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ANNALS
OF
CARNEGIE
MUSEUM
31 December 2008
Vol. 77, Number 3, Pp. 321–402
ON THE CRANIAL OSTEOLOGY OF THE HISPANIOLAN SOLENODON,
SOLENODON PARADOXUS BRANDT, 1833
(MAMMALIA, LIPOTYPHLA, SOLENODONTIDAE)
John R. Wible
Curator, Section of Mammals
Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, Pennslyvania 15206
[email protected]
Abstract
The external and internal surfaces of the skull of the Hispaniolan solenodon, Solenodon paradoxus Brandt, 1833, are described and illustrated in
detail based on five museum specimens (one from Carnegie Museum of Natural History, the remainder from American Museum of Natural History).
Two of the specimens are juveniles that preserve sutural information that is lacking in the adults; one adult is bisected and exposes the interior of
the nasal and cranial cavities. A sixth specimen is a serially sectioned ear region of a juvenile from the Max-Planck-Institut für Hirnforschung that
provides information on soft-tissue structures.
The Hispaniolan solenodon has a peculiar skull with numerous highly unusual features among mammals. Included are: a small, oval ossification, the os proboscidis, jutting anteriorly from the ventral rim of the external nasal aperture; zygoma incomplete and jugal bone absent; exposure
of the ethmoid bone in the orbit but palatine not; numerous orbital foramina accommodating veins communicating across the midline ventrally in
the presphenoid and dorsally in the frontal; tiny optic foramina; venous transverse canal through basisphenoid; entoglenoid process of squamosal
but no postglenoid process; abutment between the rostral process of the malleus and the crista parotica of the petrosal; rostral and caudal tympanic
processes of the petrosal fused; large occipital emissary vein foramen endocranially between supraoccipital and parietal; large venous condyloid
canal through exoccipital; paired interparietals; mandible with two “angles,” a true angular process and a process for the digastric muscle. Perhaps
the most unexpected finding is that of a prootic canal for the lateral head vein in the petrosal bone. Prootic canals are broadly distributed in Mesozoic
mammals including a few Cretaceous eutherians, but heretofore are unknown in placentals.
Key Words: cranial foramina, Hispaniolan solenodon, osteology, Solenodon paradoxus, skull
INTRODUCTION
Solenodon paradoxus Brandt, 1833, is a peculiar placental insectivoran from Hispaniola with some well-known
cranial oddities. Among others, its submandibular (submaxillary) glands produce toxic saliva (Rabb 1959), and
the duct of the gland ends at the base of a deeply grooved
lower second incisor (Nowak 1991); its upper molars are
zalambdodont, V-shaped in occlusal view with a large central cusp and two buccally extending ridges (Gill 1883;
Asher and Sánchez-Villagra 2005); and it has an elongate
proboscis (more than 10% of head length) that is supported
at its base by a small, round bone, which is absent (Ottenwalder 2001) or “merely represented by somewhat hardened ‘horny’ cartilage” in the other living species, Solenodon cubanus Peters, 1864, from Cuba (Allen 1908:515).
Nesophontes Anthony, 1916, is another placental insectivoran (with dilambdodont molars, W-shaped in occlusal
view) endemic to the islands of the West Indies that was
thought (e.g., MacFadden 1980; Morgan and Woods 1986)
to have survived into the 20th century, but has probably
been extinct for several hundred years (MacPhee et al.
1999). A long history of phylogenetic hypotheses concerns
Solenodon, Nesophontes, the African zalambdodonts (tenrecs and golden moles), and dilambodont insectivorans
(shrews and moles) (see summaries in McDowell 1958;
Whidden and Asher 2001). Prior to the discovery of Nesophontes (Anthony, 1916), most authors (e.g., Mivart 1871;
Dobson 1882–1890; Allen 1910) allied solenodons with
African tenrecs, in part because of shared zalambdodonty
Wible.indd 321
(Fig. 1). After 1916, authors either continued to support
that relationship, including Nesophontes with solenodons
and tenrecs (e.g., Allen 1918) or excluding Nesophontes
(e.g., Simpson 1945; Van Valen 1967), or allied solenodon and Nesophontes with shrews (e.g., McDowell 1958;
McKenna and Bell 1997). Recent DNA sequence analyses
have excluded a close relationship between S. paradoxus
and tenrecs, linking the former with shrews and moles and
the latter with golden moles in Afrosoricida within Afrotheria along with elephant shrews, aardvarks, hyraxes, elephants, and sirenians (Stanhope et al. 1998; Emerson et al.
1999). S. paradoxus grouped with shrew, hedgehog, and
mole in a recent study sequencing portions of 16 nuclear
and three mitochondrial genes (Roca et al. 2004).
The cranial osteology of the solenodon has been the
subject of considerable attention, with the most confounding recurring issue being the early fusion of many cranial
sutures. The most extensive treatment of the skull in the
nineteenth century was the original description of S. paradoxus by Brandt (1833). It includes individual descriptions
of most cranial bones in Latin based on a single imperfect
skull (lacking the occiput) preserving sutures on the rostrum
and skull base in the Zoological Museum in St. Petersburg.
The original description of S. cubanus by Peters (1864),
based on a single skull in the Museum für Naturkunde
in Berlin, noted differences between the two species in
some cranial osteological features. Mivart (1868) included
some additional observations on the skull of the type of
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S. cubanus. For his monograph on insectivorans, Dobson
(1882–1890) dissected a solenodon specimen from Paris
that had the skin and integumentary muscles removed and
made observations on several cranial muscles.
The first decade of the twentieth century witnessed four
significant contributions to our understanding of solenodon cranial anatomy. Leche (1907) restudied the types of
the two species, detailing the milk dentition in S. cubanus
and the adult dentition in S. paradoxus and providing more
detail on the skulls of both. J.A. Allen (1908) described
and photographed the skulls of three S. paradoxus at the
American Museum of Natural History in New York and
one S. cubanus at the National Museum of Natural History
in Washington, D.C. In “The Orders of Mammals”, Gregory (1910) restudied the specimens of S. paradoxus in New
York, providing individual descriptions of most cranial
bones and foramina and the first line drawings with cranial
bones, processes, and foramina labeled (see also Gregory
1920b: fig. 110). These illustrations of a juvenile Hispaniolan solenodon with most cranial sutures represent the
most comprehensive for the genus to this day (reproduced
here as Fig. 1). For his comprehensive monograph on the
anatomy of S. paradoxus, G.M. Allen (1910) studied a series of at least 12 individuals at the Museum of Comparative Zoology in Cambridge; he reported on soft- as well as
hard-tissue anatomy including the cranial musculature and
brain, although his treatment of cranial osteology is not as
extensive as that of Gregory (1910). A more complete description of the solenodon brain is found in Boller (1969).
Perhaps the best known contribution on solenodon cranial osteology is McDowell’s (1958) monograph comparing the Greater Antillean insectivorans, Solenodon and Nesophontes. He included detailed, labeled line drawings of
the orbit, ear region, squamosal-dentary articulation, and
occiput of both forms and an oft-cited glossary on cranial
osteology and basicranial vasculature. However, the vast
majority of the illustrated specimens of S. paradoxus are
adults that lack much sutural information; for example, the
orbit illustration does not show a single suture. Another
oft-cited monograph with observations on solenodon is
MacPhee’s (1981) ontogenetic treatment of the ear region
in insectivorans, strepsirhine primates, tree shrews, elephant shrews, and tenrecs. He studied a serially sectioned
ear region of a juvenile solenodon and described the hardand soft-tissue auditory anatomy. More recently, Asher
(2001) reported on the solenodon vomeronasal organ; Ottenwalder (2001) has revised solenodon systematics at the
generic level with a multivariate analysis of largely craniodental measures; and Whidden (2002) dissected muscles of
the solenodon snout.
In addition to the above works, photographs and drawings of the solenodon skull have been published elsewhere
(e.g., Thenius 1989: fig. 143; MacPhee 1994: figs. 6, 8;
Starck 1995: fig. 208). However, as plagued most of the
classic works on this taxon, the early fusion of cranial
sutures constrains the utility of these illustrations. Study
of the single solenodon specimen in the Section of
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Mammals of Carnegie Museum of Natural History as part
of a phylogenetic project (Wible et al. 2007, in press) confirmed this early fusion. To facilitate that project, several
specimens were borrowed from the American Museum of
Natural History, including the juvenile figured by Gregory
(1910: fig. 18A1, A2) and McDowell (1958: fig. 1C), a
second slightly older juvenile, and a bisected adult skull.
These specimens and the dearth of detailed figures that include sutural information of the cranial osteology of this
unusual placental are the impetus for the current report. It
also represents another contribution in the author’s series
on cranial osteology of Recent mammals (see Wible 2003,
2007; Wible and Gaudin 2004; Giannini et al. 2006).
Materials and methods
The following specimens form the basis for the anatomical
descriptions.
(1) AMNH 28272 Solenodon paradoxus, Haiti, collected 1907. This juvenile with many open cranial sutures
retains some deciduous teeth, which have been described
by Allen (1908) and redescribed with corrections by McDowell (1958). Currently, the skull of the specimen is in
three principal pieces: right premaxilla, maxilla, lacrimal,
and part of the vomer; left premaxilla, maxilla, and part
of the vomer; and the braincase and posterior nasal cavity
consisting of the ethmoid and part of the vomer. The palatines, middle ear ossicles, and right ectotympanic are missing, and the nasals are represented by the posterior part
wedged between the maxillary processes of the frontals
and an isolated 21 mm long piece of the right nasal with
all but the posterior border preserved intact. The skull was
more complete previously, because Allen (1908: pl. 33)
published photographs of the whole skull in dorsal, ventral,
right lateral, and occipital views; the palatines, right ectotympanic, and nasals are complete in these photographs.
However, Allen (1908: pl. 32) also published photographs
of the isolated right premaxilla, maxilla, and lacrimal in
lateral and ventral views as they are preserved currently.
Gregory (1910: fig. 18A1, A2) published line drawings of
the whole skull in ventral and left lateral views and addressed aspects of its cranial osteology, including nerve,
arterial, and venous foramina (see Fig. 1). Ten year later,
Gregory (1920b: fig. 110) published a line drawing of the
right snout and orbit in lateral view. McDowell (1958: figs.
1C, 11C, 12B, 14B, 15B, 16B, 18C, 19B) published a line
drawing of the whole skull in lateral view and numerous
line drawings of the dentition with descriptions and observed that one of the palatines was missing at that time; as
noted above, both are missing currently.
(2) AMNH 185012 Solenodon paradoxus, New York
Zoological Society, received June 2, 1960. This juvenile
has the adult dentition, although the upper first incisor is
not fully erupted and the teeth are generally unworn. Many
cranial sutures are open, but fewer than in AMNH 28272.
(3) AMNH 212912 Solenodon paradoxus, New York
Zoological Society. This adult has few open sutures and
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Fig. 1.—Reproduction of Gregory (1910: fig. 18) with the abbreviation list repositioned from the original. Included are lateral and ventral views in A1
and A2 respectively of the juvenile Solenodon paradoxus, AMNH 28272 that is restudied here. See text for various amendments (e.g., deciduous tooth
loci; identification of orbitotemporal foramina; ethmoid orbital exposure rather than vomer). The long-tailed tenrec Microgale dobsoni, part of the original figure, is retained to illustrate the superficial dental and cranial resemblances to the solenodon.
moderately worn dentition.
(4) CM 18069 Solenodon paradoxus, male?, La Vega,
Haiti, collected 1908. This adult has few open sutures and
moderately worn dentition. Some soft tissues, including
the proboscis, are still attached. Carnegie Museum of
Natural History received this specimen on exchange from
the Museum of Comparative Zoology (MCZ number was
34839). It was most likely part of the collection studied by
Allen (1910), but he did not include specimen numbers.
(5) AMNH 28271 Solenodon paradoxus, female, Haiti,
collected 1907. This adult has few open sutures and the
heaviest worn dentition of all specimens. Allen (1908:
pl. 29–31) published photographs of the skull in dorsal, ventral, and left lateral views. According to Gregory
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(1910:241), “the skull of an adult female (No. 28271)
has been sectioned in the median line and affords valuable morphological details.” Currently, the skull is in three
pieces: the right half, which includes the nasal septum; the
left premaxilla and part of the left maxilla and nasal (to
just behind the ultimate premolar); and the remainder of
the left half.
(6) MPIH 6863 Solenodon paradoxus, sectioned juvenile skull base including both ear regions. MacPhee
(1981) described the tympanic surface of this specimen’s
ear region in considerable detail, including the course of
the basicranial arteries and nerves, and the origins of the
tensor tympani and stapedius muscles. MacPhee (1981)
listed this specimen as Solenodon sp., but it is identified as
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S. paradoxus on the individual slides.
During the past few years, the author and collaborators
have published a number of treatments on cranial osteology of extinct and extant mammals (e.g., Wible and Rougier
2000; Wible 2003, 2007; Wible and Gaudin 2004; Wible
et al. 2004, 2005a, in press; Giannini et al. 2006). The first
choice in the evolving terminology in those publications is
English equivalents of the Nomina Anatomica (fifth edition
1983) and Nomina Anatomica Veterinaria (fourth edition
1994), the latter abbreviated NAV in this paper. However,
in instances where the Nominae are inadequate, employed
are the most appropriate terms from the current comparative literature. Appendix 1 is a table of anatomical terms
employed here along with references and equivalents; preceding the appendix is a glossary of cranial structures associated with nerves, arteries, and/or veins. An anatomical
domain included in this report that is not included in the
prior ones, except Giannini et al. (2006), is the nasal cavity. Terminology for the structures of the olfactory recess
in the Nomina Anatomica Veterinaria (1994) and Evans
(1993) follows that of Paulli (1900a, 1900b, 1900c) and
Moore (1981). However, this terminology does not adequately discriminate the numerous scrolls and accessory
scrolls of the olfactory recess. Therefore, principally Smith
and Rossie (2006) and Maier (1993) are followed here. A
similar circumstance involves the terminology for the auditory ossicles, where the Nomina Anatomica Veterinaria
is supplemented by Henson (1961). For characterization
of sutural types, Evans (1993:219) is followed: (1) serrated suture, “one that articulates by means of reciprocally
alternating processes and depressions;” (2) squamous suture, “one that articulates by overlapping of reciprocally
beveled edges;” (3) plane suture, “one in which the bones
meet at an essentially right-angled edge or surface;” and
(4) foliate suture, “one in which the edge of one bone fits
into a fissure or recess of an adjacent bone.”
The dentition of Solenodon, both deciduous and permanent, has been well documented (e.g., Leche 1907; Allen 1908; McDowell 1958; Thenius 1989) and will not be
treated here. Nevertheless, a system of tooth identification
is needed for descriptive purposes. Published dental formulae for Solenodon (e.g., Leche 1907; McDowell 1958;
Thenius 1989) include three incisors, one canine, three
premolars, and three molars in the upper and lower dentition. There are two principal ways to number the teeth
in the classes with multiples in Solenodon, i.e., incisors,
premolars, and molars. The first way is a simple numbering from one to three for each class (e.g., I1, I2, I3 for upper incisors; i1, i2, i3 for lower incisors); the second way
employs a hypothesis of tooth position homology with
reference to an ancestral condition with more teeth. The
second is only relevant for the incisors and premolars of
Solenodon, because its molar count of three is the same as
that in basal eutherians (Ji et al. 2002; Kielan-Jaworowska
et al. 2004).
Regarding the premolars, two different hypotheses of
reduction in the premolar series of Solenodon have already
Wible.indd 324
been published. Without comment, some workers (e.g.,
Leche 1907; Gregory 1910) have referred to the premolars of Solenodon as P2, P3, P4, with the loss of the P1
from an ancestral condition of four premolars. In contrast,
McDowell (1958) used P1, P2, P4, with the loss of the P3
from an ancestral condition of four premolars. McDowell
(1958) identified the first upper premolar of Solenodon as
P1, because it lacks a deciduous predecessor as is usually
the case for the placental P1 (see also Luckett 1993); based
on the morphology of the deciduous predecessors for the
remaining two upper premolars, McDowell (1958) believed the permanent teeth to be most likely P2 and P4. A
major complication to these two numbering systems (P2,
P3, P4 and P1, P2, P4) is the current widespread view that
the primitive eutherian premolar count is five and not four
(McKenna 1975; Novacek 1986a, 1986b; Rougier et al.
1998; Nessov et al. 1998; Cifelli 2000; Kielan-Jaworowska et al. 2004; Wible et al. 2007, in press). The extra premolar of the five is in the middle of the series, and some
authors (e.g., Cifelli 2000; Kielan-Jaworowska et al. 2004)
number the five as P1, P2, Pc, P3, P4 to avoid confusion
with the older literature, whereas others (e.g., Nessov et al.
1998; Wible et al. 2007, in press; this report) employ P1,
P2, P3, P4, P5.
A third hypothesis of homology for the three premolars
of Solenodon based on a five premolar ancestral condition
is proposed here: P1, P4, P5. In agreement with McDowell
(1958), the upper first and third premolars of the three in
Solenodon are the first and ultimate of the ancestral eutherian condition and, therefore, should be numbered P1 and
P5 respectively. However, McDowell’s (1958) referral of
the upper second premolar of Solenodon as the P2 of the
ancestral eutherian condition is not accepted. Rather than
the P2, the upper second premolar of Solenodon is the penultimate premolar of the ancestral five or P4. In the juvenile S. paradoxus AMNH 185012, the newly erupted upper
second premolar is a tall, trenchant tooth set obliquely in
the maxilla (Figs. 6, 11), resembling the penultimate premolar of most Cretaceous eutherians (Kielan-Jaworowska
et al. 2004; Wible et al. 2004, 2007, in press); in the adults
studied here this tooth is considerably worn (Fig. 7). Additionally, there is no instance within Eutheria in which the
penultimate premolar is lost while a more anterior premolar
is retained. Although McDowell (1958) ultimately identified the upper second premolar of Solenodon as P2 and the
missing premolar as P3 (in the four premolar scheme), he
stated (p. 160) that “there is some doubt whether the missing premolars in both upper and lower jaws are the second
or the third.” The upper premolar formula hypothesized
here for Solenodon (P1, P4, P5) is repeated in the lowers (p1, p4, p5) with the lower first premolar a permanent
tooth without deciduous predecessor as occurs in the upper
jaw (McDowell 1958).
Regarding the incisors, no authors have offered explicit
hypotheses of incisor homologies for Solenodon compared
to the ancestral eutherian condition, which is widely held
(e.g., Rougier et al. 1998; Ji et al. 2002; Kielan-Jaworowska
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Fig. 2.—Solenodon paradoxus, AMNH 185012, drawings of skull in dorsal view. Abbreviations: fplac, facial process of lacrimal; fr, frontal; iof, infraorbital foramen; ip, interpariteral; M1r, anterolabial root of upper first molar; M3r, roots of upper third molar; me, mastoid exposure; mpfr, maxillary
process of frontal; mx, maxilla; mxf, maxillary foramen; na, nasal; nc, nuchal crest; P5r, anterolabial root of upper ultimate premolar; pa, parietal;
pdp, posterodorsal process of premaxilla; pmx, premaxilla; rt, foramina for rami temporales; sc, sagittal crest; so, supraoccipital; spp, septal process of
premaxilla; sq, squamosal; zplac, zygomatic process of lacrimal; zpmx, zygomatic process of maxilla; zpsq, zygomatic process of squamosal.
et al. 2004; Wible et al. 2007, in press) to be five uppers
and four lowers. In fact, few authors have commented
on the general problem within Placentalia caused by the
presence of no more than three upper or lower incisors in
all living group members (Thenius 1989). Ziegler (1971)
suggested that the general pattern of incisor reduction in
early therians was posteroanterior. There may be support
for this in the upper incisors of some Late Cretaceous eutherians that have the posteriormost incisor either entirely
within the maxilla (e.g., the fourth in the zalambdalestid
Kulbeckia Nessov, 1993, according to Archibald and Averianov [2003]) or in the suture between the premaxilla and
Wible.indd 325
maxilla (e.g., the fifth in the asioryctithere Asioryctes Kielan-Jaworowska, 1975, according to Kielan-Jaworowska
[1981]). In contrast, placentals tend to have the posteriormost incisor entirely within the premaxilla, suggesting that it was the posteriormost incisors of the Late
Cretaceous taxa that have been lost. Solenodon, for one,
is an exception as its third incisor lies on the premaxillamaxilla suture (see Descriptions). At least one exception
to Ziegler’s (1971) proposed posteroanterior direction of
loss for early therians has been posited for the lower incisors of Cretaceous Zalambdalestidae by Archibald and
Averianov (2003), who hypothesized that the i2 is lost in
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with figures placed near the bone(s) of greatest relevance.
For terms and synonyms, see Appendix 1; for information
on cranial structures associated with nerves, arteries, and/
or veins, see Glossary before Appendix 1.
Nasal (“na” in figures)
Fig. 3.—Solenodon paradoxus, AMNH 185012, photograph of skull in
anterior view. The anterodorsal aspect of the right premaxilla is damaged
and exposes the opening into the alveolus of the I1. Also damaged are
the tips of the right nasoturbinate and maxilloturbinate. Scale = 5 mm.
Abbreviations: I1, upper first incisor; iof, infraorbital foramen; mt, maxilloturbinate; na, nasal; nt, nasoturbinate; pmx, premaxilla; spn, septal
process of nasal; spp, septal process of premaxilla.
Zalambdalestes Gregory and Simpson, 1926 (preserving
i1, i3, i4) based on comparisons with the slightly older
Kulbeckia (preserving i1–4).
For now, the upper and lower incisors of Solenodon
are numbered as I1–3 and i1–3 respectively without a hypothesis of incisor loss from the ancestral condition. The
relevant extinct and extant forms considered to factor in
the relationships of Solenodon provide no bases for hypothesizing incisor reduction, because these taxa have the
same number of or fewer incisors than the solenodon (see
Asher et al. 2002). Additionally, there does not appear to
be a general rule of incisor reduction that can be applied to
Solenodon or to any other placental at this point in time.
Institutional Abbreviations
AMNH—Department of Mammalogy, American Museum
of Natural History, New York, New York
CM—Section of Mammals, Carnegie Museum of Natural
History, Pittsburgh, Pennsylvania
MPIH—Neurobiologische Abteilung, Max-Planck-Institut
für Hirnforschung, Frankfurt am Main, Germany
DESCRIPTIONS
The order of bone description follows that of previous
publications on other mammals by the author (e.g., Wible
2003, 2007; Wible and Gaudin 2004; Giannini et al. 2006)
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In dorsal view in AMNH 185012 (Fig. 2), the narrow,
elongated paired nasal is more than 41% of greatest skull
length with the maximum posterior extent just in front
of the anterior orbital rim. The lateral margins are near
parallel, widest posteriorly at their narrow contact where
they underlie the maxillae (“mx” in Fig. 2). Anterior to
the maxilla, the lateral margin has a lengthy contact where
it underlies the premaxilla (“pmx” in Fig. 2). Posteriorly,
the nasals form an elongate V underlying the frontals (“fr”
in Fig. 2). The anterior margin of each nasal is concave,
roughly U-shaped with a very short midline spine and a
much longer lateral arm in contact with the premaxilla.
The lateral arm does not reach as far forward as the premaxilla, and the nasals are recessed posteriorly at the osseous external nasal aperture. The nasals have half a dozen
tiny foramina, asymmetrically arranged, mostly in the posteriormost surface.
In anterior view (Fig. 3), the ventral surface of each nasal is concave, forming the roof of the dorsal nasal meatus.
On the midline of the anterior ventral surface, the nasals
produce a low, sharp ridge, the septal process (“spn” in
Fig. 3). The septal process does not extend the length of the
nasal, but is confined to roughly the anterior one-third. The
medial ventrolateral surface has a raised rounded ridge, the
nasoturbinate crest (“nt” in Fig. 3). Based on the left side
of AMNH 28271 (Fig. 21B), the bisected skull, the nasoturbinate crest extends the length of the nasal. Posteriorly,
it continues as the ethmoidal part of the nasoturbinate (Endoturbinale I of Paulli 1900a, 1900b, 1900c).
In CM 18069 and AMNH 212912, the internasal suture
is fused and the sutures between the nasals and their neighbors are barely perceptible (Fig. 4). The nasal is 40% of
total skull length in the former and 37% in the latter.
Os Proboscidis (“op” in Fig. 21B)
CM 18069 and AMNH 28271 preserve the os proboscidis,
the small bone at the posteroventral base of the proboscis
named by Brandt (1833). In CM 18069, the bone is nearly
circular (4.7 mm wide and 4.3 mm long) and held by ligaments to the premaxillae, anteroventral to the external nasal aperture (Fig. 12). Its ventral surface is more or less flat
with anterior and posterior rims slightly elevated; a small
foramen appears to the right of the midline. In AMNH
28271, the bone has been cut parasagittally (Fig. 20). The
ventral surface resembles that of CM 18069, including the
presence of a small foramen to the right of the midline.
The dorsal surface has a rounded midline eminence that is
roughly triangular in dorsal view, widest anteriorly where
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Fig. 4.—Solenodon paradoxus, AMNH 185012, CM 18069, and AMNH 212912, photographs of skulls in dorsal views. Scale = 10 mm. Carat points are
on the faint temporal lines in the bottom two specimens.
