Annals of the South African Museum = Annale van die Suid-Afrikaanse Museum

VOLUME 104 PART 4 AUGUST 1994 ISSN 0303-2515

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FiscHer, P. H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie
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Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of
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(continued inside back cover)

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

Volume 104 Band
August 1994 Augustus
Part 4 Deel

EARLY PLIOCENE GREBES, BUTTON-QUAIL,
AND KINGFISHERS FROM SOUTH-WESTERN
CAPE PROVINCE, SOUTH AFRICA
(AVES: PODICIPEDIDAE, TURNICIDAE,
HALCYONIDAE)

By
STORRS L. OLSON

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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EARLY PLIOCENE GREBES, BUTTON-QUAIL, AND
KINGFISHERS FROM SOUTH-WESTERN CAPE PROVINCE,
SOUTH AFRICA
(AVES: PODICIPEDIDAE, TURNICIDAE, HALCYONIDAE)

By
STORRS L. OLSON

Department of Vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560*

(With 3 figures and 1 table)

[MS accepted 9 September 1993]

ABSTRACT

Two species of grebes (Podicipedidae) occur sparingly among the abundant fossil
remains of birds found in the early Pliocene Varswater Formation at Langebaanweg, south-
western Cape Province. The most abundant of these is a small species that is indistinguish-
able from the modern dabchick Tachybaptus ruficollis, which ranges from South Africa
through Eurasia, as far as New Guinea. In their smaller size, these fossils agree with the
African subspecies T. r. capensis, as opposed to larger forms in Europe and Asia, and appear
to indicate a 5-million-year period of stasis in this lineage in southern Africa. The second
species of grebe is much rarer, being known from three bones only. These are robust and
superficially most similar to the New World genus Podilymbus, but are sufficiently different
to be tentatively referred to the widespread genus Podiceps. These fossils represent a new,
extinct lineage of grebes for Africa.

A single fossil of a button-quail is most similar to the living species Turnix hottentotta
and provides the first Tertiary occurrence of the family Turnicidae.

Three bones of kingfishers are referable to a species of Halcyon similar in size to
H. albiventris, and to a species of Ceryle about the size of the New World C. alcyon, which
represents a size-class of Ceryle that is now extinct in Africa.

CONTENTS

PAGE
ETUC OUI CEL OTN Re eer a Ieee ee ee moa Rr tata dead ana coe REI TST VL Seal ra 50
SVS(CINALICS ote irae eee nec nackrerke cen Me ce Sua eens Niet telsuaa element ste mates 50
ID SREY Coy ay Bb en a a sates SAS SO CAE a EE i OE aor Ea HRM GEM Gea OOAEE 60
ALCO WICUSEINENILS recente eer eee rime ee tc a ucareiinetelname cies eecee deem et nceutecer 60
FRE TETETI CES tet ea pe ESI Re NERY os NAVA CSE OND S/o LRT Se Ea Rear 60

49

* Research Associate, Percy FitzPatrick Institute of African Ornithology, University of Cape
Town, Rondebosch 7700, South Africa

Ann. S. Afr. Mus. 104 (4), 1994: 49-61, 3 figs, 1 table.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM
INTRODUCTION

The deposits of the early Pliocene Varswater Formation near Langebaan-
weg, Cape Province, have yielded one of the largest Tertiary avifaunas yet dis-
covered (Rich 1980; Hendey 1981a, 1981b, 1982). The elucidation of this vast
amount of material continues slowly, with the avian taxa treated so far including
the Procellariiformes (Olson 1985a), Scopidae (Olson 1984), Plataleidae (Olson
1985c), Ciconiidae (Haarhoff 1988), Rostratulidae (Olson & Eller 1989), and
Coliidae (Rich & Haarhoff 1985).

