VOLUME 104 PART 4 AUGUST 1994 ISSN 0303-2515 _ANNALS OF THE SOUTH AFRICAN | MUee iM . CAPE TOWN INSTRUCTIONS TO AUTHORS 1. MATERIAL should be original and not published elsewhere, in whole or in part. 2. LAYOUT should be as follows: (a) Centred masthead to consist of Title: informative but concise, without abbreviations and not including the names of new genera or species Author’s(s’) name(s) Address(es) of author(s) (institution where work was carried out) Number of illustrations (figures, enumerated maps and tables, in this order) (b) Abstract of not more than 200 words, intelligible to the reader without reference to the text (c) Table of contents giving hierarchy of headings and subheadings (d) Introduction (e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents (f) Summary, if paper is lengthy (g) Acknowledgements (h) References (i) Abbreviations, where these are numerous. 3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with 3 cm margins all round. First lines of paragraphs should be indented. Tables and a list of captions for illustrations should be typed separately, their positions indicated in the text. All pages should be num- bered consecutively. Major headings of the paper are centred capitals; first subheadings are shouldered small capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further subdivisions should be avoided, as also enumeration (never roman numerals) of headings and abbreviations. Footnotes should be avoided unless they are short and essential. Only generic and specific names should be underlined to indicate italics; all other marking up should be left to editor and publisher. 4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18 cm (19 cm including caption); the reduction or enlargement required should be indicated (and preferably uniform); orig- inals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and final size. A metric scale should appear with all illustrations, otherwise magnification or reduction should be given in the caption; if the latter, then the final reduction or enlargement should be taken into consideration. All illustrations, whether line drawings or photographs, should be termed figures (plates are not printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items of composite figures should be designated by capital letters; lettering of figures is not set in type and should be in lower-case letters. If Letraset is used authors are requested to use Helvetica-style letter- ing, if possible. The number of the figure should be lightly marked in pencil on the back of each illustration. 5. REFERENCES cited in text and synonymies should all be included in the list at the end of the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable): (a) Author’s name and year of publication given in text, e.g.: ‘Smith (1969) describes...” “Smith (1969: 36, fig. 16) describes...” ‘As described (Smith 1969a, 1969b; Jones 1971)’ ‘As described (Haughton & Broom 1927) .. .” “As described (Haughton et al. 1927)...” Note: no comma separating name and year Pagination indicated by colon, not p. names of joint authors connected by ampersand et al. in text for more than two joint authors, but names of all authors given in list of references. (b) Full references at the end of the paper, arranged alphabetically by names, chronologically within each name, with suffixes a, b, etc., to the year for more than one paper by the same author in that year, e.g. Smith (1969a, 1969b) and not Smith (1969, 1969a). For books give title in italics, edition, volume number, place of publication, publisher. For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part number in parentheses, pagination (first and last pages of article). Examples (note capitalization and punctuation) Bu.touGu, 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., Duvat, 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. Konn, A. J. 19605. 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. Jn: ScHuLTzE, L. Zoologische und anthro- pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgefiihrt in den Jahren 1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270. (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 are issued in parts at irregular intervals as material becomes available Obtainable from the South African Museum, P. O. Box 61, Cape Town 8000 Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM word uitgegee in dele op ongereelde tye na gelang van die beskikbaarheid van stof Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000 OUT OF PRINT/UIT DRUK I 21235, 5=8), 3124-5, Bet epae). 4 5a ses eo) G(s De-pa.)s 74), 8922) 7), 10=3), 12s) 7 te 14(123), 15G4=5)) 2405); 27) 33), 326), 33" 36(2), 43(1), 45(1), 49(1), 67(5, 11), 84(2) Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 86813 154 7 Printed in South Africa by In Suid-Afrika gedruk deur The Rustica Press (Pty) Ltd, Die Rustica-pers (Edms) Bpk, Old Mill Road, Ndabeni, Cape Old Mill-weg, Ndabeni, Kaap 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. ‘SYJOIUSIU “J “D “AZSEOST ‘ds gsdaz1pod “A “(LLSOI9 WNSN) Sdaoipod ‘d "gq :(Z xX) “MAIA JOIIa\SOd UT IsIvJeJoWOSIe) JYSIY “S77709181U “d “q ‘O8L99r1 ‘ds gsdaoipog “DQ “690081 ‘ds ¢sdarIpod “@ “(8S800S WNSN) Sdaoipod -q ‘vy ‘(¢°] x) MIA JeuOoUe UT LOWNY IYysTY “(POESSS INNSM) iMawins syjoousiu sdaoipog pue sdasipod snqucjipog yim poreduiod Zamuereqasuey wo ‘ds 7 sdao1pog JO S[Issoy ‘7 ‘314 (7) (a4 aay ao D ce oe) Zz Z QO Zz < & < =) 4 Zz E =) ina] a eal \o) ea] Z aa) oO g — Ay : —f aa 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 BRODKORB, P. 1985. Preliminary report on Pliocene/Pleistocene birds of East Africa. Acta XVIII Congressus Internationalis Ornithologici 1: 174-177. HAARHOFF, P. J. 1988. A new fossil stork (Aves, Ciconiidae) from the late Tertiary of Langebaanweg, South Africa. Annals of the South African Museum 97: 297-313. 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, Langebaanweg, South Africa, and a reinterpretation of their geological context. Annals of the South African Museum 84: 1-104. 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 Society of Washington 97: 736-740. 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., etc. An author’s name when cited must follow the name of the taxon without intervening punctuation and not be abbreviated; if the year is added, a comma must separate author’s name and year. The author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is 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 semicolon separates more than one reference by the same author full stop separates references by different authors figures of plates are enclosed in parentheses to distinguish them from text-figures dash, not comma, separates consecutive numbers. Synonymy arrangement according to chronology of bibliographic references, whereby the year is placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable. In describing new species, one specimen must be designated as the holotype; other specimens mentioned in the original description are to be designated paratypes; additional material not regarded as paratypes should be listed separately. The complete data (registration number, depository, descrip- 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. 7. SPECIAL HOUSE RULES Capital initial letters (a) The Figures, Maps and Tables of the paper when referred to in the text e.g. °. . . the Figure depicting C. namacolus ...’: ‘. . . in C. namacolus (Fig. 10)...’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by initials or full names e.g. DuToit but A.L. du Toit; Von Huene but F. von Huene (c) Scientific names, but not their vernacular derivatives e.g. Therocephalia, but therocephalian Punctuation should be loose, omitting all not strictly necessary Reference to the author should preferably be expressed in the third person Roman numerals should be converted to arabic, except when forming part of the title of a book or article, such as ‘Revision of the Crustacea. Part VIII. The Amphipoda.’ Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial capital letter, provided the same generic name is used consecutively. The generic name should not be abbreviated at the beginning of a sentence or paragraph. 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. wilting STORRS L. OLSON EARLY PLIOCENE GREBES, BUTTON-QUAIL, AND KINGFISHERS FROM SOUTH-WESTERN CAPE PROVINCE, SOUTH AFRICA (AVES: PODICIPEDIDAE, TURNICIDAE, HALCYONIDAE)