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Fig. 5.—Solenodon paradoxus, AMNH 28272, photograph and line drawing of braincase in dorsal view; rostrum is preserved separately. Abbreviations:
fr, frontal; ip, interparietal; me, mastoid exposure of petrosal; mx, maxilla (broken); mxfc, facet for maxilla on frontal; na, nasal (broken); nc, nuchal
crest; pa, parietal; pet, petrosal; rt, foramina for rami temporales; so, supraoccipital; sq, squamosal; zpsq, zygomatic process of squamosal.
the eminence continues laterally as a rounded, raised rim
that turns posteriorly to form a lateral rim. The midline
eminence is flanked on either side by a depression that
contains several small foramina. In medial view (Figs. 20,
21A, B), the section of the os proboscidis is cigar shaped
with the ventral side slightly concave and the dorsal
convex.
Premaxilla (“pmx” in figures)
In AMNH 185012 (Figs. 6, 11), the paired premaxilla
houses three incisors that decrease in size posteriorly, the
I1, I2, and I3. The I1 is set transversely, whereas the I2 and
I3 are longitudinally arranged. The I1 is separated from
the I2 by a diastema equivalent in length to the I3. In the
posterior half of this diastema is a small, empty alveolus
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that likely housed the dI2. AMNH 28271 and 212912 have
three incisors (Fig. 7), with the diastema between I1 and I2
longer than the length of the I2. CM 18069 differs in that
it has four incisors (Figs. 7, 12); the first two are clearly
the I1 and I2, but the identity of the third and fourth teeth
is unclear. The extent of wear is comparable on the two
teeth and not significantly different from their neighbors,
leading the author to posit that both are likely adult teeth.
There is some asymmetry: on the left side the third tooth is
subcircular and smaller than the subtriangular fourth tooth,
whereas on the right the third tooth is compressed anteroposteriorly and the subequal fourth tooth is subcircular.
Given that published tooth formulas for Solenodon include
three upper incisors (e.g., Leche 1907; Thenius 1989), it is
assumed here that CM 18069 represents a supernumerary
condition.
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Fig. 6.—Solenodon paradoxus, AMNH 185012, drawings of skull in left lateral view. Abbreviations: aof, antorbital fossa; as, alisphenoid; C, upper canine;
eo, exoccipital; eth, ethmoid; fr, frontal; gf, glenoid fossa; ham, pterygoid hamulus; hf, hypoglossal foramina; I1, upper first incisor; I2, upper second incisor; I3, upper third incisor; imc, incisivomaxillary canal; iof, infraorbital foramen; ip, interparietal; lac, lacrimal; lacf, lacrimal foramen; lacfe, lacrimal
fenestra; M1, upper first molar; M2, upper second molar; M3, upper third molar; M3r, roots of upper third molar; me, mastoid exposure of petrosal; mpfr,
maxillary process of frontal; mx, maxilla; na, nasal; nc, nuchal crest; oc, occipital condyle; os, orbitosphenoid; otcr, orbitotemporal crest; P1, upper first
premolar; P4, upper penultimate premolar; P5, upper ultimate premolar; pa, parietal; pal, palatine; pdp, posterodorsal process of premaxilla; pmx, premaxilla; rt, foramina for rami temporales; so, supraoccipital; spf + mpf, sphenopalatine and minor palatine foramina; sq, squamosal; zm, zygomaticus
minor attachment.
Sutures delimiting the premaxillae are well preserved in
the juveniles AMNH 28272 and 185012, with the following based on the latter. In lateral view (Fig. 6), the facial
process is roughly rectangular, longer than tall, and taller
anteriorly than posteriorly to accommodate the large I1.
The alveolus for the I1 is more ventrally positioned than
that of the posterior incisors, with AMNH 212912 showing
the most extreme differential in that feature (Fig. 7). The
I1 in AMNH 185012 is still erupting and is roughly perpendicular to the postcanine occlusal surface, whereas the
fully erupted tooth in CM 18069 and AMNH 28271 and
212912 is slanted posteroventrally (Fig. 7). The facial process has a tail (the posterodorsal process) extending from
the posterodorsal corner to just beyond the level of the
posterior root of the canine (“pdp” in Fig. 6). Not including the posterodorsal process, the facial process represents
more than 45% of the pre-orbital length, measured at the
lacrimal foramen. The ventral border of the posterodorsal
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process underlies the maxilla; the posterior border of the
facial process has a plane suture with the maxilla dorsally
and underlies that bone ventrally. The anterior border of
the facial process is convex. The dorsal half of that convexity is the premaxilla’s contribution to the external nasal
aperture; the ventral half is the extensive suture between
the right and left premaxillae, which slopes anterodorsally.
The surface of the facial process has a bulge over the root
of the I1, but this is flatter in CM 18069, AMNH 28171
and 212912. In all four specimens, the facial process contains half a dozen tiny foramina, asymmetrically arranged,
mostly in the posteriormost surface (Fig. 6).
In ventral view (Fig. 11), the premaxilla includes the
alveolar process housing the incisors and the palatine
surface, with the palatine process on the midline (“ppp”
in Fig. 11). In AMNH 185012, the alveolus for the I3 is
not entirely within the premaxilla; its posterior border is
formed by maxilla. Sutures delimiting the premaxillae and
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Fig. 7.—Solenodon paradoxus, AMNH 185012, AMNH 28271, CM 18069, and AMNH 212912, photographs of skulls in lateral views. Scale = 10 mm.
maxillae are all but obliterated in CM 18069 and AMNH
28271 and 212912 (Fig. 7). Nevertheless, there is faint indication at the alveolar margin that the premaxilla in these
adults is as in the juvenile and does not completely enclose
the alveolus of I3 (or the fourth incisor in CM 18069). The
palatine surface is a small contributor to the surface area of
the hard palate. It occupies the area immediately behind the
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anteriormost incisor and medial to the diastemata between
the first and second incisors. At the level of the diastema is
the oval incisive foramen, whose length approximates that
of the third incisor (“inf” in Fig. 11). The incisive foramen is almost completely within the premaxilla, with the
palatine process of the maxilla forming the narrow posteriormost border. The maxillae overlap the premaxillae
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Wible—On the Cranial Osteology of Solenodon paradoxus
at the M-shaped palatal suture, with the narrow paired
palatine processes of the premaxillae forming the central
arm, the base of which is opposite the middle of the third
incisor.
In anterior view (Fig. 3), the premaxillae’s contribution
to the external nasal aperture has the shape of a cursive W,
with the middle arm of the W much lower than the lateral
arms. The top half of each lateral arm is much thicker than
the lower half. Behind the middle arm of the W, in the
floor of the nasal cavity is a narrow gap and then a raised
longitudinal crest on each premaxillae, angled roughly
30° from the sagittal plane. This crest, the septal process
(“spp” in Fig. 3), extends the length of the premaxilla,
from the dorsal surface of the premaxillary body anteriorly
to the dorsal surface of the posterior extent of the narrow
palatine process discussed above (Fig. 9). In life, the Vshaped space between these crests, the sulcus septi nasi,
housed the bottom of the nasal septum, the bulk of which
was cartilaginous in this region (see Fig. 20). In AMNH
28272, the posterior ends of the septal processes of the premaxillae are sandwiched between the lateral laminae of the
vomer medially (“llv” in Fig. 9) and the septal processes
of the maxillae laterally. Between the septal process of the
premaxilla and the nasoturbinate crest is the maxilloturbinate or ventral nasal concha (“mt” in Fig. 3). In the medial wall of the nasal cavity in AMNH 28271, 28272, and
212912, there is a sharp conchal crest (“ccp” in Figs. 9,
21B), upon which the anterior part of the maxilloturbinate
rests. The conchal crest does not extend the full length of
the premaxilla, because it is recessed slightly from the anterior margin.
Maxilla (“mx” in figures)
The paired maxilla has an alveolar process housing the
canine and postcanine dentition, which includes three premolars (P1, P4, P5) and three molars (M1–3) (Figs. 6, 11);
as noted by Gregory (1910: fig. 18A1, A2; see Fig. 1­) in
the juvenile AMNH 28272 the second and third premolars
are deciduous teeth (Figs. 8, 9). Each maxilla has a facial,
palatal, orbital, and nasal surface, of which the last is visible only in AMNH 28271 and 28272. Sutures delimit the
maxilla from its neighbors in the juveniles AMNH 28272
and 185012, except that in the latter the orbital suture with
the lacrimal is partially fused (“lac” in Figs. 6, 8, 10, 11).
In CM 18069, AMNH 28271 and 212912, the maxilla is
delimited from the palatine on the palate (Fig. 12) and, in
the first only, from the lacrimal on the face and partially
within the orbit (Fig. 7).
In lateral view in AMNH 185012 (Fig. 6), the vertical
anterior border of the facial surface contacts the premaxilla, the horizontal dorsal border contacts the premaxilla,
nasal, and frontal, and the concave posterior border contacts the frontal and lacrimal. The maxilla overlaps the
premaxilla, nasal, and frontal, but most of the suture with
the lacrimal is plane. The most prominent feature of the
facial surface is the large infraorbital foramen (“iof” in
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331
Fig. 6), the anterior opening of the short infraorbital canal
(slightly less than the length of M1). The infraorbital foramen is positioned dorsal to the anterolingual root of M1
in all specimens. In anterior view (Fig. 3), the infraorbital
foramen is oval, slightly taller than wide. Anterior to the
infraorbital foramen, dorsal to the anterolingual root of the
ultimate premolar, as well as in the floor of the infraorbital
canal, are several alveolar foramina transmitting superior
alveolar nerves and vessels (“imc” in Fig. 6; “af” in Fig.
10). The course of the anteriormost alveolar canal can be
traced through the thin bone in the juvenile AMNH 185012
nearly to the premaxilla-maxilla suture because dark tissue has dried within the canal. In light of this course, it is
equivalent to the incisivomaxillary canal of the dog (Evans
1993). Anterodorsal to the anteriormost alveolar canal is a
tiny foramen that appears to emanate from the nasolacrimal canal. Between the infraorbital foramen and the frontal suture is a broad, shallow, ovoid antorbital fossa (“aof”
in Fig. 6) for the origin of levator labii superioris proprius
(Whidden 2002). The posteroventral margin of the facial
surface forms the anteroventral orbital rim and the short
anterior zygomatic process (“zpmx” in Figs. 2, 11). The
outer edge of the orbital rim has a raised, rounded orbital
crest, anteroventral to which in the adults is a pronounced
depression (“zm” in Fig. 6), for the zygomaticus minor
(Whidden 2002). The zygomatic process tapers to a thin
rod that projects posterodorsally opposite the back of M3
(Figs. 4, 7, 12).
In ventral view, the palatine processes of the maxillae
are the major components of the hard palate, extending
from the back of the incisive foramina at the level of the
anterior edge of I2 to the narrow wall posterior to M3 with
only a small, shield-shaped cutout for the palatine bones
posterior to M1 (“pal” in Fig. 11). The maxilla underlies
the palatine in their palatal suture. Each palatine process
of the maxilla contains many small foramina, in particular
medial to P1 and P4 and in the interdental spaces between
P5/M1, M1/M2, and M2/M3 (Figs. 11, 12). In addition,
AMNH 185012 has a small foramen in the palatomaxillary suture opposite the M2 protocone (“mapf” in Fig.
11); AMNH 28271 has two such foramina, both opposite
the M2 protocone on the right side but with the posterior
one on the left side opposite the anterior edge of the M3;
AMNH 212912 has three such foramina, the anterior two
opposite M2 and the posteriormost opposite the M2/M3
interdental space; and CM 18069 has an additional posterior one opposite the M3 postcingulum (the naming of
these foramina is considered with the palatine bone). The
intermaxillary suture is flat in AMNH 185012, slightly
raised in AMNH 28272, fused and slightly raised in CM
18069 and AMNH 28271, and fused and with a noticeable
crest in AMMH 212912 (Fig. 12). Also visible in ventral
view is the short zygomatic process of the maxilla, the
posterior root of which is opposite the M3 paracone in all
specimens.
Gregory (1910:244, fig. 18A2; see Fig. 1) observed
that “in the young skull (AMNH 28272) the palatal plates
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Fig. 8.—Solenodon paradoxus, AMNH 28272, photograph and line drawing of right rostrum (premaxilla, maxilla, and lacrimal) in lateral view. Abbreviations: C, upper canine; dI1, upper first deciduous incisor; dI2, upper second deciduous incisor; dP4, upper penultimate deciduous premolar; dP5,
upper ultimate deciduous premolar; I1, upper first incisor; iof, infraorbital foramen; lac, lacrimal; lacf, lacrimal foramen; lacfe, lacrimal fenestra; M1,
upper first molar; M3r, roots of upper third molar; mx, maxilla; mxf, maxillary foramen; P1, upper first premolar; P5, upper ultimate premolar; pdp,
posterodorsal process of premaxilla; pmx, premaxilla; zpmx, zygomatic process of maxilla.
of the maxillary do not come together completely at the
posterior end in the mid line and in the adult skulls is a
small fenestra at this point. This was also noted in Mivart
(1868, p. 124) in S. cubanus.” Allen (1908:515) noted in
AMNH 28270 and 28271 that “both adult skulls show a
small longitudinal vacuity in the palatine, but they look
more like accidental fractures than normal vacuities.” Regarding AMNH 28271, the specimen has been bisected off
the midline and there is no sign of a fenestra on the larger
right hand side. The other adult S. paradoxus studied here
show absolutely no fenestra at this point (Fig. 12), nor does
the specimen photographed in Thenius (1989: fig. 143). In
the juvenile AMNH 185012 the two maxillae are clearly
separated along their entire midline suture (Fig. 12), but
this is separation resulting from the immature stage of the
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specimen and not a palatal vacuity.
Based on sutural information from AMNH 185012
and 28272, the maxilla within the orbit contributes (Figs.
6, 10), to the anteroventral wall, the floor, and the ventral
orbital rim. The maxillary foramen, the posterior opening
into the infraorbital canal, is entirely within the maxilla
(“mxf” in Figs. 8, 10); the lacrimal approaches the dorsal
border of the foramen but is excluded but a 1–2 mm sliver
of maxilla. In the anteroventral wall, the maxilla contacts
the lacrimal dorsally at a horizontal plane suture, overlaps
the frontal posterodorsally at an irregular serrated suture
that is slanted posteroventrally, and overlaps the ethmoid
at an irregular suture that is slanted anteroventrally (“eth”
in Figs. 6, 10). In the pterygopalatine fossa, posteromedial
to M3, the maxilla has a narrow squamous suture with the
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Fig. 9.—Solenodon paradoxus, AMNH 28272, photograph and line drawing of right rostrum (premaxilla, maxilla, lacrimal, and part of the vomer) in
medial view. Abbreviations: ccm, conchal crest of maxilla; ccp, conchal crest of premaxilla; cP5, crypt for upper ultimate premolar; dI1, upper first
deciduous incisor; dI2, upper second deciduous incisor; dP4, upper penultimate deciduous premolar; dP5, upper ultimate deciduous premolar; eths,
surface for ethmoid bone; frs, surface for frontal bone; I1, upper first incisor; I2, upper second incisor; lac, lacrimal; llv, lateral lamina of vomer; M1,
upper first molar; mx, maxilla; nlc, nasolacrimal canal; P1, upper first premolar; pdp, posterodorsal process of premaxilla; pmx, premaxilla; rm, recessus maxillaris; ronlc, rostral opening of nasolacrimal duct; spp, septal process of premaxilla.
palatine (Fig. 10). A tiny sliver of maxilla in the anterodorsal part of the palatomaxillary suture contributes to
the dorsal margin of the sphenopalatine foramen, which
is closed by the palatine and ethmoid (“spf” in Fig. 10).
In the ventral orbital rim, the maxilla is overlapped by the
lacrimal (“zplac” in Fig. 2).
In the juvenile AMNH 185012 (Figs. 6, 10), some roots
of the ultimate premolar and the molars are exposed on the
face and in the orbit, because the maxilla fails to contain
them: specifically the anterolabial root of P5 in front of
the infraorbital foramen, the anterolabial root of M1 ventrolateral to the infraorbital foramen, and within the orbit the lingual root of M2 and all three roots of M3. In
the adults (Figs. 4, 7), roots are not exposed on the face
and the exposure differs within the orbit. Exposed in CM
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18069 are all three roots of M2 and the two labial roots of
M3, in AMNH 212912 are all three roots of M3 and the
lingual root of the right M2, and in AMNH 28271 are only
the labial roots of the left M3 and the posterolabial root
of the right M3. In addition to the exposed molar roots,
the orbit floor is riddled with numerous small openings
in the juvenile AMNH 185012 (Fig. 2). Most of these are
not preserved in the adults, although several foramina are
evident in all specimens between the lingual roots of M2
and M3, which may transmit additional superior alveolar
branches.
Exposed in the juvenile AMNH 28272 is the medial
surface of the disarticulated maxillae (Fig. 9), which
contributes to the floor and lateral wall of the nasal cavity.
The most salient feature is the conchal crest for the ventral
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lateral to the lateral lamina of the vomer (not visible in the
figures). The septal process is not of uniform height, but
is low anteriorly and posteriorly and nearly as tall as the
vomer in between.
Ventral Nasal Concha (“mt” in figures)
Fig. 10.—Solenodon paradoxus, AMNH 185012, photograph of right
orbit in oblique posterior view. Carats points outline the fusing suture between the lacrimal and maxilla. Scale = 5 mm. Abbreviations: af, alveolar
foramen; eth, ethmoid; fr, frontal; lac, lacrimal; lacf, lacrimal foramen;
lacfe, lacrimal fenestra; M3, upper third molar; mpf, minor palatine foramen (dorsal opening); mx, maxilla; mxf, maxillary foramen; pal, palatine; spf, sphenopalatine foramen; y, bar of bone forming posterior border
of minor palatine foramen, which is absent in CM 18069 (Fig. 13B).
nasal concha (maxilloturbinate), which extends from the
premaxillary conchal crest to the level of the M1 (“ccm”
in Fig. 9). The maxillary conchal crest is much sharper
and protuberant than that on the premaxilla. The crest is
concave ventrally, more dorsal and lateral anteriorly and
more ventral and medial posteriorly. Also visible between
the level of the P1 and dP5 is the bulge produced by the
nasolacrimal canal (“nlc” in Fig. 9). This bulge begins
posteriorly at the suture between the maxilla and lacrimal
and runs obliquely in an anteroventromedial direction. The
bulge intersects the conchal crest opposite the middle of
P1, and the nasolacrimal canal opens beneath the conchal
crest into the ventral nasal meatus anterior to that (“ronlc”
in Fig. 9). In the interval posterior to the intersection of the
conchal crest and nasolacrimal canal is an opening into the
crypt of the P5 (“cP5” in Fig. 9). At the posterior margin
of the medial maxillary surface ventral to the lacrimal is a
boot-shaped surface, with the toe pointing anteriorly. The
heel and throat (crural part) of the boot overlap the frontal
bone in the orbit (“frs” in Fig. 9) and the toe overlaps the
lateral wall of the ethmoid (“eths” in Fig. 9). The surface
anterior to the boot contributes to the lateral wall of the
maxillary recess (see Ethmoid; “rm” in Fig. 9); a maxillary
sinus is lacking. The maxilla’s contribution to the floor of
the nasal cavity is flat, except along the dorsal surface of
the intermaxillary suture, where there is a septal process
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The paired ventral nasal concha or maxilloturbinate attaches via a basal lamina to the conchal crests on the medial wall of the maxilla and premaxilla. The left side of the
bisected skull AMNH 28271 preserves much of the ventral
nasal concha (Fig. 21B); missing are small segments near
the external nasal aperture and opposite the P4 and P5.
Anteriorly, the basal lamina of the ventral nasal concha is
perpendicular to the lateral nasal wall, but posteriorly it is
near vertical. The pointed anterior end of the ventral nasal concha is visible through the external nasal aperture in
AMNH 185012 (Fig. 3). Immediately distal to the pointed
end, the ventral nasal concha is roughly C-shaped in cross
section, with horizontal and vertical arms, the former, the
basal lamina, attached to the lateral nasal wall. Proceeding distally into the nasal cavity, four primary scrolls are
identified, dorsoventrally, from one to four, based on the
left side of the bisected skull AMNH 28271 (Fig. 21B).
The first leaves the dorsal surface of the basal lamina at the
level of the I2 and is displaced posterolaterally by ethmoturbinate I. The second arises from the dorsal surface at the
pointed anterior end, but assumes a dorsomedial position
posteriorly, ending medial to ethmoturbinate II. The third
arises from the medial surface between the canine and P4,
but its full posterior extent is uncertain because of damage.
The fourth scroll is only visible at the level of P4 where it
lies beneath the maxillary conchal crest; the anterior and
posterior ends of this scroll are unknown. Three spaces
within the nasal cavity are associated with the ventral nasal concha: dorsally is the middle nasal meatus, medially
is the common nasal meatus, and ventrally is the ventral
nasal meatus.
Palatine (“pal” in figures)
The paired palatine is a small element contributing to the
back of the hard palate and the floor of the nasopharyngeal
meatus via a horizontal process, and to the medial wall of
the pterygopalatine fossa and the lateral wall of the choanae and basipharyngeal canal via a perpendicular process.
The horizontal processes of the palatines are delimited
from the maxillae by sutures in all specimens (Figs. 11,
12); the interpalatine suture is fused in CM 18069 and
AMNH 28271; and the sutures with other elements in the
pterygopalatine fossa and basipharyngeal canal are fusing
or fused in the adults (Figs. 6, 7).
On the palate (Fig. 11), the horizontal processes are
shield-shaped, with each specimen showing some leftright asymmetry in the outer contour of the palatomaxillary
suture. The palatal surface of the palatine is fairly flat
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Fig. 11.—Solenodon paradoxus, AMNH 185012, drawings of skull in ventral view. Abbreviations: as, alisphenoid; bo, basioccipital; bs, basisphenoid;
C, upper canine; e, ectotympanic; eo, exoccipital; fm, foramen magnum; gf, glenoid fossa; ham, pterygoid hamulus; I2, upper second incisor; inf, incisive foramen; M1, upper first molar; mapf, major palatine foramen; me, mastoid exposure of petrosal; mpf, minor palatine foramen; mx, maxilla; oc.
occipital condyle; os, orbitosphenoid; P4, upper penultimate premolar; pa, parietal; pal, palatine; pet, petrosal; pmx, premaxilla; ppp, palatine process
of premaxilla; ppt, postpalatine torus; pr, promontorium of petrosal; ps, presphenoid; pt, pterygoid; so, supraoccipital; sof, sphenorbital fissure; v,
vomer; zpmx, zygomatic process of maxilla; zpsq, zygomatic process of squamosal.
except along its posteriormost border, which sports a pronounced postpalatine torus (“ppt” in Fig. 11), especially in
the adults with parts showing considerable rugosity (Fig.
12). At the lateral limit of the postpalatine torus is the minor
palatine foramen (“mpf” in Fig. 11), which shows variability in the studied specimens related to the passage of two
principal neurovascular bundles from the pterygopalatine
fossa, one directed anteriorly onto the hard palate and the
other posteriorly into the soft palate. The juvenile AMNH
185012 and the left side of AMNH 212912 have a narrow
gap in the postpalatine torus (gap lies between the asterisks
in Fig. 13A); dorsal to this gap is a foramen (“dor” in Fig.
13A), which connects to the pterygopalatine fossa (“mpf”
in Fig. 10). On the right side of AMNH 212912, the narrow
gap in the postpalatine torus described above is closed by
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bone (“x” in Fig. 13A), and three separate foramina are delimited, which for descriptive purposes are called dorsal,
anteroventral, and posteroventral minor palatine foramina
(“dor”, “ant”, and “post” in Fig. 13A). The dorsal is interpreted as transmiting the minor palatine nerve and vessels
from the pterygopalatine fossa (“mpf” in Fig. 10); the anteroventral transmits branches to the back of the hard palate;
and the posteroventral transmits branches to the soft palate.
CM 18069 has no gap in the postpalatine torus (that is, “x” is
present in Fig. 13B), but it differs in that the bar of palatine
bone forming the posterior border of the dorsal minor palatine foramen in the other specimens (“y” in Figs. 10, 13A)
is absent (“y absent” in Fig. 13B); it has only one minor
palatine foramen equivalent to the anteroventral foramen of
the right side of AMNH 212912 (“ant” in Fig. 13).
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Fig. 12.—Solenodon paradoxus, AMNH 185012, CM 18069, and AMNH 212912, photographs of skulls in ventral views. Scale = 10 mm.
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In addition to the foramina in the palatomaxillary suture
described with the maxillary bone, the horizontal process
of the palatine is pierced by a variable number of small foramina showing considerable left-right asymmetry in number and position (Fig. 12). CM 18069 has the fewest with
one on the right and two on the left; AMNH 212912 has
four on both sides; and AMNH 185012 has four on the left
and five on the right (cf. Figs. 11, 13). In the dog (Evans
1993), the accessory palatine nerves exit via one or more
openings in the horizontal process (accessory palatine foramina of Wible and Gaudin 2004) and the major palatine
nerve exits via one opening in the anterior palatomaxillary suture (major palatine foramen of Evans 1993). Those
criteria are followed here for Solenodon and the openings
in or adjacent to the anterior palatomaxillary suture are
called the major palatine foramina and all others accessory
palatine foramina.
Based on the bisected skull AMNH 28271 (Fig. 21B),
the dorsal surface of the horizontal process, which forms
the floor of the nasopharyngeal meatus anterior to the
choanae, is essentially featureless. The left side has two
foramina, a larger anterior one opposite the M2/M3 embrasure and a smaller posterior one opposite the M3 (“mapf”
and “apf” in Fig. 22). The former is likely for the major
palatine nerve and the latter for the accessory palatine
nerve. Posterolateral to these openings is a large foramen
with a groove extending anteriorly from it (“spf” in Figs.