The three families covered in the present report have little in common other
than the uncertain ordinal affinities of the first two. The grebes (Podicipedidae)
are usually accorded their own order Podicipediformes, placed near the loons
(Gaviiformes), but are probably most closely related to families included in the
Gruiformes (Olson 1985b), whereas the button-quails (Turnicidae), which have
traditionally been placed in the Gruiformes, actually have no close relatives
within that varied assemblage (Olson & Steadman 1981).

Fossils from Langebaanweg are from the collections of the Department of
Cenozoic Palaeontology, South African Museum, Cape Town, and are prefixed
SAM-PQ, which has generally been omitted below.

Comparative material examined

Material of Podicipedidae included skeletons representing all living species
except Podiceps pelzelni of Madagascar (see list in Olson in press). Material of
Tachybaptus ruficollis included the subspecies T. r. ruficollis (1 female from
Germany, USNM); T. r. poggei (5 males and 7 females from China, USNM);
T. r. capensis (1 male from Zimbabwe, USNM; 1 female from South Africa,
USNM;; and an unsexed specimen each from Gabon, FMNH, and Tanganyika,
UMMZ); T. r. philippensis (1 male from Luzon, USNM); and T. r. tricolor
(1 female from Celebes, USNM). Of African Turnicidae, 3 skeletons of Turnix
sylvatica lepurana (USNM 429078, 429079, 430658) and 1 skeleton of T. hot-
tentotta nana (CM 1160) were examined. Representative skeletons of each of
the species of Halcyonidae mentioned were all from the USNM collections.

SYSTEMATICS

Order ‘PODICIPEDIFORMES’
Family Podicipedidae

Only three species of grebes occur in Africa today, all of which are wide-
spread Palaearctic forms. The lack of endemism among African grebes contrasts
rather markedly with South America or even Australia. Because none of the
modern African grebes have differentiated at the specific level within Africa, it
becomes of interest to determine when each of the species may have become
established in the continent. Although the grebes are osteologically more homo-
geneous than reflected by modern generic usage, I have followed the nomen-
clature of Storer (1979).

EARLY PLIOCENE GREBES, BUTTON-QUAIL, AND KINGFISHERS 51

Genus Tachybaptus Reichenbach, 1853
Tachybaptus ruficollis (Pallas, 1764)
Fig. 1

Material

Coracoid: complete right—L53324; complete left—L22417D, LSOOSOE;
right scapular end—L25514BC. Humerus: proximal right—L22755, L50342E,
L56221B; proximal left—L22941, L50069A, L55048, L56221A; distal
right—L42745H, L50582J, L55049, L56222B; distal left—L56222A, L56222C.
Carpometacarpus: left proximal—L17060B, L34956, L34957. Femur: complete
right—L50056A; complete left—L22422K, L50056B, L53322; proximal
left—L42853G; distal right—L50012WV, L53323. Tibiotarsus: right lacking
proximal end—L56220B; left lacking proximal end—L56220A, L61672,
L22450Q; distal right—L23646, L24125N; distal left—L22422AA, L25601LF,
L25602EN, L50048G, L50379F; juv.—L42854G; juv.—L50048J. Tarsometa-
tarsus: complete left—L57854, L50063B; proximal right—L25292E,
L25293EC, L50012AG, L50062E; proximal left—L28423DV, L50577AG,
L50577R; distal right—L50062F1, L50062F2, L50577W;; distal left—L17006F,
L50577AL, L50577N; lacking inner trochlea—L22420I.

The total number of specimens is 56 and, if each site in the mine is con-
sidered separately, the minimum number of individuals involved would be 16.

Measurements
See Table 1.

Stratigraphic provenance

Early Pliocene, Varswater Formation: Quartzose Sand Member (18 speci-
mens, minimum number of individuals 7); Pelletal Phosphorite Member
Bed 3aN (36 specimens, minimum number of individuals 8); Pelletal Phospho-
rite Member Bed 3aS (3 specimens, minimum number of individuals 1).

