J Mammal Evol (2015) 22:307342

DOI 10.1007/s10914-014-9284-3

ORIGINAL PAPER

Systematics and Phylogeny of Paleocene-Eocene Nyctitheriidae

(Mammalia, Eulipotyphla?) with Description of a new Species
from the Late Paleocene of the Clarks Fork Basin, Wyoming, USA
Carly L. Manz & Jonathan I. Bloch

Published online: 9 December 2014

# Springer Science+Business Media New York 2014

Abstract Nyctitheriidae is a diverse group of small, insectivorous mammals from the Paleogene of Asia, North America,
and Europe that have alternately been linked to Eulipotyphla
(shrews, moles, hedgehogs, solenodons), Euarchonta (primates, tree shrews, dermopterans), or Chiroptera (bats).
Even intrafamilial relationships are poorly understood,
resulting in ambiguity regarding morphological character polarity critical for evaluating supraordinal relationships and
paleobiogeographic patterns. To help address this issue, we
performed a cladistic analysis of 51 North American,
European, and Asian nyctitheriid species, including a new
nyctitheriid, Plagioctenodon thewisseni sp. nov. from the late
Paleocene of Wyoming, using 66 characters derived from
dental morphology. Although the oldest nyctitheriids are
found in North America, the resulting most-parsimonious
cladograms support an Asian origin of the family with dispersal into North America by the early Paleocene. Among
North American and European groups, the subfamilies
Nyctitheriinae and Amphidozotheriinae, and the genera
Leptacodon and Saturninia are not monophyletic and require
future study and revision. The multi-species genera
Nyctitherium, Plagioctenodon (including P. thewisseni),
Plagioctenoides, Cryptotopos, and Euronyctia are found to
be monophyletic, whereas Wyonycteris is paraphyletic, having
Pontifactor bestiola nested within it. The earliest known
European nyctitheriids (Leptacodon nascimentoi,
Placentidens lotus, Plagioctenodon dormaalensis,

C. L. Manz (*)
Department of Geological Sciences, University of Florida,
Gainesville, FL 32611, USA
e-mail: clmanz@ufl.edu
C. L. Manz : J. I. Bloch
Florida Museum of Natural History, University of Florida,
Gainesville, FL 32611, USA

Wyonycteris richardi) appear in the early Eocene and are each

found in an otherwise strictly North American clade
consisting of either solely Paleocene or a combination of
Paleocene and Eocene taxa, suggesting at least four earliest
Eocene dispersals between North America and Europe.
Keywords Nyctitheriidae . Leptacodon . Plagioctenodon .
Paleogene . Systematics . Biogeography

Introduction
Nyctitheriidae is an extinct family of small-bodied mammals
with sectorial teeth known from the Paleocene-Eocene of
North America and Asia, and the Eocene-Oligocene of
Europe. Nyctitheriids were likely insectivorous and may have
had similar ecological adaptations to those of extant shrews
(Lopatin 2006; Rose et al. 2012). The fossil record of
nyctitheriids consists largely of isolated teeth and jaw fragments, providing limited morphological information for
reconstructing phylogenetic relationships between
nyctitheriids and other mammalian clades. Consequently,
nyctitheriids have been hypothesized to be closely related to
a diverse array of clades, including Chiroptera, Eulipotyphla,
and Euarchonta, by several authors without a clear consensus
being reached. This diverse range of suggested relationships,
along with their early Paleocene appearance in the fossil
record and primitive dental morphology, suggests that
nyctitheriids may be important taxa for understanding early
boreoeutherian (Euarchontoglires + Laurasiatheria) evolution.
Suprafamilial Relationships
Marsh (1872) classified the first nyctitheriid, Nyctitherium
velox, as a chiropteran based on its dental morphology. In
the following decades, some taxa first classified as

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chiropterans (i.e., Paradoxonycteris soricodon [Revilliod

1922] and Wyonycteris chalix [Gingerich 1987]) were later
recognized as nyctitheriids, further illustrating the morphological similarities between early bats and nyctitheriids. These
similarities in dental morphology include: 1) a buccal cingulid
on the lower molars in N. velox (Robinson 1968); 2) Wshaped ectoloph on the upper molars of Pontifactor bestiola
(West 1974), W. chalix (Gingerich 1987), Wyonycteris
richardi (Smith 1995), Euronyctia (Sig 1997), and
Paradoxonycteris (Hooker and Weidman 2000); 3) skewed
lower molars with higher lingual cusps than labial present in
most species of Plagioctenodon (Bown and Schankler 1982;
this publication), Nyctitherium krishtalkai (Christiansen and
Stucky 2013), Ceutholestes dolosus (Rose and Gingerich
1987), P. bestiola (West 1974), Wyonycteris taxa (Gingerich
1987; Smith 1995; Secord 2008; Beard and Dawson 2009),
Plagioctenoides taxa (Rose et al. 2012), and Placentidens
lotus (Russell et al. 1973); and 4) a simple P4 with a reduction
in size and number of cusps of the talonid basin. This last trait
occurs throughout Nyctitheriidae, from the primitive Asian
taxa near the base of the family (i.e., Missiaen and Smith
2005; Lopatin 2006) to more derived North American and
European taxa, such as Plagioctenoides (Rose et al. 2012) and
Amphidozotherium (Sig 1976). However, many of these
characteristics are found throughout the placental mammal
radiation, suggesting that they might be easily acquired
through parallel or convergent evolution. Furthermore, characteristics likely to be synapomorphies for early bats, such as a
buccal cingulid and reduced para- and metaconules on the
upper molars, do not occur in most nyctitheriids (Hand et al.
1994). Those nyctitheriids that do possess these chiropteran
synapomorphies, such as N. velox and some late Eocene
European nyctitheriids, have other derived traits, such as large
postcingula and hypocones on the upper molars that are quite
different from early bats, indicating that they are unlikely
directly related to that radiation.
Less controversial has been a suggested link between
Nyctitheriidae and Eulipotyphla, a group that includes
modern shrews, moles, hedgehogs, and solenodons, either with Soricomorpha (Simpson 1928; Butler 1988;
McKenna and Bell 1997) or Erinaceomorpha
(McKenna 1968; Robinson 1968; Sig 1976). These
hypotheses are less focused on derived synapomorphies,
and instead are based on observations that nyctitheriids
exhibit a dental morphology intermediate to primitive
eutherians and extant eulipotyphlans and thus make
likely candidates for the ancestral stock of Eulipotyphla
(Dawson and Krishtalka 1984; Butler 1988). For example,
Butler (1988: 132) notes the advances in nyctitheriids,
including multicuspid incisors, reduction in canine size, and
widening of the upper molar postcingulum that could be the
first steps in the highly derived dilambdodonty and
zalambdodonty of some eulipotyphlans.

J Mammal Evol (2015) 22:307342

In contrast to these dentition-based hypotheses, a

third supraordinal relationship for Nyctitheriidae was
proposed based upon the first described nyctitheriid
postcrania. Hooker (2001) observed that isolated
calcanea and astragali attributed to the European
nyctitheriid Cryptotopos sp. exhibit two characteristics
proposed to be euarchontan synapomorphies (e.g.,
Szalay and Drawhorn 1980; Silcox et al. 2005): a distal
sustentacular facet on the calcaneum and confluent ectal
and navicular facets on the plantar surface of the astragalus. Consequently, results from a phylogenetic analysis that included tarsal characters suggested that
Nyctitheriidae might be stem euarchontans, the order
that includes primates, treeshrews, and dermopterans
(Hooker 2001). More recently, a calcaneum has been
attributed to a second European nyctitheriid, Plagioctenodon
dormaalensis (as Leptacodon dormaalensis), which strongly resembles the calcaneum of Cryptotopos sp. (Coillot et al.
2013).

Intrafamilial Relationships
A major impediment to understanding the relationships
of nyctitheriids to higher eutherian clades is ambiguity
of character polarity due primarily to poorly-understood
intrafamilial relationships. Currently, the clade includes
roughly 70 species classified in 20 genera, with several
taxa under dispute over their validity or even inclusion
within the family. In the last century, there have been
many sub-clades proposed within the family
Nyctitheriidae (e.g., Simpson 1928; Krishtalka 1976;
Sig 1976; Bown and Schankler 1982; McKenna and
Bell 1997; Lopatin 2006) but there is little consensus
among authors (for review see Robinson 1968; Sig
1976; Gunnell et al. 2008).
The most recent classification of Nyctitheriidae includes
recognition of five subfamilies (Lopatin 2006): Asionyctiinae,
Praolestinae, Eosoricodontinae, Nyctitheriinae, and
Amphidozotheriinae. The former three subfamilies solely include Asian taxa and the latter two consist of a mixture of
European and North American taxa with the exception of a
single Asian nyctitheriid included in the Nyctitheriinae.
Nyctitheriinae is thought to be the most primitive nyctitheriid
subfamily and includes species in the North American genera
Nyctitherium, Leptacodon, and Pontifactor, the European
genera Saturninia, Scraeva, and Euronyctia (although this
genus is considered an amphidozotheriine by Hooker and
Weidmann [2000]), and the Asian genus Yuanqulestes
(McKenna and Bell 1997; Lopatin 2006). These taxa exhibit
a broad range of dental morphologies and are mostly differentiated from the other subfamilies based on the presence of a
submolariform lower fourth premolar (P4) (Lopatin 2006).

J Mammal Evol (2015) 22:307342

The Nyctitheriinae includes two speciose and problematic genera: Leptacodon and Saturninia from North
America and Europe, respectively. These genera have
been treated as wastebasket taxa for plesiomorphic
nyctitheriid species from their respective continents.
Although some recent efforts have been made to revise
Leptacodon and Saturninia (e.g., Sig 1997; Hooker and
Weidmann 2000; Beard and Dawson 2009), they are
both likely to be paraphyletic or even polyphyletic as
currently defined (see Results and Discussion).
Reconstructing the primitive condition of nyctitheriids
is complicated in part because the oldest members of
the family are poorly represented in the fossil record.
The earliest definitive nyctitheriids, Leptacodon tener
and Leptacodon munusculum, appear in the Torrejonian
NALMA. The oldest known proposed nyctitheriid,
Leptacodon proserpinae, was recovered from just after
the Cretaceous-Paleogene boundary in the Puercan
North American Land Mammal Age (NALMA) of
Montana (Van Valen 1978) with only the P4 described.
The classification of this taxon is somewhat questionable, though, due to the paucity of fossils attributed to
it (Gunnell et al. 2008). Meanwhile, the proposed
Cretaceous nyctitheriid Paranyctoides has since been
argued to be more closely related to the Cretaceous
eutherian family Zhelestidae (Archibald et al. 2001;
Averianov and Archibald 2013).
Species classified as Leptacodon are widely considered to be the most primitive nyctitheriids and are
therefore crucial to understanding the phylogenetic relationship of the family to higher clades. However, since
the description of the type species, Leptacodon tener,
more than ten species have been classified in this genus,
many of which have been challenged by later authors
and the monophyly of the group has been called into
question (Bown and Schankler 1982; Gunnell et al.
2008). Species classified as Leptacodon are predominantly found in North America, outside of which only
Leptacodon nascimentoi from Portugal is still considered valid. Whereas Smith (1996) suggested that
Gypsonictops dormaalensis from Belgium was better
classified in Leptacodon (Leptacodon dormaalensis),
which he considered a senior synonym of
Plagioctenodon, we follow Beard and Dawson (2009)
who recognized the validity of Plagioctenodon as distinct
from Leptacodon, with Plagioctenodon dormaalensis classified in that genus.
Most of the diagnostic characters for Leptacodon are
considered to be primitive for Nyctitheriidae and there
is little agreement on what, if any, synapomorphies exist
for the genus. Matthew and Granger (1921) described
Leptacodon tener from the Tiffanian NALMA and originally placed it within Leptictidae. Recognizing the

309

primitive morphology of L. tener, they called it the

least specialized in molars and premolars of any member of Leptictinae (Matthew and Granger 1921: 3). The
proposed diagnostic characters for Leptacodon included
molars with lower trigonids than those of the leptictid
Prodiacodon, distinct paraconids, and a reduction in
molar size from first to third, characters that can be
found in most other nyctitheriid taxa. The meaning of
another character, the protoconid overtopping inner
cusp, is unclear, but if it refers to a taller protoconid
than metaconid, the only other species of Leptacodon
with this state is Leptacodon packi, whereas a taller
protoconid than metaconid is frequently found in other
nyctitheriids not classified as Leptacodon (See character
38, state 1, in Appendix 3).
Simpson (1935) later revised the generic diagnosis, adding
characteristics of the P4 and the third lower molar (M3). He
noted that the P4 has a small, highly-positioned metaconid
partially joined with the protoconid and a low, yet distinct
paraconid, whereas the M3 talonid exhibits three subequal
cusps with a projecting hypoconulid and a subequal or
slightly higher protoconid than metaconid (Simpson
1935). McKenna (1968) described an upper dentition
associated with the type of Leptacodon tener and argued
for its classification in Nyctitheriidae rather than
Leptictidae. Rose et al. (2012) cited a low, crest-like
paraconid, centrally positioned hypoconulid, and absence of a mesostyle as diagnostic features of
Leptacodon; however, they also pointed out that these
characters could be plesiomorphic. Another problem
challenging the monophyly of Leptacodon is the morphological similarities of Leptacodon and Plagioctenodon,
causing authors to either synonymize the two (Smith 1996)
or to suggest new taxonomic attributions for some of the
species (Bown and Schankler 1982; Beard and Dawson
2009).
Here, we describe a new nyctitheriid species,
Plagioctenodon thewisseni, from the late Paleocene of North
America, provide the first description of parts of the anterior
dentition of Plagioctenodon rosei, and perform the most exhaustive phylogenetic analysis to date for Nyctitheriidae
using an original character-taxon matrix spanning 51
species of nyctitheriids. Although our study provides a
better understanding of intrafamilial relationships and
character polarities within Nyctitheriidae and compelling
evidence that many of the current generic attributions
for nyctitheriid species are in need of revision, we are
limiting the systematic focus of this paper to the composition of a monophyletic Plagioctenodon. Naming and
diagnosing new genera is outside of the scope of this
paper. Although it is likely that some genera, particularly Leptacodon and Saturninia, are paraphyletic or
even polyphyletic, we will not be placing the generic

310

names in quotation marks until there is a greater consensus on the composition of these genera within
Nyctitheriidae so as to avoid confusion with the existing
literature.

Materials and Methods

J Mammal Evol (2015) 22:307342

this species, including those with dentally associated crania

and postcrania. Additionally, palates and dentaries of
P. rosei have been recovered with most or all anterior
teeth preserved. All skeletal and dental elements of
P. thewisseni and P. rosei were extracted from limestone
by etching with 7 % formic acid buffered with calcium
phosphate tribasic and careful documentation of associated
bones through photography, similar to the methodology of
Bloch and Boyer (2001).

Fossil Recovery
Imaging and Measurements
Fossils of Plagioctenodon thewisseni and Plagioctenodon
rosei were recovered from a freshwater limestone nodule
in the Clarkforkian (Cf) NALMA, Cf-3 faunal zone, of
the Willwood Formation, Wyoming, USA (Fig. 1).
Paleocene and Eocene freshwater limestones from the
Willwood Formation preserve fossils of small vertebrates
not often represented in collections made predominantly
by surface prospecting, including rare or new species of
birds (Houde 1986, 1987; Gingerich 1987) and mammals
(Gingerich 1987; Rose and Gingerich 1987; Bloch et al.
1998, 2007; Bloch and Boyer 2001). Freshwater limestones in the Willwood Formation are often found as
lenses at discrete levels associated with drab paleosols,
possibly forming in depressions on the distal floodplain
where the water table was high or after periodic flooding
(Bloch and Bowen 2001; Bowen and Bloch 2002). The
limestone nodule containing the holotype for the new
species P. thewisseni, UM 86725, was collected at SC117, one of the University of Michigan Museum of
Paleontology Sand Coulee vertebrate localities. Additional
fossils recovered from limestones found at other latest
Clarkforkian Sand Coulee localities are also referred to

High-resolution images were generated from threedimensional digital reconstructions using CT data obtained
from either the Yale University Core Center for
Musculoskeletal Disorders microCT facility using a Scanco
Medical CT 35 machine or the Shared Materials
Instrumentation Facility (SMIF) at Duke University using a
Nikon XT H 225. Specimens were adhered to wax-covered
discs or mounted in foam to prevent movement during scanning and were scanned at resolutions less than 2m.
Three-dimensional digital reconstructions and twodimensional still images were created using Aviso 7 or
8 (http://www.vsg3d.com/avizo). Unfortunately, accurate
measurements could not be directly obtained from scan
data from the Duke SMIF lab due to an unrecognized
detector misalignment. Instead, all measurements were
performed using a Gaertner Scientific Corporation micrometer, which was checked for accuracy using a pair
of digital calipers.
Measurements for all teeth (Tables 1 and 2) are maximum
lengths and widths. The lower premolar lengths were measured parallel to the dentary and widths were perpendicular.

