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Original article

A South American snake lineage from the Eocene Greenhouse of North

America and a reappraisal of the fossil record of “anilioid” snakes

Jason J. Head

PII: S0016-6995(20)30097-8
DOI: https://doi.org/10.1016/j.geobios.2020.09.005
Reference: GEOBIO 935

To appear in: Geobios

Received Date: 24 March 2020

Revised Date: 25 July 2020
Accepted Date: 15 September 2020

Please cite this article as: J.J. Head, A South American snake lineage from the Eocene Greenhouse of North
America and a reappraisal of the fossil record of “anilioid” snakes, Geobios (2020), doi: https://doi.org/10.1016/
j.geobios.2020.09.005

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A South American snake lineage from the Eocene Greenhouse of
North America and a reappraisal of the fossil record of “anilioid”
snakes 

Jason J. Head *

Department of Zoology and University Museum of Zoology. Downing St. University of

Cambridge. CB2 3EJ Cambridge, UK

* E-mail address: jjh71@cam.ac.uk


Corresponding editor: Thierry Smith.

Abstract
“Anilioidea” is a likely paraphyletic assemblage of pipe snakes that includes extant
Aniliidae from equatorial South America, Uropeltoidea from South and Southeast
Asia, and a fossil record that consists primarily of isolated precloacal vertebrae
ranging from the earliest Late Cretaceous and includes geographic distributions in
North America, South America, Europe, and Africa. Articulated precloacal vertebrae
from the middle Eocene Bridger Formation of Wyoming, attributed to Borealoilysia
nov. gen., represent an unambiguous North American aniliid record and prompts a
reconsideration of described pipe snakes and their resultant biogeographic histories.
On the basis of vertebral apomorphies, the vast majority of reported fossils cannot be
assigned to “Anilioidea”. Instead, most records represent stem taxa and
macrostomatans erroneously assigned to anilioids on the basis of generalized
features associated with fossoriality. A revised fossil record demonstrates that the
only extralimital distributions of fossil “anilioids” consist of the North American aniliid
record, and there is no unambiguous fossil record of Old World taxa. The occurrence
of aniliids in the mid-high latitudes of the late early Eocene of North America is
consistent with histories of northward shifts in equatorial ecosystems during the early
Paleogene Greenhouse.
Keywords:
Serpentes
Aniliidae
Pipe snake
Systematics
Eocene
North America

1. Introduction
Snakes are a highly diverse, successful vertebrate clade occupying nearly all
non-polar terrestrial, as well as tropical marine, habitats. As such, they are
extensively studied with respect to phylogenetics, evolutionary tempo and mode,
evolutionary development, evolutionary ecology, and origins (Cohn and Tickle, 1999;
Pyron and Burbrink, 2012; Head and Polly, 2015; Hsiang et al., 2015; Leal and Cohn,
2016; Streicher and Wiens, 2016; Myers et al., 2019). Fossils play a central role in
providing temporal, paleoenvironmental, and biogeographic data for such studies
(e.g., Burbrink et al., 2020), which requires rigorous, testable phylogenetic
hypotheses for the identity of fossil taxa in order for them to provide accurate
information (Parham et al., 2012).
The vast majority of the snake fossil record consists of isolated or associated
precloacal vertebrae, and descriptions and interpretations of the record have
historically been inconsistent in justification for, and meaning of, taxonomic
assignments based on vertebral morphology. As a result, the described fossil record
of snakes consists of taxonomic histories that are a conflation of phylogenetically
justified assignments and generalized “form taxon” assignments that are not explicitly
phylogenetic (Head, 2015). This inconsistency has resulted in inflated temporal and
geographic distributions for extant snake clades, which becomes positively
misleading for neontological studies (Noonan and Chippindale, 2006).
The fossil record of pipe snakes represents a prominent example of the
problems with taxonomic identification. Extant pipe snakes include monotypic
Neotropical Anilius scytale and Southern Asian Cylindrophis and Anomochilus
species. They are fossorial and occasionally semi-aquatic taxa that inhabit wet
lowland forests and specialize in preying primarily on elongate vertebrates and
invertebrates (Greene, 1983; Himstedt et al., 2003). Traditional morphological
phylogenetic hypotheses place pipe snakes with uropeltids (shield-tailed snakes) in a
monophyletic clade historically named Aniliidae and subsequently Anilioidea (Bellairs
and Underwood, 1951; Hecht, 1959; McDowell, 1975), which is either sister-taxon to
booid snakes in the clade Henophidia or sister-taxon to Macrostomata, the clade that
includes booids and caenophidians (Rieppel, 1988; Kluge, 1991; Tchernov et al.,
2000; Lee and Scanlon, 2002; Wilson et al., 2010; but see Cundall et al., 1993;
Gauthier et al., 2012). Conversely, molecular phylogenetic analyses find strong
support for paraphyly of Anilioidea with respect to the majority of snakes: Anilius is
consistently recovered as the sister taxon to New World tropidophiid dwarf boas,
whereas Cylindrophis, Anomochilus, and uropeltids either form a poorly resolved
clade with more derived taxa or form the monophyletic Uropeltoidea as sister-taxon
to the clade composed of pythonids, Loxocemus, and Xenopeltis (Gower et al., 2005;
Vidal et al., 2007, Vidal and Hedges, 2009; Vidal et al., 2009; Wiens et al., 2012;
Pyron et al., 2013a, 2013b; Reynolds et al., 2014, Figueroa et al., 2016). Combined
molecular-morphological analyses either recover the generalized molecular topology
with Anilius+Tropidophiidae as sister taxon to all other alethinophidians and
Cylindrophis+Uropeltidae as henophidians (Harrington and Reeder, 2017) or recover
either the morphological or molecular topology, depending on constraint methods
(Hsiang et al., 2015).
The fossil record has the potential to inform on these radically different
hypotheses in terms of timing and geographic distribution. Rochebrune (1880) first
identified a fossil aniliid, “Scytalophis” lafonti from the Eocene of Quercy (see
Hoffstetter, 1955 for reidentification of the taxon as booid), but Hecht (1959) was the
first study to explicitly list vertebral characters considered unique to aniliids in
assigning fossils. Fossil aniliid/“anilioid” taxa have been subsequently described from
the early Late Cretaceous through Pliocene on all continents except for Australia
(e.g., Auffenberg, 1963; Rage, 1974; Holman, 1976; Szyndlar, 1994; Rage, 1998;
Gardner and Cifelli, 1999; Rage and Werner, 1999). Most assignments have used
character combinations similar to Hecht (1959), but many of these characters are
generalized fossorial ecomorphologies present in other snakes and non-snake
squamates (Hoffstetter and Gasc, 1969). Although a few anilioid fossil taxa have
been reidentified as boid (Rage, 1984), reliance on coarse, generalized characters in
assignment persists (e.g., Syromyatnikova et al., 2019), and there has been no
previous attempt to examine which, if any, vertebral characters are diagnostic for
pipe snakes and their relatives.
Here I describe an aniliid snake lineage from the middle Eocene Bridger Fm.,
Green River Basin, Wyoming, based on description of new material, reassessment of
previously described specimens, and re-examination of diagnostic vertebral
morphology within a molecular phylogenetic topology. I then discuss the implications
of the revised record for inferring evolutionary histories of pipe snakes. Because the
molecular phylogenetic topologies I use to infer both vertebral character histories and
the relationships of fossil material support paraphyly of the traditional concept of
Anilioidea with respect to other alethinophidians, I follow the taxonomy of Vidal et al.
(2009) in restricting Aniliidae to Anilius scytale and its fossil relatives and identifying
the clade of Old World taxa as Uropeltoidea. Subsequent discussion of “Anilioidea”
refers explicitly to the traditional, inclusive monophyletic hypothesis.

