7 ; b * VOLUME 103 PART 4 JUNE 1993 ISSN 0303-2515 H OF THE SOUTH AFRICAN MUSEUM 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. 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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. 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- pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgeftihrt 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 103 +#£2Band June 1993 Junie Part 4 Deel LATEST PLIOCENE MOUSEBIRDS (AVES, COLITIDAE) FROM OLDUVAI GORGE, TANZANIA By PHILIPPA J. HAARHOFF 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 1 2(=315=8); 322) 4-518 at. pis) ole Sacse 729) 6(l tepals T(=4)) 8.92.7). 10G1=3)) (LD Satpal): 14(1-3), 15(4-5), 24(2, 5), 27, 31(1-3), 32(5), 33, 36(2), 43(1), 45(1), 67(5), 84(2) Copyright enquiries to the South African Museum Kopieregnavrae aan die Suid-Afrikaanse Museum ISBN 0 86813 146 6 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 D2221 LATEST PLIOCENE MOUSEBIRDS (AVES, COLIIDAE) FROM OLDUVAI GORGE, TANZANIA By PHILIPPA J. HAARHOFF South African Museum, Department of Cenozoic Palaeontology, Cape Town (With 8 figures and 4 tables) [MS accepted 26 October 1992] ABSTRACT Two mousebird species, Colius cf. C. striatus and Urocolius sp., have been recognized amongst the fossil bird remains of Olduvai. This is the first fossil record of the genus Urocolius. The only other African fossil record of the order Coliiformes is Colius hendeyi from Langebaanweg, south-western Cape. Several other fossil species are known from Europe. The extant Coliiformes now occur only in Africa. CONTENTS PAGE | Ur VHiofe LYS COT Tha AR AUR ca cE ARO TEE NSSAe a ENR Oe tre inet Rape ae RET A ACORN Une 191 Comparativomateniales meson ecco see Se teers heie i ees ana he epee ea ane 193 | DYSSErTTOLHG Sil: tals See ats A SIR mr amet eet ati ae ae ane MPAN RCNA My nan. Wine eee 194 Biome tricalramalysise scissor ors ds, sesh Ve heres re RUE ete 206 IAC Eel S yer etre repens set cyes cise kai cet arr siiet ce rors ou ouah ont oa sine isthe MLE 206 RE SUES epetertere a ewe Meee ea ie eh leo c Sled Rel Saicth lec UNMNRR nhc ae U oR ee areas 206 | DSS TISS (0) 1 Sele ito Acie cane CAEN eRe ene pea A ea RCE Cer nee Rea 208 PNCKMOWICAOCMENtSemrey cc cieiaiset te it cistern le ateade ede eye Rec aac te octane elie anbcane 210 FRELCECHCES err cre ye ce cves cnet repre oneaes Meets Potent RMR cay ROR TO 211 INTRODUCTION The mousebird remains from Olduvai Gorge in Tanzania form a minor part of the entire avian assemblage from that area. Out of a total of some 30 000 bird bones belonging to about 46 different taxa (D. Matthiesen pers. comm.), only 57 bones have been assigned to the family Coliidae. These are none the less important because the fossil record of this family is poor. The only other fossil mousebird recorded from Africa is the extinct species Colius hendeyi Rich & Haarhoff, 1985, from the Early Pliocene site of Langebaanweg in South Africa. Two extinct species occurred in Europe during the Late Eocene and at least a further six species (requiring revision) in the Miocene (Ballmann 1969; Olson 1985; Mourer-Chauviré 1988) (Table 1). Today, four species of mousebirds in the genus Colius and two species in the genus Urocolius occur in sub-Saharan Africa (Schifter 1985; Fry et al. 1988). Compari- sons between the fossils and five of the six living species (skeletons of Colius castanotus were unobtainable) indicate that most of the Olduvai specimens are readily distinguishable from the genus Urocolius. Morphological differences at the species level were more difficult to ascertain. However, it appears that all but three of the specimens belong to a single species that is closely related to the extant speckled mousebird, Colius striatus, which inhabits the vicinity of Olduvai Gorge today, along 1911 Ann. S. Afr. Mus. 103 (4), 1993: 191-211, 8 figs, 4 tables. ANNALS OF THE SOUTH AFRICAN MUSEUM 192 Idapuay snijod SNIIPUL SNYOIOAN SNIDLIS SNYOD snyoo snyod VOIddV NYAHLNOS 2 TVALNAO AOUNL SNYOIOW Ld cee! 10318 SNOIOWLd aLV'I ANHOOF LIDIYIAD SHYOT SNULIGOSUOD SHYOD ATYVA snjooipnyod snyod ae echt ANHOOIN siajsnjod “dQ “49 snyogd sae : sLusnjpod snyog ds snijop STACI suusnjod “dQ “Jo snyog ALVT ATUAVA ANHAOOI Id ‘ds snyoo0iQ rete SNIDLUS “DQ “JO SNYOD SNAHOAIDU SHTOIOL() SNIDIAIS SNIOD JINOLSTH snjoydado0ona] snyjogd VOINAV LSVaA HONVaA ANVW&H9D ‘POUIWILXS SPIIQasNOU! SUIAT] PUR [ISSOJ JO UOTNQIIsSIP pur ady 1] T1adVL PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE 193 with the white-headed mousebird, C. leucocephalus, and the blue-naped mousebird, Urocolius macrourus. One proximal end of an ulna, OLD FLK NI 19721, one distal end of an ulna, OLD FLK NI 19765, and the incomplete sternal end of one coracoid, OLD FLK NI 26010, are somewhat larger than the other fossil specimens and are morphologically more similar to Urocolius, the genus to which they have been refer- red in this report. With such a small sample size, the species determination is uncertain. | The most commonly preserved elements for the genus Colius are the ulna, tarso- metatarsus, femur and humerus. Bones that are almost complete include one