Fig. 1 Map of the Bighorn and Clarks Fork basins in north central Wyoming. All fossils of Plagioctenodon thewisseni were recovered from the Sand
Coulee localities in the Clarks Fork Basin located north of the Bighorn Basin

J Mammal Evol (2015) 22:307342

Table 1

Measurements of the upper teeth of Plagioctenodon rosei

UM 39843

P1 l
P1 w
P2 l
P2 w
P3 l
P3 w
P4 l
P4 w
M1 l
M1 w
M2 l
M2 w
M3 l
M3 w

311

0.84
0.45
0.98
0.49
1.17
0.78
1.57
1.81
1.67
1.93

UM 39875

1.71
1.92
1.74
2.39
1.56
2.34

UM 76895

1.99
1.73
2.17
1.55
2.07
1.27*
1.71*

UM 77032

1.71
1.89
1.59
2.08
1.46
2.03

Mean
0.84
0.45
0.98
0.49
1.17
0.78
1.66
1.90
1.68
2.14
1.52
2.14
1.27
1.71

l length, w width. *Measurements may not be representative of a typical

P. rosei M3 because UM 76895 contains a supernumerary M4 (Gingerich
1987), which may have affected the development and dimensions of the
M3

Measurements of the lower molars were obtained by

drawing a line through the metaconid and protoconid
and taking the maximum length and width that were
perpendicular and parallel to that line, respectively, because of the oblique position of some nyctitheriid molars with respect to the dentary. The simple P1-P2 were
oriented so that the length was taken along the long
axis of the teeth and the width was perpendicular to that
measurement. Measurements of P3-M3 were obtained by
drawing a line through the paracone and metacone and taking
the maximum length and width that were parallel and perpendicular to that line, respectively, following the Type 1 methodology of Secord (2008).
Choice of Taxa Used in the Phylogenetic Analysis
The nyctitheriids in this matrix include the five currently
proposed Plagioctenodon species, eight Leptacodon species
(only excluding Leptacodon proserpinae), the fourteen remaining North American taxa, nineteen European
nyctitheriids, and five Asian species. The taxa were coded
from specimens, casts, and the literature (Appendix 1).
European nyctitheriid taxa that were not included are either
from the later (latest Eocene to Oligocene) portion of the
familys lineage (Cryptotopos communis, Darbonetus
aubrelongensis, Darbonetus tuberi, Euronyctia belgica,
Euronyctia franconica, Euronyctia recta, Euronyctia
saturninensis, Euronyctia tobieni, Oligonyctia hoffmani,
Signyctia oligocoena) or are systematically questionable as
to their inclusion in Nyctitheriidae (Clinopternodus gracilis,

Remiculus). Most of the Asian-endemic nyctitheriids have

been proposed to form a monophyletic clade resulting from
a single migration event of nyctitheriids from North America
during the early Tiffanian NALMA (Missiaen and Smith
2005). Consequently, the inclusion of Asian nyctitheriids
was limited to well-known representatives of the three strictly
Asian subfamilies (Asionyctinae, Eosoricodontinae, and
Praolestinae) proposed by Lopatin (2006) to test their monophyly relative to included North American and European taxa.
A single Asian species, Yuanqulestes qiui, was classified in
the otherwise North American and European subfamily
Nyctitheriinae, but this hypothesis could not be assessed and
the species could not be included in the matrix using the
images from the primary literature because they were line
drawings of four isolated teeth solely in occlusal view (Tong
1997).
Inclusion of Leptacodon proserpinae hindered the resolution of our phylogenetic results in early iterations of our
analysis due to the limited morphological information available for this taxon and the destabilizing effect on our strict
consensus tree. Leptacodon proserpinae is a rare taxon and
only a single P4 has been described and figured in the literature
(Van Valen 1978). Because of this, only eight characters could
be coded for the taxon. If L. proserpinae is a nyctitheriid, it
would be the oldest one known and would likely occupy a
basal position in the family. But until more fossils of
L. proserpinae are known, it is impossible to provide a more
detailed phylogenetic hypothesis for the species in relation to
the rest of Nyctitheriidae.
Maelestes gobiensis, a Cretaceous eutherian, was used as
an outgroup to approximate an early eutherian morphology in
light of the primitive dental morphology of nyctitheriids. The
teeth of Macrocranion junnei and Adunator minutus have
many morphological similarities with those of nyctitheriids
and were also included in this analysis as outgroups that more
closely approximate the sister group to nyctitheriids than
M. gobiensis. However, M. junnei and A. minutus were not
treated as forced outgroup taxa in the analysis. Some differences in character states are likely derived and their inclusion
in the analysis was used to test whether the taxa with questionable placement in Nyctitheriidae would fall outside the
family. The Paleocene genus Adunator (including
Diacocherus; see Novacek et al. 1985; Secord 2008), with
the species A. minutus, A. martinezi, A. fredericki, A. lehmani,
A. amplus, and A. abditus, and the Eocene genus
Macrocranion, with the species M. junnei, M. germonpreae,
M. nitens, M. robinsoni, M. tenerum, M. tupaiodon, and
M. vandebroeki, are known from North America and Europe
and are commonly placed in the eulipotyphlan suborder
Erinaceomorpha (Novacek et al. 1985; McKenna and Bell
1997; Gunnell et al. 2008). Recent studies have questioned
this placement and instead suggested an affinity with the
afrotherian macroscelideans for both genera, as well as several

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Table 2

P1 l
P1 w
P2 l
P2 w
P3 l
P3 w
P4 l
P4 w tri
P4 w tal
M1 l
M1 w tri
M1 w tal
M2 l
M2 w tri
M2 w tal
M3 l
M3 w tri
M3 w tal
P1 l
P1 w
P2 l
P2 w
P3 l
P3 w
P4 l
P4 w
M1 l
M1 w
M2 l
M2 w
M3 l
M3 w

J Mammal Evol (2015) 22:307342

Measurements of the upper and lower teeth of Plagioctenodon thewisseni
UM 86725
Left

UM
86725
Right

0.75
0.31

0.80
0.38
1.20
0.59
0.47

0.69
0.30
0.88
0.35
0.85
0.38
1.17
0.52
0.47

1.23
0.73
0.75
1.16
0.74
0.73
1.16
0.68
0.57

1.32
0.73
0.66
1.20
0.80
0.73
1.15
0.70
0.57

0.67
0.32
0.94
0.59
1.24
1.43
1.22

0.85
0.38
0.93
0.60
1.22
1.48
1.28

1.52
1.13
1.57
0.86
1.41

1.65
1.15
1.56
0.83
1.36

UM
39873

UM
83931

1.05
0.45
1.15
0.52
0.49

0.75
0.38
1.16
0.50
0.42

1.29
0.74
0.73
1.32
0.75
0.67

1.30
0.73
0.83
1.24
0.82
0.74
1.00
0.69
0.52

UM
82576

UF 294696
Left

UF 294696
Right

UF
289746
Left

UF 289746
Right

0.75
0.40
0.90
0.36
0.87
0.33
1.10
0.49
0.40

1.04
0.44
0.40
1.22
0.76
0.78
1.10
0.78
0.71
1.06
0.59
0.56

1.16
0.74
0.73
0.63
0.62
0.29
0.76
0.37
0.99
0.74
1.21
1.39
1.18
1.53
1.14
1.58
0.78

1.16
1.41

UF
289747

1.07
0.55
0.47

1.22
0.72
0.70
1.17
0.73
0.75
1.10
0.66
0.57

Mean

0.73
0.33
0.89
0.35
0.86
0.38
1.12
0.51
0.44

1.25
1.36
1.22

0.96
0.74
1.18
1.36
1.24

1.26
0.73
0.74
1.19
0.76
0.72
1.09
0.66
0.56
0.62
0.29
0.76
0.35
0.95
0.67
1.21
1.40
1.23

1.61
1.16
1.56
0.89
1.25

1.55
1.13
1.54
0.89
1.33

1.57
1.14
1.56
0.85
1.34

l length, w width, w tri width of the trigonid, w tal width of the talonid, h height. The P1- P3 does not have a talonid, so the total width can be found in the
w tri row

other taxa traditionally considered to be erinaceomorphs

(Penkrot et al. 2008; Hooker and Russell 2012; Rose
et al. 2013). Regardless of the proper ordinal phylogenetic position of Macrocranion and Adunator, their geographic and temporal proximity and morphological similarities to nyctitheriids suggest that they are good genera to test the placement of taxa within Nyctitheriidae.
Adunator minutus and M. junnei were chosen to represent these two genera because both taxa are well represented in the fossil record and are not considered derived within their respective clades.

Body Mass Calculations

Estimated weight in grams was calculated for all species of
nyctitheriids included in the cladistic matrix for which either
the M1 or M1 was known. Two separate equations based on
the size of M1 or M1 (Bloch et al. 1998) were used to estimate
body mass with 95% confidence intervals. The maximum
length and maximum width of complete teeth were compiled
from the literature for each species (Appendix 1) and then
averaged for each tooth position. Tooth sizes of the M1 and M1
were then calculated by multiplying the averaged maximum

J Mammal Evol (2015) 22:307342

length by the averaged maximum width of each tooth.

Incomplete teeth with an estimated length or width were
included because in some species they represent the only M1
or M1 published in the literature.
Some species are represented mostly by isolated teeth in
the fossil record and the M1 and M1 could not be differentiated
from the M 2 or M 2 , respectively (e.g., Sig 1976,
1997). Only teeth that could be definitively attributed
to M1 or M1 were used in the body mass equations. For
this reason, we were unable to calculate an estimated
weight for Saturninia intermedia or Saturninia grandis.
Plagioctenoides tombowni and Saturninia pirenaica
could not be included because neither has a complete
M1 or M1 published in the literature.
The dimensions of the upper teeth of Plagioctenodon rosei
have not been published, so measurements of the upper teeth
of P. rosei are provided here (Table 1) taken from specimens
UM 39843 and UM 39875 and casts UM 76895 and UM
77032. We were unable to include measurements from many
of the specimens recovered from the limestones because most
of them are occluded or otherwise unable to be measured using a micrometer. The measurements from
Table 1 were used to calculate the estimated weight of
P. rosei based on the M1.
The dimensions for the M1 of Leptacodon nascimentoi
were taken from SV3-300 (Estravis 1996: Fig. 10), which is
identified as the M2 (?) in the literature (Estravis 1996). This
tooth, along with SV2-12 (identified as an M1, but we believe
to be an M2) are isolated teeth. Based on the upper molar
morphology of other nyctitheriid taxa, SV3-300 appears to be
an M 1 because of the distinctive anteriorly projecting
parastylar lobe that is positioned almost directly anterior to
the paracone. This is a feature commonly found in the M1 of
other nyctitheriids (e.g., Leptacodon packi [Secord
2 0 0 8 ] , Wy o n y c t e r i s c h a l i x [ G i n g e r i c h 1 9 8 7 ] ,
Nyctitherium velox [Robinson 1968], Pontifactor
bestiola [West 1974]) but not in the M2. SV2-12 instead
has a more labially directed parastylar lobe, which is
the state of the M2 in most other nyctitheriids. SV2-12
also has a more rectangular trigon basin with a greater
width to length ratio than SV3-300, which is another
feature that distinguishes the M 2 from the M 1 in
nyctitheriids (e.g., Leptacodon tener [McKenna 1968],
P. rosei [Gingerich 1987], P. bestiola [West 1974]).
Phylogenetic Analysis
The phylogenetic analysis is based on a character-taxon matrix
that includes 66 dental characters coded for 51 nyctitheriids, two
fossil erinaceomorphs, and the outgroup Maelestes gobiensis
(Appendix 1). It should be noted that several characters in our
matrix are similar to those recently published by Christiansen
and Stucky (2013). However, wording and character states vary

313

between the two matrices, primarily due to differences in scope

of study and issues surrounding outgroup choice. The analysis
presented here is focused on determining character polarity in
early nyctitheriids and how species classified in the genera
Leptacodon and Plagioctenodon are related to other North
American taxa, contemporary European taxa, and the endemic
Asian subfamilies. The character-taxon matrix of Christiansen
and Stucky (2013) was more limited in scope, testing relationships of Nyctitherium and Acrodentis species from the late early
Eocene (Wasatchian-7) Wind River Formation in relation to
other select North American nyctitheriids.
A phylogenetic analysis was performed in TNT v. 1.1
(Goloboff et al. 2008) using maximum parsimony. The
New Technology Search was first utilized with the
Sectorial Search, Ratchet, and Tree Fusing options selected and allowing the consensus to stabilize five times.
These trees were then used as the starting point in a
Traditional Search that searched for additional trees using
tree bisection and reconnection (TBR) branch-swapping. All
characters were treated as unordered and equally weighted.
Bremer values were calculated from 50,000 suboptimal trees
up to ten steps longer than the most parsimonious tree using
the Bremer script that is included in the TNT download.
Retention and consistency indices were calculated using the
stats available on the TNT wiki (http://tnt.insectmuseum.org).
Institutional Abbreviations
UF, Vertebrate Paleontology collection, Florida Museum of
Natural History, University of Florida, Gainesville, FL; UM,
University of Michigan, Museum of Paleontology, Ann
Arbor, MI; YPM, Yale Peabody Museum, New Haven, CT;
YPM-PU, Princeton University collections, Yale Peabody
Museum, New Haven, CT.
Systematic Paleontology
Class Mammalia Linnaeus, 1758
Order Eulipotyphla Waddell, Okada, and Hasegawa, 1999
Family Nyctitheriidae Simpson, 1928
Genus Plagioctenodon Bown, 1979
Type speciesPlagioctenodon krausae Bown, 1979
Included speciesPlagioctenodon krausae Bown, 1979;
Plagioctenodon savagei Bown and Schankler, 1982;
Plagioctenodon dormaalensis (Quinet, 1964);
Plagioctenodon rosei (Gingerich, 1987); Plagioctenodon
thewisseni sp. nov.
DistributionClarkforkian and early Wasatchian (late
Paleocene-early Eocene) of the Clarks Fork and Bighorn
basins in Wyoming; early Wasatchian (early Eocene) of the
Gulf Coastal Plain of Mississippi; upper Landenian (early
Eocene) of Dormaal, Belgium.

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Differential DiagnosisUnique among nyctitheriids in

having the combination of 1) a relatively high paraconid on
the P4, 2) a P3 that is smaller than the P2 and P4, and 3) an
anteriorly canted protoconid on the P3 (see Beard and Dawson
2009). Further differs from Asian nyctitheriids (except the
possible nyctitheriine Yuanqulestes qiui [Tong 1997]) in having a talonid basin and more than one talonid cusp on
the P4. All Plagioctenodon species (except
Plagioctenodon dormaalensis) differ from all species
classified as Leptacodon in having a significantly taller
entoconid than hypoconid on the lower molars, rather
than the two cusps being subequal. Plagioctenodon further differs from Leptacodon donkroni and Leptacodon
munusculum in having cuspate rather than crestiform
paraconids on P4-M1 and a distinct metacone on the
P4. Further differs from Leptacodon packi in having a
distinct metacone on the P4 and distinct hypocones on
the M12 (but see Secord 2008). Further differs from
Leptacodon nascimentoi in having subequal metaconids
and protoconids on the lower molars, rather than the
former being taller and larger than the latter as in
L. nascimentoi, and in lacking a pericone on the M12
precingula. Further differs from Leptacodon tener in
having a distinct metacone and protocone on the P3.
Further differs from Leptacodon catulus in having a
much greater length to width ratio of the P4.
Plagioctenodon differs from Leptacodon choristus in having a smaller P3 relative to the other premolars, although it is
similar to L. choristus in having an anteriorly canted
protoconid on the P3 and a relatively high paraconid on the
P4. Plagioctenodon further differs from L. choristus in having
a much greater length to width ratio of the P4, a more developed talonid basin that is longer relative to the rest of the P4, a
smaller M 3 relative to the M 12 , and a shallower
dentary. Plagioctenodon differs from Leptacodon
acherontus in having a smaller P3 relative to the other
premolars and a higher paraconid on the P4, although it
is similar in having an anteriorly inclined protoconid on
the P3. Plagioctenodon further differs from
L. acherontus in having less inflated cusps and more
lingually positioned paraconids on the lower molars.
Plagioctenodon further differs from Ceutholestes dolosus
in lacking a metaconid on the P3 and having a less molariform
P4. It further differs from Placentidens lotus in lacking a
metaconid on the P3, having a distinct hypoconulid on the
lower molars, and lacking an extension of the cristid obliqua
to the tip of the metaconid on the lower molars. Further differs
from Acrodentis rosenorum and Nyctitherium in having a
metacone on the P3, wider P4 talonids, less anteriorly compressed trigonids on the lower molars, less linguallypositioned hypoconulids on the lower molars, and in lacking
postcingulids on the lower molars. Differs from
Limaconyssus habrus in having a greater P4 talonid

J Mammal Evol (2015) 22:307342

width relative to the trigonid width, having lowercrowned molars, lacking postcingulids on M12 and
lacking an extension of the cristid obliqua to the tip
of the metaconid on the lower molars.
Plagioctenodon differs from all Plagioctenoides,
Wyonycteris, and late Eocene European nyctitheriid taxa with
the relevant premolar morphology in possessing a P 4
paraconid positioned high on the anterior trigonid, although
it is similar to those taxa in having a reduced P 3
relative to the other premolars and, with the exceptions
of Plagioctenoides microlestes, Saturninia grandis, and
Euronyctia grisollensis, an anteriorly inclined P 3
protoconid. Plagioctenodon further differs from
Plagioctenoides and Wyonycteris in having more
molariform P4 s, lacking an extension of the cristid
obliqua to the tip of the metaconid in the lower molars,
and lacking the nyctalodont condition (sensu Menu and
Sig 1971) in the lower molar talonids. Further differs
from Wyonycteris (except for Wyonycteris primitivus)
and Pontifactor in lacking mesostyles and dilambdodonty on
M12. Further differs from Wyonycteris in lacking pericones
on M12 and differs from Pontifactor in lacking a stylocone
and expanded postcingula on M12.
Plagioctenodon differs from all late Eocene European
nyctitheriids (including Amphidozotherium, Cryptotopos,
Euronyctia, Paradoxonycteris, and all Saturninia taxa except
Saturninia ceciliensis) in lacking greatly expanded
postcingula on the P4-M2. Further differs from Cryptotopos
in having less anteroposteriorly compressed trigonids on the
lower molars, lacking a notch halfway along the length of the
cristid obliqua on the lower molars, and lacking lower molar
postcingulids. Further differs from Paradoxonycteris and
Euronyctia in lacking upper molar dilambdodonty, lacking
an extension of the cristid obliqua to the tip of the metaconid
in the lower molars, and lacking a postcingulid on the lower
molars. Further differs from Amphidozotherium cayluxi in
lacking a notch in the cristid obliqua of the lower molars,
lacking a mesoconid on the cristid obliqua of the lower molars, having a less reduced M3 relative to the M2 size, and
having more strongly winged para- and metaconules on the
M12. Further differs from Scraeva hatherwoodensis in lacking a mesoconid on the cristid obliqua of the lower molars and
having less anteroposteriorly compressed trigonids on the
lower molars.
Further differs from the middle Eocene European
Saturninia ceciliensis in having the cristid obliqua of the
lower molars meet the protocristid lingual to the protocristid
notch, rather than more labially.
Dis cussi on B o w n (1 979 ) in i t i a l l y c l a s s i f i e d
Plagioctenodon in Adapisoriculidae and the original diagnosis for the genus included: (1) molars that decrease in size
posteriorly; (2) a P4 with a relatively high paraconid arising on
the anterior surface of the trigonid and a well-developed

J Mammal Evol (2015) 22:307342

talonid heel; (3) trigonids of the M2 and M3 more constricted

relative to the trigonid of the M1; (4) the P2 and P3 are
procumbent and not separated by a diastema; (5) the P2 is
double-rooted and considerably taller than P3; (6) a more
medially situated hypoconulid on the M1; (7) the P4 is shorter
anteroposteriorly relative to the M1; and (8) the protocones on
the upper molars are relatively steep posteriorly. Bown and
Schankler (1982) determined that Plagioctenodon was better
classified in Nyctitheriidae and that the original hypodigm for
the type species, Plagioctenodon krausae, actually included
several other taxa. Bown and Schankler (1982) emended the
diagnosis to: (1) P2 is larger than P3 with both teeth being
procumbent and not separated by a diastema; (2) P4 is
semimolariform and relatively elongated anteroposteriorly,
with a large, anteriorly-projecting paraconid situated high on
the anterior surface of the trigonid and a well-developed
talonid with two or three cusps; (3) M1 and M2 are not
significantly different in size, whereas M3 is slightly smaller;
(4) entoconids on M1 and M2 are relatively taller than the
hypoconids and located on the posterolingual margin of the
molars; and (5) molars have a shallow hypoflexid and exhibit
no vespiform constriction between the trigonids and talonids.
Beard and Dawson (2009) later argued that the diagnosis
should focus only on the third and fourth lower premolars,
as they felt these teeth contained the most significant morphological differences between Plagioctenodon and Leptacodon.
They identified Plagioctenodon by having: (1) a P3 with an
anteriorly canted protoconid that is shorter relative to the other
premolars and (2) a more molariform trigonid on the P4 than in
Leptacodon, on which the paraconid is positioned higher and
is less anteriorly shifted (Beard and Dawson 2009).
There has been much uncertainty with the genus
Plagioctenodon concerning its relation to Leptacodon. Smith
(1996) synonymized the two genera, whereas Beard and
Dawson (2009) retained the separate genera but transferred
Leptacodon rosei and Leptacodon dormaalensis into
Plagioctenodon based upon their updated diagnosis. This
created the first occurrence of a Plagioctenodon species outside of North America, with the presence of Plagioctenodon
dormaalensis in the early Eocene of Belgium. Beard and
Dawson (2009) further suggested that Leptacodon is a primitive genus restricted to the Paleocene and only includes
Leptacodon tener, Leptacodon packi, and Leptacodon
munusculum. Rose et al. (2012) noted that variation can be
found in the diagnostic traits of Beard and Dawson (2009)
within a single species and also pointed out that the lower
premolars are often missing in specimens, making identification of those fossils impossible. The derived traits listed in
previously published diagnoses of Plagioctenodon imply the
genus is monophyletic. The majority of traits used to diagnose
Leptacodon, such as sectorial cusps, relatively low trigonids
with a centrally-positioned hypoconulid, distinct paraconids,
and absence of a mesostyle (Matthew and Granger 1921;

315

McKenna 1968; Bown and Schankler 1982; Rose et al.

