Early guenon from the late Miocene Baynunah Formation, Abu … · 2014. 7. 14. · Early guenon...

Early guenon from the late Miocene BaynunahFormation, Abu Dhabi, with implications forcercopithecoid biogeography and evolutionChristopher C. Gilberta,b,c,1, Faysal Bibid,e,f, Andrew Hillg, and Mark J. Beechh

aDepartment of Anthropology, Hunter College of the City University of New York, New York, NY 10065; bDepartments of Anthropology and Biology,Graduate Center of the City University of New York, New York, NY 10016; cNew York Consortium in Evolutionary Primatology, New York, NY; dMuseum fürNaturkunde, Leibniz Institute for Evolution and Biodiversity Science, 10115 Berlin, Germany; eDepartment of Mammalogy, American Museum of NaturalHistory, New York, NY 10024; fInstitut International de Paléoprimatologie, Paléontologie Humaine: Evolution et Paléoenvironnements, IPHEP UMR CNRS7262, Université de Poitiers, 86022 Poitiers Cedex, France; gDepartment of Anthropology, Yale University, New Haven, CT 06520-8277; and hCoastal Heritageand Palaeontology Section, Historic Environment Department, Abu Dhabi Tourism and Culture Authority, Abu Dhabi, United Arab Emirates

Edited by Leslea J. Hlusko, University of California, Berkeley, CA, and accepted by the Editorial Board June 2, 2014 (received for review December 21, 2013)

A newly discovered fossil monkey (AUH 1321) from the BaynunahFormation, Emirate of Abu Dhabi, United Arab Emirates, is im-portant in a number of distinct ways. At ∼6.5–8.0 Ma, it representsthe earliest knownmember of the primate subfamily Cercopithecinaefound outside of Africa, and it may also be the earliest cercopithecinein the fossil record. In addition, the fossil appears to represent theearliest member of the cercopithecine tribe Cercopithecini (gue-nons) to be found anywhere, adding between 2 and 3.5 million y(∼50–70%) to the previous first-appearance datum of the crownguenon clade. It is the only guenon—fossil or extant—known out-side the continent of Africa, and it is only the second fossil monkeyspecimen so far found in the whole of Arabia. This discovery sug-gests that identifiable crown guenons extend back into the Mio-cene epoch, thereby refuting hypotheses that they are a recentradiation first appearing in the Pliocene or Pleistocene. Finally, thenewmonkey is a member of a unique fauna that had dispersed fromAfrica and southern Asia into Arabia by this time, suggesting thatthe Arabian Peninsula was a potential filter for cross-continentalfaunal exchange. Thus, the presence of early cercopithecines onthe Arabian Peninsula during the late Miocene reinforces the prob-ability of a cercopithecoid dispersal route out of Africa throughsouthwest Asia before Messinian dispersal routes over the Medi-terranean Basin or Straits of Gibraltar.

cercopithecid | Old World monkeys | Bab el Mandeb | Mesopithecus | Sinai

Cercopithecine monkeys (Order Primates, Superfamily Cer-copithecoidea, Family Cercopithecidae, Subfamily Cercopi-thecinae), also known as cheek-pouch monkeys, are the mostspeciose and widely distributed group of living Old World pri-mates. Recent molecular estimates date the divergence of Cer-copithecinae from Colobinae (leaf-eating monkeys) to between17.6 Ma (range 21.5–13.9 Ma) and 14.5 Ma (range 16.2–12.8 Ma)and the origin of crown Cercopithecinae to around 11.5 Ma (range13.9–9.2 Ma) (1, 2). However, the earliest known fossil cercopi-thecines only appear much later, around 7.4 Ma in the TurkanaBasin of East Africa (3, 4).Cercopithecine monkeys are divided into two tribes: Cerco-

pithecini, including African guenons (Allenopithecus, Miopithecus,Chlorocebus, Erythrocebus, Allochrocebus, Cercopithecus), and Papio-nini, which includes African and Eurasian macaques (Macaca) aswell as African papionins (Papio, Lophocebus, Rungwecebus,Theropithecus, Mandrillus, Cercocebus). Of the living cercopithe-cines, only two genera are known outside of the African continent,both of them papionins: Papio (found on the Arabian Peninsula)andMacaca (found throughout Southern and Southeast Asia, andintroduced inGibraltar). The earliest fossil cercopithecines knownoutside ofAfrica are attributed to the genusMacaca and appear tobe latest Miocene or early Pliocene in age (∼6.0–5.0 Ma) (Fig. 1)

(5–9). Until now, no guenons, extant or extinct, have ever beenknown outside of the African continent.Three possible routes can be reasonably hypothesized for

cercopithecine (and cercopithecoid) dispersal out of Africa andinto Europe and Asia during the late Miocene: (i) over theMediterranean Basin or Straits of Gibraltar to the north/north-west, (ii) across the Arabian Sinai Peninsula to the northeast, or(iii) across the Arabian Straits of Bab el Mandeb to the east (Fig.1). Fossil Macaca specimens from the terminal Miocene of Spainand Italy have been suggested to provide evidence for the use ofa route across the Mediterranean Basin or the Straits of Gibral-tar via an ephemeral land bridge either immediately before—orperhaps associated with—the drop in Mediterranean sea levelsduring the Messinian (∼6.0–5.3 Ma) (8–19). Paleontologicalevidence for an Arabian route has been lacking, but paleoge-ographic and paleoenvironmental work on circum-Arabia sug-gests that the region did not present a persistent ecologicalbarrier to some amount of intercontinental exchange during thelate Miocene (20). In fact, an established land connection throughSinai was probably present during this time period, and oceanicspreading is not estimated to have begun in the southern Red Seauntil around 5 Ma, with progressive development of open marineconditions throughout the Pliocene. Thus, before 6.5 Ma, a southern

Significance

The primate subfamily Cercopithecinae represents the mostdiverse and successful living Old World primate group, witha current distribution throughout Africa and Asia. However,how and when these monkeys dispersed out of Africa is notwell understood. This paper is significant in its description ofa ∼6.5–8.0 million-y-old fossil guenon from Arabia represent-ing the earliest cercopithecine (and only guenon) yet knownoutside of Africa. Furthermore, this specimen extends theguenon fossil record by at least 2.3 million y and may representthe earliest known cercopithecine as well. Because Old Worldmonkeys appear to have dispersed out of Africa sometimeduring the Late Miocene, the Arabian fossils also have impli-cations for dispersal scenarios in Old World monkey bio-geography and evolution.

Author contributions: C.C.G., F.B., A.H., and M.J.B. designed research; C.C.G., F.B., A.H.,and M.J.B. performed research; C.C.G., F.B., and A.H. contributed new reagents/analytictools; C.C.G., F.B., and A.H. analyzed data; and C.C.G., F.B., and A.H. wrote the paper.

The authors declare no conflict of interest.

This article is a PNAS Direct Submission. L.J.H. is a guest editor invited by the EditorialBoard.1To whom correspondence should be addressed. E-mail: [email protected].

This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental.

www.pnas.org/cgi/doi/10.1073/pnas.1323888111 PNAS Early Edition | 1 of 6

ANTH

ROPO

LOGY

http://crossmark.crossref.org/dialog/?doi=10.1073/pnas.1323888111&domain=pdf&date_stamp=2014-06-25mailto:[email protected]://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplementalhttp://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplementalwww.pnas.org/cgi/doi/10.1073/pnas.1323888111

route in the region of the Straits of Bab el Mandeb was also possible(Fig. 1) (21).Although Arabia is a large area of the earth, fossil monkeys

have so far been represented by only a single specimen, an iso-lated male lower canine (AUH 35), discovered in 1989 by A.H.and Peter Whybrow in the late Miocene Baynunah Formation,Abu Dhabi, United Arab Emirates (22–25). The specimen camefrom Jebel Dhanna, site JDH-3 (JD-3 in refs. 24 and 26) (Fig. 2),a locality now lost to industrial development. Because malecercopithecid lower canines are not metrically identifiable be-yond the Family level of classification (23), AUH 35 was de-scribed as a cercopithecid with indeterminate affinities. Here wereport the discovery of a second monkey specimen from theBaynunah Formation in Abu Dhabi (AUH 1321), found almost20 y after the first. AUH 1321 clearly represents a cercopithecineand, because it is dated to between 6.5 and 8.0 Ma, it is the oldestcercopithecine yet known outside of Africa and possibly theoldest cercopithecine in the fossil record. Thus, the discovery ofAUH 1321 provides the earliest paleontological evidence ofcercopithecine dispersal out of continental Africa and possiblyhints at an Arabian cercopithecoid dispersal route into Eurasiaduring the Late Miocene (Fig. 1). Furthermore, we believe AUH1321 can be attributed to the Cercopithecini (guenons) and,therefore, it represents the only record of this tribe, living orfossil, yet known outside of Africa.

