Carnivora from the Kanapoi hominid site, Turkana Basin, northern Kenya

18 Pages·2003·14.5 MB·English

by  Lars Werdelini

Carnivora from the Kanapoi Hominid Turkana
Site,
Northern Kenya
Basin,
Lars Werdelini
ABSTRACT. Kanapoi is the earliest Pliocene site yet described in eastern Africa with a substantialcarni-
voran record. It includes eight species in as many genera, representing five families. The material is dom-
inated by the hyaenid Parahyaena howelli n. sp., but also includes a new Enhydriodon species, E. ekeca-
man, thelutrinecf. Torolutrasp.,thefelidsDinofelispetteri,Homotheriumsp.,andFelissp.,theherpestid
Helogalesp., andtheviverrid Genetta n. sp.TheKanapoi Carnivora includestheremainsofthefirstpost-
Miocene radiation ofendemic African Carnivora.
INTRODUCTION CONVENTIONS AND ABBREVIATIONS
The earliestPliocene (heretakenasca. 5.2-4.0Ma) Kanapoi fossils in the collections of the National
has provided relatively few eastern African locali- Museums of Kenya are formally accorded the ac-
ties containing mammalian fossils compared with ronym KNM-KP before the accession number; to
the million years that followed. Most of the local- save repetition, this acronym is omitted from the
ities thatdo existfrom thistime intervalhaveeither lists of Kanapoi specimens and abbreviated to KP
few carnivores associated with them or are as yet in the body ofthe text.
undescribed. Thus, the Apak Member atLothagam Tooth abbreviations in upper case (I, C, P, M)
includes only fourcarnivore taxa (Werdelin,2003), indicate upper teeth; those in lowercase (i, c, p, m)
the Lonyumun Member ofthe Koobi Fora Forma- indicate lower teeth. Hence Ml is the upper first
tion three (Werdelin and Lewis, unpublished), the molar, p4 is the lower fourthpremolar. Thefollow-
Kataboi Member of the Nachukui Formation and ing abbreviations are used in the text and tables:
Kosia (also West Turkana) also three (personal ob-
a-p anteroposterior
servations). Outside ofKenya, localitiesofthistime
Dist distal
period from Uganda have also yielded few carni-
vores (Fetter et ah, 1991). A richer site is Aramis, E anatomical length (long bones); mesio-
distal length (teeth)
in Ethiopia, although the carnivores there have yet
LpP, length of main cusp ofP
to be described (Howell, personalcommunication). MC 4
metacarpal
Kanapoi, with its somewhat larger sample of car- MT
metatarsal
nivores, thus addssignificantlyto ourknowledgeof max maximum
carnivoranevolution intheearliestPlioceneofeast- min minimum
ern Africa.
Prox proximal
The bulk of the material discussed herein was
Sust sustentaculum
obtained by the National Museums of Kenya ex-
transv transverse
peditions to Kanapoi in the early 1990s. However, W
buccolingual width
a few carnivore specimens were recovered by the
WaP" anterior width (including protocone) of
earlier American expeditions. These have been
P4
mentioned a few times in the literature under var- WblP^ minimum blade width of P^
ious guises (Behrensmeyer, 1976; Savage, 1978;
Howell and Fetter, 1980; Turner et ah, 1999). In SYSTEMATIC DESCRIPTION
particular, Behrensmeyer (1976) lists Enhydriodon
n. sp.. Hyaena sp., and Machairodontinae indet. as Order Carnivora
present at Kanapoi. Of these, the first is here still Family Mustelidae
referredtoEnhydriodonFalconer, 1863,thesecond
to Parahyaena Hendey, 1974, while the third is Generally speaking, mustelids are rare in the fossil
here listed as Carnivora indet., as it cannot be de- record of eastern Africa. For the most part, this
termined if the specimen belongs to Dinofelis may reflect a preservational bias against smaller
Zdansky, 1924, or Homotherium Fabrini, 1890. species ofCarnivora. However, those localitiesthat
The remaining taxa described below are newtothe are particularlyrichinsmallerCarnivoraalsodiffer
Kanapoi fauna. among themselves in the richness and diversity of
their mustelids. Thus, Hadar is rich in small mus-
1. Department of Palaeozoology, Swedish Museum of telids (personal observations), while Olduvai has
Natural History, Box 50007, S-104 05, Stockholm, Swe- fewer taxa, despite the smaller Carnivora being
den. well represented at the latter site (Fetter, 1973).
Contributionsin Science, Number498, pp. 115-132
Natural HistoryMuseum ofLos Angeles County, 2003
116 CS 498, Harris and Leakey: Kanapoi
basin andrelatively smallerparaconid, morerobust
and larger Ml hypocone(s) and more stoutly built
i
P4.
