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Naming and rediagnosing the Cretaceous
gekkonomorph (Reptilia, Squamata) from Öösh
(Övörkhangai, Mongolia)
ab
Jack L. Conrad
& Juan D. Daza
c
a
Department of Anatomy, New York Institute of Technology, College of Osteopathic
Medicine, Old Westbury, New York 11568, U.S.A.,
b
Division of Paleontology, American Museum of Natural History, Central Park West at 79th
Street, New York, New York 10024, U.S.A.
c
Click for updates
Department of Biological Sciences, Sam Houston State University, 1900 Avenue I,
Huntsville, Texas 77341, U.S.A.,
Published online: 28 Aug 2015.
To cite this article: Jack L. Conrad & Juan D. Daza (2015): Naming and rediagnosing the Cretaceous gekkonomorph (Reptilia,
Squamata) from Öösh (Övörkhangai, Mongolia), Journal of Vertebrate Paleontology
To link to this article: http://dx.doi.org/10.1080/02724634.2015.980891
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Journal of Vertebrate Paleontology e980891 (5 pages)
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DOI: 10.1080/02724634.2015.980891
SHORT COMMUNICATION
NAMING AND REDIAGNOSING THE CRETACEOUS GEKKONOMORPH
€ OSH
€
€ ORKHANGAI,
€
(REPTILIA, SQUAMATA) FROM O
(OV
MONGOLIA)
JACK L. CONRAD*,1,2 and JUAN D. DAZA3; 1Department of Anatomy, New York Institute of Technology, College of
Osteopathic Medicine, Old Westbury, New York 11568, U.S.A., jack.conrad@gmail.com; 2Division of Paleontology, American
Museum of Natural History, Central Park West at 79th Street, New York, New York 10024, U.S.A.3Department of Biological
Sciences, Sam Houston State University, 1900 Avenue I, Huntsville, Texas 77341, U.S.A., juand.daza@gmail.com
Downloaded by [University of Manitoba Libraries] at 14:21 28 August 2015
http://zoobank.org/urn:lsid:zoobank.org:pub:F16D073A-A0E2-4CA6-B797-756847D7C5AD
Although AMNH FR 21444 is an important early and relatively basal lizard whose internal skull structure was first
described more than eight years ago (Conrad and Norell, 2006),
this specimen has never been formally diagnosed or named. Several papers (Conrad, 2008; Daza et al., 2008, 2012, 2013; Daza
and Bauer, 2010; Gauthier et al., 2012) have used AMNH FR
21444 to help reconstruct ancestral states for various squamate
groups, especially Gekkonomorpha, its constituent clades, and
basal squamate anatomy.
Here, we formally describe, diagnose, and name AMNH FR
21444.
METHODS
The specimen was scanned at the High-Resolution X-ray CT
(HRXCT) Facility at The University of Texas at Austin. Details
of the equipment used are described in Conrad and Norell
(2006). For this study, we resliced the data set using Avizo Standard Edition 8.0.1 (FEI Visualization Sciences Group, Burlington, Massachusetts, U.S.A.). Three-dimensional reconstructions
and slice-by-slice animations are available online (see Conrad
and Norell, 2007).
Institutional Abbreviations—AMNH—American Museum of
Natural History, New York, U.S.A.; ZPAL, Zaklad Paleobiologii, Polska Akademia Nauk (Paleobiological Institute, Polish
Academy of Sciences), Warsaw, Poland.
SYSTEMATIC PALEONTOLOGY
REPTILIA SQUAMATA
Oppel, 1811
GEKKONOMORPHA F€
urbringer, 1900
NORELLIUS NYCTISAUROPS, gen. et sp. nov.
(Figs. 1, 2)
Holotype—AMNH FR 21444 is a nearly complete skull with
mandibles and partial hyoid, lacking the snout and mandibular
tips, the lateral parts of the otooccipitals, the squamosal, and the
mandibular suspensorium.
€ osh Basin, Altai region of CenType Locality—Red Mesa, O€
tral Mongolia, Mongolian People’s Republic.
Formation and Age—Although the age remains somewhat
uncertain, correlated exposures that appear to be of the same
age have been dated at around 130 Ma (Samiolov et al., 1988;
Watabe and Suzuki, 2000; Rougier et al., 2001; Prieto-M
arquez
et al., 2012).