Geological and Paleoenvironmental ContextThe Baynunah Formation (27) is a sequence of sandstones andmudstones that are mainly fluvial in origin and that containfossils at several levels. Up to some 60 m thick, the Formation isexposed along more than 200 km of the Abu Dhabi coast in thewestern Al Gharbia region, and extends more than 30 km inland.From 1988 to 1995 a joint research project conducted by A.H.

and Peter Whybrow recovered a diverse fossil fauna from the

Baynunah Formation (24). More recently, a team led by F.B.,A.H., and M.J.B. conducted annual field expeditions from 2006 to2011 (28–30). The Baynunah sediments indicate the presence ofa large meandering river system, with flanking woodlands, pos-sibly tapering off into grasslands and even desert laterally (31).The accompanying fauna is unique, but largely represented bya typical African wooded grassland assemblage with aquaticelements, including hippopotamids, bovids, giraffids, suids, a rangeof carnivores including a mustelid, a large felid, and two hyaenids,equids, a number of rodents, an insectivore, and no less than threeproboscidean taxa. The Baynunah rivers were well populated withdiverse fish, turtles, crocodiles, gavials, and molluscs, and there areinteresting birds and land reptiles as well. For a full current faunallist, see refs. 29 and 30.

Fig. 1. Hypothesized cercopithecoid dispersal routes out of Africa in relation to the known late Miocene fossil record. The oldest cercopithecine, Parapapiolothagamensis (light blue circles), is known from ∼7.4–6.1 Ma in the Turkana Basin and Tugen Hills, Kenya (3, 4, 41). An unnamed fossil papionin (purple circle)is known from the late Miocene of Ongoliba, Congo (5, 57). Macaca spp. (dark blue circles) are located throughout North Africa at sites ranging in age from∼6.5–5.5 Ma (5, 8, 58, 59), and Macaca spp. first appear in Europe ∼6.0–5.3 Ma and in China in the early Pliocene (5–9). The oldest colobine outside of Africa,Mesopithecus (green circles), is known from a number of late Miocene sites securely dated between ∼8.5 and 5.3 Ma in Greece, Macedonia, Italy, Ukraine,Iran, Afghanistan, possibly Pakistan, and China (46–48). Three dispersal routes for cercopithecoids can be hypothesized: route 1 imagines a dispersal eventover the Straits of Gibraltar or Mediterranean Basin into Europe and Asia; route 2 postulates a dispersal event through the Arabian Sinai Peninsula; and route3 suggests a migration over the Arabian Straits of Bab el Mandeb. The discovery of AUH 1321 and AUH 35 in Abu Dhabi at >6.5–8 Ma (red circle), con-temporaneous with the first appearance of Mesopithecus sp. in Eurasia and ∼1–2 million y earlier than the appearance of Macaca spp. in Eurasia, suggestsscenarios 2 and 3 were possible before scenario 1. None of these scenarios is mutually exclusive and may have occurred in combination or succession.

Fig. 2. Map illustrating the location of the two fossil sites in the BaynunahFormation that have produced fossil monkeys. Top Right Inset shows the lo-cation of the SHU 2–2 excavation (kite aerial photography by Nathan Craig).

2 of 6 | www.pnas.org/cgi/doi/10.1073/pnas.1323888111 Gilbert et al.

www.pnas.org/cgi/doi/10.1073/pnas.1323888111

Overall, the Baynunah fauna is taxonomically most similar tocontemporaneous sites in Libya, Chad, Ethiopia, and Kenya, andvery different from those in Greece, Turkey, and Iran (e.g., refs. 24,29, and 30). Most Baynunah fossil taxa appear African in originand are not known in Asia. However, there are some southernAsian forms (i.e., Siwaliks) that are never recorded in Africa, suchas Propotamochoerus hysudricus (32) and Pachyportax latidens,implying that the Arabian late Miocene fauna was not merelya biogeographic extension of Africa, but was a unique combinationof southern Asian and African elements, possibly reflecting anintermittent and not totally permeable corridor of faunal exchangebetween these neighboring continents. Very few Baynunah speciesare shared with Europe and Asia north of the Himalayan-Zagros-Tauride mountain belt, although Prostrepsiceros aff. vinayaki andpossibly some rodents may be examples (33, 34). Thus, given theevidence at hand, Arabia in the late Miocene appears to haveserved as a cross-continental corridor for only a few taxa, but for themajority it was a filter through which only those taxa with suitableadaptations were occasionally able to disperse into, although notalways beyond.The fossil tooth AUH 1321 was found in 2009 at site SHU 2-2

[24.1148N 52.4320E (WGS84)], a sublocality of SHU 2 (S2 inrefs. 24 and 26) on the island of Shuwaihat (Fig. 2). The fossilcame from a small area (∼4 × 4 m) of sedimentary exposure thathas produced a large number of microvertebrate fossils throughdry sieving. The specimen was spotted by F.B. on a surface thathad been scooped and sieved the previous day. The color andrelatively good preservation of the specimen—in a coastal areawhere sun, humidity, gypsum, and halite quickly attack fossilspecimens as they approach the surface—attests to very recentsurface exposure and negligible postexposure transport. Site SHU2-2 has also produced a diverse microvertebrate fauna, includingnew specimens of the murid rodent Abudhabia baynunensis (35),a new thryonomyid rodent, Protohummus dango (33), and severalfish and squamate taxa currently under study.

Geological AgeRadiometric dates are not available for the Baynunah Forma-tion, and paleomagnetic estimates are not well resolved (22, 36).Biochronological comparisons with African and Asian faunas in-dicate an age between 8 Ma and 6.5 Ma, and most probably around7 Ma (29, 30, 37, 38). The upper limit is suggested by the suidP. hysudricus (32), which is not known from the Siwaliks in Pakistanlater than 6.8 Ma, with an estimated last appearance datum of 6.5Ma (39). The presence of the hippopotamid Archaeopotamus andthe ratite Diamantornis laini in both the Baynunah and the LowerMember of the Nawata Formation (7.4–6.54 Ma) at Lothagam,Kenya also indicates an age greater than 6.5 Ma (38, 40, 41). Thelower limit is not so well resolved, but is unlikely to be greater than8 Ma. This age estimate puts the Baynunah specimens in the timerange of the earliest known cercopithecine fossils in Africa, allfrom Kenya: Parapapio lothagamensis, known to occur in the LowerNawata Formation at Lothagam (3, 4) and also at around 6.1 Main the Lukeino Formation, Tugen Hills (42).

Description and Comparison of AUH 1321AUH 1321 is a small lower left molar, most likely an M1, pre-serving the majority of the crown and the distal root; the mesialroot is broken off (Fig. 3 and Fig. S1). The mesial and distallophids are complete and transversely oriented with the distallophid being slightly wider than the mesial lophid. Comparisonswith extant and late Miocene cercopithecoids are given in Table 1and Tables S1 and S2. Given available M1 crown measure-ments, AUH 1321 represents a small cercopithecine, similar insize to extant vervet monkeys or blue monkeys, (estimated bodymass ∼3.9–5.8 kg, based on cercopithecine regressions of femaleM1 mesial buccolingual width and male M1 mesiodistal lengthfrom ref. 7), but much smaller than any known cercopithecine

from the late Miocene of Africa or the Miocene-Pliocene ofEurasia (Table 2).The crown of AUH 1321 is moderately worn and in good

condition overall, although it is slightly damaged and abraded inplaces (Fig. 3 and Fig. S1). The crown is high relative to the lowcusps, as is typical of cercopithecines (Fig. 3, Fig. S1, and TableS1). A mesial contact facet is present at the mesial end of thetooth. The mesial shelf is relatively elongated and slightly largerthan the distal shelf/fovea, another feature shared with extantcercopithecines (43). The median buccal cleft is relatively wideand of moderate depth, not nearly as deep as typically seen incolobines. There is a very small and shallow mesial buccal cleftpresent, but no visible distal buccal cleft. However, because thedistobuccal portion of the enamel is damaged, it is impossible tosay with certainty whether a distal buccal cleft or distal contactfacet was present or absent (Fig. 3 and Fig. S1).The tooth is relatively narrow, as shown by the mesiodistal

length divided by the mesial or distal width (Table 1), but not asnarrow as is typically seen in cercopithecine dP4s, confirmingthat the specimen most likely represents an adult molar (TablesS1 and S2). The preserved distal root is moderate in length androbust at its proximal and distal ends, also indicating a perma-nent rather than deciduous tooth. The distal root appears slightlybroken at the apex, exposing a small opening. An alternativeexplanation might be that the root’s apex was still slightly open;however, given the moderate wear of the tooth combined with an

Fig. 3. Photographs of AUH 1321 in (A) occlusal, (B) mesial, (C) distal, (D)buccal, and (E) lingual views. Mesial is to the right in A and E; mesial is to theleft in D. (Scale bars, ∼1 mm) (photos by Eric Lazo-Wasem).