Institution Abbreviations: AMNH, American Museum of Natural History; BMNH,

Natural History Museum, London. DMNH, Denver Museum of Nature and Science;
TMM, Texas Memorial Museum, University of Texas; UCMP, University of California
Museum of Paleontology; UMZC-R, University Museum of Zoology, University of
Cambridge, Reptile collection; UNSM, United States National Museum (National
Museum of Natural History), Smithsonian Institution.

2. Geology and palaeoenvironments of the Bridger Formation

The Bridger Fm. crops out in the Green River Basin on the western side of the
Greater Green River Basin in Southwest Wyoming (Fig. 1). It is a ca. 842 m
sequence of predominately volcaniclastic mudrock with sandstone, limestone, and
ashfall tuff representing fluviolacustrine depositional environments (Murphy and
Evanoff, 2007, 2011). The Bridger Fm. is dated from 49.09 to 45.57 Ma and is
faunally zonated into four biochrons, from oldest to youngest, Br1b, Br2, Br3, and
Ui1a, within the Bridgerian and Uintan NALMAs (Gunnell et al., 2009; Murphy et al.,
2011). Palaeobotanical, palaeosol, and mammalian trophic structure and body size
distributions indicate decreasing temperatures through the Bridgerian from the high
of the Early Eocene Climatic Optimum at ca. 50-52 Ma (Woodburne et al., 2009), but
with persistent closed forest habitats, flooded soils, and paludal and lacustrine
deposition up to the top of Br3 (Gunnell, 1997; Murphy and Evanoff, 2007). Fossil
attributed to the taxon described here are from localities in Br2 and Br3 (Fig. 1).

3. Material and methods

In addition to the fossil specimen described here, I examined multiple
specimens of extant taxa for comparative purposes, including: Acrochordus javanicus
(UMZC-R.7.2-2 ), Anilius scytale (BMNH 56.10.16; BMNH 1855.5.28.3;
BMNH 1923.11.7.6; TMM 8281), Boedon lineatus (UMZC-R.7-430-1), Calabaria
reinhardti (UMZC-R3.41/1, UMZC-R3.41/2), Cylindrophis lineatus
(BMNH 1901.5.17.1), Cylindrophis maculatus (UMZC-R.4 11-2), Cylindrophis
melanotus (BMNH 1872.4.6.123), Cylindrophis ruffus (UCMP 136995; UMZC-R.4 12-
1; UMZC-R.4 12-2; UMZC-R.4 12-3; UNSM 523565), Epicrates cenchria (UMZC-
R.3.51-1), Epictia tenella (UMZC-R2.5/1, UMZC-R2.5/2), Malayopython reticulatus
(UMZC-R.3.24-4), Platyplectrurus madurensis (BMNH 1930.5.8.114), Plecturus
perroteti (UMAZ-R5.56-1), Python molurus (UMZC-R.3.25-7), Rhinophis philippinus
(UMZC-R.5.7-1), Trachyboa boulengeri (BMNH 1923.10.12.9), Tropidophis
haetianus (BMNH 1897.12.31.6), Tropidophis melanurus (BMNH 1932.11.11.33),
Typhlops punctatus (BMNH-R 1911.6.9.2), Uropeltis maculata (UMZC-R5.22-1),
Uropeltis woodmasoni (TMM-8663), and Xenopeltis unicolor (BMNH 1912.6.26.3,
UMZC-R6.1/1, UMZC-R6.1/2).
The fossil specimen was scanned using a Zeiss microXCT 400 micro-
computed tomographic scanner at the University of Texas High-Resolution X-ray
Computed Tomography Facility (UTCT), University of Texas at Austin, at a voltage of
70 kV and amperage of 0.833 A. Extant specimens were micro C-T scanned on a
Nikon Metrology XT H 225 ST High Resolution CT Scanner using a 225 kV X-ray
tube at the Cambridge Biotomography Centre (CBC), University of Cambridge. Scan
processing and visualization was conducted in VG StudioMax. I took measurements
for extant taxa from 2D images of precloacal vertebrae in anterior view from the data
set of Head and Polly (2015). I measured vertebrae at 5% intervals along the
precloacal vertebral column beginning at 25-30% and ending at 95% along the
anteroposterior axis of the column to avoid the higher morphological variation of
anterior and posteriormost precloacal vertebrae, which can be distinguished and
excluded from comparison by discrete regional characters. I took all measurements
using ImageJ v.2.0.0.

