and continuous, without apical

process. Posterior tibia consistently with a sharp lateral tooth in middle.

Aedeagus of asymmetrical type. Penis strongly sclerotized and asymmetrical.

Right paramere armed at inner margin; apex (distal) truncate, acute-angled; left

paramere narrower than in C. izardi, apical portion parallel with roundly

truncate apex.

Thorax, body, elytra, and head (except vertex), black; vertex, mandibles and

legs (except tarsi), reddish brown.

Female. Head conspicuously small (Fig. 22B). Black, but coxae, humeri,

tibiae and parts of head reddish brown. More convex than in male, lateral

corrugation of elytra less distinct. Prosternal process similar; anal sternite sharply

punctate, punctures with fine and short hairs; anal sternite in male only finely

punctured and bare.

Length

Male 28-35 mm (lectotype 35 mm); length of mandibles 8-10,5 mm

(lectotype 10,5 mm); female 20-22 mm.

Distribution

Central section of Swartberg range, northern range of Cape mountains.

Material examined

5 3 @ fresh, 2 damaged) (2 dissected), 7 2 (5 fresh, one damaged, one

fragmented). Lectotype ¢, 2 2 paralectotypes, and additional 3 ¢ and 2 9

specimens in South African Museum, Cape Town; 1 6 and 2 2 paralectotypes

in British Museum (Natural History), London. 1 2 in Transvaal Museum,

Pretoria.

Lectotype. 3: Seven Weeks Poort Berg, Ladysmith District, 7 000 ft.

[2 134 m], K. H. Barnard, 1928; in South African Museum, Cape Town.

Paralectotypes. 1 36, 2 2: Seven Weeks Poort Berg, Ladysmith District,

7 000 ft. [2 134 m], K. H. Barnard, 1928; in British Museum, London. 2 2 (one

badly fragmented): Seven Weeks Poort Berg, Ladysmith District, 7 000 ft.

[2 134 m], K. H. Barnard, 1928; in South African Museum, Cape Town.

Further material. 1 fresh 6, 2 damaged 6, 2 fresh 2: Blue Berg, Zwartberg

range, 1930, F. Berrisford; in South African Museum, Cape Town. 1 2: Seven

Weeks Poort Berg, 6 300 ft. [1 921 m], 2 Feb. 1986, coll J. Vlok; in Transvaal

Museum, Pretoria.

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ANNALS OF THE SOUTH AFRICAN MUSEUM

406

REVISION OF THE GENUS COLOPHON GRAY 407

Remarks

Colophon primosi is the most peculiar species of the genus, with mandibles

not readily derived from any other forms. Other characters, however, secure its

place in the genus and also in the apomorphic lineage (asymmetrical aedeagus,

raised clypeus).

All specimens of the type series were found dead, some of them very

distinctly bleached, leaving Barnard (1929: 176) in doubt in respect of their

natural coloration. Three fresh specimens (1 6, 2 ¢) of the Blue Berg series were

probably found live and confirm the unusual coloration described by him. Male

mandibles, anterior part of head, gula and all femora and tibiae light yellow-

brown; rest of body, antennae and tarsi black. In females the coloration is similar,

except mandibles are black in small specimens.

There is no noteworthy difference between specimens of the Seven Weeks

Poort Mountain and Blue Berg samples, which raises some interesting historic

biogeographic problems (see p. 362).

Colophon westwoodi Gray, 1832

Figs 8, 20B, 23, 24A

Colophon westwoodi Gray, 1832: 533, pl. 46 (fig. 5). Castelnau, 1840: 173. Westwood,

1855: 194. Parry, 1864: 70. Péringuey, 1901: 3. Van Roon, 1910: 8. Boileau, 1913: 218.

Handlirsch, 1924: 695. Barnard, 1929: 166, fig. 1; 1932a: 19, figs. Didier & Séguy, 1953: 77.

Brinck, 1956: 309, 320.

Diagnosis

Black, medium to large size. Most of anterior margin of pronotum

marginate. Mandibles short and strongly curved inward, main arm broadly bifid

with rounded protuberance dorsally; ventral and dorsal process in basal position,

small and roundly pointed. Anal sternite asymmetrical, aedeagus of asymmetrical

type. Anterior tibia finely arcuate, dilated towards apex, with four or five lateral

teeth not quite regularly increasing in size distally.

Description

Male. Head broadly oblong, genal margin in front of eyes semi-parallel,

occasionally finely emarginate, anterior angle sharp. Anterior margin moderately

sinuate between mandibles, a canthus above clypeus sometimes more or less

protuberant in middle. Vertex deflected to pre-clypeal canthus, overhanging the

clypeus; latter is flush with the base of flat, triangular labrum. A somewhat similar

canthus is present only in C. primosi. Ocular canthi flat. Vertex with a pair of

anterior protuberances and an even more distinct median protuberance.