2012), are probably primitive. Consequently, Leptacodon
may be paraphyletic with Plagioctenodon nested within it,
or a polyphyletic grouping distributed throughout the family
Nyctitheriidae.
Plagioctenodon rosei (Gingerich, 1987)
(Fig. 2)
Cf. Leptacodon packi Rose, 1981; Leptacodon rosei
Gingerich, 1987
HolotypeUM 71650, left dentary with P2- M3, described
in Rose (1981: Fig. 11).
DistributionClarkforkian and possibly early
Wasatchian.
DiagnosisSee Rose (1981) and Gingerich (1987).
Emended descriptionSeveral specimens of P. rosei
have been recovered from the freshwater limestone with complete dentitions, which allows for description of their anterior
teeth (e.g., UF 303728; UF 303729; UF 303730). The dental
formula is 3-14-3/3-14-3.

Fig. 2 Micro-CT scan images of anterior dentitions of Plagioctenodon

rosei in labial view. a, Right dentary (I1-P4) and palate (I1-P4) of UF
303728. b, Left dentary (I1-M1)and palate (I2-P4) of UF 303729
(reversed), lingual view of R M13 visible

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The upper incisors of P. rosei are small and bi-lobed, with

one main anterior cusp and a small posterior cusp (Fig. 2).
They show a slight decrease in size posteriorly (Fig. 2a). The
anterior-most cusp of the I1 curves posteriorly near the tip and
comes to a rounded point. The smaller posterior cuspule of I1
points ventrally and is crestiform. The I23 are similar in
morphology to that of C1 and P1 in having larger rectangular
anterior cusps and considerably smaller posterior
cuspules in labial view. In I23, however, the anterior
cusp is less curved and has a more dorsoventral orientation. The upper canine is significantly larger than the
surrounding teeth and has a single broad root, a large
conical cusp that curves slightly posteriorly, and a much
smaller and shorter posterior cusp relative to the anterior
cusp. The P12 are both double-rooted, labiolingually
compressed, and consist of a triangular anterior cusp
and a much smaller posterior cusp in labial view. The
P2 is slightly taller than the P1. The P3 has three roots
and is dominated by a large, central paracone. It has a
small parastyle anterior to the paracone and a relatively
larger metastyle posteriorly that is connected to the
paracone by the postparacrista. There is also a small
protocone that is directly lingual to the paracone. The
metacone shows up variably as a small swelling on the
postparacrista about midway between the paracone and
metastyle (UM 39843, UF 303728) and is completely
absent in some specimens (UF 303729).
The lower incisors are small, procumbent, roughly equal in
size, medial-laterally compressed, and multi-lobed (Fig. 2b).
The I1 is the tallest of the lower incisors but is shorter in length
than the I2. It has three lobes and is flat on its medial surface,
which tapers dorsoanteriorly to form the anterior-most lobe of
the tooth. This lobe continues to curve posterolaterally,
becoming broad and rounded. A second lobe is posterolateral to the first at about the same height, but is
shorter in length. The tooth ends posteriorly in the third
cuspule, which is much smaller and shorter than the
other two. The I2 is lower-crowned than the I1 and is
the longest anteroposteriorly of the lower incisors. As
previously described by Rose et al. (2012), it has five
lobes, the first of which is the longest anteroposteriorly;
the following three lobes are rectangular in shape and
about the same size, whereas the fifth and final lobe is
small and is sometimes absent, possibly due to wear.
The third incisor has three lobes, although the separation between the first two is difficult to distinguish in
specimens exhibiting wear. The gap between the second
and third lobes is much larger, making the smaller,
posterior lobe easier to distinguish from the first two.
The lower canine is larger than the incisors and is about
twice the size of the P1. The C1 and P1 are similar in
morphology in being dominated by a large, rectangular
cusp that is oriented about 45 to the long axis of the

J Mammal Evol (2015) 22:307342

jaw and possessing a very small posterior cuspule at the

base of the tooth. The P2-M3 were previously described
by Rose (1981) as cf. Leptacodon packi.
ComparisonsIncisor and canine morphology is known
for few nyctitheriid taxa. North American taxa with associated
incisors or canines include the late Paleocene P. rosei (Rose
et al. 2012; this publication), Plagioctenodon thewisseni (this
publication), Ceutholestes dolosus (Rose and Gingerich
1987), and possibly the earliest Eocene Leptacodon donkroni,
although the isolated I2 attributed to it is from a locality that
also contains the small-bodied Plagioctenoides microlestes
and Plagioctenoides tombowni (Rose et al. 2012). Several
European taxa have also been documented with incisors or
canines, including the middle to late Eocene Saturninia
carbonum (Sig and Storch 2001), Saturninia grandis (Sig
1976), Cryptotopos beatus (as Saturninia beata) (Sig 1976),
and Euronyctia grisollensis (as Saturninia grisollensis) (Sig
1976), the late Eocene through early Oligocene Saturninia
gracilis (Sig 1976) and Amphidozotherium cayluxi (Sig
1976), and the early Oligocene Cryptotopos communis
(Ziegler 2007).
There are very few nyctitheriid species with documented
upper incisors. Sig (1976: Fig. 60) attributed a possible I23
to C. beatus (as Saturninia beata). The teeth described by Sig
(1976) are similar in morphology to the I23 of P. rosei (Fig. 2),
supporting their attribution to a nyctitheriid taxon.
Plagioctenodon thewisseni is the only other nyctitheriid for
which upper incisors have been recovered, with a referred I1I3. Among these three taxa, all upper incisors have a main
labiolingually-flattened cusp with a much smaller posterior
cusp and in terms of overall size, the I2 is slightly smaller than
the I1 but considerably larger than the I3. The main difference
between P. rosei, C. beatus, and P. thewisseni is the shape of
the large, anterior cusps of the incisors. The anterior cusps of
C. beatus have a more triangular shape in labial view, whereas
those of P. rosei are more squared off posteriorly and look like
an oblique rectangle, and in P. thewisseni they are like an
anteriorly canted triangle, with a convex anterior face and a
concave posterior face.
In P. rosei, there is some variability seen in the presence of
the upper canine posterior cuspuleit is absent in at least one
specimen, UF 303729, but present in UF 303728 and UF
303730. The upper canines of P. rosei specimens UF
303728 (Fig. 2a) and UF 303730 are similar to the upper
canine referred to P. thewisseni (Fig. 3) in having a large
anterior cusp with a small, posterior cuspule. Five teeth anterior to the P4 are illustrated for the holotype of Leptacodon
tener, but due to crushing and displacement, the positions of
those teeth are not certain. The two anterior teeth, A and B
(McKenna 1968: Fig. 2), are both large and single-rooted and
could potentially be the canine. A does not have a posterior
cuspule and is morphologically similar to the upper canine of
the P. rosei specimen UF 303729 (Fig. 2b), whereas B does

J Mammal Evol (2015) 22:307342

317

Fig. 3 Micro-CT scan images of left maxilla of Plagioctenodon

thewisseni with C1-M3 (UF 294696). a, Labial view. b, Occlusal view

have a posterior cuspule and resembles the upper canines of

both other P. rosei specimens and P. thewisseni.
The morphology of the anterior upper premolars is very
similar between P. rosei (Fig. 2) and P. thewisseni
(Figs. 3 and 4), with the P1 and P2 both double-rooted
and having a large, triangular, labio-lingually flattened
anterior cusp and a small posterior cuspule. There is
also a significant increase in size between the P1 and P2
in both taxa. In the upper anterior teeth described by
McKenna (1968), C and D are most similar in
morphology to the P1 and P2, respectively, of known
nyctitheriid taxa, as was predicted (McKenna 1968).
The P 3 morphology does not appear to be highly
constrained in nyctitheriid species. In P. rosei (Fig. 2), the
presence of a metacone is variable; it is present as a small
swelling on the postparacrista in UF 303728 (Fig. 2a) and UF
303730, whereas it is absent in UF 303729 (Fig. 2b). In the
two specimens with a P3 known for P. thewisseni (Figs. 3 and
4), the metacone is consistently present, but the morphology
of the protocone varies. The P3 protocone of the holotype
(UM 86725: Fig. 4) is considerably smaller and less lingually
extended than the protocone in UF 294696 (Fig. 3). Multiple
P3s are documented for C. beatus (Sig 1976: Fig. 61) that all
lack metacones but vary greatly in shape. The P 3 of
Cryptotopos hartenbergi (Sig 1976: Fig. 47) is dissimilar in
morphology to those of C. beatus and strongly resembles the P3s seen in most Plagioctenodon specimens,
with a clearly defined metacone and a small protocone
that is not lingually extended.
At least one P3 is documented for Acrodentis rosenorum
(Christiansen and Stucky 2013: Fig. 2a-b), Nyctitherium
krishtalkai (Christiansen and Stucky 2013: Fig. 2c-d), and
S. gracilis (Sig 1976: Figs. 3 and 11). Like C. beatus, these
three taxa do not possess a metacone on the P3. Tooth E of

Fig. 4 Micro-CT scan images of left (P2-M3) and right (P2-M3) maxillae
of Plagioctenodon thewisseni (UM 86725). a, Left and right palate
articulated in occlusal view. b, Left palate in occlusal view. c, Left
palate in lingual view. d, Right palate in occlusal view. e, Right palate
in lingual view

L. tener (McKenna 1968: Fig. 2) is almost certainly the P3; it is

dominated by the paracone and lacks a metacone, but it is
unusual in that it does not appear to have a distinct protocone.
The lack of a protocone on the P3 separates L. tener from most
other nyctitheriids and may represent the ancestral state, because this is the oldest nyctitheriid known with a P3. However,
it is important to note that the P3 of the L. tener holotype was
destroyed in preparation and it is impossible to verify this
morphology other than through examination of a photograph
(McKenna 1968: Fig. 1). The only other nyctitheriids that lack
a protocone on the P3 likely lost that cusp secondarily. The P3
of the European Cryptotopos communis from the Oligocene
consists mostly of a large paracone with little to no parastyle
or metastyle and a small lingual bulge (Ziegler 2007: Fig. 3.3).
The late Eocene European Paradoxonycteris soricodon is

318

reported as having a single-rooted, unicuspid P3 (Revilliod

1922), although that tooth has since been lost (Hooker and
Weidmann 2000).
The morphology of the P. rosei I1 (Fig. 2b) differs from that
of the other two nyctitheriid species with known I1s, A. cayluxi
and Ceutholestes dolosus. The I1 of A. cayluxi is longer and
taller than its I2 and I3 and it appears to have a single
large anterior lobe with a small accessory labial cuspule
(Sig 1976: Fig. 95A-C), whereas in C. dolosus, the I1
is shorter in both length and height than the I2 and it
has four lobes (Rose and Gingerich 1987: Figs. 1, 2 and
3).
The morphology observed in the I2 is the same as that
reported for P. rosei by Rose et al. (2012), in which they
document five lobes with the posterior-most one being small
and acute. We have found in our sample size that the fifth cusp
is sometimes absent, although this is likely due to wear or
damage. Whereas other nyctitheriids exhibit a very similar
morphology, all other taxa outside the Plagioctenodon genus
with known I2s, including Ceutholestes (Rose and Gingerich
1987: Figs. 1, 2 and 3), S. carbonum (Sig 2001: Fig. 1),
E u ro n y c t i a g r i s o l l e n s i s ( S i g 1 9 7 6 : F i g . 2 7 ) ,
Amphidozotherium (Sig 1976: Fig. 95), and tentatively
L. donkroni (Rose et al. 2012: Fig. 21I-J), only possess four
lobes on that tooth, lacking the acute, posterior-most cuspule.
A possible exception is C. communis, which was attributed I2s
with both four and five lobes, although it was also suggested
that the lower incisor with five lobes could represent either the
not-yet-identified I1 morphology or is an I2 exhibiting an
autapomorphy (Ziegler 2007). The incisor with five cusps
(Ziegler 2007: Fig. 2.5) does not look very similar to the
morphology of I1s from other nyctitheriid taxa and bears a
striking resemblance to the I2 of P. rosei. It is much more
likely that the I2 of C. communis is polymorphic or the fourand five-lobed incisors represent I 2 s of two different
nyctitheriid taxa. Rose et al. (2012) commented that the line
drawing of A. cayluxi in Sig (1976: Fig. 95) appears to
exhibit a short, fifth cuspule at the posterior base of the tooth,
but the text states that the taxon only has four cuspules. The
only two nyctitheriids with definitive five-lobed I2s are
P. rosei and P. thewisseni, representing a potential synapomorphy for the genus.
The I3 of S. gracilis (Sig 1976: Fig. 4) and S. grandis (Sig
1976: Fig. 69) is nearly identical to that of P. rosei (Fig. 2) in
having three lobes, but C. beatus (Sig 1976: Fig. 56),
C. communis (Ziegler 2007: Fig. 2.7), A. cayluxi (Sig
1976: Fig. 95), and S. carbonum (Sig 2001: Fig. 1) are
all more similar to that of P. thewisseni (Fig. 5) in
having just two lobes. The I3 of P. thewisseni is very
similar to that of the more worn P. rosei specimens,
suggesting that its unworn state might have three lobes
in similar configuration to those seen in P. rosei. Therefore, the
two lobes found in some of the taxa may be a true

J Mammal Evol (2015) 22:307342

morphological state or it could be due to wear on the anterior

two lobes that make them appear to be merged.
The large, procumbent anterior cusp and small posterior
cuspule of the lower canine of P. rosei (Fig. 2b) is shared with
S. gracilis (Sig 1976: Fig. 5), S. grandis (Sig 1976: Fig. 70),
S. carbonum (Sig 2001: Fig. 1), C. beatus (Sig 1976:
Fig. 55), C. hartenbergi (Sig 1976: Fig. 55), C. communis
(Ziegler, 2007: Fig. 2.8), and A. cayluxi (Sig 1976: Fig. 95).
Ceutholestes dolosus also has a similar morphology, although
the area in which the basal cuspule is located is more of a
flattened shelf (Rose and Gingerich 1987: Fig. 1). The canines
of these nyctitheriids are unlike those of Maelestes gobiensis
and Adunator minutus, which both lack the posterior shelf.
The lower canines of Nyctitherium velox and Nyctitherium
krishtalkai have been described as primitive (didelphoid)
(Robinson 1968: 130) and round in cross-section, primitive,
and single-rooted (Christiansen and Stucky 2013: 7), respectively. Both of these teeth appear to be damaged and therefore
we cannot assess whether they were more similar in morphology to the other nyctitheriids or the outgroup taxa used in our
cladistic analysis.
DiscussionLeptacodon rosei was transferred to
Plagioctenodon by Beard and Dawson (2009) based on characters of the third and fourth lower premolars including a
relatively smaller, anteriorly canted P3 and a relatively high
paraconid on the P4 (see genus discussion of Plagioctenodon in
this paper). The species is primarily known from the
Clarkforkian (Gunnell et al. 2008) but it may be present in the
Tiffanian (Secord 2008) and the Wasatchian (Rose et al. 2012).
The largest sample of P. rosei, including the holotype, was
recovered from the Cf-2 locality SC-188. Although most of
the fossils from SC sites were collected via surface
prospecting, this site was screen-washed, which led to the
recovery of some of the most complete jaws known for this
species (Rose 1981; Gingerich 1987), including at least one
dentary with I1-P4 (UM 77030). Lower incisors are also
reported in the specimens USNM 539484 and UM 82389 by
Rose et al. (2012). Although some anterior teeth of P. rosei
have been found by screen-washing and surface prospecting,
those recovered from limestones are unique in being part of
exceptionally complete specimens, often with the upper and
lower jaws in association and the full dental formula preserved
(e.g., Fig. 2).
Plagioctenodon thewisseni sp. nov.
(Figs. 2 and 3, Table 2)
Cf. Plagioctenodon krausae: Rose, 1981:41, Figs. 12
and 13; Gingerich, 1987:304
Plagioctenodon bowni: nomen nudum, Bloch, 2001:65,
Fig. 5
EtymologyNamed for Professor J. G. M. Thewissen of
Northeast Ohio Medical University who in 1986 as a graduate

J Mammal Evol (2015) 22:307342

319

Fig. 5 Micro-CT scan images of

left (I2-P1, P3-M3) and right (C1M3) dentary of Plagioctenodon
thewisseni (UM 86725). a, Left
dentary in labial view. b, Left
dentary in lingual view. c, Left
dentary in occlusal view. d, Right
dentary in labial view. e, Right
dentary in lingual view. f, Right
dentary in occlusal view

student working as a member of the University of Michigan

field party, discovered the freshwater limestone nodule from
which the holotype was extracted. Other notable fossils from
that same nodule include a remarkable skull and skeleton of
Plesiadapis cookei (Gunnell and Gingerich 1987; Gingerich
and Gunnell 1992, 2005). The holotype (UM 86725) of
P. thewisseni was first mentioned as an exceptionally
complete nyctitheriid by Gingerich (1987) and referred
to Cf. Plagioctenodon krausae. It was later discussed
in the dissertation of J. I. Bloch (2001) in which it was referred
to a new species Plagioctenodon bowni as a nomen nudum.
Since that time, the specific name bowni has been used in
another closely related nyctitheriid species name
(Plagioctenoides tombowni; Rose et al. 2012) and thus not
used again here to avoid confusion.
HolotypeUM 86725: Left dentary with crowns of I2-P1
and P3-M3; right dentary with crowns of C1-M3; right and left
maxillae with crowns of P2-M3 (Figs. 2 and 3).
Type LocalityUniversity of Michigan (UM) Locality
SC-117, Willwood Formation, Clarks Fork Basin, Wyoming