4. Systematic paleontology
Serpentes Linnaeaus, 1758 sensu Head et al., 2020
Alethinophidia Nopcsa, 1923
Aniliidae Fitzinger, 1826
Genus Borealoilysia nov. gen.
Derivation of the name: “Boreal” (Gr. “boreas”), meaning north, and “Ilysia”, a junior
synonym of Anilius, meaning “Northern Ilysia”, referring to the occurrence of an aniliid
in North America.
Type species: Borealoilysia carinata nov. comb.
Other included species: Borealoilysia gunnelli nov. gen., nov. sp.
Diagnosis (modified in part from Hecht, 1959): Aniliid snake possessing the
following characters: neural spine consisting of low lamina with concave dorsal
margin anteriorly connecting to a short, flat-topped, subtriangular process at the
posterodorsal edge the neural arch; posterior edge of neural spine bifid; accessory
ridges present lateral to the neural spine, terminating posteriorly at the edge of the
neural arch, resulting in a roughly tricarinate posterior edge of the arch in dorsal view;
posteromedian margin of neural arch thick and straight at posterior margins of neural
spine and accessory ridges, posteromedian notch absent; centrum anteriorly wide;
and anterior margin of zygosphene broadly concave.

Borealoilysia carinata nov. comb.

Holotype: AMNH 3826, incomplete precloacal vertebra (type specimen of Coniophis
carinatus; Hecht, 1959: pl. 52).
Type locality and age: “Hyposodus Hill”, Tabernacle Butte Locality 5, Bridger
Formation, Sublette Co., Wyoming, 42°25’N, 109°21’W (McGrew et al., 1959; West
and Atkins, 1970; Fig. 1). Late Bridgerian NALMA (Br3), 46.92  0.17 Ma (Robinson
et al., 2004).
Diagnosis: As for genus.
Borealoilysia gunnelli nov. gen., nov. sp.
Fig. 2
Derivation of the name: Species nomen in honor of vertebrate paleontologist Gregg
Gunnell for his extensive contributions to the paleontology and geology of the
Western Interior Paleogene, and for his mentorship and friendship.
Holotype: DMNH EVP 57990A, two articulated precloacal vertebrae.
Type locality and age: DMNH loc. 727, “Gatortown”, Bridger Formation, Black’s
Fork Member, Sweetwater Co., Wyoming, 41°17’N, 110°0’W (Fig. 1). Middle
Bridgerian NALMA (Br2), 47.96  0.13 Ma (Robinson et al., 2004).
Diagnosis: Borealoilysia species differing from B. carinata in possessing:
comparatively thick, short prezygapophyses; prezygapophyseal inclination at a
higher angle (~20° in B. carinata, ~23° in B. gunnelli); and comparatively reduced
accessory ridges on the neural arch roof.
Description: DMNH EVP 57990A consists of two nearly complete, articulated
precloacal vertebrae, missing only small portions of the cotylar margin on the anterior
vertebra and the left postzygapophysis and neural arch on the posterior vertebra. In
anterior view (Fig. 2(A)), the cotyle is large and slightly ovoid. The cotylar rim forms a
thin edge around most of the articular surface. Small paracotylar ventrolateral
processes (LaDuke, 1991; Head, 2005) are present as slight expansions of the
cotylar rim just medial to the synapophyseal bases. Lateral to the cotyle, the surface
of the vertebra is flat and lacks any form of paracotylar fossae or foramina.
Synapophyses are broad, with a bulbous dorsal diapophyseal articular surface and a
weakly defined, concave ventromedial parapophyseal articular surface. The ventral
surfaces of the synapophyses are shallowly concave and diverge from the centrum at
a low angle. There is no excavation on the ventral surfaces of the synapophyses to
form channels for the paralymphatic vessels, as seen in posterior precloacal
vertebrae (LaDuke, 1991). The ventral edge of the parapophyseal articular facets is
approximately level with the ventral edge of the cotyle, as in mid- to posterior thoracic
vertebrae of scolecophidians, Anilius, and uropeltoids, but unlike other
alethinophidian snakes, in which the synapophyses are more differentiated and
extend ventrally below the base of the centrum.
Dorsal to the synapophyses, the prezygapophyses are laterally short and
vertically thick and diverge from the body of the centrum at a high angle of
approximately 23°. Despite being laterally short, the high angle of the
prezygapophyses extends their lateral edge dorsal to the top of zygosphene. Small,
convex accessory process are present at the lateral edges of the prezygapophyses.
Medially, the neural canal is trefoil with a broadly convex dorsal margin. The
zygosphene is large with prominent zygantral articular facets.
In dorsal view (Fig. 2(B)), the anterior margin of the zygosphene is strongly
concave. The neural spine is present as a low, poorly-defined ridge-like lamina that
originates just posteriorly to the zygosphene and extends posteriorly to where it
expands to form a slightly wider, subtriangular process with a bifid posterior margin.
Lateral to the neural spine are two accessory ridges that originate at the approximate
mid-length of the element and extend posteriorly where they diverge laterally to reach
the posterior margin of the neural arch.
The prezygapophyseal articular facets are large, ovoid and anterolaterally
oriented and the long axes of the accessory processes are oriented in line with the
articular facets. A small, strongly concave interzygapophyseal ridge is present
between the pre- and postzygapophyses. Posteriorly, the lateral margins of the
postzygapophyseal articular facets extend as thin edges beyond the body of the
neural arch. The posterior margin of the neural arch is broadly concave, with the
median region approximately straight and thickened at the posterior edge of the
neural spine and the accessory ridges. Development of the posterior spine and ridge
appears to have inhibited development of the posteromedian notch present in most
alethinophidian and some stem snakes. The bifid posterior edge of the neural spine
extends slightly beyond the median margin.
In lateral view (Fig. 2(C)), the elements are dorsoventrally short. Anteriorly, the
diapophyseal articular facet of the synapophysis is hemispherical and posterodorsally
angled over the parapophyseal facet. Small lateral foramina are present posterior to
the diapophysis, just ventral to the interzygapophyseal ridge. Dorsally, the neural
spine is low with a sinuous to increasing concave dorsal edge of the lamina,
posteriorly. The lamina terminates at the dorsal edge of a short process at the
posteromedian edge of the neural arch. The process is anteroposteriorly narrow with
a flat top that is slightly angled posteroventrally. Lateral to the spine, the accessory
ridges are elevated and mound-like with a rounded dorsal edge. Ventrally, the
surface of the centrum is broadly concave.
 In ventral view (Fig. 2(D)), the centrum is wide anteriorly and narrows
posteriorly as in B. carinata nov. comb., but to a greater degree than in Anilius,
Cylindrophis, and uropeltids. The haemal keel is a poorly-defined flattened surface
that is triangular in outline with a narrow anterior vertex widening to a broad posterior
edge. Large, paired subcentral foramina are present toward the posterior end of the
central body at the lateral edges of the keel. Posteriorly, the condyle is differentiated
from the body of the centrum by a pronounced constriction.
In posterior view (Fig. 2(E)), the neural arch is depressed with a shallowly
convex dorsal edge. The posterior edge of the arch is rounded and inflated, forming a
continuous ridge to the carinae of the accessory ridges. The zygantrum is well-
defined and deep above the neural arch. Ventral to the arch, the condyle is wide and
subcircular in outline.