Punctation strong, dense and rather uniform. Mandible short and stout, main arm

broad and short, evenly curved in about 90° towards equally broad and bifid apex.

There is a large and rounded protuberance of different elevations in the middle of

the dorsal surface, close to inner margin. As a small dorsal process is present near

the base, it is thought that the dorsal protuberance is not analogous with the

408 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. Colophon westwoodi Gray, male; length

20 mm.

dorsal processes of other species (Fig. 24A). Mentum almost flat, about twice as

broad as long, slightly converging anteriorly, apex rounded. Transverse gular

sulcus flat.

Pronotum even in large males only moderately broader than elytra, rather

convex. Broadest in front of basal third, lateral margins evenly arcuate and

converging anteriorly, emarginate posteriorly to sharp and rectangular posterior

angle. All edges margined, margination only interrupted in middle of anterior

margin. Punctation very fine, but sharp, uniformly dense; hardly more distinct

laterally.

Elytra about as long as combined breadth, semi-parallel in anterior half of

length; do not appear as disproportionately small in relation to the pronotum, as

in most other species. Surface smooth and matt-shagreened with eight irregular

rows of fine punctures. Lateral submarginal depression more densely, but not

more coarsely, punctate.

REVISION OF THE GENUS COLOPHON GRAY 409

Anal sternite very distinctly asymmetrical.

Anterior tibia moderately arcuate at inner margin, evenly dilated from base

to apex. Lateral margin with four or five progressively larger triangular processes;

last two (largest) processes do not appear as a bifid unit (Fig. 20B). Ventral crest

continuous from base to apex, not forming an apico-ventral process. Lateral

median process of posterior tibia as large and sharp as in intermediate tibia (in

other species usually reduced or missing).

Aedeagus of asymmetrical type. Penis strongly sclerotized, apical portion

asymmetrically dilated. Right paramere produced to a large inner process; its

anterior margin to apex is straight, posterior margin evenly and deeply

emarginate towards base. Left paramere narrow with rounded apex.

Female. Distinguished by anteriorly marginate pronotum with sharply

pointed rectangular posterior angle.

Fig. 24. Male mandibular structure in Colophon species. A. C. westwoodi Gray. B. C. thunbergi

Westwood. C. C. barnardi sp. nov.

Length

Male 21-25 mm, female 19-24 mm.

Distribution

Table Mountain, Cape Town.

Material examined

14 3 and 19 2. Holotype and three additional specimens in British Museum

(Natural History), London; 24 specimens in South African Museum, Cape Town,

and 5 specimens in Transvaal Museum, Pretoria.

Holotype. 3: no data, ex Vigin’s collection, British Museum, London.

Additional material. The other material listed above has been recorded from

Devil’s Peak, Constantiaberg, Kalk Bay Mountain, and Echo Valley. Live

specimens were collected in December, January and February.

Remarks

This was the first species of the genus to be described, and ever since it has

been the most frequently collected.

410 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colophon barnardi sp. nov.

Figs 20E, 24C, 25

Diagnosis

Small black species. Mandibles with ventral process only. Anterior tibia with

three apico-lateral teeth. Clypeus raised above labrum. Aedeagus of asymmetri-

cal type.

Description

Male. Medium size, completely black. Head broadly oblong with sharp genal

canthus and with rounded and arcuate elevations at the inner margins of eyes.

Clypeus slightly produced and sharply elevated over labrum. Surface densely and

sharply punctate, integument matt. Mandibles short, sickle-shaped, arcuate

through 90 degrees. Dorsal process entirely absent, ventral processes large, their

inner margins parallel; apex of processes forward-pointing, acute-angled. Apex of

main arms simple (Fig. 24C). Mentum flat, hardly convex transversely, apical

margin broadly arcuate. Pregular sulcus deep.

Pronotum evenly convex transversely, more deflected towards posterior than

towards anterior margin. All around marginate; submarginal lateral depression

very narrow anteriorly, slightly dilated at lateral angle of pronotum. Pronotum

broadest at posterior third of length, gradually more arcuately contracted towards

anterior lateral angle; pronotum considerably contracted posteriorly, but lateral

margin straight to obtuse-angled posterior lateral angle. Disc smooth and matt

with scattered punctures at lateral and anterior margins. Base of prosternal

apophysis sharply dentate.