Gilbert et al. PNAS Early Edition | 3 of 6

ANTH

ROPO

LOGY

http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST2http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST2

irregular shaped opening (Fig. 3 and Fig. S1), we believe that theformer is more likely. In either case, the distal root would havebeen even longer than is preserved, and long robust roots with nosign of lingual resorption are typical of permanent rather thandeciduous teeth (Figs. S2 and S3 and Tables S2 and S3). Al-though the distal root of AUH 1321 is angled posteriorly, rem-iniscent of a deciduous premolar, it is within the variation seenamong M1s of living cercopithecines, and a discriminant func-tions analysis of five crown and root shape variables demon-strates that AUH 1321 is most likely an M1 rather than a dP4,with a >99% probability (more specifically a guenon M1, witha >97% probability) (Figs. S2 and S3 and Tables S1–S3).AUH 1321 displays low-to-moderate flare, in contrast to most

papionin permanent molars and cercopithecine dP4s, and ismore similar to the condition seen in the permanent molars ofcolobines, some extant macaques, and some cercopithecin taxa(Table 1 and Tables S1–S3) (44, 45). The overall size and shapeof the tooth—small, narrow, low-to-moderately flaring, elon-gated basin, and a distally expanded lophid—is differentiatedfrom extant small-bodied and fossil macaques and is most similarto non-Allenopithecus cercopithecins among extant cercopithe-cine taxa. A phylogenetic analysis of 13 dental characters con-firms that AUH 1321 is most similar to the tribe Cercopitheciniamong extant cercopithecine taxa (Fig. S4 and Tables S4 andS5). Thus, we assign AUH 1321 to Cercopithecini sp. indet untilmore material becomes available. Although it appears that AUH1321 most probably represents a new taxon, we refrain fromnaming a new genus and species in the absence of a more di-agnostic type specimen. The size of the lower male canine AUH35, the only other fossil monkey known from Abu Dhabi, is alsosmaller than any known fossil cercopithecine from North Africaand within the range of living guenons, being closest in size toCercopithecus mitis among extant Cercopithecus taxa and makingit possible that AUH 1321 and AUH 35 represent the same

taxon (Table S6). However, AUH 35 is also within the size rangeof modern macaques and the fossil colobine Mesopithecus,making it equally possible that multiple cercopithecoid taxa andsubfamilies are being sampled (Table S6).

DiscussionAlthough sparse, the cercopithecoids of the Baynunah Forma-tion in Arabia are both chronologically and biogeographicallyhighly significant. AUH 1321 represents the earliest appearanceof the subfamily Cercopithecinae outside of Africa and, together,AUH 1321 and AUH 35 provide tantalizing evidence for a po-tential primate dispersal route through Arabia and into southernAsia and Europe during the latest Miocene (Fig. 1). Althoughguenons do not appear to have ever dispersed into Europe andAsia, it is clear that other cercopithecines (and colobines) did,and the presence of two monkey specimens in the Late Mioceneof Arabia suggests that Arabia provided suitable environments forcercopithecoid primates dispersing out of Africa (see reconstructedwoodlands, above) (23, 29–30). Thus, although the appearance ofthese monkeys in the Arabian Peninsula does not refute a Gibraltaror Mediterranean dispersal route for cercopithecoids during theMessinian Crisis, it does suggest that an Arabian dispersal route forOld World monkeys was possible earlier, in pre-Messinian times(Fig. 1). In fact, the earliest securely dated cercopithecoids knownfrom Europe and Asia are late Miocene specimens placed in thecolobine genus Mesopithecus, ∼8.5–5.3 Ma (46–48), roughly con-temporaneous with the Baynunah Formation and consistent witha pre-Messinian route through Arabia in the Late Miocene (Fig. 1).Because AUH 35 is within the size range ofMesopithecus pentelicuslower canines (Table S6), it is possible that the Baynunah Formationcaptures a snapshot of both cercopithecines and colobines exitingAfrica during the late Miocene. Additional evidence of limited lateMiocene cross-continental faunal exchange between Africa, Arabia,and Asia may also be provided by the bovid Prostrepsiceros cf.

Table 1. Selected comparative measurements (mm) of the dentition in the Abu Dhabi cercopithecines and other Old World monkeys

Taxon Source Specimen no. M1 MD M1 MBL M1 DBL M1 NH/NR M1 FlareM1

(MD/MBL)M1

(MD/DBL)M1

(MBL/DBL)

Cercopithecinisp. indet.

Present study AUH 1321 6.2 4.5 4.6 67 66 138 135 98

Late Miocenefossil macaquesMacaca libyca (59) Average 7.6 7.5 7.3 — 53* 101.0 103.8 102.7Macaca sp. Menacer (59) Average 9.1 7.6 6.8 — 65* 120.5 133.8 111.0Macaca sp. Almenara (8) IPMC 11676 8.0 6.2 6.1 — — 129.0 131.1 101.6As Sahabi (59) Average 7.6 6.4 6.3 — 51* 119.0 121.2 101.9

(8, 59) Fossil macaqueaverage

8.1 6.9 6.6 — 57 117 122 104

(8, 59) Fossil macaquespecies range

7.6–9.1 6.2–7.6 6.1–7.3 — 51–65 101–129 104–134 102–111

Extant macaquesMacaca (45, 60) Extant macaque

average7.0 5.4 5.3 49 68 129 130 101

(45, 60) Extant macaquespecies range

6.4–7.5 4.9–5.8 4.7–5.8 44–61 66–71 120–137 119–140 98–104

Extant guenonsCercopithecus/Chlorocebus

(60, PRIMO,present study)

Extant guenonaverage

5.6 4.0 4.2 61 61 139 133 96


Extant guenonspecies range

5.0–6.3 3.6–4.4 3.8–4.7 39–77 55–63 130–144 130–137 89–100

In addition to measurements collected during the course of this study, comparative measurements are derived from refs. 8, 43, 45, 59, and 60, as well asfrom the PRIMO (Primate Morphology) database, access courtesy of E. Delson. —, unavailable measurement; *, estimated from M2 values in ref. 59; DBL,maximum distal buccolingual width measured across the distal lophid; MBL, maximum mesial buccolingual width measured across the mesial lophid; MD,maximum mesiodistal length; NH/NR, notch height/crown height below the notch [see Benefit (43, 45, 61) for details]; species range, range of values amongspecies averages within a given group. Flare measured as in Benefit (43, 45, 61) and lower values indicate higher degrees of flare. Note that AUH 1321 is mostsimilar to extant small-bodied guenons in M1 size and shape. See also Table 2 and Fig. S4. For a more complete list of taxa, see Tables S1–S3.


http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF2http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF3http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST2http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST3http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF2http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF3http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST3http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST3http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF4http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST4http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST5http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST6http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST6http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST6http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF4http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST1http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST3www.pnas.org/cgi/doi/10.1073/pnas.1323888111

vinayaki, the fish Clarias sp., the carnivore genus Plesiogulo, and therodent genera Myocricetodon and Abudhabia (30, 33–35).A phylogenetic analysis of 13 cercopithecine dental characters

demonstrates the affinities of AUH 1321 to Cercopithecini, es-tablishing its position as the earliest known guenon in the fossilrecord by at least 2.3 million y (49), and the only guenon yetknown outside of Africa (Fig. S4 and Tables S4 and S5). On bothmolecular and morphological grounds, nearly all authorities rec-ognize Allenopithecus as the most basal extant member of theCercopithecini (e.g., refs. 1, 44, 50–54). Allenopithecus is primi-tive relative to other cercopithecin taxa in that it retains basallyflaring molars, a feature shared with both papionins and moreprimitive cercopithecoids, such as Victoriapithecus. In contrast,AUH 1321 clearly displays relatively nonflaring molars, a distinctivefeature shared exclusively with non-Allenopithecus guenons amongmodern cercopithecins. Thus, the available morphological evi-dence suggests that AUH 1321 is a crown guenon nested withinthe modern non-Allenopithecus radiation (Fig. S4). Among non-Allenopithecus guenons, molecular estimates place the divergencebetween the terrestrial Chlorocebus/Erythrocebus/Allochrocebusclade and the arboreal Cercopithecus clade between 8.8 and7.6 Ma (55), making it possible that AUH 1321 is an earlymember of one of these two clades as suggested by our phylo-genetic analysis of cercopithecine dental characters (Fig. S4).In any case, the fact that the Baynunah Formation predates theestimated appearance of the Cercopithecus common ancestor (4.9–4.3 Ma) and the Chlorocebus/Erythrocebus/Allochrocebus commonancestor (5.5–5.0 Ma) (55) suggests that the Arabian specimenbelongs to a distinct genus. In addition, to our knowledge, it

provides the first paleontological evidence refuting the hypoth-esis that the crown guenons are only a recent Pliocene-Pleis-tocene radiation (56).The discovery of AUH 1321, in addition to AUH 35, em-

phasizes the problem of determining the existence of rare taxa ingeographically isolated fossil situations, and making paleoge-ographical inferences in the absence of more comprehensivefossil data. If it were not for these two teeth we would not knowthat monkeys existed in the late Miocene of Arabia. Perhaps forsimilar reasons we are currently ignorant of other possible im-portant yet rare elements of the fauna, including other primates(e.g., other monkeys, apes, and early hominins). Further sus-tained paleontological exploration of Arabia and northern Africais necessary if we are to clarify primate and mammalian evolu-tion and biogeography during the late Miocene.