(See [Rose 1981] for more detailed locality information) from

the late Clarkforkian part of the Paleocene epoch (Copecion
interval zone [Cf-3], approximately 56 Ma; Secord 2008).
Referred SpecimensSC 117: UM 39873, right (R)
dentary with C1, P3-M2. SC 62: UM 83931, R dentary with
broken P2, P3-M3; UM 82576, R edentulous dentary with
roots for M1 and P3, associated with loose R C1, broken R
P1, R M2-M3 (trigonid), R I1, and R maxilla with I2-I3. SC 29:
UM 76906, left (L) I2; UM 76920, R M2 labial fragment. SC
327: UF 289746, R I2, R P4, L C1, partial skeleton; UF
289747, R maxilla with P3-M3, L maxilla with P4-M3, L
dentary with P1-M3, partial skeleton; UF 294696, L maxilla
with C1-M3, R P4, R dentary with M1-M3.
DiagnosisDiffers from all other Plagioctenodon species
in being smaller, having larger and more anteriorly projecting
paraconids on P34, a more anteroposteriorly elongated P4,
and a more reduced precingulum on the upper molars. It
further differs from P. dormaalensis in having more acute
cusps, larger, more anterolingually shifted molar paraconids,
taller entoconids than hypoconids, and two mental foramina

320

rather than one. It further differs from P. rosei in having more

elongate lower premolars with a greater exaggeration in the
anterior cant of the protoconids. It further differs from
P. krausae and P. savagei in having narrower molar talonids
relative to the trigonids, and more anteriorly positioned
entoconids in relation to the hypoconulid.
Age and Distribution All localities are located in the
Willwood Formation, late Clarkforkian North American
Land Mammal Age Copecion interval zone (Cf-3),
which spans 56.22-55.8 Ma (Secord 2008). UM localities SC-117 (at the stratigraphic level of 1370 m above
the Cretaceous-Paleogene boundary in the Bighorn
Basin), SC-62 (1380 m); SC-29 (1385 m); SC-327
(1420 m) of the Clarks Fork Basin, Wyoming, United
States of America.
Measurementssee Table 2.
DescriptionThe holotype is relatively complete and
shows that P. thewisseni has small, sectorial teeth set in a
shallow, elongate dentary. The lower dental formula is 3-14-3 and the lower molar series shows a slight decrease in size
from M1 to M3. The dentary contains two mental foramina;
the anterior foramen is located beneath the P2 and the posterior
foramen is below the posterior root of the P3.
The left dentary preserves the crowns of I23 and an alveolus for I1. The lower incisors appear to be similar in size and
are procumbent and multi-lobed or scalloped. The tip of I2 is
broken but there is a small, bulbous cuspule at the
posterolingual extent of the tooth crown with two lobes situated anterobucally above it that curve anteriorly. A complete
I2 referred to P. thewisseni (UM 76906) was first described by
Gingerich (1987) as Cf. Plagioctenodon krausae in which
he stated that it has five cusps and is similar in appearance to
that of the European nyctitheriids Saturninia and
Amphidozotherium. The tooth has since been broken, but after
examination, we agree with this assessment. The third
incisor is more worn than the second, but it appears to
have been slightly smaller than I2 and had at least two
lobes.
The lower canine is procumbent and only slightly
enlarged relative to the incisors and P1. It is dominated
by one large cusp that is laterally compressed and more
spatulate than conical, but also has a small posterior
basal cuspule.
The P1 is similar in shape to that of the canine; it is
procumbent with a large, squared-off spatulate anterior cusp
and a much smaller posterior basal cusp. The P2 and P3 are
similar in shape but the former is larger than the latter. They
are both double-rooted and dominated by a large, anteriorly
canted protoconid. There is a small, anteriorly projecting
paraconid located on the anterior surface of the protoconid,
situated relatively high on the P23, about halfway between the
base of each tooth and the tip of the protoconid. A crest runs
posteriorly from the tip of the protoconid to the base of the

J Mammal Evol (2015) 22:307342

crown, where it ends in a single posterior cusp. There is no

talonid basin on either the P2 or P3.
The P4 is elongate, with a length almost twice that of the
width. It is semi-molariform with a large, anteriorly projecting
paraconid that arises high on the anterior face of the trigonid,
similar to the condition in other species of Plagioctenodon.
The paraconid is joined to the protoconid by a v-shaped
paracristid below which, on the buccal surface of the crown,
there is a slight anterior bulge that occupies a similar area of
the precingulid as on the lower molars. The metaconid is
relatively large, being only slightly smaller and shorter than
the protoconid. The talonid is nearly equal in width to that of
the trigonid, although slightly shorter anteroposteriorly. It has
three distinct cusps. The hypoconid and entoconid are
subequal in size but the latter is slightly taller. The
hypoconulid is smaller than the other talonid cusps and is
situated midway between them and slightly posterior. The
cristid obliqua is strong, extending from the hypoconid and
climbing about halfway up the posterior wall of the trigonid
where it ends immediately below the notch in the protocristid.
The trigonid is slightly narrower than the talonid on the M1,
subequal in width on the M2, and wider than on the M3. The
paraconids of the lower molars project anteriorly and are
situated lingual to the midline on the trigonid. A robust
precingulid is present on the anterolabial surface of the molars,
terminating posteriorly below the protoconid. The metaconid
and protoconid are subequal in height. The base of the
protoconid has a larger circumference than that of the
metaconid of M1, whereas the protoconid and metaconid are
subequal in circumference for the M2 and M3. The talonids of
the lower molars have three distinct cusps; the entoconid is
similar in length and width to the hypoconid, whereas the
hypoconulid is somewhat smaller than both. The entoconid is
taller than the hypoconid and is positioned anterolingually to
the hypoconulid. The hypoconulid is positioned slightly lingual to the midline. The cristid obliqua extends from the
hypoconid and climbs a short way up the posterior surface
of the protoconid, just labial to the notch in the protocristid.
Associated crowns of I13 were recovered from a limestone
in association with lower teeth that are referred to P. thewisseni
(UM 82576). The upper incisors decrease in size posteriorly
and are similar to each other in morphology, with a large
anterior cusp that is labiolingually compressed and asymmetrically shaped with the anterior edge bulging anteriorly, and a
small posterior cuspule. The C1 of P. thewisseni is singlerooted and the crown is about twice the height of the P1, has
a large primary cusp with a slight posterior curve at the crest,
and a small posterior basal cuspule. Other than its smaller size,
the P1 is nearly identical to the P2; both have a large, erect
paracone and a much smaller posterior basal cuspule. The P3
has a conical paracone with a strong postparacrista from
which a small metacone swells. A crista continues posteriorly
from the metacone and extends to a metastyle that is only

J Mammal Evol (2015) 22:307342

slightly smaller than the metacone. There is a distinct parastyle

that is not connected to the paracone. The protocone is small
and similar in size to the metacone; it is situated closely to the
posterolingual side of the paracone.
The P4 paracone and metacone are appressed to one another and similar in size, but the paracone is taller. There is a
distinct, conical parastyle that is separated from the paracone
and a similar-sized metastyle that is connected to the metacone
by a postmetacrista. The stylar shelf is shallow with no discernible ectoflexus and oriented anterolinguallyposterobuccally. The protocone is shifted anteriorly to the
paracone. A small trigon basin is present but there are no
conules. There is a small precingulum and postcingulum,
whereas the hypocone is vestigial to absent.
The M1 and M2 are similar in form with the M1 being
slightly larger. The paracone and metacone are subequal in
size and height, joined by a rectilinear centrocrista. The stylar
shelf is shallow with the parastyle and metastyle as the only
stylar cusps. The ectoflexus of the M2 is deeper than that of
M1. There is a small conical parastyle that is anterolabial to the
paracone and connected to it by a small preparacrista. The
parastyle projects anteriorly on the M1 and more anterolabially
on the M2. The metastyle is similar in size to the parastyle and
is posterolabial to the metacone, connected to it by a strong
postmetacrista. The paraconule and metaconule are present
and strongly winged by their respective pre- and post- conule
cristae. The paraconule is located anterior to the paracone and
protocone and situated closer to the latter. The preparaconule
crista forms an anterior ridge on the crown that extends
labially, almost reaching the parastyle, whereas the
postparaconule crista terminates at the base of the
paracone. The metaconule is located about halfway between the metacone and protocone and the
premetaconule crista ends at the anterior edge of the
metacone, whereas the postmetaconule crista extends
posterolabially, stopping just short of the metastyle.
The protocone is situated lingual to the paracone and
projects anteroventrally. A small precingulum and
postcingulum are present, the former extends from the
protocone to the paraconule and the latter from just
lingual to the protocone to the metaconule. The
postcingulum has a small, conical hypocone that is
situated posterolingual to the protocone. The M3 has a
taller paracone than metacone, a prominently projecting
parastylar shelf, no metastyle, distinct but small paraand metaconules, a small precingulum, and no
postcingulum.
Some of the referred specimens (UF 294696, UF
289746, UF 289747) include cranial and/or postcranial
elements outside of the scope of this study and are not
described here.
ComparisonFor the most part, nyctitheriids for which it
can be documented have a 3-1-4-3/3-1-4-3 dental formula. It

321

is possible that the holotype of L. tener had five upper premolars (McKenna 1968; Krishtalka 1976), but the specimen is
badly crushed and not fully developed (McKenna 1968). The
condition of two mental foramina in the dentary appears to be
the primitive condition for Nyctitheriidae; the reduction to a
single foramen is derived in some clades. For those species
that have at least one dentary complete enough to be assessed,
the Asian nyctitheriids, all species classified as Leptacodon,
Wyonycteris, Plagioctenoides, Ceutholestes, and
Limaconyssus, and most species of Nyctitherium and
Plagioctenodon, including P. thewisseni, have two mental
foramina. One of the foramina has been lost in Nyctitherium
velox, Acrodentis rosenorum, Plagioctenodon dormaalensis,
and all of the late Eocene European nyctitheriids, including
Amphidozotherium, Saturninia, Cryptotopos, Scraeva, and
Euronyctia. Although most of the specimens of P. rosei have
a similar condition to that of P. thewisseni, with mental
foramina below P2 and the posterior root of P3, at least
two of them (UM 76498 and UF 303729) only have a
single, larger mental foramen centered under the anterior
root of the P3. The nyctitheriid taxa with only one
foramen all have it similarly positioned below the P3
in contrast to the variable position of the mental foramina in nyctitheriid taxa with two.
For comparisons of I1-P3 and I1-P1, see the corresponding
section for P. rosei. The morphologies of the P2 and P3 (Fig. 5)
are typical for nyctitheriids in having a single large protoconid
with a smaller anterior and posterior cusp. Nyctitheriids that
vary from this basic configuration are Ceutholestes dolosus
(Rose and Gingerich 1987: Fig. 1) and Placentidens lotus
(Russell et al. 1973: Fig. 12D), which both have an additional
cusp, a small metaconid appressed to the protoconid on the P3.
Some nyctitheriids, including Wyonycteris richardi (Smith
1995: Fig. 2), both Plagioctenoides species (Rose et al.
2012), Amphidozotherium cayluxi (Sig 1976: Fig. 95), and
Eosoricodon terrigena (Lopatin 2005: Fig. 2), exhibit a more
simplified morphology in which the P2, and sometimes the P3,
may lack the anterior cusp or is single-rooted rather than
double-rooted. The asymmetrical, anterior bulging of the
protoconid on both premolars is a variable trait in the family,
however. This condition is seen in several clades, including all
Plagioctenodon species (Fig. 5; Beard and Dawson 2009),
Leptacodon acherontus (Secord 2008: Fig. 33), Leptacodon
choristus (Secord 2008: Fig. 36), W. richardi (Smith 1995:
Fig. 2), most Saturninia and Cryptotopos species (e.g., Sig
1976), and in Oedolius perexiguus (Lopatin 2006: plate 4,
Fig. 1), whereas many other nyctitheriids and the outgroup
taxa have a more symmetrical, erect protocone. The smaller
size of the P3 compared to that of the surrounding premolars
appears to be a derived trait that occurs only in the
Plagioctenodon, Wyonycteris, Plagioctenoides, and late-occurring European nyctitheriid clades and may have
evolved just once in the familys history (see Node L

322

J Mammal Evol (2015) 22:307342

Fig. 6 Strict consensus of two

most parsimonious trees for the
phylogenetic analysis of
Nyctitheriidae. Tree length = 330
steps; CI = 0.282; RI = 0.601.
Bremer supports are indicated by
the number above the lines and
the letter below the lines represent
key nodes listed in Appendix 2

in Fig. 6 and Appendix 2). In all other nyctitheriids, the

lower premolars generally increase in size posteriorly.
The P4 of Plagioctenodon exhibits one of the only
definitive synapomorphies for the clade, a relatively
high P4 paraconid, but this trait does occur in the other
non-Plagioctenodon nyctitheriid taxa Leptacodon
choristus (Secord 2008: Fig. 36), Ceutholestes dolosus
(Rose and Gingerich 1987: Fig. 1), Limaconyssus
habrus (Gingerich 1987: Fig. 20), Asionyctia guoi
(Missiaen and Smith 2005: Fig. 1), and in some
Leptacodon munusculum specimens (e.g., Gingerich
et al. 1983: Fig. 2c; Krause and Gingerich 1983:
Fig. 11). In other nyctitheriids the P4 paraconid is lower
on the anterior surface of the trigonid. The P 4 of
P. thewisseni is semimolariform, with a relatively large
metaconid and a clearly defined talonid basin that is
nearly the same width as the trigonid, just like the P4s
of the other Plagioctenodon species. Some variation
exists in the number of the talonid cusps, however.
Although all known specimens for P. thewisseni,
P. dormaalensis (Smith 1996), and P. savagei (Bown
and Schankler 1982) have three talonid cusps, the samples for P. rosei and P. krausae include P 4 s with
talonids that have both two and three cusps (see Bown

1979; Rose 1981). The last two species also have the
largest sample sizes in Plagioctenodon, so it is possible there
may also be some two-cusped P4 talonids found for the other
Plagioctenodon species upon an increase in sample size. The
P4 of P. thewisseni is unique among other Plagioctenodon
species and most other nyctitheriids in having a noticeably
lower width to length ratio: 0.47 and 0.46 for the holotype and
averaged across all referred specimens, respectively (calculations made from Table 2). This makes it one of the few
nyctitheriid species with a length to width ratio below 0.5;
the only other nyctitheriids are Leptacodon proserpinae (Van
Valen 1978), Leptacodon tener (McKenna 1968: Fig. 4),
Wyonycteris primitivus (Beard and Dawson 2009), Edzenius
lus (Lopatin 2006: table 18), and Oedolius perexiguus
(Lopatin 2006: table 15). A low P4 width to length ratio is
likely a primitive trait for Nyctitheriidae, because this is the
state of the outgroup (Maelestes gobiensis [Wible et al. 2009:
table 1]), the oldest proposed nyctitheriid (L. proserpinae),
one of the oldest definitive nyctitheriids (L. tener), a
nyctitheriid exhibiting unusually primitive characters for its
clade (W. primitivus, see Beard and Dawson 2009), and two
Asian nyctitheriids (E. lus and O. perexiguus). The presence
of this trait in P. thewisseni is not necessarily retention of a
primitive feature, however, because the extreme anterior

J Mammal Evol (2015) 22:307342

projection of the paraconid on the P4 accounts for part of the

unusual length of the tooth.
The lower molars are generally similar between the
Plagioctenodon species. P. thewisseni has larger and more
anteriorly projecting paraconids than the other species
(Fig. 5), but otherwise its molars look nearly identical to those
of P. rosei (Rose 1981: Fig. 11). The lower molars of
P. thewisseni, P. rosei, P. krausae, and P. savagei differ from
P. dormaalensis in having taller entoconids than hypoconids,
whereas they are equal in height in P. dormaalensis (Smith
1996). The lower molars of P. krausae and P. savagei differ
from those of other Plagioctenodon species in having the
entoconid located along the posteriormost edge of the
crown (Bown and Schankler 1982); it is more anteriorly
located in the other taxa. Although the lower molars of
nyctitheriids can generally be classified as having
subequal (within 5%) trigonid and talonid widths, there
are subtle differences in the proportions of the trigonid
and talonid within the tooth row. The M1 and M2 of
P. thewisseni are very similar in morphology, but the
trigonid is slightly narrower than the talonid in the M1,
whereas it is subequal in the M2. This trend is shared
with P. rosei (Rose 1981: table 4) and some
P. dormaalensis (Smith 1996: Figs. 2 and 3) specimens,
as well as Nyctitherium velox (Krishtalka 1976: table 3),
Ceutholestes dolosus (Rose and Gingerich 1987: table
1), Placentidens lotus (Russell et al. 1973: Fig. 12),
Wyonycteris chalix (Gingerich 1987), and Wyonycteris
richardi (Smith 1995: tableau p. 928).
The P4 of P. thewisseni is nearly identical to those of
P. rosei and P. dormaalensis. All three have a small
postcingulum, but P. dormaalensis has a clearly-defined,
small hypocone, whereas P. thewisseni and P. rosei do
not.
The upper molars of P. thewisseni differ from those of
P. rosei, P. krausae and P. dormaalensis in having a smaller
precingulum, but the size of the precingulum is quite variable
across the family (Appendices 34: Character 55).
Otherwise, the species classified as Plagioctenodon all
have very similar upper molars that can be differentiated
from other nyctitheriid clades mostly by their lack of
derived features, including 1) the mesostyle otherwise
f o u n d i n a l l Wy o n y c t e r i s s p e c i e s o t h e r t h a n
W. primitivus, 2) the dilambdodonty otherwise seen in
Pontifactor bestiola, Wyonycteris chalix, Wyonycteris
richardi, Paradoxonycteris, and the Euronyctia species,
3) the pericone otherwise found in Wyonycteris,
Cryptotopos, Placentidens lotus, and several Saturninia
species, 4) a reduction in para- and metaconule size and
winging otherwise seen in Nyctitherium and many late
Eocene European nyctitheriids, and 5) a greatly expanded postcingulum otherwise found in Nyctitherium,
Pontifactor, and the late Eocene European nyctitheriids.