HOLOTYPE. KNM-KP 10034, A = right P4
(Fig. lA), B = right ml (Fig. IB), C = right Ml
(Fig. 1C), D = right C). Written documentationre- ii'
garding the association in the field of these four
specimens was provided by Dr. J. C. Barry (in cor- i
respondence). Given this information, it is reason-
i
able to assume that all four belonged to a single
individual and therefore the entire hypodigm is j
chosen as the holotype of this new species.
ETYMOLOGY. After the Turkana “ekecaman,”
meaning fisherman. The diet of this animal was |
i
composed at least in part of fish.
KANAPOI MATERIAL. 10034; associated teeth
,
(holotype).
Figure 1 Enhydriodon n. sp., A, KP 10034A, right P4 in MEASUREMENTS. KP 10034A, buccal length
occlusal view; B, KP 10034B, right ml in (top) occlusal = 16.5; KP 10034B, total length = 21.2, trigonid
and (bottom) lingual view; C, KP 10034C, right Ml in length = 11.5, talonid length = 9.3, trigonidwidth
occlusal view. In A and C, anterior is to the right; in B, = 13.3, talonid width = 13.5; KP 10034C, buccal
anterior is to the left length = 12.1, lingual length = 15.8, anterior
width = 19.8, posterior width = 19.0. Measure-
ments defined as in Willemsen (1992).
/ The rightuppercarnassial, KP 10034A (Fig. lA),
This variation may reflect real differences in paleo- is missing the protocone and most ofthe hypocone
ecology between the sites. Nevertheless, the major- and posterior shelf. The parastyle is short and low
ity of mustelids found in the eastern African fossil but has a distinct anterior cusp. The paracone is
record are larger species of the genera Mellivora high and much the largest buccal cusp. There are
Storr, 1780, Enhydriodon and Torolutra Fetter, blunt crests leading anteriorly to the parastyle, an-
Pickford, and Howell, 1991. Kanapoi is no excep- terolingually toward the protocone and posteriorly
tion to this pattern, with both of the latter genera toward the metacone. The lattercrestis interrupted
represented. by a shallow valley before it meets the metacone.
The latter cusp is set far posteriorly and is much
Enhydriodon Falconer, 1863 lower and smaller than the paracone. Therewas an
extensive basin formed posterobuccally to the hy-
The genus Enhydriodon encompasses a number of pocone and the posterobuccal corner and posterior
species of Enhydrini (sea otters) of large to very margin ofthe tooth lie at nearly right angles to the
large size. The genus is known from the Siwaliks anteroposterior axis ofthe tooth. Thecingulumex-
i
of Pakistan and India, where it was first described tends along the entire preserved part of the tooth
(Lydekker, 1884; Willemsen, 1992). However, the except for the posterobuccal corner.
main diversity within Enhydriodon is found in Af- The right lower carnassial, KP 10034B (Fig. IB)
rica, with several species of varying size known is low, broad, and stoutly built. The three trigonid
from a number of localities dating between ca. 6.0 cusps are all low and pyramidal with their major
Ma (Lukeino Formation; Pickford, 1978) and ca. axes directed either anteroposteriorly (protoconid,
2.5 Ma (Omo, Shungura Formation Members metaconid) or transversely (paraconid). In occlusal
E+F; Howell and Fetter, 1976). The Kanapoi ma- view, the paraconid is the smallest of the three
terial represents one of the earlier members ofthis cusps but all three are worn down to about the
lineage. same height. The apexoftheparaconidissetslight-
ly lingual to the middle of the cusp and it is also
Enhydriodon ekecaman sp. nov. slightly broader lingually than buccally. The apex
(Figure 1) ofthe protoconid is set slightly anteriorto themid-
EEnnhhyyddrriiooddoonn np.atstpe.rsBoenhireSnasvmaegyee,r,19179876(nomen nu- ddloewonftthheelciungsupa.lTshiedereofisthaisbcluusnpt tcroewsatrdthatthertuanls-
onid, making the cusp almost diamond-shaped,
dum)
EnhydriodonpattersoniTurner, Bishop,Denys,and thoughwitha gentlycurved buccalside. Anteriorly,
McKee, 1999 (nomen nudum) the protoconid is separated from the paraconid by
a shallow valley. Posteriorly, the accessory proto-
DIAGNOSIS. Differs from Siwalik Enhydriodon conid cusp is very weakly developed, merely form-
in its smaller size. Differs from E. africanus Strom- ing a low bulge on the posterior face of the cusp.
er, 1931 (the other described AfricanEnhydriodon) The metaconid is set slightly posterior to the pro-
by having a broader ml with a more open talonid toconid. Its apex is at the middle of the cusp. The
Werdelin: Carnivorans 117
cusp is triangular in occlusal view, with the apex of Kanapoi Enhydriodon belongs to a hitherto un-
the triangle directed toward the buccal side. Ante- known species of that genus, a species that is not
riorly, the metaconid is separated from the para- at present known from any other site. All four En-
conid by a deep valley. Between the trigonid cusps hydriodon teeth from Kanapoi originate from the
mm
is formed a shallow, flat basin that is about 3 American expeditions (1966-72) and were collect-
wide and 2 mm long. The talonid basin is low and ed in 1967.