5. Discussion
5.1. Taxonomic validity of Borealoilysia nov. gen. and vertebral morphological
diagnosis of Aniliidae and Uropeltoidea
The type species of Borealoilysia nov. gen., B. carinata nov. comb., was
originally assigned to Coniophis (Hecht, 1959) but, as discussed below, that genus is
a taxonomic wastebasket of fossorial ecomorphs whose monophyly has long been
considered dubious, and to whom taxonomic referrals have been for convenience
(e.g., Rage et al., 2004; Head and Holroyd, 2008; Smith, 2013; Albino and Brizuela,
2014). The type specimen of Borealoilysia carinata nov. comb., an incomplete
precloacal vertebra, does not share any unique characters with Coniophis
precedens, the type species of the genus, but does possess unique apomorphies of
the posterior neural arch. The species is therefore retained and used to anchor the
new generic designation. Referral of Borealoilysia nov. gen. to Aniliidae requires a
review of vertebral apomorphies of Aniliidae and Uropeltoidea based on
reassessment of previous studies and specimen examination.
Hecht (1959) provided the first explicit vertebral character list for his Aniliidae,
consisting of Anilius, Cylindrophis, and Anomochilus: “(1) the lack of a well-
developed neural spine; (2) the presence of an upward-flaring zygosphene; (3) the
lack of a posterior notch in the neural arch; (4) the lack of a well-developed haemal
carina; and (5) the outline of the vertebrae, square and not broader than long.” Rage
(1984) (reiterated in Holman, 2000, and Ikeda, 2007), diagnosed his superfamily
Anilioidea in part on vertebral characters: “vertebrae depressed; zygapophysial
facets slanting; mid and posterior trunk vertebrae without hypapophyses; paracotylar
foramina absent”, and included within it the family Aniliidae, diagnosed in part on:
“vertebrae rather depressed; neural arch flattened; neural spine poorly developed;
caudal vertebrae without hemapophyses.” Hsiou et al. (2010) included a list of
characters shared by Anilius and Cylindrophis: “a clearly depressed neural arch;
prominent and strongly inclined zygapophyses; short prezygapophyseal process; a
shallow median notch in the posterior border of the neural arch; and a centrum not
markedly widened anteriorly.” A precise angle of greater than 20° for the
prezygapophyses was considered a shared character of Anilius, Cylindrophis, and
fossil taxa referred to Anilioidea sensu lato (Hsiou et al., 2010). Hsiou et al. (2010)
additionally listed a uniform centrum width and small prezygapophyses, as
characters shared by Anilius, Cylindrophis, and the fossil taxon Colombophis.
The coarse degree of morphological specificity provided in these diagnoses
encompasses characters present in a wider sample of taxa than those grouped into
Aniliidae, Uropeltoidea, or “Anilioidea”. For example, reduction of the neural spine
and depression of the neural arch are characters associated with fossoriality among
scolecophidians and macrostomatans (e.g., Hoffstetter and Gasc, 1969; Rage, 1984;
Lawing et al., 2012), and the absence of characters such as a haemal keel,
hypapophyses on mid-trunk vertebrae, hemapophyses, and paracotylar foramina, as
well as a possession of a reduced or absent posteromedian notch, uniform centrum
width, and small prezyagpophyseal accessory processes are plesiomorphies for
crown Serpentes (e.g., Hoffstetter, 1959; Lee and Scanlon, 2002; Wilson et al.,
2010). Additionally, these diagnoses either implicitly assume monophyly of Anilius
with Old World uropeltoids or treat “Anilioidea” as a form taxon (Hsiou et al., 2010;
see also Smith, 2013 for discussion). No study has examined vertebral morphology
of Aniliidae or Uropeltoidea (sensu Vidal et al., 2007) for diagnostic characters under
a hypothesis of “anilioid” paraphyly.
Examination of Anilius, Cylindrophis, and uropeltid species, as well as
representative scolecophidians and fossorial macrostomatans, provides revision of
these characters in order to determine the interrelationships of Borealoilysia nov.
gen. and the published fossil record of anilioids sensu lato. The following list
examines the most common characters used to identify “anilioids”:
 High-angled prezygapophyses (Fig. 3): High-angled zygapophyses with
angles exceeding 30° occur in taxa across Squamata (e.g., Hoffstetter and
Gasc, 1969: fig. 42), and measuring variation of prezygapophyseal angle within
the mid to posterior precloacal vertebral column in representative crown snakes
does not support a high angle as a diagnostic feature shared by Anilius and
uropeltoids to the exclusion of other taxa (Fig. 3). Uropeltoidea does possess
higher average angles than other taxa (>24°, represented by Cylindrophis and
Uropeltis); however, the approximate 20° angle in Anilius is lower than in at
least some typhlopid taxa (Fig. 3(D)) and is approached by boid taxa including
Charina and Eryx species (Fig. 3(H, I)). Intracolumnar variation within sampled
taxa is approximately 5-7°, which additionally limits the ability to identify or
distinguish taxa based on a limited sample size. Anilius and uropeltoids do
share a dorsal extension of the prezygapophyses; however, there is variation in
this character as well. In uropeltoids, the prezygapophyses will reach or extend
dorsally to the level of, or beyond, the zygosphenial roof due to the high angle
of the processes, whereas they generally extend just ventral to the roof in
Anilius due to the lateral elongation of lower-angled processes (compare Fig.
3(A-C));
 Neural spine (Fig. 4): The neural spine in squamates generally consists of a
thin lamina that extends along the anteroposterior midline of the neural arch
and is continuous with a more thickened posterior process that arises from the
posteromedian edge of the arch. In Anilius (Fig. 4(A.1, A.2)), the lamina is
reduced with a concave to straight anterodorsal margin that joins with the short
posterior process. There is anterior-posterior intracolumnar variation in spine
height, and the spine is more distinct in anterior precloacal vertebrae, becoming
progressively reduced posteriorly. Among uropeltoids, Cylindrophis completely
lacks any anterior lamina (Fig. 4(C.1, C.2)), and only a small ridge is present at
the midpoint of the neural arch in some uropeltids (Fig. 4(D.1, D.2));
 Depression of the neural arch (Figs. 3, 4): In Anilius and uropeltoids, the