323

Discussion Plagioctenodon thewisseni is similar to other

Plagioctenodon species in having an anteriorly inclined
protoconid on P2, a reduced P3 with an anteriorly canted
protoconid and a P4 paraconid that is positioned relatively high on the trigonid. Plagioctenodon thewisseni is
one of the smallest nyctitheriids known, which may
explain its limited sample size in the otherwise fossiliferous localities of the Clarks Fork Basin. The body size
of P. thewisseni is estimated to be 5.01 g and 7.49 g
(Table 3) based on the M1 and M1 crown areas, respectively, using the regression functions of Bloch et al.
(1998). The true body mass is likely to be closer to
the estimate provided by the M1 because that tooth is
argued to be a better mass estimate than the M 1
(Gingerich 1982). Plagioctenodon thewisseni is the
smallest Plagioctenodon species, with a body mass
~30% less than that of the next smallest species,
P. dormaalensis, which is estimated here to be 7.17 g
using M 1 dimensions. The only North American
nyctitheriids with smaller estimated body masses than
P. thewisseni based on the M1 are Leptacodon donkroni,
Plagioctenoides microlestes, and Saturninia mamertensis.
Based on the M1, Saturninia gracilis and Wyonycteris microtis
are smaller, although S. gracilis is estimated to be larger using
the M1 equation and W. microtis may not be a nyctitheriid
based on the results of our cladistic analysis. Plagioctenoides
tombowni is likely to be smaller than P. thewisseni, as well;
although there is not a complete M1 or M1 known for
P. tombowni, other teeth are nearly the same size as those of
P. microlestes (Rose et al. 2012).
Fossils similar to those of P. thewisseni have been in
museum collections for over thirty years. Rose (1981)
described an isolated P4 (UM 69942), M1 (UM 71689),
and M2 (UM 71686) from late Clarkforkian localities in
the Clarks Fork Basin and suggested that they represented a slightly smaller Clarkforkian predecessor of
early Wasatchian P. krausae. These teeth were referred
to as cf. Plagioctenodon krausae (Rose 1981:42) and
they could be included in the P. thewisseni hypodigm
based on their similar temporal range, size, and morphology (see Rose 1981: Fig. 1213). Additionally,
Gingerich (1987) referred half of an upper molar and
an I2 to Cf. Plagioctenodon krausae, stating that it
resembles Leptacodon as much as it does
Plagioctenodon, and definitive placement of all of these
specimens must await study of the new material
(Gingerich 1987:304). Finally, Secord (2008) referred
several isolated teeth from the Y2K Quarry (Ti-5b) to
Plagioctenodon sp. and noted that they closely resemble
the Cf. Plagioctenodon krausae of Rose (1981) and
Gingerich (1987). We agree that the lower molars have
strong similarities to P. thewisseni, particularly UM
109346, but the lack of the diagnostic P4, the overly

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Table 3 Size of nyctitheriid species in area of the M1 and M1 and in

estimated weight. Tooth size of the M1 and M1 were calculated by
multiplying the averaged lengths and widths of each those teeth

published in the literature for a given nyctitheriid taxon. Estimated

weight and 95% confidence intervals for nyctitheriid species are given
in grams and calculated based on equations from Bloch et al. (1998)
M1

M1
n

Tooth size (mm2)

Estimated weight (g)

with 95% CI

Tooth size (mm2)

Estimated weight (g)

with 95% CI

Plagioctenodon krausae

1.75

13.95 1.36

Plagioctenodon savagei
Plagioctenodon rosei

1
3

2.70*
1.49

28.30 2.36
10.72 1.67

0
4

3.60

21.74 1.58

18

1.16

7.17 1.30

10

2.12

8.77 1.38

Red Hot P. dormaalensis

Plagioctenodon thewisseni

1
5

1.06
0.93

6.20 2.40
5.01 1.54

0
3

1.93

7.49 1.72

Leptacodon tener

1.31

8.71 1.68

Leptacodon munusculum
Leptacodon catulus

3
1

1.18
1.17

7.38 1.68
7.25 2.39

0
0

Leptacodon packi
Leptacodon nascimentoi

5
3

1.56
0.99

11.62 1.50
5.53 1.70

1
1

2.57
2.02

12.20 2.48
8.10 2.49

Leptacodon acherontus

2.82

30.38 1.84

Leptacodon donkroni
Leptacodon choristus

1
1

0.70*
3.58

3.14 2.43
44.83 2.36

1
0

1.49

4.78 2.51

3
10
5

1.94
1.60
1.19

16.57 1.66
12.08 1.35
7.48 1.52

6
12
2

3.52
2.28
2.38

20.96 1.45
9.96 1.34
10.70 1.91

Nyctitherium christopheri
Acrodentis rosenorum

0
1

1.13

6.81 2.40

1
1

4.08
2.30

27.01 2.46
10.08 2.48

Ceutholestes dolosus
Limaconyssus habrus

2
1

3.28
1.30

38.88 1.83
8.61 2.39

0
0

Pontifactor bestiola
Wyonycteris chalix
Wyonycteris richardi

0
2
21

1.06
1.00

6.20 1.88
5.65 1.30

10
2
4

2.88
2.18
1.71

14.87 1.35
9.19 1.92
6.07 1.63

1
2
0
1
6

1.52
2.06

11.14 2.38
18.19 1.84

1.36

4.09 2.52

0.83
2.86

4.11 2.42
31.10 1.42

0
0
1
0
2

2.55

12.04 1.91

Saturninia gracilis
Saturninia mamertensis
Saturninia rigasii
Saturninia ceciliensis
Saturninia pelissiei
Cryptotopos carbonum
Crypototopos beatus
Cryptotopos woodi
Cryptotopos hartenbergi
Scraeva hatherwoodensis

14
2
1
1
0
1
15
3
5
1

1.23*
0.85*
2.05
1.75*

7.85 1.32
4.34 1.90
18.03 2.37
13.94 2.37

1.90
1.35

7.32 1.50
4.05 2.52

3.43

20.04 2.46

1.94*
2.45*
2.38
1.82
1.65

16.48 2.37
24.17 1.26
23.10 1.65
14.86 1.49
12.70 2.38

3.90

24.96 1.69

Placentidens lotus
Euronyctia montana
Euronyctia grisollensis
Paradoxonycteris soricodon
Asionyctia guoi
Bumbanius rarus

1
0
1
0
16
6

1.30

8.61 2.39

2.80*
2.88

14.16 2.47
14.88 2.47

1.31

8.75 2.39

1.18*
2.27

7.39 1.31
21.28 1.43

2.53
2.46
5.27

11.87 2.48
11.32 1.39
41.84 2.45

Plagioctenodon dormaalensis

Nycitherium velox
Nyctitherium serotinum
Nyctitherium krishtalkai

Wyonycteris primitivus
Wyonycteris galensis
Wyonycteris microtis
Plagioctenoides microlestes
Amphidozotherium cayluxi

6
1
0
0
1
0
3
0
0
0
1
1
0
1
9
1

J Mammal Evol (2015) 22:307342

325

Table 3 (continued)
M1

M1
Eosoricodon terrigena
Oedolius perexiguus
Edzenius lus

0
10
2

1
1.11
1.31

6.64 1.38
8.68 1.87

2.30

10.07 2.48

0
0

n number of specimens used in the calculations

*denotes values that were calculated using a specimen either with an estimated length or width

elongate P3, and the large temporal range extension that

would occur if the Y2K Quarry specimens were included in the new species causes us to refrain from adding
the Y2K Plagioctenodon to the referred specimens. The
fossils included in Cf. Plagioctenodon krausae and
Plagioctenodon sp. consist mostly of isolated teeth and
it has only been with the preparation of fresh water
limestone that there has been sufficient fossils to diagnose this new species of Plagioctenodon.
Beard and Dawson (2009) reported a small
Plagioctenodon species from the Red Hot Local Fauna of
the early Eocene of Mississippi and referred it to
P. dormaalensis based on the morphology of an isolated M1
and M2. We estimate the mass of the Red Hot Plagioctenodon
species to be 6.20 g based on the M1, a mass intermediate of those for P. thewisseni and P. dormaalensis
(Table 3). Although a similarly sized Plagioctenodon
is now known in North America, the Plagioctenodon
from Mississippi still appears to share closer affinities
with P. dormaalensis. The M1 paraconid is less lingually shifted and less anteriorly projecting than the condition seen in P. thewisseni and all of the cusps are less
acute. The M2 of P. dormaalensis and the Red Hot
Plagioctenodon exhibits less expanded parastylar and
metastylar lobes and a larger hypocone situated on a
more expanded postcingulum in comparison to
P. thewisseni. Larger sample sizes and associated dentitions, particularly from the Red Hot Local Fauna, are
necessary to determine if the significant morphological
differences between P. thewisseni and the Red Hot
Plagioctenodon represent intraspecific variation. If the
taxon from the Red Hot Local Fauna is P. dormaalensis, it
would represent the only nyctitheriid species on two separate
continents. Such a broad paleobiogeographic distribution is
not inconsistent with the hypothesis discussed below of multiple dispersals of nyctitheriid clades between North America
and Europe.
Recognition of P. thewisseni increases the late Paleocene
diversity of nyctitheriids in the Clarks Fork Basin to six
species: P. thewisseni, P. rosei, Wyonycteris chalix,
Ceutholestes dolosus, Limaconyssus habrus (Gunnell et al.
2008), and an unnamed taxon alternately called cf. Pontifactor

bestiola (Rose 1981) or cf. Wyonycteris sp. (Gingerich 1987).

Rose et al. (2012) noted several other sympatric occurrences
of nyctitheriids during the Paleogene, drawing a comparison
with the overlapping geographic ranges of modern shrew
species. Although it might be reasonable to expect some
overlap in use of resources in such a speciose, sympatric
assemblage of small-bodied, insectivorous mammals, the dental morphologies of these six taxa are remarkably similar with
no obvious evidence for niche-partitioning and, therefore,
cranial and postcranial evidence may be necessary to address
this paleoecological question.

Results
The new technology search in TNT yielded four most parsimonious trees (MPTs) of 330 steps and the traditional search
was unable to find any additional trees. The strict consensus
(Fig. 6) contains four nodes with polytomy. The consistency
index is 0.282 and the retention index is 0.601. The Bremer
supports for all of the clades are very low (Fig. 6), which is not
surprising because of the low number of characters relative to
number of taxa.
Results from this analysis suggest that Tiffanian
Wyonycteris microtis shares a closer relationship with
taxa included in the outgroup, and thus does not support
its identification as a nyctitheriid. Instead, two
erinaceomorph taxa, Macrocranion junnei and
Adunator minutus, are more closely related to the rest
of a monophyletic Nyctitheriidae. The most primitive
nyctitheriids are the five included Asian taxa, consecutively nested in a paraphyletic clade outside the North
American and European nyctitheriids.
Among European and North American Nyctitheriidae,
the results show some support of generic monophyly,
including a monophyletic Plagioctenodon, consisting of
P. dormaalensis, P. rosei, P. thewisseni, P. krausae, and
P. savagei. This supports the proposal by Beard and
Dawson (2009) to include P. dormaalensis and P. rosei
in Plagioctenodon, rather than Leptacodon. Other monophyletic genera found were Nyctitherium, Plagioctenoides,
Euronyctia, and Cryptotopos, if Saturninia carbonum is

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J Mammal Evol (2015) 22:307342

Fig. 7 Strict consensus of the phylogenetic analysis of Nyctitheriidae

with stratigraphic and biogeographic information. The dark gray and light
gray boxes denote Asian and European taxa, respectively, and nonshaded taxa are North American. Stratigraphic ranges are either in black
or are striped. The European ranges could only be constrained to MP
level, except in the case of the two taxa from Dormaal, which are thought
to be earliest Eocene and coincide with the PETM (e.g., Smith and Smith

1996). The diagonally striped stratigraphic ranges represent a locality or

formation with a highly uncertain age. The vertically striped stratigraphic
ranges represent uncertain identifications of taxa outside of their known
ranges. The medium gray vertical lines represent, from left to right, the
Cretaceous-Paleogene boundary, the Paleocene-Eocene Thermal
Maximum, and the Grande Coupure or Eocene-Oligocene Boundary
(Woodburne 2004)

subsumed into the latter and classified as Cryptotopos

carbonum. The Wyonycteris clade is represented as
paraphyletic with respect to Pontifactor bestiola. The
two most diverse genera, Leptacodon (eight species)
and Saturninia (nine species), are both likely polyphyletic as currently classified, with most of their species

widely separated from the type species L. tener and

S. gracilis.
Although there does seem to be a geographic signal
in the data (Fig. 7), separating the Asian nyctitheriids at
the base of the tree, the European nyctitheriids in a deeply
nested clade, and the North American nyctitheriids between

J Mammal Evol (2015) 22:307342

the two groups, there appears to be some intermixing between

the European and North American taxa. Several early and
middle European taxa are found in predominantly
North American clades, including Saturninia ceciliensis,
Leptacodon nascimentoi, Placentidens lotus,
Plagioctenodon dormaalensis, and Wyonycteris richardi.
A single North American nyctitheriid, Limaconyssus
habrus, is found in the clade including all late Eocene
European taxa. These results suggest that there was
considerable interchange between the nyctitheriid taxa
of Europe and North America.

Discussion
Outgroup
Our results suggest that Wyonycteris microtis is best
classified outside of Nyctitheriidae. Wyonycteris
microtis is the most basal taxon of the ingroup. Sister
to W. microtis is a group consisting of two taxa traditionally classified as erinaceomorphs and Nyctitheriidae
(Node A in Fig. 6), which is distinguished by the presence of a hypocone and the loss of a stylocone on the
upper molars (See Appendix 2 for a list of all unambiguous synapomorphies at key nodes). The basal position
of W. microtis might be driven in part by the absence
of hypocones and pericones on the upper molars.
Although these features are apparently absent on the type
specimen, it is at least possible that a hypocone and
pericone could have been present on less damaged or
worn specimens (Secord 2008; Beard and Dawson 2009).
Regardless, the relatively long upper molar shape, presence of small pre- and postcingula, and the presence of a
stylocone on M1 of W. microtis support its basal position in the cladogram. Given the current state of knowledge for this taxon, known only from a single specimen
preserving M13, claims regarding the interfamilial relationships of this taxon should be viewed as tenuous at
best.
Also in the outgroup relative to Nyctitheriidae, Adunator
minutus and Macrocranion junnei were recovered as sister
taxa (Node B) representing Erinaceomorpha (e.g.,
Novacek et al. 1985; although see Hooker and Russell
2012). The two taxa share several characteristics of that
clade including relatively wider talonids than trigonids
on the lower molars and relatively reduced parastylar
lobes on the upper molars.
Asian Nyctitheriidae
Results suggest that the five taxa from Asia included in
the analysis are consecutively nested at the base of the

327

Nyctitheriidae, with the three basal-most taxa, Edzenius

lus, Oedolius perexiguus, and Asionyctia guoi, consistent
with a paraphyletic subfamily Asionyctiinae (between
Nodes C and D). Asionyctiinae was first proposed by
Missiaen and Smith (2005) to include the genera
A s i o n y c t i a , B u m b a n i u s , O e d ol i u s, Volt ai a, an d
Bayanulanius, but was later modified by Lopatin (2006)
to not include Bumbanius. Although this subfamily is said
to be distinguished in part by a premolariform P4 with a
small to absent metaconid and highly transverse upper
molars (Missiaen and Smith 2005; Lopatin 2006), these
are also characters seen in primitive eutherians, including
the outgroup taxon Maelestes gobiensis. Synapomorphies
uniting Bumbanius rarus with the rest of Nyctitheriidae to
the exclusion of the basal Asionyctiinae (Node D) include
the presence of a metaconid on the P4 and a reduction in
the upper molar length to width ratio. Bumbanius has
been classified in the subfamily Praolestinae with
Praolestes (not included in this study) and Eosoricodon
terrigena is classified in the monospecific subfamily
Eosoricodontinae (Lopatin 2006). Our results suggest that
those subfamilies might be nested consecutively within
Asionyctiinae and outside the clade consisting of the
rest of the North American and European nyctitheriids,
but this result could be further tested with inclusion of
other Asian taxa in the analysis, which is beyond the
scope of the current study.
Asian nyctitheriids have previously been classified as a
single, monophyletic subfamily likely derived from a primitive Leptacodon-like morphology (Missiaen and Smith
2005: 520). One of the most significant changes from
the presumed primitive condition to the Asionyctiinae
was the reduction of the P4 from semimolariform to
premolariform, with the losses of the talonid basin,
two talonid cusps, and the metaconid. A premolariform
P4 is also present in many early eutherians, but it was
argued that in the Asionyctiinae, it was secondarily
derived (Missiaen and Smith 2005). With the recognition of greater diversity in the Asian nyctitheriids and
the addition of two more Asian subfamilies, Lopatin
(2006) challenged the monophyly of Asian nyctitheriids,
but still considered the subfamily including Leptacodon,
the Nyctitheriinae, to be closest to the primitive condition in the family. This position is supported by stratigraphic evidence. The oldest purported nyctitheriid,
known from a single P4 (Van Valen 1978), is from the
Puercan NALMA from North America. By the
Torrejonian NALMA, the first definitive nyctitheriids
L. tener and L. munusculum appear, and by the early
Tiffanian NALMA, a modest diversity is present with
L. packi, L. acherontus, and L. choristus making their
first appearances (Gunnell et al. 2008). In contrast,
Asian nyctitheriids do not appear until the Gashatan

328

ALMA, the beginning of which is concurrent with the

Tiffanian NALMA (Ti5a; Missiaen 2011). Thus,
nyctitheriids appear in North America 37 million years
before they do in Asia. The conflicting stratigraphic and
cladistic evidence suggests two possibilities: 1) the cladistic results are reflecting a hidden diversity of early
Paleocene Asian nyctitheriids not yet recovered from the
fossil record, or 2) Asian nyctitheriids originated from a
Leptacodon-like morphology, later migrating from North
America to Asia, and their primitive-looking characters
are actually secondarily derived. Paleontological investigations of the early Paleocene in Asia, together with a
cladistic analysis with a more exhaustive taxonomic
treatment of Asian nyctitheriids have the potential to
provide substantial insight into this issue.
North American and European Nyctitheriidae
Several characteristics differentiate North American and
European nyctitheriids from those recovered from Asia
(Node E). These include semi-molarization of P4, possibly derived from the more premolariform state found
in Asionyctiinae, specifically in having a basined talonid
and more than one talonid cusp. The lower molar
talonids are also notably different: the talonid basin is
wider so that it is subequal to the trigonid width, the
hypoconulid is shifted to occupy a position on the
midline, and the entoconid is subequal in height to the
hypoconid in contrast to its smaller and lower position
in most Asian taxa.
Monophyly of the North American and European
subfamilies, Nyctitheriinae and Amphidozotheriinae, is
not supported by this study. According to McKenna
and B ell (1997) and modified by Hooker and
Weidmann (2000), the Amphidozotheriinae includes
Amphidozotherium, Paradoxonycteris, Euronyctia,
Plagioctenoides, and the Oligocene Darbonetus (not included in this cladistic analysis). The former three taxa
are nested deeply within a late Eocene European clade
(Node S) with proposed nyctitheriines Saturninia rigasii
and Scraeva hatherwoodensis and the unclassified
Limaconyssus habrus. The group represented by Node
S is sister to Node O, which includes two North
American clades. The first represents the sole North
American genus classified as an amphidozotheriine,
Plagioctenoides (Node P), and the second is a clade that
includes Wyonycteris and Pontifactor (Node Q).
These results of this analysis strongly suggest that
Leptacodon and Saturninia, as they are currently understood, are not monophyletic. Although these genera have
been modified over the years, it is apparent that several
species still classified with them, likely due to a lack of
distinguishing derived features, should be reconsidered.