ACKNOWLEDGMENTS. We thank the American Museum of Natural Historyfor the use of CT Scan imaging equipment; Eric Delson, Steve Frost, JohnFleagle, Peter Ungar, and Mike Plavcan for their assistance and advice; SarahElton, two anonymous reviewers, and the editor for their helpful comments;Eileen Westwig for access to primate specimens at the American Museum ofNatural History; Darrin Lunde and Nicole Edmison for access to primatespecimens at the Smithsonian National Museum of Natural History; AndreaBaden for help with Fig. 1; Eric Lazo-Wasem for the photographs formingFig. 3; Emily Goble for biostratigraphical discussions; and B. Kraatz,E. Moacdieh, M. Al Neyadi, and staff from the Historic Environment De-partment at the Abu Dhabi Tourism and Culture Authority, who participatedin fieldwork efforts. Our research in Abu Dhabi has been supported by theHistoric Environment Department of the Abu Dhabi Tourism and CultureAuthority (formerly Abu Dhabi Authority for Culture and Heritage); anInternational Research Fellowship Award (National Science FoundationGrant 0852975) (to F.B); the Revealing Hominid Origins Initiative (NationalScience Foundation Grant BCS-0321893); the Institute International de

Table 2. Body mass estimates (kg) for selected extant and fossil cercopithecine taxa

Taxon Average male mass Average female mass

Cercopithecini sp. indet.AUH 1321 5.8 3.9

Late Miocene/Plio-Pleistocene CercopithecinesParapapio lothagamensis 10.5 8.0Papionin sp. indet., Ongoliba 9.5 6.5Macaca libyca 10.0 9.5As Sahabi papionin 11.0 8.5Macaca sp., Menacer 10.0 7.0Macaca sp., Almenara 11.5 8.0cf Macaca sp., Moncucco 12.0 8.0Macaca majori 9.5 6.0Macaca sylvanus ?prisca 12.0 —Macaca sylvanus ?florentina 13.0 10.5Macaca palaeindica 13.0 —Macaca sylvanus ?pliocena 14.0 7.5Macaca anderssoni 14.0–17.0 10.5Procynocephalus subhimalayanus 13.0 24.0Paradolichopithecus avernensis 31.0 19.0Paradolichopithecus sushkini 36.0 35.0

Extant small cercopithecinesCercopithecus cephus 3.8 2.7Macaca fascicularis 5.0 3.0Chlorocebus aethiops 5.1 3.6Cercopithecus ascanius 5.5 2.5Macaca mulatta 6.2 —Cercopithecus neglectus 6.9 4.2Macaca nigra 7.5 4.7Cercopithecus mitis 8.0 3.9Macaca nemestrina 8.0–11.5 5.0–6.5Macaca sylvanus 15.0 10.0

Fossil cercopithecine taxonomy and body mass estimates from Delson et al. (7) or derived from equationstherein. Note that AUH 1321 is much smaller than any known Late Miocene cercopithecine and most similar insize to small-bodied guenons among living taxa. Taxa in bold are most similar in size to AUH 1321.

Gilbert et al. PNAS Early Edition | 5 of 6

ANTH

ROPO

LOGY

http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF4http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST4http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=ST5http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF4http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1323888111/-/DCSupplemental/pnas.201323888SI.pdf?targetid=nameddest=SF4

Paléoprimatologie et Paléontologie humaine; the Agence Nationale de laRecherche (ANR-09-BLAN-0238); the Yale Peabody Museum; the Richard

Gilder Graduate School and the Gerstner Scholars Program; and a grant fromthe Yale University President’s Office (to A.H.).

1. Perelman P, et al. (2011) A molecular phylogeny of living primates. PLoS Genet 7(3):e1001342.

2. Ting N (2008) Mitochondrial relationships and divergence dates of the African colobines:Evidence of Miocene origins for the living colobus monkeys. J Hum Evol 55(2):312–325.

3. McDougall I, Feibel CS (2003) in Lothagam: the Dawn of Humanity in Eastern Africa,eds Leakey MG, Harris JM (Columbia Univ Press, New York), pp 43–64.

4. Leakey MG, Teaford MF, Ward CV (2003) in Lothagam: The Dawn of Humanity inEastern Africa, eds Leakey MG, Harris JM (Columbia Univ Press, New York), pp 201–248.

5. Delson E (1980) in The Macaques: Studies in Ecology, Behavior and Evolution, edLindburg DG (Van Nostrand, New York), pp 10–30.

6. Delson E (1996) in Abstracts, International Symposium: Evolution of Asian Primates,ed Takenaka O (Primate Research Institute, Inuyama, Japan), p 40.

7. Delson E, et al. (2000) Body mass in Cercopithecidae (Primates, Mammalia): Estimationand scaling in extinct and extant taxa. Anthropol Pap Am Mus Nat Hist 83:1–159.

8. Köhler M, Moyà-Solà S, Alba DM (2000) Macaca (Primates, Cercopithecidae) from thelate Miocene of Spain. J Hum Evol 38(3):447–452.

9. Alba DM, et al. (2014) First joint record of Mesopithecus and cf. Macaca in the Mio-cene of Europe. J Hum Evol 67:1–18.

10. Delson E (1975) in Approaches to Primate Paleobiology. Volume 5: Contributions toPrimatology, ed Szalay F (Karger, Basel), pp 167–217.

11. Gautier F, Clauzon G, Suc J-P, Cravatte J, Violenti D (1994) Age et durée de la crise desalinité messinienne. CR Acad Sci II A 318:1103–1109.

12. Clauzon G, Suc J-P, Gautier F, Berger A, Loutre M-F (1996) Alternate interpretation ofthe Messinian salinity crisis: Controversy resolved? Geology 24(4):363–366.

13. Krijgsman W, Hilgen FJ, Raffi I, Sierro FJ, Wilsonk DS (1999) Chronology, causes andprogression of the Messinian salinity crisis. Nature 400:652–655.

14. McKenzie JA (1999) From desert to deluge in the Mediterranean. Nature 400:613–614.

15. Sierro FJ, et al. (1999) Messinian pre-evaporite sapropels and precession induced os-cillations in western Mediterranean climate. Mar Geol 153:137–146.

16. Van Couvering JA, Castradori D, Cita MB, Hilgen FJ, Rio D (2000) The base of theZanclean Stage and of the Pliocene Series. Episodes 23(3):179–187.

17. Lourens LJ, Hilgen FJ, Laskar J, Shackleton NJ, Wilson DS (2004) in A Geologic Time-scale 2004, eds Gradstein F, Ogg J, Smith A (Cambridge Univ Press, Cambridge), pp409–440.

18. Rouchy J-M, Caruso A (2006) The Messinian salinity crisis in the Mediterranean basin:A reassessment of the data and an integrated scenario. Sediment Geol 188–189:35–67.

19. Gibert L, et al. (2013) Evidence for an Africa-Iberian mammal dispersal during the pre-evaporitic Messinian. Geology 41(6):691–694.

20. Kingston JD, Hill A (1999) Late Miocene palaeoenvironments in Arabia: a synthesis.Fossil Vertebrates of Arabia: With Emphasis on the Late Miocene Faunas, Geology,and Palaeoenvironments of the Emirate of Abu Dhabi, eds Whybrow PJ, Hill A (YaleUniv Press, New Haven, CT), pp 389–407.

21. Bosworth W, Huchon P, McClay K (2005) The Red Sea and Gulf of Aden Basins. J AfrEarth Sci 43:334–378.

22. Whybrow PJ, Hill A, Yasin W, Hailwood E (1990) Late Miocene primate fauna, floraand initial paleomagnetic data from the Emirate of Abu Dhabi, United Arab Emirates.J Hum Evol 19(4-5):583–588.

23. Hill A, Gundling T (1999) A monkey (Primates, Cercopithecidae) from the Late Mio-cene of Abu Dhabi, United Arab Emirates. Fossil Vertebrates of Arabia: With Emphasison the Late Miocene Faunas, Geology, and Palaeoenvironments of the Emirate ofAbu Dhabi, eds Whybrow PJ, Hill A (Yale Univ Press, New Haven, CT), pp 198–202.

24. Whybrow PJ, Hill A, eds (1999) Fossil Vertebrates of Arabia: With Emphasis on theLate Miocene Faunas, Geology, and Palaeoenvironments of the Emirate of AbuDhabi, United Arab Emirates (Yale University Press, New Haven and London), 566 pp.

25. Gee H (1989) Fossils from the Miocene of Abu Dhabi. Nature 338:704.26. Whybrow PJ, Clements D (1999) Late Miocene Baynunah Formation, Emirate of Abu

Dhabi, United Arab Emirates. Fossil Vertebrates of Arabia: With Emphasis on the LateMiocene Faunas, Geology, and Palaeoenvironments of the Emirate of Abu Dhabi, edsWhybrow PJ, Hill A (Yale Univ Press, New Haven, CT), pp 317–333.

27. Whybrow PJ (1989) New stratotype; The Baynunah Formation (Late Miocene), UnitedArab Emirates: Lithology and palaeontology. Newsl Stratigr 21:1–9.

28. Bibi F, et al. (2012) Early evidence for complex social structure in Proboscidea froma late Miocene trackway site in the United Arab Emirates. Biol Lett 8(4):670–673.