neural arch is low with an approximately straight posterodorsal margin that
diverges ventrolaterally from the midline of the element at a low angle between
 ca. 10° and 20° in mid-posterior precloacal vertebrae. This differs from the
condition in most alethinophidian and examined scolecophidian snakes, in
which the neural arch is high and ventrolaterally curved or bi-angled (Fig. 4(B.1,
B.2)) where it roofs a comparatively elevated zygantrum, and the straight-line
angle of the arch, measured from the lateral edge of the postzygapophysis to
the base of the neural spine or midline of the element, is generally between 20°
and 30°;
 Posteromedian notch of the neural arch (Fig. 4): In most squamates, the
posterior margin of the neural arch is transversely straight. In alethinophidians,
the posterior margin of the neural arch is deeply medially emarginated forming
an acutely-angled notch that exposes the dorsal surface of the condyle and
posterior central body in disarticulated specimens. In Anilius and uropeltoids,
the posterior edge of the arch ranges from nearly straight in the most anterior,
and sometimes posterior vertebrae to shallowly emarginated in most elements.
The emarginated edges are broadly convex and well-developed consistently in
Anilius (Fig. 4(A.1)) and variably in uropeltid species (e.g., Uropeltis
madurensis). The notch is most weakly developed in Cylindrophis, where it is
nearly absent in some specimens, and most uropeltid species among examined
alethinophidians (Fig. 4(C.1, D.1));
 Development of the haemal keel (Fig. 4): A keel is present in Anilius and
variably present in uropeltoids. In Anilius, the keel is a narrow ridge with a wider
spatulate posterior extent in the anterior region of the precloacal vertebral
column and becomes slightly wider and medially constricted in mid and
posterior precloacal vertebrae (Fig. 4(A.3)) as the ventral surface becomes
progressively more excavated by subcentral paralymphatic fossae. In
Cylindrophis species, the pattern of anterior-posterior variation is similar to
Anilius, but the keel is wider and more poorly developed (Fig. 4(C.3)). In
uropeltids, development of the keel ranges from a similar extent as in Anilius to
practically absent (Fig. 4(D.3)), being only weakly defined by paralymphatic
fossae.
To test the systematic utility of these characters, I discretized different states
based on specimen observation (see above, Material and methods), and optimized
character states onto a tree topology that represents the consensus of molecular
studies (e.g., Pyron et al., 2013a, 2013b; Reynolds et al., 2014) (Fig. 5). I included
Borealoilysia nov. gen., based on character states of both B. carinata nov. comb. and
B. gunnelli nov. gen., nov. sp., and included Cylindrophis and uropeltid species to
represent Uropeltoidea because specimens of Anomochilus were not available for
study. I used parsimony ancestral state reconstruction in Mesquite 3.61 (Maddison
and Maddison, 2019) to infer potentially diagnostic characters for Aniliidae and
Uropeltoidea. The following is the list of character codings I used in analysis based
on the aforementioned character observations. The taxon-character matrix is
provided in Table 1:
1. Prezygapophyseal angle: > 25° = 0; 20°-25° = 1; <20° = 2;
2. Prezygapophyses extend to level of top of neural canal: present = 0;
absent = 1;
3. Neural spine lamina: present = 0; reduced = 1; absent = 2;
4. Angle of neural arch in posterior view: 20°-30° = 0; 10°-20° = 1;
5. Posteromedian neural arch: absent or shallowly concave = 0; present, weakly
defined = 1; strongly defined with acute edges = 2;
6. Haemal keel: absent = 0; broad, poorly defined = 1; narrow = 2.
Under the molecular phylogenetic hypothesis of pipe snake paraphyly,
traditional characters are homoplasies associated with fossorial habits between
Anilius and uropeltoids, as additionally indicated by the variable occurrence of
several of these characters in scolecophidians. Conversely, optimization of refined
character states reduces the amount of inferred homoplasy and provides
differentiating diagnoses based on vertebral morphology. Of the six characters
examined, only elevation of prezygapophyses to the level of the top of the neural
canal and a depressed neural arch optimize as homoplastic between Aniliidae and
Uropeltoidea (Fig. 5).
Aniliidae, based on Anilius, is diagnosable on the combination of: elongate
prezygapophyses elevated to just shorter than zygosphene, and angled at an
average of 20°-25°; neural spine with short anterior lamina running much of neural
arch length; and depressed neural arch. A narrow, medially emarginated haemal keel
on mid and posterior precloacal vertebrae additionally distinguishes Anilius from
uropeltoids but is present in the greater diversity of alethinophidians. Uropeltoidea is
diagnosable on the combination of: high-angled prezygapophyses (an average of
>24°); neural spine lamina absent or greatly reduced, spine restricted to posterior
edge of neural arch resulting in a saddle-shaped dorsal margin of the neural arch in
lateral view; depressed neural arch; and shallow concave posteromedian notch.
Within Uropeltoidea, examined uropeltid species all possess relatively narrow
zygosphenes with strongly concave dorsal margins, to the exclusion of Cylindrophis.
Borealoilysia nov. gen. shares the orientation and elevation of the
prezygapophyses, presence of a reduced neural spine lamina, and a depressed
neural arch with Anilius (Fig. 5), and B. carinata nov. comb. additionally shares a
narrow haemal keel. The neural spine lamina is more prominent in the type
specimens of B. carinata nov. comb. and Coniophis platycarinatus (see below)
(Hecht, 1959: pls. 52, 53) than in B. gunnelli nov. gen., nov. sp., though this may
represent taphonomic abrasion on DMNH EVP 57990A. The posteromedian notch of
the neural arch is more weakly developed in Borealoilysia nov. gen. and is more
superficially similar to the condition in uropeltoids. However, the posteromedian
margin of arch is uniquely thick and expanded in Borealoilysia nov. gen., suggesting
that the notch is apomorphically shallow relative to other taxa. Placement of
Borealoilysia nov. gen. as sister taxon to Anilius requires one less character
transformation than as sister to uropeltoid taxa, and the shared diagnostic characters
with Anilius provides justification for assigning Borealoilysia nov. gen. to Aniliidae.