21B, 22), the anterior opening of a short canal leading
from the sphenopalatine foramen in the pterygopalatine
fossa. Based on the juveniles AMNH 28272 and 185012,
this opening is between the palatine and ethmoid. The right
side of AMNH 28271 (Fig. 20), which includes the midline is flat, with any foramina present covered by connective tissue.
Posterior to the postpalatine torus, the back edge of the
horizontal processes form the floor of the choanae (Fig. 11).
The back edge of each horizontal process is concave and
they meet on the midline to form a short posterior nasal
spine (“pns” in Fig. 13), although in AMNH 212912 they
meet off the midline and most of the spine is on the right
palatine (Fig. 13A). Most of the lateral walls of the choanae
and of the anterior half of the basipharyngeal canal behind
the choanae is formed by the medial side of the perpendicular process of the palatine, with the vomer completing the
walls and forming the roof (“v” in Fig. 11). The vomer overlaps the palatine anteriorly, but the vomeropalatine suture is
plane posteriorly. Forming the walls of the basipharyngeal
canal posterior to the palatines are the pterygoids (“ham” in
Fig. 11). The lateral side of the perpendicular process forms
the medial wall of the pterygopalatine fossa and is entirely
excluded from the orbit (Figs. 6, 14). The lateral side of the
perpendicular process has squamous sutures with the ethmoid and orbitosphenoid dorsally and the alisphenoid and
pterygoid posteriorly (“eth”, “os”, “as”, and “pt” in Fig. 14).
The anterior end of the lateral side of the palatine’s perpendicular process contributes to the formation of two foramina
in the pterygopalatine fossa, a larger more oval dorsal one,
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Fig. 13.—Solenodon paradoxus, photographs of rear aspect of palate in
ventral view. A, AMNH 212912; B, CM 18069. Scale = 5 mm. In A, on
the specimen’s right side, the course of the minor palatine nerve is via
three conduits: a dorsal one, the only complete foramen, that opens into
the pterygopalatine fossa in the orbit (Fig. 10); an anteroventral one that
opens onto the back of the hard palate; and a posteroventral one that opens
onto the soft palate. The bar of bone (x) separating the anteroventral and
posteroventral foramina on the right side is incomplete on the left side
(*). In B, the bar of bone (y) forming the posterior margin of the dorsal
foramen in A is lacking. Anterior to the postpalatine torus are a number of
foramina in the palatine or in the palatomaxillary suture; those in or near
the anterior palatomaxillary suture are termed major palatine foramina
and the remainder are accessory palatine foramina (see Palatine). Abbreviations: ant, anteroventral minor palatine foramen; dor, dorsal minor
palatine foramen; pal, palatine; pns, posterior nasal spine; post, posteroventral minor palatine foramen; ppt, postpalatine torus.
the sphenopalatine foramen (which includes the sphenopalatine and caudal palatine foramina of the dog (Evans 1993),
and a circular ventral one that is slightly more posterior, the
(dorsal) minor palatine foramen already mentioned above
(“spf” and “mpf” in Fig. 10). The palatine forms the ventral
half of the sphenopalatine foramen, which is completed by
the ethmoid and a sliver of maxilla.
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Fig. 14.—Solenodon paradoxus, AMNH 185012, line drawing of left orbit in oblique lateral view. Position of ethmoid is based on AMNH 28272 (see
“oleth” in Fig. 16). Abbreviations: aef, anteroventral ethmoidal foramen; as, alisphenoid; asc, alisphenoid canal; bs, basisphenoid; eth, ethmoid; fdv,
foramen for frontal diploic vein; fo, foramen ovale; fr, frontal; gf, glenoid fossa; ham, pterygoid hamulus; mpf, minor palatine foramen (dorsal opening);
mx, maxilla; of, optic foramen; os, orbitosphenoid; otc, anterior opening of orbitotemporal canal; pa, parietal; pal, palatine; pef, posterodorsal ethmoidal
foramen; pf, piriform fenestra (of opposite side); pt, pterygoid; sof, sphenorbital fissure; spf, sphenopalatine foramen; sq, squamosal; zpsq, zygomatic
process of squamosal.
Lacrimal (“lac” in figures)
The paired lacrimal is a small bone placed high in the anterior orbital rim (Figs. 2, 6). It has a very narrow, crescentic
facial process that is overlapped anteriorly by the maxilla
(“fplac” in Fig. 2). The posterior aspect of the facial process is thickened and forms the anterior orbital rim. The
lacrimal orbital crest is thickest in its central part, but not
enough to identify as a lacrimal tubercle. The orbital crest
curves posteroventrally as a zygomatic process of the lacrimal (“zplac” in Fig. 2) onto the dorsal root of the zygomatic process of the maxilla. The lacrimal is fully delimited
from its neighbors in AMNH 28272 (Figs. 8, 9), nearly so
in AMNH 185012 and CM 18069, with the orbital suture
with the maxilla fusing (Fig. 10), and not differentiated at
all in AMNH 28271 and 212912 (Figs. 4, 7).
In AMNH 185012, immediately posteromedial to the
orbital rim within the orbit is the large, oval lacrimal foramen (in essence, the fossa for the lacrimal sac), taller than
wide, which narrows anteriorly into the circular nasolacrimal canal for the nasolacrimal duct (“lacf” in Fig. 10).
The orbital surface of the lacrimal is narrow posteromedial and posteroventral to the lacrimal foramen where the
lacrimal contacts the maxilla at a plane suture and widest
posterodorsally where the lacrimal overlaps the frontal.
Within the orbit, the lacrimal overlaps the frontal dorsally and posterodorsally, contacts the orbital surface of the
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maxilla posteriorly, and overlaps the zygomatic process
of the maxilla posteroventrally (Figs. 2, 10). The suture
with the zygomatic process of the maxilla is nearly completely fused and that with the orbital surface is fusing.
Within the latter suture (also in AMNH 28272), dorsal to
the maxillary foramen is a narrow slit that likely represents
a lacrimal fenestra for the inferior oblique muscle (“lacfe”
in Figs. 6, 8, 10; Wible and Gaudin 2004). Dorsal to the
lacrimal fenestra in AMNH 28272 and 185012 is a small
vascular foramen reminiscent of that described in Pteropus
Brisson, 1762, by Giannini et al. (2006). In AMNH 28271,
212912, and CM 18069, the lacrimal fenestra is replaced
by a narrow pit with the closure of the suture between the
lacrimal and maxilla (Fig. 7).
The lacrimal is not as fully formed in the juvenile
AMNH 28272 as in the remaining specimens. Unlike the
adults, its lacrimal does not completely enclose the nasolacrimal canal, with the ventrolateral margin formed by
the maxilla (Fig. 8).
The medial surface of the lacrimal is visible on the right
side of AMNH 28272 (Fig. 9). It is wedge shaped, with
the point directed anteriorly, and dorsally and ventrally
has plane sutures with the maxilla. At the posterior limit
is a small triangular surface that overlaps the frontal in the
orbit. The remainder of the medial surface of the lacrimal
contributes to the maxillary recess (see Ethmoid).
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Jugal
The absence of a paired jugal bone has long been accepted
as a feature of the Hispaniolan solenodon (Gregory 1910;
Muller 1934), and all specimens examined here agree.
Frontal (“fr” in figures)
The paired frontal bone contributes to the braincase roof
via the squama frontalis and to the medial orbital wall via
the pars orbitalis. AMNH 28272 is the only specimen with
sutures delimiting the entire frontal (Figs. 5, 16); in AMNH
185012 some sutures within the orbit are not visible (Figs.
6, 14) and most of the interfrontal suture on the skull roof
is obliterated (Fig. 2).
The squama frontalis is described based on AMNH
185012 (Fig. 2); differences from the other specimens are
noted below. At the midline, the squama comprises less
than 20% of total skull length; at its greatest length, which
includes the elongate maxillary process (“mpfr” in Figs. 2,
6), the squama is more than 30% of total skull length. The
squama can be visualized as an ice cream cone, with the tip
of the cone projecting anteriorly as the maxillary process.
The maxillary process tapers to a point reaching the level
of the anterior (mesial) root of P4; it is separated from the
posterodorsal process of the premaxilla by approximately
4 mm. The maxillary process contacts the nasal anteromedially and the maxilla anterolaterally. The posterior interfrontal suture is raised slightly as the low, anteriormost
sagittal crest. Half a dozen small foramina are asymmetrically arranged just off the midline of the interfrontal suture. At its lateral limit, the squama bends ventrally into the
orbit as the pars orbitalis; a distinct orbital rim is lacking
and there is virtually no postorbital constriction. Posteriorly, the squama is overlapped by the parietal (“pa” in Fig.
2) along an asymmetric suture. The bulk of the suture is
roughly transverse, but at the midline is a small, irregular
U-shaped exposure of frontal between the parietals. The
surface of the squama is generally flat, with a slight bulging towards the pars orbitalis.
The squama frontalis of the adult specimens (Fig. 4)
differs in that the sagittal crest is slightly more raised and it
continues forward into low temporal lines that arch ventrolaterally into the anterior orbital rim at the lacrimal and are
too subtle to see in direct dorsal view. With this forward
position for the temporal line, the length of the temporal
fossa is more than 50% of total skull length. The squama
frontalis of the more immature juvenile AMNH 28272 differs in that, as noted above, the interfrontal suture is retained, the midline foramina and sagittal crest are absent,
and the sutures with the parietals are entirely transverse
(Fig. 5).
The pars orbitalis in AMNH 28272 is fully delimited
from its neighbors (Fig. 11), whereas in AMNH 185012
the ventral borders are unclear (Figs. 6, 14). The anterior
sutures with the lacrimal and maxilla have been described
above. Posterodorsally, the pars orbitalis overlaps the
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parietal at a vertical suture. Ventral to the parietal, the frontal underlies a narrow anteriorly directed process of the
alisphenoid (Fig. 6), which is twice as tall on the right side
as on the left in AMNH 185012. In AMNH 28272 (Fig. 16),
the frontal inferior to its contact with the alisphenoid has
two short, quadrangular processes, one directed ventrally
and the other posteriorly. The ventral process contacts the
maxilla anteriorly, the ethmoid (the vomer of Gregory
1910; see Fig. 1) ventrally, and the orbitosphenoid posteriorly, and the posterior process contacts the alisphenoid
dorsally and the orbitosphenoid posteriorly and ventrally.
The frontal’s sutures with the orbitosphenoid and ethmoid
are obliterated in AMNH 185012 (Figs. 6, 14).
In the ventroposterior pars orbitalis are a variable number of small to medium sized foramina, ranging from four
in AMNH 185012 to nine in AMNH 28272 (right side).
The bisected skull AMNH 28271 is the only specimen in
which the internal connections of these foramina can be
traced, and based on it, four categories of foramina are
identified here.
(1) Anterior opening of orbitotemporal canal (“1” in
Fig. 15). All specimens consistently have a posterodorsalmost foramen that is directed posteriorly. Previous authors
(e.g., Gregory 1910; McDowell 1958; MacPhee 1994)
agree that this medium sized aperture is the anterior opening of the sinus canal (Parker 1886) or the orbitotemporal
canal (Rougier et al. 1992; Wible et al. 2004), which in
other placentals transmits the ramus superior of the stapedial artery and accompanying vein (Wible 1984, 1987).
This aperture is called variously the supraorbital foramen
(Gregory 1910:274), the sinus canal foramen (McDowell
1958), the cranio-orbital foramen (MacPhee 1994), and the
anterior opening of the orbitotemporal canal (Wible and
Gaudin 2004). The last term (“otc” in Figs. 14, 16) is preferred, because it is the best descriptor of the position and
best reflects the homology of this structure across a broad
spectrum of cynodonts (Rougier et al. 1992; Wible and
Gaudin 2004). The identification of this foramen in Solenodon is confirmed by establishing continuity between the
foramen and the orbitotemporal groove and canal on the
endocranial surface of AMNH 28271 (“otg” and “otc” in
Fig. 21C). AMNH 28272 reveals that the foramen is not
entirely within the frontal; the orbitosphenoid forms most
of the medial border (Fig. 16). The alisphenoid appears to
contribute but merely overlies the frontal.
(2) Ethmoidal foramina (“2a” and “2p” in Fig. 15).
AMNH 28271 has two foramina that empty endocranially into a depression posterolateral to the cribriform plate
and represent ethmoidal foramina. The larger posterodorsal foramen lays anteroventrally in a common depression
with the orbitotemporal canal opening and is directed anteromedially into the skull, and the smaller anteroventral
foramen is directed posteromedially into the skull and
has a faint groove extending anteroventrally from it. Internally, the posterodorsal foramen opens posteromedial
to the anteroventral foramen (“pef” and “aef” in Fig. 22).
These same two ethmoidal foramina can be identified by
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Fig. 15. Solenodon paradoxus, CM 18069, photograph of right orbit in
oblique lateral view. Abbreviations: 1, anterior opening, orbitotemporal
canal; 2a, anteroventral ethmoidal foramen; 2p, posterodorsal ethmoidal
foramen; 3a, anterior frontal diploic vein foramen; 3p, posterior frontal
diploic vein foramen; 4, supra-optic foramen; asc alisphenoid canal; ch,
choanae; fo, foramen ovale; ham, left pterygoid hamulus; of, optic foramen; sbof, suboptic foramina; sof, sphenorbital fissure; spf, sphenopalatine foramen; tcf, transverse canal foramen.
position and direction in the orbits of the remaining specimens (“aef” and “pef” in Figs. 14, 16, 19). In the juvenile
AMNH 28272 (Figs. 16, 19), the anteroventral foramen is
entirely within the frontal but the posterodorsal one is not;
the orbitosphenoid contributes to the medial wall. AMNH
212912 differs from the other specimens in that the posterodorsal foramen is not in a common depression with
the orbitotemporal canal opening. The dog also has two
ethmoidal foramina with the larger posterodorsal one for
the external ethmoidal artery and companion vein and the
anteroventral one for the ethmoidal nerve (Evans 1993).
Gregory (1910: fig. 18A2; see Fig. 1) identified only
one ethmoidal foramen in AMNH 28272, the anteroventral ethmoidal foramen of this report; he appears to have
labeled this foramen incorrectly as a venous foramen in
another figure of the same specimen (1910: fig. 18A1; see
Fig. 1). Gregory (1910: fig. 18A2; see Fig. 1) labeled the
posteroventral ethmoidal foramen as the supra-optic foramen, a venous foramen. McDowell (1958: fig. 10B) correctly labeled two ethmoidal foramina; the only oddity is
that the groove extending from the anteroventral foramen
is directed anterodorsally, whereas it is anteroventral in the
specimens studied here. MacPhee (1994: figs. 6 [p. 63],
8) identified one ethmoidal foramen in AMMH 35330,
the posterodorsal foramen of this report; however, in another view of the same specimen (fig. 6 [p. 61]) he labeled
a foramen more anteriorly positioned as the ethmoidal
foramen. This foramen likely belongs to the third category
discussed below.
(3) Frontal diploic vein foramen (“3a” and “3p” in Fig.
15). MacPhee (1994: fig. 6) noted a third foramen situated anterolaterally in the same depression as the orbitotemporal canal opening (his cranio-orbital foramen) and
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posterodorsal ethmoidal foramen (his ethmoidal foramen)
in AMNH 35330 and identified it as the frontal diploic foramen. In the dog (Evans 1993), the frontal diploic vein
is an emissary vein of the diploë of the frontal bone to the
dorsal external ophthalmic vein occupying an unnamed
foramen in the orbital surface of the postorbital process of
the frontal (for incidence in other placentals, see Thewissen 1989). MacPhee’s identification is accepted, but there
are two frontal diploic vein foramina directed dorsally into
the substance of the frontal in the specimens studied here,
anterior and posterior. The former is anterolateral to the
anteroventral ethmoidal foramen and was misidentified
as the ethmoidal foramen in AMNH 28272 by Gregory
(1910: fig. 18A1; see Fig. 1); the latter corresponds to the
foramen identified by MacPhee. CM 18069 and AMNH
28271, 28272 (right side), and 212912 have both anterior
and posterior frontal diploic vein foramina (Fig. 15), although the posterior one is tiny in AMNH 28271 and the
anterior one is tiny in AMNH 212912. AMNH 185012 has
only the posterior foramen (“fdv” in Fig. 14), and AMNH
28272 (left side) only the anterior one. It was not possible
to pass a probe through either foramen in the bisected skull
AMNH 28271 in order to trace the endocranial connection; nevertheless, both foramina likely lead dorsally into
a thick column of frontal bone that demarcates the rostral
and middle cranial fossae (see below). In addition to the
anterior and posterior foramina in AMNH 28272 (right
side) are three small, dorsally directed foramina posterior
to the anteroventral ethmoidal foramen that are identified
here as middle frontal diploic vein foramina (Fig. 19); the
left side has two such foramina immediately lateral to the
anteroventral ethmoidal foramen. All of the frontal diploic vein foramina in AMNH 28272 are entirely within the
frontal bone. Gregory (1910: fig. 18A2; see Fig. 1) labeled
one of the middle frontal diploic vein foramina on the left
side of AMNH 28272 as supra-ethmoid foramen, noting
(p. 248) correctly that it “leads dorsad, between the inner
and outer tables of the frontal.”
(4) Supra-optic foramen (“4” in Fig. 15). Gregory (1910:
fig. 18A2; see Fig. 1) labeled a foramen medial to the orbitotemporal canal opening (his sinus canal) in AMNH 28272
as the supra-optic foramen. He stated (p. 248) that it “runs
downward and backward and joins the suboptic” foramen,
which “runs downward and backward to the transverse sinus in the presphenoid.” However, the opening that Gregory labeled in AMNH 28272 is clearly the posterodorsal
ethmoidal foramen of this report (Fig. 16). Nevertheless,
the right side of the bisected skull AMNH 28271 does have
a supra-optic foramen as defined by Gregory. The medium
sized foramen is positioned immediately ventromedial to
the posterodorsal ethmoidal foramen, and a probe passed
through it reaches a transverse sinus in the presphenoid.
The left side of AMNH 28271 has two tiny foramina where
the supra-optic foramen occurs on the right. CM 18069
has a supra-optic foramen on both sides (Fig. 15). AMNH
28272 has a small foramen within the frontal anteromedial to the posterodorsal ethmoidal foramen that likely is
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341
Fig. 16.—Solenodon paradoxus, AMNH 28272, photograph and line drawing of braincase in oblique right lateral view. Palatine is entirely absent from
specimen, whereas premaxilla and most of maxilla and nasal are preserved separately. Scale = 5 mm. Abbreviations: aef, anteroventral ethmoidal foramen; as, alisphenoid; asc, alisphenoid canal; bs, basisphenoid; fr, frontal; gf, glenoid fossa; mx, maxilla (broken); mxfc, facet for maxilla on frontal;
na, nasal (broken); nt, nasoturbinate (ethmoid part); of, optic foramen; oleth, orbital lamina of ethmoid; os orbitosphenoid; otc + fdv, anterior opening,
orbitotemporal canal and posterior frontal diploic vein foramen; pa, parietal; pef, posterior ethmoidal foramen; pet, petrosal; pf, piriform fenestra;
pl, primary lamina (for ethmoturbinate I and II); ppe, perpendicular plate of ethmoid; ps, presphenoid; psw, presphenoid wing (of opposite side); pt,
pterygoid; rm, recessus maxillaris; sof, sphenorbital fissure; sq, squamosal; tcf, transverse canal foramen; tl, transverse lamina (of opposite side); tleth,
tectorial lamina of ethmoid; v, vomer.
a supra-optic foramen (“s-of” in Fig. 19). In light of the
sutural relations and position of the supra-optic foramen
in AMNH 28272, it is doubtful that the supra-optic foramen in AMNH 28271 and CM 18069 are entirely within
the frontal; they are more likely within the orbitosphenoid
or between the orbitosphenoid and frontal. AMNH 185012
and 212912 do not have comparable foramina.
The internal surface of the frontal can be studied in
the bisected skull AMNH 28271 (Figs. 20, 21A, C) and
through the foramen magnum in the juvenile AMNH
28272, with only the latter providing sutural information.
AMNH 28272 (see Fig. 21C) reveals that the frontal has
only a small contribution in the posterolateral roof and
posterodorsal wall of the nasal cavity, with the remaining
posterior roof and wall formed by the underlying ethmoid.
This internal exposure of frontal corresponds to the lateral
half of the triangular maxillary process on the exterior. The
frontals form the roof and lateral walls of the rostral cranial
fossa in AMNH 28272, with the underlying ethmoid completing that space ventrally. The frontals also form the roof
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and dorsolateral walls of the anteriormost middle cranial
fossa, with the overlying parietals forming the roof and
lateral walls posterior to the frontals. The orbitosphenoids
form the ventrolateral walls of the anteriormost middle cranial fossa, with the frontal underlying the orbitosphenoid
throughout most of their suture. A frontal sinus appears to
be lacking.
Along the midline roof in the rostral cranial fossa in
AMNH 28271 and 28272 is a broad crest for the attachment of the falx cerebri, the crista frontalis, which anteriorly turns ventrally into a short crest at the anterodorsal aspect of the cribriform plate. Perpendicular to the
crista frontalis is a more pronounced crest on the frontal
demarcating the rostral and middle cranial fossae, which
presumably lies opposite the circular fissure, which separates the olfactory bulb from the cerebral hemisphere
(“ar” in Fig. 21C). This is the annular ridge of the frontal of Rowe et al. (2005). Occupying the space between
the inner and outer table of the frontal within the annular
ridge is the frontal diploic vein, based on the position and
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direction of the frontal diploic vein foramina in the orbit.
Also, a large canal for the frontal diploic vein is visible just
off the midline in the braincase roof in the bisected skull
AMNH 28271 (see Fig. 21C), suggesting that this vein extends the height of the annular ridge and likely communicates with its fellow of the opposite side. Asymmetrically
arranged foramina on the annular ridge presumably drain
from the endocranium into the frontal diploic vein.
Parietal (“pa” in figures)
The paired parietal forms the bulk of the braincase roof
and lateral walls. AMNH 28272 is the only specimen with
sutures delimiting the entire parietal (Fig. 5); in AMNH
185012 the sutures with the interparietals are obliterating
(“ip” in Fig. 2).
The external surface is described based on AMNH
185012 (Fig. 2); differences from the other specimens are
noted below. At the midline, the parietal comprises more
than 25% of total skull length; at its greatest length, the parietal is 35% of total skull length. At the level of the glenoid
fossa, the parietals comprise more than 66% of skull width.
The parietals meet on the midline and have fairly straight
contacts with their other neighbors, except posteriorly. Anteriorly, the parietal overlaps the frontal at a roughly transverse suture, except for a midline U-shaped incursion of
the frontals (see above). On the lateral border, based on
AMNH 28272, the parietal has a straight suture with the
alisphenoid anteriorly and underlies the squamosal posteriorly (“as” and “sq” in Fig. 16). The anterior half of the
alisphenoid suture is plane and the posterior half is foliate;
the parietosquamosal suture is curved with the parietal the
concave member. Based on AMNH 185012, between the
back of the squamosal and the front of the supraoccipital
(“so” in Fig. 2) is an interval where the parietal forms the
entire braincase roof, its posterolateral edge forming the
nuchal crest (“nc” in Fig. 34) and articulating with the supraoccipital and mastoid exposure of the petrosal (left side
only) on the occiput (“me” in Fig. 34). Posteriorly (Fig. 2),
the parietal overlaps the supraoccipital and the interparietal
at a roughly L-shaped suture.
At the level of the glenoid fossa, the midline parietal
suture is fairly flat in AMNH 185012, but is slightly raised
anteriorly and posteriorly as a weak sagittal crest (“sc” in
Fig. 2). A more pronounced sagittal crest runs the extent
of the midline parietal suture in the adults CM 18069 and
AMNH 28271 and 212912 (Figs. 4, 7). A sagittal crest is
lacking in the juvenile AMNH 28272 (Fig. 5). The parietal in all specimens except AMNH 28272 has a distinct
orbitotemporal crest that curves anterodorsally from the
front of the parietosquamosal suture (“otcr” in Fig. 6). In
the posterior parietosquamosal suture are located several
posterodorsally directed foramina for rami temporales (“rt”
in Figs. 2, 6), which carry blood to and from the temporalis
muscle based on MPIH 6863 and lead into posterodorsally
directed sulci on the parietal. AMNH 28272 has two per
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side in the suture (Fig. 5); AMNH 185012 has one in the
left suture and three in the right (Fig. 2). More of these
foramina are located entirely within the squamosal near to
those in the suture. Consequently, it is not possible to identify the bony relationships of the rami temporales foramina in the adults AMNH 28271, 212912, and CM 18069,
because the parietosquamosal suture is obliterated (Fig.
7). The juvenile AMNH 185012 and the adults also have a
variable number of asymmetrically positioned tiny parietal
foramina, emissary foramina along the midline (Fig. 2).
