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Early evolution and biogeography of lorisiform strepsirrhines.

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American Journal of Primatology 69:27–35 (2007)
Early Evolution and Biogeography of Lorisiform
Department of Earth Sciences and Museum of Natural History, University of Oxford,
Oxford, United Kingdom
This brief review summarizes new paleontological and molecular data
that together support a late middle Eocene Afro-Arabian origin for crown
Lorisiformes. Phylogenetic analysis indicates that late Eocene Karanisia
is a possible stem lorisiform, late Eocene Saharagalago and Wadilemur
and Miocene Komba are stem galagids, and early Miocene Mioeuoticus
and Progalago may be crown lorisids. Character evolution along the
lorisid and galagid stem lineages is reconstructed as having occurred
primarily in postcranial and dental morphology, respectively. These
patterns have important implications for interpreting an early lorisiform
fossil record that is still composed primarily of jaws and isolated teeth.
Am. J. Primatol. 69:27–35, 2007. c 2006 Wiley-Liss, Inc.
Key words: Eocene; Miocene; Galagidae; Lorisidae; Lorisiformes
Our understanding of early lorisiform evolution has changed quite radically
since the turn of the century, thanks to new fossil discoveries that have
unexpectedly doubled the temporal range of crown Lorisiformes [Seiffert et al.,
2003], the publication of molecular divergence estimates that are remarkably
congruent with this expanded lorisiform fossil record [Yoder & Yang, 2004], and
the long-awaited identification of molecular markers that have clarified the
interrelationships of extant lorisid genera [Roos et al., 2004]. This review briefly
summarizes these recent developments and discusses the implications of these
data for our understanding of the origin and historical biogeography of crown
Morphological and genetic data have consistently supported a monophyletic
Galagidae, but the molecular evidence for lorisid monophyly has long been
unstable, with immunodiffusion and nucleotide data tending to support the
Correspondence to: Erik R. Seiffert, Department of Earth Sciences and Museum of Natural
History, University of Oxford, Parks Road, Oxford OX1 3PR, UK.
Received 7 July 2005; revised 6 February 2006; revision accepted 15 February 2006
DOI 10.1002/ajp.20324
Published online in Wiley InterScience (
r 2006 Wiley-Liss, Inc.
28 / Seiffert
paraphyly of lorises with respect to galagos [Dene et al., 1976; Goodman, 1967;
Goodman et al., 1998; Masters et al., 2005; Porter et al., 1997; Poux & Douzery
2004; Roos et al., 2004; Sarich & Cronin 1976; Yoder et al., 2001], despite an
impressive array of postcranial morphological specializations that appear to unite
Lorisidae as a well-defined clade [Gebo, 1989; Rasmussen & Nekaris, 1998; Yoder
et al., 2001]. Possible resolution of this conflict between molecular and
morphological data has been only recently provided by Roos et al. [2004], who
detected a number of Alu short interspersed nuclear elements (SINEs) that
solidify support for the existence of African (Arctocebus-Perodicticus) and Asian
(Loris-Nycticebus) clades within a monophyletic Lorisidae. Unlike the nucleotide
characters that were previously used to estimate lorisiform phylogeny, SINEs
should, in the absence of incomplete lineage sorting, be essentially immune to
homoplasy and reversal [Shedlock et al., 2000]. The interrelationships of extant
galagid genera remain controversial [Masters & Brothers, 2002], and unfortunately no SINEs have been detected that could convincingly resolve the
interrelationships of Galago, Galagoides, and Otolemur. However, Roos et al.’s
[2004] analysis of the mitochondrial locus cytochrome b has provided compelling
support for a Galago-Otolemur clade to the exclusion of Galagoides, and this
result is in general agreement with DelPero et al. [2000] and Yoder et al.’s [2001]
analyses of the same locus, as well as DelPero et al.’s [2000] analyses of two other
mtDNA loci (12S and 16S rRNA) and Yoder et al.’s [2001] analysis of the nuclear
IRBP gene.
The fossil record documenting early lorisiform evolution has long been a
source of great interest for paleoprimatologists, largely because Tertiary lorisiforms have, for many decades, been the oldest representatives of the entire crown
strepsirrhine radiation. Recent paleontological work in Egypt has significantly
expanded the temporal range of fossil lorisiforms and crown strepsirrhines
with the recovery of a 37-million-year-old (Ma) stem galagid (Saharagalago)
and a loris-like strepsirrhine (Karanisia) whose dental morphology resembles
that of the extant lorisid Arctocebus [Seiffert et al., 2003]. These taxa are
approximately twice as old as the oldest Miocene lorisiforms from east Africa, and
suggest that crown Lorisiformes had appeared by at least the late middle Eocene.