5.2. Taxonomic validity of Borealoilysia nov. gen. species

There are numerous sources of variation in snake vertebral morphology that
can potential confound taxonomic assignment, including intracolumnar position,
ontogenetic growth, and individual variation. The diagnostic apomorphies of
Borealoilysia gunnelli nov. gen., nov. sp., relatively thick, short prezygapophyses,
higher-angled prezygapophyses, and comparatively reduced accessory ridges, could
simply be characteristic of subadult B. carinata nov. comb. or result from sampling
different regions of the vertebral column. DMNH EVP 57990A and AMNH 3826 are
both from the vertebral column posterior to the anterior, hypapophyseal-bearing
region, and possess the same degree of paralymphatic excavation on the ventral
margin of the centrum, indicating similar positions along the length of the posteriorly-
expanding paralymphatic system and the axial skeleton (Ottaviani and Tazzi, 1977;
LaDuke, 1991). Other differences, such as the lateral extension of the
prezygapophyses and the degree of neural arch carination, could be ontogenetic, but
the relative proportions of vertebral apophyses, the neural canal, and the thickened
zygosphene of the smaller B. gunnelli nov. gen., nov. sp. type specimen are not
consistent with subadult morphologies. One differentiating character between the two
species, the angle of the prezygapophyses, can only be considered a tentative
hypothesis until larger sample sizes are recovered from the Bridger Fm.
Hecht (1959) erected a second Coniophis species, C. platycarinatus, from the
same locality as Borealoilysia carinata nov. comb., and distinguished it on the
absence of the accessory processes of the neural arch and in lacking any haemal
keel on the centrum. Although poorly-described, the specimen referred to C.
platycarinatus, AMNH 3827, is distinct from Borealoilysia gunnelli nov. gen., nov. sp.,
because the latter possesses small accessory ridges and a distinctly bifid posterior
neural spine, which is clearly absent in AMNH 3827. Additionally, the posterior
margin of the neural arch is more emarginated in B. gunnelli nov. gen., nov. sp., the
subcentral foramina are more anteriorly positioned in the type specimen of C.
platycarinatus, and the angle of the prezygapophyseal accessory processes is
parallel to the long axis of the articular facet in B. gunnelli nov. gen., nov. sp., but
perpendicular in AMNH 3827. Smith (2013) considered the type of C. platycarinatus
to be a posterior precloacal vertebra referable to C. carinatus, which is corroborated
by the presence of deep subcentral paralymphatic channels in AMNH 3827. In the
absence of recognizable apomorphies, the validity of Coniophis platycarinatus is not
maintained here, and the taxon is referred to Borealoilysia sp. indet. (Table 2).