Remarks

Despite a good sample from Langebaanweg, including highly diagnostic ele-
ments, I was unable to distinguish any differences whatever between the fossils
and the living dabchick Tachybaptus ruficollis. Furthermore, although the com-
parative sample sizes were small in some cases, the measurements of the fossils
are consistently small, and thus appear to conform to the size of the small Afri-
can subspecies T. r. capensis, as opposed to the larger European and Asian
races (Table 1). This implies an extraordinary period of stasis of some 5 million
years (Hendey 1981: 95) within this species’ lineage in southern Africa. It might
also imply that the size differences associated with subspecific differentiation
may have originated millions of years ago in this lineage. On the other hand, it
is equally possible that the smaller size is primitive and that the larger size of
more northern forms was derived subsequent to the early Pliocene. Regardless,
the dabchick seems to have been in South Africa for at least 5 million years,
during which it shows no osteological changes.

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Comparison of fossils from Langebaanweg referred to Tachybaptus ruficollis with modern speci-

mens of the African subspecies T. r. capensis and the Asian subspecies T. r. poggei.

(Measurements following in parentheses are from a single specimen each of T. r. ruficollis,
T. r. philippensis, and T. r. tricolor, respectively.)

Measurement n Range Mean Population
CORACOID
Greatest length 2 22.5-22.7 22.6 T. r. capensis
3 21.0-22.6 21.8 fossils
12 22.6-26.9 24.7: (25.1, 25:25 2351) leer pOeees
HUMERUS
Proximal width 3 8.3-9.0 8.7 T. r. capensis
6 8.2-9.6 8.8 fossils
12 8.9-10.2 9.5 (10.4, 9.7, 9.4) T. r. poggei
Distal width 4 5.2=5.5 533 T. r. capensis
6 4.8-5.4 en fossils
12 5.3-6.1 a (Sts) Slod/5 Ds)) T. r. poggei
CARPOMETACARPUS
Proximal depth ~ 4.6-4.8 4.7 T. r. capensis
3 4.7-4.8 4.7 fossils
12 4.5-5.1 Asi Os OE2 0) T. r. poggei
FEMUR
Length 3 27.7-29.2 28.3 T. r. capensis
4 25.2-27.4 26.5 fossils
12 27.9-31.3 29.6 (31.4, 31.4, 28.6) T. r. poggei
Proximal width 4 6.5-7.3 6.8 T. r. capensis
5 6.3-7.3 6.7 fossils
12 7.0-7.8 TAA Te leee8) T. r. poggei
Distal width 4 6.7-7.4 51 T. r. capensis
of 6.0-7.4 6.6 fossils
11 7.3-8.4 Tey Us Sets Unis) T. r. poggei
TIBIOTARSUS
Distal width 4 5.2-5.7 5.4 T. r. capensis
13 5.0-6.0 S35) fossils
12 5.4-6.2 5.9 (5.8, 6.0, 6.0) T. r. poggei
TARSOMETATARSUS
Length 4 33.1-34.8 33.8 T. r. capensis
2 33.1-32.7 B29. fossils
12 33.1-37.4 35.1 (38.6, 38.7, 35.7) T. r. poggei
Proximal width 4 5.6-6.2 6.0 T. r. capensis
8 5.4-6.0 2 y31/ fossils
12 5.8-7.1 6:51(6:6; 6554655) T. r. poggei
Width trochleae 3, 4 4 4.1-4.8 4.5 T. r. capensis
9 4.2-4.7 4.4 fossils
12 4.4-5.3 4:9'(6:2. 915 522) T. r. poggei

EARLY PLIOCENE GREBES, BUTTON-QUAIL, AND KINGFISHERS

Fig. 1. Fossils of Tachybaptus ruficollis from Langebaanweg compared to a modern
specimen of the same species (USNM 430777—on the right in each pair). A. Humeri in
anconal view (proximal end—L50342E; distal end—L50582J). B. Tibiotarsi in anterior
view (L61672). C. Coracoids in ventral view (L53324). D. Femora in anterior view
(L50056B). E. Tarsometatarsi in posterior view (L57854). All figures x 1.8.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

The dabchick in southern Africa is mainly confined to still or slow-moving
bodies of fresh water, usually with emergent vegetation. It invariably nests
in such situations, but non-breeders may occur rarely in marine environments
on the west coast (Maclean 1985). The greater number of specimens and
individuals in the Pelletal Phosphorite Member (PPM) of the Varswater
Formation is probably a reflection of the fluviatile origins of those deposits, as
opposed to the more estuarine or marine environments of the Quartzose Sand
Member.