29. Hill A, Bibi F, Beech M, Yasin al-Tikriti W (2012) in Fifty Years of Emirates Archaeology,eds Potts D, Hellyer P (Ministry of Culture, Youth and Community Development, AbuDhabi), pp 20–33.

30. Bibi F, Hill A, Beech M, Yasin W (2013) in Neogene Terrestrial Mammalian Bio-stratigraphy and Chronology in Asia, eds Wang X, Fortelius M, Flynn LJ (ColumbiaUniv Press, New York), pp 583–594.

31. Friend PF (1999) Rivers of the Lower Baynunah Formation, Emirate of Abu Dhabi,United Arab Emirates. Fossil Vertebrates of Arabia: With Emphasis on the Late Mio-cene Faunas, Geology, and Palaeoenvironments of the Emirate of Abu Dhabi, edsWhybrow PJ, Hill A (Yale Univ Press, New Haven, Ct), pp 39–49.

32. Bishop L, Hill A (1999) Fossil Suidae from the Baynunah Formation, Emirate of AbuDhabi, United Arab Emirates. Fossil Vertebrates of Arabia: With Emphasis on the Late

Miocene Faunas, Geology, and Palaeoenvironments of the Emirate of Abu Dhabi, edsWhybrow PJ, Hill A (Yale Univ Press, New Haven, CT), pp 254–270.

33. Bibi F (2011) Mio-pliocene faunal exchanges and African biogeography: The record offossil bovids. PLoS ONE 6(2):e16688.

34. Kraatz B, Bibi F, Hill A, Beech M (2013) A new fossil thryonomyid from the LateMiocene of the United Arab Emirates and the origin of African cane rats. Natur-wissenschaften 100(5):437–449.

35. de Bruijn H (1999) A late Miocene insectivore and rodent fauna from the BaynunahFormation, Emirate of Abu Dhabi, United Arab Emirates. Fossil Vertebrates of Arabia:With Emphasis on the Late Miocene Faunas, Geology, and Palaeoenvironments of theEmirate of Abu Dhabi, eds Whybrow PJ, Hill A (Yale Univ Press, New Haven, CT), pp186–197.

36. Hailwood EA, Whybrow PJ (1999) Palaeomagnetic correlation and dating of the Bay-nunah and Shuwaihat Formations, Emirate of Abu Dhabi, United Arab Emirates. FossilVertebrates of Arabia: With Emphasis on the Late Miocene Faunas, Geology, andPalaeoenvironments of the Emirate of Abu Dhabi, eds Whybrow PJ, Hill A (Yale UnivPress, New Haven, CT), pp 75–87.

37. Hill A (1999) Late Miocene sub-Saharan vertebrates, and their relation to the Bay-nunah fauna, Abu Dhabi, United Arab Emirates. Fossil Vertebrates of Arabia: WithEmphasis on the Late Miocene Faunas, Geology, and Palaeoenvironments of theEmirate of Abu Dhabi, eds Whybrow PJ, Hill A (Yale Univ Press, New Haven, CT), pp420–429.

38. Bibi F, Shabel AB, Kraatz BP, Stidham TA (2006) New fossil ratite (Aves: Palae-ognathae) eggshell discoveries from the Late Miocene Baynunah Formation of theUnited Arab Emirates, Arabian Peninsula. Palaeontol Electronica 9(1):1–13.

39. Badgley C, et al. (2008) Ecological changes in Miocene mammalian record show im-pact of prolonged climatic forcing. Proc Natl Acad Sci USA 105(34):12145–12149.

40. Weston EM (2003) in Lothagam: The Dawn of Humanity in Eastern Africa, eds LeakeyMG, Harris JM (Columbia Univ Press, New York), pp 441–483.

41. Boisserie J-R (2005) The phylogeny and taxonomy of Hippopotamidae (Mammalia:Artiodactyla): A review based on morphology and cladistics. Zool J Linn Soc 143(1):1–26.

42. Gilbert CC, Goble ED, Hill A (2010) Miocene cercopithecoidea from the Tugen Hills,Kenya. J Hum Evol 59(5):465–483.

43. Benefit BR (1993) The permanent dentition and phylogenetic position of Victor-iapithecus from Maboko Island, Kenya. J Hum Evol 25(2):83–172.

44. Szalay FS, Delson E (1979) Evolutionary History of the Primates (Academic, New York).45. Benefit BR (1987) The Molar Morphology, Natural History, and Phylogenetic Position

of the Middle Miocene Monkey Victoriapithecus. PhD Dissertation (New York Univ,New York).

46. Delson E (1994) in Colobine Monkeys: Their Ecology, Behaviour, and Evolution, edsDavies AG, Oates JH (Cambridge Univ Press, Cambridge, UK), pp 11–43.

47. Ji X, et al. (2013) Juvenile hominoid cranium from the terminal Miocene of Yunnan,China. Chin Sci Bull 58(31):3771–3779.

48. Koufos GD (2013) in Fossil Mammals of Asia: Neogene Biostratigraphy and Chronol-ogy, eds Wang X, Flynn LJ, Fortelius M (Columbia Univ Press, New York), pp 595–621.

49. Plavcan JM, Ward CV, Manthi FK (2013) Diminutive cercopithecine teeth from Ka-napoi, Kenya, and implications for the evolution of body size and diversity in gue-nons. Am J Phys Anthropol S56:222.

50. Strasser E, Delson E (1987) Cladistic analysis of cercopithecid relationships. J Hum Evol16(1):81–99.

51. Groves CP (2000) in Old World Monkeys, eds Whitehead PF, Jolly CJ (Cambridge UnivPress, Cambridge, UK), pp 180–213.

52. Groves CP (2001) Primate Taxonomy (Smithsonian Institution Press, Washington, DC).53. Tosi AJ, Detwiler KM, Disotell TR (2005) X-chromosomal window into the evolution-

ary history of the guenons (Primates: Cercopithecini). Mol Phylogenet Evol 36(1):58–66.

54. Xing J, et al. (2007) A mobile element-based evolutionary history of guenons (tribeCercopithecini). BMC Biol 5:5.

55. Hart JA, et al. (2012) Lesula: A new species of Cercopithecus monkey endemic to theDemocratic Republic of Congo and implications for conservation of Congo’s centralbasin. PLoS ONE 7(9):e44271.

56. Leakey MG (1988) in A Primate Radiation: Evolutionary Biology of the African Gue-nons, eds Gautier Hion A, Bourliere F, Gautier J-P (Cambridge Univ Press, Cambridge,UK), pp 7–12.

57. Hooijer DA (1963) Miocene Mammalia of the Congo. Annales Musee Royal del’Afrique Central, 8th ser. 46:1–71.

58. Geraads D (1987) Dating the northern African cercopithecid fossil record. Hum Evol2(1):19–27.

59. Benefit BR, McCrossin ML, Boaz NT, Pavlakis P (2008) New fossil cercopithecoids fromthe Late Miocene of As Sahabi, Libya. Garyounis Scientific Bulletin Special Issue 5:265–282.

60. Swindler DR (2002) Primate Dentition: An Introduction to the Teeth of Non-HumanPrimates (Cambridge Univ Press, Cambridge, UK).

61. Benefit BR (1999) Victoriapithecus, the key to Old World monkey and catarrhineorigins. Evol Anthropol 7(5):155–174.


www.pnas.org/cgi/doi/10.1073/pnas.1323888111

Supporting InformationGilbert et al. 10.1073/pnas.1323888111

Fig. S1. CT Scans of AUH 1321. Clockwise from Top Left: mesial, occlusal, distal, oblique root, buccal, and lingual views. Note that the distal root of AUH 1321appears either broken or slightly open at its distal apex, indicating it would have been longer than preserved. The irregularity of the opening combined withthe moderate wear of the tooth suggests a small amount of breakage rather than a slightly open apex.

Gilbert et al. www.pnas.org/cgi/content/short/1323888111 1 of 9

www.pnas.org/cgi/content/short/1323888111

Fig. S2. Comparison of cercopithecine dP4 and M1 crown and root morphology. (Upper) From left to right: cercopithecine teeth in distal view scaled toapproximately the same buccolingual width: Chlorocebus aethiops lower left dP4, Macaca fascicularis lower left dP4, Cercopithecus mitis lower left dP4, AUH1321, and Allochrocebus lhoesti lower left M1. Note that typical cercopithecine dP4s are characterized by a high degree of basal crown flare and a proximal rootthat is constricted underneath the crown, leading to clear enamel “bulging” on the buccal and lingual sides at the base of the crown. Typical guenon M1s, onthe other hand, do not exhibit a high degree of flare and are straight-sided at the base of the crown with robust proximal roots that do not display any obviousconstriction. Furthermore, typical dP4s with a moderate-to-high degree of wear demonstrate clear root absorption on the lingual side, a feature not found inAUH 1321, whose distal root is either slightly broken (most likely) or exhibits a slightly open apex (less likely) and would have been even longer than preserved.(Lower) From left to right: guenons permanent lower left molars in buccal view scaled to approximately the same mesiodistal length: Cercopithecus cephusM1,AUH 1321, and Cercopithecus ascanius M2. Note that similar degrees of distal flare of the distal root can be found in a small sample of extant guenonscompared with AUH 1321. Finally, again note that the distal end of AUH 1321 appears to be slightly broken, making it highly likely that the distal root wouldhave been even longer, more typical of permanent molars rather than dP4s. See also Figs. S1 and S3 and Tables S1–S3.