J Mammal Evol (2015) 22:307342

Furthermore, the type species of Leptacodon, L. tener, is

far removed from the other Leptacodon species, with
some uncertainty as to the specific relationships of that
taxon. In the strict consensus it is part of a polytomy
(Node G) with two large clades (Nodes H and K) and in
the four MPTs, it falls out as either the sister taxon to a
clade including both Nodes H and K or as sister only to
Node K, the clade comprised of L. acherontus and all
taxa nested more deeply. Beard and Dawson (2009)
suggested that Leptacodon is a relatively primitive
Paleocene clade that includes L. tener, L. packi, and
L. munusculum. Several morphological steps separate
these taxa, but a particularly significant character separates L. tener from L. packi, L. munusculum, and other
basal nyctitheriids: the presence of a metacone on P4. In
taxa with the relevant tooth available for study, this cusp
is absent in all basal nyctitheriids and present in all
nyctitheriids at Node G and more highly nested. Therefore,
we propose that at least five of the Leptacodon species merit
reattribution to a new or different genus: L. donkroni,
L. munusculum, L. nascimentoi, L. choristus, and L. packi.
Most likely L. proserpinae, upon discovery of additional
fossils, might also merit reattribution. Unfortunately this
study faces the same problems that authors in the past
had in that there are few, if any, good derived characters
grouping these taxa. For that reason, we do not rename
any of these taxa or create new clades. Some authors
have suggested synonymizing some of the Leptacodon
species with other nyctitheriid genera. Krishtalka (1976)
suggested that L. packi may be synonymized with
Nyctitherium, but Bown and Schankler (1982) instead
hypothesized that it shows relatively primitive, generalized morphology for early soricomorphs. Our phylogenetic analysis supports the latter theory. Krishtalka
(1976) further suggested that L. munusculum may best
be classified within Pontifactor, but our results do not
support this hypothesis, either.
Results from the cladistic analysis also strongly suggest that species currently classified in Saturninia are
polyphyletic with respect to each other. Among these,
S. ceciliensis is the most problematic, falling out with
the more primitive Leptacodon species and far removed
from the other Saturninia species. Although it was
mentioned that S. ceciliensis has some significant differences from other Saturninia species, such as lower
molars with a labial termination of the cristid obliqua
and a shorter entoconid than hypoconid (Storch and
Haubold 1989), the extremely basal position it occupies
in the cladogram seems incongruous with its temporal
distribution. Saturninia ceciliensis is the stratigraphically
oldest Saturninia species, but its middle Eocene age is
still much younger than the Paleocene L. munusculum
and earliest Eocene L. donkroni that it shares a clade

J Mammal Evol (2015) 22:307342

with. It is only known from one specimen consisting of

lower teeth, however, and therefore many of the
distinguishing characters for the later Eocene European
nyctitheriids, particularly the expanded postcingular
shelves of the P4 and upper molars, cannot be assessed.
The other Saturninia taxa are paraphyletic with the type
species S. gracilis in a clade of mostly late Eocene
European taxa, but other than possibly S. mamertensis,
they should probably be reassigned to a different genus.
Similar to the situation with Leptacodon, many taxa
were placed in Saturninia based on a very broad diagnosis of the group (Sig 1976). Some taxa were subsequently removed from the genus and placed in morphologically distinct genera (Sig 1997; Hooker and
Weidmann 2000), but that has resulted in Saturninia
being defined by an amalgamation of characters present
in the remaining species, rather than by unique, derived
traits.
The strict consensus shows a clade including Leptacodon
catulus, Placentidens lotus, Ceutholestes dolosus, Acrodentis
rosenorum, and all of the Nyctitherium species (Node H),
united by having a relatively wide P4, a longer M1 than M2,
lower molars with a cristid obliqua terminating lingually
on the protocristid, and a postcingulid on the lower
molars. Within that clade, European P. lotus falls out
as sister taxon to North American C. dolosus (Node I),
a result foreshadowed by Rose and Gingerich (1987)
who noted the similarities between the two taxa. At
the time, Placentidens was thought to be a plagiomenid
and Ceutholestes was described as an insectivore with
uncertain affinities, but it was noted that the two shared
lingually canted, exodaenodont molars with acute cusps
(Rose and Gingerich 1987). It has also been observed
that the morphology of P. lotus is very similar to that of
Wyonycteris (Gingerich 1987; Hand et al. 1994; Beard
and Dawson 2009). Smith (1995) suggested that
Wyonycteris is closely related to Pontifactor and
Remiculus, but argued against a close relationship with
Placentidens. Beard and Dawson (2009) proposed the
subfamily Placentinae, a clade consisting of
Placentidens, Remiculus, Wyonycteris, and Pontifactor.
The inclusion of Placentidens in that clade was based
on the more basal morphology of Wyonycteris primitivus
acting as an intermediate between Placentidens and the
more derived Wyonycteris species. Our analysis does not
support a close relationship of Placentidens with
Wyonycteris or Pontifactor, although a close relationship
between Wyonycteris and Pontifactor is supported (Node
Q). Remiculus was not included in this clade because it
is likely an adapisoriculid (Gheerbrant and Russell
1989; de Bast et al. 2012).
The synapomorphies uniting Ceutholestes dolosus and
Placentidens lotus (Node I) are the presence of a

329

metaconid on the P3 and lower molars with a greater

talonid width compared to that of the trigonid, a larger
metaconid than protoconid, and significantly taller
entoconids than hypoconids. The synapomorphies
concerning the lower molars are present in various
nyctitheriid taxa throughout the cladogram, but the P3
metaconid is unique to Ceutholestes and Placentidens.
There is a possibility that the isolated tooth identified as
the P3 of P. lotus is actually the P4; Russell et al.
(1973) hypothesized that Placentidens was a primitive
plagiomenid and their allocation of the tooth to the P3
position was based on comparisons to plagiomenid premolars that exhibit a much more highly molariform P4.
If Placentidens is a nyctitheriid, the P3 identification is
much less certain because some nyctitheriids have a
premolariform P4. Although we dont feel we can confidently ascribe the tooth in question to either the P3 or
P4 position, we did test whether the phylogenetic placement of P. lotus changes dependent on the tooth position. When the tooth was coded as a P4 rather than a
P3, the topology of the strict consensus tree did not
change, but the MPTs did increase in length by three
steps. One of the most distinguishing features of
Ceutholestes is that it has a fully molariform P4, whereas other insectivorans and nyctitheriids do not (Rose
and Gingerich 1987). If the P3 of Placentidens is
truly a P3, the more molariform characters it share with
Ceutholestes, such as the acquisition of a metaconid,
suggests that the unknown Placentidens P4 may also
be fully molariform and provide further synapomorphies
for the two taxa.
Sister to Placentidens lotus + Ceutholestes dolosus is
a monophyletic clade comprised of Acrodentis
rosenorum and all Nyctitherium species. Acrodentis is
in a sister relationship with Nyctitherium and
Nyctitherium krishtalkai is found to be the most primitive Nyctitherium species. This echoes the result found
in a recent analysis that tested the relationship of
Nyctitherium to other North American nyctitheriids
(Christiansen and Stucky 2013). We were unable to
determine the phylogenetic relationship between
Nyctitherium christopheri, Nyctitherium serotinum, and
Nyctitherium velox because a direct comparison between
N. christopheri and N. serotinum was not possible for
this analysis; there are no lower teeth known for
N. christopheri and the upper teeth of N. serotinum
have not been described or figured in the literature.
Plagioctenodon (sensu Beard and Dawson 2009, plus
P. thewisseni) is found to be a monophyletic clade.
Beard and Dawsons (2009) proposed synapomorphies
for the genus, however, (an anteriorly canted, relatively
small P3 and a more molariform P4 trigonid with a
relatively high paraconid) appear to have accumulated

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gradually in the evolutionary history of Nyctitheriidae.

Although the relatively high P4 paraconid appears as a
synapomorphy for the genus, the distinguishing P3 characters evolved earlier in the tree. The anterior cant of
the P3 protoconid first appears at Node K and is seen in
Leptacodon acherontus and many of the more deeply
nested taxa in that clade. The reduction in size of the P3
relative to the surrounding molars is a synapomorphy
for Node L and is present in all of the taxa that make
up that clade with the relevant morphology known,
including species in Plagioctenodon, Plagioctenoides,
Wyonycteris, and the late Eocene European nyctitheriids.
Although this P3 morphology is present in Plagioctenodon, it
should be recognized that those characters are gained at
an earlier stage in nyctitheriid evolution and that definitive diagnoses for the genus should be based on the
combination of the P 3 characteristics with the P 4
paraconid height. Within Plagioctenodon, the North
American taxa are separated from the basalmost
European species, P. dormaalensis, by having taller
entoconids than hypoconids on the lower molars. The
most nested species, P. krausae and P. savagei, are also
the youngest species and are differentiated from the older
species by having the entoconid positioned on the
posterolingual margin of the tooth, rather than being
anterolingual to the hypoconulid, a character formerly
included in the generic diagnosis (Bown and Schankler
1982). The position of the sole European Plagioctenodon
species at the base of the clade suggests that the genus
may have originated in Europe and then later migrated to
North America. The fact that P. dormaalensis is from the
earliest Eocene and therefore younger than the more
nested late Paleocene taxa, P. rosei and P. thewisseni,
suggests that older Plagioctenodon might have been
present outside of North America. Alternatively, but not
supported by our results, P. dormaalensis might have
evolved from one of the earlier species in North
America and, after migrating, diverged sufficiently in
morphology from the rest of the genus (re-evolving
primitive morphology) to appear more basal in this cladogram. A North American origin is further supported by
the greater diversity of Plagioctenodon in North
America, suggesting the family arose on that continent
and a single member later migrated to Europe.
Node N includes all species classified in Plagioctenoides,
Wyonycteris, and Pontifactor, as well as the late Eocene
European nyctitheriids, in which a single Paleocene
North American nyctitheriid, Limaconyssus habrus, is
deeply embedded. Synapomorphies supporting Node N
are a crestiform paraconid on the lower molar, a cristid
obliqua ascending the protocristid to end high on the
metaconid on the lower molar, and a precingulum containing a pericone on the upper molar. The crestiform

J Mammal Evol (2015) 22:307342

paraconid is a character that is present in the basalmost

nyctitheriids, including all of the Asian taxa and the
basalmost clade of European and North American taxa
that includes Saturninia ceciliensis, Leptacodon
munusculum, and Leptacodon donkroni. In most other
nyctitheriids the paraconid is cuspate, but at Node N
the crestiform state reoccurs and is present in
Plagioctenoides, Wyonycteris primitivus, some species
of Saturninia, Paradoxonycteris, and Euronyctia. This
character seems to have evolved and reversed at least
twice in the familys history. The termination of the
cristid obliqua high on the metaconid of the lower molar
and the pericone on the upper molar have also occurred
several times in the familys history. In fact, there are
many hypotheses of reversals throughout the cladogram,
likely playing a strong role in the difficulty in determining intrafamilial nyctitheriid relationships over the last
century.
A monophyletic group that includes Wyonycteris,
Pontifactor, and Plagioctenoides is present (Node O),
in which Wyonycteris is paraphyletic in a clade with
Pontifactor. A monophyletic Plagioctenoides is sister to
the paraphyletic Wyonycteris. Node O has five unambiguous synapomorphies, one of which, nyctalodonty, is
only present in this clade. A nyctalodont condition refers
to the relationship between the lower molar talonid
cusps, in which the hypoconid and hypoconulid are
connected by a crest, while there is a fissure between
the hypoconulid and entoconid (Menu and Sig 1971).
This clade also has two significant characteristics that are
rare within the rest of Nyctitheriidae: a mesostyle and
dilambdodonty in the upper molars. These two character
states do not show up as unambiguous synapomorphies
at Node O because Plagioctenoides does not have any
upper teeth attributed at the species level and therefore
the timing at which these characters evolved is uncertain.
Although upper molars have been described attributed to
the genus (Rose et al. 2012) that have both a mesostyle
and the dilambdodont condition, they were not included
in the phylogenetic analysis because of specific uncertainty. That said, if they belong to either Plagioctenoides
species, they could support the acquisition of the two
character states at Node O. In nyctitheriids, a mesostyle
is only present in Wyonycteris species excluding
Wyonycteris primitivus, Pontifactor, Plagioctenoides sp.,
Nyctitherium velox, and the questionably attributed
Wyonycteris microtis and dilambdodonty is only found
in Wyonycteris species excluding W. primitivus,
Pontifactor, Plagioctenoides sp., Paradoxonycteris, and
Euronyctia.
Close relationships of Plagioctenoides, Wyonycteris, and
Pontifactor were predicted by previous authors (Gingerich
1987; Smith 1995; Rose et al. 2012). In the initial

J Mammal Evol (2015) 22:307342

description of the first Wyonycteris species, Gingerich

(1987) noted similarities between it and Pontifactor, but
concluded that the differences between the two taxa were
great enough to merit generic level distinction. Our results support these conclusions by finding Pontifactor
bestiola within a paraphyletic Wyonycteris, but
P. bestiola has four autapomorphies in the upper molars
differentiating it from Wyonycteris: lack of pericone,
more labially situated conules, subequal heights of the
paracone and metacone, and an expanded postcingulum.
Concerning Wyonycteris and Plagioctenoides, Rose et al.
(2012) noted that the similarities were so great between
the two genera that they may not be generically distinct.
Our study does show a separation of the two, but it is
only supported by a single synapomorphy for each genus. The synapomorphy for Plagioctenoides is a single
prominent cusp on the P4 talonid (Rose et al. 2012),
whereas the Wyonycteris species with a known P4 have
three minute cusps (e.g., Gingerich 1987) and the synapomorphy for Wyonycteris is a more anteroposteriorly
compressed trigonid on the lower molar than is seen in
Plagioctenoides. Although it seems questionable to separate genera based on such small morphological differences, we hesitate to combine the two when our analysis
does show a clear separation of two monophyletic
groups.
Within Wyonycteris (Node Q), W. primitivus is found
to be at the base, in accordance with Beard and Dawson
(2009). Many of the characteristics used to differentiate
Wyonycteris in its original description, such as a twinned
metaconid on the lower molar, a mesostyle, and
dilambdodonty (Gingerich 1987) are actually found at
node R, nested within W. primitivus. Beard and
Dawson (2009) based their generic attribution of
W. primitivus on other morphological similarities, particularly the nyctalodont talonids, and hypothesized that the
lack of mesostyle and dilambdodonty in W. primitivus
was due to its status as a basal species of the genus,
which this study corroborates.
The clade of late Eocene European nyctitheriids (Node
S) is marked by an expansion of the postcingulum on the
P4 and upper molars into a large shelf; this is seen
throughout the entire clade in the taxa with known upper
teeth. In the nyctitheriids outside of Node S, expanded
postcingula on the upper molars only occur in the terminal genera Nyctitherium and Pontifactor. This suggests
that expanded postcingula evolved three times within
Nyctitheriidae and that it is a fairly stable character once
acquired, as no lineages show the secondary loss of the
character. There are two possible nodes representing a
monophyletic Cryptotopos: Node T or Node U. Since
Cryptotopos was first described (Crochet 1974), its status
as a genus and the species that are classified within it

331

have been in flux. Sig (1976) synonymized Cryptotopos

with Saturninia and renamed the type species of
Cryptotopos beatus to Saturninia beata. Hooker and
Weidmann (2000) resurrected the genus and reallocated
several species to it: C. beatus, C. woodi, C. hartenbergi,
and potentially Saturninia pirenaica and Saturninia
pelissiei. Ziegler (2007) added a new Oligocene species,
Cryptotopos communis, to the genus and also suggested
that Saturninia carbonum may best be classified within
Cryptotopos, as well. This analysis supports the allocation of S. carbonum to Cryptotopos because that species
is deeply nested within the other Cryptotopos species.
Node U includes C. beatus, C. woodi, C. hartenbergi,
and the newly allocated Cryptotopos carbonum. It is
supported by three synapomorphies: an anteroposteriorly
compressed lower molar trigonid, a cristid obliqua on the
lower molars whose distal portion is convex occlusally,
and an absent to weak centrocrista on the upper molars.
Of these characters, the first two traits have been used in
the diagnosis of Cryptotopos (Hooker and Weidmann
2000; Ziegler 2007). In a broader sense of the genus,
Node T could represent Cryptotopos; this expanded clade
would include the previously suggested Cryptotopos species S. pelissiei, and also Saturninia intermedia. The
three synapomorphies for Node T are a relatively wide
P4, a cuspate paraconid on the lower molars, and a notch
located halfway down the cristid obliqua on the lower
molars. Of these, only the last character has been used to
diagnose the genus (Hooker and Weidmann 2000;
Ziegler 2007). This indicates that the diagnostic characters for Cryptotopos are accumulated gradually in the
clade and that the species classified in the genus differ
depending on which characters are used for diagnosis.
Saturninia pirenaica, the only other species suggested to
be classified in Cryptotopos, is part of a polytomy at
Node S. Although its classification as a species of
Cryptotopos is not supported, it should be noted that
few characters could be coded for S. pirenaica because
it is not well known and only a few figures exist in the
literature (Gibert Clols and Agust Ballester 1979; Sig
1997).
A monophyletic Euronyctia as sister to Paradoxonycteris
is supported at Node X. This is in agreement with the
view in Hooker and Weidmann (2000) that the two genera
are almost indistinguishable. The clade including both
genera shows another acquisition of dilambdodonty in
the upper molars, but unlike the Wyonycteris taxa associated with Node R, Euronyctia and Paradoxonycteris do
not have a mesoconid or nyctalodont lower molar
talonids. The only difference between Euronyctia and
Paradoxonycteris in this analysis is that Euronyctia has
a more lingually situated hypoconulid. Smith (2004) argued that the holotype of Paradoxonycteris soricodon is