humerus, two ulnae, three carpometacarpi, one phalanx I of digit II and one femur. Reports on other fossil birds from Olduvai Gorge include those of Brodkorb & Mourer-Chauviré (1982, 1984a, 1984b) and of Harrison & Walker (1976, 1979). The specimen numbers are catalogue numbers of the Olduvai fossil birds, and the data are kept at the Department of Zoology, University of Florida, Gainesville. The fossils belong to the Tanzanian Ministry of Antiquities, Dar es Salaam. They were collected by Mary Leakey in 1960-1962. The following abbreviations are used in this paper: FLK Frida Leakey Korongo (Leakey 1965) L Langebaanweg M University of Miami NMB_ National Museum, Bloemfontein OLD = Olduvai PB Osteological collection of Pierce Brodkorb SAM South African Museum YPM _ Peabody Museum, Yale University Anatomical abbreviations are: (e complete d distal l left p proximal I right COMPARATIVE MATERIAL Recent Colius leucocephalus: 1 unsexed; C. colius: 10 males, 5 females, 6 unsexed; C. striatus: 15 males, 11 females, 30 unsexed; Urocolius macrourus: 3 males, 3 females, 1 unsexed; U. indicus: 11 males, 11 females, 11 unsexed. Fossil Colius hendeyi Rich & Haarhoff, 1985. The entire assemblage of 124 bones from Langebaanweg, South Africa, was available. Data for the other described fossil mouse- birds including Colius paludicolus, C. consobrinus, C. archiaci, C. palustris, Colius cf. C. palustris, Colius sp., Primocolius sigei and P. minor were obtained from the litera- ture (Milne Edwards 1871; Ballmann 1969; Brodkorb 1971; Olson 1985; Mourer- Chauviré 1988). 194 ANNALS OF THE SOUTH AFRICAN MUSEUM DESCRIPTION Order COLIIFORMES Murie, 1872 Family Coliidae Swainson, 1837 Genus Colius Brisson, 1760 For ordinal, family and generic diagnoses, see Ballmann (1969) and Rich & Haar- hoff (1985). For additional differences between the genera Urocolius Bonaparte, 1854, and Colius, see Table 2. Species of Urocolius generally have more strongly sculptured features than species of Colius in all the elements examined. The fossils from Olduvai were also compared with, and found to be different from, the extinct genus Primocolius Mourer-Chauviré, 1988, from the Upper Eocene Phosphorites du Quercy in France. Colius cf. C. striatus Figs 1A-B, K, 2A, E, J, 3A, F, G, 4A, F, G Material Olduvai. Coracoids: FLK NI 12567 (Ip with some shaft); FLK NNI 2007 (rp with shaft). Humeri: FLK NI 10195 (Ic deltoid crest slightly damaged); FLK NI 29627 (Ip broken at base of bicipital crest); FLK NNI 3136 (Id); FLK NI 12875 (ap); FLK NI 12401 (rp); FLK NI 28439 (rp small fragment with head and bicipital crest missing); FLK NI 27534 (rd); FLK NNI 2482 (rd with most of shaft); FLK I 3643 (rd). Ulnae: FLK NI 18301: (cc); FLK_NI 27330 (ac); FLK NNI 20669" ap); FLK NI 19721 (Ip); FLK NI 29531 (Ip, olecranon damaged); FLK NI 24547 (Ip, ole- cranon missing); FLK NI 22756 (Id); FLK NI 25063 (ld); FLK NI 28923 (ld); FLK NI 18295 (rp, olecranon missing); FLK NI 22660 (rp); FLK NI 29532 (tp); FLK NI 13895, FLK I 5003 (rp, olecranon damaged); FLK I 5004 (rp, lacking olecra- non); FLK NI 18269 (rp slightly damaged); FLK NI 19766 (rd); FLK NI 19765 (rd); FLK NI 13895 (rd internal condyle damaged). Radius: FLK NI 21182 (Ip). Carpometacarpi: FLK NI 10373 (1 lacking metacarpal III); FLK NI-7985 (1 lacking metacarpal III and posterior carpal trochlea); FLK NNI 15514 (Id incom- plete); FLK NI 8031 (r lacking metacarpal III, metacarpal I and posterior carpal trochlea). Phalanx I of digit Il: FLK NNI 15886 (rc). Femora: FLK NNI 15356 (rc); FLK NI 18520 (rp); FLK NI 29596 (rp); FLK NI 19949 (rd); FLK NI 19950 (rd); FLK NI 21843 (rd); FLK NI 19973 (Ip); FLK NI 23510 (Ip). Tibiotarsi: FLK I 4906 (Ip rotular crest damaged); FLK NI 18432 (rp rotular crest and inner cnemial crest damaged); FLK NNI 16690 (ld); FLK NI 1356 (rd); FLK NNI 2035 (rd external condyle missing). Tarsometatarsi: FLK NI 12867 (Ip); FLK NI 18425 (Ip hypotarsus damaged); FLK NI 1128 (Ip hypotarsus damaged); FLK NI 7421 (Id); FLK NI 1034 (ld); FLK I 11274 (rd); FLK NI 12866 (rd); FLK NI 7499 (rd internal trochlea missing). PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE TABLE 2 Osteological differences between Colius and Urocolius additional to Ballmann (1969) and Rich & ELEMENT CORACOID ULNA RADIUS CARPOMETACARPUS PHALANX I of DIGIT II Haarhoff (1985). COLIUS Coracohumeral surface and furcular facet relatively compressed and slightly sloping Furcular facet circular Sternal end slightly expanded Sternal facet regularly shaped and relatively broad Shaft relatively straight Scapular facet relatively small No scar present on surface external to humero-ulnar depression Olecranon relatively rounded Fossa under external cotyla, palmar view, absent or poorly developed External condyle rounded at base of shaft, internal view Carpal tuberosity relatively rounded Internal condyle relatively small in relation to external condyle Ridges on either side of distal tendinal groove not very pronounced Metacarpal I relatively short and not very recurved proximally Metacarpal facet rounded in shape Anterior internal edge not noticeably flattened on to shaft FEMUR Relatively robust Internal condyle relatively