Genus Podiceps Latham, 1787
Podiceps? sp.
Fig. 2

Material

Humerus: complete right—L50069; distal left—L46678C. Tarsometatarsus:
distal right—L50352E.

The total number of specimens is three and the minimum number of
individuals is one.

Measurements (in mm)

Humerus: length, 72.2; proximal width, 12.3; distal width, 7.3, 7.8.
Tarsometatarsus: distal width through inner and outer trochleae, 5.9.

Stratigraphic provenance

Early Pliocene, Varswater Formation: all three specimens are from the
Pelletal Phosphorite Member Bed 3aN.

Remarks

These three bones are from a grebe quite unlike any hitherto known from
Africa, being much too small for Podiceps cristatus, and larger and much more
robust than either P. nigricollis or Tachybaptus ruficollis. Brodkorb (1985)
reported fossils of a medium-sized grebe from the Upper Pliocene and Lower
Pleistocene Beds I and II of Olduvai, Tanzania. I compared Brodkorb’s material
directly with the fossils from Langebaanweg and found that two very different
species were represented, with the Olduvai bird being considerably smaller and
more gracile, and thus more like P. nigricollis.

The length of the single complete humerus from Langebaanweg is near the
maximum for P. nigricollis or the minimum for P. auritus, and falls well within
the range of the New World pied-billed grebe Podilymbus podiceps (see Storer
1976). The relative robustness of the shaft is greater than in any of those species
and the width of the distal end of the other humeral specimen exceeds that of
any of the 38 specimens of P. podiceps measured by Storer (1976). In the
complete specimen, the scar for M. scapulohumeralis anterior is very deep,
although this tends to be somewhat variable individually in modern grebes.

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56 ANNALS OF THE SOUTH AFRICAN MUSEUM

The tarsometatarsus of the fossil is much more robust than in modern
species of Podiceps of comparable size and is superficially similar to Podi-
lymbus. It differs from Podilymbus in having trochlea II situated farther distally,
and trochlea IV extending distally to about the same extent as trochlea III, rather
than being more proximal as in Podilymbus. The fossil is evidently less special-
ized than modern species of Podiceps of roughly comparable size (e.g.
P. auritus or P. nigricollis) in having the shaft less laterally compressed and
trochlea II less retracted proximally and medially, but is not too unlike certain
smaller taxa such as P. occipitalis. It differs from any of these species in having
the distal foramen smaller.

The general resemblance of these fossils is closest to that of Podilymbus, a
genus that is altogether unknown in the Old World. On the other hand, the
differences between Podilymbus and Podiceps, in the skeletal elements rep-
resented as fossils, are not particularly trenchant, so that given the age of the
fossils one could not be certain that these three bones are not from a very
robust, perhaps highly localized endemic form of Podiceps with a circumscribed
range, such as Podiceps gallardoi of Southern Argentina is today. Another
possibility is that the rarity of this species at Langebaanweg may indicate that it
was only a casual wanderer to the area.

Whatever the generic relationships or the previous distribution of this
enigmatic grebe may have been, it definitely represents a lineage of Podiciped-
idae hitherto unknown in Africa. We can only hope that more diagnostic
material, such as a femur—which in Podilymbus is much longer and more
gracile than in other grebes—will surface and help to clear up taxonomic
uncertainties.

Order ‘GRUIFORMES’
Family Turnicidae

Genus Turnix Bonnaterre, 1791

Turnix cf. T. hottentotta Temminck, 1815
Fig. 3B

Material

Humerus: distal right—L43366A.
The total number of specimens and minimum number of individuals is one.