Elytra distinctly longer than combined breadth, not particularly small

compared to pronotum, narrower than the latter. Humeral angle rectangular,

narrowly rounded. Disc convex, submarginal depression broad anteriorly;

gradually narrower behind; suture not elevated, marginate or smoother than disc.

Surface entirely smooth, without any costal elements, matt with indistinct traces

of scattered punctation.

Anterior tibia straight, narrowest at basal quarter of length; dilated in apical

half, forming three lateral teeth; inner apical angle sharply produced (Fig. 20E).

Ventral crest moderately elevated in basal half, gradually diminishing anteriorly;

apico-ventral process absent.

Anal sternite asymmetrical at apical margin, forming a narrow process to the

right of median line (ventral view).

Aedeagus of asymmetrical type.

Female. Pronotum slightly narrower, but of similar shape to that of male;

disc of elytra similarly smooth (paratype) or with short, subhumeral costal

element (allotype).

Length

Male 20 mm, female 20,2—21,0 mm.

REVISION OF THE GENUS COLOPHON GRAY 411

Fig. 25. Colophon barnardi sp. nov., male; holotype;

length 20 mm.

Material examined

16,2 2, and1 6 thorax with right leg attached. Holotype ¢ and allotype @

in South African Museum, Cape Town; 2 paratypes (1 ¢ thorax, 1 @ lacking

abdomen) in Transvaal Museum, Pretoria.

Holotype. 3 (dead): Leeuw River Peak, Swellendam, April 1941, K. H.

Barnard.

Allotype. 2 (dead): Leeuw River Peak, Swellendam, April 1941, K. H.

Barnard.

Paratypes. 1 6 thorax, 1 2: Leeuw River Peak, Swellendam, April 1941,

K. H. Barnard.

All specimens were labelled by Barnard as C. thunbergi.

Etymology

This species is named in honour of Dr K. H. Barnard, collector of most of

the known specimens of Colophon and first reviser of this genus.

412 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colophon thunbergi Westwood, 1855

Figs 24B, 26

Colophon thunbergi Westwood, 1855: 198. Parry, 1864: 70; 1870: 71; 1872: 83. Boileau,

1913: 217. Barnard, 1929: 167, fig. 2; 1932b: 170, fig. 1. Didier & Séguy, 1953: 77. Brinck,

1956: 310, 320.

Diagnosis

Rather large, black species. Mandible with dorsal and ventral processes.

Clypeus raised above labrum. Anterior tibia slender and bent at base, dilated

with bifid or trifid apico-lateral armature.

Description

The description below is based on the holotype and points out, where

necessary, the differences shown by the fragments.

Male. Head broad and short, angularly oblong, anterior angle of gena only

slightly obtuse-angled, narrow, anterior margin not indented or emarginate near

lateral angle. Anterior margin evenly sinuate in middle. Disc rather sharply raised

at inner margin of eye, moderately depressed anteriorly in middle, with a pair of

round and indistinct elevations at either side of depression. Punctation consists of

rather uniform, moderately large, well-spaced punctures in the middle; integu-

ment densely shagreened. Mandibles short and stout, bent more or less at a right

angle; dorsal process near base conical; in the fragments less forward-pointing (as

figured). Ventral process very broad, with inner corner roundly produced

anteriorly, reaching about two-thirds of mandibular length (Fig. 24B). Labrum

flat and pointedly triangular, clypeus sharply elevated above its base. Mentum

large and flat, oblong, anterior margin truncate. Pregular sulcus evenly concave,

not sharply incised.

Fig. 26. Colophon thunbergi Westwood,

male; length 15mm. An estimated total

length of the specimen could be 25 mm.

REVISION OF THE GENUS COLOPHON GRAY 413

Pronotum broadest at about basal third; margins very slightly emarginate or

straight, contracting to obtuse-angled basal corner. Margination also complete

anteriorly. Pronotum in the holotype evenly rounded disciform, lateral margins

evenly arcuate, converging to perfectly rectangular anterior angle. Surface

densely shagreened, also very finely punctured at sides.

Elytra semi-parallel at anterior portion, very slightly contracted towards

rectangular humeral angle. Surface of disc smooth, densely and finely shagreened

and indistinctly punctate. A much reduced subhumeral costa is slightly elevated at

basal fifth of length (some further); hardly discernible longitudinal lines are

shown only by smoother integument. Dimensions in holotype 11,5 x 11,8 mm.

Anal sternite distinctly asymmetrical, off-centre process flatly rounded.