5.3. Reassessing the fossil record of aniliid and “anilioid” snakes

Examination of the published “anilioid” fossil record in the context of revised
diagnostic characters results in a reconsideration of the evolutionary and
biogeographic histories of uropeltoids and aniliids but also indicates a greater past
record of fossorial ecomorphologies among crown and stem snakes (Table 2). The
oldest South American record, Australophis anilioides, can be assigned to Aniliidae
based on the combination of neural spine morphology, prezygapophyseal height, and
shallow development of the posteromedian notch (Gomez et al., 2008); however, the
majority of past extralimital occurrences of Aniliidae and “Anilioidea” are either
taxonomically ambiguous or are referable to other snake clades. None of the North
American Coniophis records aside from the specimens here reassigned to
Borealoilysia nov. gen. can be referred to Aniliidae. The type species of Coniophis,
C. precedens, is represented by a single anterior precloacal vertebra from the
Maastrichtian (Lancian) Laramie Fm. of Wyoming (Marsh, 1892; Gilmore, 1938).
Hecht (1959) assigned it to Aniliidae based on his aforementioned list of characters,
however, the short, low-angled zygapophyses, minute prezgapophyseal accessory
processes, and vaulted and smoothly concave posterior margin of the neural arch of
the type specimen of C. precedens (USNM 2134) are more similar to other fossorial
alethinophidians or stem snakes.
Other North American records referred to Coniophis range from stem snakes
from the Cedar Mountain Fm. (latest Albian-Cenomanian) of Utah (Gardner and
Cifelli, 1999) which represents the geologically oldest record of snakes based on
unambiguous vertebral morphology, to uniquely elongate fossorial vertebral
morphologies from the late Eocene of North Dakota (Smith, 2013), to multiple
records of C. precedens from the Late Cretaceous through Paleogene (Estes, 1964;
Estes et al., 1969; Fox, 1975; Armstrong-Ziegler, 1978; Longrich et al., 2012). The
identity and validity of many of the C. precedens records have been questioned (e.g.,
Rage, 1984; Holman, 2000); however, the most recent analysis of Coniophis
precedens includes a hypodigm constructed on screen-washed records from multiple
Lancian-aged localities with some, but not all, vertebrae that are anatomically
consistent with the type of C. precedens (Longrich et al., 2012).
Reports of Coniophis outside of North America consist of records from the
Campanian of Sudan (Rage and Werner, 1999, after Salih et al., 2016; see also
Head, 2015; Head et al., 2016), Maastrichtian of India (Rage et al., 2004), late
Paleocene of Morocco (Augé and Rage, 2006), early Eocene of Brazil and Peru
(Rage, 1981, 1998; Albino, 1990; Albino, 1996; Albino and Brizuela, 2014). These
records are indeterminate specimens possessing fossorial characteristics (Table 2).
The genus Eoanilius and the type species E. europae was erected by Rage
(1974) on the basis of isolated vertebrae and a single quadrate from the upper
Eocene locality of Malpérié, Perrière (MP17b) Reference Level, France. Referral to
Aniliidae was based on a depressed neural arch and low, short spine. Quadrate
morphology is also similar to aniliids in being robust with a wide distal condyle for
articulation with the mandible. A second species, Eoanilius oligocenicus was erected
on the basis of isolated precloacal vertebrae from the upper Oligocene (MP28) of
Herrlingen 8, Baden-Württemberg, Germany (Szyndlar, 1994). Eoanilius oligocenicus
was distinguished from E. europae on the basis of a relatively more well-developed
neural spine, trilobate zygosphene, greater differentiation of the haemal keel, larger
synapophyses, and distinct subcentral and lateral foramina (Szyndlar, 1994).
Subsequent descriptions of Paleogene through Neogene herpetofaunas have
reported the genus throughout Europe and western Asia on the basis of precloacal
vertebrae (e.g., Szyndlar and Böhme, 1993; Szyndlar and Schleich, 1993; Duffaud
and Rage, 1997; Venczel and Sanchíz, 2006; Böhme, 2008; Szyndlar, 2009; Rage,
2013; Čerňanský et al., 2017; Syromyatnikova et al., 2019), making it the most
geographically and temporally widespread “anilioid” fossil record after Coniophis.
Despite the relative high density of the referred fossil record, and doubts about
the aniliid affinities of Eoanilius (e.g., Smith, 2013; Syromyatnikova et al., 2019), the
interrelationships and identity of the taxon remain poorly understood. Vertebral
elements of the genus, including types of both species, lack any unique apomorphies
of Aniliidae, Uropeltoidea, or “Anilioidea” under any phylogenetic hypothesis. Instead,
the low-angled zygapophyses, short but well-defined and elongate neural spines,
well-developed posteromedian notch, and small, circular cotylar-condylar
articulations are characteristic of alethinophidian taxa traditionally considered
Macrostomata (e.g., booids, caenophidians). Among these taxa, Eoanilius lacks any
caenophidian apomorphies (Head et al., 2016), and most likely represents a small-
bodied, fossorial booid (Smith, 2013). The quadrate referred to E. europae from
Malpérié does possess a similar morphology to extant Anilius and Cylindrophis in
being short with a wide mandibular condyle including a broad lateral malleolus
(Rage, 1974). However, some aspects of aniliid and uropeltoid quadrate morphology
are plesiomorphic for Serpentes (Zaher et al., 2009), and the specimen lacks other
features, including a suprastapedial process, which makes assignment to either
Aniliidae or Uropeltoidea tentative at best. Regardless, the Malpérié quadrate
remains the only potential evidence for a past dispersal of aniliids or uropeltoids into
Europe (Table 2).
Other potential aniliid/uropeltoid taxa such as Michauxophis occitanus from
the late Pliocene of Hérault, France (Bailon, 1988) and Hoffstetterella brasiliensis
from the early Eocene of Brazil (Rage, 1998) either completely lack apomorphies of
Aniliidae or Uropeltoidea, as in Michauxophis, or possess a mélange of
aniliid/uropeltoid and macrostomatan characters, as in Hoffstetterella.
Reassessing the fossil record in the context of apomorphic characters greatly
reduces the number of remains assignable to Anilioidea s.l. (Table 2). Among taxa
possessing apomorphies, Australophis and Borealoilysia nov. gen. can be referred to
Aniliidae based on the presence of a thin, low neural spine lamina, whereas
Colombophis species possess a suite of aniliid and uropeltoid characters (e.g., Hsiou
et al., 2010). As a result, the only definitive referral of the “anilioid” fossil record is to
New World Aniliidae. The crown clade of Old World Uropeltoidea has no
unambiguous fossil record, and biogeographic and paleoenvironmental histories
inferred from the interpretation of the European fossil record as a prior expansion of
the Old-World clade should be reconsidered.