Measurements (in mm)
Humerus: distal width, 4.0.

Stratigraphic provenance

Early Pliocene, Varswater Formation: Pelletal Phosphorite Member
Bed 3aS or possibly the Quartzose Sand Member (Dump 7).

EARLY PLIOCENE GREBES, BUTTON-QUAIL, AND KINGFISHERS a)

Remarks

This specimen is one that I encountered among some unsorted avian
material, the family Turnicidae not having previously been noted at Langebaan-
weg (Rich 1980; Hendey 19815). The skeleton in the Turnicidae presents a
mosaic of unique characters and those of several other orders. It is not unlikely,
therefore, that more material of Turnix is included among the fossils referred to
the Rallidae, Phasianidae, or even Charadriiformes. The apparent scarcity of
button-quails at Langebaanweg is thus to some extent probably an artefact.

The fossil was compared with the two African species of Turnix but, as only
one specimen was available anywhere for T. hottentotta, the results are incon-
clusive. The bone from Langebaanweg seems to be more like 7. hottentotta in
having the ectepicondylar process more robust and laterally (as opposed to
proximally) oriented. It differs from either of the modern species in having the
proximal end of the radial condyle sharply demarcated from the shaft.

Turnix hottentotta is the only button-quail in the western Cape Province
today. It prefers short, moist grasslands, in contrast to T. sylvatica, which
occurs in ranker grasses (Snow 1978). If the fossil is correctly associated with
the 7. hottentotta lineage, it would accord well with the inferred environmental
conditions at Langebaanweg in the early Pliocene (Hendey 19815; Olson
1985c).

Order CORACIIFORMES

In her preliminary account, Rich (1980) indicated the presence of the order
Coraciiformes at Langebaanweg. Based on her identifications, Hendey (1981b)
listed one species of Coraciiformes of undetermined family and at least two
species of kingfisher (Alcedinidae = Halcyonidae), neither identified to genus.
The amount of coraciiform material in the collections from Langebaanweg is
very scant and, in looking over the few fossils that had previously been assigned
to this group, I found that the majority had been misidentified to order. When
the bones of falcons, doves, and parrots had been removed, only three speci-
mens remained. These represent two species in two genera of kingfishers, no
other taxa of Coraciiformes (or Bucerotiformes) as yet being indicated. The
material at present is insufficient for the characterization of new species,
although it is likely that more fossils of kingfishers will become available with
further sorting of the Langebaanweg collections.

Family Halcyonidae
Genus Halcyon Swainson, 1821
Halcyon sp.
Fig. 3D
Material

Tibiotarsus: distal right—L24593H.
The total number of specimens and minimum number of individuals is one.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Measurements (mm)
Tibiotarsus: distal width, 3.2.

Stratigraphic provenance
Early Pliocene, Varswater Formation: Quartzose Sand Member.

Fig. 3. Button-quail and kingfishers from Langebaanweg. A-C. Distal ends
of right humeri of Turnix in palmar view (x 3). A. Turnix hottentotta nana
(CM 1160). B. Fossil Turnix cf. T. hottentotta (L43366A). C. Turnix
sylvatica lepurana (USNM 429079). D-F. Leg elements of kingfishers
(x 4). D. Distal end of right tibiotarsus of Halcyon sp. (L24593H), anterior
view. E. Distal end of left tibiotarsus of Ceryle sp. (L24001JR), anterior
view. F. Distal end of right tarsometatarsus of Ceryle sp. (L24000FL),
posterior view.