Anterior tibia bent inward in dorsal view. In holotype almost the whole basal

third narrow, apical dilation disciform. Apical armature bifid (Fig. 26). In all

fragments the narrow basal portion is shorter, apical dilation more gradual, and

the apical armature consists of three triangular processes (in one tibial fragment

the proximal process is clearly broken off). Ventral crest continuous between

base and apex, evenly and moderately elevated, not raised at base as in

C. barnardi. In the latter species the tibia is more slender and much straighter.

Aedeagus of asymmetrical type.

Female. The single female in the British Museum, from Zuurbrak Peak, is

herein tentatively retained in C. thunbergi. It is somewhat different from the two

females from Leeuw River Peak that are herein referred to C. barnardi (see

p. 411). The Zuurbrak Peak female differs from the Leeuw River Peak females in

the following characters: ocular canthus sharper, punctation of head coarser and

sparser. Elytron in the Zuurbrak specimen is marked by two punctured costae of

common subhumeral origin; they reach to about the middle of the elytral length.

Length

Male (holotype) 26 mm.

Material examined

1 3 (holotype); 1 @ (identity doubtful), in British Museum (Natural

History), London. 5 6 heads and 5 6 anterior legs, South African Museum,

Cape Town.

Holotype. 3: no locality, with labels “Type Col. 247; C. thunbergi and

C. westwoodi Gray i[n]. litt. in the Hope Museum, Oxford’.

Additional material. 1 @ (identity uncertain): Zuurbrak Peak, in British

Museum, London. The 5 ¢ heads and 5 ¢ anterior legs are from Zuurbrak Peak,

Langeberg Range, near Swellendam; coll. Barnard & Thorne, February 1932;

South African Museum, Cape Town. These latter fragments indicate the possible

locality of the holotype.

Remarks

The locality of the holotype in the Hope Museum, Oxford, is not known, but

morphologically it agrees sufficiently with the fragments collected by Barnard at

414 ANNALS OF THE SOUTH AFRICAN MUSEUM

Zuurbraak Peak, where Colophon thunbergi is sympatric with C. izardi. The

slight differences found between the holotype and the fragments do not appear to

be greater than the range observed among the fragments—with the possible

exception of bifid tibial armature in holotype.

Colophon whitei Barnard, 1932

Figs 27D-E, 28, 29A

Colophon whitei Barnard, 1932b: 171, fig.3; 1932a: 20, figs; 1936: 8. Didier & Séguy, 1953: 17.

Brinck, 1956: 311, 320.

Diagnosis

Large black species. Mandible with dorsal and ventral processes. Anterior

tibia bent inward, dilated lateral margin deflected and deeply excised before

reaching bifid apical armature. Anal sternite and aedeagus of asymmetrical type.

Description

Male. Head broadly oblong with sharp genal canthus. Surface longitudinally

elevated at inner margins of eyes, elevations rounded and slightly inward slanting

from frons towards base; a similar, broadly funnelling elevation occupies the

whole frons. Frons steeply deflected to clypeus. Clypeus broad, flatly emarginate

at anterior margin, showing some variation in breadth and depth of emargination;

sharp anterior edge of clypeus raised above the level of labrum. Labrum flat,

elongately triangular, sometimes briefly bifid at apex. Integument of head more

densely and coarsely punctate in depressions than on elevations. Mandibles with

dorsal and ventral processes (Fig. 29A); main arm prognathous, usually only

A 7 B C D E F

Fig. 27. Male anterior tibiae in Colophon species. A. C. neli Barnard. B.C. montisatris

sp. nov. C.C. montisatris sp. nov. in lateral view. D.C. whitei Barnard. E. C. whitei

Barnard, lateral view. F. C. izardi Barnard.

REVISION OF THE GENUS COLOPHON GRAY 415

apical third curved inward through 90 degrees; apices, particularly that of the left

mandible, narrow and truncate. In the inner margin of each mandible two

processes appear. The ventral process seems to be homologous with that of

C. cassoni, C. eastmani and C. cameroni, but the more dorsal one is probably

not. This tooth lies in the midline of the mandible’s thickness in C. whitei,

whereas in the other three species it lies flush with the dorsal surface. This

mandibular process in semi-dorsal position appears as a unique character among

the known species of the genus. The most closely related species, C. montisatris

sp. nov., entirely lacks this semi-dorsal tooth, although its ventral one is

homologous with that of C. whitei. The rudimentary semi-dorsal process of

C. barnardi might be homologous with that of C. whitei. Mentum slightly

concave, lateral margins parallel, anterior lateral angles narrow; truncate anterior

margin slightly produced in middle. Pregular sulcus deep.