Etymology—Named for our friend Mark A. Norell in honor of
his extensive work on Asian paleontology and squamates
*Corresponding author.
worldwide. Species epithet from ‘nychta’ (Greek: night), ‘sauros’
(Greek: reptile), ‘opsi’ (Greek: face).
Diagnosis—Norellius nyctisaurops (Figs. 1, 2) is a gecko-like
basal squamate, as identified by its braincase morphology. It differs from the basal gekkotan Hoburogekko suchanovi (Fig. 3A)
in possessing paired frontals and in lacking ventrally fused cristae
cranii encircling the olfactory stalk and tracts (Fig. 2A). Norellius nyctisaurops differs from Gobekko cretacicus (Fig. 3B) in
possessing a broader maxillary overlap of the prefrontal, an anteromedial process of the frontal that extends to about the same
anterior level as the anterolateral processes, a jugal with extensive participation on the postorbital bar, a broader vomer-palatine contact, a pineal foramen, presence of pterygoid teeth, and
an open Meckelian canal (Fig. 1C). Norellius nyctisaurops differs
from Bavarisaurus macrodactylus in possessing posteromedial
processes of the parietal and straight and tightly packed marginal
teeth (as compared with larger and recurved teeth in Bavarisaurus macrodactylus). Norellius nyctisaurops differs from Eichstaettisaurus schroederi in possessing posteromedial parietal
processes and paired frontals and parietals.
Description—Although one recent phylogenetic analysis of
relatively narrow scope has suggested an unresolved position for
AMNH FR 21444 (Daza et al., 2013), we follow the consensus of
studies, including two very inclusive treatments of Squamata, in
considering it a member of Gekkonomorpha (Conrad and Norell, 2006; Conrad, 2008; Daza et al., 2012; Gauthier et al., 2012).
The specimen is small, measuring only 15.39 mm in length. It has
large orbits, a complete postorbital bar and supratemporal arch,
and a broad pyriform recess (Fig. 1). The skull is broadest at the
level of the orbits and, apparently, tapered anteriorly. The lacrimal is absent; the maxilla and prefrontal form the margins of the
lacrimal foramen.
The prefrontal is robust and laterally overlapped by a tall nasal
process of the maxilla, as indicated by a facet on the prefrontal.
This facet also indicates that the maxilla reached the orbital rim.
There is no indication that the maxilla contacted the frontal. The
frontal process of the prefrontal does not approach the postfrontal (Fig. 1A) as in pygopodids and some modern geckos, such as
Phelsuma. The maxillary suborbital process and tooth row
extend beyond the level of the anterior one-fourth of the orbit.
There is no palatine process and no posterior emargination of
the maxilla for the ectopterygoid.
The large jugal is gently curved (Fig. 1A, B) rather than
strongly angled, and contacts the postorbital. The small postorbital is limited to the posterodorsal margin of the orbit (Fig. 1A,
B, D) and laterally overlaps the postfrontal. The postfrontal is
triradiate (with frontal, parietal, and postorbital processes) and
clasps the frontoparietal suture. The paired frontals bear deep
cristae cranii, but these do not approach one another (Fig. 2A).
The frontoparietal suture is anteriorly curved (Fig. 1A, D). The
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Conrad and Daza— Cretaceous gekkonomorph (Reptilia, Squamata) from O€
FIGURE 1. Norellius nyctisaurops, gen. et sp. nov. (AMNH FR 21444). A, dorsal view of the skull as preserved; B, left lateral view of the reconstructed skull and mandible; C, reconstructed mandible in medial view (reversed). Reconstructed skull in C, dorsal and D, ventral views. Unpreserved
areas appear as semiopaque layers. Abbreviations: a, angular; bo, basioccipital; c, coronoid process; cc, crista cranii; d, dentary; e, epipterygoid; ec,
ectopterygoid; f, frontal; j, jugal; m, maxilla; mfo, mandibular fossa; oo, otooccipital; p, parietal; pa, palatine; pbs, parabasisphenoid; pf, postfrontal;
pif, pineal foramen; po, postfrontal; prf, prefrontal; pra, prearticular; pro, prootic; pt, pterygoid; ptt, pterygoid tooth; rap, retroarticular process; sa, surangular; so, supraoccipital; sp, splenial; t, tooth; V, path of the trigeminal nerve or one of its branches; v, vomer.