5.4. Biogeographic implications from Borealoilysia nov. gen.

The occurrence of Borealoilysia nov. gen. in the middle Eocene of North
America represents the only unambiguous continental-scale extralimital occurrence
of Aniliidae or Uropeltoidea relative to extant distributions. The Northward distribution
of an otherwise equatorial clade during the Eocene is grossly consistent with patterns
seen in other taxa (Wing et al., 2005; Smith, 2009; Bourque et al., 2015); however,
these records demonstrate Northward range shifts coinciding with hyperthermal
events during the earliest Eocene, between six to seven million years older than the
fossils assigned to Borealoilysia nov. gen. The early Eocene snake record from the
Western Interior remains very poorly studied, and it is very likely that the first
occurrence of Borealoilysia nov. gen. predates the records described here. As a
result, it is possible that the Borealoilysia nov. gen. lineage moved Northward
concomitantly with other herpetofaunal and flora components of Paleogene
ecosystems, with earlier records remaining to be discovered. Conversely, Western
Interior climates, especially during the earliest Eocene hyperthermal, were dry
relative to the middle Eocene (Gunnell, 1997; Krause and Riggins, 2007). The humid,
wet environments of the Bridger Fm. (Gunnell, 1997) are most similar to habitats of
extant Anilius scytale among Western Interior paleoenvironments, and the
geographic distribution of Borealoilysia nov. gen. may have instead been regulated
by humidity more than the temperature-associated dispersals of early Eocene taxa.
Although lower than thermal maximum values (Woodburne et al., 2009),
paleotemperatures of the middle Eocene were still high enough (Zachos et al., 2008)
to support diverse subtropical to tropical ecosystems (e.g., Gunnell and Bartels,
1994, 2001; Gunnell, 1997, 1998), including a member of wet-tropically restricted
Aniliidae.
The dispersal routes required to explain the occurrence of an aniliid in North
America remain enigmatic. There is no geological evidence to support terrestrial
immigration between North and South America prior to the assembly of Central
America and the earliest formation of the Panamanian Isthmus by the early Miocene
(Farris et al., 2011). Thus, some over-water component of dispersal would have been
required to establish aniliids in North America. This route remains unknown, but the
occurrence of other South American clades in the Eocene of the Western Interior, as
well as Europe, demonstrates successful intercontinental dispersal during the early
Paleogene (Smith, 2006, 2011; Conrad et al., 2007; Scanferla and Smith, 2020).

6. Conclusions
The fossil record of “anilioid” snakes is not nearly as diverse as previously
described based on revision of systematically informative vertebral morphology. The
North American Paleogene record of Borealoilysia nov. gen. represents the only
continental-scale extralimital distribution of the clade and is a component of the larger
pattern of Northward expansion of equatorial faunas during the Paleogene
Greenhouse. There is no unambiguous record of Old World Uropeltoidea, despite a
wide geographic distribution, ecological specialization, and relative species richness
among extant taxa. Studies that use the fossil record to inform on evolutionary
histories, such as calibrating evolutionary rates and estimating biogeographic mode,
of extant snake taxa should rely only on fossil records that have been critically
evaluated (e.g., Parham et al., 2012).

Acknowledgements
Research was funded by a Natural Environmental Research Council Award
(NE/S000739/1) to JJH. For access to specimens, I thank R. Stucky, J. Sertich, and
T. Lyson (DMNS), M.T. Carrano (NMNH), D. Gower and P. Campbell (NHM-UK),
C.J. Bell (TMM), and N. Jones, M. Lowe, and R. Stebbings (UMZC). I thank J.
Maisano (UTCT) for C-T scanning, postprocessing, and image construction of
DMNH EVP 57990A and K. Smithson (CBC) for C-T scanning of extant specimens. I
thank J.I. Bloch and T. Smith for discussion and T. Smith for the invitation to
contribute. I thank A. Folie, K. Smith, and an anonymous reviewer for comments that
greatly improved the manuscript. Most of all, I thank Gregg Gunnell for his
mentorship and support through our many years of friendship.

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Table and Figure captions

Table 1. Taxon-character matrix for optimization of vertebral characters in

determining phylogenetic relationships of Borealoilysia nov. gen.

Table 2. Described fossil record of aniliid and “anilioid” snakes and their systematic
reassessments based on this study.

Figure 1. Map of Wyoming showing location of Greater Green River Basin (dark
grey). Fossil localities: 1, “Hyposodus Hill”; 2, DMNH loc. 727 “Gatortown”.

Figure 2. Borealoilysia gunnelli nov. gen., nov. sp. Articulated precloacal vertebrae in
anteroventral (A), dorsal (B), left lateral (C), ventral (D), and posterior (E) views.
Anatomical abbreviations: ar, accessory ridges; hk, haemal keel; nspl, neural spine
lamina; pcvlp, paracotylar ventrolateral processes; prz, prezygapophyses; psp,
posterior spine process; tpm, thickened posterior margin of the neural arch. Scale
bar: 5 mm.

Figure 3. Intracolumnar variation in size and orientation of prezygapophyses in

fossorial snakes. A. Anilius scytale (BMNH 56.10.16) showing measurements taken.
B. Cylindrophis ruffus. C. Uropeltis woodmasoni. D. Typhlops punctatus. E. Rena
dulcis. F. Xenopeltis unicolor. G. Calabaria reinhardti. H. Charina bottae. I. Eryx
johni. J. Micrurus fulvius. Drawn angles represent average values for intracolumnar
variation in prezygapophyseal angle. Angles and vertebral heights were
superimposed to the midline of the element laterally and average midpoint of height
vertically.

Figure 4. Precloacal vertebral morphology in representative aniliid, uropeltoid, and

tropidophiid taxa. High-Resolution C-T reconstructions of articulated precloacal
vertebrae in dorsal (1), left lateral (2), and ventral (3) views. A. Anilius scytale
(BMNH 1855.5.28.23), vertebrae 90-95. B. Tropidophis haetianus
(BMNH 1897.12.31.6), vertebrae 99-103. C. Cylindrophis ruffus (UMZC R4.12-3),
vertebrae 101-105. D. Rhinophis philippinus (UMZC R.5.7-1), vertebrae 58-62.
Anatomical abbreviations: pmn, posteromedian notch. All other abbreviations follow
Fig. 2.

Figure 5. Parsimony optimization of vertebral characters (Table 1) for ancestral

states on molecular topology of snakes. Taxon names follow Reynolds et al. (2014).
Numbers refer to characters and their state changes () for each clade.
Table 1.

Taxon 1 2 3 4 5 6
Squamate outgroup 0 0 0 0 0 0
Scolecophidia 1&2 1 2 0 0 0
Anilius scytale 1 0 1 1 1 2
Borealoilysia 1 0 1 1 0 1&2
Tropidophiidae 2 1 0 0 2 2
Boidae 2 1 0 0 2 2
Pythonidae 2 1 0 0 2 2
Cylindrophis 0 0 2 1 0&1 1
Uropeltidae 0 0 1&2 1 0&1 0&1&2
Bolyeridae 2 1 0 0 2 2
Caenophidia 2 1 0 0 2 2