Pronotum evenly convex transversely, more deflected to posterior than to

anterior margin; broadest at the basal third of length where lateral margins are

parallel; margins evenly curved towards produced anterior angle; distinctly

emarginate behind; posterior angle obtuse. All margins evenly and sharply

bordered, margination only interrupted in a spot in the middle of the anterior

margin. Integument smooth but not shiny, with a few fine punctures laterally; disc

without, or occasionally with, faint scattered punctures. Base of prosternal

apophysis with a small callus. Scutellum triangular with arcuate margins showing

some variation in breadth and shape.

Elytra narrower than the widest part of the pronotum but broader than the

pronotum at its posterior angles. Humerus narrowly rounded, almost rectangular.

Disc convex, submarginal depression broad, integument here more or less

distinctly corrugated. Surface otherwise smooth, matt with fine and scattered

punctation. Suture line not conspicuously raised, although sometimes smoother

and shinier than rest of surface.

Anterior tibia gradually dilated from base, dilation turns gradually into a

latero-ventral position; dilation is abruptly reduced at apical quarter where it

forms a sharp rectangular corner; apico-lateral corner of tibia is formed by a bifid

process (Fig. 27D, E). Median (primary) ventral crest distinct. Inner margin of

anterior tibia almost straight. Intermediate tibia with a single, sharply pointed

median process; posterior tibia unarmed.

Apical margin of anal sternite strongly asymmetrical in middle.

Aedeagus asymmetrical; right paramere with large inner process; left

paramere simple, comparatively broad.

Female. Pronotum narrower than in males, disc similarly smooth, only

punctate near lateral margins; a faint longitudinal median impression appears in

some specimens. Humeral angle rounded, lateral margins finely arcuate. Surface

smooth and matt with scattered, very fine punctures. No distinct costal elements.

No reliable differences could be found between females of C. whitei and of

C. montisatris sp. nov.

416 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 28. Colophon whitei Barnard, male; lectotype;

length 25,5 mm.

Length

Male 26-31 mm, female 19-22 mm.

Distribution

Central section of the Swartberg (northern range of the Cape mountains),

west of Meiringspoort.

Material examined

15 3 (3 damaged, 3 bleached, 1 anterior part of body), 5 2, intact

specimens; 5 ¢ and 2 heads.

Lectotype. 3, Meiringspoort Berg, Zwartberg Range, Febr. 1932, K. H.

Barnard and C. W. Thorne; South African Museum, Cape Town.

REVISION OF THE GENUS COLOPHON GRAY 417

Paralectotypes. 8 6 and 5 &, Meiringspoort Berg, Zwartberg Range, Febr.

1932, K. H. Barnard and C. W. Thorne. (Of these 7 ¢ and 1 2 in South African

Museum, Cape Town, and 1 6 and 2 2 in British Museum, London.) 6 ¢ and

5 ¢ heads: Meirings Poort Berg, Zwartberg Range, [6 500 ft. (= 1 982 m),

March] 1931, A. B. Berrisford; 1 @ (data as for holotype) does not belong to this

species; 1 d, 2 2: Blesberg, Zwartberg Range, 1930, F. Berrisford (locality

probably erroneous; see below), in South African Museum.

Remarks

This and the following species (C. montisatris sp. nov., p. 418) are closely

related and their separation might have occurred only after the present

populations retreated to the high altitudes of the Swartberg Range, on either side

of Meirings Poort. It is evident that the common ancestor had a distribution area

on the flat at the level of the foot of the mountain, since Meirings Poort cuts

through the latter and serves to drain water from the north to the south.

The locality date for the specimens recorded from Blesberg must be regarded

as doubtful. These three specimens formed part of Barnard’s (1932b) type series.

However, Blesberg is on the east side of Meiringspoort. The identity of the

females could not be ascertained, but the male is certainly conspecific with the

rest of the type series of C. whitei from the west side of Meiringspoort. Blesberg is

the type locality of C. montisatris sp. nov., where 10 6 specimens (2 anterior

parts of body) of that species were collected in 1979 and 1981 without finding any

fragments of C. whitei. It seems to be almost certain that at least the male

specimen of C. whitei recorded from Blesberg is incorrectly labelled, and was

actually collected on the west side of Meiringspoort. Similar problems were

encountered with material of C. stokoei (see p. 393), where the possibility of

erroneous locality labelling made the final evaluation of C. stokoei morphs

impossible. It has to be taken into consideration that a considerable portion of

Colophon specimens were collected by non-entomologist mountaineers.

A female specimen labelled as C. whitei, and with the same locality data as

the lectotype, does not belong to this species.

Specimens mentioned in the Barnard’s (1932b) description as collected at

Blou Punt (Blaauw Punt) and Spitzkop (both on the west side of Meiringspoort),

could not be found in the South African Museum, nor in the British Museum,

although there is no record that Barnard disposed of any specimens to any other

museums.