expanded Angle between head and trochanter cn anterior side relatively wide UROCOLIUS Coracohumeral surface and furcular facet erect and elongated Furcular facet more linear Sternal end greatly expanded Sternal facet irregularly shaped and narrow Shaft more curved Scapular facet relatively large Scar present on surface external to humero-ulnar depression Olecranon relatively pointed Fossa under external cotyla, palmar view, generally well developed External condyle tapers to a point at base of shaft, internal view Carpal tuberosity relatively pointed Internal condyle relatively large in relation to external condyle Ridges on either side of distal tendinal groove are more pronounced Metacarpal I elongated and pointed proximally Metacarpal facet horseshoe-shaped Anterior internal edge flattened on to shaft Relatively gracile Internal condyle not as expanded Angle between head and trochanter on anterior side more acute 195 196 ANNALS OF THE SOUTH AFRICAN MUSEUM y Fig. 1. A-—J. Humeri. - A, B. Colius cf. C. striatus, OLD FLK NI 10195. C,D. C. striatus, PB25226. E, F. C. colius, SAM-ZOT.26. G,H. C. leucocephalus, YPMS5797. I, J. C. hendeyi, L24001 IF. x3. K-N. PhalanxI of digit II. K.Colius cf. C. striatus, OLD FLK NNI 15886. L. C. striatus, PB19300. M. C. colius, SAM-—ZOT.26. N. C. leucocephalus, YPM5797. x4. PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE 197 Age and distribution Late Pliocene, about 1,72 to 1,83 m.y. (Curtis & Hay 1972; Hag et al. 1977; Savage & Russell 1983; Kappelman 1986). This species is known only from sites FLK, FLK NN and FLK N in Bed I at Olduvai Gorge, north-western Tanzania. Dating methods used include potassium-argon and geomagnetic time scales. The accuracy of the dates given have an average co-efficient of variation of between one and two per cent (Curtis & Hay 1972). A Plio—Pleistocene boundary of 1,64 m.y. is taken from Harland ef al. (1990). Measurements See Rich & Haarhoff (1985) and Table 3 for measurements of species of Colius. Skeletons of Colius castanotus were unavailable for comparison. Description The following description differentiates the Olduvai material, here assigned to Colius cf. C. striatus, from the extant species C. colius and C. leucocephalus, and from the extinct species C. hendeyi, C. paludicolus, C. consobrinus, C. archiaci, C. palustris and Colius cf. C. palustris. Most preserved elements show the characteristic features of the species Colius striatus and most are within the size range of that species. Coracoid (Fig. 2J). (1) Area between furcular facet and glenoid facet, in internal view, is wide, not narrow; (2) coracohumeral surface not medially constricted; (3) coracohumeral surface rises gradually from glenoid facet in external view; (4) shaft robust; (5) external margin of dorsal surface between glenoid facet and coracohumeral surface deeply indented, forming acute angle. Characters 1—4 separate Colius striatus (Fig. 2K) and Colius cf. C. striatus (Fig. 2J) from C. colius (Fig. 2L) and C. leuco- cephalus (Fig. 2M); character 5 is unique to Colius cf. C. striatus. The coracoid is unknown for other fossil species. Humerus (Fig. 1A, B). (1) Relatively robust proximal end; (2) head globular in anconal view; (3) deltoid crest curves relatively abruptly palmarly in anconal view; (4) median crest slightly notched; (5) entepicondyle rounded in anconal view and does not project beyond internal condyle; (6) internal condyle well rounded in palmar view and not obviously directed toward external condyle. Characters 1-3 separate Colius striatus (Fig. 1C,D) and Colius cf. C. striatus (Fig. 1A,B) from C. colius (Fig. 1E, F), C. leucocephalus (Fig. 1G, H), C. hendeyi (Fig. 11, J) and C. paludico- lus. Character 4 separates C. striatus (Fig. 1C,D) and Colius cf. C. striatus (Fig. 1A,B) from C. hendeyi (Fig. 11, J), C. leucocephalus (Fig. 1G,H) and C. colius (Fig. 1E, F). Character 5 separates Colius striatus (Fig. 1C, D) and Colius cf. C. striatus (Fig.1A,B) from C.colius (Fig.1E,F), C. leucocephalus (Fig. 1G, H), C. hendeyi (Fig. 11, J), Colius cf. C. palustris and C. paludicolus. Character 6 separates C. striatus (Fig. 1C, D) and Colius cf. C. striatus (Fig. 1A, B) from C. colius (Fig. 1E,F), C. leucocephalus (Fig.1G,H) and_ C. hendeyi (Fig. 11, J). Ulna (Fig. 3A). (1) Proximal end, robust in internal view; (2) carpal tuberosity erect in distal view, not orientated over internal condyle. Both characters separate 198 ANNALS OF THE SOUTH AFRICAN MUSEUM TABLE 3 Measurements (mm) of living and fossil species of Colius. C. leucocephalus C. colius C. striatus Colius cf. C. striatus mean mean n range mean n range mean n range Measure- ment no. CORACOID Vena’ a) oom vo NANN ANTM RUS HUME ~~, — ae ES o ~~, mer ATTN N Seni seeniihaeen ithe een ihe! m= AN xt OO NADONe Sener FONSTEISS mom tN < EASPEN ON AnaAN —_— Stag Saag i CDIDIG IS on —N N n = — 1,8 Levy 1,92 155=159 211 53 1,5-2,1 Se ih eee ee | CO = CA GD) 13,1-13,2 2 3 —Notror ZHNtO AN FGHnOt =) < = =) [a4 Oo Pal Diall =—aAD FEMUR SESRENSREN eoranann AQ ee 20 20,24 4,29 2,83 4,03 2.93 21 21 21 21 oroort ~MnA—O = TNO nN | 5 4 Ay (al 4 =ANMOrM PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE COCONC SCAICAICA moon Om wt © wt aOANtrets Von eon ana) Wot + als oe oe) Foeliced Reet nna FOMN ™ OM ™\ CO ANN S eniihameniihonen tian! mnwoeo ANN CO aaa ANN st NMOMr~ ENMOr faa) = a te minimum shaft width; maximum head depth; 9 = maximum middle maximum proximal depth; 4 maximum distal depth; 8 maximum proximal width; 3 maximum distal width; 7 depth; CMC = carpometacarpus; PuI Dicll = phalanx I of digit I]; TIB = tibiotarsus; TMT = tarsometatarsus. greatest length; 2 minimum shaft depth; 6 I Measurements are: 5 199 200 ANNALS OF THE SOUTH AFRICAN MUSEUM f ¢ + z Wa Fig. 2. A-D. Radii. A. Colius cf. C. striatus, OLD FLK NI 21182. B.C. striatus, PB36207. C. C. colius, SAM-—ZOT.26. D. C. leucocephalus, YPM5797. x 4. E-I. Carpometacarpi. E. Colius ef. C. striatus, OLD FLK NI 10373. F.C. striatus, PB36209. G.C. colius, SAM-—ZO57160. H. C. hendeyi, L20733. I. C. leucocephalus, YPM5797 (transposed). x4. J—M. Coracoids. J. Colius — cf. C. striatus, OLD FLK NNI 2007. K. C. striatus, PB19300. L. C. colius, NMB03275. M. C. leucocephalus, YPMS797. x3. PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE 201 Fig. 3. A-E. Ulnae. A. Colius cf. C. striatus, OLD FLK NI 27330. B. C. striatus, PB36207. C. C. colius, SAM-ZOT.26. D.C. leucocephalus, YPM5797. EE. C. hendeyi, 123163. x3. F-K. Tibiotarsi. F. Colius cf. C. striatus, OLD FLK I 4906. G. Colius cf. C. striatus, OLD FLK NNI 16690. H. C. striatus, PB19300. I. C. colius, SAM-ZO57160. J. C. leucocephalus, YPM5/97. “KC. hendeyi,. 117139" “<3: 202 ANNALS OF THE SOUTH AFRICAN MUSEUM C. striatus (Fig. 3B) and Colius cf. C. striatus (Fig. 3A) from C. colius (Fig. 3C), C. leucocephalus (Fig. 3D) and C. hendeyi (Fig. 3E). Radius (Fig. 2A). (1) Capital tuberosity, medially situated in anconal view; (2) ulnar facet relatively shallow in palmar view. Character 1 separates C. striatus (Fig. 2B) and Colius cf. C. striatus (Fig. 2A) from C. colius (Fig. 2C). Character 2 separates Colius cf. C. striatus (Fig. 2A) from C. striatus (Fig. 2B), C. colius (Fig. 2C) and C. leucocephalus (Fig. 2D). Radius of other extinct species is unknown. Carpometacarpus (Fig. 2E). (1) Process of metacarpal I and (2) facet for digit II relatively robust. Both characters separate C. striatus (Fig. 2F) and Colius cf. C. stria- tus (Fig. 2E) from C. colius (Fig. 2G), C. leucocephalus (Fig. 21) and C. hendeyi (Fig. 2H). (It was not possible to make adequate comparisons with Colius cf. C. palustris in these characters without examining the actual specimen.) Phalanx I of digit IT (Fig. 1K). (1) Proximal view, internal margin of metacarpal facet rounded, not pointed at posterior end. This character separates C. striatus (Fig. 1L) and Colius cf. C. striatus (Fig. 1K) from C. colius (Fig. 1M) and C. leucoce- phalus (Fig. 1N). Femur (Fig. 4A). (1) Fibular condyle not as deeply notched as in C. colius (Fig. 4C), but more deeply notched than in C. leucocephalus (Fig. 4E); (2) tubercle above external condyle, posterior view, more raised and prominent than in either C. colius (Fig. 4C) or C. leucocephalus (Fig. 4E). Proximal end not diagnostic, no distal ends known for C. hendeyi or any other fossil species. Tibiotarsus (Fig. 3F, G). (1) Interarticular area has a single, deep depression at the base of the rotular crest in proximal view; (2) inner cnemial crest relatively reduced in proximal view; (3) outer cnemial crest much reduced; (4) rotular crest lacking indentation; (5) rotular crest not as erect as in Colius cf. C. palustris, but more erect than in C. archiaci and C. consobrinus (difficult to compare with Colius cf. C. palustris, for same reason as above); (6) distal end with external condyle not deflected externally in anterior view; (7) internal ligamental prominence aligned with anterior, not posterior, shaft edge; (8) condyles not well rounded posteriorly in exter- nal view. Character 1 is shared only with C. striatus (Fig. 3H) and C. consobrinus. Character 2 separates C. striatus (Fig. 3H) and Colius cf. C. striatus (Fig. 3F) from C. colius (Fig. 31) and C. archiaci. Characters 3 and 4 separate C. striatus (Fig. 3H) and Colius cf. C. striatus (Fig. 3F) from C. consobrinus and C. archiaci. Character 6 separates Colius cf. C. striatus (Fig. 3G) and all extant mousebirds (this report) and C. hendeyi (Fig. 3K) from C. archiaci. Character 7 separates C. striatus (Fig. 3H) and Colius cf. C. striatus (Fig. 3G) from C. consobrinus. Character 8 separates Colius cf. C. striatus (Fig. 3G) from all extant mousebirds (this report) and C. hendeyi (Fig. 3K) from C. palustris. Tarsometatarsus (Fig. 4F, G). (1) Cotylae more oval than circular in proximal view; (2) cotylae with lateral edges more or less even or internal edge not more raised than external edge in anterior view; (3) external cotyla projects farther anteriad and dips slightly toward the distal end (most marked in C. striatus (Fig. 4H)); (4) internal trochlea almost same length as internal ridge of middle trochlea, and also relatively larger and not as close to medial trochlea. Characters 1-4 separate C. striatus (Fig. 4H) and Colius cf. C. striatus (Fig. 4F, G) from C. colius (Fig. 41), C. leuco- PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE 203 Fig. 4. A-E. Femora. A. Colius cf. C. striatus, OLD FLK NNI 15356. B. C. striatus, PB25289. C. C. colius, SAM-ZOT.26. D. C. hendeyi, L24593F. E. C. leucocephalus, YPM5797 (transposed). x3. F-K. Tarsometatarsi. F. Colius cf. C. striatus, OLD FLK NI 12867. G. Colius cf. C. striatus, OLD FLK NI 1034. H.C. striatus, PB25226. I. C. colius, SAM-ZO57160. J. C. hendeyi, L28423FZ. K. C. leucocephalus, YPM5797 (transposed). X 2.56. cephalus (Fig. 4K) and C. hendeyi (Fig. 4J). Character 4 separates C. striatus (Fig. 4H) and Colius cf. C. striatus (Fig. 4G) from C. palustris. Genus Urocolius Bonaparte, 1854 Urocolius sp. Fig. 5A, E, 1, M Material Olduvai. Coracoid: FLK NI 26010 (r, sternal end missing sterno-coracoidal process and internal distal angle). Ulna: FLK NI 19721 (lp, with part of shaft); FLK NI 19765 (rd, with part of shaft). 204 ANNALS OF THE SOUTH AFRICAN MUSEUM Fig. 5. A-D. Coracoids. A. Urocolius sp., OLD FLK I 26010. B. U. indicus, SAM-—ZO57546. C. U. macrourus, M3184. D. Colius striatus, SAM-—ZOT.592. E-P. Ulnae. E, M. Urocolius sp., OLD FLK NI 19721. F,N. U. indicus, PB27428. G, O. U. macrourus, PB27520. H, P. Colius striatus, PB25289. I. Urocolius sp., OLD FLK NI 19765. J. U. indicus, PB27428. K. U. macrourus, PB27519. L. Colius striatus, PB25289. All figures x 3. PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE 205 Age and distribution Late Pliocene, about 1,72 to 1,83 m.y. (Curtis & Hay 1972; Haq et al. 1977; Savage & Russell 1983; Kappelman 1986). This species is known only from sites FLK N in Bed I at Olduvai Gorge, north-western Tanzania. Measurements The maximum proximal width of OLD FLK NI 19721 (ulna) is 3,9 mm and its least shaft width is 1,6 mm. The maximum distal width of OLD FLK NI 19765 (ulna) is 3,3 mm. The width and depth of the shaft just anterior to the attachment of the coraco-brachialis of OLD FLK NI 26010 (coracoid) is 1,7 mm and 1,2 mm respect- ively. (See Table 4.) TABLE 4 Measurements (mm) of living and fossil species of Urocolius. Urocolius sp. U. indicus U. macrourus Measure- is a. n mean range n mean range n mean CORACOID 10 ] le 1,1-1,6 26 ; 1 O=1 3) 22. 11 1 52 1,0-1,3 26 1,15 OF 2 5 1,08 HUMERUS 225-242, 33) 923513 22256. s0™ =22 13 2 ue 338} 8,28 7,7-8,5 6 7,90 8 2.0=255 33 2,18 2 O=223\a9 i) 2,10 4 = 25233 1,96 LO=1975 5 1,74 6 4,7-5,4 33 a) V2 AS =552 na, 2,10 7 259-353) 33 3,06 PRTIEBYA Oa Tl 2,87 ULNA ] 22,4-24,8 33 23,80 DOD 23,3" On 2215 2 1 379 353-3,1.. 33 3,54 359-5507) mil Syovll 4 ] 1,6 L216) 33 1,42 1,2-1,5 6 1,30 6 323 3,0-3,5 33 3,16 3,0-3,3 W Sal5 Measurements are: | = greatest length; 2 = maximum proximal width; 4 = minimum shaft width; 6 = maximum distal width; 7 = maximum distal depth; 8 = maximum head depth; 10 = shaft width sternal end just proximal to expansion point; 11 = shaft depth sternal end just proximal to expansion point. Remarks These specimens are larger and more robust than the modern species of mouse- birds, except for Urocolius indicus. Unfortunately, the sterno-coracoidal process, which is missing on the fossil specimen (Fig. 5A), is one of the most diagnostic fea- tures of the coracoid at the generic level in the family Coliidae. However, the pronounced attachment of the coraco-brachialis and the presence of a small nutrient foramen close to this attachment (dorsal view) on the fossil specimen are more charac- teristic of the genus Urocolius (Fig. 5B, C) than the genus Colius (Fig. 5D). These features, in addition to the overall robust nature of this specimen, favour its place- ment in the genus Urocolius. It is not possible to assign it to a species. 206 ANNALS OF THE SOUTH AFRICAN MUSEUM The slightly more pointed olecranon, the well-developed fossa under the external cotyla (palmar view) and the scar on the surface external to the humero-ulnar depres- sion on specimen OLD FLK NI 19721 (proximal ulna) (Fig. 5E, M) are features it shares with the genus Urocolius (Fig. 5F, G, N, O) rather than with the genus Colius (Fig. 5H, P). Likewise, the external condyle tapering to a point, rather than being rounded, at the base of the shaft in internal view and the pointed rather than rounded carpal tuberosity and the slightly larger internal condyle, are features that specimen OLD FLK NI 19765 (distal ulna) (Fig. 51) has in common with the genus Urocolius (Fig. 5J, K) rather than with the genus Colius (Fig. 5L). The depth of the fossa under the external cotyla is greatest in the fossil specimen (Fig. 5E) and rather variable in the extant species of Urocolius (Fig. 5F, G). However, it is generally better developed in the genus Urocolius than in the genus Colius (Fig. 5H) and is therefore considered to be diagnostic at this level. These specimens fit better within the size range of Urocolius indicus than that of U. macrourus but morphologically they share features with both species. Conse- quently, without a larger sample, it is not possible to assign them to a species. They provide the first Tertiary record of the genus Urocolius. BIOMETRICAL ANALYSIS METHODS The mensural data were analysed using the co-variance biplot technique, which is one of a family of data analytic techniques that displays the rows and columns of a data matrix as points in a low-dimensional space, usually consisting of two or three axes (Greenacre & Underhill 1982). The analysis can be reduced to three steps: 1. Defining two clouds of points on their corresponding two multidimensional spaces; here the points of each cloud represent the specimens and the skeletal measurements respectively. 