wide. The entoconid is well developed but low. In
occlusal view, it is about the size ofthe paraconid. Torolutra Petter, Pickford, and Fiowell, 1991
None ofthe other talonid cusps is well defined. In- Torolutra is a genus of otters similar in size to the
stead, they form a low, broad, gently undulating living European otter Eutra (Linnaeus, 1758),
ridge surroundingthecentral basin. The buccalcin- or slightly larger. Only a single species, T. ougan-
gulum is strong and extends from the anteriormost densis, has been described (Petteretah, 1991).This
part of the tooth to the posterior end of the hypo- material is from Nyaburogo and Nkondo in Ugan-
conid. da, while the species has also been tentativelyiden-
The right Ml, KP 10034C (Fig. 1C), is broad, tified from Ethiopia in the Usno Formation of the
low, and robust, with a very broad lingual basin. Omo Group. These localities bracket Kanapoi in
The paracone is small, and buccal to it, there is a age.
large parastyle, which is in fact larger than the
paracone itself. The paracone is low and worn and cf. Torolutra sp.
connected to the metacone by a narrowvalley. The
pmaertaaccoonneeaisncdonhsaisdearpabylryamhiidgahlerbaasned.lIatrgisersetthaant tthhee rigKhtANP4APfrOaIgmMenAtTs,ERpIreAmLo.la3r01f5ra5g,mreingth,tm13l,lteaftloannidd
posterior margin ofthe tooth. Lingual to the meta- fragment, proximal left radius fragment, proximal
cone and separated fromit bya narrowvalleythere right tibia fragment, humerus shaft fragment, par-
is a metaconule that is almost as large as the meta- tial cervical, thoracic and caudal vertebrae, assort-
ed indeterminate fragments.
cone itself. This cusp is also set at the posterior
margin of the tooth. Lingual to the metaconule The 13 is strongly recurved and has a short
there is a wear facet for occlusion with m2. This crown. The enamel reaches farther down on the
wear facet is confined to the posterior margin of lateral than on the medial side. There is a promi-
the tooth. At the posterolingual corner ofthe tooth nent cingulum surrounding this tooth. This cingu-
there is a large swelling ofthe cingulum, forminga lum is best developed on the medial side. The root
low cusp. The protocone is double, with one cusp of13 is relatively straight. The P4 preservestheme-
set slightly anterobuccal to the other. These two tastyle, a partial paracone and a part ofthe lingual
cusps are of about equal size. The cingulum runs protocone shelf. There is no carnassial notch. The
around the entire tooth, except anteromedially, protocone shelf is not as long as in Enhydriodon,
wahnedrpeositterioslwinogrunalldyo,wwnhebrye tthheeawfeoarremfeancteitonfoerdmP42, b1e7i6n2g.iTnhsteeamdlmoprreesseirmvielsarthteo tphoastteorfioErutpraartBroifsstohne,
wear facet is located. The cingulum is otherwise talonid, with a prominent hypoconid and low
weakest around the metacone. bumps indicating entoconulid and entoconid. The
The right C, KP 10034D, is short and straight, cingulum is prominent around the posterior end of
with a crown that is only slightly longer than it is the tooth. The proximal radius fragment is small
wide. The tip is worn flat. There is a strong medial and broken. The proximal tibia fragment shows
cingulum but no lateral one. only parts ofthe proximal articular surface. All the
DISCUSSION. The material of Enhydriodon vertebrae are relatively robust, the proximalcaudal
from Kanapoi is limited, but can nevertheless be vertebra extremely so, while the cervical vertebrae
distinguished from other African Enhydriodon of are relatively much smaller.
similar age. The lower carnassial, for example, dif- DISCUSSION. These specimenscomparewell,as
fers from that of E. africanus from Langebaanweg far as comparisons can be made, with specimensof
amongotherfeaturesinbeingshorterandrelatively Torolutra ougandensis described by Petter et al.
wniidderc,uspins,thaendsoinmetwhehaftlatmteorraendwimdeolryesoppaecnedtatlroingiod- f(1r9o9m1)U.gTahndeay,arbeutamliatttlcehlatrhgeerOtmhaonstpheecsipmeecnismefings-
basin (cf. Hendey, 1978b). The development ofthe ured by those authors quite well.