Variability

In evaluating the characters of 14 complete males, one pronotum and head,

and seven heads, the variability in main characters, such as mandible, mentum

and anterior tibia, was found to be minimial. No allometric variation was found

between the biggest and smallest specimens, except the relatively greater breadth

of pronotum in the larger specimens. In the type series there is a specimen with

rather distinct longitudinal striation of elytra, which gives the impression of an

418 ANNALS OF THE SOUTH AFRICAN MUSEUM

ontogenetic anomaly. A certain ontogenetic anomaly is shown in another

specimen in which the right mandible is about half normal size, and having

reduced but distinct male characters—and not partially hermaphrodite as is

found occasionally in Lucanidae. Would the two malformations in 14 specimens

indicate the genetic effect of in-breeding in a tiny extant population?

Fig. 29. Male mandibular structures in Colophon species. A. C. whitei Barnard.

B. C. montisatris sp. nov. C. C. izardi Barnard, semi-lateral view.

Colophon montisatris sp. nov.

Figs 1, 4, 27B—C, 29B, 30A-B

Remarks

Colophon montisatris is a sister species of C. whitei. In many respects the

new species seems to be the more ancestral and, in its characters, closer to the

hypothetical common ancestor. Characters not described below were found to be

identical with those described for C. whitei.

Description

Male. Declivity of frons towards clypeus not quite vertical; clypeus broader,

distinctly emarginate. Labrum similar. Mandibles with ventral process only (the

most diagnostic specific character). Main arms of mandibles slightly shorter than

in C. whitei, not tapering towards broader apex. Ventral process sharply pointed,

less produced, and altogether closer to base than in C. whitei (Fig. 29B).

Mentum, pronotum and elytra very similar to those of C. whitet.

Anterior tibia angularly nicked inward in middle, in dorsal view semi-

parallel, not dilated towards apex. Lateral crest more deflected ventrally,

narrower, anterior emargination towards bifid apico-lateral process deeper

Aedeagus similar to that of C. whitei, but left unarmed paramere narrower.

Female. No characters could be found that enable one to distinguish between

females of this species and those of C. whitei.

REVISION OF THE GENUS COLOPHON GRAY 419

Length

Male 23-27,8 mm (breadth 11-14 mm), female 23 mm.

Material examined

10 6, 1 2 and 2 6 anterior bodies. Holotype ¢, allotype 2 and 6 ¢

paratypes in Transvaal Museum, Pretoria; 1 d paratype each is deposited in the

South African Museum, Cape Town, British Museum (Natural History),

London, and Natural History Museum, Budapest.

Holotype. 3, Swartberg Range, Blesberg, 2.III.1979, leg. Endrédy-Younga,

E-Y:1556.

Allotype. 2 (dead), Swartberg Range, Blesberg, 6.III.1981, E-Y:1755.

Paratypes. 4 3, Swartberg Range, Blesberg, 2.III.1979, leg. Endrédy-

Younga, E-Y:1556. 1 6 (dead): Swartberg Range, Blesberg, 6.III.1981,

Fig. 30. Colophon moniisatris sp. nov. A. Holotype, male; length 25 mm. B. Allotype, female; length

23 mm.

420 ANNALS OF THE SOUTH AFRICAN MUSEUM

E-Y:1755. 1 ¢ (dead) and 1 ¢ anterior body: Swartberg Range, Blesberg,

17.XII.1978, E-Y:1534. 1 ¢ live and 1 d dead, Swartberg Range, Blesberg,

5.111.1981, E-Y:1754. 1 3 live, 1 6 anterior body, Swartberg Range, Blesberg,

7 III.1981, E-Y:1756, (3 further d seen).

Variation

Allometric variation is evident from the specimens examined. The lengths (in

mm) of the ten complete males are: 22,3; 23,9: 24,9: 25,0: 26,9: 27/02 27-42 705-

27,5; and 27,5. All specific characters can be found on both the smallest and

largest specimens, though moderate accentuation of secondary male sexual

characters was observed in large specimens. With increase in size mandibles

become larger with sharper edges; ventral process, flush with main arm at ventral

surface in small specimens, is produced ventrally in large specimens; pronotum

becomes broader and proportionately bigger than elytra. There is no distinct

difference in anterior tibia between small and large specimens.

Colophon izardi Barnard, 1929

Figs 2, 3, 27F, 29C, 31A-B

Colophon izardi Barnard, 1929: 173, fig. 8; 1932a: 20, figs. Didier & Séguy, 1953: 77. Brinck,

1956: 310, 320.