EARLY PLIOCENE GREBES, BUTTON-QUAIL, AND KINGFISHERS 59

Remarks

This specimen is referable to Halcyon, as contrasted to Ceryle, by the con-
stricted shaft proximal to the condyles, especially noticeable on the posterior
face; also the markedly narrower medial condyle and wider tendinal bridge. In
Alcedo, the lateral condyle is much narrower and the tendinal bridge is smaller
than in the fossil. The specimen is from a species about the size of H. albiven-
tris. The living African species of Halcyon are, for the most part, inhabitants of
open savanna and bushlands, but occur in nearly every ecotype other than
desert. No species of Halcyon occurs in the Langebaanweg area today, the
nearest approach being made by H. albiventris, which extends along the south-
eastern coast of South Africa to the vicinity of Cape Town.

Genus Ceryle Boie, 1828
Ceryle sp.
Fig. 3E-F
Material

Tibiotarsus: distal left—L24001JR. Tarsometatarsus: right, lacking proximal
end and inner and outer trochleae—L24000FL.

The total number of specimens is two and the minimum number of indivi-
duals is one.

Measurements (mm)

Tibiotarsus: distal width, 3.8. Tarsometatarsus: no meaningful measure-
ments possible.

Stratigraphic provenance
Early Pliocene, Varswater Formation: Quartzose Sand Member.

Remarks

The tarsometatarsus in the ceryline kingfishers differs greatly from that in
Halcyon in being markedly shorter and stouter. The fossil tarsometatarsus from
Langebaanweg agrees with Ceryle in these respects but differs from any of the
species examined in having the scar for the hallux more deeply excavated,
creating much more of an indentation in the medial margin of the bone. Also,
the distal foramen is more proximally situated.

The tibiotarsus is from a species much larger than the preceding species of
Halcyon, and possesses the characters of Ceryle as outlined above, although it
has a wider tendinal bridge than in living forms of that genus. It would appear
to be from a species similar in size to that represented by the tarsometatarsus,
and both are tentatively referred to the same species.

This kingfisher was considerably larger than C. rudis but much smaller than
C. maxima, and would more closely have approximated the size of the New
World species C. alcyon or Chloroceryle amazona. The proportions of the
tarsometatarsus are intermediate between the very short, stout bone of Ceryle

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

alcyon and the more elongate tarsometatarsus in Chloroceryle amazona. The
two fossils from Langebaanweg are clearly from an undescribed, extinct species
of Ceryle belonging to a size-class that is now absent from Africa. The living
species of Ceryle are all piscivorous and hence are almost invariably found near
water. Both of the extant species in South Africa (C. rudis and C. maxima) also
frequent the seashore and coastal lagoons, in addition to freshwater streams and
lakes.

DISCUSSION

The inferred habitat requirements of the grebes, button-quails and king-
fishers from Langebaanweg are in agreement with environmental reconstruc-
tions based on a variety of data, including other taxa of birds, and indicate a
stream in rather open savanna. As with other groups of birds, the present
selection consists of species that are almost certainly on a direct line to existing
species, aS well as species belonging to lineages that have become entirely
extinct. The Tachybaptus at Langebaanweg, which is represented by a consider-
able number of specimens, cannot be distinguished from the existing species
T. ruficollis. The button-quail (Turnix) and one of the kingfishers (Halcyon) are
too poorly known to be certain of their specific relationships but are not very
different from living taxa, whereas the remaining grebe (Podiceps) and king-
fisher (Ceryle) belong to size-classes within their respective genera that have
vanished from Africa since the Pliocene.

ACKNOWLEDGEMENTS

I continue to be indebted to Q. Brett Hendey and Philippa Haarhoff for my
introduction to the Langebaanweg avifauna and to the South African Museum
for access to the fossil material. Comparative material was from the collections
of the Field Museum of Natural History, Chicago (FMNH); Royal Ontario
Museum, Toronto (ROM); Museum of Vertebrate Zoology, University of Cali-
fornia, Berkeley (MVZ); University of Michigan Museum of Zoology, Ann
Arbor (UMMZ); and National Museum of Natural History, Smithsonian Institu-
tion, Washington (USNM). I thank D. Scott Wood and Diana Matthiesen for
making available a skeleton of Turnix hottentotta nana from the Carnegie
Museum of Natural History, Pittsburgh (CM). For comments on the manuscript
I thank R. Brooke, T. Crowe, Q. B. Hendey, and C. Mourer-Chauviré.
P. Haarhoff also provided comments and the measurements of the kingfisher
fossils. The photographs are by Victor E. Krantz, Smithsonian Institution.