pKoasntaeprooliinfgouraml.cTuhspeleMtslisdimfoferres fprroomnoaunpcaertdiailnMthle Family Hyaenidae
from Kosia (West Turkana, ca. 4.0 Ma) in that the Hyenas are common elements in eastern African
anterolingual corner of the Kanapoi Ml forms a Mio-Pliocene faunas. As in Eurasia, there is an ex-
nearly right angle, while the homologous angle of tinction event at the end of the Miocene that elim-
the Kosia tooth is closer to 135 degrees (personal inates the dominant ‘dog-like’ hyenas that are still
observations). The Kanapoi Enhydriodon further present at localities such as Lothagam (Werdelin,
differs fromyoungerEnhydriodon fromformations 2003). AtPliocenelocalities,hyenasaremainlyrep-
such as Hadar, Koobi Fora, and Shungura in its resented by relatives ofthe living hyenas. ThesePli-
much smaller size. Hence, we can infer that the ocene forms had adaptations to a scavenging life-
118 CS 498, Harris and Leakey: Kanapoi
style similar to those of living hyenas, but less ac- tial alveolus for c, alveolus for p2, roots ofp3, an-
centuated. At Laetoli, hyenas are abundant, andin- terior root of p4; 29297, left mandibular ramus
clude fossil relatives of three of the four living fragment with roots ofp4 and anterior root ofml;
hyenas (Werdelin, unpublished). One of these spe- 29299, proximal, shaft, and distal fragments of
cies is also present at Kanapoi. right femur; 29301, left mandibular ramus frag-
ment with p4 and anteriorroot ofml, separate p3;
Parahyaena Hendey, 1974 29302, left P4; 30229, right femur lacking distal
end, proximal right tibia; 30234, associated partial
There has been some debate regarding the validity
left forelimb with left ulna lacking olecranon, left
of Parahyaena as a genus distinct from Hyaena radius, left humerus lacking proximal articulation,
l(iWne,rd19e9l8i)n.aOndnStohleouonnieash,an1d9,91t;oJdeanykstheasnedaWreertdweo- cleifftosrcma;p3h0ol2u3n5a,r,aslseoftcimaategdnupamr,tialelftskpeilseitfoonrmi,nclelfutduinn-g
mgeonneorsipcecdiifstiicncstiistoenrmtaaxya,beanddeefmreodmutnhinsecpeesrssapreyc.tiOven, rleifgthtdimsatnaldihbuumlaerrursamfursagwmietnht,i2rainghdtcp-rmolxi(mFiagl.2aBn)d,
the other hand, the split between the two, as in- distal humerus fragments, rightradius (Figs. 3A-B),
ferred from both molecular and paleontologicalev- right calcaneum, right tibia lacking distal articula-
idence, extends down into the Miocene, and most
tion, right ulna lacking distal articulation, frag-
generic splits among carnivores are of about this ments of the right and left scapulae, left cuboid,
age or even younger. From this perspective, generic right and left navicular, right scapholunar, left lat-
separation is valid. Here I follow the latter path, as eral cuneiform, damaged right lateral cuneiform,
wIbiebtceholimweevheicrctihthiacttaol.gerUoenlstooigllivcenaolrwa,angkPeianrigashityshaseeuenosanlwwyhasecnrkitnteohreiwsonen rtdiiiscgtuhaltlatuainrotcni,icfupolrramot,ixoinmP,arirlgihgpthatrptMatoCefllIarI,iglhaltecfktMinMCgCpIrIIoI,Ixillemfaatclkaiannrdg-
from the single extant species F. brunnea (Thun- right MC V lacking proximal articulations, right
gbeeorlg,ogi1c8a2l0)a,gewh(JiecnhkshaasndaWelrimdietleidn,di1s9t9r8i)b.utAionsinagnlde MinCgpIr,odxiismtaallpaarrtticouflartiigohnt,MleTftaIIn,drirgihgthtMMTTIVV llaacckk--
record extends the range ofF. brunnea into eastern ing proximal articulations, proximal, middle and
Africa in the middle Pleistocene (Werdelin and Bar- distal phalanges includingproximalphalanxofdig-
thelme, 1997). This makes the presence of an an- it 1 of the manus, pisiform, sternebrae, and cervi-
cestral species ofParahyaena atKanapoihighlysig- cal, thoracic, lumbar, and caudal vertebral frag-
nificant. ments; 30272, associatedpartialskeletonwithright
mandible fragmentwithc, damagedp3,rootsofp2
Parahyaena hoivelli sp. nov. and p4, left mandible fragment with roots of p2~
(Figures 2-5) p3, rightmaxilla fragmentwithCroot,PI alveolus,
Hyaena sp. Behrensmeyer, 1976 nearly complete P2, roots of P3, anterior root of
Pachycrocuta sp. Howell and Fetter, 1980 P4, leftmaxilla fragmentwith alveolus for13,dam-
aged C, roots of P1-P2, anterior root of P3, right
DIAGNOSIS. Hyaenid of large size (larger than zygomatic, proximal end ofright and left scapulae,
Hyaena hyaena Linnaeus, 1758). Mandibular ra- fragment of distal right humerus shaft, right tibia,
mus robust, premolars moderately developed for proximal, shaft and distal fragments of left tibia,