2. Defining a metric structure on each cloud of points that refers to how distances between specimens and between measurements are defined. 3. Defining the fit of each cloud of points to a low-dimensional space on to which the points are projected for subsequent display. These two or three dimensions represent, as accurately as possible, the points’ true high-dimensional positions. A full descrip- tion of the analysis can be found in Greenacre & Underhill (1982), Greenacre (1984) and Underhill (1990). Prior to analysis, the data were standardized by subtracting from each measurement its corresponding column mean. This renders all column means equal to zero but keeps the respective variances unchanged. RESULTS Using the above method, Figures 6, 7 and 8 help to demonstrate similarities between the fossil and living species of mousebirds studied for this report. Figure 6 depicts the results using six measurements of the humeri (greatest length, GLE; maximum proximal width, MPW; depth of head, HD; maximum distal width, MDW; maximum distal depth, MDD; minimum shaft width, MSW) of 114 specimens representing seven species of mousebirds. Axis 1 accounts for 85 per cent of the variance of the data matrix. It is defined by variables GLE and MPW. PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE 207 AXIS 2 6 aes eS —— 66 \ ie 6 lo 7. S ~ fe Cre NG 6 AXIS: S Fig. 6. Result of co-variance biplot analysis based on mensural data of the humeri of seven species of living and fossil mousebirds. 1 = Colius striatus (n = 51); 2 = C. colius (n = 19); 3 = C. leucocepha- lus (n = 1); 4 = Colius cf. C. striatus (n = 1); 5 = Urocolius macrourus (n = 5); 6 = U. indicus (n = 33); 7 = Colius hendeyi (n = 4). A = greatest length (GLE); B = maximum proximal width (MPW); C = maximum head depth (MHD); D = maximum distal width (MDW); E = maximum distal depth (MDD); F = minimum shaft width (MSW). Specimens with large values for these two variables are pulled towards them, e.g. group 4 (Colius cf. C. striatus), group 6 (Urocolius indicus), and some specimens of group 1 (Colius striatus). Conversely, those specimens with small GLE and MPW are plotted on the opposite side, e.g. group 7 (Colius hendeyi), group 2 (C. colius), and part of group 1 (C. striatus). Axis 2, which accounts for 19 per cent of the variance, is defined by MPW and GLE. Specimens with large MPW are plotted in the same direc- tion of that variable (B), e.g. groups 5 and 6 (Urocolius macrourus and U. indicus). Specimens with small MPW and large GLE are on the lower half of the plot, e.g. group 2 (Colius colius), group 1 (C. striatus), and group 4 (Colius cf. C. striatus). The extinct species Colius hendeyi (group 7) is very clearly separated from all the other species on account of it having both small GLE and MPW. Figure 7 shows the results using three measurements (GLE, MPW and MDW) of the ulnae of 118 specimens representing seven species of living and fossil mousebirds. Axis 1 accounts for 98,8 per cent of the variance. It is defined by the variable GLE. Axis 2 is defined by variables MPW and MDW. The seven species are separated simi- larly as in Figure 1, with a certain amount of overlap between group 1 (Colius striatus) and group 2 (C. colius). The fossil species group 4 (Colius cf. C. striatus) falls well within the distribution of C. striatus. The extinct species C. hendeyi (group 7) is again clearly separated from all the other species. 208 ANNALS OF THE SOUTH AFRICAN MUSEUM AXIS 1 Fig. 7. Result of co-variance biplot analysis based on mensural data of the ulnae of seven species of living and fossil mousebirds. 1 = Colius striatus (n = 53); 2 = C. colius (n = 21); 3 = C. leucocepha- lus (n = 1); 4 = Colius cf. C. striatus (n = 2); 5 = Urocolius macrourus (n = 6); 6 = U. indicus (n = 33); 7 = Colius hendeyi (n = 2). A = greatest length (GLE); B = maximum proximal width (MPW); C = maximum distal width (MDW). Figure 8 displays the results using three different measurements (MPW, MDD, MSW) of the ulnae of 117 specimens representing eight species of living and fossil mousebirds. Axis 1 accounts for 82 per cent of the variance. It is defined by variables MDW and MPW. Axis 2 is also defined by variables MDW and MPW. Although the species are plotted in associations similar to those in Figures 6 and 7, there is consider- ably more overlap. However, group 4 (Colius cf. C. striatus) still falls within the range of group | (C. striatus). Group 8, which represents Urocolius sp. from Olduvai, is indeed placed closest to the Urocolius species complex (groups 6 and 5). It must be emphasized that the measurements for Urocolius sp. were combined from two differ- ent specimens for the purpose of this analysis. DISCUSSION Most of the questions raised by Rich & Haarhoff (1985) with regard to