332

sufficiently damaged for that taxon to be considered a

nomen nudum or dubium and that only Euronyctia is a
valid genus. The holotype of Paradoxonycteris (as illustrated by Hooker and Weidmann [2000]) seemed complete
enough to be coded into the analysis and it was therefore
retained as a distinct species, but it may be advisable to
synonymize Paradoxonycteris and Euronyctia. A revision
of these genera, however, is beyond the scope of this
study.
A clade including Amphidozotherium cayluxi, Saturninia
rigasii, Scraeva hatherwoodensis, and Limaconyssus habrus
(Node Y) is supported by having lower molars with
narrower talonids than trigonids, a cuspate paraconid,
and a larger paraconid than metaconid. Our results suggest that the relatively narrow talonids of this clade
might represent a character reversal, as this is the state
also seen in the Asian nyctitheriids. Within this clade,
Scraeva and Limaconyssus are sister taxa (Node Z) with
the following synapomorphies: a large, cuspate
paraconid on the P4 and lower molars with a trigonid
at least twice as high as the talonid. The former character state is also seen in the basalmost nyctitheriids,
whereas the latter is the state seen in the outgroup taxon
Maelestes, suggesting two more examples of likely
character reversals. The placement of L. habrus in this
tree is suspect, however, because it is the only North
American taxa within Node S, it is only known from
lower teeth, which means the presence of important
synapomorphies for Node S such as the expansion of
the postcingular shelf on the P4 and upper molars cannot be assessed, and its late Paleocene age is more than
ten million years older than the late Eocene age of the
other taxa in the clade. Thus, it is possible, although not
supported by our cladistic results, that Limaconyssus
and the late Eocene nyctitheriids may have independently regained some primitive characteristics that, combined with an assortment of derived traits, caused the
strange placement of Limaconyssus as a highly nested
taxon in a late Eocene European clade.
Biogeography of Nyctitheriidae
Nyctitheriidae is roughly grouped into three major
clades that correspond to three separate continents
(Fig. 7). The five Asian species in the analysis are
consecutively nested at the base of the tree in a
paraphyletic relationship. Although there are several
more Asian nyctitheriids not included in our study (see
Lopatin 2006), species from all three Asian subfamilies
were included in an attempt to capture much of the
morphological disparity on the continent. Because of
this, the position of the Asian nyctitheriids outside of
the European and North American clades, is likely to be

J Mammal Evol (2015) 22:307342

a true pattern, even if the specific relationships of the

Asian taxa are less certain. Results from the analysis
support an Asian origin for the family with dispersal
into North America by at least the Torrejonian and
possibly the Puercan. But as was noted in the discussion of the phylogenetic results, the temporal data conflict with the cladistic results, in that the oldest
nyctitheriids are North American. This conflicting evidence can be reconciled in the future either by the
discovery of early Paleocene Asian nyctitheriids or
through a more exhaustive test of character polarity in
the Asian taxa demonstrating that the primitive traits
in those taxa are actually character reversals following
dispersal and isolation on the Asian continent.
Regardless, the monophyly of the European and North
American nyctitheriids, to the exclusion of the Asian
taxa, would seem to indicate that there was no
dispersal of nyctitheriids to or from Asia after the
early Paleocene. Russell and Dashzeveg (1986) noted
the similarities of early Eocene Asian nyctitheriids and
other insectivores to late Paleocene forms in North
America and suggested that the Asian taxa were not
strongly endemic, but Missiaen and Smith (2005) proposed a single migration event between North America
and Asia in the late Paleocene, after which the Asian
nyctitheriids evolved in isolation. The results of this
analysis better support the idea of a single migration
event, whether it is from Asia to North America in the
early Paleocene or from North America to Asia in Ti5a, but more Asian nyctitheriids should be included in
the phylogenetic analysis before this can be said
definitively.
All of the North American and European taxa (only
excluding those that were latest Eocene and strictly
Oligocene) were included in this analysis, which gives
us a more confidant view of the patterns seen between
those two continents. The earliest nyctitheriids are found
in the Paleocene of North America in the form of
several species classified as Leptacodon (found to be
polyphyletic here). By the late Paleocene the family is
relatively diverse, with the presence of Ceutholestes,
Limaconyssus, and representatives of Plagioctenodon
and Wyonycteris. There appears to have been a large
dispersal event between North America and Europe
around the Paleocene-Eocene Boundary, when the
stratigraphically earliest European nyctitheriids appear
in four morphologically distinct clades. These species,
Leptacodon nascimentoi, Placentidens lotus,
Plagioctenodon dormaalensis, and Wyonycteris richardi
all appear in the earliest Eocene, which suggests immigration in multiple clades at this time. There is also
evidence for dispersal between North America and
Europe at the Paleocene-Eocene boundary in other

J Mammal Evol (2015) 22:307342

mammalian genera, including a proposed North America

to Europe route for Palaeonictis (Chester et al. 2010) and
Europe to North America routes for both Teilhardina (Rose
et al. 2011) and Macrocranion (Smith et al. 2002). This
coincides with the Holarctic dispersal events in many
other clades near the Paleocene-Eocene boundary, including hyaenodontid creodonts (Gingerich and
Deutsch 1989), as well as the modern orders of primates, artiodactyls, and perissodactyls (Gingerich 2006;
Smith et al. 2006). Although the dispersal of these
groups included an Asian componentand possible
origination on that continent (Krause and Maas 1990;
Beard and Dawson 1999; Bowen et al. 2002)we do
not see evidence for a second instance of migration
either to or from Asia in the Nyctitheriidae.
The North American nyctitheriids as a group are
paraphyletic with respect to the Eocene European taxa
that are spread throughout it. With the exception of
some Nyctitherium species and Pontifactor bestiola, the
stratigraphic ranges of North American Nyctitheriidae
do not extend past the early Eocene. A diverse clade
of late Eocene nyctitheriids can be found in Europe, but
their specific origins are unclear. The cladistic results
suggest that this group evolved from a common ancestor it shares with its sister clade (comprised of
Plagioctenoides, Wyonycteris, and Pontifactor), diversified in Europe, and experienced a single migration event
to North America by the deeply nested Limaconyssus
habrus. A late Paleocene timing of all of these events is
required with the inclusion of L. habrus in the clade,
but as was discussed in the phylogenetic results section,
several phylogenetically informative characters for late
Eocene European nyctitheriids could not be coded for
this taxon. If L. habrus is not included in the late
Eocene European clade, the ghost lineage for that
clade would be nearly 20 million years, with
diversification not occurring until the late Eocene. It is
therefore possible that the late Eocene nyctitheriids may
have arisen from one of the earlier Eocene European
taxa or from a middle Eocene North American taxon
after a later immigration event from North America, but
with great enough divergence that it is not being
captured in the cladistic analysis. Potentially the
discovery of more Saturninia ceciliensis fossils,
particularly the upper molars, may transfer this taxon
to the late Eocene group and would represent a
transitional middle Eocene European taxon. It also
seems significant that one of the uniting features of all
late Eocene European nyctitheriids, the expanded
postcingulum on the upper molars, is also present in
the middle Eocene P. bestiola, a member of the sister
clade to the late Eocene European clade. Smith (1996)
suggests that some nyctitheriid fossils from the early

333

Eocene of Europe are likely to be Pontifactor or

Wyonycteris; he notes two P 4 s, attributed to cf.
Leptacodon (RI 218 from Rians, Sparnacian, Provence,
France) by Godinot (1981) and to Saturninia sp. or
Leptacodon sp. (FDN 1401 from Fordones, middle
Ilerdien, Bas-Languedoc, France) by Marandat (1991),
as being very similar in morphology to Pontifactor and
a lower molar (RI 382 from Rians) attributed to cf.
Leptacodon by Godinot (1981) with a nyctalodont condition that would ally it with Wyonycteris. Although
these teeth have not been formally diagnosed, it is
possible that they are part of the ancestral stock of the
late Eocene European clade because of their early
Eocene presence in Europe and their similar morphology to the sister clade of late Eocene nyctitheriids. It is
apparent that several clades of nyctitheriids dispersed
between North America and Europe in the earliest
Eocene and that there may have been at least one
further immigration event into Europe later in the
Eocene from a Pontifactor-like nyctitheriid.

Conclusions
The Nyctitheriidae is a diverse and disparate fossil
family that spanned the early Paleocene through early
Oligocene and in the early Eocene, could be found on
three different continents. The stratigraphically earliest
nyctitheriids are found in North America, and the family
later appeared in Asia in the late Paleocene and in
Europe in the earliest Eocene. By the late Eocene, the
family was solely found in Europe and then became
extinct in the early Oligocene. Nyctitheriidae contains
over twenty genera, but two of the most diverse,
Saturninia and Leptacodon, are almost certainly polyphyletic and need to be revised. The monophyly of
other multi-species genera are supported by our cladistic
analysis, including Nyctitherium, Plagioctenodon,
Plagioctenoides, Cryptotopos, and Euronyctia, whereas
Wyonycteris is found to be paraphyletic with Pontifactor.
The new species of Plagioctenodon, P. thewisseni, is found
to be in a clade with P. dormaalensis, P. rosei, P. krausae, and
P. savagei, but the previously suggested synapomorphies
for Plagioctenodon in the anteriorly canted P 3
protoconid and reduction in size of the P3 are gradually
accumulated in the nodes nested below the genus. Only
the high position of the paraconid on the P4 is found to
be a synapomorphy at the Plagioctenodon node. The
monophyly of the subfamilies Nyctitheriinae and
Amphidozotheriinae is not supported and these subdivisi on s m ay ne e d t o b e ab an do ne d . Th e A sia n
nyctitheriids are found to be consecutively nested at

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the base of Nyctitheriidae due to many morphological

similarities with the outgroup, but it is unlikely that the
family actually originated in Asia because nyctitheriid
species are found much earlier in North America.
Instead, the position of the Asian taxa in the cladogram
could be explained by a single immigration event from
North America to Asia in the late Paleocene, and then
isolation of the Asian nyctitheriids with convergence
upon the primitive condition in some characters.
Within the North American and European taxa, there
is a dispersal event at the earliest Eocene that includes
at least four clades and coincides with Holarctic dispersal events of several other mammalian clades, including hyaenodontid creodonts, Primates, Artiodactyla, and
Perissodactyla.
This is the most complete phylogenetic analysis of the
family to date and although the position of some of the
taxa and clades are likely to change upon addition of
more taxa and characters or discovery of more fossils for
already included taxa, many of the phylogenetic relationships suggested in the literature (e.g., Wyonycteris +
Pontifactor [Gingerich 1987]; Placentidens lotus +
Ceutholestes dolosus [Rose and Gingerich 1987];
Euronyctia + Paradoxonycteris and reassessment of
Cryptotopos [Hooker and Weidmann 2000]; further reassessment of Cryptotopos [Ziegler 2007]; reassessment of
Plagioctenodon [Beard and Dawson 2009];
Plagioctenoides + Wyonycteris [Rose et al. 2012];
Acrodentis + Nyctitherium [Christiansen and Stucky
2013]) are corroborated by this analysis and it can be
used as a good approximation of the intrafamilial relationships as they are currently understood. The
intrafamilial relationships of Nyctitheriidae are complex,
with the evolution and reversal of many characters multiple times, but it is necessary to understand the phylogeny within the family before we can understand the
relationships of the family to higher level taxa with
confidence.
Acknowledgments We thank Richard Hulbert, Kristen MacKenzie, and Jason Bourque (UF) for help in cataloguing and preparation of the UF specimens. We thank Philip Gingerich (UM) and
Gregg Gunnell (now of Duke Lemur Center) for access to specimens and comparative casts and for help with loans at UM, as
well as William Sanders (UM) for help with preparation of
specimens. We thank Peter Houde (New Mexico State University)
for access to limestones containing invaluable fossils. We thank
Stephen Chester (now of Brooklyn College) and Josh Van Houten
(Department of Internal Medicine, Yale University) for micro-CT
imaging done at the Yale University Core Center for

J Mammal Evol (2015) 22:307342

Musculoskeletal Disorders microCT facility; and Doug Boyer,
Jimmy Thostenson, and Judit Marigo (Duke University) for access
to and assistance with the microCT facilities at the Shared Materials Instrumentation Facility at Duke University. We thank Jason
Bourque, Stephen Chester, Jerry Hooker, Paul Morse, Ken Rose,
and Aaron Wood for helpful discussions. We thank John Wible
and an anonymous reviewer for providing helpful comments that
improved the manuscript. Research was in part funded by the
Miss Lucy Dickinson Fellowship, the Geological Sciences Graduate Award, and a Geological Society of America Student Research Grant to C.L.M. The version of TNT used in the cladistic
analysis was produced with support by the Willi Hennig Society.
This is University of Florida Contribution to Paleobiology 679.

Appendix 1. Taxa Selected for Analysis and Sources

of Data
Outgroup
Maelestes gobiensis Wible et al., 2007Wible et al. (2007,
2009)
Erinaceomorpha
Adunator minutus (Jepsen, 1930)Jepsen (1930);
Gingerich (1983); Secord (2008); YPM-PU 19463 (cast)
Macrocranion junnei Smith et al., 2002Smith et al.
(2002); Rose et al. (2012); UM 93378 (cast)
Nyctitheriidae
Plagioctenodon krausae Bown, 1979Bown (1979);
Bown and Schankler (1982)
Plagioctenodon savagei Bown and Schankler, 1982
Bown and Schankler (1982)
Plagioctenodon dormaalensis (Quinet, 1964)Smith
(1996) as Leptacodon dormaalensis
Plagioctenodon rosei (Gingerich, 1987)Rose (1982) as
cf. Leptacodon packi; Rose and Gingerich (1987) as cf.
Leptacodon packi; Gingerich (1987) as Leptacodon rosei;
UF 303728; UF 303729; UF 303730; UM 71650 (cast); UM
76895 (cast); UM 77032 (cast)
Plagioctenodon thewisseni this publication UM 39873;
UM 76906; UM 76920; UM 82576; UM 83931; UM 86725;
UF 289746; UF 289747; UF 294696
Leptacodon tener Matthew and Granger, 1921
Simpson (1935); McKenna (1968); Krishtalka (1976);
Scott (2003)
Leptacodon munusculum Simpson, 1935Krishtalka
(1976); Gingerich et al. (1983); Krause and Gingerich
(1983); Scott (2003)
Leptacodon catulus Krishtalka, 1976Krishtalka (1976);
Gheerbrant and Hartenberger (1999)

J Mammal Evol (2015) 22:307342

Leptacodon packi Jepsen, 1930Jepsen (1930);

Krishtalka (1976); Secord (2008)
Leptacodon nascimentoi Estravis, 1996Estravis(1996)1
Leptacodon acherontus Secord, 2008Secord (2008);
YPM-PU 19957 (cast)
Leptacodon donkroni Rose et al., 2012Rose et al. (2012)
Leptacodon proserpinae2 Van Valen, 1978Van Valen
(1978)
Leptacodon choristus Secord, 2008Secord (2008)
Nyctitherium velox Marsh, 1872Robinson (1968); YPM
13510 (cast)
Nyctitherium serotinum (Marsh, 1872)Robinson (1968)
Nyctitherium krishtalkai Christiansen and Stucky, 2013
Christiansen and Stucky (2013)
Nyctitherium christopheri Krishtalka and Setoguchi,
1977Setoguchi (1973)as Nyctitherium robinsoni 3 ;
Krishtalka and Setoguchi (1977)
Acrodentis rosenorum Christiansen and Stucky, 2013
Christiansen and Stucky (2013)
Ceutholestes dolosus Rose and Gingerich, 1987Rose
and Gingerich (1987); UM 82503 (cast)
Limaconyssus habrus Gingerich, 1987Gingerich (1987);
UM 86724 (cast)
Pontifactor bestiola West, 1974West (1974); Bown
(1979)
Wyonycteris chalix Gingerich, 1987Gingerich (1987)
Wyonycteris richardi Smith, 1995Smith (1995)
Wyonycteris primitivus Beard and Dawson, 2009Beard
and Dawson (2009)
Wyonycteris galensis Secord, 2008Secord (2008); YPMPU 14138 (cast)
Wyonycteris microtis Secord, 2008Secord (2008); YPMPU 19479 (cast)
Plagioctenoides microlestes Bown, 1979Bown (1979);
Rose et al. (2012)

The codings for the M1 and M2 of Leptacodon nascimentoi were based

on Fig. 10 (SV3-300) and Fig. 9 (SV2-12), respectively, which is opposite
to how they are identified in Estravis (1996). See the reasoning in the
Body Mass Calculations subsection of the Materials and Methods section.
2
Leptacodon proserpinae was taken out of the final analysis. It is only
mentioned in one publication with a single tooth, the P4, illustrated. Only
a small fraction of the characters in this analysis could be coded (8 of 66),
which caused a great deal of instability in the placement of the taxon and
resulted in a large polytomy in the strict consensus tree.
3
The holotype specimen number in Setoguchi (1973) is given as TTM
1219 but based on the figures, the specimen (broken P4, M1-M3) is the same
as the holotype for Nyctitherium christopheri in Krishtalka and Setoguchi
(1977), although that specimen number is reported as CM 16996.

335

Plagioctenoides tombowni Rose et al., 2012Rose et al.