Nearly the entire endocranial surface of the parietal is
visible through the foramen magnum and piriform fenestra
in AMNH 28272. The parietal is the major element of the
roof and lateral walls of the middle cranial fossa and also
contributes to the roof and walls of the caudal cranial fossa
(see Fig. 21C). In the anterior part of the middle cranial
fossa, the parietal overlaps the frontal in the roof and upper lateral wall and the orbitosphenoid in the lower lateral
wall. Posterior to the orbitosphenoid, the parietal forms the
lateral wall, overlying the alisphenoid, which is confined
largely to the floor. At roughly a right angle to the alisphenoid contact, the parietal contacts the squamosal at a plane
suture (much of this suture is preserved in AMNH 28271);
the squamosal contributes only a narrow strip to the wall
of the middle cranial fossa anterior to the petrosal. Posterodorsal to the squamosal, the parietal overlaps the dorsal
border of the petrosal (“pet” in Fig. 21C), contributing to
the upper lateral wall of the caudal cranial fossa (this suture is preserved in AMNH 28271). The parietal overlaps
the supraoccipital posteroventrally and the interparietal
posterodorsally. Dorsal to and in line with the crista petrosa (“crp” in Fig. 21C) is a low eminence on the parietal
that demarcates the middle and caudal cranial fossae and is
attachment for the tentorium cerebelli, which contains the
transverse sinus, the main distributary of the dorsal sagittal
sinus. This eminence is less distinct than the annular ridge
on the endocranial surface of the frontal in AMNH 28271
and 28272.
The endocranial surface of the parietal has several welldeveloped grooves associated with arteries and veins, described here based on AMNH 28271. Spanning 11 mm in
the wall of the middle cranial fossa is the orbitotemporal
groove for the ramus superior and accompanying veins,
roughly 1 mm in width (“otg” in Fig. 21C). Posteriorly,
the orbitotemporal groove begins at the anterodorsal corner of the petrosal, where based on AMNH 28272 the
squamosal forms a narrow segment of the wall of the initial groove; the remainder of the groove is in the parietal.
The groove arches anteroventrally and disappears at the
posterior edge of the orbitosphenoid. Anterior to this is a
short orbitotemporal canal, at least initially between the
parietal and orbitosphenoid, which leads to the anterior
opening of the orbitotemporal canal between the frontal
and orbitosphenoid in the orbit (Fig. 14). Spanning 5 mm
in the wall of the caudal cranial fossa is the sulcus for the
sigmoid sinus, roughly 2 mm in width (“sss” in Fig. 21C).
For the anterior 3 mm, the sulcus is principally on the
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parietal, with the petrosal forming the ventral quarter; this
relationship is reversed posteriorly. The part of the sulcus
on the parietal is directed posteroventrally; at the posterodorsal corner of the petrosal the sulcus turns ventrally toward the condyloid canals in the exoccipital (“eo” in Fig.
21C). At the bend the supraoccipital contributes to the posteriormost part of the sulcus and the petrosal the remainder. Extending anteriorly from the sulcus for the sigmoid
sinus is a 2 mm long canal between the parietal and petrosal for the prootic sinus (“ecps” in Fig. 21C), the other major distributary of the transverse sinus, that leaves the skull
primarily via the postglenoid foramen as the postglenoid
vein. Anterior to the canal, a sulcus for the prootic sinus in
the squamosal runs anteroventrally toward the postglenoid
foramen (“sps” in Fig. 21C).
Interparietal (“ip” in figures)
As noted by Gregory (1910:242), the juvenile AMNH
28272 “possesses a pair of good sized interparietal bones
separated by a median but slightly asymmetrical suture,
and partly overlaid by the parietals” (Figs. 1, 5). The juvenile AMNH 185012 also preserves the interparietals
as separate elements (Fig. 2), but in the adult specimens
the interparietals are fused with each other and with their
neighbors (Fig. 4).
In dorsal view (Fig. 5), the right interparietal of AMNH
28272 is essentially the shape of a thumbnail, with the
curved part directed anteriorly. The base, posterior side,
has a serrated suture with the supraoccipital, and the right
and anterior sides are overlaid by the right parietal. The
midline interparietal suture, as noted by Gregory, is asymmetrical, with the suture listing to the right anteriorly and
posteriorly but on the midline in between. Considerably
more of the left interparietal is currently exposed along
the supraoccipital margin, but most of the extra exposure
sports a facet for a missing part of the left parietal. Damage to the left parietal postdates Gregory because his illustration (1910: fig. 18A1; see Fig. 1) of the left interparietal resembles the right one. The interparietals in AMNH
185012 are also in the shape of thumbnails (Fig. 2). Sutures with the supraoccipital are present, but those with the
parietals are obliterating, more so along the anterior sides
than the lateral. AMNH 28272 has no sagittal crest along
the midline interparietal suture (Fig. 5), but the remaining
specimens do (Figs. 2, 4, 7). AMNH 28272 and 185012
lack any foramina in the interparietal (Figs. 2, 5), whereas
the adults AMNH 28271 and 212912 have several small,
asymmetrically arranged foramina; CM 18069 has several
foramina in the position where the interparietal-supraoccipital suture was likely located (Fig. 4).
The endocranial surface of the interparietals of AMNH
28272 is visible through the foramen magnum. The interparietals form most of the roof of the caudal cranial
fossa, and the endocranial surface is considerably larger
anteriorly and laterally than the external exposure. Each
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interparietal is roughly quadrangular, but only the medial
side is straight, the others being convexly curved. A distinct depression for the vermis of the cerebellum occupies
the posteromedial half of the right and left interparietals
and continues posteriorly onto the supraoccipital. This depression is much less distinct in the adult AMNH 28271.
Pterygoid (“pt” in figures)
The paired pterygoid is not fully delimited as a separate
element from the basi- and alisphenoid in any specimen.
The juvenile AMNH 28272 and to a lesser extent AMNH
185012 have seams that are interpreted as marking the line
of fusion with the basi- and alisphenoid along the anteromedial, medial, posterior, and posterolateral borders of the
pterygoid (“>” in Fig. 19). However, the anterolateral borders with the alisphenoid are not indicated. The pterygoid
is described here based on these seams. The serially sectioned juvenile MPIH 6863 has sections preserving only
the posterior part of the pterygoid; this bone is for the most
part independent of the overlying basi- and alisphenoids,
but some sections do show fusion.
The pterygoid is a long, narrow element lying ventral to
the sphenoid complex, separated by the pre- and basisphenoid from the pterygoid of the opposite side (Fig. 19). Posteriorly, the main axis of the pterygoid is in the same plane
as the skull base; anteriorly the main axis is perpendicular
to the skull base, forming the parasagittal posterolateral wall
of the basipharyngeal canal behind the palatine. The transition between these two parts is gradual rather than abrupt.
The horizontal part of the pterygoid is the caudal process,
and the perpendicular part is the entopterygoid process, at
the ventral extent of which is the pterygoid hamulus (“cpp”,
“enp”, and “ham” in Fig. 19).
The anterior face of the entopterygoid process contacts
the back of the perpendicular process of the palatine and a
suture indicating this is preserved both medially within the
basipharyngeal canal and laterally within the orbital floor
only in AMNH 185012 (Figs. 11, 14). Only part of the medial pterygopalatine suture is preserved in AMNH 212912
and CM 18069, and the suture is obliterated in AMNH
28271. The anteromedial base of the entopterygoid process
is separated from the overlying presphenoid by a suture in
the juveniles AMNH 28272 and 185012 (Figs. 11, 19), but
these are fused in the adults. The entopterygoid process approximates but does not contact the vomer in AMNH 28272
(Fig. 19), whereas sutural contact of these elements is preserved in AMNH 185012 (Fig. 11), 212912, and CM 18069.
The entopterygoid process of the juvenile AMNH
28272 differs from the remaining specimens in that it projects only slightly more ventrally than the caudal process
(Fig. 19); that is, there is not as great a height differential
between the two as occurs in the older skulls (Figs. 6, 7,
14). In addition to being relatively shorter, the entopterygoid process of AMNH 28272 is mediolaterally thicker
except at its ventral margin (Fig. 19) and not the laminar
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Fig. 17.—Solenodon paradoxus, AMNH 28272, photograph and line drawing of braincase (nasal cavity) in anterior view. Both sides have been damaged
with most structures more complete on the specimen’s right side, ethmoturbinal I being an exception. Damage to the septum frontomaxillare on the left
side exposes the recessus frontalis. Abbreviations: ec 1, ectoturbinal 1; ec 2, ectoturbinal 2; et I, ethmoturbinal I; et II, ethmoturbinal II; et III, ethmoturbinal III; fnf, fundus of nasal fossa; nt, nasoturbinate (ethmoid part); pl, primary lamina of ethmoturbinal I and II; ppe, perpendicular plate of ethmoid;
rf, recessus frontalis; rm, recessus maxillaris; sfm, septum frontomaxillare; tl, transverse lamina; v, vomer (broken).
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plate that occurs in the older skulls (Figs. 11, 12). The two
millimeter long ventral edge of the entopterygoid process
in AMNH 28272 is the pterygoid hamulus. This edge is
uniformly thin mediolaterally on the left side, but twice as
wide anteriorly as posteriorly on the right side (Fig. 19).
The pterygoid hamulus is damaged bilaterally in AMNH
185012 and on the right side of CM 18069 (Fig. 12). In
AMNH 212912 (Fig. 12), the hamulus is oriented obliquely anterolaterally, with a thickened ventral border some
three millimeters in length; on the right side of CM 18069,
the thickened part of the ventral edge is half as long and
straight (Fig. 12); and in AMNH 28271, the ventral border
is not thickened at all, but has a slight medial inflection at
its posterior edge.
Posterior to the level of the hamulus, the caudal process begins the gradual transition from a near vertical element to a near horizontal one. In the seam between the
medial edge of the caudal process and the basisphenoid in
AMNH 28272 are two foramina: an anterior one slightly
posterior to the level of the foramen ovale (“fo” in Fig. 19)
and a posterior one medial to the posterior opening of the
pterygoid canal (vidian canal) (“pc” in Fig. 19). The older
specimens have a variable number (two to six) of similarly
situated foramina. In AMNH 28272 these foramina can be
traced into the basisphenoid’s transverse canal (“tsc” in
“bs” in Fig. 21C), and veins occur in similar foramina in
the sectioned juvenile MPIH 6863.
The posterior border of the caudal process in AMNH
28272 nearly reaches as far posterolaterally as does the
overlying basisphenoid, falling just short of contacting
the anterior pole of the promontorium and contributing to
the border of the piriform fenestra (“pr” and “pf” in Fig.
19). In the posterolateral margin of the caudal process is
the entrance into the pterygoid canal (Fig. 19): the caudal
process forming the floor and the basisphenoid the roof
as in the sectioned juvenile MPIH 6863. Anterior to the
pterygoid canal opening, a faint seam delimits the caudal
process from the overlying tympanic process of the alisphenoid (“tpas” in Fig. 19), but anterior to the level of the
foramen ovale, the pterygoid and alisphenoid are seamlessly fused (not visible in Fig. 19).
Ethmoid (“eth” in figures)
The unpaired ethmoid forms the back of the nasal cavity
and the front of the rostral cranial fossa. With the exception of a small triangular exposure in the orbit, based on
AMNH 28272 (Fig. 16), the ethmoid is hidden in external
view. Aspects of the ethmoid can be studied in the bisected
adult and partially disarticulated juvenile skulls, AMNH
28271 and 28272. The ethmoid consists of four parts: a
median perpendicular plate or lamina, a cribriform plate,
and two lateral masses covered by the external lamina of
the ethmoid; the four ethmoturbinates, two ectoturbinates,
and the ethmoid part of the nasoturbinate are attached to
the lateral masses and/or cribriform plate.
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The major elements of the nasal septum are the perpendicular plate of the ethmoid behind and the cartilaginous nasal septum in front. The proportions of these two
elements differ dramatically in the two specimens. In the
younger AMNH 28272, the perpendicular plate reaches
forward to the level of the maxillary process of the frontal
opposite dP5 (“ppe” in Figs. 16, 18). In contrast, in AMNH
28271 more of the cartilaginous nasal septum is ossified
(and sutures do not delimit the contributing elements),
presumably with the ethmoid reaching I2 (Fig. 20). In the
former, AMNH 28272 (Fig. 16), the perpendicular plate
stretches between the posterior nasal cavity roof, which is
the external laminae of the ethmoid (the tectorial lamina or
tectum nasi), and the posterior sagittal part of the vomer
(Fig. 17). Based on AMNH 28271, which lacks sutural information, the perpendicular plate also likely contacts the
nasals dorsally and the anterior sagittal part of the vomer
and the septal processes of the premaxillae ventrally (Fig.
20).
The cribriform plate is a deeply concave, sieve-like partition between the posterior nasal cavity and rostral cranial
fossa (“cpl” in Figs. 22B, C, 23). It is angled such that its
dorsal margin is anterior to its ventral margin. Approximately 300 variably sized foramina perforate the plate
on the right and left sides in AMNH 28271 (Fig. 23) and
transmit olfactory nerve bundles. A broad central ridge
(“cr” in Fig. 23) separates the right and left ethmoidal fossae, which housed the olfactory bulbs in life. The ridge is
pierced by numerous foramina but has no crista galli in all
specimens exposing this surface (CM 18069 is unknown
because dried meninges cover the plate). Within each ethmoidal fossa, the cribriform foramina are arranged in cauliflower-like clusters, which surround the attachment of the
ethmoturbinals (see below) as reported in other placentals,
e.g., dog (Evans 1993), Monodelphis Burnett, 1830 (Rowe
et al. 2005), and Pteropus (Giannini et al. 2006). AMNH
28272 is the only specimen to preserve sutures delimiting the ethmoid in the rostral cranial fossa, but the view
through the foramen magnum does not provide access to
the entire circumference of the bone. The dorsal and lateral
sutures with the frontal follow closely the outer margin of
the cribriform plate (see Fig. 21C). Regarding the ventral
suture, only the midline area is visible. It shows a broad,
non-cribriform eminence of the ethmoid in contact with
the presphenoid. The posterior face of this eminence faces
the middle cranial fossa. In AMNH 28271, a shallow, horizontal sulcus (“hs” in Fig. 23) near the outer margin of the
cribriform plate demarcates two major clusters of foramina, a larger anterodorsal and smaller posteroventral. In the
anterodorsal cluster are the foramina for the nasoturbinate and the two ectoturbinals (“nt”, “ec 1”, and “ec
2” in Fig. 23) as well as the cribroethmoidal foramen for
the passage of the ethmoidal nerves (“cef” in Fig. 23); in
the posteroventral cluster are the foramina for ethmoturbinal II, III, and IV (“et II”, “et III’, and “et IV” in Fig.
23); and sandwiched in between the two clusters are foramina for ethmoturbinal I (“et I” in Fig. 23). At the outer
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Fig. 18.—Solenodon paradoxus, AMNH 28272 photograph and line drawing of braincase in ventral view. Palatine is absent from specimen and rostrum is preserved separately. Nasal cavity is damaged on both sides, but more so on the specimen’s left side (see Fig. 17). Abbreviations: as, alisphenoid; bo, basioccipital; bs, basisphenoid; e, ectotympanic, ec 1, ectoturbinal 1; ec 2, ectoturbinal 2; egp, entoglenoid process; eo, exoccipital; eth, ethmoid; fr, frontal; gri, groove
for ramus inferior; na, nasal (broken); nt, nasoturbinate (ethmoid part); oc, occipital condyle; oleth, orbital lamina of ethmoid; os, orbitosphenoid; pa, parietal;
pf, piriform fenestra; pl, primary lamina of ethmoturbinal I and II; ppe, perpendicular plate of ethmoid; pr, promontorium of petrosal; ps, presphenoid; pt,
pterygoid; rm, recessus maxillaris; so, supraoccipital; sq, squamosal; tl, transverse lamina; tpas, tympanic process of alisphenoid; v, vomer (broken).
margin of the cribriform plate, the horizontal sulcus turns
posteroventromedially for 5 mm and ends at two foramina posterolateral to the cribriform plate (“aef” and “pef”
in Fig. 22). The smaller anterolateral one connects with
the anteroventral ethmoidal foramen in the orbit, and the
larger posteromedial one with the posterodorsal ethmoidal
foramen. Unfortunately, the positional relationship of these
endocranial foramina to the frontoethmoidal or sphenoethmoidal suture cannot be ascertained.
As viewed in AMNH 28272, the outer covering of the
lateral masses is the external lamina, which includes a roof
or tectorial lamina (“tleth” in Fig. 16), lateral wall or orbital lamina (“oleth” in Fig. 16), and a floor or basal lamina,
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which is fused to the vomerine wing to form the transverse
lamina (“tl” in Fig. 16). Dorsal to the tectorial lamina are
the posterior parts of the nasal and maxilla and the maxillary processes of the frontals and ventral to the roof is
the perpendicular plate. Most of the orbital lamina is covered by the frontal with a narrow strip along the anterior
margin covered by the maxilla. However, there is a triangular
exposure of the orbital lamina in the orbital wall between
the sphenopalatine and optic foramina in AMNH 28272
(Figs. 1, 16, 19). The short, vertical anterior side of the
triangle is covered by the maxilla; the long, horizontal ventral side is covered by the palatine; and the long, inclined
dorsal side is covered by the frontal; the posterior tip of the
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Fig. 19.—Solenodon paradoxus, AMNH 28272, photograph of braincase in oblique right ventral view. Carat points outline the suture separating the right
pterygoid and presphenoid and the seams separating the right pterygoid from the basisphenoid and alisphenoid. Scale = 5 mm. Abbreviations: acc, alicochlear commissure; aef, anteroventral ethmoidal foramen; as, alisphenoid; asc, alisphenoid canal; bs, basisphenoid; cpc, craniopharyngeal canal; cpp,
caudal process of pterygoid; enp, entopterygoid process; ew, epitympanic wing of petrosal; fr, frontal; gf, glenoid fossa; ham, pterygoid hamulus; mxfc,
facet for maxilla on frontal; of, optic foramen; oleth, orbital lamina of ethmoid; orf, oribital fissure; os, orbitosphenoid; otc + fdv, anterior opening, orbitotemporal canal and posterior frontal diploic vein foramen; pa, parietal; pc, posterior opening, pterygoid canal; pef, posterodorsal ethmoidal foramen;
pf, piriform fenestra; ps, presphenoid; psw, presphenoid wing; pt, pterygoid; rtp, rostral tympanic process of petrosal; sbof, suboptic foramen; s-of,
supraoptic foramen; sq, squamosal; tcf, transverse canal foramen; tl, transverse lamina; tpas, tympanic process of alisphenoid; v, vomer (broken).
triangle has a plane contact with the orbitosphenoid (Fig.
1). Immediately ventromedial to the orbital exposure of the
ethmoid is the transverse lamina (Fig. 16), which is concealed in external view by the missing palatine.
In anterior view, two distinct spaces are visible on the
right ethmoid of AMNH 28272; the larger medial space is
the fundus of the nasal fossa or olfactory recess (“fnf” in
Fig. 17), which contains the ethmoturbinals and ends posteriorly at the cribriform plate, and the smaller, crescentic
lateral space is a dead end diverticulum off the main nasal
passage (“rm” in Fig. 17). The latter space is part of the
recessus lateralis of the pars intermedia of the nasal capsule, which lies between the pars anterior and pars posterior
(Smith and Rossie 2006); the pars posterior includes the olfactory recess and the pars anterior lies between the external
nasal aperture and olfactory recess. The recessus lateralis is
often subdivided in therians by a septum frontomaxillare
into a recessus maxillaris ventrally and a recessus frontalis dorsally (Smith and Rossie 2006). The crescentic lateral
space in AMNH 28272 is the recessus maxillaris (maxillary recess). The ethmoid forms most of the boundaries of
the recessus maxillaris, with the maxilla contributing the
anterior part of the lateral wall (“rm” in Fig. 9). The medial wall of the recessus maxillaris has several longitudinal
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depressions and rounded prominences (Fig. 16), the former
marking the attachments of ethmoturbinals and ectoturbinals. The largest of these prominences is centrally located
and is delimited by the attachment of ectoturbinal 2 above
and the primary lamina of ethmoturbinal I and II below
(“pl” in Fig. 17). At the anterior end of this prominence is
a distinct, curved, lip-shaped thickening that connects the
anterior base of ectoturbinal 1 to the primary lamina (Fig.
17).
Part of the dorsal wall between the recessus maxillaris
and the fundus of the nasal fossa is damaged on the left
side, revealing more of a third space between the ethmoid
part of the nasoturbinate and the recessus maxillaris; this
is the recessus frontalis (“rf” in Fig. 17). Consequently, the
wall between the recessus maxillaris and recessus frontalis is the septum frontomaxillaris (“sfm” in Fig. 17). The
recessus frontalis houses the two ectoturbinals (Fig. 17).
In AMNH 28272, the base of the ectoturbinal 1 is thick,
extends medially and slightly ventrally, and thins to a
scroll directed dorsally and slightly laterally; the base of
the ectoturbinal 2 is thin, thickens medially, and then thins
to dorsolateral and ventrolateral scrolls. Two ectoturbinals
are present in AMNH 28271 (although not in the figures),
but the base of ectoturbinal 1 is thin, in contrast to the
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Fig. 20.—Solenodon paradoxus, AMNH 28271, photographs of left and right sides of bisected skull in medial views. Bisection occurred off the midline
such that the right side includes the osseous nasal septum; the cartilaginous nasal septum above the premaxilla is not preserved. Posterior to the osseous
nasal septum the posterior part of ethmoturbinal III and all of ethmoturbinal IV are visible. The left side is broken in two pieces; an oblique seam is
visible at the break directed anterodorsally from just posterior to the penultimate premolar. Scale = 10 mm.
condition in AMNH 28272.
The fundus of the nasal fossa contains the ethmoid part
of the nasoturbinate and the four ethmoturbinals (Fig. 17).
In identifying ethmoturbinals, followed here is the terminology of Smith and Rossie (2006), which includes a useful table of synonyms with authors such as Moore (1981)
and Maier (1993).
The ethmoid part of the nasoturbinate (endoturbinate I
of Moore 1981) is completely preserved on the left side of
AMNH 28271 (Fig. 21B). It arises from the tectorial lamina and is continuous anteriorly with the nasoturbinate crest
on the nasal. It is a simple ridge, low anteriorly and posteriorly, and with a triangular prominence opposite P5 and
M1 that is partially hidden by ethmoturbinal I in medial
view. The ethmoid part of the nasoturbinate extends to the
posterior wall of the nasal fundus. The triangular prominence of the ethmoid part of the nasoturbinate is preserved
on the left side of AMNH 28272 (Fig. 17). The triangle’s
anterior side is damaged or incompletely ossified, and the
posterior side is slightly thickened. The triangle extends
roughly a third of the way down the nasal fundus, medial
to ectoturbinal 1.
Ethmoturbinal I and II are nearly completely preserved
on the left side of AMNH 28271 (Fig. 21B). They arise
from a common base, the primary lamina, opposite M2,
that is very thick, directly slightly dorsomedially from
the horizontal, and divides into two scrolls, both of which
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extend to the posterior wall of the nasal fundus. The longer
dorsal scroll, ethmoturbinal I, is most prominent opposite
P5 and M2 and the ventral scroll, ethmoturbinal II, opposite M2 and M3. Both scrolls are relatively simple and
concave ventromedially. The base of the ethmoturbinal I
is thickened and divides the scroll into anterior and posterior halves, with the tip of the anterior half overlying the
ventral nasal concha or maxilloturbinate and the posterior
half overlying ethmoturbinal II. Ethmoturbinal I and II are
preserved incompletely and completely, respectively, on
both sides of AMNH 28272 (Fig. 17).
Ethmoturbinal III is preserved bilaterally in AMNH
28271, but only the anterior part is visible on the left side
(Fig. 21B) and the posterior part on the right (Fig. 20). It
arises from the orbital lamina slightly posterior and ventral
to the base of ethmoturbinal II, opposite the back of M2.
The ventral prominence of its scroll extends from the level
of M2 to the minor palatine foramen and it contacts the
posterior wall of the nasal fundus behind the level of that
foramen. Ethmoturbinal III is also preserved bilaterally in
AMNH 28272, but only its anteriormost aspect is visible
(Fig. 17).
Ethmoturbinal IV can only be seen in AMNH 28271.
On the right side (Fig. 20), the ventral prominence of its
scroll extends from the level of the sphenopalatine to the
optic foramen. Its posterodorsal border contacts the posterior wall of the nasal fundus. The narrow space posterior to
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Fig. 21.—Solenodon paradoxus, AMNH 28271, left side of bisected skull in medial view. A, shaded drawing; B, line drawing of nasal cavity; C, line drawing of cranial cavity. Note specimen is broken in nasal cavity posterior to P4. Dashed lines in cranial cavity are sutures based on AMNH 28272; numbers
1 and 2 refer to foramina that transmit the postglenoid vein to the postglenoid foramen on the skull base. Abbreviations: ar, annular ridge; as, alisphenoid;
av, aqueductus vestibuli; bs, basisphenoid; C, upper canine; ccp, conchal crest of premaxilla; ch, choanae; coca, condyloid canals; cpl, cribriform plate;
crp, crista petrosa; ecps, endocranial canal for prootic sinus; eff, ethmoidal foramina; enpc, entopterygoid crest; eo, exoccipital; et I, ethmoturbinal I; et II,
ethmoturbinal II; et III, ethmoturbinal III; et IV + sphf, membrane covering space for ethmoturbinal IV and sphenoidal fossa; eth, ethmoid; fac facial canal;
fai, foramen acusticum inferius; fdv, frontal diploic vein; fo, foramen ovale; fr, frontal; ham, pterygoid hamulus; hf, hypoglossal foramen; I2, upper second
incisor; js jugum sphenoidale; M1, upper first molar; mt, maxilloturbinate; nt, nasoturbinate; oev, occipital emissary vein; op, os proboscidis; os, orbitosphenoid; otc, orbitotemporal canal; otg, orbitotemporal groove; P4, upper penultimate premolar; pa, parietal; pet, petrosal; ps, presphenoid; rt, foramen
for ramus temporalis; saf, subarcuate fossa; so, supraoccipital; sof, sphenorbital fissure; spf, sphenopalatine foramen; spp, septal process of premaxilla; sps,
sulcus for prootic sinus; sq, squamosal; sss, sulcus for sigmoid sinus; sva, superior vestibular area; tl, transverse lamina; tsc, transverse sinus canal.