This hypothesis gains additional support from recently recovered dental remains
of the 34 Ma species Wadilemur elegans, which was previously thought to be an
anchomomyin adapiform [Simons, 1997]. These specimens indicate that that
species is also likely to be a primitive stem galagid [Seiffert et al., 2005b].
A proximal femur of W. elegans is also clearly galagid-like and bears greatest
similarity to the early Miocene stem galagid Komba, although the polarities of the
proximal femoral characters that Wadilemur shares with younger galagids are
currently unclear. Bayesian analyses of nuclear and mitochondrial nucleotide
sequences have independently recovered congruent middle Eocene estimates for
the origin of crown Lorisiformes [Yang & Yoder, 2003; Yoder & Yang, 2004],
although some other divergence estimates within crown Strepsirrhini indicate a
much later (early or middle Miocene) origin for crown lorisiforms [Porter et al.,
1997; Poux & Douzery, 2004].
Phylogenetic analysis of morphological data supports a stem galagid
placement for Saharagalago and Wadilemur, but the position of Karanisia is
unstable (Fig. 1a and c) [Seiffert et al., 2005b]. With increased taxon and
Am. J. Primatol. DOI 10.1002/ajp
Early Lorisiform Evolution / 29
character sampling, and the constraint of Arctocebus-Perodicticus monophyly,
Karanisia is placed as either a stem lorisiform (Fig. 1a) or a crown strepsirrhine of
uncertain affinities (Fig. 1b), given different treatments of certain multistate
characters. In light of these results, it appears increasingly likely that the
Arctocebus-Karanisia clade recovered by Seiffert et al. [2003] was based on
homoplasious and/or plesiomorphic features of the upper and lower dentition, and
that Karanisia is not a crown lorisid.
Although numerous studies have focused on the early Miocene east African
lorisiforms Komba, Mioeuoticus, and Progalago [Gebo, 1986, 1989; Le Gros Clark,
1956; Le Gros Clark & Thomas, 1952; Leakey, 1962; MacInnes, 1943; McCrossin,
1992; Phillips & Walker, 2002; Simpson, 1967; Szalay & Katz, 1973; Walker, 1970,
1974, 1978], specialists have never reached a consensus as to whether these taxa
are stem or crown members of Galagidae and Lorisidae [McCrossin, 1992; Phillips
& Walker, 2002], or possibly advanced stem, or very basal crown, lorisiforms
[Rasmussen & Nekaris, 1998]. When Mioeuoticus bishopi and Progalago
songhorensis are scored for craniodental features and added to the matrix of
Seiffert et al. [2005a], these species are either nested within Lorisidae [cf.,
McCrossin, 1992] or placed as crown lorisiforms of uncertain affinities [cf.,
Fig. 1. A: Relationships among living and extinct crown strepsirrhines, based on a strict consensus
of 18 most parsimonious trees (MPTs) recovered from a parsimony analysis with the Miocene
lorisiforms Progalago songhorensis and Mioeuoticus bishopi added to Seiffert et al.’s [2005a] matrix
of 106 taxa and 360 morphological features (with some multistate characters ordered and scaled).
The poorly known adapiform Europolemur dunaifi, which was included in Seiffert et al.’s [2005a]
analysis, was excluded from this analysis. Methods are as described in Seiffert et al. [2005a]. Branch
color/shading reflects biogeographic history, as optimized in Mesquite v. 1.06 [Maddison &
Maddison, 2005]. B: Proportion of dental, postcranial, and cranial characters unequivocally and
equivocally optimized (ACCTRAN) as having evolved along the lorisiform, lorisid, and galagid stem
lineages in the phylogram depicted in A. C: Adams consensus of 1,160‘MPTs, derived from analysis
of the Seiffert et al. [2005a] matrix with all characters unordered. D: The same as B, with
optimizations derived from the tree in C.
Am. J. Primatol. DOI 10.1002/ajp
30 / Seiffert
Rasmussen & Nekaris, 1998], depending on the treatment of certain multistate
characters. Regardless of where Mioeuoticus and Progalago are placed,
morphological evolution along the galagid and lorisid stem lineages is reconstructed as having occurred primarily in dental and postcranial morphology,
respectively (Fig. 1b and d). This pattern implies that stem galagids should be
easy to identify from the most common elements in the primate fossil record (i.e.,
mandibles, maxillae, and isolated teeth); however, as it is unlikely that the dental
morphology of the ancestral crown lorisid differed much from that of the
ancestral crown lorisiform, it appears that advanced stem lorisiforms, stem
lorisids, and extinct crown lorisids may be much more difficult to place as such on
the basis of dental morphology alone.