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Conrad and Daza— Cretaceous gekkonomorph (Reptilia, Squamata) from O€
FIGURE 2. Details of the anatomy of Norellius nyctisaurops, gen. et sp. nov. (AMNH FR 21444). A, HRXCT transverse section through the frontals
demonstrating the cristae cranii extent; B, HRXCT, ventromedial view of the left pterygoid showing the tiny pterygoid teeth; C, digital reconstruction
of the braincase in ventral view; D, digital braincase endocast showing the semicircular canals and the tiny accessory endolymphatic duct. Abbreviations: aar, anterior auditory recess; aed, path of the accessory endolymphatic duct; asc, ascending semicircular canal; bo, basioccipital; bpt, basipterygoid processes; cc, cristae cranii; cpr, prootic crest; f, frontal bones; fo, fenestra ovalis; hsc, horizontal semicircular canal; lgr, lagenar recess; lhv, path
of the lateral head vein through the basipterygoid process; par, posterior auditory recess; pbs, parabasisphenoid; ptt, pterygoid teeth; V, path of the trigeminal nerve or one of its branches.
pineal foramen lies within the parietal. The parietal has short
posteromedial processes that may contact the supraoccipital.
The supratemporal processes are less than one-half the length of
the main body of the parietal. Each supratemporal process has a
small posterolateral facet that would have received either the
supratemporal or the squamosal (neither of which is preserved).
Although the vomers are incomplete anteriorly, their posterior
parts show that they are broad and plate-like, with wide palatine
contacts (Fig. 1E), contrasting with the reduced contact or hypokinetic joint of more deeply nested gekkonomorphs. The broad
palatines have a robust maxillary contact and a broad pterygoid
contact. There is no ectopterygoid-palatine contact. The narrow
ectopterygoids are anterolaterally oriented. The pterygoid has
ectopterygoid, palatine, and quadrate processes. Ventrally, it
bears a distinct dental ridge with several tiny teeth (Figs. 1E, 2B).
A well-developed columellar fossa supports the epipterygoid.
Conrad and Norell (2006) provided an extensive description of
the braincase, but we will offer some details here (Fig. 2C, D).
The fused nature of the otooccipital suggests that the animal is a
late subadult or adult (Rieppel, 1993). The elongate basipterygoid processes have lateral head vein canals, an unusual feature
also found in many gekkotans and xantusiids. The parabasisphenoid has narrow posterolateral processes that extend onto the
bases of the spheno-occipital tubercles (Figs. 1E, 2C). The
Vidian and cranial carotid canals are completely housed by
the parabasiphenoid. The prootic possesses an elongate alar
crest. The bifurcated supratrigeminal process has a facet that
probably contacted the inferior process, forming a foramen prootico (Daza et al., 2013) when in natural articulation. A trigeminal
foramen is present in the wide prootic crest (Figs. 1E, 2C). The
basioccipital forms the ventral part of the occipital condyle and
elongate spheno-occipital tubercles. The fine preservation of the
braincase allowed generation of an osseous labyrinth endocast.
There is no indication of a statolithic mass in the vestibule. The
semicircular canal arrangement (Fig. 2D) is intermediate
between the compact pattern around the vestibule of fossorial
squamates (Olori, 2010) and the well-spaced arrangement of
other lizards (Oelrich, 1956; Bever et al., 2005). There is a welldefined constriction between the vestibule and endosseous
cochlear duct (Witmer et al., 2008). An accessory endolymphatic
duct is connected to the anterior semicircular canal and opens
dorsal to the alar crest, similar to that in the pygopodid Aprasia
repens (Daza and Bauer, 2015).
The mandible is robust, with a postdentary part that is subequal in length to the dentary (Fig. 1B, C). The dentary lacks a
coronoid process and has an open Meckel’s canal. Posteriorly,
the dentary is deeply emarginated such that it is developed into
surangular and angular processes. The splenial extends for most
of the length of the dentary tooth row and extends posteriorly to
a point at the same level as the coronoid apex. The coronoid possesses a dorsally concave ventral margin. The anterior descending process partly invades the dentary-splenial suture. The
posterior descending process anteriorly borders the mandibular
fossa. The coronoid process is subtriangular. It possesses a weak
lateral flange lacking posterior development.
The surangular forms the entire margin of the anterior surangular foramen. An anteriorly oriented groove extends from the
anterior surangular foramen onto the posterior part of the dentary. The retroarticular process is posteriorly oriented without
the medial notch or lateral offset or spoon shape present in some
gekkotans.