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Fig. 22.—Solenodon paradoxus, AMNH 28271, photograph of posterior part of left half of bisected skull in oblique medial view. Scale = 5 mm. Abbreviations: ace, anterior crus of ectotympanic; aef, anteroventral ethmoidal foramen; apf, accessory palatine foramen; ar, annular ridge; asc, alisphenoid canal;
av, aqueductus vestibuli; cpl, cribriform plate; et IV + sphf, membrane covering space for ethmoturbinal IV and sphenoidal fossa; fo, foramen ovale; fpgv,
endocranial foramen for postglenoid vein; gica, faint endocranial groove for internal carotid artery; hyf, hypophyseal fossa; js, jugum sphenoidale; mapf,
major palatine foramen; of, optic foramen; pf, piriform fenestra; saf, subarcuate fossa; sof, sphenorbital fissure; spf, sphenopalatine foramen.
ethmoturbinal IV is the sphenoidal fossa. On the left side it
is hidden by a connective tissue septum separating the left
and right sides (“et IV + sphf” in Fig. 21B).
Vomer (“v” in figures)
The unpaired vomer extends nearly the length of the nasal
cavity and through the roof of the choanae into the anterior
part of the basipharyngeal canal. The vomer is described
(e.g., Evans 1993) as having sagittal and horizontal parts,
with the latter at right angles to the former. In the juvenile
AMNH 28272, the sagittal part is delimited from its neighbors by sutures anteriorly and posteriorly (“llv” in Fig. 9),
but the horizontal part is fused to the ethmoid to form the
transverse lamina separating the nasal fundus from the
nasopharyngeal meatus (“tl” in Figs. 16, 18). In AMNH
185012, 212912, and CM 18069 the posterior sagittal part
in the basipharyngeal canal is delimited (Figs. 11, 12).
AMNH 28272 preserves the vomer in three broken
pieces. Attached to the right and left premaxillae and maxillae are two pieces 10 mm in length (between C and dP5)
of the anterior sagittal part of the vomer (Fig. 9). These
two pieces represent the right and left lateral laminae of the
vomer, which when rearticulated ventrally at their base on
the midline between them form the V-shaped sulcus septi
nasi for the perpendicular plate of the ethmoid. The notch
between the anterior tips of the two laminae is the incisive
incisure.
The third piece of vomer is attached to the floors of the
posterior nasal cavity and anterior braincase and includes
the posterior sagittal and horizontal parts. The front of the
posterior sagittal part is broken and at least a 6 mm section in front of the break is missing beneath the perpendicular plate of the ethmoid (Fig. 18); in addition there are
longitudinal cracks and a narrow wedge of bone missing
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posteriorly along the midline (Fig. 19). In ventral view
(Fig. 18), the posterior sagittal part has a midline rodshaped eminence with narrow lateral wings that posteriorly in the basipharyngeal canal would have contacted the
perpendicular plates of the palatines (see Fig. 11). Posteriorly, the posterior sagittal part extends just over a millimeter ventral to the presphenoid and sends narrow, sharp
lateral processes off more posteriorly, with the notch between those processes being the sphenoidal incisure (Fig.
18). The lateral laminae of the posterior sagittal part are
visible in anterior view (Fig. 17); the sulcus septi nasi between them is distinctly U-shaped.
Extending laterally from all but the posterior five mm
of the posterior sagittal part are the vomerine wings or
alae, the horizontal part. It is only near the midline that the
wings are horizontal, because the bulk of the wing angles
ventrolaterally at roughly 30°. The vomerine wings are
fused seamlessly with the external laminae of the ethmoid
forming the transverse lamina. Roux (1947) reported the
same condition in the adult shrew Suncus orangiae (Roberts, 1924) and in embryos the vomer was confined to the
floor of the posterior nasal cavity. In light of this and the
general morphology of the vomer in other placentals (e.g.,
dog, Evans 1993; horse, Sisson 1910), the vomerine wings
in AMNH 28272 are reconstructed in the floor of the posterior nasal cavity and the ethmoid in the lateral wall of
the nasal cavity, part of which is exposed externally in the
orbit (also suggested by Saban 1956:219). In contrast, in
Gregory’s (1910) illustration of AMNH 28272 (Fig. 1), he
identified the orbital exposure of the lateral nasal cavity
wall as the vomerine wing.
In the three intact skulls, CM 18069 and AMNH
185012 and 212912, the posterior sagittal part of the
vomer exposed in the basipharyngeal canal is variable. It
is flat in CM 18069, with a rounded eminence that has a
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short, sharp, low crest anteriorly just behind the choanae in
AMNH 185012, and with a low, narrow, rounded crest in
AMNH 212912 (Figs. 11, 12).
Sphenoid Complex
The sphenoid complex contributes to the roof of the mesocranium, the walls of the orbitotemporal region, and to
the floor of the middle cranial fossa. In AMNH 28272, the
sphenoid is separated into anterior and posterior parts by a
transverse midline gap at a level just anterior to the glenoid
fossa (Figs. 18, 19). Each part of the sphenoid complex
includes an unpaired element in the mesocranial roof and
a pair of wings in the orbitotemporal walls: unpaired presphenoid and paired orbitosphenoid for the anterior part,
and unpaired basisphenoid and paired alisphenoid for the
posterior part. It is unknown how many ossification centers contribute to the sphenoid complex, and the number
varies in different mammals (De Beer 1937; Moore 1981).
The serially sectioned juvenile MPIH 6863 preserves only
the caudal end of the posterior part, and its basisphenoid
and paired alisphenoid are a single ossification as in all the
skulls. AMNH 28272 preserves a short midline suture on
the endocranial surface of the anteriormost presphenoid,
which is suggestive that the anterior part of the sphenoid
complex is composed of at least two ossifications. Early
developmental stages are needed to fully address whether
there is an additional midline ossification, for example, as
in Monodelphis (Clark and Smith 1993).
The paired pterygoid bone described above underlies
the anterior part of the sphenoid complex and is fused to
the posterior part. The sphenoid complex is only fully delimited from its neighbors in AMNH 28272 (Figs. 16, 18,
19), with AMNH 185012 preserving all sutures except part
of that between the orbitosphenoid and ethmoid (Figs. 11,
14). These two specimens provide the bases for the descriptions of the individual sphenoid elements below.
Presphenoid (“ps” in figures)
In ventral view in AMNH 28272 (Fig. 18), the presphenoid
is roughly trapezoidal, longer than wide, and wider anteriorly than posteriorly, but this entire surface is not exposed
on the skull base in the other specimens. The ventral surface of the presphenoid in AMNH 28272 has a paired, low
longitudinal ridge off the midline that represents the point
of abutment of the (missing) palatines anteriorly and the
pterygoids posteriorly. The presphenoid between these two
faint ridges is the part exposed in the roof of the basipharyngeal canal. Anteriorly, the presphenoid is underlain by
the vomer; posteriorly, it is separated from the basisphenoid
(“bs” in Fig. 19) by a gap, the remnant of a synchondrosis.
Along the anterior two-thirds of the presphenoid’s lateral
edge is a ventrally projecting crest that is entirely hidden
by the palatine and pterygoid bones in the other skulls; this
crest is termed the presphenoid wing (“psw” in Fig. 19).
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Fig. 23.—Solenodon paradoxus, AMNH 28271, photograph of left cribriform plate in posterior view. Scale = 3 mm. Abbreviations: ar, annular
ridge; cef, cribroethmoidal foramen; cr, central ridge; ec 1, foramina to
ectoturbinal 1; ec 2, foramina to ectoturbinal 2; et I, foramina to ethmoturbinal I; et II, foramina to ethmoturbinal II; et III, foramina to ethmoturbinal III; et IV, foramina to ethmoturbinal IV; hs, horizontal sulcus; js,
jugum sphenoidale; nt, foramina to nasoturbinate; of, optic foramen; sof,
sphenorbital fissure.
This crest reaches its maximum height anteriorly where
it has a plane suture with the transverse lamina as well as
its maximum separation from its fellow of the opposite
side. The lateral surface of this crest is exposed in the orbit as the part of the orbitosphenoid ventral to the level of
the optic foramen (“of” in Fig. 19). The presphenoid in
AMNH 185012 is delimited from the vomer, palatines, and
pterygoids, and there is a faint trace of the presphenoidbasisphenoid (intersphenoidal) synchondrosis (Fig. 11). In
AMNH 28271, 212912, and CM 18069, only part of the
suture with the vomer is preserved (Fig. 12).
The endocranial surface of the presphenoid is visible
through the foramen magnum in AMNH 28272. Its surface
is gently rounded posteriorly, but anterior to the level of
the optic foramina the jugum sphenoidale (yoke) is flattened and angled anterodorsally (“js” in Figs. 21C, 23). A
midline suture extends posteriorly from the ethmoid suture
halfway to the level of the optic foramina. The low posterolateral wall of the presphenoid’s endocranial surface
is also visible through the sphenorbital fissure and is in
contact with the alisphenoid. In the bisected skull, AMNH
28271, there is a large transverse canal that crosses the
midline of the presphenoid ventral to the optic foramina
(“tsc” in “ps” in Fig. 21C).
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Orbitosphenoid (“os” in figures)
The orbital surface of the orbitosphenoid in AMNH 28272
can be divided into two parts: ventral and dorsal. The ventral part is the main orbital exposure; the dorsal part is
largely covered by the alisphenoid (Figs. 16, 19).
The ventral part is roughly rectangular, longer than
high (Figs. 16, 19). On its anterior border, it overlaps the
ethmoid ventrally and the frontal dorsally. On its ventral
border, it has plane sutures with the (missing) palatine anteriorly and the pterygoid posteriorly. Its posterior border
is hidden in lateral view by the alisphenoid; it forms the
medial wall of the sphenorbital fissure (“sof” in Figs. 16,
19), which is closed laterally by the alisphenoid, and tapers
ventrally into the presphenoid wing. On its dorsal border,
it has a plane suture with the frontal anteriorly and is overlapped by the alisphenoid posteriorly; between these two
bones is the narrow dorsal part of the orbitosphenoid (see
below). AMNH 185012 preserves the sutures on the ventral border with the palatine and pterygoid, and parts of the
sutures on the anterior and posterodorsal borders with the
fused ethmoid + frontal and alisphenoid, respectively (Fig.
14).
Centrally located in the ventral part is the anteriorly directed optic foramen, roughly half a millimeter in diameter
(Figs. 16, 19). Extending anteriorly from the optic foramen
to the border with the frontal is a groove of similar dimensions. Three specimens show left-right asymmetry in the
optic foramen. In AMNH 185012, the right foramen is half
the diameter of the left and also placed more ventrally in
the orbitosphenoid; in AMNH 212912, the left foramen
opens at a level about two millimeters anterodorsal to the
right; and in CM 18069, the right foramen opens about a
millimeter anterodorsal to the left. Penetrating the orbitosphenoid anteroventral and posteroventral to the optic foramen in AMNH 28272 are small foramina (six on the left
and four on the right) that resemble the suboptic foramen
identified by Gregory (1910) (“subof” in Fig. 16) as lying just below the optic foramen with its canal running posteroventrally to join the transverse sinus in the presphenoid.
This is confirmable in AMNH 28271, where the single
suboptic foramen on the right and double on the left join
the transverse sinus canal. There is considerable variation
within and between specimens in the number of suboptic
foramina.
The dorsal part of the orbitosphenoid in AMNH 28272
is mainly hidden in direct lateral view, but in oblique ventrolateral view is roughly Y-shaped (Figs. 16, 19). The base
is between the frontal anteriorly and alisphenoid posteriorly. The anterior arm contributes to the medial border of the
posterodorsal ethmoidal foramen, with the frontal closing
the foramen. The posterior arm contributes to the medial
border of the anterior opening of the orbitotemporal canal,
with the frontal closing the foramen.
Much of the endocranial surface of the orbitosphenoid
is visible through the foramen magnum in AMNH 28272
(see Fig. 21C). It is a broad wing that climbs roughly
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halfway up the lateral wall of the anterior part of the middle cranial fossa. The anterior border of the orbitosphenoid
is a curved edge that delimits the rostral and middle cranial
fossae ventrolaterally. The dorsal border of the orbitosphenoid is pyramidal with the anterior edge having a plane
suture with the frontal and the posterior edge overlapped
by the parietal. The posterior border is straight and overlapped by the alisphenoid laterally and has a free concave
edge medially, which forms the medial wall of the sphenorbital fissure. Near the juncture of the posterior border
with the presphenoid is the small optic foramen, and near
the posterior extent of the suture with the parietal, the orbitotemporal canal is formed between the orbitosphenoid
and parietal. On the left side of the bisected skull AMNH
28271 posterior to the optic foramen is a small foramen
that opens into the transverse canal.
Basisphenoid (“bs” in figures)
In ventral view in AMNH 28272 (Figs. 18, 19), the basisphenoid is roughly trapezoidal. It has a narrow, straight
anterior side separated from the presphenoid by a gap,
oblique lateral sides that overlie the pterygoid and continue into the orbitotemporal fossa as the alisphenoid, and a
wide, straight posterior side that has a plane suture with the
basioccipital (“bo” in Figs. 18, 19). The central part of the
basisphenoid is essentially flat; the posterolateral corners
opposite the anterior pole of the petrosal promontoria are
bent slightly dorsally, contributing to the medial border of
the piriform fenestrae. AMNH 28272 and 185012 have a
small midline foramen positioned just rostral to the center
of the bone that likely is the craniopharyngeal canal (“cpc”
in Figs. 18, 19, 25, 27A) and the remaining specimens a
tiny slit.
There is variability among the studied skulls in the part
of the basisphenoid in the piriform fenestra, variability
both between and among the represented developmental
stages. In the juvenile AMNH 28272, the basisphenoid has
two tiny prongs of bone that cup but do not surround the internal carotid artery at its endocranial entrance. The lateral
of these two prongs is visible in ventral view (“acc” in Fig.
19), but the medial is more dorsally positioned. Because
the lateral prong lies lateral to the internal carotid artery it
represents an ossification of the alicochlear commissure,
the cartilaginous bar of the chondrocranium that connects
the processus alaris of the ala temporalis with the front of
the auditory capsule (De Beer 1937). As figured by McDowell (1958: fig. 7B), the internal carotid artery in the
juvenile solenodon enters the cranial cavity via the piriform fenestra. The right side of the other juvenile, AMNH
185012, resembles the condition in AMNH 28272, but on
the left side the lateral of the two prongs has expanded
dorsal to the artery and contacts the medial prong to enclose a carotid foramen within the basisphenoid (also on
right side of CM 18069); the lateral prong approximates the
petrosal and excludes the artery from the piriform fenestra
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(“cf” in Fig. 25). A similar arrangement occurs bilaterally
in AMNH 212912 except that a small gap remains between the medial and lateral prongs. In contrast, prongs
are lacking entirely in AMNH 28271 and the left side of
CM 18069.
The dorsal surface of the basisphenoid is accessible in
the bisected skull AMNH 28271 (Figs. 20, 21A, B) and
through the foramen magnum in AMNH 212912. The basisphenoid is not delimited from either the presphenoid in
front or the basioccipital behind, and the bony limits are
inferred from the juveniles AMNH 28272 and 185012.
Anteriorly, the basisphenoid is gently rounded as was the
presphenoid, although the basisphenoid is slightly wider.
Posteriorly, there is a shallow hypophyseal fossa, three
millimeters in length in AMNH 28271, occupying the
space between the paired pars cochlearis of the petrosal
bone (“hyf” in Fig. 21C). The anterior and posterior walls
of the fossa are only slightly raised in AMNH 28271 (see
right side in Fig. 20) and 212912. From its position, the
hypophyseal fossa cannot be entirely on the basisphenoid,
but must be on the basioccipital as well, and in MPIH
6863, the hypophysis is centered over the spheno-occipital
synchondrosis, extending anteriorly over the basisphenoid
and posteriorly over the basioccipital. Anteromedial to the
hypophyseal fossa in AMNH 28271 is a faint groove for the
internal carotid artery running in a broad arc from the piriform fenestra toward the presphenoid (“gica” in Fig. 22).
The medial cut edge of the basisphenoid in AMNH
28271 reveals a transverse sinus canal at the level of the
foramen ovale (“tsc” in “bs” in Fig. 21C). MPIH 6863 has
three sorts of venous foramina that connect to the venous
transverse canal: in the ventral surface of the basisphenoid
in or near the pterygoid suture (described above), in the
lateral aspect of the basisphenoid in a common depression
with the alisphenoid canal (described with the alisphenoid
below; “tcf” in Figs. 16, 19), and in the dorsolateral aspect
of the basisphenoid within the cranial cavity. The last could
not be verified in the skulls examined. However, AMNH
28271 has an additional connection to the transverse sinus canal not present in MPIH 6863; a probe pushed into
the caudal opening of the pterygoid canal emerges in the
transverse sinus canal and then out an aperture into the rear
of the cavum epiptericum (Gaupp 1902, 1905), the extradural endocranial space housing the trigeminal ganglion.
This aperture, the rostral opening of the pterygoid canal,
is within the dorsal surface of the basisphenoid at the level
of the transverse canal foramen and is visible through the
sphenorbital fissure in all specimens. In MPIH 6863, the
pterygoid canal is separated from (passes ventral to) the
transverse sinus canal.
Alisphenoid (“as” in figures)
In ventral view in AMNH 28272 (Figs. 16, 18, 19), the
alisphenoid is an irregular shaped quadrangular element.
Its medial side is delimited by the entopterygoid and
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caudal processes of the pterygoid (Fig. 19). Its anterior side
overlaps the orbitosphenoid ventrally and the frontal dorsally, and slopes anterodorsally such that the anterodorsal
corner is rostral to the anteroventral corner. The anterior
side (Figs. 16, 19) has a U-shaped cutout ventrally where
the alisphenoid forms the lateral wall of the sphenorbital
fissure and a L- or J-shaped cutout (left versus right side)
dorsally where the alisphenoid rims the depression housing the anterior opening of the orbitotemporal canal, the
frontal diploic vein foramen, and the posterodorsal ethmoidal foramen. On its dorsal margin (Figs. 16, 19), the
alisphenoid has a straight suture with the parietal and is the
concave member of a curved suture with the squamosal.
The posterior side is the shortest and forms the anteromedial border of the piriform fenestra, lateral to the caudal
opening of the pterygoid canal (Figs. 16, 19).
The surface adjacent to the piriform fenestra slopes anteroventrally to the low alisphenoid tympanic process, the
preotic crest of McDowell (1958) (“tpas” in Figs. 18, 19,
25). The alisphenoid tympanic process forms the posterior
margin of the foramen ovale (“fo” in Figs. 19, 25) and is
continuous posterolaterally with the entoglenoid process
of the squamosal (“egp” in Figs. 16, 25) and anteromedially with the entopterygoid process, although it does not
project as far ventrally as either neighboring process. The
bulk of the alisphenoid tympanic process has its main axis
oriented coronally, but it bends anteriorly to merge with the
back of the parasagittal entopterygoid process. The posterodorsal aspect of the slope posterior to the alisphenoid
tympanic process includes an oval depression identified by
McDowell (1958) as for the tensor tympani muscle (“ttf”
in Fig. 25). Medial to this depression is a faint groove for
the auditory tube and lateral to it is a broad groove for
the ramus inferior of the stapedial artery (McDowell 1958;
MacPhee 1981) and lesser petrosal nerve (“gri” in Figs. 18,
25, 27B), based on MPIH 6863. The composition of this
sulcus differs in AMNH 28272 and 185012, the only forms
preserving a suture here; in the former the alisphenoid and
squamosal contribute equally (Fig. 18), but in the latter
it is nearly entirely alisphenoid (Fig. 25). In CM 18069,
a dense connective tissue closes the sulcus for the ramus
inferior to form a canal; posterior to the alisphenoid, this
dense connective tissue canal continues posteriorly below
the piriform fenestra and ends at the anterior border of the
petrosal.
Several foramina penetrate the alisphenoid medial to
the glenoid fossa. Anterior to the tympanic process is the
round, anteroventrally directed foramen ovale (Fig. 25).
Although the foramen ovale is entirely in the alisphenoid,
its lateral border has a seam in AMNH 28272 suggesting
the foramen was formed by anterior and posterior arms of
the alisphenoid meeting lateral to the mandibular nerve
(Fig. 19). Anteromedial to the foramen ovale is an oval
depression (Fig. 16) that in its anterior half includes the
caudal opening of the alisphenoid canal and in its posterior
half a smaller opening for the transverse canal foramen
(transverse canal of McDowell 1958); the latter opening
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Fig. 24.—Solenodon paradoxus, AMNH 185012, line drawing of left braincase in lateral view. Abbreviations: as, alisphenoid; asc, alisphenoid canal;
bo, basioccipital; csm, crista supramastoideus; ctpl, lateral section of caudal tympanic process of petrosal; ctpm, medial section of caudal tympanic process of petrosal; e, ectotympanic; egp, entoglenoid process; gf, glenoid fossa; m, malleus (manubrium); me, mastoid exposure of petrosal; pet, petrosal;
pp, paroccipital process of petrosal; ptc, posttympanic crest; ptp, posttympanic process; sf, stapedius fossa; sh, stylohyal; smc, suprameatal crest; smef,
suprameatal foramen; sq, squamosal; th, tympanohyal; zpsq, zygomatic process of squamosal.
is double on the left side and single on the right side of
AMNH 28272 (also in CM 18069 and AMNH 212912 with
the foramen quite small in both). Rather than an oval depression, there is a round depression in AMNH 28271 and
185012; a transverse canal foramen is absent from the left
side of AMNH 185012 (Fig. 14), small and single on the
right of AMNH 28271 and 185012, and small and double
on the left side of AMNH 28271. These various openings
are confirmed as transverse canal foramina based on MPIH
6863.
Only the anterior aspect of the endocranial surface of
the alisphenoid is visible through the foramen magnum
in AMNH 28272 (see Fig. 21C). The alisphenoid has a
straight transverse suture with the orbitosphenoid anteriorly and a straight suture angled posterolaterally with
the parietal laterally. The anterolateral corner of the alisphenoid is roughly one millimeter medial to the posterior
opening of the orbitotemporal canal. What is visible of the
alisphenoid is confined to the braincase floor. Sutures between the alisphenoid and its endocranial neighbors are not
preserved in the bisected skull AMNH 28271.
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Squamosal (“sq” in figures)
The paired squamosal is a major contributor to the external
sidewall of the posterior braincase and the anterolateral basicranium; it also has a narrow exposure in the posterolateral floor of the middle cranial fossa and on the ventrolateral occiput. The squamosal is separated by sutures from its
neighbors externally in AMNH 28272 (Figs. 5, 16, 18) and
185012 (Figs. 6, 11), and only along the posterior suture
with the petrosal in AMNH 28271, 212912, and CM 18069
(Fig. 12). Most of the sutures between the squamosal and
neighbors endocranially are preserved in the bisected skull
AMNH 28271 (Fig. 21C).
In dorsal view, the most visible part of the squamosal
is the triangular zygomatic process (“zpsq” in Figs. 2, 5).
The zygomatic process ends in a short, anteriorly directed
point, which is broken on the left side of AMNH 185012
(Fig. 2), the right side of AMNH 28272 (Fig. 5), and CM
18069 (Fig. 4); a sharp anterior point seems to be lacking
naturally in AMNH 28271. Posterior to the tip, the dorsolateral edge of the zygomatic process, the crista supramastoidea of the Nomina Anatomica Veterinaria (1994), has
a raised crest roughly over the anteroposterior length of
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the glenoid fossa (“csm” in Fig. 24). Behind the crest, the
dorsolateral edge of the squamosal represents the top of the
suprameatal crest and posttympanic process (see below).
The relatively flat, horizontal surface medial to the dorsolateral edge is penetrated by a variable number (between
three and six) of tiny to small foramina in all specimens.
Based on MPIH 6863, these foramina drain veins into the
substance of the squamosal and the horizontal surface provides attachment for the temporalis muscle.
In lateral view in AMNH 28272 (Fig. 16) and 185012
(Fig. 6), the squamosal’s contribution to the lateral braincase has an irregular arrowhead shape with the point directed anteriorly. The squamosal can be divided into two
parts by their relation to the glenoid fossa (“gf” in Figs.
6, 16), the squama rostrally and the caudal process. The
tip of the squama, the point of the arrowhead, contacts the
alisphenoid (Figs. 14, 16). Behind the tip, the squamosal
overlaps the ventral margin of the parietal along a fairly
straight suture that first is angled posterodorsally and then
turns posteriorly before ending at the nuchal crest (Fig. 6).
The posterior border of the squamosal is more completely
preserved in AMNH 185012 (Figs. 24, 34); it forms the anteroventral part of the nuchal crest along with the mastoid
exposure of the petrosal. The nuchal crest is not vertical
but is angled anteroventrally (Figs. 6, 7). Near its midpoint
the posterior border of the squamosal has a small concavity that accommodates an oval exposure of the mastoid on
the lateral braincase wall (Fig. 24). The ventral border of
the squamosal is free, inflected medially on the basicranium. The posterior two-thirds represent the crista supramastoidea over the glenoid fossa, the suprameatal crest
over the external acoustic meatus (“smc” in Fig. 24), and
posttympanic process (“ptp” in Fig. 24). A well-developed,
laterally directed foramen for ramus temporalis penetrates
the caudal process near the parietal suture dorsal to the
suprameatal crest in AMNH 28272 and 185012 (“rt” in
Figs. 2, 4, 6). A foramen in a comparable position occurs
in the other specimens, but the parietosquamosal suture is
obliterated (Fig. 7). Additional smaller, posterodorsally directed foramina for rami temporales occur nearer or within
the parietosquamosal suture in AMNH 28272 (Fig. 5) and
185012 (Figs. 2, 6). The left side of AMNH 28272 (Fig.