Fig. S3. Results of discriminant functions analyses (DFA) on a sample of cercopithecine dP4s and M1s using five crown and root shape variables: (i) mesiodistalcrown length/mesial buccolingual crown breadth, (ii) mesiodistal crown length/distal buccolingual crown breadth, (iii) mesial buccolingual crown breadth/distal buccolingual crown breadth, (iv) distal root apex breadth/mesiodistal crown length, and (v) distal root + crown height (total distal height)/mesiodistalcrown length. Sample included 43 guenon M1s, 11 guenon dP4s, 30 macaque M1s, and 8 macaque dP4s sampled from the American Museum of Natural History(AMNH) and US Smithsonian National Museum of Natural History (USNM) Collections, including wild-shot and nonpathological zoo specimens to maximizesample sizes. (Left) DFA correctly classifying 94.6% (88% cross-validated) of cercopithecine specimens as either M1s or dP4s. The indices with the highest loadingon discriminant function 1 are the root height index (index 5) and the crown shape indices (indices 1 and 2), indicating that dP4s are most easily distinguishedfrom M1s by their shorter roots and narrower crowns. Note that AUH 1321 plots well within the M1 range (and outside of the dP4 range) on discriminantfunction 1 and is classified as an M1 with a probability of 99.2%. (Right) DFA correctly classifying 79.3% (73.9% cross-validated) of cercopithecine specimens asguenon M1s, guenon dP4s, macaque M1s, and macaque dP4s. The indices with the highest loading on discriminant function 1 are the overall narrowness of thecrown (index 1), width of the distal lophid compared with the mesial lophic (index 3), and the root height index (index 5). The index with the highest loadingon discriminant function 2 is the distal root breadth index (index 4). Note that AUH 1321 plots well within the range of guenon M1s (and outside the range ofguenon or macaque dP4s) and is classified as a guenon M1 with a probability of 97.5%. In sum, although there is overlap between AUH 1321 and cercopithecinedP4s in some features, there is no dP4 in our sample that has all of the characteristics found in AUH 1321 and AUH 1321 is considered to be an M1 with a highdegree of probability.



Fig. S4. Most parsimonious trees derived from a cladistic analysis of 13 dental characters including stem cercopithecoids (Victoriapithecus), small-bodiedextant papionins, small-bodied extant cercopithecins, and AUH 1321: (A) AUH 1321 is reconstructed as the sister taxon of Chlorocebus and (B) AUH 1321 isreconstructed as the sister taxon of Cercopithecus. An exhaustive search was performed with Aegyptopithecus and Proconsul assigned as outgroups; bootstrapsupport values from a 10,000 replication branch and bound search with replacement are provided above various nodes. Tree length = 26, CI = 0.7308,HI = 0.2692, RI = 0.7586, RC = 0.5544; see Tables S4 and S5 for additional details.



Table S1. Selected comparative measurements (mm) of the dentition in the Abu Dhabi cercopithecines and other Old World Monkeys


(MD/MBL)M1

(MD/DBL)M1

(MBL/DBL)

Cercopithecini sp. indet. (Present study) AUH 1321 6.2 4.5 4.6 67 66 138 135 98Late Miocene

fossil macaquesMacaca libyca (1) Average 7.6 7.5 7.3 — 53* 101 104 103Macaca sp. Menacer (1) Average 9.1 7.6 6.8 — 65* 121 134 111Macaca sp. Almenara (2) IPMC 11676 8.0 6.2 6.1 — — 129 131 102As Sahabi (1) Average 7.6 6.4 6.3 — 51* 119 121 102

(1, 2) Average 8.1 6.9 6.6 — 57 117 122 104(1, 2) Species Range 7.6–9.1 6.2–7.6 6.1–7.3 — 51–65 101–129 104–134 102–111

Late Miocenefossil colobinesMicrocolobus

tugenensis(3) KNM-BN 1740 5.5 4.4 4.3 80 — 125 128 102

Mesopithecus sp. (3) Average 6.8 6.0 6.4 137 — 114 107 94Extant cercopithecidsCercopithecinae† (4) Average — — — 54† 60† 125† — 101†

(4) Species range — — — 53–78† 53–71† 115–130† — 97–102†

(4) Ind. range — — — 29–78† 42–83† 106–161† 106–174† 83–127†

Colobinae (4) Average — — — 124 70 135 — 96(4) Species range — — — 83–143 64–80 114–140 — 93–102(4) Ind. range — — — 66–196 49–87 100–154 100–174 79–118

Extant macaquesM. fascicularis (male) (5) Average 6.5 5.1 5.0 — — 128 130 102M. fascicularis (female) (5) Average 6.4 5.1 4.9 — — 126 131 104M. fascicularis (6) Average — — — 61 66 131 135 103Macaca mulatta (male) (5) Average 7.2 5.6 5.5 — — 129 131 102M. mulatta (female) (5) Average 7.0 5.5 5.4 — — 127 130 102Macaca nemestrina

(male)(5) Average 7.5 5.8 5.8 — — 129 129 100

M. nemestrina (female) (5) Average 7.3 5.6 5.4 — — 130 135 104M. nemestrina (6) Average — — — 44 70 131 128 98Macaca nigra (male) (5) Average 7.1 5.2 5.3 — — 137 134 98M. nigra (female) (5) Average 6.6 4.9 4.7 — — 135 140 104M. nigra (6) Average — — — — — 120 119 99M. maura (6) Average — — — 42 71 127 122 100Macaca radiata (6) Average — — — — — 128 129 101Macaca silenus (6) Average — — — 50 67 131 133 101

(5, 6) Macaqueaverage

7.0 5.4 5.3 49 68 129 130 101

(5, 6) Species range 6.4–7.5 4.9–5.8 4.7–5.8 44–61 66–71 120–137 119–140 98–104(6) Ind. range — — — 21–114 40–89 107–161 113–174 83–127

Extant guenonsCh. aethiops (male) (5) Average 5.9 4.4 4.5 — — 134 131 98Ch. aethiops (female) (5) Average 5.6 4.2 4.2 — — 133 133 100Ch. aethiops (PRIMO,

present study)Average 5.7 4.4 4.4 66 62 130 130 100

C. ascanius (male) (5) Average 5.2 3.8 4.0 — — 137 130 95C. ascanius (female) (5) Average 5.0 3.6 3.8 — — 139 132 95C. ascanius (PRIMO,


Cercopithecusneglectus (male)

(5) Average 6.3 4.4 4.6 — — 143 137 96

C. neglectus (female) (5) Average 5.9 4.4 4.5 — — 134 131 98C. neglectus (present study) Average 5.9 4.1 — — 62 144 — —C. mitis (male) (5) Average 6.2 4.3 4.6 — — 144 135 93C. mitis (female) (5) Average 6.0 4.3 4.5 — — 140 133 96C. mitis (PRIMO,


Cercopithecus mona(male)

(5) Average 5.4 3.9 4.0 — — 138 135 98

C. mona (female) (5) Average 5.2 3.7 3.9 — — 141 133 95



Table S1. Cont.


(MD/MBL)M1

(MD/DBL)M1

(MBL/DBL)

Ceropithecus nictitans(male)

(5) Average 5.7 4.0 4.2 — — 143 136 95

C. nictitans (female) (5) Average 5.5 3.9 4.2 — — 141 131 93C. cephus (male) (5) Average 5.6 4.1 4.2 — — 137 133 98C. cephus (female) (5) Average 5.4 3.8 4.0 — — 142 135 95A. lhoesti (PRIMO,



Guenonaverage

5.6 4.0 4.2 61 61 139 133 96


Species range 5.0–6.3 3.6–4.4 3.8–4.7 39–77 55–63 130–144 130–137 89–100

(PRIMO,present study)

Ind. range — — — 5–164 43–77 110–169 103–149 85–111

In addition to measurements collected during the course of this study, comparative measurements are derived from refs. 1–6, as well as from the PRIMO(Primate Morphology) database, access courtesy of E. Delson. —, unavailable measurement; DBL, max distal buccolingual width measured across the distallophid; Ind. range, range of values among individuals within a given group; MBL, maximum mesial buccolingual width measured across the mesial lophid; MD,maximum mesiodistal length; NH/NR, notch height/crown height below the notch [see Benefit (4, 6) for details]; species range, range of values among speciesaverages within a given group. Flare measured as in Benefit (4, 6) and lower values indicate higher degrees of flare.*Estimated from M2 values in ref. 1.†Cercopithecine values include papionin taxa only [see Benefit (4, 6)].