DISCUSSION AND CONCLUSIONS
There are relatively few squamate fossils prior to the Late Cretaceous (Estes, 1983; Evans, 2003). Norellius nyctisaurops from
the Early Cretaceous of Asia represents a very basal gekkonomorph (Conrad and Norell, 2006; Conrad, 2008; Gauthier et al.,
2012; but see Daza et al., 2013, 2014). Given recent evidence suggesting that Gekkonomorpha is basal within Squamata (e.g.,
Townsend et al., 2004; Vidal and Hedges, 2004, 2009; Pyron
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Conrad and Daza— Cretaceous gekkonomorph (Reptilia, Squamata) from O€
FIGURE 3. Reconstructions of A, Hoburogekko suchanovi (derived from data in Alifanov, 1989; Daza et al., 2012, 2014) and B, Gobekko cretacicus
(derived from data in Borsuk-Bialynicka, 1990; Daza et al., 2013, 2014) in left lateral view. Unpreserved areas appear as semiopaque layers. Abbreviations: c, coronoid; d, dentary; e, epipterygoid; f, frontal; m, maxilla; oo, otooccipital; p, parietal; pm, premaxilla; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; rap, retroarticular process; sa, surangular.
et al., 2013), fossils of basal gekkonomorphs are especially
important for reconstructing and dating squamate phylogeny.
The importance of Norellius nyctisaurops is further
highlighted given the recent reexamination of the anatomy of
two other gekkonomorphs (Fig. 3), each of which represents a
modern-looking gekkotan or proximal gekkotan outgroup (Daza
et al., 2012, 2013). Thus, Norellius nyctisaurops and Parviraptor,
as basal gekkonomorphs or proximal outgroups to gekkotans
and higher squamates, are especially important. Unfortunately,
Parviraptor is known only from disarticulated and partial specimens (Evans, 1994).
Norellius nyctisaurops shows no gekkotan characteristics in its
dermatocranium (Fig. 1). Indeed, the roofing elements and palate are like those of basal squamates in exhibiting paired frontals
and parietals, a pineal foramen, complete postorbital and supratemporal arches, and ventrally open olfactory canals. By contrast, the braincase is gekkotan-like in having an enclosed lateral
head vein canal, perforate prootic crest, and a well-developed
supratrigeminal process; character states uniting it with modern
gekkotans. Thus, Norellius nyctisaurops demonstrates a distinct
decoupling of the evolution of braincase morphology and external skull morphology (Conrad and Norell, 2006).
Norellius nyctisaurops, as a basal gekkonomorph, may be close
to the base of Squamata (Conrad and Norell, 2006; Conrad, 2008;
Daza et al., 2012, 2013, 2014; Gauthier et al., 2012). Even so, the
elongate postorbital skull, curved and elongate jugal, long postdentary part of the jaw, and very gecko-like braincase differ
strikingly from the morphology seen in basal rhynchocephalians
(see Evans, 1980). Clearly, more Jurassic and Triassic squamates
are needed to help bridge the morphological gap between basal
lepidosaurs and modern Squamata.
ACKNOWLEDGMENTS
We are grateful to all those from AMNH and other institutions who have sought out and recovered Gobi fossils. Digital
scans of Norellius nyctisaurops were performed at the University
of Texas, Austin, HRXCT laboratory by J. A. Maisano and M.
Colbert. We thank the Maisanos for hosting J.L.C. on his visits
to Austin, Texas. For specimen access, we thank C. M. Mehling
(AMNH, Paleontology), M. Borsuk-Bialynicka (ZPAL), and D.
Frost, D. Kizirian, R. Pascocello, and M. G. Arnold (AMNH,
Herpetology). We thank A. M. Balcarcel and A. M. Bauer for
comments on an earlier draft of the manuscript. The Department
of Anatomy at NYIT College of Osteopathic Medicine, Division
of Paleontology at the American Museum of Natural History,
and the Department of Biological Sciences and the Office of
Research at Sam Houston State University supported this study.
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Submitted May 10, 2014; revisions received July 20, 2014;
accepted October 5, 2014.
Handling editor: Emily Rayfield.
Citation for this article: Conrad, J. L., and J. D. Daza. 2015. Naming and
rediagnosing the Cretaceous gekkonomorph (Reptilia, Squamata) from
€ osh (Ov€
€ orkhangai, Mongolia). Journal of Vertebrate Paleontology.
O€
DOI: 10.1080/02724634.2015.980891.