5) has two in the squamosal and one in the parietosquamosal suture; and the right side has two in the suture. The
right side of AMNH 185012 (Fig. 2) has one within the
squamosal and three in the parietosquamosal suture; the
left side has one in the suture. Similar foramina occur in
the other specimens (Figs. 4, 7), but their bony relationship
cannot be ascertained, because as noted above the parietosquamosal suture is gone.
In ventral view in AMNH 28272 (Fig. 18) and 185012
(Figs. 11, 25), the squamosal can be divided front to
back into preglenoid, glenoid, and postglenoid areas. The
pre- and postglenoid areas, the ventral aspects of the squama and caudal process respectively, are positioned dorsal
to the glenoid. Based on AMNH 28272 (Fig. 18), the triangular preglenoid and oval glenoid areas have a medial
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foliate suture at which the squamosal underlies the alisphenoid, and the quadrangular postglenoid area has a plane suture along its posteromedial and posterior borders with the
petrosal.
The oval glenoid area, which includes the glenoid fossa,
zygomatic process, and entoglenoid process (“egp” in Fig.
25), has its long axis directed from posteromedial to anterolateral. McDowell (1958:142) described the glenoid fossa
in detail:
“On specimens of Solenodon paradoxus with the glenoid articular apparatus dried in place, two separate ovoid spots of cartilage
can be seen on the glenoid surface of the squamosal. One is on
the external portion of the glenoid fossa (the portion of the fossa
borne by the vestige of the zygomatic portion of the squamosal);
this, the external capitular facet, faces directly ventrad. The other
capitular facet lies more mesially, on the projecting postglenoid
process [entoglenoid of this report]; this the postglenoid [entoglenoid] capitular facet, faces forward and dorsad. The external and
postglenoid [entoglenoid] capitular facets are quite separate on
the squamosal.”
These two facets articulate with corresponding facets
on the mandible (see below). The pronounced entoglenoid
process, which buttresses the medial and posteromedial
glenoid fossa, is tongue shaped and angled obliquely from
posterolateral to anteromedial (Fig. 25). It is continuous
medially with the low tympanic process of the alisphenoid (Fig. 25), best seen in the adults (Fig. 12). A variety of terms have been employed for this process on the
squamosal: postglenoid process (Allen 1910), postglenoid
(entoglenoid) process (Gregory 1910), modified entoglenoid process (McDowell 1958; MacPhee 1981), pseudopostglenoid (entoglenoid) process (Novacek 1986a),
and pseudoglenoid process (MacPhee 1994). McDowell
(1958:172) concluded that the solenodon process is not
equivalent to the postglenoid process of most mammals,
but confusingly throughout the descriptions and illustrations of Solenodon he employed the term postglenoid process, which he meant in a generic way for “any kind of
projection of the squamosal forming a posterior brace for
the squamosal-dentary articulation.” Following McDowell
(1958), the term entoglenoid process is used here, because
this process is medial to the main glenoid surface. Additionally, near the midpoint of the posterior surface of the
entoglenoid process is a narrow groove for the chorda tympani nerve (McDowell 1958; “gct” in Fig. 25); the chorda
tympani nerve lies medial or dorsomedial to a true postglenoid process (McDowell 1958; MacPhee 1981).
Posteroventral to the glenoid fossa and entoglenoid
process is the narrow, quadrangular postglenoid area (Fig.
25). The most conspicuous feature of the postglenoid area
is the large postglenoid foramen (“pgf” in Figs. 25, 27A),
which is situated posterior to the lateral margin of the
entoglenoid process. Extending ventrally from the postglenoid foramen onto the posterior face of the entoglenoid
process is a broad groove for the postglenoid vein. In the
intact skull, the surface anteromedial to the postglenoid foramen is covered by the anterior crus of the ectotympanic
(“e” in Fig. 25) and the anterior process of the malleus.
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In fact, the anterior crus appears to form the medial wall
of the postglenoid foramen, but actually contributes to the
medial wall of the groove for the postglenoid vein immediately ventral to the postglenoid foramen (cf. right and left
sides in Fig. 25). The ectotympanic is missing from one
side in four specimens (Figs. 12, 18; right side of AMNH
28271, 28272, 185012; left side of CM 18069). In two of
these (AMNH 185012; CM 18069), there is a narrow facet
on the squamosal anteromedial to the postglenoid foramen
for the ectotympanic and rostral process of the malleus
(“efc” in Fig. 25). Bilaterally in the juvenile AMNH 28272
is a foramen medial to the postglenoid foramen that opens
endocranially (Figs. 26, 27); in MPIH 6863, this foramen
is an additional route for venous blood to the postglenoid
vein. Much smaller versions of this opening occur bilaterally in AMNH 185012 and on the right side of AMNH
28271, but in the latter the foramen is within the medial
aspect of the groove for the postglenoid vein. Posteromedial to and of similar girth to the postglenoid foramen is
a depression, the epitympanic recess, the space over the
incudomallear joint (Klaauw 1931; “er” in Figs. 25, 26B,
28). The anterolateral half of the epitympanic recess is on
the squamosal and the posteromedial half on the petrosal.
The remaining postglenoid area posterolateral to the
postglenoid foramen and epitympanic recess includes a
central depression and two eminences at the posterior and
posteromedial aspects connected by a ridge. The central
depression, which housed the external acoustic meatus, has
an irregular surface medially and smooth surface laterally
(“eam” in Figs. 25, 26B). The eminence along the posterior
border is the posttympanic process of the squamosal (“ptp”
in Figs. 24, 25, 26B, 29B), the back of which has a large
depression that based on MPIH 6863 is for the common
tendon of the sternomastoid and cleidomastoid muscles of
Allen (1910). The eminence along the posteromedial border is the posttympanic crest (“ptc” in Figs. 24, 25, 26B,
29B), which Wible et al. (2004) named for a more pronounced process in the Late Cretaceous eutherian Zalambdalestes. The solenodon posttympanic crest contacts the
tympanohyal (“th” in Figs. 25, 29B) and the posterior crus
of the ectotympanic (Fig. 25); Wible et al. (2004:82) reported the former contact in Zalambdalestes and speculated the latter as well. In AMNH 212912, the posttympanic
crest also has a narrow contact with the stylohyal (see “sh”
in Fig. 25).
All solenodon specimens have one or more small foramina either in the anterior base of the posttympanic crest
or in the central depression for the external acoustic meatus. Gregory (1910; “f.sb.sq.” in Fig. 1) identified the one
on the left side of AMNH 28272 as a subsquamosal foramen transmitting a vein; the right side of AMNH 28272
has seven foramina across this part of the squamosal (three
of which are labeled “smef” in Fig. 27A). MPIH 6863 has
a similar tiny opening that transmits a branch off the ramus
posterior of the stapedial artery. Because these openings
lie inferior to the suprameatal crest, they are identified as
suprameatal foramina.
Wible.indd 356
The endocranial surface of the squamosal is fully exposed
in the bisected skull AMNH 28271 (Fig. 21C) and partially
visible through the piriform fenestra in AMNH 28272.
The latter preserves sutures delimiting the squamosal
entirely from its neighbors, and the former preserves the
petrosquamous suture and parts of the sutures with the
parietal (Fig. 21C). Only a narrow endocranial exposure
of the squamosal anterior to the petrosal is present; this
represents the inner surface of the caudal process of the
squamosal and an endocranial exposure of the squama is
lacking. The squamosal contacts the parietal anteriorly and
the alisphenoid ventrally. The bulk of the squamosal exposure is a broad sulcus transmitting the prootic sinus (“sps”
in Fig. 21C) to its exit on the skull base at the postglenoid
foramen. In the ventral half of this groove are two foramina
(“1” and “2” in Fig. 21C), one posterodorsal and the other
anteroventral, leading to the single external postglenoid foramen. On the right side of AMNH 28271, the two internal
apertures are next to each other, but on the left side they
are separated. Both apertures can be seen from the postglenoid foramen on the exterior bilaterally in AMNH 28271,
28272, and on the left side of AMNH 212912. In the dorsal
part of the sulcus for the prootic sinus is the internal orifice
for a foramen for ramus temporalis (“rt” in Fig. 21C).
Petrosal (“pet” in figures)
The paired petrosal is usually divided into the anteroventromedial pars cochlearis, enclosing the cochlear duct and
saccule, and the posterodorsolateral pars canalicularis, enclosing the utricle and semicircular canals. The therian petrosal presents four surfaces: tympanic or ventral, cerebellar
or dorsal, squamosal or lateral, and lambdoid or mastoid
(MacIntyre 1972; Wible 1990). Much of the squamosal
surface is visible in the juvenile AMNH 28272, because
the petrosal and squamosal bones are narrowly separated.
The remaining three surfaces can be viewed in all specimens. The dorsal surface is best viewed in the bisected
skull of AMNH 28271, but part of the endocranial surface
is visible through the foramen magnum in the remaining
specimens.
Ventral View (Figs. 25–28).—In ventral view, the pars
cochlearis is represented by the ovoid cochlear housing,
whose long axis is oblique, from posterolateral to anteromedial. The pars cochlearis abuts the basioccipital and basisphenoid medially and anteromedially, and is continuous
with the pars canalicularis posteriorly and laterally; most of
the anterior border of the pars cochlearis contributes to the
posterior edge of the piriform fenestra. Centrally located on
the ovoid pars cochlearis and recessed from its medial and
anterior borders is a raised, round hill, the promontorium
of the petrosal (“pr” in Figs. 25, 26B), reflecting the coiling
of the cochlear duct through three or slightly more turns in
MPIH 6863. The juvenile AMNH 28272 has a flatter, less
protuberant promontorium than the other specimens (Figs.
19, 26, 27).
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Fig. 25.—Solenodon paradoxus, AMNH 185012, line drawing of basicranium in ventral view. Tympanohyal is fused to lateral section of the caudal tympanic process of petrosal (not labeled; see Figs. 26B, 28, 29). Abbreviations: as, alisphenoid; asc, alisphenoid canal; bo, basioccipital; bs, basisphenoid;
cf, carotid foramen; cpc, craniopharyngeal canal; ctpm, medial section of caudal tympanic process of petrosal; e, ectotympanic; eam, roof of external
acoustic meatus; efc, ectotympanic facet on squamosal; egp, entoglenoid process; eo, exoccipital; er, epitympanic recess; fm, foramen magnum; fo,
foramen ovale; fs, facial nerve sulcus; gct, groove for chorda tympani nerve; gf, glenoid fossa; gri, groove for ramus inferior of stapedial artery; hf, hypoglossal foramina; ips, inferior petrosal sinus groove; jf, jugular foramen; m, malleus (manubrium); mc, mastoid canaliculus; me, mastoid exposure of
petrosal; oc, occipital condyle; on, odontoid notch; pc, pterygoid canal; pcop, paracondylar process; pf, piriform fenestra; pgf, postglenoid foramen; pp,
paroccipital process of petrosal; pr, promontorium of petrosal; pt, pterygoid; ptc, posttympanic crest; ptp, posttympanic process; rtp, rostral tympanic
process of petrosal; sf, stapedius fossa; sh, stylohyal; smn, stylomastoid notch; sq, squamosal; th, tympanohyal; tpas, tympanic process of alisphenoid;
ttf, tensor tympani muscle fossa; vcof, ventral condyloid fossa.
In the posterolateral surface of the promontorium is an
elliptical opening, the oval window or fenestra vestibuli,
which accommodates the footplate of the stapes (“fv” in
Figs. 26B, 28). The length and width of five oval windows
were measured to calculate the stapedial ratio of Segall
(1970) with the average being 3.02; the remainder was obscured from view. The values are 2.9 and 3.0 on the left
and right side of AMNH 28272; 3.1 and 3.0 on the left
and right side of AMNH 185012; and 3.1 on the right side
of AMNH 28271. The fenestra vestibuli is nearly vertical, with the dorsal rim only slightly lateral to the ventral
rim. The posterior half of the fenestra vestibuli is recessed
slightly from the neighboring promontorial surface, producing a narrow vestibular fossula, best developed posterodorsally.
Centrally positioned in the posterior surface of the
promontorium is a kidney-bean shaped opening that is
hidden from view by tympanic processes described below,
except on the right side of AMNH 28271 and left side of
CM 18069 where the processes are broken off (“cfo” in
Fig. 28). McDowell (1958) identified this opening in Solenodon as the fenestra rotunda or fenestra cochleae, but
MacPhee (1981) correctly identified it as the aperture of
the cochlear fossula. The fenestra cochleae, the aperture
closed by the secondary tympanic membrane, is recessed
Wible.indd 357
from the exterior of the promontorium; this is well shown
on the right side of AMNH 28271 because the secondary tympanic membrane is preserved in situ. Between the
fenestra cochleae and the aperture of the cochlear fossula
lies the cochlear fossula, a “funnel-shaped niche in auditory capsule which contains fenestra cochleae and hides it
from view” (MacPhee 1981:51).
Medial to the promontorium is a rostral tympanic process of the petrosal that extends the length of the pars cochlearis and varies in height (“rtp” in Figs. 25, 26B, 29A).
In its anterior two-thirds, the rostral tympanic process is a
low ridge (not projecting as far ventrally as the promontorium) that decreases slightly in height anteriorly (Figs.
26A, B, 29A); the ventral margin of this ridge is sharp in
AMNH 28271 and 28272 but blunter in the other skulls.
In contrast, in its posterior one-third, the rostral tympanic
process is expanded into a high, laminar, quadrangular
element that contacts the posterior crus of the ectotympanic ring and is continuous with another high, laminar,
quadrangular element originating from the pars canalicularis, a caudal tympanic process of the petrosal (“ctpm”
in Fig. 25). The line of origin for the solenodon rostral
tympanic process is strikingly similar to that of MacPhee’s
(1981: fig. 2) rostral tympanic process in his schematic
placental ear region including its posterior origin at the
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posteromedial margin of the aperture of the cochlear
fossula. In his study of the juvenile solenodon serial sections (MPIH 6863), MacPhee (1981) reported the rostral
tympanic process as absent, and upon restudy of this specimen the oversight is understandable. The rostral tympanic
process is present, but much less pronounced in height
and length in MPIH 6863 than in the studied skulls. Near
the posterior limit of the rostral tympanic process medial
to the aperture of the cochlear fossula is a small opening
through the process (“tca” in Figs. 27A, 29A). Based on
MPIH 6863, this opening transmits the tympanic nerve, a
branch of the glossopharyngeal nerve, and, therefore, is a
tympanic canaliculus.
Dorsomedial to the rostral tympanic process is a shallow vascular sulcus running along the medial face of the
pars cochlearis between the anterior pole and the jugular
foramen (“ips” in Figs. 25, 29A). This sulcus can be seen
in all specimens except the juvenile AMNH 28272 where
glue and tight approximation of the petrosal and basioccipital hinder observation. Based on MPIH 6863, the occupant of this sulcus is the inferior petrosal sinus. Dorsal
to the sulcus for the inferior petrosal sinus, the pars cochlearis overlaps the basioccipital at a squamous suture based
on the right side of AMNH 28271, forming a roof over
the sulcus for the inferior petrosal sinus. There is also a
partial floor to the groove formed by a lamina on the pars
cochlearis, again visible in all specimens except AMNH
28272, although it is barely perceptible in AMNH 28271.
The extent of this lamina varies; it runs the length between
the anterior pole and jugular foramen in AMNH 185012
but is confined to the rostral half of the pars cochlearis in
CM 18069 and AMNH 28271 and 212912. In MPIH 6863,
the inferior petrosal sinus exits the skull via the piriform
fenestra, receiving the pterygoid vein running posteriorly
ventral to the basisphenoid, then runs posteriorly ventral to
the lateral edge of the basioccipital and medial edge of the
pars cochlearis, and joins the internal jugular vein beneath
the jugular foramen.
Anterior to the promontorium the pars cochlearis is
prolonged into a shelf, which following MacPhee’s (1981)
terminology is an epitympanic wing of the petrosal (“ew”
in Figs. 19, 26B). The solenodon epitympanic wing is not
flat, but undulates with the central part a rounded column
directed anteromedially and positioned ventral to concavities on either side of it. The lateral concavity is a fossa for
the tensor tympani muscle based on MPIH 6863, and the
smaller, shallower medial concavity is bordered medially
by the rostral tympanic process. The central column on its
lateral surface bears a groove for the internal carotid artery
(see below) directed anteromedially toward the basisphenoid. The maximum length of the epitympanic wing varies
between one-third in the juvenile AMNH 28272 and onehalf in the adult AMNH 28271 of the maximum length of
the promontorium.
The ventral surface of the solenodon promontorium bears
grooves for the internal carotid circulation, which have been
described elsewhere (McDowell 1958; MacPhee 1981).
Wible.indd 358
At the posterior end of the low anterior part of the rostral
tympanic process is a shallow millimeter wide concavity that
indicates the entrance of the internal carotid artery into the
middle ear (“gica” in Figs. 27A, 28). A similar size groove
runs laterally from the entrance point towards the fenestra
vestibuli. About a millimeter medial to the fenestra vestibuli,
this groove divides into an anteromedially directed groove
for the continuation of the internal carotid artery and a laterally directed groove for the stapedial artery (“gsa” in Figs.
26B, 28). The former runs the length of the promontorium
and epitympanic wing; the latter ends at the fenestra vestibuli, specifically at the posterior half of the ventromedial
border of the fenestra vestibuli. Stapes are preserved within
the fenestra vestibuli bilaterally in AMNH 212912 and CM
18609; on the right side of the latter dried blood representing
the stapedial artery passes through the large stapedial (intercrural) foramen. MacPhee (1981) noted that the stapedial
artery at its origin was about twice the diameter of the rostral continuation of the internal carotid artery in MPIH 6863.
The grooves are hard to measure at their division in most of
the studied skulls, because they are obscured by the petrosal
tympanic processes and ectotympanic. However, the right
side of AMNH 28271 (Fig. 28) and left of CM 18069 are
missing the tympanic processes and ectotympanic; here the
groove for the stapedial artery is 0.9 mm and that for the rostral continuation of the internal carotid artery is 0.5 mm.
The pars canalicularis is essentially a horizontal shelf
with various processes, sulci, and spaces on its ventral
surface lying posterior and lateral to the pars cochlearis.
The pars canalicularis has squamous sutures posteromedially with the exoccipital and laterally with the squamosal.
The juvenile AMNH 28272 serves as the basis for description, because its auditory ossicles and ectotympanic are
removed, fully exposing the pars canalicularis. However,
some ossification features are not as developed as in the
older skulls.
Dominating the lateral part of the pars canalicularis is
a longitudinal crest, which represents the crista parotica
(“cp” and “pcp” in Fig. 26B) and its derivatives. Near the
middle of this longitudinal crest is the attachment of the
tympanohyal (“th” in Fig. 26B), which for the purposes of
description is used to divide the crest into anterior and posterior halves. By definition, the point of attachment of the
tympanohyal on the auditory capsule is the crista parotica
(De Beer 1937; MacPhee 1981). The tympanohyal is a 1.3
mm long, rod-shaped, near vertical projection in AMNH
28272, directed slightly anteriorly and medially. Its ventral
end has an oval facet for the missing stylohyal, which is
preserved in situ on the left side of AMNH 185012 (“sh”
in Figs. 25, 29) and 212912. The stylohyal facet is directed
posteroventrally. The lateral surface of the tympanohyal
base contacts the posttympanic crest of the squamosal in all
specimens except AMNH 28272, but this absence appears
artificial, produced by shifting of the bones after death. In
the skulls preserving the ectotympanic, the posterior crus
contacts the tip of the tympanohyal medial to the stylohyal
facet (Figs. 25, 29B).
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Fig. 26.—Solenodon paradoxus, AMNH 28272, drawings of right ear region in ventral view. A, shaded drawing; B, line drawing; C, line drawing with
internal carotid artery and main branches in place based on MacPhee (1981) and MPIH 6863. Arrow in B points to hidden base of lateral section of the
caudal tympanic process on the crista interfenestralis, the bar of bone between the fenestra vestibuli and the aperture of the cochlear fossula. Scale is
for B and C. Abbreviations: as, alisphenoid; bo, basioccipital; bs, basisphenoid; cp, crista parotica; ctpl, lateral section of caudal tympanic process of
petrosal; ctpm, medial section of caudal tympanic process of petrosal; eam, roof of external acoustic meatus; eo, exoccipital; er, epitympanic recess;
ew, epitympanic wing of petrosal; fs, facial nerve sulcus; fv, fenestra vestibuli; gct, groove for chorda tympani nerve; gsa, groove for stapedial artery
(posterior part); hf, hypoglossal foramina; ica, internal carotid artery; jf, jugular foramen; mb, muscular branch; mfc, mallear facet on crista parotica;
mnb, meningeal branch; oc, occipital condyle; pa, posterior auricular artery; pcp, posterior continuation of crista parotica; pf, piriform fenestra; pp,
paroccipital process of petrosal; pr, promontorium of petrosal; ptc, posttympanic crest; ptp, posttympanic process; ri, ramus inferior of stapedial artery;
rio, ramus infraorbitalis; rm, ramus mandibularis; rp, ramus posterior of stapedial artery; rs, ramus superior of stapedial artery; sa, stapedial artery; sf,
stapedius fossa; sma, suprameatal artery; th, tympanohyal; tpas, tympanic process of alisphenoid; tt, tegmen tympani; vcf, ventral condyloid foramen.
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Fig. 27.—Solenodon paradoxus, AMNH 28272, drawings of right ear region in oblique ventral view. A, line drawing; B, line drawing with lateral head vein,
facial nerve, and auricular branch of the vagus nerve in place based on MPIH 6863. Abbreviations: abX, auricular branch of vagus nerve; cpc, craniopharyngeal canal; efc, ectotympanic facet on squamosal; fn, facial nerve; fo, foramen ovale; gf, glenoid fossa; gg, geniculate ganglion; gica, groove for internal carotid artery; gpn, greater petrosal nerve; gri, groove for ramus inferior of stapedial artery; gsa, groove for stapedial artery (anterior part); hF, hiatus Fallopii;
lhv, lateral head vein; lhvs, sulcus for lateral head vein; lwfi, lateral wall of fossa incudis; mc, mastoid canaliculus; pc, pterygoid canal; pcop, paracondylar
process; pgf, postglenoid foramen; prc, prootic canal; smef, suprameatal foramina; smn, stylomastoid notch; tca, tympanic canaliculus.
The posterior half of the longitudinal crest (“pcp” in
Fig. 26B) is the posterior continuation of the crista parotica
or lateral section of the caudal tympanic process situated
lateral to the stapedius fossa (MacPhee 1981); the former
term is employed here. The posterior continuation of the
crista parotica is near vertical and uniform in height. It is
thinnest immediately posterior to the tympanohyal, with a
faint concavity on the medial surface marking the stylomastoid notch (“smn” in Fig. 27A), the exit of the facial
nerve from the ear region (Fig. 27B). It is thickened posteriorly where it abuts the posttympanic process of the squamosal at the base of the nuchal crest; this represents the
paroccipital process of the petrosal (see Wible and Gaudin
2004; “pp” in Figs. 26B, 29B). The surface dorsolateral to
the posterior continuation of the crista parotica is angled
ventromedially and abuts the squamosal, with the pars
canalicularis slightly concave, as visible in AMNH 28272
where an artificial narrow gap separates the two bones.
The anterior half of the longitudinal crest has a medial
section extending to the piriform fenestra and a shorter but
higher lateral section. The lateral section (“lwfi” in Fig.
27A) arises from the anterolateral base of the tympanohyal
and extends forward anterolaterally for a millimeter. Its lateral surface is in contact with the posttympanic crest of the
Wible.indd 360
squamosal. The lateral section is a thin, sharp, vertical crest
that is low posteriorly and ends in an anteroventrally projecting prong based on the right sides of AMNH 185012
and 212912; this process is blunt on the left side of AMNH
28272 and broken on the right. The bulk of this crest forms
the lateral wall of the fossa incudis (not visible in ventral
view), housing the crus breve of the incus, with the anterior prong contributing to the posterolateral wall of the
epitympanic recess (described above in Squamosal). The
fossa incudis is a small, shallow, anteromedially directed,
near vertical depression set about a half millimeter posterior to (and continuous with) the epitympanic recess immediately anterior to the tympanohyal attachment.
The medial section of the longitudinal crest’s anterior
half is the crista parotica (“cp” in Fig. 26B). It follows a
sinuous course between the anteromedial base of the tympanohyal and the piriform fenestra. It has a near vertical
base attached to the overlying horizontal shelf of the pars
canalicularis and a sharp ventral end that is bent medially
to form the lateral edge of an incomplete horizontal floor
of the lateral head vein sulcus and facial sulcus (see below). Posteriorly, it forms the low-lying medial wall of the
fossa incudis and anteriorly, the medial wall of the epitympanic recess. In AMNH 28272, the crista parotica ends in
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an anteroventromedially directed prong, posterior to which
is a millimeter long facet (“mfc” in Fig. 26B) and a second
prong (double on the right side). Based on the left side of
AMNH 185012 and both sides of AMNH 212912, this facet contacts a thick process on the rostral (anterior) process
of the malleus. Also, based on MPIH 6863, the tendon of
the tensor tympani muscle runs behind the prong posterior
to the mallear facet.