(2012)
Amphidozotherium cayluxi Filhol, 1877Sig (1976)
Saturninia gracilis Stehlin, 1940Crochet (1974); Sig
(1976)
Saturninia mamertensis Sig, 1976 Sig (1976)
Saturninia grandis Sig, 1976 Sig (1976)
Saturninia rigasii Hooker and Weidman, 2000Hooker
and Weidmann (2000)
Saturninia ceciliensis Storch and Haubold, 1989Storch
and Haubold (1989)
Saturninia intermedia Sig, 1976 Sig (1976)
Saturninia pirenaica Gibert Clols and Agust Ballester,
1979Gibert Clols and Agust Ballester (1979); Sig
(1997); Ziegler (2007)
Saturninia pelissiei Sig, 1997Sig (1997); Ziegler (2007)
Saturninia carbonum Sig and Storch, 2001Sig
(2001)
Cryptotopos beatus Crochet, 1974Crochet (1974); Sig
(1976) as Saturninia beata; Hooker and Weidmann (2000);
Ziegler (2007)
Cryptotopos woodi (Cray, 1973)Cray (1973); Ziegler
(2007)
Cryptotopos hartenbergi (Sig, 1976) Sig (1976);
Ziegler (2007)
Scraeva hatherwoodensis Cray, 1973Cray 1973
Placentidens lotus Russell et al.1973Russell et al. (1973)
Euronyctia montana Sig, 1997 Sig (1997)
Euronyctia grisollensis (Sig, 1976)Sig (1976); Sig
(1997)
Paradoxonycteris soricodon Revilliod, 1922Hooker and
Weidman (2000)
Asionyctia guoi Missiaen and Smith, 2005Missiaen and
Smith (2005)
Bumbanius rarus Russell and Dashzeveg, 1986Russell
and Dashzeveg (1986); Lopatin (2006)
Eosoricodon terrigena Lopatin, 2005Lopatin (2005)
Oedolius perexiguus Russell and Dashzeveg, 1986
Russell and Dashzeveg (1986); Lopatin (2006)
Edzenius lus Lopatin, 2006Lopatin (2006)

Appendix 2. Unambiguous Synapomorphies for Key

Nodes
Node A
52 (1) upper molar stylocone absent
61 (1) upper molar hypocone present

336

Node B: Adunator minutus + Macrocranion junnei

22 (2) upper ultimate premolar postcingulum absent
25 (2) M1 trigonid less wide than talonid
26 (2) M2 trigonid less wide than talonid
50 (1) M1 parastylar lobe anterolabially projecting
63 (1) M3 parastylar lobe small
Node C: Nyctitheriidae
14 (1) lower ultimate premolar paraconid lingual to
midline
34 (1) lower molar paraconid anteriorly projecting
37 (2) lower molar protoconid larger than metaconid
46 (2) lower molar entoconid smaller than hypoconid
66 (0) posteriormost mental foramen below penultimate
premolar or more anterior
Node D
15 (2) lower ultimate premolar metaconid present
48 (1) upper molar length between 0.7-0.95 of the width
Node E: North American and European nyctitheriids
17 (1) lower ultimate premolar talonid basin present
18 (1 or 2) lower ultimate premolar talonid has two or three
cusps
26 (1) M2 trigonid width subequal to talonid
44 (0) lower molar hypoconulid positioned on midline
46 (0) lower molar entoconid subequal in height to
hypoconid
58 (2) upper molar paraconule lingual to metaconule and
closer to protocone
Node F
14 (0) lower ultimate paraconid on midline
21 (0) upper ultimate premolar precingulum present
33 (0) lower molar paraconid cuspate in shape
37 (1) lower molar protoconid and metaconid subequal in size
Node G
24 (1) upper ultimate premolar metacone present
49 (0) M1 ectoflexus present
54 (1) upper molar centrocrista strong and rectilinear
55 (1) upper molar precingulum strong, reaching past
paraconule
Node H
11 (3) lower ultimate premolar width greater than 0.8 of the
length
14 (1) lower ultimate premolar paraconid lingual to midline
29 (1) M1 length greater than that of M2
40 (0) lower molar cristid obliqua meets the protocristid
lingual to notch
47 (1) lower molar postcingulid present
Node I: Placentidens lotus + Ceutholestes dolosus
6 (1) lower penultimate premolar metaconid present
37 (0) lower molar metaconid larger than protoconid
46 (1) lower molar entoconid taller than hypoconid
Node J: Nyctitherium

J Mammal Evol (2015) 22:307342

20 (1) anterodorsal shift in upper ultimate premolar visible

from labial view
23 (1) upper ultimate premolar hypocone present
56 (0) upper molar paraconule small, not strongly winged
60 (1) upper molar postcingulum greatly expanded into shelf
Node K
7 (1) lower penultimate premolar anteriorly canted
Node L
8 (1) lower penultimate premolar smaller than surrounding
premolars
Node M: Plagioctenodon
12 (2) lower ultimate premolar paraconid present and high
on anterior of trigonid
59 (1) upper molar paracone and metacone height subequal
Node N
33 (1) lower molar paraconid crestiform in shape
40 (2) lower molar cristid obliqua ascends the protocristid
to the tip of the metaconid
55 (2) upper molar precingulum strong with pericone
Node O
11 (1) lower ultimate premolar width between 0.5-0.65 of
the length
43 (1) cristid obliqua is deeply concave occlusally
44 (2) lower molar hypoconulid positioned lingually in
nyctalodont condition
46 (1) lower molar entoconid taller than hypoconid
66 (1) posteriormost mental foramen below ultimate premolar or more posterior
Node P: Plagioctenoides
18 (0) lower ultimate premolar has one talonid cusp
Node Q: Wyonycteris + Pontifactor
27 (0) lower molar trigonid anteroposteriorly compressed
Node R: W. galensis + W. chalix + W. richardi + P. bestiola
16 (0) lower ultimate premolar talonid narrower than
trigonid or absent
31 (1) lower molar metaconid twinned
33 (0) lower molar paraconid cuspate in shape
53 (1) upper molar mesostyle present
54 (2) upper molar centrocrista dilambdodont
Pontifactor bestiola autapomorphies
55 (1) upper molar precingulum strong, reaching past
paraconule
58 (1) upper molar paraconule lingual to metaconule and
halfway to protocone
59 (1) upper molar paracone and metacone subequal
60 (1) upper molar postcingulum greatly expanded into shelf
Node S
22 (1) upper ultimate premolar postcingulum greatly expanded into shelf
23 (1) upper ultimate premolar hypocone present
60 (1) upper molar postcingulum greatly expanded into shelf

J Mammal Evol (2015) 22:307342

64 (1) dentary has one mental foramen

Node T: Cryptotopos?
11 (3) lower ultimate premolar width greater than 0.8 of the
length
33 (0) lower molar paraconid cuspate in shape
42 (1) lower molar cristid obliqua has notch halfway along
the length
Node U: Cryptotopos?
27 (0) lower molar trigonid anteroposteriorly compressed
43 (2) cristid obliqua is convex occlusally
54 (0) upper molar centrocrista absent or weak
Node V
55 (0) upper molar precingulum absent or weak
56 (0) upper molar paraconule small, not strongly winged
57 (0) upper molar metaconule small, not strongly winged
Node W
47 (1) lower molar postcingulid present
Node X: Paradoxonycteris + Euronyctia
54 (2) upper molar centrocrista dilambdodont
56 (2) upper molar paraconule large, not strongly winged
57 (1) upper molar metaconule small, strongly winged
Node Y
25 (0) M1 trigonid wider than talonid
26 (0) M2 trigonid wider than talonid
33 (0) lower molar paraconid cuspate in shape
37 (2) lower molar protoconid larger than metaconid
Node Z: Scraeva + Limaconyssus
13 (1) lower ultimate premolar paraconid is large cusp
28 (0) lower molar trigonid height twice that of the talonid

Appendix 3. Characters Used in Cladistic Analysis

1. Lower incisor shape: conical (0), spatulate (1), or pectinate (2)
2. Second lower incisor number of cusps: one (0), two (1),
three (2), four (3), or five (4)
3. Lower canine posterior shelf or basal cusp: absent (0) or
present (1)
4. Second lower premolar number of roots: two (0) or one (1)
5. Lower penultimate premolar4 number of roots: two (0)
or one (1)
6. Lower penultimate premolar metaconid: absent (0) or
present (1)
7. Lower penultimate premolar protoconid orientation:
dorsoventral (0) or anteriorly canted (1)
4
Posterior premolars are referred to as penultimate and ultimate to avoid
confusion because Maelestes gobiensis has five premolars and
nyctitheriids have four. It is understood that the third premolar is eventually lost in eutherians when the count goes down to four, so that the P4 and
P5 of Maelestes are homologous to the P3 and P4 of nyctitheriids.

337

8. Lower penultimate premolar size relative to other premolars: similarly sized to large (0) or small (1)
9. Upper penultimate premolar protocone: present (0) or
absent (1)
10. Upper penultimate premolar metacone: absent (0) or
present (1)
11. Lower ultimate premolar shape (width/length):
less than 0.5 (0), 0.5-0.65 (1), 0.65-0.8 (2), or more
than 0.8 (3)
12. Lower ultimate premolar paraconid position: absent
(0), present and low on trigonid (1), or present and high on
trigonid (2)
13. Lower ultimate premolar paraconid size: absent
or very small cuspule on low anterior cingulid (0) or
large cusp (1)
14. Lower ultimate premolar paraconid or anterior-most
projection of tooth position: on midline or buccal to midline
(0) or lingual to midline (1)
15. Lower ultimate premolar metaconid: absent (0), swelling (1), or present (2)
16. Lower ultimate premolar width of talonid: narrower
than trigonid (0) or as wide as or wider than trigonid (1)
17. Lower ultimate premolar talonid basin: absent (0) or
present (1)
18. Lower ultimate premolar talonid number of cusps: one
(0), two (1), or three (2)
19. Upper ultimate premolar highly waisted: present (0) or
absent (1)
20. Upper ultimate premolar anterodorsal shift (from labial
view): absent (0) or present (1)
21. Upper ultimate premolar precingulum: present (0) or
absent (1)
22. Upper ultimate premolar postcingulum size: small (0),
greatly expanded (1), or absent (2)
23. Upper ultimate hypocone: absent (0) or present (1)
24. Upper ultimate premolar metacone: absent (0) or present (1)
25. M1 trigonid width: wider than talonid (0), equal to
talonid (1), or narrower than talonid (2)
26. M2 trigonid width: wider than talonid (0), equal to
talonid (1), or narrower than talonid (2)
27. Lower molar5 trigonid length: anteroposteriorly compressed (0) or about equal to talonid or longer (1)
28. Lower molar trigonid height: nearly twice the
height of the talonid (0), less than twice the height of

5
For characters that just specify lower molar or upper molar, all taxa
were coded using the second molar, if possible. If that position was
unknown, the first molar was used.

338

the talonid (1), or nearly the same height as the talonid

(2)
29. M1 and M2 length: subequal, within 5% (0), M1 longer
than M2 (1), or M2 longer than M1 (2)
30. M2 and M3 length: M3 longer than M2 (0), subequal,
within 5% (1), or M2 longer than M3 (2)
31. Lower molar (more visible on the M1) metaconid
twinned, or a groove runs alongside the cristid
obliqua up the posterior surface of the metaconid: absent (0) or present (1)
32. Lower molar precingulid: strong, anteriorly projecting
(0) or reduced, not projecting (1)
33. Lower molar paraconid shape: cuspate (0) or crestiform
(1)
34. Lower molar paraconid anteriorly projecting: absent (0)
or present (1)
35. Lower molar paraconid position: on midline (0) or
lingually positioned (1)
36. Lower molar protoconid position relative to metaconid:
protoconid anterior to metaconid (0) or the cusps are transverse (1)
37. Lower molar protoconid and metaconid size:
metaconid is larger (0), subequal (1), or protoconid is larger
(2)
38. Lower molar protoconid and metaconid height:
subequal (0), protoconid is taller (1), or metaconid is
taller (2)
39. Lower molar labial cingulid: absent (0) or present along
length of tooth (1)
40. Lower molar cristid obliqua position: ends lingual to
notch in protocristid (0), ends labial to notch in protocristid
(1), or ascends to apex of metaconid (2)
41. Lower molar mesoconid on cristid obliqua: absent (0)
or present (1)
42. Lower molar notch in cristid obliqua: absent
(0), present, located halfway along cristid obliqua
(1), or present, located on distal third of cristid
obliqua (2)
43. Lower molar cristid obliqua shape in labial view:
straight or slightly concave occlusally (0), deeply concave
occlusally (1), or convex occlusally (2)
44. Lower molar hypoconulid position: on midline
(0), lingual to midline (1), close to entoconid in
nyctalodont condition6 (2), or incipient on postcingulid
to absent (3)
45. Lower molar entoconid position: anterolingual to
hypoconulid (0) or on posterolingual margin of tooth and even
posteriorly with hypoconulid (1)

6
sensu Menu and Sig 1971: The entoconid and hypoconulid are closely
appressed but separated by a fissure and the hypoconid and hypoconulid
are connected by a crest.

J Mammal Evol (2015) 22:307342

46. Lower molar entoconid height: subequal to hypoconid

(0) or taller than hypoconid (1)
47. Lower molar postcingulid: absent (0) or present
(1)
48. Upper molar shape (length/width): less than 0.7 (0),
0.7-0.95 (1), or greater than 0.95 (2)
49. M1 ectoflexus: present (0) or absent (1)
50. M1 parastylar lobe orientation: anteriorly projecting (0)
or anterolabially projecting (1)
51. M2 ectoflexus: present, relatively deep (0) or absent to
very shallow (1)
52. Upper molar stylocone (stylar cusp B): present (0) or
absent (1)
53. Upper molar mesostyle: absent (0) or present
(1)
54. Upper molar centrocrista (between protocone and
metacone): absent or weak (0), strong, rectilinear (1), or
strong, dilambdodont (2)
55. Upper molar precingulum: unlobed and small (0),
unlobed and large, reaching labially past paraconule (1),
or with pericone (2)
56. Upper molar paraconule: small and not strongly winged
(0), large and strongly winged (1), or large and not strongly
winged (2)
57. Upper molar metaconule: small and not strongly
winged (0), large and strongly winged (1), or large and not
strongly winged (2)
58. Upper molar paraconule and metaconule positions: level with each other transversely, closer to
paracone and metacone than protocone (0), paraconule
more lingual than metaconule and located halfway between paracone and protocone, metaconule closer to
metacone than protocone (1), or paraconule more lingual than metaconule and located closer to protocone
than paracone, metaconule halfway between metacone
and protocone (2)
59. Upper molar paracone and metacone height: paracone
taller than metacone (0) or subequal (1)
60. Upper molar postcingulum size: absent or small (0) or
expanded into a large shelf (1)
61. Upper molar hypocone: absent (0) or present (1)
62. Upper molar postcingulum shape: flat posteriorly (0) or
rounded posteriorly (1)
63. M3 parastylar lobe size: large, expanded anterolabially
(0) or small (1)
64. Number of mental foramina: two (0) or one (1)
65. Location of anteriormost mental foramen: below P1 or
anterior (0), below P2 (1), or below penultimate premolar or
posterior
66. Location of posteriormost mental foramen (this character is coded if only one mental foramen): below penultimate
premolar or more anterior (0), below ultimate premolar or
more posterior (1)

J Mammal Evol (2015) 22:307342

Appendix 4. Data Matrix Used in Cladistic Analysis

MAELESTES_GOBIENSIS 000000000000000000000000000000000000000000000000000000000000000011
ADUNATOR_MINUTUS 100000000112101011101201221102001011000100000000010100011100101011
MACROCRANION_JUNNEI ????????102100100010120022011?011010100100000100010100022100101011
PLAGIOCTENODON_KRAUSAE
???00011??1210211[12]??000?1111010?0111?001000011???0??0?1112?01??020
PLAGIOCTENODON_SAVAGEI ????0011??12102112???????111??0?0111?001000011????????????????????
PLAGIOCTENODON_DORMAALENSIS
???00??1??22002112100011[12]111010001111001000000010001011112101101-0
PLAGIOCTENODON_ROSEI
24100011012210211[12]100001211101000111100100000101[01]00101111210110[01]10
PLAGIOCTENODON_THEWISSENI
241000110102102112100001211101000111100100000101[01]00101011210110010
LEPTACODON_TENER ???0000010011021111000012111010001112101000000010?01011112[01]0110010
LEPTACODON_MUNUSCULUM
???00?????2[12]112012???00011110200100110010000100???????????????????
LEPTACODON_CATULUS ???00???0?31?120121?0001111111001?01100000?00?100001011112?0110???
LEPTACODON_PACKI ???00?00??2110201110000011111100011111110001000110010?011200010010
LEPTACODON_NASCIMENTOI ??????????1110201???????1111??0001110201000000011001002012?0110???
LEPTACODON_ACHERONTUS ????0010??21102112???????1110100010110010000000??????????????????0
LEPTACODON_DONKRONI 23??0??0??1????111011000?1112?00100120010000000?10?100011200010??0
LEPTACODON_CHORISTUS ????0010??32102011??????11110000010112010001000???????????????????
NYCITHERIUM_VELOX ???00000??1111201[12]11011121010100011110120011001101011110011111?1-0
NYCTITHERIUM_SEROTINUM ??????????1111201110011111010101011110020011001??????1?????????010
NYCTITHERIUM_KRISHTALKAI
???000000011112011110011100111010111120000?101?1010100101101110000
NYCTITHERIUM_CHRISTOPHERI
??????????????????111111???????????????????????0010101000111110???
ACRODENTIS_ROSENORUM ???00???00111020111000011101210101111200001100?10101011111001101-0
CEUTHOLESTES_DOLOSUS 23100100??32112112??????21011101011102000011011????????????????010
LIMACONYSSUS_HABRUS ???00?????12102012??????00101200011121020011001????????????????010
PONTIFACTOR_BESTIOLA ??????????????????1?0001???????????????????????1101012111111110???
WYONYCTERIS_CHALIX ??????????11?02012??????21010?110111110200120101101112211200100??1
WYONYCTERIS_RICHARDI ???10011??11102012??????210112100111100200120101101012211200110021
WYONYCTERIS_PRIMITIVUS ??????????01102112??????2?01??011111100200?2011?10110121120011????
WYONYCTERIS_GALENSIS ???00?????3110201???????11010211011111120012010????????????????011
WYONYCTERIS_MICROTIS ???????????????????????????????????????????????00000100112000-0???
PLAGIOCTENOIDES_MICROLESTES
???11001??11102110??????2?11??10111110020012010????????????????011
PLAGIOCTENOIDES_TOMBOWNI
???10?????11102110???????111?200111110020012010????????????????011
AMPHIDOZOTHERIUM_CAYLUXI
23111011??3100211[012]1111110011020000012102021[01]001110?10000030111?1-0
SATURNINIA_GRACILIS
2?100011002110211[012]1101111111020[01][01]101[12]100000000110001000002011101-0
SATURNINIA_MAMERTENSIS
??100??1??111?21121101111111?20[01][01]11???020000000000110?0002?11101-0
SATURNINIA_GRANDIS 2?1?0001??21002?1???????1?11??001101100010000000???10?2112?1110???
SATURNINIA_RIGASII ??????????1100201???????00110?00010121001000001??????????????????0
SATURNINIA_CECILIENSIS ????????????????????????1111010011012?010000000???????????????????
SATURNINIA_INTERMEDIA ??????????31102112????????11??000101??02?1000?11???1012212?1110???
SATURNINIA_PIRENAICA ??????????2????????????????1??0?111???0????0?00?????012212?111????
SATURNINIA_PELISSEI ??????????3????????????????????????????2??????0?0001012212?111????
SATURNINIA_CARBONUM 23100011??31002?1????????101?001?1012101?001000????????????????1-0
CRYPOTOTOPOS_BEATUS
2?10001100[23]1[01]0211[12]1001111101010001011002112000010001002212011101-0
CRYPTOTOPOS_WOODI ???00?????11112012??????1?010101010121021120000????????????????1-0
CRYPTOTOPOS_HARTENBERGI
??10001101[23]100211[12]1101111101010[01][01]10121021120000[01]??01002212?11101-0
SCRAEVA_HATHERWOODENSIS ???00?????21102012??????00100?00010021021??0000????????????????10
PLACENTIDENS_LOTUS_P3 ????0100????????????????2111?2100110020200?3011?000101211[01]?011????
EURONYCTIA_MONTANA ??1000????310021121?11112?11?21011011?02021100111001020212?1110???
EURONYCTIA_GRISOLLENSIS
23?00001??[23]1[01]021121??1111111020[01][01]100100202110011???1020212?11101-0
PARADOXONYCTERIS_SORICODON
??????????????????1?1111??????????????????10001??001020212?111????
ASIONYCTIA_GUOI ???00000??121100001010001011010011112101000100001001010111001000[12]1
BUMBANIUS_RARUS ???00000??11112000??????10110100111[01]00010011020110010001110010???0
EOSORICODON_TERRIGENA
???10000??21111000??????001101001111210[12]0011020110010?020100110010
OEDOLIUS_PEREXIGUUS ???0001???01010000??????1011010011101?020010020????????????????0[01]0
EDZENIUS_LUS ???00?????01011000??????0?01??01011120010000020??????????????????0

339

340

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