Diagnosis

Male anterior tibia narrow and bent inward at 90 degrees. Armature of male

mandibles forms a funnel in closed position. Pronotum in both sexes usually with

a pair of red dots.

Description

Male. Head broadly oblong, gena emarginate in front of eye, anterior angle

narrow, obtuse-angled. Ocular canthus distinct but rounded, disc broadly

depressed in between. Punctation denser and coarser laterally. Frons only slightly

deflected towards clypeus, which is visible in dorsal view. Clypeus broad, finely

emarginated and sharply raised above level of labrum; labrum flat and broadly

triangular. Mandibles very characteristic: main arms simple, sickle-shaped, with

flat dorsal surface. Ventral process reaches maximum development in this

species; in dorsal view as much produced as main arms, broad from the base, the

whole mandibular structure forming a funnel towards clypeus (not towards the

mouth, which is beneath) (Fig. 29C). Gula broadly rounded, gular sulcus sharp

and deep.

Pronotum as long as elytra, slightly broader than latter, bearing a pair of

large, round centrally bilateral reddish-brown patches. Broadest at basal third,

flatly and evenly curved anteriorly, evenly tapering behind, not emarginate

prebasally. All edges margined, only briefly interrupted in middle of anterior

margin. Punctation very fine and scattered but distinct, stronger laterally.

Elytra with fine subhumeral crest, extending from base to middle of elytron.

REVISION OF THE GENUS COLOPHON GRAY 421

Anal sternite asymmetrical, produced apex not calloused.

Anterior tibia apomorphic in shape: anterior third bent 90° inward; ventral

crest low and continuous, lateral crest dilated pre-apically, then sharply

interrupted, emargination between lateral crest and undivided apico-lateral

process deeply U-shaped (Fig. 27F).

Aedeagus of asymmetrical type. Penis strongly sclerotized, asymmetrical.

Right paramere armed inwards; inner process similar, but apex rectangular and

not acute-angled as in C. whitei. Left paramere angularly emarginate at inner

margin.

Female. Punctation of head, pronotum and elytra much more distinct than in

male. Inner margin of anterior tibia evenly curved. Anal sternite symmetrical,

densely punctate and finely pubescent, whereas in male it is smooth and bare.

Discoidal red patches of pronotum similar to male.

Length

Male 21,8—25,3 mm, female 20-21 mm.

Fig. 31. Colophon izardi Barnard. A. Male; length 22,5 mm. B. Female; length 19,5 mm. The

first live specimen found during this project.

422 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Central section of Langeberg Range, the southern range of the Cape

mountains.

Material examined

5 6,5 2; 1764 heads, 1 2 head; 17 6 anterior tibia, 2 2 anterior tibia.

Complete specimens are in the British Museum (Natural History), London;

South African Museum, Cape Town; and Transvaal Museum, Pretoria.

Lectotype. 6: Lemoenshoek, Heidelberg, C.P., 5 000 ft. [= 1 524 m], K. H.

Barnard; British Museum (Natural History), London.

Paralectotypes. 3 2: Lemoenshoek, Heidelberg, C.P., 5 000 ft. [= 1 524 ml],

K. H. Barnard (1 2 in British Museum, London; 2 ¢ in South African Museum,

Cape Town).

Additional material. 1 3: Lemoenshoek, January 1957, T. P. Stokoe. 2 6

pronota: Grootberg, Langeberg, 1.XI.1978, Endrédy-Younga, E-Y:1499. 1 9:

Grootberg, Langeberg, 7.12.1978, E-Y:1527. 3 d, 1 2: Grootberg, Langeberg,

SMES 7 OSE NGIS 02:

Remarks

Some females, due to their entirely black pronota, were excluded from the

type series by Barnard, as were some fragments from localities from where no

complete males were known. These included 3 2, 8 d heads and 7 ¢ anterior

legs, Riversdale Mountains, tops of peaks (Kampsche Berg in Barnard’s

description); 1 2, Zuurbrak Peak, Swellendam Dist., 5 000 ft. [= 1 524 ml],

K. H. Barnard, Oct. 1925; 9 ¢ heads, 1 2 head, 10 6 and 2 @ anterior legs:

Tradouw Peak, Swellendam Distr., 4 400 ft. [= 1 341 m], K. H. Barnard, Oct.

1928.