Medial to the crista parotica is a longitudinal space
running the length of the lateral part of the pars canalicularis. This space is widest in its posterior half between
the posterior continuation of the crista parotica and caudal tympanic process (see below) where it accommodates
the fossa for the stapedius muscle (“sf” in Figs. 26B, 28).
The ovoid stapedius fossa has a long axis of some one and
a half millimeters set obliquely anterolateral to posteromedial. The raised posterior rim of the stapedius fossa is
formed by the lateral semicircular canal. The narrower
anterior half of the space medial to the crista parotica is
bordered medially by the pars cochlearis and provides passage for three principal structures, the facial nerve (“fn” in
Fig. 27B), the lateral head vein (“lhv” in Fig. 27B), and the
stapedial artery (“sa” in Fig. 26C). This space is described
by tracing the course of these three structures, based on
MPIH 6863.
The facial nerve leaves the cranial cavity on the endocranial surface of the petrosal via the facial canal in the
foramen acusticum superius of the internal acoustic meatus (described below; “fac” in Fig. 21C). After a short
course through the facial canal, the nerve enters a space
within the pars cochlearis hidden from external view, the
cavum supracochleare, where the nerve expands into the
geniculate ganglion (“gg” in Fig. 27B). There are two conduits from the cavum supracochleare, neither of which is
fully visible in ventral view in AMNH 28272, the hiatus
Fallopii for the greater petrosal nerve (“gpn” in Fig. 27B)
and the secondary facial foramen for the continuation of
the facial nerve (“fn” in Fig. 27B). The smaller hiatus Fallopii (“hF” in Figs. 27A, 28) lies near the middle of the
anterior border of the tensor tympani fossa in the posterior
border of the piriform fenestra and opens anteromedially;
the secondary facial foramen lies anterior to the fenestra
vestibuli and opens posterolaterally. The hiatus Fallopii is
partially hidden by an anteroventromedially directed spine
(“tt” in Figs. 26B, 28), which continues posteriorly to the
fenestra vestibuli as a low crest that fully conceals the secondary facial foramen. The spine, crest, and bone between
the hiatus Fallopii and secondary facial foramen represents
the tegmen tympani (see below) and forms the floor of the
cavum supracochleare and the roof of the lateral part of the
tensor tympani fossa.
Several authors (e.g., Rougier et al. 1998; SánchezVillagra and Wible 2002) have considered the position of
the hiatus Fallopii as a character in phylogenetic analyses. Three states are usually identified with the principal
difference being the relative proportions of the ventral
floor and dorsal roof of the hiatus to one another: a dorsal
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361
Fig. 28.—Solenodon paradoxus, AMNH 28271, photograph of right ear
region in ventral view. Dashed lines circumscribe the broken bases of the
posterior part of the rostral tympanic process of the petrosal and the medial and lateral sections of the caudal tympanic process of the petrosal. The
opening in the midst of these broken bases is the aperture of the cochlear
fossula; the cochlear fossula is the depression just inside the aperture.
Anterior to and in a plane perpendicular to the cochlear fossula is the preserved secondary tympanic membrane (not visible here), which defines
the fenestra cochleae. Scale = 2 mm. Abbreviations: as, alisphenoid; bo,
basioccipital; bs, basisphenoid; cfo, cochlear fossula; ctpl(br), broken
lateral section of caudal tympanic process of petrosal; ctpm(br), broken
medial section of caudal tympanic process of petrosal; er, epitympanic
recess; fs, facial sulcus; fv, fenestra vestibuli; gica, groove for internal carotid artery; gsa, groove for stapedial artery (posterior part); jf, jugular foramen; pf, piriform fenestra; pps, postpromontorial tympanic sinus; psc,
posterior semicircular canal; ptc, posttympanic crest; sq, squamosal; tt,
tegmen tympani.
hiatus position has the floor projecting anterior to the roof;
an intermediate position has the floor and roof subequal
in their anterior projection; and a ventral position has the
roof projecting anterior to the floor. The solenodon specimens exhibit all three states: a dorsal position occurs in
AMNH 212912, an intermediate in AMNH 185102, and
a ventral in AMNH 28271 (Fig. 28). Technically, the juvenile AMNH 28272 has a ventral position, but it is not
far removed from an intermediate one (Fig. 27). It is
cautioned that in instances where the hiatus Fallopii is at
the anterior border of the petrosal, such as the solenodon
and Monodelphis (see Wible 2003), characters concerning
the position of the aperture may not be very informative.
Posterior to the secondary facial foramen, between the
crista parotica and the pars cochlearis on the right side of
AMNH 28272 is a roughly V-shaped space, with the base
pointing posteriorly toward the stapedius fossa. The longer
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medial leg of the V is the facial sulcus (“fs” in Figs. 26B,
28), which transmits the facial nerve from the secondary
facial foramen to the stylomastoid notch posterior to the
tympanohyal. The shorter lateral leg of the V is a sulcus
(“lhvs” in Fig. 27A) for a vessel in MPIH 6863 that is
identified here as the lateral head vein (“lhv” in Fig. 27B),
because of its resemblance to the vein of the same name
known for monotremes and some marsupials among extant
mammals (Wible 1990; Wible and Hopson 1995; Rougier
and Wible 2006). The lateral head vein sulcus ends at two
small, anterolaterally directed foramina opposite the fenestra vestibuli (the anterior one at the level of the back of the
epitympanic recess; “prc” in Fig. 27A); it is possible to
see anterolaterally through these two foramina to the squamosal surface of the pars canalicularis (described below).
The lateral head vein sulcus is much shorter on the left side
of AMNH 28272, because the two small foramina are more
posteriorly place (the anterior one at the level of the fossa
incudis). Between the legs of the V (i.e., between the facial
sulcus and lateral head vein sulcus) is a rounded eminence
that appears to represent the ampulla of the lateral semicircular canal.
A lateral head vein sulcus is lacking in the remaining
examined specimens, but a foramen comparable in size to
the hiatus Fallopii resembling the double ones in AMNH
28272 is visible bilaterally in AMNH 28271 and on the
right side of AMNH 212912 at the level of the tympanohyal. Such foramina appear to be lacking on the left side of
AMNH 212912 and bilaterally in AMNH 185012, but
might be hidden from view by the anterior continuation of
the crista parotica. On the left side of AMNH 28271, the
bisected skull, a hair was passed through the foramen and
was traced to a small opening in the squamosal within the
sulcus for the prootic sinus just dorsal to the postglenoid
foramen. The same course is present in MPIH 6863, that is,
a connection between the lateral head vein and prootic sinus through a canal in the pars canalicularis and a small foramen in the squamosal. In its position, contents, and connections, the venous conduit through the pars canalicularis
represents a prootic canal and the opening on the ventral
surface of the pars canalicularis is a tympanic aperture of
the prootic canal. Since it original description in the echidna by Gaupp (1908), a prootic canal has been found among
extant mammals in the platypus and some marsupials and
is commonly present among Mesozoic mammaliaforms
(Wible 1990; Wible and Hopson 1995; Rougier and Wible
2006), including the Late Cretaceous eutherian Maelestes
Wible et al., 2007 (see Wible et al. 2007, in press) and isolated petrosals referred to the Early Cretaceous eutherian
Prokennalestes Kielan-Jaworowska and Dashzeveg, 1989
(see Wible et al. 2001) and Late Cretaceous zhelestids
(Ekdale et al. 2004). The discovery of a prootic canal in the
solenodon is the first for any placental, extant or extinct.
The third neurovascular structure to consider is the stapedial artery (see also McDowell 1958). After penetrating
the stapedial foramen in the stapes, the stapedial artery
(“sa” in Fig. 26C) turns anteriorly ventral to the facial
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nerve in the facial sulcus. At the secondary facial foramen,
where the facial nerve bends anteromedially to enter the
cavum supracochleare, the stapedial artery continues forward in a millimeter long sulcus before reaching the piriform fenestra (“gsa” in Fig. 27A). The higher lateral wall
of this sulcus is formed by the anterior continuation of the
crista parotica and the medial wall is formed by the tegmen
tympani.
A last issue to consider with the lateral part of the pars
canalicularis is the tegmen tympani, which generally is
considered a neomorphic component of the chondrocranium in mammals (DeBeer 1937; Moore 1981), although
it is reported by some (e.g., Kuhn and Zeller 1987) to be
lacking in monotremes. The tegmen tympani is produced
forward from the crista parotica and forms a new side wall
of the cranial cavity lateral to the primary side wall represented by the prefacial (suprafacial) commissure (DeBeer
1937; Moore 1981). As a result, a new space, the cavum
supracochleare, is formed between the pars cochlearis and
tegmen tympani. The facial nerve pierces the primary side
wall beneath the prefacial commissure via the facial canal in the internal acoustic meatus and enters the cavum
supracochleare where the geniculate ganglion is located.
In addition to its usage in the mammalian chondrocranium, the term tegmen tympani is also used in adult human
anatomy for the “thin, translucent plate of bone [petrous
temporal=petrosal]. . .forming the roof of the tympanic cavity” (Terry 1942:147) and is found in the Nomina
Anatomica (1983) and Nomina Anatomica Veterinaria
(1994). Applying the term in the adult is often complicated
by late ontogenetic fusions between the tegmen tympani
and the pars cochlearis. Following De Beer’s (1937) and
Moore’s (1981) usage, the tegmen tympani in ventral view
in AMNH 28272 includes the floor of the cavum supracochleare, which is produced forward as the spine beneath
the hiatus Fallopii described above, and the adjacent parts
of the facial sulcus and the sulcus for the stapedial artery
(“tt” in Figs. 26B, 28).
Dominating the posterior aspect of the pars canalicularis is the caudal tympanic process, which as stated above
is a high, laminar, quadrangular element continuous anteromedially with the rostral tympanic process. The caudal tympanic process has two separate points of origin on
the pars canalicularis (“ctpm(br)” and “ctpl(br)” in Fig.
28), which following MacPhee (1981) are termed the medial and lateral sections of the caudal tympanic process.
MacPhee (1981) further recognized two types of lateral
sections based on their position lateral or medial to the
stapedius fossa; the lateral section of the caudal tympanic
process in the solenodon is of the type medial to the stapedius fossa. The base of the medial section (“ctpm” in Fig.
26B) is situated about a millimeter posterior to the aperture
of the cochlear fossula, is about 1.5 mm wide in AMNH
28272, and lies in a coronal plane. Extending anteroventrally from the base at about a 45º angle, the medial section
is a quadrangular lamina about three millimeters high in
AMNH 28272. Near its anteroventral terminus, the medial
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363
Fig. 29.—Solenodon paradoxus, AMNH 185012, photographs of left ear region. A, oblique medial view; B, oblique posterior view. 1 = the laminar
posterior part of the rostral tympanic process of petrosal; 2 = the medial section of the caudal tympanic process of petrosal; and 3 = the lateral section of
the caudal tympanic process of petrosal. Scale = 2 mm. Abbreviations: bo, basioccipital; bs, basisphenoid; e, ectotympanic; egp, entoglenoid process;
fn, aperture for facial nerve; gf, glenoid fossa; ips, inferior petrosal sinus; m, malleus (manubrium); mc, mastoid canaliculus; me, mastoid exposure of
petrosal; oc, occipital condyle; pgf, postglenoid foramen; pp, paroccipital process of petrosal; ptc, posttympanic crest; ptp, posttympanic process; rtp,
rostral tympanic process of petrosal; sh, stylohyal; sm, aperture for stapedius muscle tendon; tca, tympanic canaliculus; th, tympanohyal.
section merges medially with the rostral tympanic process
(described above) and laterally with the lateral section of
the caudal tympanic process. The base of the lateral section (“ctpl” in Fig. 26B) is situated about a millimeter lateral to the aperture of the cochlear fossula on the crista
interfenestralis (the bar of bone between the round and
oval windows), is about a half millimeter wide in AMNH
28272, and lies obliquely posteromedial to anterolateral.
Extending ventrally from the base, the lateral section is a
near vertical quadrangular lamina about a millimeter high
that merges with the medial section, as noted above.
The base of the medial section of the caudal tympanic
process is about a millimeter posterior and a millimeter
dorsal to both the base of the lateral section and the base
of the rostral tympanic process (Fig. 28). The separation
of these tympanic processes at their bases produces two
openings: a medial one (“mc” in Figs. 25, 26B, 29A) between the rostral tympanic process anteriorly, the pars
canalicularis dorsally, and the medial section of the caudal
tympanic process ventrally, and a lateral one between the
lateral section of the caudal tympanic process anteriorly,
the pars canalicularis dorsally, and the medial section ventrally. These two openings lead into the same space, between the aperture of the cochlear fossula and the medial
section of the caudal tympanic process, and provide a passageway for two structures running between the jugular foramen and stylomastoid notch in MPIH 6863: the auricular
ramus of the vagus nerve (also noted by McDowell 1958
and MacPhee 1981; “abX” in Fig. 27B) and the lateral
head vein draining into the internal jugular vein. On the
left side of AMNH 185012, the medial opening is divided
by a thin bar into a smaller dorsal opening and larger ventral one (Fig. 29A). Based on MPIH 6863, the dorsal opening is for the auricular ramus of the vagus and the ventral
one for the lateral head vein. Rather than a complete bar
separating two openings, the right side of AMNH 185012,
left side of CM 18069, and both sides of AMNH 212912
Wible.indd 363
have an incomplete bar with a figure eight-shaped medial
opening; AMNH 28272 and the left side of AMNH 28271
merely have an oval medial opening. The passageway for
the auricular ramus of the vagus is the mastoid canaliculus
(NAV 1994).
The anteroventral margin of the merged rostral and
caudal tympanic processes contacts the posterior crus of
the ectotympanic at a squamous suture, three millimeters long on the left side of AMNH 185012 (Fig. 25). The
isolated ectotympanic of AMNH 28272 shows a distinct
facet where it abuts the tympanic processes (“tpfc” in
Fig. 31; see also McDowell 1958: fig. 5C, D). MacPhee
(1981:216) reported that the suture between the ectotympanic and tympanic processes (his caudal tympanic process) for MPIH 6863 was “in the process of disappearing”
and McDowell (1958) noted that fusion of these elements
occurs in large solenodon skulls. The skulls examined here
that preserve contact between the ectotympanic and tympanic processes show no indication of sutural fusion: the
left sides of AMNH 28271 and 185012, the right side of
CM 18069, and both sides of AMNH 212912. The left side
of AMNH 185012 preserves part of the stylohyal in place
(Figs. 25, 29) and the dorsal surface of that element has a
small abutment with the medial section of the caudal tympanic process, near its suture with the ectotympanic.
In addition to its fusion with the medial section of the
caudal tympanic process and sutural relationship to the
ectotympanic, the lateral section of the caudal tympanic
process has an additional osseous contact with the tympanohyal, except in the juvenile AMNH 28272 where
these elements are separated by a narrow gap (Fig. 26).
In the older specimens, the lateral section is fused to the
tympanohyal dorsomedial to the facet for the stylohyal on
the left side of AMNH 28271 and on both sides of AMNH
185012 (Figs. 25, 29B) and 212912; this relationship is
obscured by breakage on the right side of AMNH 28271
and bilaterally in CM 18069.
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Lateral to the lateral section of the caudal tympanic
process is the confluence of two spaces discussed above
medial to the crista parotica, the stapedius fossa posteriorly and the facial sulcus anteriorly. Based on MPIH 6863,
four structures pass through this confluence (Figs. 26C,
27B); positioned from dorsolateral to ventromedial these
are the lateral head vein, the facial nerve, the tendon of the
stapedius muscle, and the ramus posterior of the stapedial
artery (“rp” in Fig. 26C). The course of the last has been
described in detail for MPIH 6863 by MacPhee (1981) as
leaving the stapedial artery medial to the fenestra vestibuli, running posteriorly in a groove along the anterolateral
margin of the lateral section of the caudal tympanic process, and dividing into two branches: one to the stapedius
muscle and the other leaving the stylomastoid notch to end
in the auricle. With the exception of the juvenile AMNH
28272, weak laminar flanges derived from the lateral section of the caudal tympanic process medially and the crista
parotica laterally serve to subdivide the confluence of the
stapedius fossa and facial sulcus to segregate some of these
four structures (Fig. 29B). AMNH 185012 and the right
side of 212912 have flanges that meet to completely wall
off a ventromedial foramen for the ramus posterior (not
visible in the figures) and nearly meet to separate a central foramen for the stapedius tendon (“sm” in Fig. 29B)
from a dorsolateral foramen for the lateral head vein and
facial nerve (“fn” in Fig. 29B). The juvenile AMNH 28272
has a broad confluence with no indication of subdivision
(Fig. 26); and this area is obscured in CM 18069, AMNH
28271, and the left side of AMNH 28271.
At the posteromedial corner of the petrosal is the jugular incisure, the petrosal’s contribution to the anterolateral
wall of the jugular foramen (“jf” in Figs. 25, 28, 29B).
Within the jugular incisure, the petrosal is thick dorsoventrally and only slightly concave. The pars cochlearis and
pars canalicularis meet at the incisure with the posterior
border of the former marked by the opening into the cochlear canaliculus (for the perilymphatic duct and vein in
MPIH 6863) barely visible near the dorsal limit of the petrosal in the middle of the incisure (not visible in ventral
view). Posterolateral to the incisure, the pars canalicularis
meets the exoccipital at a foliate suture, with the petrosal
the concave member.
Dorsal View (Figs. 20, 21A, C, 22).—The endocranial
surface of the petrosal is only fully visible in the bisected
skull AMNH 28271, the basis for the following. Two large
apertures, the internal acoustic meatus and subarcuate
fossa, dominate the endocranial surface and represent the
principal elements in the pars cochlearis and pars canalicularis, respectively.
The elliptical internal acoustic meatus is only recessed
slightly from the endocranial surface of the pars cochlearis. A transverse crest (“tc” in Fig. 22), slightly more recessed than the meatus, delimits an inferior fossa, the foramen acusticum inferius (“fai” in Fig. 21C), from a superior
fossa, the foramen acusticum superius. The distribution
Wible.indd 364
of openings within these two fossae is similar to that in a
newborn human (Terry 1942: fig. 137), which provides the
basis for the terminology below.
The foramen acusticum inferius is round and dominated
by a deep, oval opening, the posterior half of which has
numerous small foramina, the spiral cribriform tract (“sct”
in Fig. 22), transmitting twigs of the cochlear nerve, and
the anterior half has an elliptical foramen, the foramen
centrale cochleare (“fcc” in Fig. 22), indicating the orifice
of the canal of the modiolus. The foramen acusticum inferius has two additional smaller openings, a larger dorsal
one and a smaller posterodorsal one, that are not as deeply
recessed as that containing the spiral cribriform tract and
central cochlear foramen. The dorsal foramen is the inferior vestibular area for nerves to the saccule (“iva” in Fig.
22); the posterodorsal foramen is foramen singulare for the
nerve to the ampulla of the posterior semicircular canal
(“fsi” in Fig. 22).
The foramen acusticum superius is subdivided by a
slightly recessed crest running perpendicular to the transverse crest into a larger round opening into the facial canal
(“fac” in Figs. 21C, 22) and a smaller oval superior vestibular area with numerous small foramina for twigs of the
nerve of the utricle (“sva” in Fig. 21C).
Dorsal to the internal acoustic meatus is the subarcuate
fossa (“saf” in Figs. 21C, 22), which housed the paraflocculus of the cerebellum. The aperture of the subarcuate
fossa is nearly square with only slight curvature at the four
angles. The posteromedial border of the aperture is formed
by the crus commune, the conjoined anterior and posterior
semicircular canals, and the dorsal and anterolateral borders by the anterior semicircular canal. The internal acoustic meatus and aperture of the subarcuate fossa are oriented
differently; the former is directed dorsomedially whereas
the latter is directed anteriorly and slightly medially. The
aperture of the subarcuate fossa is constricted with respect
to the fossa itself, except along the ventral border where
the aperture and fossa are in the same plane. In the lateral
wall of the fossa are half a dozen small foramina, transmitting venous blood based on MPIH 6863.
Anterior to the aperture of subarcuate fossa is a thin,
raised crest, the crista petrosa (“crp” in Figs. 21C, 22),
which continues anteroventromedially to form the slightly raised anterior border of the internal acoustic meatus
(equal to the prefacial commissure). The crista petrosa is
the attachment of the tentorium cerebelli and, thus, delimits the middle cranial fossa in front from the caudal cranial
fossa behind. Anterior to the crista petrosa is the narrow,
near vertical petrosal contribution to the posterior wall of
the middle cranial fossa. Based on MPIH 6863, this faintly
concave surface housed the back of the cerebrum. The majority of this surface is pars canalicularis with the ventral
end the tegmen tympani (“tt” in Fig. 22). The anterior face
of this surface abuts the squamosal; the ventral half of the
petrosquamous suture is plane and the dorsal half foliate.
Running the length of the dorsal margin of the pars
canalicularis is the petrosal’s contribution to the sulcus for
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the sigmoid sinus and anterior to that the petrosal’s contribution to the canal for the prootic sinus (both described
above with the Parietal; Fig. 21C). Between the sulcus for
the sigmoid sinus and the crus commune is the aqueductus vestibuli for the endolymphatic duct and vein based on
MPIH 6863 (“av” in Figs. 21C, 22). This aperture is dorsally directed and hidden in medial view by an irregular,
low process in AMNH 28271. The sulcus for the sigmoid
sinus bends ventrally posterior to this process and ends at
the condyloid canals in the exoccipital (Fig. 21C). MPIH
6863 has some venous blood exiting the jugular foramen
as the internal jugular vein; sections posterior to this with
the condyloid canals are not preserved. However, based
on the size of the condyloid canals in AMNH 28271, they
represent the primary drainage for the sigmoid sinus. Most
of the posterior margin of the pars canalicularis contacts
the exoccipital at a squamous suture, but based on AMNH
28272, there is also a narrow contact with the supraoccipital (see Fig. 21C).
Squamosal View.—As noted above, much of the squamosal surface is visible in the juvenile AMNH 28272, because of an artificial separation between the petrosal and
squamosal bones, more so on the right side than left. However, the separation is not wide enough to allow illustration of the squamosal surface. Nevertheless, the exposed
squamosal surface of the pars canalicularis is remarkably
similar to that of some extant didelphid marsupials, such
as Metachirus nudicaudatus (É. Geoffroy, 1803), AMNH
72551 illustrated in Sánchez-Villagra and Wible (2002: fig.
5c; this figure is erroneously attributed to Marmosa murina (Linnaeus, 1758), AMNH 99980 in the figure legend)
and re-illustrated here (Fig. 30). As in M. nudicaudatus,
the squamosal surface of the solenodon pars canalicularis
has two principal parts: a broader posterior part overlaid
by the squamosal bone and a narrower anterior part not in
direct contact with the squamosal bone.
The surface of the posterior part of the squamosal surface of the juvenile solenodon pars canalicularis is convex dorsoventrally, and its posterior margin is thickened
ventrally, representing the base of the paroccipital process. Near the middle of the posterior part in M. nudicaudatus is a longitudinal posttemporal sulcus (“pts” in Fig.
30), which accommodates the arteria and vena diploëtica
magna based on personal observations (Wible 1990, 2003)
of Didelphis marsupialis Linnaeus, 1758 and Monodelphis
domestica (Wagner, 1842). In other M. nudicaudatus with
the petrosal in place, some (e.g., AMNH 96621) have a
posttemporal foramen between the squamosal and mastoid
exposure of the petrosal transmitting the posttemporal vessels to the occiput and others (e.g., CM 52728) do not. The
juvenile solenodon also has a shallow posttemporal sulcus,
less than half a millimeter in width, but only on the caudal
half of the posterior part (the incidence of the posttemporal foramen is considered with the Mastoid view). Based
on MPIH 6863, the primary occupant of the juvenile solenodon posttemporal sulcus is venous; this specimen has
Wible.indd 365
365
Fig. 30.—Metachirus nudicaudatus, AMNH 72551, right petrosal in lateral view, redrawn from Sánchez-Villagra and Wible (2002: fig. 5c; this
figure was attributed erroneously to Marmosa murina, AMNH 99980,
in the figure legend). Abbreviations: cp, crista parotica; fc, fenestra cochleae; fv, fenestra vestibuli; hF, hiatus Fallopii; prc, prootic canal; pts,
posttemporal sulcus; rtp, rostral tympanic process of petrosal; sps, sulcus
for prootic sinus; tgf, trigeminal ganglion fossa; th, tympanohyal.
a tiny arteria diploëtica magna arising from the ramus
superior dorsal to the piriform fenestra and well-developed
accompanying veins.