bone cracking (weaker than in PliocrocutaKretzoi, pelvic fragment, left navicular, proximal fragment
MT
1938, Pachycrocuta Kretzoi, 1938 and Crocuta ofleft V, tuberfragmentofleftcalcaneum,par-
Kaup, 1828). Masseteric fossa clearly subdivided tial distal right femur, proximal and distal frag-
by a ridge into a ventral and dorsal part (unlikeH. ments ofleft femur, fragmentofleftpisiform,prox-
MC MT
hyaena). Metastyle of P4 clearly longer than para- imal end of right III, pathological left II,
MT MT
cone (unlike Ikelohyaena abronia (Hendey, 1974) distal end of left III, right III, proximal
MT
and H. hyaena). Metacarpals shortand robust (un- fragment of right and left IV, proximal end of
like all modern hyenas). right MT V, proximal phalanx of left manus digit
HOLOTYPE. KNM-KP 30235, associated par- 3, proximal phalanx of right manus digit 4, prox-
tial skeleton (Figs. 2, 3). imal phalanx of right manus digit 5, middle pha-
ETYMOLOGY. After Dr. F. Clark Howell, lead- lanxofrightmanusdigit4,middlephalanxofright
ing scholar ofAfrican fossil carnivores. and left pes digit 4, middle phalanx of right and
KANAPOI MATERIAL. 10033, complete right left pes digit 5, sternebrae, fragments of cervical,
mandibular ramus with c-ml (Fig. 2A, Hyaena sp. thoracic, lumbar, and caudal vertebrae; 30306, left
in Behrensmeyer 1976; Pachycrocuta sp. inHowell distal femur fragment; 30463, rightmandibularra-
and Fetter 1980); 29249, right mandibular ramus mus fragment with roots of p2-p4; 30482, associ-
MC
fragmentwithp4; 29280,proximalfragmentofleft ated complete left III-V (Fig. 4); 30487, prox-
MT
radius; 29290, right mandibular ramus fragment imal part ofleft III; 30495, proximalfragment
MT MT
with p4, distal left radius fragment, right radius of II; 30534, proximal right III; 30536,
shaft fragment; 29293, left distal radius fragment; left p3; 30540, right 13; 30541, right lower canine;
29294, right mandibular ramus fragment with p2; 30544, left mandibular ramus fragment with p3
29296, right mandibular ramus fragmentwithpar- and ml, p2 and p4 crowns separate; 31734, prox-
Werdelin: Carnivorans 119
imal right ulna; 31735, proximal left ulna lacking mandible rises in a shallow S shape to the angular
olecranon; 32538, rightml; 32550, leftP3; 32552, process. The mandibular condyle is relatively slen-
mandible fragments with associated left c and p3- der in comparison with modern Hyaena and con-
ml; 32813, proximal right ulna fragment; 32822, sists oftwo semidistinct areas, a lateral, higher one
left lower canine; 32865, postcranial fragments in- and a medial, lower one. The latter does not taper
MC
cluding a fragment ofa proximal II, distal me- medially as is the case in Hyaena. The coronoid
tapodials, a distal humerus fragment, and vertebral process is tall and slender compared with that of
fragments. modern Hyaena, and has a distinct backward tilt.
MEASUREMENTS. See Tables 1 and 2 The anterior margin of the coronoid process is
The following is a composite description of the steeply S shaped. The masseteric fossa is deep and
material. The craniodental material shows limited flat. Theventralpart (insertionarea fortheM. mas-
variability, except in size, but where there is varia- seterintermedins) is delimited bya strongridgeand
tion, this is noted. There is limited duplication be- is set distinctly lateral to the more dorsal parts of
tweenpostcranial elements,whichallowsforavery the masseteric fossa (insertion area for the M. mas-
limited grasp of variation in the taxon, but does seter profundis). This is in contrast with the situa-
mean that a significant proportion of the skeleton tion in Hyaena, where the ridge separating these
of Parahyaena hoivelli is actually known (Fig. 5). two insertion areas is lower and less distinct and
Some comparisons with representative morpholo- the two areas are located inthe sameverticalplane.
gies of extant Hyaena hyaena are made. There is a single, large mental foramen situated be-
SKULL AND UPPER DENTITION. The skull is neath p2. In KP 10033, the symphysis is broken
represented only by some very small fragments and all the incisors lost. The lowercanineis broken
from KP 30272, which unfortunately are too small and chipped, but can be seen to have been a rela-
and damaged to provide any informationaboutthe tively robust tooth whose anteroposterior axis is
morphology ofthe species. This specimenhassome angled relative to the main axis of the ramus. The
very damaged teeth and tooth roots that indicate diastema is ofabout the same length as in Hyaena.