the origin and systematics of the Coliiformes remain unanswered. Although the oldest mouse- bird fossils have been found in the Upper Eocene of France (Mourer-Chauviré 1988), it cannot be said that Europe is the place of origin for the Coliidae, because the fossil record for the early and mid-Tertiary of Africa and Asia is still so poorly known. The Miocene specimens from Europe still require revision. Material referred to the genus Colius by Ballmann (1969) may represent another extinct genus, whereas some other PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE 209 AXIS 2 aan / \ 7 - Cc Vie = ‘ us a 111 x A SS 1 1 Ness 116666 8 Me < ee 6 N aN \ 1 Fig. 8. Result of co-variance biplot analysis based on mensural data of the ulnae of eight species of fossil and living mousebirds. 1 = Colius striatus (n = 52); 2 = C. colius; (n = 18); 3 = C. leucoce- phalus (n = 1); 4 = Colius cf. C. striatus (n = 2); 5 = Urocolius macrourus (n = 6); 6 = U. indicus (n = 33); 7 = Colius hendeyi (n = 4); 8 = Urocolius sp. (n = 1). A = maximum proximal width (MPW); B = maximum distal width (MDW); C = minimum shaft width (MSW). material from Europe, examined by Ballmann (pers. comm.), may belong to the genus Urocolius. How the living and the fossil species are related to each other is not understood. This, and the fact that only skeletal material can be studied, precludes a more definitive statement other than that the Olduvai species Colius cf. C. striatus might be ancestral to the living Colius striatus. More fossil material of the other Olduvai species, Urocolius sp., would probably help to clarify its taxonomic status. Some of the remaining questions therefore pertain to the phylogenetics of the Collii- formes, when and why they became restricted to Africa, and what limits their present distribution. The five extant species of mousebirds studied in this report show that there is some overlap in terms of size but, generally, Colius striatus has the largest and most robust skeleton of that genus. The wing, tarsus and weight measurements given in Fry et al. (1988) suggest, however, that, of the four living species of Colius, the skeleton of C. castanotus should, in fact, be the most robust. Because of a lack of comparative specimens of this species, it was not possible to confirm this. In the genus Urocolius, the skeleton of the red-faced mousebird, U. indicus, tends to be more robust than that of the blue-naped mousebird, U. macrourus. A comparison between the measure- ments given in Rich & Haarhoff (1985) and in this paper shows that the considerably larger sample of specimens used herein has produced a wider range of variation in the 210 ANNALS OF THE SOUTH AFRICAN MUSEUM elements measured; more sexed specimens were available, but there is no evidence of sexual dimorphism in any of the species studied. The morphological features used to differentiate the species are difficult to inter- pret in that they tend to be differences of degree (e.g. shape, angle, etc.), rather than simply being present or absent. A better understanding of their functional relevance would help to determine whether they are phylogenetically useful. The coracoid, humerus and tibiotarsus have the most easily defined morphological differences at the species level. The co-variance biplot analyses shown in Figures 6-8 tend to support the results of the morphological data presented herein and in Rich & Haarhoff (1985). Colius cf. C. striatus is shown to be consistently associated with Colius striatus in all three figures; the position of Urocolius sp. is closest to that genus in Figure 8; and the extinct species Colius hendeyi is shown to be well separated from all the other species but is clearly associated with the genus Colius. In Figure 6, the distance of the fossil species Colius cf. C. striatus from group 1 (C. striatus) is possibly due to the MPW measurement of the fossil being smaller than the norm in proportion to its GLE. This, in turn, could be due to the wear on the fossil bone. Although the fossil sample size is very small, the results of this type of analysis indicate that similarities between the dif- ferent groups/species can be demonstrated in the form of loose associations. However, it should also be noted that whereas Colius cf. C. striatus falls consistently within the range of C. striatus, so also does the single specimen of the extant species C. leucoce- phalus (group 3). Thus, the problem of having such a small sample is also highlighted. It is evident that the species are separated more clearly when the data used are a combination of both small and large measurements, for example, when GLE, MPW and MDW are used, as in Figure 7. Where only small measurements (MPW, MSW, MDW) have been analysed, as in Figure 8, the overlap between the different species is noticeably greater. Unfortunately, when fossil bones form part of the data base, the most useful measurements cannot always be taken, due to the incomplete nature of some of the specimens. The fossil mousebirds from Olduvai add one more small piece to the puzzle of the history and biogeography of this curious avian order. ACKNOWLEDGEMENTS This paper is dedicated