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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

507,68

VOLUME 78 BAND

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1979

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

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

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ANNALS OF THE ANNALE VAN DIE

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VOLUME 78 BAND 78

us “

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{

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van

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—

LIST OF CONTENTS

BERRY, P. F.

A new species of deep-water palinurid lobster (Crustacea, Decapoda, Palinuridae)

from the east coast of southern Africa. (Published June 1979.) ..

COETZEE, D. J. see DE DECKER, A. H. B.

Cooper, M. R.

A new species of Myophorella (Bivalvia, Trigoniidae) from the Sunday’s River

Formation, South Africa. (Published April 1979.)

Cooper, M. R.

Cretaceous Trigoniidae (Mollusca, Bivalvia) from the Brenton Formation, Knysna.

(Published May 1979.)

DE Decker. A. H. B. & COETZEE, D. J.

Indicator copepods and oil yield fluctuations in pelagic fish in the Benguela Current

system. (Published June 1979.)

DINGLE, R. V., Lorp, A. R. & HENDEY, Q. B.

New sections in the Varswater Formation (Neogene) of Langebaan Road, south-

western Cape, South Africa. (Published June 1979.)

HENDEY, Q. B. see DINGLE, R. V.

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

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

Cretaceous faunas from Zululand and Natal, South Africa. A new genus and

species of tuberculate desmoceratacean ammonite from the Mzinene Forma-

tion (Albian). (Published April 1979.) ..

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

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

Cretaceous faunas from southern Africa. Lower Cretaceous ammonites, including

a new bochianitid genus, from Umgazana, Transkei. (Published April 1979.)

Lorp, A. R. see DINGLE, R. V.

PRINS, A. J.

Discovery of the Oriental latrine fly Chrysomyia magacephala (Fabricius) along

the south-western coast of South Africa. (Published April 1979.) Z

SCHWEITZER, F. R.

Excavations at Die Kelders, Cape Province, South Africa. The Holocene deposits.

(Published August 1979.)

SIMPSON, G. G.

A new genus of late Tertiary penguin from Langebaanweg, South Africa. (Published

April 1979.) .. ‘ a me eye oe 7 s - te)

WRIGHT, C. W. see KENNEDY, W. J.

Page

101

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NEW GENERIC AND SUBGENERIC NAMES PROPOSED

IN THIS VOLUME

Dege Simpson, 1979 .. ae ous

Umgazaniceras Klinger & Kennedy, 1979 ..

Umsinenoceras Kennedy & Klinger, 1979 ..

VOLUME 78 PART 1 APRIL 1979 ISSN 0303-2515

567, 68

OF THE SOUTH AFRICAN

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

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

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

Zool. exp. gén. 74: 627-634.

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

Ann, Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 78 ~ Band

April 1979 April

Part 1 Deel

A NEW GENUS OF LATE TERTIARY PENGUIN

FROM LANGEBAANWEG, SOUTH AFRICA

GEORGE GAYLORD SIMPSON

Cape Town Kaapstad

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A NEW GENUS OF LATE TERTIARY PENGUIN FROM

LANGEBAANWEG, SOUTH AFRICA

GEORGE GAYLORD SIMPSON

The Simroe Foundation, 5151 East Holmes Street, Tucson, Arizona 85711

(With 5 figures and 3 tables)

[MS. accepted 11 December 1978]

ABSTRACT

Additional specimens of penguin bones from the ‘E” Quarry in the latest Miocene/early

Pliocene Varswater Formation make possible the identification of the larger of the two penguin

taxa in that fauna and its definition as a new genus and species, Dege hendeyi.

CONTENTS

PAGE

Introduction - 1

Systematics 3

Acknowledgements 8

References 8

INTRODUCTION

In two previous papers (Simpson 1971, 1975) an extinct genus and species

of penguins, Inguza predemersus, was described and named from the late

Tertiary of Langebaanweg. It was noted that a second, clearly distinct taxon

occurred in the same deposit, but the specimens then at hand did not suffice

for adequate identification or, if new, definition of that taxon. Specimens

subsequently collected, although still not ideal, do suffice for those purposes

and are here described and named as a new genus and species.

The specimens are all from the ‘E’ Quarry in the Varswater Formation in

the vicinity of Langebaanweg, Cape Province. The Varswater Formation is of

very late Miocene and/or early Pliocene age (Hendey 1978). The quarry and

its stratigraphy have been carefully described by Hendey (1976), and those

data need not be repeated here. Hendey has shown that the rich faunas of this

formation represent a number of different environments—marine, estuarine,

fluviatile, and terrestrial. Almost all known fossil penguins have been found

in marine beds but generally in those deposited near shore or in distinctly

littoral conditions. The Varswater penguins also have some marine associates,

but the overall fauna of this varied formation, especially as regards the mammals

except for cetaceans, a seal and an otter, is predominantly nonaquatic.