that the upper canine was slightly larger than the The cheek tooth row curves gently to buccal from
lower, that the PI was small and single rooted, and p2 to p3, then curves gently back from p4 to ml,
that P2 was considerably smaller than P3. All these much as in modern Hyaena. Unlike the latter,how-
features are normal in hyenas and the only fact of ever, the p2 of KP 10033 is set at an angle to the
interest is the presence ofPI. The 13 is represented main axis of the ramus.
by specimen KP 30540, which is worn, but shows The i2 is presentonly in KP 30235A. Itisheavily
the distinctive derived hyaenid subcaniniformmor- worn, but is clearly longer (anteroposteriorly) than
phology of this tooth. The P3 is also represented wide (mesiodistally). The tip is worn flat, though
by a damaged specimen, KP 32550. This tooth the wear facet is angled from distal (higher) to me-
probably had a small anterolingual accessorycusp, sial. On the mesial side, the wear has reached the
though damage in this area makes this somewhat enamel-dentine juncture. The lingual face is also
uncertain. The main cusp is high and conical, as is worn and there is an angle of about 70° between
typical of derived hyenas, and the posterior acces- the apical and buccal wear facets. The p2 is the
sory cusp is substantial, thoughpreciselyhowlarge most variable tooth in both size and shape. There
cannotbe determinedduetospecimendamage.The is no anterior accessory cusp, but a small swelling
tooth is similar to P3 of modern Hyaena, but the atthe anteriorendofthe toothispresent.Themain
main cusp is larger and stouter. The upper carnas- cusp is narrow and conical, with slightly convex
sial is represented by the isolated tooth KP 29302. anterior and posterior margins. The posterior ac-
It is typically hyaenid in morphology,withsubstan- cessory cusp is low and narrow. The posteriorshelf
tial parastyle and protocone, a large and relatively of this tooth is variable in width. In KP 10033, it
narrow paracone and a long metastyle. The par- is narrow in comparison with modern Hyaena,
astyle isrobustwitha distinctanteriorridgeleading which has a small shelf that is not present in this
down to an anterior cingulum with a hint ofa pre- Kanapoi specimen. In KP 30235A and KP 30544,
parastyle. The protocone is large but low and set p2 is much broader posteriorly and more similarto
in line with the anterior margin of the parastyle. the condition in Hyaena. In KP 29296, the p2, as
The paracone is tall and trenchant and the metas- judged from the alveolus, must have been notice-
tyle long. There is a lingual cingulum that reaches ably shorter than in the other specimens that pre-
fromthe posteriorrootoftheprotoconetothepos- serve traces of this tooth. The p3 has no anterior
terior quarter of the metastyle. accessory cusp. Instead, the anterior margin ofthe
MANDIBLE AND LOWER DENTITION. The main cusp is formed into a low crest that reaches
mandible and associated material are known from the anterior end ofthe tooth. The anteriorandpos-
sixteen specimens, most of which are fragmentary terior margins ofthe robust, conical main cusp are
and damaged, but include two nearly complete slightly convex. The posterioraccessorycuspis low
rami, KP 10033 and KP 30235A (Figs. 2A-B). The and round and setcentrally in a posteriorcingulum
mandible increases gradually in depth from anteri- shelf. The tooth is variable in size and shape,
or to posterior and is deepest just posterior to ml. though not to the extent seen in p2. KP 29301 has
Posterior to this point, the ventral margin of the a p3 that is nearly identical to that of KP 10033,
120 CS 498, Harris and Leakey: Kanapoi
Figure 2 A, Parahyaena howelli, KP 10033, right mandibular ramus in (top to bottom) buccal, lingual, and occlusal
view; B, Parahyaena howelli, KP 30235A, rightmandibular ramus in (top to bottom) lingual, buccal, andocclusalview
whereas in KP 30235A and KP 30544, the p3 is has a small, round anterior accessory cusp ap-
narrower and has a more distinct waist. KP 30536 pressed to a narrow, conical main cusp with more
and KP 32552 are intermediateinmorphology.The or less straight anterior and posterior margins. The
tooth is broadly similar to p3 in Hyaena exceptfor posterior accessory cusp is relatively high and tren-
the absence of an anterior accessory cusp. The p4 chant. The posterolingual partofp4 has beendam-
Werdelin: Carnivorans 121
Figure 2 Continued
aged and it is not possible to determine the width Theradius isknownfromKP30235 (Figs. 3A-B)
of the posterior part of the tooth. The lower car- and KP 29280. It is in general very similar to that
nassial is long and relatively low. The paraconid is of Hyaena, but is shorter for the same robusticity.