to the late Professor Pierce Brodkorb. He made it poss- ible for me to study the material and I am honoured that I have been afforded this opportunity. I regret most sincerely that we have been unable to publish the work together as originally planned. I am most grateful to Diana Matthiesen who identified these fossils as Colius spp. (sensu lato) during her preliminary identification of the Olduvai fossil bird collection and provided provenance data and background information. René Navarro (PFIAO) is greatly thanked for his considerable contribution to the section on the methods and results. I thank James Dean and Dr Storrs Olson (Smithsonian Institution), Dr Alison Andors (American Museum of Natural History), Tamar Cassidy and Meg Kemp (Transvaal Museum), Johann Welman (National Museum, Bloemfontein), Dr Gra- ham Avery and Denise Drinkrow (South African Museum), Eleanor Stickney and PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE Dla Professor Charles Sibley (Peabody Museum) for the loan of comparative material. Dr Graham Avery also helped with a preliminary statistical analysis for which I am grateful. I had useful discussions and/or comments from Dr Peter Ballmann, Richard Brooke, Professor Tim Crowe, Richard Dean, Diana Matthiesen, Terry Oatley, Sally Price, Dr Pat Vickers-Rich and much appreciated guidance from Drs Margaret Avery and Gillian King. Clive Booth, Cedric Hunter and Bill van Rijssen are greatly thanked for their work on the figures. Jacqueline Blaeske and Mike Wilson aided in the preparation of the manuscript. The South African Museum is thanked for allowing me the time to work on this project. REFERENCES BALLMANN, P. 1969. Les oiseaux miocénes de la Grive-Saint-Alban (Isére). Geobios 2: 157-204. Bropxors, P. 1971. Catalogue of fossil birds. Part 4. (Columbiformes through Piciformes). Bulletin of the Florida State Museum: Biological Sciences 15: 163-266. Bropkors, P. & Mourer-CuHauvirE, C. 1982. Fossil anhingas (Aves: Anhingidae) from early man sites of Hadar and Omo (Ethiopia), and Olduvai Gorge (Tanzania). Geobios 15 (4): 505-515. Bropkors, P. & Mourer-CHAUVIRE, C. 1984a. A new species of cormorant (Aves: Phalacrocoraci- dae) from the Pleistocene of Olduvai Gorge, Tanzania. Geobios 17 (3): 331-337. Bropxors, P. & Mourer-CHAuvirE, C. 19846. Fossil owls from early man sites of Olduvai Gorge. Ostrich 55 (1): 17-27. Curtis, G. H. & Hay, R. L. 1972. Further geological studies and potassium-argon dating at Olduvai Gorge and Ngorongoro Crater. Jn: Bishop, W. W. & MILLER, J. A. eds. Calibration of homi- noid evolution: 289-301. Edinburgh: Scottish Academic Press. Fry, C. H., KerrH, S. & URBAN, E. K. 1988. The birds of Africa 3. London: Academic Press. GREENACRE, M. J. 1984. The theory and application of correspondence analysis. London: Academic Press. GREENACRE, M. J. & UNDERHILL, L. G. 1982. Scaling a data matrix in a low-dimensional euclidean space. In: Hawkins, D. M. ed. Topics in multivariate analysis: 183-268. Cambridge: Cambridge University Press. Haq, B. U., BERGGREN, W. A. & VAN COUVERING, J. A. 1977. Corrected age of the Pliocene/Pleisto- cene boundary. Nature 269 (5628): 483-488. HARLAND, W. B., ARMSTRONG, R. L., Cox, A. V., Craic, L. E., SmitrH, A. G. & Situ, D. G. 1990. A geologic time scale 1989. Cambridge: Cambridge University Press. Harrison, C. J. O. & WALKER, C. A. 1976. A new fossil pelican from Olduvai. Bulletin of the British Museum of Natural History (Geology) 27 (4): 315-320. Harrison, C. J. O. & WaLker, C. A. 1979. A recent and an extinct cormorant species from the Middle Pleistocene of Tanzania. Ostrich 50 (3): 182-183. KAPPELMAN, J. 1986. Plio-Pleistocene marine-continental correlation using habitat indicators from Olduvai Gorge, Tanzania. Quaternary Research 25 (2): 141-149. Leakey, L. S. B. 1965. Olduvai Gorge 1951-1961 1. A preliminary report on the geology and fauna. Cambridge: Cambridge University Press. MILNE Epwarps, A. 1871. Recherches anatomiques et paléontologiques pour servir a histoire des oiseaux fossiles de la France 2. Paris: Masson. Movrer-Cuavuvir£, C. 1988. Le gisement du Bretou (Phosphorites du Quercy, Tarn-et-Garonne, France) et sa faune de vertebres de Eocene Superieur. Palaeontographica (A) 205: 29-50. Otson, S. L. 1985. The fossil record of birds. In: FARNER, D., Kinc, J. & PARKES, K. C. eds. Avian biology 8: 79-238. New York: Academic Press. Rico, P. V. & Haaruorr, P. J. 1985. Early Pliocene Coliidae (Aves, Coliiformes) from Langebaan- weg, South Africa. Ostrich 56 (1-3): 20-41. Savace, D. E. & Russet, D. E. 1983. Mammalian paleofaunas of the world. Massachusetts: Addison-Wesley Publishing Company. ScuirTer, H. 1985. Systematics and distribution of mousebirds (Coliidae). Proceedings of the Inter- national Symposium on African Vertebrates, Bonn 1985: 325-347. UNDERHILL, L. G. 1990. The co-efficient of variation biplot. Journal of Classification 7: 241-256. 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°S1’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. SMITHSONIAN INSTITUTION LIBRARIES “ICAI ANIA 3 9088 01206 7054 PHILIPPA J. HAARHOFF LATEST PLIOCENE MOUSEBIRDS (AVES, COLHDAE) FROM OLDUVAI GORGE, TANZANIA