1. Benefit BR, McCrossin ML, Boaz NT, Pavlakis P (2008) New fossil cercopithecoids from the Late Miocene of As Sahabi, Libya. Garyounis Scientific Bulletin Special Issue 5:265–282.2. Köhler M, Moyà-Solà S, Alba DM (2000) Macaca (Primates, Cercopithecidae) from the late Miocene of Spain. J Hum Evol 38(3):447–452.3. Benefit BR, Pickford M (1986) Miocene fossil cercopithecoids from Kenya. Am J Phys Anthropol 69(4):441–464.4. Benefit BR (1993) The permanent dentition and phylogenetic position of Victoriapithecus from Maboko Island, Kenya. J Hum Evol 25(2):83–172.5. Swindler DR (2002) Primate Dentition: An Introduction to the Teeth of Non-Human Primates (Cambridge Univ Press, Cambridge, UK).6. Benefit BR (1987) The Molar Morphology, Natural History, and Phylogenetic Position of the Middle Miocene Monkey Victoriapithecus. PhD Dissertation (New York Univ, New York).



Table S2. Selected comparative measurements of the cercopithecine dP4 crown and AUH 1321

Taxon Source Specimen no.Samplesize MD MBL DBL Flare (MD/MBL) (MD/DBL) (MBL/DBL)


(Present study) AUH 1321 n = 1 6.2 4.5 4.6 66 138 135 98

Extant CercopithecinedP4 Total

(PRIMO,present study)

Average n = 120 — — — 55 150 146 98

Fossil macaques dP4 MD dP4 MBL dP4 DBL dP4 Flare dP4(MD/MBL)

dP4(MD/DBL)

dP4(MBL/DBL)

Macaca sylvanus?pliocena

(PRIMO) Average n = 3 7.1 5.2 5.2 48 138 136 99

M. sylvanus?florentina

(PRIMO) Average n = 2 7.7 4.7 5.3 58 164 146 90

M. sylvanus?prisca

(PRIMO) Average n = 5 8.1 5.2 5.4 49 155 151 98

Macaca majori (PRIMO) Average n = 12 6.5 4.5 4.5 44 146 145 99Fossil macaque

average7.4 4.9 5.1 50 151 145 96

Species range 7.6–9.1 6.2–7.6 6.1–7.3 44–58 138–164 136–151 90–99Extant macaques dP4 MD dP4 MBL dP4 DBL dP4 Flare dP4

(MD/MBL)dP4

(MD/DBL)dP4

(MBL/DBL)M. sylvanus (PRIMO,

present study)Average n = 6 6.9 4.7 4.7 63 147 145 98

M. fascicularis (Present study) Average n = 4 6.0 4.4 4.4 58 138 137 99M. mulatta (Present study) Average n = 13 6.4 4.5 4.5 53 142 142 101M. nemestrina (Present study) Average n = 4 6.9 4.7 4.7 48 148 148 100M. nemestrina (1) Average n = 40 6.8 4.2 4.2 — 162 162 —M. silenus (Present study) Average n = 3 6.1 4.3 4.2 55 141 144 102Macaca sinica (Present study) Average n = 1 5.8 4.0 3.9 50 145 149 103Macaca tonkeana (Present study) Average n = 2 7.2 5.4 5.3 53 134 136 102

Extant macaqueaverage

6.5 4.5 4.5 54 145 145 101

Species range 5.8–7.2 4.0–5.4 3.9–5.3 50–63 134–162 136–162 98–103Extant baboons dP4 MD dP4 MBL dP4 DBL dP4 Flare dP4

(MD/MBL)dP4

(MD/DBL)dP4

(MBL/DBL)Papio ursinus (PRIMO) Average n = 30 9.7 6.5 6.7 — 149 145 97Papio

cynocephalus(1) Average n = 35 9.5 — 6.1 — — 156 —

Baboon average 9.6 6.5 6.4 — 149 151 97Species range 9.5–9.7 — 6.1–6.7 — — 145–156 —

Extant guenons dP4 MD dP4 MBL dP4 DBL dP4 Flare dP4(MD/MBL)

dP4(MD/DBL)

dP4(MBL/DBL)

Ch. aethiops (Present study) Average n = 14 5.6 3.8 3.9 54 146 143 98A. lhoesti (Present study) Average n = 2 6.0 3.5 3.9 57 170 153 90Cercopithecus

albogularis(Present study) Average n = 6 5.6 3.3 3.7 59 169 153 91

Cercopithecushamlyni

(Present study) Average n = 1 5.1 3.7 3.7 57 138 138 100

C. ascanius (Present study) Average n = 1 4.3 2.8 3.0 — 154 143 93C. mitis (Present study) Average n = 11 5.5 3.4 3.6 57 161 153 95C. mitis (1) Average n = 12 5.5 — 3.4 — — 162 —C. neglectus (Present study) Average n = 5 5.9 3.7 3.9 55 162 152 94C. mona (Present study) Average n = 2 4.6 3.4 3.4 — 137 135 98

Guenon average 5.3 3.5 3.6 57 155 148 95Species range 4.3–6.0 2.8–3.8 3.0–3.9 54–59 137–170 135–162 90–100

—, unavailable measurement; DBL, maximum distal buccolingual width measured across the distal lophid [see Benefit (2, 3) for details]; Group averagesrepresent averages of all of the species averages within a given group; MBL, maximum mesial buccolingual width measured across the mesial lophid; MD,maximum mesiodistal length; Species range, range of values among species averages within a given group. Flare measured as in Benefit (2, 3) and lower valuesindicate higher degrees of flare. Note that cercopithecine dP4s are more flaring and significantly narrower than M1s as measured by MD/MBL and MD/DBL;AUH 1321 is most similar in shape to a typical cercopithecine M1, specifically a guenon M1. See also Table 1, Figs. S2–S4, and Tables S1 and S3.

1. Swindler DR (2002) Primate Dentition: An Introduction to the Teeth of Non-Human Primates (Cambridge Univ Press, Cambridge, UK).2. Benefit BR (1993) The permanent dentition and phylogenetic position of Victoriapithecus from Maboko Island, Kenya. J Hum Evol 25(2):83–172.3. Benefit BR (1987) The Molar Morphology, Natural History, and Phylogenetic Position of the Middle Miocene Monkey Victoriapithecus. PhD Dissertation (New York Univ, New York).



Table S3. Selected comparative measurements (mm) of cercopithecine M1 and dP4 distal roots with AUH 1321

Taxon SourceSpecimen

no.Samplesize MD

Distal rootinferior breadth

Distal rootmax height

Distal rootBR index

Distal rootHT index


(Present study) AUH 1321 n = 1 6.2 2.4 9.6* 39 155*

CercopithecinedP4 Total

(Present study) Average n = 19 — — — 23 152†

CercopithecineM1 Total

(Present study) Average n = 73 — — — 26 181†

Extant macaques dP4 MD dP4 distalroot inferior

breadth

dP4 distalroot max height

dP4 distalroot BR index

dP4 distalroot HT index

M. fascicularis (Present study) Average n = 4 6.0 1.6 8.8 27 144M. mulatta (Present study) Average n = 1 7.0 1.2 9.4 17 134M. nemestrina (Present study) Average n = 1 6.6 1.4 10.6 21 161M. silenus (Present study) Average n = 1 6.0 1.1 10.8 18 180M. sinica (Present study) Average n = 1 5.8 0.7 11.0 12 190

(Present study) Average n = 8 6.2 1.3 9.6 22 155(Present study) Ind. Range n = 8 5.7–7.0 0.7–2.8 5.0–11.3 12–49 88–190

Extant guenons dP4 MD dP4 distalroot inferior

breadth

dP4 distalroot max height

dP4 distalroot BR index

dP4 distalroot HT index

Ch. aethiops (Present study) Average n = 7 5.7 1.3 8.9 22 156A. lhoesti (Present study) Average n = 1 6.1 1.3 8.5 21 139C. mitis (Present study) Average n = 1 5.6 1.4 7.2 25 129C. neglectus (Present study) Average n = 2 6.1 2.1 8.5 34 142

(Present study) Average n = 11 5.8 1.4 8.6 24 150(Present study) Ind. range n = 11 5.0–6.6 0.8–2.9 7.2–12.2 16–45 114–185

Extant macaques M1 MD M1 distalroot inferior

breadth

M1 distalroot max height

M1 distalroot BR index

M1 distalroot HT index

M. fascicularis (Present study) Average n = 12 6.5 1.6 11.9 25 185M. mulatta (Present study) Average n = 6 7.1 1.7 13.2 23 185M. nemestrina (Present study) Average n = 5 7.6 1.8 14.6 24 191M. silenus (Present study) Average n = 3 6.7 1.7 12.5 25 187M. tonkeana (Present study) Average n = 4 7.9 1.7 14.4 22 182

(Present study) Average n = 30 7.0 1.7 13.1 24 187(Present study) Ind. range n = 30 5.2–8.1 1.1–2.6 9.8–15.3 16–40 165–234