Most of the specimens here studied come from the middle member of the

Varswater Formation, formerly (Hendey 1974) designated as “Bed 2’ but now

(Hendey 1976) as the Quartzose Sand Member. Some, however, may be either

Ann. S. Afr. Mus. 78 (1), 1979: 1-9, 5 figs, 3 tables.

y ANNALS OF THE SOUTH AFRICAN MUSEUM

from that member or from one of two fossiliferous beds, 3aS and 3aN of Hendey,

in the overlying Pelletal Phosphorite Member (formerly ‘Bed 3’). The field

data provided by Hendey are given below for each specimen listed in the

hypodigm of the new species.

All specimens are in the South African Museum. All bear catalogue

numbers with the prefix (SAM-—PQ). As that prefix is uniform throughout, only

the distinctive locality prefix (L) and numerical part of the catalogue designation

will be given hereafter.

All measurements are in millimetres. In Tables 2 and 3 the following

abbreviations are used:

N number of specimens in sample.

OR _ observed range.

X-_ sample mean and its standard error.

S+ sample estimate of standard deviation and its standard error.

V+ sample estimate of coefficient of variation and its standard error.

Most of the designations and definitions of the dimensions involved in

the tables are among those used in Simpson (1975) as follows:

Humerus:

a. Maximum longitudinal dimension.

b. Width of shaft about one-third of the distance distal to head.

c. Width of shaft about two-thirds of distance distal to head.

d. Distance from radial condyle to longest distal process.

Femur:

a. Distance from the hollow between head and trochanter to that

between distal condyles.

b. Proximal width.

c. Distal width.

Tibiotarsus:

a. Distance from the proximal articulation, excluding the crest, to the

hollow between distal condyles.

6. Distal width.

The fragmentary nature of the three partial tarsometatarsi prevents use of

standard dimensions (a—c in Simpson 1975) for this bone but the following

ad hoc dimension is used:

Tarsometatarsus:

d. Width across distal ends (phalangeal articulations) of metacarpals

IJ-II.

A NEW GENUS OF LATE TERTIARY PENGUIN FROM LANGEBAANWEG 3

SYSTEMATICS

Order SPHENISCIFORMES

Family Spheniscidae

Dege gen. nov.

Etymology

Dege, pronounced approximately day-gay, is an augmentative form of

Swahili ndege, ‘bird’, hence “big bird’. It is hoped that, as with Inguza, this will

be an interesting change from the now rather stereotyped names of fossil

penguins based on classical roots. There are, of course, larger African birds,

but ‘big bird’ is appropriate because the type-species of Dege is larger than that

of Inguza and also larger than the living African penguin, Spheniscus demersus.

There seems to be no simple Swahili word for ‘penguin’ and in that language a

penguin is designated descriptively, for example, as ndege ya nchi za baridi,

approximately ‘cold country bird’. |

Dege is arbitrarily considered masculine in taxonomic usage.

Type-species

Dege hendeyi sp. nov.

Included species

Type only.

Known distribution

Langebaanian latest Miocene/early Pliocene in the Quartzose Sand Member

and perhaps the Pelletal Phosphorite Member of the Varswater Formation near

Langebaanweg.

Diagnosis

Tarsometatarsus short and stout. Medial intermetatarsal foramen present,

relatively proximal and small, plantar opening more radial than in Inguza or

Aptenodytes and Pygoscelis but less medial than in Spheniscus or most other

Recent penguins. Medial calcaneal ridge not entire on known specimens but

apparently weakly developed. Humerus with strongly bipartite tricipital fossa,

dorsal part deeper but smaller, ridge between the two parts of the fossa parallel

to long axis of bone. Latissimus dorsi insertion small, circular, convex. Shaft

slightly bowed, not distinctly sigmoid, slightly wider distally than proximally.

Preaxial angle feeble or absent. Femur and tibiotarsus sphenisciform, without

marked peculiarities. (Figs 1-5.)

Discussion

Apart from size and proportions, the tarsometatarsus is especially distinc-

tive in the position and course of the medial intermetatarsal foramen (the inner

proximal foramen of Zusi 1975). The plantar opening is somewhat broken on

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Dorsal and plantar views of Dege hendeyi holotype (partial

left tarsometatarsus, L28455) from Langebaanweg. (Twice natural

S1Ze.)

specimen L28455, holotype of the type-species, but seems clearly to have been

in a position unlike that in any other penguin known to me. Zusi (1975) and |

others have noted that this opening is on the medial side of a prominent calcaneal |

ridge in most Recent genera, but less medial and below a less prominent ridge |

in Aptenodytes and Pygoscelis. In Dege it is apparently in a position more or |

less intermediate between those two states. It is not known which, if either,

Fig. 2. Dorsal, ventral and posterior views of Dege hendeyi left

humerus (L55005) from Langebaanweg. (Natural size.)

