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Konn, A. J. 19606. 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 103 +#£2Band

June 1993 Junie

Part 4 Deel

LATEST PLIOCENE MOUSEBIRDS

(AVES, COLITIDAE)

FROM OLDUVAI GORGE, TANZANIA

PHILIPPA J. HAARHOFF

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

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D2221

LATEST PLIOCENE MOUSEBIRDS (AVES, COLIIDAE)

FROM OLDUVAI GORGE, TANZANIA

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

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VOIddV NYAHLNOS

2 TVALNAO

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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

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

range

mean

range

mean

range

Measure-

ment no.

CORACOID

Vena’ a)

oom

NANN

ANTM

RUS

HUME

~~, — ae ES o ~~,

mer ATTN

Seni seeniihaeen ithe een ihe!

m= AN xt OO

NADONe

Sener

FONSTEISS

mom tN

EASPEN ON

AnaAN

—_—

Stag

Saag

CDIDIG IS

on —N

—

1,8

Levy

1,92 155=159 211

1,5-2,1

Se ih eee ee |

CO = CA GD)

13,1-13,2

—Notror ZHNtO AN FGHnOt

=) < =

[a4

Pal Diall

=—aAD

FEMUR

SESRENSREN

eoranann

20 20,24

4,29

2,83

4,03

2.93

oroort

~MnA—O

= TNO

Ay (al

=ANMOrM

PLIOCENE MOUSEBIRDS FROM OLDUVAI GORGE

COCONC

SCAICAICA

moon

Om 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) =

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

Measurements are:

199

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

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

aes

—— 66

\ ie 6 lo

7. S

~ fe Cre NG

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

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.

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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-

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Savace, D. E. & Russet, D. E. 1983. Mammalian paleofaunas of the world. Massachusetts:

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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

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.