Extant guenons M1 MD M1 distalroot inferior

breadth

M1 distalroot max height

M1 distalroot BR index

M1 distalroot HT index

Ch. aethiops (Present study) Average n = 19 6.0 1.7 10.4 29 174A. lhoesti (Present study) Average n = 2 6.4 1.4 11.7 21 183C. albogularis (Present study) Average n = 9 6.1 1.5 11.2 24 188C. cephus (Present study) Average n = 2 5.5 1.4 10.1 26 167C. hamlyni (Present study) Average n = 1 5.4 1.4 8.7 26 161C. ascanius (Present study) Average n = 2 5.2 1.4 9.2 27 179C. mitis (Present study) Average n = 4 6.4 2.0 10.3 31 161C. neglectus (Present study) Average n = 1 5.8 1.3 12.4 22 214C. mona (Present study) Average n = 3 5.7 1.3 10.4 22 184

(Present study) Average n = 43 5.9 1.6 10.5 27 178(Present study) Ind. Range n = 43 5.0–6.9 1.1–2.8 8.2–13.0 20–43 140–216

—, unavailable measurement; Average, average of all individual specimens measured; Distal root BR index, Distal root inferior breadth/MD; Distal root HTindex, Distal root max height/MD; Distal root inferior breadth, maximum buccolingual breadth across the apex of the distal root; Distal root max height,maximum height from the distal root apex to the top of the distal portion of the molar crown (entoconid); Ind. range, range of all individual specimensmeasured; MD, maximum mesiodistal length. See text for details. Note that cercopithecine M1 distal roots are typically broader and significantly longer thandP4 distal roots and the distal root of AUH 1321, if fully preserved, would have been broader and longer than a typical dP4. See also Table 1, Figs. S2–S4, andTables S1 and S2.*This value is almost certainly an underestimate due to breakage at the apex of the distal root.†Denotes a significant difference at the P < 0.05 level between cercopithecine dP4s and M1s.



Table S4. Cladistic characters used in this study

Character Definition Character Type

1 Adult molar bilophodonty; States: 0 = absent, 1 = variably complete, 2 = complete QL2 Adult molar crista obliqua/cristid obliquid; States: 0 = present, 1 = variably present, 2 = absent QL3 Adult molar cingulae; States: 0 = present, 1 = absent QL4 Adult M1–2 hypoconulid; States: 0 = present, 1 = variably present/reduced, 2 = absent QL5 Adult M3 hypoconulid; States: 0 = present, 1 = variably present/reduced, 2 = absent QL6 Adult molar flare; States: 0 = high, 1 = intermediate, 2 = low QN7 Adult M1 crown shape (MD/MBL); States: 0 = wide crown, 1 = intermediate, 2 = narrow crown QN8 Adult M1 mesial shelf length/crown length; States: 0 = short, 1 = long QN9 Adult M1 basin length/crown length; States: 0 = short, 1 = long QN10 Adult molar cusp height (NH/NR); States: 0 = low, 1 = intermediate QN11 Adult M1 Shearing (SUMS/length); States: 0 = low, 1 = intermediate QN12 Adult M1 Crown waisting (MBL/DBL); States: 0 = wide distally, 1 = narrow distally QN13 Body mass as estimated from M1 length; States: 0 = small, 1 = intermediate QN

QL, Qualitative character; QN, Quantitative character. Quantitative characters scored using gap-weighted coding. All characterswere considered ordered. All character states were assessed using our own measurements and observations, measurements from thePRIMO database (access courtesy of E. Delson), as well as those found in the following references (1–11). Abbreviations as in Table 1and Table S1.

1. Benefit BR (1993) The permanent dentition and phylogenetic position of Victoriapithecus from Maboko Island, Kenya. J Hum Evol 25(2):83–172.2. Swindler DR (2002) Primate Dentition: An Introduction to the Teeth of Non-Human Primates (Cambridge Univ Press, Cambridge, UK).3. Benefit BR (1987) The Molar Morphology, Natural History, and Phylogenetic Position of the Middle Miocene Monkey Victoriapithecus. PhD Dissertation (New York Univ, New York).4. Delson E, et al. (2000) Body mass in Cercopithecidae (Primates, Mammalia): estimation and scaling in extinct and extant taxa. Anthropol Pap Am Mus Nat Hist 83:1–159.5. Szalay FS, Delson E (1979) Evolutionary History of the Primates (Academic, New York).6. Strasser E, Delson E (1987) Cladistic analysis of cercopithecid relationships. J Hum Evol 16(1):81–99.7. Benefit BR (1999) Victoriapithecus, the key to Old World monkey and catarrhine origins. Evol Anthropol 7(5):155–174.8. Rossie JB, Gilbert CC, Hill A (2013) Early cercopithecid monkeys from the Tugen Hills, Kenya. Proc Natl Acad Sci USA 110(15):5818–5822.9. Kay RF (1981) The nut-crackers: A new theory of the adaptations of the Ramapithecinae. Am J Phys Anthropol 55(2):141–151.10. Kay RF, Fleagle JG, Simons EL (1981) A revision of the Oligocene apes of the Fayum Province, Egypt. Am J Phys Anthropol 55(3):293–322.11. Simons EL, Rasmussen DT, Gebo DL (1987) A new species of Propliopithecus from the Fayum, Egypt. Am J Phys Anthropol 73(2):139–147.

Table S5. Matrix used in cladistic analysis

Taxon Matrix

Aegyptopithecus 00000001?1010Proconsul 00000110?0011Victoriapithecus 1111001101110AUH 1321 2212?22111100Macaca 2212021100111Cercocebus 2212001100111Allenopithecus 2212201110110Cercopithecus 2212212110100Chlorocebus 2212212111110



Table S6. Selected male cercopithecoid C1 areas compared with AUH 35

Taxon Reference Sample size C1 area average C1 area range

Microcolobus tugenensis (PRIMO) n = 1 10.3 —Cercopithecus ascanius (1) n = 16 20.9 14.9–31.9Cercopithecus ascanius (PRIMO) n = 8 14.8 11.1–18.1Cercopithecus pogonias (PRIMO) n = 7 18.2 13.0–20.3Cercopithecus mona (1) n = 16 23.7 10.9–52.8Chlorocebus aethiops (1) n = 24 24.5 18.8–43.5Chlorocebus aethiops (PRIMO) n = 8 22.1 14.4–27.7Cercopithecus cephus (1) n = 6 25.2 22.4–28.0Mesopithecus monspessulanus (PRIMO) n = 5 29.8 27.9–33.9Allochrocebus lhoesti (PRIMO) n = 3 30.1 23.2–40.2Cercopithecus mitis (PRIMO) n = 1 31.0 —Cercopithecus neglectus (1) n = 8 31.1 21.5–38.3Macaca nigra (1) n = 5 32.8 13.8–55.1Macaca majori (PRIMO) n = 7 33.6 29.6–36.0Mesopithecus delsoni (PRIMO) n = 4 34.0 29.3–37.8Cercopithecus nictitans (1) n = 9 34.3 24.4–54.4Mesopithecus pentelicus (PRIMO) n = 25 37.3 27.3–46.0Cercopithecus mitis (1) n = 24 38.7 24.6–48.1Macaca fascicularis (1) n = 40 42.3 19.6–108.0AUH 35 (2) n = 1 42.6 —Macaca fuscata (PRIMO) n = 1 44.2 —Macaca arctoides (PRIMO) n = 1 45.8 —As Sahabi (3) n = 1 46.4 —Macaca mulatta (1) n = 31 47.3 11.4–88.2Macaca sylvanus pliocena (PRIMO) n = 1 48.0 —Erythrocebus patas (PRIMO) n = 4 48.1 42.7–53.0Macaca sylvanus prisca (PRIMO) n = 1 47.6 —Macaca sylvanus (PRIMO) n = 6 48.3 42.3–54.2Macaca sylvanus florentina (PRIMO) n = 5 62.8 51.3–69.6Macaca nigra (PRIMO) n = 1 64.3 —Macaca thibetana (PRIMO) n = 3 67.0 62.7–71.0Macaca nemestrina (PRIMO) n = 1 71.7 —Macaca tonkeana (PRIMO) n = 2 72.7 67.2–78.3Macaca nemestrina (1) n = 15 78.5 57.3–109.2

—, unavailable measurement. C1 area computed as C1 mesiodistal length x C1 buccolingual breadth. Becauseindividual values are not given in Swindler (1); range values derived from this source represent approximationsbased on multiplying the smallest C1 length and breadth values (minimum area) and the largest C1 length andbreadth values (maximum area) in the sample. Taxa in boldface represent those species that include the AUH 35C1 area value in their range and include extant guenons, extant macaques and the late Miocene colobineMesopithecus. Access to PRIMO database courtesy of E. Delson.

1. Swindler DR (2002) Primate Dentition: An Introduction to the Teeth of Non-Human Primates (Cambridge Univ Press, Cambridge, UK).2. Hill A, Gundling T (1999) Fossil Vertebrates of Arabia, eds Whybrow PJ, Hill A (Yale Univ Press, New Haven, CT), pp 198–202.3. Benefit BR, McCrossin ML, Boaz NT, Pavlakis P (2008) New fossil cercopithecoids from the Late Miocene of As Sahabi, Libya. Garyounis Scientific Bulletin Special Issue 5:265–282.


Date post:	15-Feb-2021
Category:	Documents
Upload:	others
View:	0 times
Download:	0 times

Early guenon from the late Miocene Baynunah Formation, Abu … · 2014. 7. 14. · Early guenon...

Documents