A NEW GENUS OF LATE TERTIARY PENGUIN FROM LANGEBAANWEG 5)

Zoy

Fig. 3. Distal view of Dege

hendeyi left humerus

(L55010) from Langebaan-

weg. (Iwice natural size.)

state is more primitive. The humerus is less distinctive and is within the rather

stereotyped penguin pattern, but in detail is not quite like that of any other

genus in which this part is known.

The only known penguins other than Inguza and Dege that may be of

comparable age are three specimens from Motunau Beach on the South Island

of New Zealand (Marples 1960; Simpson 1972). This Pliocene locality is about

10° farther south and almost 156° farther east than Langebaanweg, but that

alone does not necessarily preclude generic relationships. Some living genera

Fig. 4. Anterior and posterior views of Dege hendeyi left femur

(L28217) from Langebaanweg. (Natural size.)

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Anterior view of Dege hendeyi right tibiotarsus (L5503) from Langebaanweg.

(Natural size.)

have comparably wide ranges. However, the known Langebaanweg and

Motunau fossil penguins are quite different. Known parts of two of the Motunau

penguins are not generically distinguishable from the living genera Pygoscelis

and Aptenodytes and are referred to those genera as extinct species. The humerus

of P. tyreei can be compared with Jnguza and Dege and the tarsometatarsus of

A. ridgeni can be similarly compared as can other parts of living Pygoscelis and

Aptenodytes. The diagnoses of Jnguza and Dege include distinctions from

Pygoscelis and Aptenodytes that are reasonably construed as of generic rank.

The third Motunau genus, Marplesornis, can be compared mainly on the basis

of a not quite perfect humerus, which differs from both /nguza and Dege in

having an extraordinarily expanded and somewhat concave latissimus dorsi

insertion and from Dege in having the shaft distinctly more expanded distally.

Dege hendeyi sp. nov.

Etymology

For Dr Q. B. Hendey, who supervised collecting of these and many other

fossils at Langebaanweg and who has done so much research on the geology

and fauna of that deposit.

Holotype

L28455, left tarsometatarsus with most of metatarsals II and III, lacking

metatarsal IV* and proximal articulation. From Quartzose Sand Member.

Hypodigm

Holotype and the following. From Quartzose Sand Member: L22985,

fragment of left tarsometatarsus with most of distal end of metatarsal III;

128456, fragment of right tarsometatarsus with distal ends of metatarsals III

and IV; L13154, right femur nearly complete; L25028, left femur nearly com-

plete; L25807, right femur nearly complete; L28216, right femur nearly com-

plete; L28217, left femur complete; L28222, left tibiotarsus complete. From

Quartzose Sand Member or bed 3aN of Pelletal Phosphorite Member: L55004,

left humerus with proximal and distal ends imperfect; L55005, left humerus

with distal end imperfect; L55003, right tibiotarsus complete. From Quartzose

* The missing metatarsal IV was found and restored to the holotype after this study was

completed.

A NEW GENUS OF LATE TERTIARY PENGUIN FROM LANGEBAANWEG i/

Sand Member or bed 3aS of Pelletal Phosphorite Member: slightly damaged

proximal end of right humerus; L55010, well-preserved distal end of left

humerus; L3656, left femur with proximal and distal ends slightly abraded.

Known distribution

As for the genus.

Diagnosis

Only known species of the genus. Measurements as in Tables | and 2.

TABLE |

Measurements of specimens of Dege hendeyi.

(Specifications of measurements are given in the text.)

Tarsometatarsus - °- °: ° d

Holotype 23018: - =: -: L270

Humerus a b Cc d

IL SSS 8 oo a6!) 12,8 14,0 —

ILSSOOS 8 8 eR Eos) 13,4 14,4 —

ILSSOI@ © 9 0 6 oe <6 == == == 20,3

Tibiotarsus a b

HESS OOS wed os eel ee 1 O6;5 16,0

ILDBWAD, 8 9S BF sg NS 15,3

TABLE 2

Statistics on femora of Dege hendeyi.

(Specifications of variates and symbols of statistics are given in the text.)

Variate N OR x S Vv

a at * eG 70,4-79,5 PS S4E130 317092 4,22+1,22

De eh eS 17,3-18,6 18,16+0,25 0,56+0,18 3,88 +0,97

Ges et, 5 (5,117.9 16,96-+0,50 1,11-40,35 6,57+2,08

Discussion

Inguza and Dege are distinct genera, not necessarily more closely related

to each other than to some other penguins, but they occur together and sorting

or preliminary identification can most easily be made by the difference in size

of their type-species. Comparable measurements of those species are given in

Table 3.