slightly longer and wider than the protoconid. The The grooves for the extensor digitis communis, ex-
metaconid is very small but is distinctly developed tensor carpi radialis, and abductor pollicis longus
and the talonid has two cusps, presumably the hy- are all more deeply incised than in Hyaena.
poconidandentoconid. ComparedwithmodernHy- The ulna is known from several fragmentary
aena, the tooth is relatively much longer, the meta- specimens, KP 30234, KP 30235, KP 31734, KP
conid smaller, and the talonid relatively shorter. 31735, and KP 32813. The first two are the most
FORELIMB. The humerus is known from KP complete and indicate that the ulna of this taxon
30235 (proximal and distal pieces) and KP 30234 was slightly shorter but more robust than that of
(shaft and distal articulation). The proximal frag- Hyaena. The shaft is more rounded than in Hyae-
ment is too worn for detailed comparisons with na, the triceps groove is wider, the ridge on thecra-
Hyaena, but is larger and appears relatively nar- nial surface of the olecranon is narrower, the at-
rower. The distal articulation is transversely broad- tachment area for the flexor carpi ulnaris is less
er than in modern Hyaena, but is relatively more distinct, andthepitbeneaththeradialnotchisshal-
slender anteroposteriorly. There is a large supra- lower. In addition, the rugosity for the articulation
trochlear foramen present in KP 30235. with the radius begins more proximally on the
122 CS 498, Harris and Leakey: Kanapoi
A
B
Figure 3 Parahyaena howelU, KP 30234, right radius in A. anterior and B, posteriorview
shaft, while the groove for the abductor pollicis for the flexor carpi radialis tendon is deeper and
longus is much more distinct than in Hyaena. boundedmedioventrallybyamoreprominentridge
Several of the carpals are known from different than in modern H. hyaena. The magnum is known
specimens. The scapholunar is known in specimens in KP 30234. It is shorter and wider than that of
KP 30234 and KP 30235AC. It is slightly larger modern H. hyaena butismorphologicallyverysim-
and more slender anteroposteriorly than that of ilar in all other respects. The unciform is also
modern H. hyaena. Specimen KP 30235AC is bro- known from KP 30234. It is more robustthanthat
ken on the medial side, but in KP 30234, it can be ofmodernH. hyaena andhasamoreopenarticular
seen that the articular face for the radius extends face for MC III. The pisiform is known from KP
down onto the medial rim, while in modern H. hy- 30234. It is larger than that ofmodern H. hyaena,
aena, there is a much more distinct ridge limiting but aside from size is practically indistinguishable
the radial articulation to the dorsal side of the from it.
bone. This may indicate greater mobility ofthis ar- All the metacarpals except MC II are known
ticulation in the fossil form. In addition, the sulcus from complete specimens. MC I, known from KP
Werdelin: Carnivorans 123
bust than that of modern H. hyaena. The two spe-
MC
cies are similar in their IVmorphology, butthe
proximal articulation in KP 30482 is somewhat
moretriangularinshape,witha broaderdorsaland
narrower plantar side than in the modern form.
The fifth metacarpal is also more robust and short-
er than that of modern H. hyaena, the difference
being more accentuated in this element. The prox-
MC
imal articular surface with IV is set less
obliquely and more directly anteroposteriorly than
in modern H. hyaena and is also very wide com-
pared with the condition in the modern species.
F4INDLIMB. The femur is represented by vari-
ous fragments from specimen KP 30272, including
a partial distal femur KP 30272N, proximal frag-
ments of specimen KP 30229, distal fragments,
specimen KP 30306, and proximal, shaftanddistal
fragments, KP 29299. These fragments suggest a
femur that is somewhat larger and more robust
than the corresponding element in modern H. hy-
aena, but otherwise do notshowanydistinguishing
features of note.
The tibia is known from fragments fromthe par-
tial skeletons KP 30235D and KP 30272P, Q, as
well as from KP 30229. In general, the tibia is not
a very diagnostic boneinFiyaenidae andthisistrue
in the presentcase aswell, the fossilspecimensonly
being distinguished from modern H. hyaena by
their greater size and by the slightly greater devel-
opment of the medial malleolus of KP 30272Q.
The navicular is known from KP 30235AE and
Figure 4 Parahyaena hoivelli, KP 30482, associated left AD, and KP 30272F. This bone is generally similar
MC III-V in dorsal view. Scale = 50 mm to that of modern H. hyaena, but is slightly larger.
It differs in that the plantar process is wider than
high, the reverse ofthe conditioninH. hyaena. The
process for the separation between the articulation
with the cuboid and the plantar side ofthe bone is
less prominent than in H. hyaena.