The absence of clear dental morphological support for the monophyly of
crown Lorisidae provides some explanation for why the phylogenetic positions of
the possible lorisids Mioeuoticus and Progalago are so unstable, since both genera
have very loris-like teeth and lack the dental synapomorphies of Galagidae, but
are not yet clearly represented by postcranial remains. Unfortunately, body mass
estimates for Mioeuoticus, Progalago, and larger species of Komba show broad
overlap, so it is not possible to confidently assign isolated postcranial elements to
any early Miocene species, other than the tiny stem galagid Komba minor.
Previous allocations of postcrania to Progalago dorae [Gebo 1989] on the basis of
size may be incorrect because the galagid-like upper dentitions that have been
attributed to P. dorae actually do not convincingly occlude with the very lorisidlike holotype mandible (containing p4 and m2). If these upper and lower
dentitions are not from the same species, then 1) P. dorae may actually be a
relatively rare lorisid, 2) an additional unnamed stem galagid species of similar
body size was present in the east African early Miocene (represented by the
‘‘P. dorae’’ upper teeth), and 3) the larger galagid-like postcrania previously
assigned to P. dorae on the basis of size [Gebo, 1989] may actually belong to that
unnamed galagid species. This hypothesis is supported by the fact that most
postcranial remains from the early Miocene resemble those that can be
confidently attributed to the tiny stem galagid species Komba minor, and are
thus most likely to be attributable to the stem galagids Komba robustus and the
galagid species represented by the ‘‘P. dorae’’ upper teeth.
On temporal grounds, it is unlikely that Mioeuoticus and Progalago are
either stem lorisiforms or very primitive stem galagids, since either placement
would require that these two lineages had independently persisted for 420 Ma
through the later Eocene and Oligocene into the early Miocene, while stem and
crown lorisid lineages that also would have existed during this interval would
be completely unsampled. The absence of loris-like postcrania attributable to
Mioeuoticus or Progalago could simply be due to the fact that these species are
very rare; alternatively, it could be the case that the unique postcranial features
shared by the African and Asian lorisids evolved in parallel in the two clades
[Walker, 1970], and that stem members of these African and Asian clades more
closely resembled primitive galagids in their postcranial morphology. The short
internal branch separating the origin of crown Lorisiformes and crown Lorisidae
[Roos et al., 2004] certainly leaves little time for all of the shared postcranial
adaptations of extant lorises to have evolved. Nevertheless, a maximum-likelihood
estimate for the ancestral mode of locomotion of crown Lorisidae, taking into
account three locomotor character states and Roos et al.’s [2004] divergence dates
within Lorisiformes (Fig. 2), indicates that the ancestral crown lorisid is likely to
have been a slow climber, while the ancestral crown lorisiform is only slightly
more likely to have been a pronograde quadruped than a slow climber. The fact
Am. J. Primatol. DOI 10.1002/ajp
Early Lorisiform Evolution / 31
Fig. 2. Maximum-likelihood optimization of locomotor style within Strepsirrhini, taking into
account the divergence dates of Roos et al. [2004] and locomotor categories of Shapiro and Simons
[2002]. Pie charts depict the proportional likelihoods for each of the locomotor styles, calculated
in Mesquite v. 1.06 [Maddison & Maddison, 2005].
that the proximal femoral morphology of 34 Ma Wadilemur is very similar to
that of Komba, and lacks the distinctive features of lorisids [Seiffert et al., 2005b],
lends additional support to the hypothesis that the ancestral crown lorisiform was
more likely to have been a pronograde quadruped (probably with some capacity
for leaping) than a slow climber. The eventual recovery of additional postcranial
remains of later Paleogene lorisiforms should help to better characterize the
locomotor adaptations of the ancestral crown lorisiform, and further inform
the likelihood that lorisids’ postcranial specializations evolved either in a short
burst along the short internal branch uniting crown lorisids, or in parallel in the
African and Asian clades.