It is remarkable that specimens with red pronotal dots came only from the

Lemoenshoek—Grootberg area (11 specimens), and specimens with the pro-

notum entirely black from Kampsche Berg and Zuurbrak Peak. A final decision

on the status of the black specimens is further complicated by the fact that they

are all females, which are in any case difficult to identify. However, male

mandibles and anterior legs from these localities do not show any difference when

compared to the type specimens. Furthermore, one of the male specimens

collected live in Grootberg (E-Y:1562) shows a very definite reduction of the red

marking of its pronotum. The dots in this specimen are hardly more than 1 mm in

diameter and dark reddish brown. Though further material would be of great

interest, I have little doubt that the black-pronotum specimens are C. izardi

Barnard.

ACKNOWLEDGEMENTS

My sincere thanks are due to my colleagues who enabled me to study the

valuable material preserved in their respective museums: Dr V. B. Whitehead,

South African Museum, Cape Town; Mr M.E. Bacchus, British Museum

REVISION OF THE GENUS COLOPHON GRAY 423

(Natural History), London; and Mrs A. Z. Smith, Hope Museum, Oxford

University. I am also grateful to Prof. H. E. H. Patterson, University of the

Witwatersrand, Johannesburg, Dr M.-L. Penrith, State Museum, Windhoek, and

Dr V. B. Whitehead, South African Museum, Cape Town, for their critical

reading and valuable comments on this paper.

REFERENCES

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163-182.

BARNARD, K. H. 1932a. The Colophon. J. Mountain Club S. Afr. 34: 19-22.

BARNARD, K. H. 19326. The rediscovery of Colophon thunbergi Westw. with descriptions of

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BARNARD, K. H. 1936. Geographical zoology. S. Afr. geogr. J. 19: 3-13.

BoiLEAu, H. 1911. Description de quelques espéces nouvelles de Lucanides appartenant aux

collections du British Museum. Trans. ent. Soc. Lond. 1911: 426-451.

BoiLEAu, H. 1913. Note sur les Lucanides conservés dans les collections de l’Université

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BRAIN, C. K. 1981. The evolution of man in Africa: was it a consequence of Cainozoic cooling?

Alex L. du Toit Memorial Lectures No. 17. J. geol. Soc. S. Afr. 84 (annexure): 1-19.

BrINcK, P. 1956. Coleoptera, Lucanidae. In: South African animal life 3: 304-335.

Stockholm: Almqvist & Wiskell.

CASTELNAU, F. L. DE Laporte. 1840. Histoire naturelle des animaux articulés coléopteéres 2.

Paris: P. Dumellin.

Coope, G. R. 1979. Late Caenozoic fossil Coleptera: evolution, biogeography and ecology.

Annu. Rev. Ecol. Syst. 10: 247-267.

Dipigr, R. & SéGuy, E. 1953. Catalogue illustré des Lucanides du Globe. Text and atlas.

Paris: Paul Lechavalier.

ELDREDGE, N. & GouLp, S. J. 1972. Punctuated equilibria. An alternative to phyletic

graduation. In: ScHopr, T. M. ed. Models in paleobotany: 82-115. San Francisco:

Freeman, Cooper & Co.

ENDRODY-YOUNGA, S. 1978. Coleoptera. In: WeERGER, M. J. A. ed. Biogeography and

ecology of southern Africa 2: 797-821. The Hague: Junk.

ENDRODY-YOUNGA, S. 1981. The concept of heteromorph speciation. Consequences of

speciation in homomorph and heteromorph populations. Ann. Transv. Mus. 32: 241-247.

ENDRODY-YOUNGA, S. 1982. Dispersion and translocation of dune specialist tenebrionids in the

Namib area. Cimbebasia (A) 5: 257-271.

Gray, G. R. 1832. New species of insects of all.orders. In: GrirFirH, E. The animal kingdom

arranged in conformity with its organization by the Baron Cuvier 14. London: Whittaker.

HAND.urscH, A. 1908. Die fossilen Insecten und die Phylogenie der rezenten Formen. Leipzig:

Wilhelm Engelmann.

HEnnIGc, W. 1966. Phylogenetic systematics. (Translation by D. Dwight and R. Zanger.)

Urbana: University of Illinois Press.

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WEstwoop, J. O. 1855. Descriptions of some new species of exotic Lucanidae. Trans. ent.

Soc. Lond. 3: 197-221.

6. SYSTEMATIC papers must conform to the International code of zoological nomenclature (particu-

larly 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.,

ete.

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 trans-

ferred 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

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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 fuil for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

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tion 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 Eliza-

beth (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.

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S. ENDRODY-YOUNGA

EVIDENCE FOR THE LOW-ALTITUDE ORIGIN OF

THE CAPE MOUNTAIN BIOME DERIVED FROM

THE SYSTEMATIC REVISION OF THE GENUS

COLOPHON GRAY (COLEOPTERA, LUCANIDAE)

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