As noted above, some of the specimens of Dege hendeyi may have come

from the Pelletal Phosphorite Member, which overlies the Quartzose Sand

Member and must be to some degree younger in age. Hendey (1976) has sug-

gested that the differences may be appreciable and that further studies of the

enlarged collection from the Pelletal Phosphorite Member may reveal some

faunal difference, although this has not yet been clearly established. In the

material here studied, it happens that specimen L3656, the only femur that

may be from the Pelletal Phosphorite Member, is the largest of the six femora

referred to Dege hendeyi. The difference is not great enough to warrant any

taxonomic distinction, and as the matter now stands the whole lot could well,

and probably does, represent a single, not even particularly variable species.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 3

Comparative dimensions of Dege hendeyi and Inguza predemersus.

(Specifications of variates and symbols of statistics are given in the text.)

Variate Species N OR xX

Tarsometatarsusd + °- ° Dege hendeyi 1 — 16,5

Inguza predemersus 1 — W257

jalumnens @ 2 9 2 9 9 D. hendeyi y} 68,5—69,0 68,75

I, predemersus 2 57,5-59,0 58,25

DD, Po fore Ba) ene as D. hendeyi D, 12,8-13,4 13,10

I. predemersus 6 8,6-10,0 9,35

Che RES ed cons D. hendeyi 2, 14,0-14,4 14,20

I. predemersus 5 10,0-11,7 10,98

(7 kee Me Melk ei D. hendeyi 1 — 20,30

I. predemersus 4 17,8-19,3 18,60

leGAI G9 8. 8 8) 9 9 D. hendeyi 6 70,4—79,5 153

I. predemersus 3 59,8-62,3 61,20

Da ae Naea ae Cp ae eis D. hendeyi 5 17,3-18,6 18,16

I. predemersus 2 13,9-14,4 14,15

(Nigh ga See Bia S027 Ay D. hendeyi 5 15,1-17,9 16,96

I, predemersus 4 13,3-14,5 13,78

All the penguin bones from this deposit are dissociated, but they fall into

two non-overlapping size groups and the ratios of measurements of non-

homologous bones are similar to those of associated bones in Recent species and

in the few fossil species in which associated bones have been found. It is highly

probable that the known Langebaanweg specimens do represent just two

species and are correctly sorted by size.

ACKNOWLEDGEMENTS

As in previous studies of South African fossil and Recent penguins, I am

greatly indebted to the South African Museum and to Dr Q. B. Hendey who

arranged the loan of the fossil specimens, provided the field data on them, and

reviewed the manuscript of this paper. Study was carried out at the Simroe

Foundation and with support from the Department of Geosciences of the

University of Arizona, Tucson, U.S.A.

The illustrations were supplied by the South African Museum.

REFERENCES

HENDEY, Q. B. 1974. The late Cenozoic Carnivora of the south-western Cape Province.

Ann. S. Afr. Mus. 63: 1-369.

HENDEY, Q. B. 1976. The Pliocene fossil occurrences in ‘E’ Quarry, Langebaanweg, South

Africa. Ann. S. Afr. Mus. 69: 215-247.

HENDEY, Q. B. 1978. The late Tertiary Hyaenidae from Langebaanweg, South Africa, and

their relevance to the phylogeny of the family. Ann. S. Afr. Mus. 76: 265-297.

Marp tes, B. J. 1960. A fossil penguin from the later Tertiary of North Canterbury. Rec.

Canterbury Mus. 7: 185-195.

Simpson, G. G. 1971. Fossil penguin from the late Cenozoic of South Africa. Science 171:

1144-1145.

A NEW GENUS OF LATE TERTIARY PENGUIN FROM LANGEBAANWEG 9

SIMPSON, G. G. 1972. Pliocene penguins from North Canterbury, New Zealand. Rec. Canter-

bury Mus. 9: 159-182.

Simpson, G. G. 1975. Notes on variation in penguins and on fossil penguins from the Pliocene

of Langebaanweg, Cape Province, South Africa. Ann. S. Afr. Mus. 69: 59-72.

Zusi, R. L. 1975. An interpretation of skull structure in penguins.* In: STONEHOUSE, B., ed.

The Biology of Penguins: 59-86. London and Basingstoke: Macmillan.

* This study also refers to postcranial bones and classification of Recent penguins.

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

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

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Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

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

Note punctuation in the above example:

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In describing new Species, one specimen must be designated as the holotype; other speci-

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Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach

Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

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Name of new genus or species is not to be included in the title: it should be included in the

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Biological Abstracts.

GEORGE GAYLORD SIMPSON

A NEW GENUS OF LATE TERTIARY PENGUIN

FROM LANGEBAANWEG, SOUTH AFRICA

VOLUME 78 PART 2 APRIL 1979

OF THE SOUTH AFRICAN

~ MUSEUM

CAPE TOWN

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