It has recently been hypothesized that primates originated on the IndoMalagasy land mass in the late Cretaceous [Marivaux et al., 2001; Martin, 2000,
Am. J. Primatol. DOI 10.1002/ajp
32 / Seiffert
2003], and that lorisiforms dispersed into Asia, and then Africa, following the
collision of Indo-Pakistan with Asia near the Paleocene-Eocene boundary about
55 million years ago. While this scenario would provide a convenient explanation
for the sudden appearance of probable crown primates on northern continents in
the earliest Eocene, it must be emphasized that there is no fossil evidence for this
hypothesis. Furthermore, this scenario is very difficult to reconcile with the fossil
record of crown primates’ closest relatives, such as plesiadapiforms and Glires
(the rodent-lagomorph clade), which make their first appearances on northern
continents in the earliest Paleocene [Clemens, 2004; Johnston & Fox, 1984;
Meng, 2004], about 10 million years before Indo-Pakistan collided with Asia. The
first undoubted records of other close primate relatives, such as Scandentia (tree
shrews) and Dermoptera (flying lemurs), are found in Asia [Ducrocq et al., 1992;
Tong, 1988] and support an Asian origin for Primates and stem Strepsirrhini
[Beard, 1998], as does the Asian stem primate Altanius, which may be late
Paleocene in age [Bowen et al., 2002] and as such could predate the India-Asia
collision. The optimization of a biogeographic character onto topologies derived from
the phylogenetic analyses presented here suggests either an Asian or European
origin for crown primates, and a European origin for stem Strepsirrhini. The only
possible crown strepsirrhine from the Asian Paleogene, early Oligocene Bugtilemur,
is unlikely to be a cheirogaleid lemuriform as originally claimed [Marivaux et al.,
2001], and is probably a primitive relative of the recently described older Asian
genus Muangthanhinius [Marivaux et al., 2006] and the younger North American
primate Ekgmowechashala (unpublished results), neither of which had a toothcomb. Thus, there is no fossil evidence for crown strepsirrhines in Asia until the
middle Miocene, with the first appearance of Nycticeboides [MacPhee & Jacobs,
1986] 40 million years after the India-Asia collision, and over 20 million years after
the first appearance of Saharagalago and Karanisia in Africa.
In this regard it is important that recent phylogenetic analyses of Paleogene
primates [Seiffert et al., 2005a,b] have recovered two Afro-Arabian lineages–
Plesiopithecidae and a Djebelemur-‘‘Anchomomys’’ milleri clade–as successive
sister taxa of crown Strepsirrhini to the exclusion of all other Laurasian
adapiforms (Fig. 1a and c). These results, combined with Karanisia’s position as
either a stem lorisiform or a crown strepsirrhine of uncertain affinities, appear to
bear decisively on the longstanding debate surrounding the geographic origin of
stem and crown lorisiforms [MacPhee & Jacobs 1986; Yoder, 1997]. The simplest
biogeographic explanation for this phylogenetic pattern, with three of the known
crown lorisiform sister taxa being Afro-Arabian, and the other Malagasy, is
clearly an endemic Afro-Arabian origin for both crown Strepsirrhini and crown
Lorisiformes (Figs. 1a and c, and 3). Interestingly, the optimization of a
biogeographic character onto the two topologies derived from the phylogenetic
analyses presented here congruently suggests that the most ancient Afro-Arabian
stem strepsirrhine population that ultimately gave rise to crown Strepsirrhini
was probably derived from a European immigrant. Mean Bayesian estimates of
divergence dates suggest that this dispersal could have occurred as early as the
very latest Cretaceous [Yoder & Yang, 2004] or as late as the middle Paleocene
[Eizirik et al., 2004]. The primate Altiatlasius is known from the late Paleocene of
Africa (Morocco), and shares some upper molar features with the strepsirrhines
Plesiopithecus and Karanisia (e.g., complete lingual cingula), but on the basis of
available evidence Altiatlasius is most likely to be a basal anthropoid [Beard,
1998; Godinot, 1994; Seiffert et al., 2005a]. The stem anthropoid placement of
Altiatlasius in the phylogeny figured by Seiffert et al., [2005a], nested alongside
Asian taxa such as eosimiids and living and extinct tarsiers, nevertheless suggests
Am. J. Primatol. DOI 10.1002/ajp
Early Lorisiform Evolution / 33
Fig. 3. Scenario for the origin and dispersal of strepsirrhine primates, superimposed on Old World
paleogeography ca. 65 Ma (map modified from Scotese [2001]).
that limited primate dispersal between Afro-Arabia and Asia was possible in the
Paleocene, notably before the India-Asia collision, and at a time when, the
mammalian fossil record in Afro-Arabia and Asia for this period of time is poorly
sampled. The African stem strepsirrhine Djebelemur may be as old as early
Eocene in age [Hartenberger et al., 1997], and by that time already shows a
number of important morphological differences from Laurasian primates,
suggesting a considerable period of endemic evolution on the Afro-Arabian land
mass. Paleontological exploration in the Paleocene and early-middle Eocene of
Afro-Arabia should help to further illuminate the nature of crown strepsirrhine
origins and the earliest phases of stem lorisiform evolution.
I thank C. Beard, P. Chatrath, L. Gordon, G. Gunnell, P. Jenkins, R. Kay,
B. Marandat, L. Marivaux, E. Simons, and particularly A. Walker for providing
access to fossils, casts, and/or skeletal material; M. Silcox and two anonymous
reviewers for their helpful comments; and A. Burrows and L. Nash for inviting me
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