REFERENCES

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HAARHOFF, P. J. 1988. A new fossil stork (Aves, Ciconiidae) from the late Tertiary of
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EARLY PLIOCENE GREBES, BUTTON-QUAIL, AND. KINGFISHERS 61

HENDEY, Q. B. 1981la. Geological succession at Langebaanweg, Cape Province, and global
events of the late Tertiary. South African Journal of Science 77: 33-38.

HENDEY, Q. B. 19815. Palaeoecology of the late Tertiary fossil occurrences in ‘E’ Quarry,
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HENDEY, Q. B. 1982. Langebaanweg. A record of past life. Cape Town: South African
Museum.

MacLean, G. L. 1985. Roberts’ birds of southern Africa. Cape Town: John Voelcker Bird
Book Fund.

Otson, S. L. 1984. A hamerkop from the Early Pliocene of South Africa (Aves: Scopidae)
from Langebaanweg, southwestern Cape Province. Proceedings of the Biological
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Otson, S. L. 1985a. Early Pliocene Procellariiformes (Aves) from Langebaanweg,
southwestern Cape Province, South Africa. Annals of the South African Museum 95:
123-145.

OLson, S. L. 19855 The fossil record of birds. Jn: FARNER, D., KING, J., & PARKES, K. C.
eds. Avian Biology 8. New York: Academic Press.

Otson, S. L. 1985c. Early Pliocene ibises (Aves, Plataleidae) from south-western Cape
Province, South Africa. Annals of the South African Museum 97: 57-69.

OLson, S. L. In press. Redescription of ‘Thiornis’ sociata Navas, a nearly complete Mio-
cene grebe from Spain (Aves: Podicipedidae). Courier Forschungsinstitut Senckenberg.

OLson, S. L. & ELLER, K. G. 1989. A new species of painted snipe (Charadriiformes:
Rostratulidae) from the Early Pliocene at Langebaanweg, southwestern Cape Province,
South Africa. Ostrich 60: 118-121.

OLson, S. L. & STEADMAN, D. W. 1981. The relationships of the Pedionomidae (Aves:
Charadriiformes). Smithsonian Contributions to Zoology 337: 1-25.

Rico, P. V. 1980. Preliminary report on the fossil avian remains from late Tertiary sedi-
ments at Langebaanweg (Cape Province), South Africa. South African Journal of
Science 76: 166-170.

Rico, P. V. & HAARHOFF, P. J. 1985. Early Pliocene Coliidae (Aves, Coliiformes) from
Langebaanweg, South Africa. Ostrich 97: 20-41.

Snow, D. W. ed. 1978. An atlas of speciation in African non-passerine birds. London:
British Museum (Natural History).

STORER, R. W. 1976. The Pleistocene pied-billed grebes (Aves: Podicipedidae).
Smithsonian Contributions to Paleobiology 27: 147-153.

STORER, R. W. 1979. Order Podicipediformes. In: Mayr, E. & COTTRELL, G. W. eds.
Check-list of birds of the world 1: 140-155. 2nd ed. Cambridge, Mass.: Museum of
Comparative Zoology.

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.,
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An author’s name when cited must follow the name of the taxon without intervening punctuation
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Synonymy arrangement should be according to chronology of names, i.e. all published scientific
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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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figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is
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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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wilting

STORRS L. OLSON

EARLY PLIOCENE GREBES, BUTTON-QUAIL,
AND KINGFISHERS FROM SOUTH-WESTERN
CAPE PROVINCE, SOUTH AFRICA

(AVES: PODICIPEDIDAE, TURNICIDAE,
HALCYONIDAE)