30235P, is a substantial element associated with at The lateral cuneiform is known only from KP
least one phalanx. This is corroborated by KP 30235BJ and BP. Itis largerthanthecorresponding
39235V, which is tentatively identified as the prox- bone in modern H. hyaena, apparently relatively
imal phalanx ofthe same digit. This phalanx bears more so than other tarsals and carpals reported
a large articular surface for an ungual phalanx, here. The proximal articular surface ofthe fossil is
which has not been identified in the material. It is more deeply indented laterallyandmediallythanin
far larger than that of any extant hyaenid, though H. hyaena and, in addition, the distal articular sur-
relatively smaller than its counterpartinIkelohyae- face (for MTIII) is concave in the fossil ratherthan
na ahronia Fiendey, 1974, from Langebaanweg, a slightly convex as in H. hyaena.
species that is otherwise smaller than the Kanapoi The cuboid is known from KP 30235W. It is the
form. The second metacarpal is known from a dis- tarsal that differs most from the corresponding el-
! tal and a proximal fragment, KP 30235AF4 and ement inmodern H. hyaena. The medial side ofthe
AO, respectively. This metacarpal is more robust proximal articular surface ofthe fossil has a medial
MC
than II in modern H. hyaena. Metacarpals III extension that probably buttressed the medial part
to V are associated in specimen KP 30482 (Fig. 4) of the sulcus for the M. peritoneus longus tendon.
and MC III is in addition known in KP 30235AG This sulcus is shallow and nondescript in modern
and KP 30235BH and m KP 30272. The third H. hyaena, deep and well developed in KP
metacarpal has larger proximal and distal articular 30235W. Thecuboid ofCrocutacrocuta (Erxleben,
MC
surfaces than III in modern H. hyaena, while 1777), on the other hand, is short and square with
the shaft is more robust but markedly shorterthan a deep, narrow sulcus.
in the extant species. The plantar side ofthe prox- The second metatarsal is known fromKP 30495,
imal articular surface is narrower relative to the 30235AM, and 30272AB. The two former are
MT
dorsal side than in the modern species. The fourth proximal ends that are more robust than II in
metacarpal is, like the third, shorter and more ro- modern H. hyaena but otherwise too worn for
124 CS 498, Harris and Leakey: Kanapoi
Figure 5 Farahyaena howelli, known skeletal parts of P. howelli (gray) superimposed on a skeleton of a dog; skeleton
adapted from Evans (1993)
meaningful comparisons to be made. The third the P4 metastyle is one feature that clearly distin-
specimen, KP 30272, is pathological in that the guishes all extant hyena species. Hyaena hyaena
bone appears to have been broken in life and sub- has a short P4 metastyle, ofabout the length ofthe
sequently healed. The distal end of this bone is paracone or slightly shorter. In C. crocuta, the me-
composed of an amorphous mass of secondary tastyle of P4 is exceptionally long and straight. In
bone suggestive of healing. Farahyaena brunnea, the metastyle of P4 is longer
The third metatarsal is known from specimens than that of H. hyaena but shorter than that of C.
KP 30272U and KP 30272AO, KP 30534, and KP crocuta. In the present case, the P4 metastyle has
30487. These specimens are more robust than the the relative length of that of F. brunnea. This con-
corresponding element in modern H. hyaena, but trasts with the condition in Ikelohyaena abronia, a
are otherwise similar except for the proximodorsal possible ancestor of H. hyaena (Fiendey, 1978a;
articular surface being concave rather than flat to Werdelin and Solounias, 1991), in which the P4
conTvheex.fourth metatarsal is known from KP motehtearstfyelaetuirsesshocrotn,traasdiicntitasspsuitgantmievnetdoefsctehnedKaannt.apNooi
30235AF and KP 30272T and AC. It differs from hyena to Farahyaena, and rather than positthe ex-
that ofH. hyaena only in its greater size and in the istence of a previously unknown hyaenid lineage, I
lesTsheexpfiafnthdemdetpatraorxsiamloipsaklnmoawrnemfirnoemncKeP.30235AK prefer to suggest a link to the living brown hyena.
and AL and KP 30272AA and KP 30272AP. Inthis Tcehsitsryreopfrtehseenbtsrotwhenfhiyrsetnad,ireacstalilndoitchaetrikonnoofwnthfeosasni-l
case, the greater size is the only clear difference
from extant striped hyaena. Farahyaena fit comfortably within the extant spe-
DISCUSSION. KP 10033 was recovered by the cies (e.g., Hendey, 1974).
American expeditions. It was referred to Hyaena
sp. by Behrensmeyer (1976) and to Pachycrocuta Family Felidae
sp. by Flowell and Petter (1980). The referral of
this species to Farahyaena rests chiefly on the Felids are common elements in the fossil faunas of
length of the metastyle of P4. This is only known eastern Africa. Both Machairodontinae and Felinae
from a single specimen, KP 29302, but can also be are present throughout the Plio-Pleistocene, butup
inferred from the length of the ml trigonid in re- to about 1.5 Ma, the former are by far the more
lation to p4 and ml talonid length. The length of common in the fossil record.