The surface of the anterior part of the squamosal surface of the pars canalicularis in M. nudicaudatus has a near
vertical sulcus for the prootic sinus (“sps” in Fig. 30), the
anterior distributary of the transverse sinus in marsupials
and monotremes (Wible 1990, 2003; Wible and Hopson
1995), at the ventral end of which is a small lateral aperture of the prootic canal (“prc” in Fig. 30). The intact squamosal bone contributes more surface area to the sulcus for
the prootic sinus, but does not fully enclose a canal. The
juvenile solenodon AMNH 28272 has the same arrangement, although the single lateral aperture of the prootic canal is more centrally placed in the ventral end of the sulcus
for the prootic sinus; a probe passed through this aperture
reaches both of the tympanic apertures of the prootic canal described above. The adult solenodon, based on the
bisected skull AMNH 28271, exhibits a major difference
from both the juvenile and M. nudicaudatus. Rather than
the lateral aperture of the prootic canal opening directly
into the prootic sinus as in the juvenile and M. nudicaudatus, the squamosal in the adult has an additional thin
layer of bone that encloses a second lateral aperture of the
prootic canal within the squamosal appressed to the opening within the petrosal. In summary, the prootic sinus in
the adult solenodon runs ventrally in an anteromedially
open sulcus between the squamosal and pars canalicularis
(Fig 21C); dorsal to the postglenoid foramen the sulcus is
entirely within the squamosal. In the posteromedial aspect
of this squamosal sulcus is a lateral aperture of the prootic
canal in the squamosal leading into the prootic canal within the pars canalicularis.
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Fig. 31.—Solenodon paradoxus, AMNH 28272, drawings of left ectotympanic and broken rostral process of malleus. A, ventral (extratympanic) view;
B, dorsal (intratympanic) view. Abbreviations: acr, anterior crus; at, sulcus for auditory tube; ct, crista tympanica; pcr, posterior crus; pep, petrosal
process of rostral process of malleus; rpm, rostral process of malleus; sp, styliform process; st, sulcus tympanicus; ti, tympanic incisure; tpfc, tympanic
process of petrosal facet.
Mastoid View (Figs. 34, 35).—The exposure of the pars
canalicularis on the occiput, the mastoid exposure, is
roughly trapezoidal. The medial and dorsal borders are
subequal and long, whereas the lateral and ventral are subequal and short. The medial border is near vertical and has
a squamous suture with the exoccipital, based on AMNH
28272, which preserves a suture between the ex- and
supraoccipital (Fig. 35). The dorsal border is oblique, following the contour of the nuchal crest; from dorsomedial
to ventrolateral, it overlaps the supraoccipital, based on
AMNH 28272 (Fig. 35), and has squamous sutures with
the parietal and the squamosal. The lateral border, the
paroccipital process, is near vertical and has a squamous
suture with the posttympanic process of the squamosal; it
forms the raised medial edge of a muscular depression on
the posterior face of the posttympanic process. The ventral
border has a central concavity, which includes the posterior face of the lateral semicircular canal, flanked by the
paroccipital process laterally and the medial section of the
caudal tympanic process medially.
Based on MPIH 6863, the lateral border, paroccipital
process, provides attachment for the common tendon of the
sternomastoideus and cleidomastoideus of Allen (1910),
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which arises primarily from the depression on the backside
of the posttympanic process of the squamosal. Also based
on MPIH 6863, the ventral end of the paroccipital process
marks the lateral limit of a broad sheet of musculature that
medially reaches the medial section of the caudal tympanic
process and exoccipital; as noted by Thiel (1955) this sheet
includes the posterior digastric and mastoideohyoideus
(mastoideostyloideus of Saban 1968). In the two longest
skulls, AMNH 28271 and 212912, the posterior face of the
posterior semicircular canal is raised from the surface of
the mastoid exposure dorsal to the medial section of the
caudal tympanic process.
Small foramina penetrate the mastoid surface in all specimens (“msf” in Fig. 34) except the juvenile AMNH 28272
and the left side of AMNH 212912. Most are along the dorsal
border near the suture with the squamosal: AMNH 185012
and 212912 right side only; AMNH 28271 and CM 18069
bilaterally. The left side of AMNH 185012 and right side
of AMNH 28271 have a more dorsomedially placed foramen, near the suture with the parietal. These openings do not
lead into a posttemporal canal between the pars canalicularis
and squamosal and, therefore, are not posttemporal foramina, but are identified as mastoid foramina. Gregory (1910;
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Wible—On the Cranial Osteology of Solenodon paradoxus
“?f.p.m.” in Fig. 1) and McDowell (1958) noted a small foramen on the lateral mastoid surface of AMNH 28272 that
they thought might be a mastoid foramen (their post-mastoid
foramen), but this is a slit rather than a foramen related to the
juvenile nature of the growing petrosal bone.
Ectotympanic (“e” in Figs. 11, 18, 24, 25, 31)
The C-shaped ectotympanic is preserved in situ bilaterally in
AMNH 212912, the left side of AMNH 28272 and 185012,
and the right side of AMNH 28271 and CM 18069 (Figs.
11, 12). McDowell (1958: fig. 5C, D) figured external and
internal views of a left ectotympanic (his tympanic) for an
unspecified S. paradoxus, but this element is from a specimen other than AMNH 28272, because the latter bone was
preserved in situ at the beginning of this project (Fig. 18)
but subsequently fell off (Fig. 19). In addition, the right ectotympanic in CM 18069 was removed for this study. Here
the isolated elements are described first, followed by those
preserved in situ.
Referring to Figure 31, the maximum dimension of the
isolated left ectotympanic of the juvenile AMNH 28272 and
the right of CM 18069 is roughly 5 mm, from the styliform
process (“sp” in Fig. 31) to the distal end of the anterior
crus (“acr” in Fig. 31); the dimension perpendicular to that
is roughly 4 mm, from the anterior surface of the anterior
crus to the posterior surface of the posterior crus (“pc” in
Fig. 31). The anterior crus is fairly straight, whereas the posterior crus is more curved, even hook-like at its distal end;
the tympanic incisure, the gap between the distal ends of the
anterior and posterior crura, is less than a millimeter (“ti” in
Fig. 31).
The anterior face of the anterior crus is flat except where
it is crossed by a millimeter wide, obliquely-oriented (from
dorsolateral to ventromedial) groove housing the rostral
process of the malleus. The groove is well shown in McDowell’s (1958: fig. 5C, D) illustrations, but is occupied by
the rostral process of the malleus in AMNH 28272 (“rpm”
in Fig. 31) and CM 18069. The rostral process is partially
fused to the anterior crus in both specimens and broke from
the rest of the malleus, which in the case of AMNH 28272
is missing. The anterior face of the anterior crus is also of
uniform height except at its distal end where it tapers to a
rod-like prong. The anteromedial corner of the ectotympanic has a short, trapezoidal styliform process (Klaauw
1931; =anteromesial spine of McDowell 1958 or anterior
process of MacPhee 1981), which is slightly more robust
in CM 18069. The dorsal surface of the styliform process
has a faint sulcus (“at” in Fig. 31B), which based on MPIH
6863 floors the auditory tube. The proximal half of the posterior crus in AMNH 28272 and proximal two-thirds in CM
18069 are smooth and uniform in width (Fig. 31). However,
the distal part bears a concave facet on its posterior surface
(“tpfc” in Fig. 31) for the ectotympanic’s abutment with the
conjoined rostral and caudal tympanic processes of the petrosal described above. In AMNH 28272, the bone around the
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facet is broader than the proximal posterior crus, whereas in
CM 18069 it is narrower. The size of the facet varies in the
other specimens: it is about half the posterior crus in AMNH
212912, but two-thirds of it in AMNH 27271 and 185012.
The inner margin of the ectotympanic has a well-developed sulcus tympanicus (“st” in Fig. 31B) except at the
distal tips of both the anterior and posterior crurae; only
the middle portion of the sulcus is visible in dorsal view.
Based on MPIH 6863, the sulcus tympanicus is the point
of attachment of the tympanum. Forming the dorsal margin of the sulcus tympanicus is a low semicircular ridge, the
crista tympanica (“ct” in Fig. 31B). McDowell (1958) noted
the presence in solenodon of a recessus meatus, “the most
proximal part of the bony external auditory meatus, set off
from the meatus proper by its broader diameter” (McDowell
1958:128). The recessus meatus (“rem” in Fig. 31B) is the
narrow gap between the sulcus tympanicus and the ventralmost margin of the ectotympanic.
The ectotympanic bones preserved in situ (AMNH
212912, the right side of AMNH 28271, and the left side
of AMNH 185012) provide information about the contacts
of the tympanic ring with the skull base. The anterior crus
abuts the posteromedial base of the entoglenoid process of
the squamosal (Figs. 12, 25), occupying a facet described
above (“efc” in Fig. 25). The posterior crus abuts the conjoined rostral and caudal tympanic process of the petrosal,
the tympanohyal, the posttympanic crest of the squamosal,
and the stylohyal in the two instances part of that element is
preserved (left side of AMNH 185012 and 212912).
MacPhee (1981) reported the inclination of the right and
left ectotympanic of MPIH 6863 to be 40º from the horizontal plane of the basicranium. The angle in the three skulls
preserving ectotympanics in situ varies: 34º bilaterally in
AMNH 212912, 43º on the left side of AMNH 185012, and
46º on the left side of AMNH 28272. The different values
may result from differences in developmental stage, but a
larger sample is needed to address that.
Auditory Ossicles
Malleus (Fig. 32).—The right malleus of CM 18069 was
removed in two pieces with some damage at their point of
union. The malleus is also preserved in situ bilaterally in
AMNH 212912 and on the right of AMNH 185012. The
malleus in Figure 32 is based on CM 18069 with the damaged portion completed from the other three elements in
AMNH 185012 and 212912.
The basic parts of the malleus are the head, neck, manubrium, osseous lamina, and rostral (anterior) process. The
head (“hm” in Fig. 32B) bears two articular surfaces for the
incus set off from each other at an angle of 110º. The much
larger superior (anterior) articular facet is flat and oval,
whereas the inferior (posterior) articular facet is triangular
and strongly convex (“suaf” and “iaf” in Fig. 32B). The
anteroventral border of the head is delimited by a weak capitular crest with a capitular spine (“cs” in Fig. 32B).
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Fig. 32.—Solenodon paradoxus, CM 18069, right malleus (rostral process broken and reconstructed based on AMNH 185012 and 212912). A, dorsal
view shaded drawing; B, dorsal view line drawing; C, ventral view line drawing. Abbreviations: cb, central buttress; cs, capitular spine; fct, foramen for
chorda tympani nerve; gct, groove for chorda tympani nerve; hm, head; iaf, inferior articular surface; ila, inner longitudinal lamella; lp, lateral process;
mn, manubrium; mnb, manubrial base; mptt, muscular process for tensor tympani; nm, neck; oa, orbicular apophysis; ol, osseous lamina; ola, outer
longitudinal lamella; pep, petrosal process of rostral process of malleus; suaf, superior articular facet.
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The neck (“nm” in Fig. 32B) runs posteriorly from the
posterior aspect of the head and turns medially and ventrally to the base of the manubrium. Just before achieving
the manubrial base, the neck is crossed by the groove for
the chorda tympani nerve on its dorsal aspect (“gct” in Fig.
32B). Immediately lateral to the groove for the chorda tympani is the large, digitiform muscular process for the attachment of the tensor tympani muscle based on MPIH 6863
(“mptt” in Fig. 32B).
The base of the manubrium (“mnb” in Fig. 32B) is at
the confluence of the neck, orbicular apophysis, lateral
process, and manubrium. The prominent orbicular apophysis (“oa” in Fig. 32B, C) projects posteriorly from the
manubrial base. The lateral process is on the ventral aspect
of the manubrial base (“lp” in Fig. 32C) and marks the lateral limit of the tympanum attachment to the manubrium.
The manubrium (“mn” in Fig. 32B, C) is fully preserved
only on the left sides of AMNH 185012 (Fig. 29A) and
212912; it is flattened anteroposteriorly and tapers to its
tip, which is slightly more spatulated in the former specimen. In both specimens, the manubrium is only slightly
less inclined than the ectotympanic from the horizontal
basicranial plane. Its tip is not straight, but has a slight
ventral curvature (Fig. 29A).
Anterior to the manubrial base and muscular process
and medial to the head is the roughly quadrangular osseous
lamina (“ol” in Fig. 32B, C). The osseous lamina is paper thin and transparent; it is convex on its dorsal aspect
and deeply concave ventrally, which sets the lamina off
distinctly from the head. The groove for the chorda tympani nerve described above on the neck continues forward
along the dorsal aspect of the osseous lamina. The lateral
margin of the groove is marked by a distinct ridge extending forward from the muscular process; this ridge is the
central buttress (“cb” in Fig. 32B); there is no indication
of the central buttress on the ventral aspect of the osseous
lamina (Fig. 32C). The anterior margin of the osseous lamina is marked by a raised ridge on the dorsal and ventral
aspects: the inner lamella dorsally (“ilm” in Fig. 32B) and
outer lamella ventrally (“olm” in Fig. 32C). These lamellae connect the head to the well-developed rostral process.
The medial margin of the osseous lamina is not thickened
to form a pars processus anterioris.
The rostral process (“rpm” in Fig. 32B, C) lies perpendicular to the head and in situ extends anteroventrally between the anterior surface of the ectotympanic’s anterior
crus and the posterior surface of the entoglenoid process
of the squamosal, occupying facets on both bones (Figs.
25, 31). The rostral process is anteroposteriorly flat and tapers to a point at its anteroventral end. Where the posterior
surface of the rostral process meets the dorsal aspect of
the osseous lamina is the foramen for the chorda tympani
nerve (“fct” in Fig. 32B, C), transmitting the nerve through
the rostral process to a groove on the posterior face of the
entoglenoid process (“gct” in Figs. 25, 26B). Projecting
posterodorsally from the dorsal edge of the rostral process
is a distinct ovoid process, called here the petrosal process
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of the rostral process (“pep” in Fig. 32B), that contacts a
facet on the crista parotica (“mfc” in Fig. 26B). When the
malleus was removed from CM 18069, connective tissue
held these elements in articulation. MPIH 6863 preserves
this contact, with no indication of a synovial joint.
Incus (Fig. 33A, B).—The left incus was removed from
CM 18069 for study and illustration; the right incus is preserved in situ in CM 18069 unobstructed by the malleus,
which was removed for this study. The incus is preserved
in situ bilaterally in AMNH 212912, but most of it is hidden by the malleus and ectotympanic.
The basic parts of the incus are the body, crus breve
(short process), crus longum (long process), and the lenticular process. The body (“bi” in Fig. 33A) bears two articular surfaces for the malleus; the larger superior articular surface is oval and flat (“sas” in Fig. 33B), whereas the
inferior articular surface is triangular and strongly concave
(‘ias” in Fig. 33A, B). The crus breve (“cb” in Fig. 33A)
is short and cylindrical, projects posterodorsally from
the body into the fossa incudis in the petrosal part of the
epitympanic recess, and is attached there by the posterior
ligament of the incus. The crus longum (“cl” in Fig. 33A,
B) extends posteriorly and slightly ventrally and laterally
from the body towards the stapes; it lies roughly in the
same plane as the neck of the malleus, which it is dorsal
to and separated from by a narrow gap. The crus longum
tapers slightly at its distal end, turns medially at a right
angle, and is connected to the flat, oval lenticular process
(“lpr” in Fig. 33A, B) by a narrow pedicle (“ped” in Fig.
33B).
Stapes (Fig. 33C).—Stapes are preserved in situ bilaterally in AMNH 212912. The presence of the ectotympanic,
malleus, and incus in this specimen partially obscures the
head and footplate of the stapes, but a reconstruction of the
left stapes was made (Fig. 33C).
The stapes includes a head, anterior and posterior crurae, and footplate. The head (“hs” in Fig. 33C) articulates
with the lenticular process of the incus at the incudostapedial joint and appears to be roughly of the same shape and
dimension. The head blends with the merged anterior and
posterior crurae (“acr” and “pcr” in Fig. 33C), separated
from each by an expansion, the shoulder of the anterior
and posterior crurae (“acs” and “pcs” in Fig. 33C). On
the shoulder of the posterior crus is a small, posteriorly
directed muscular process for the insertion of the stapedius muscle based on MPIH 6863 (“mps” in Fig. 33C).
From the shoulders, the anterior and posterior crurae bow
outward and then inward to similar extents before connecting to the footplate (“fp” in Fig. 33C), which occupies the fenestra vestibuli on the petrosal promontorium.
The space outlined by the shoulders, crurae, and footplate
is the stapedial (intracrural) foramen (“stf” in Fig. 33C),
through which the stapedial artery passes based on MPIH
6863. The anterior crus is uniformly narrow, whereas the
posterior crus is expanded at the footplate end. Both the
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Fig. 33.—Solenodon paradoxus, CM 18069, drawings of left incus and stapes. A, left incus, oblique lateral view; B, left incus, oblique ventral view; C,
left stapes, slightly oblique anteroventral view. Abbreviations: acr, anterior crus; acs, shoulder of anterior crus; bi, body; cb, crus breve; cl, crus longum;
crs, crural sulcus; cst, crista stapedis; fp, footplate; hs, head of stapes; ias, inferior articular surface; lpr, lenticular process; mps, muscular process for
stapedius muscle; pcr, posterior crus; pcs, shoulder of posterior crus; ped, pedicle; sas, superior articular surface; stf, stapedial foramen.
anterior and posterior crurae have a crural sulcus facing
the stapedial foramen, but only the sulcus on the posterior
crus is visible in the figure because of the slightly oblique
view (“crs” in Fig. 33C). The edges delimiting the crural
sulci extend across the tympanic surface of the footplate as
a paired, low crista stapedis (“cst” in Fig. 33C).
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Occipital Complex
The occipital complex contributes to the basicranium, occiput, braincase roof, and the caudal cranial fossa. Based
on MPIH 6863 and AMNH 28272, the occipital complex
comprises four ossifications: the median basioccipital on
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the skull base, the paired exoccipital posterolateral to the
basioccipital on the skull base and on the occiput, and the
median supraoccipital dorsal to the foramen magnum on the
occiput and on the braincase roof. MPIH 6863 is the only
specimen delimiting the basioccipital and exoccipitals, and
AMNH 28272 is the only specimen delimiting the exoccipitals and supraoccipital (Fig. 35). The individual components
of the occipital complex are described based on the sutural
positions in MPIH 6863 and AMNH 28272.
Basioccipital (“bo” in figures)
The basioccipital occupies the midline of the basicranium
posterior to the basisphenoid, medial to the pars cochlearis of the petrosal, and anterior to the foramen magnum
(“fm” in Fig. 25). The basioccipital is fused to the exoccipitals in all skulls examined (Figs. 11, 12, 18, 25, 26A,
B), but these ossification centers are separated by cartilage
in MPIH 6863.
In the juveniles AMNH 28272 and 185012, the basioccipital is separated from the basisphenoid by a narrow
transverse gap, the open spheno-occipital synchondrosis,
at the level of the anterior limit of the rostral tympanic
process of the petrosal (Figs. 25, 26A, B); in the adults,
the basioccipital and basisphenoid are fused (Fig. 12). The
lateral contact with the pars cochlearis of the petrosal is
described above. As interpreted from MPIH 6863, the posterolateral limit of the basioccipital is at the anterior end of
the jugular foramen (“jf” in Fig. 25). The basi- and exoccipital juncture then runs obliquely posteromedially to the
anteromedial rim of the foramen magnum. The odontoid
notch (intercondyloid incisure of Evans 1993; “on” in Fig.
25) and part of the adjacent occipital condyles (“oc” in Fig.
25) are on the basioccipital, although the exact extent is not
known.
Ventral surface features of the basioccipital differ
across the age range examined. In the juvenile AMNH
28272 (Fig. 18), the entire surface is relatively flat; immediately anterior to the occipital condyles are faint ovoid
depressions for the rectus capitis ventralis based on MPIH
6863. In the older juvenile AMNH 185012 (Figs. 11, 12),
the central third of the anterior half is convex and flanked
by concavities for the longus capitis based on MPIH 6863;
the ovoid depressions for the rectus capitis ventralis are
more pronounced. The adult CM 18069 (Fig. 12) differs
in that it has a midline keel in the anterior half, rather than
a rounded eminence, that extends anteriorly onto a fainter
midline keel on what positionally must be the posterior half
of the basisphenoid, these two bones being fused. Finally,
the longest adult skull, AMNH 212912 (Fig. 12), differs in
that the midline keel on the basioccipital and basisphenoid
is more pronounced and there is a pronounced obliquely
oriented muscular tubercle on either side of the keel at
or near the juncture of the basioccipital and basisphenoid. Based on MPIH 6863, this process is for the longus
capitis, because that muscle arises from the basioccipital,
basisphenoid, and the spheno-occipital synchondrosis.
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The odontoid notch also differs with age. In the juveniles AMNH 28272 and 185012, the odontoid notch is
shallow and without facets (Figs. 11, 12, 18). In the adult
CM 18069 (Fig. 12), the notch is deeper and with faint facets for the odontoid process of the axis facing posteromedially on the posteromedial face of the occipital condyles;
the left and right facet come within a millimeter of each
other and are a millimeter at their thickest. Finally, in the
longest adult skull, AMNH 212912 (Fig. 12), the facets are
more pronounced, within a half millimeter of each other
and nearly two millimeters at their thickest.
In AMNH 28271 (Figs. 20, 21A, C), the endocranial
surface of the basioccipital on the right side is essentially
flat except anteriorly where it appears to contribute to the
shallow hypophyseal fossa (see Basisphenoid); most of
the basioccipital is missing from the left side (Fig. 21C). A
dorsum sellae and posterior clinoid processes are lacking.
Exoccipital (“eo” in figures)
The paired exoccipital has two quadrangular parts: a horizontal one in the skull base (Fig. 25) and a vertical one
in the occiput (Fig. 35); both contribute to the closure of
the foramen magnum and to the occipital condyles. The
exoccipitals are fused to the basioccipital in all skulls (Fig.
12) and to the supraoccipital in all (Fig. 34) except AMNH
28272 (Fig. 35).
The anteromedial border of the exoccipital’s horizontal
part with the basioccipital is described above. The oblique
lateral border includes the exoccipital’s contribution to the
walls of the jugular foramen in front and the foliate suture with the pars canalicularis of the petrosal behind. At
this suture with the petrosal, the edge of the exoccipital is
slightly raised (“pcop” in Figs. 25, 27A), more so in the
adults (Fig. 12); in MPIH 6863 this faint process provides
attachment for the rectus capitis lateralis and the broad
sheet of musculature that includes the mastoideohyoideus
and posterior digastric. In the dog (Evans 1993), the stout
jugular (paracondylar) process of the exoccipital has attachment for the same muscles (mastoideohyoideus=jugulohyoideus), and, therefore, the solenodon eminence is
identified as a very weak paracondylar process. The posterior border of the exoccipital’s horizontal process is on
the free posterior edge of the skull base; it is formed by
the occipital condyle and a depression between the condyle
and the faint paracondylar process, the ventral condyloid
fossa (“vcof” in Fig. 25). The posteromedial border is the
foramen magnum.
Between the occipital condyle and jugular foramen are
the hypoglossal foramina (“hf” in Figs. 25, 26B), which are
paired except for the right side of AMNH 185012, which
has three. The position and size of the paired foramina varies: they are subequal with the posterior one slightly lateral
to the anterior one in AMNH 28271, 28272, and 185012
(left side); subequal with the anterior one slightly lateral
to the posterior one in AMNH 212912 (left side); and the
anterior one smaller and slightly lateral to the posterior
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Fig. 34.—Solenodon paradoxus, AMNH 185012, drawings of skull in occipital view. Abbreviations: bcf, basicapsular fissure; bo, basioccipital; ctp,
caudal tympanic process of petrosal; dcf, dorsal condyloid foramen; e, ectotympanic; egp, entoglenoid process; eo, exoccipital; eoc, exoccipital crest;
me, mastoid exposure of petrosal; msf, mastoid foramen; nc, nuchal crest; oc, occipital condyle; pa, parietal; pet, petrosal; pp, paroccipital process
of petrosal; ptp, posttympanic process; rt, foramina for rami temporales; rtp, rostral tympanic process of petrosal; sc, sagittal crest; sh, stylohyal; so,
supraoccipital; soc, supraoccipital crest; socf, supraoccipital foramina; sq, squamosal; zpsq, zygomatic process of squamosal.
one in AMNH 212912 (right side) and CM 18069. The
hypoglossal foramina open into a common depression in
CM 18069 and AMNH 28271, 212912, and 185012 (right
side), whereas a common depression is lacking on the left
side of AMNH 185012 and both sides of AMNH 28272
(see Fig. 25). The posterior hypoglossal foramen shares
a common depression with one or more openings that
connect to the condyloid canal (described below with the
endocranial surface of the exoccipital) in AMNH 28271,
185012, 212912, and CM 18069, whereas the anterior hypoglossal foramen shares a common depression with such
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openings in AMNH 28271 and 28272.
The shallow ventral condyloid fossa lies posterior to
the hypoglossal foramina and contains a variable number of small to large-sized foramina that connect to the
condyloid canal (“vcf” in Fig. 26B). The fewest ventral
condyloid foramina are the two small ones on the left side
of AMNH 185012 and the most the seven small ones on
the left side of AMNH 212912. Along with several small
openings, AMNH 28271 has bilateral large openings that
are a millimeter and a half in maximum diameter.
The vertical process of the exoccipital (Fig. 35) has a
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373
Fig. 35.—Solenodon paradoxus, AMNH 28272, photograph and line drawing of skull in occipital view. Abbreviations: as, alisphenoid; bo, basioccipital; bs, basisphenoid; ctp, caudal tympanic process of petrosal; dcf, dorsal condyloid foramen; egp, entoglenoid process; eo, exoccipital; fm, foramen
magnum; ham, pterygoid hamulus; hf, hypoglossal foramina; me, mastoid exposure of petrosal; nc, nuchal crest; oc, occipital condyle; pet, petrosal;
pp, paroccipital process of petrosal; ptp, posttympanic process; so, supraoccipital; socf, supraoccipital foramina; sq, squamosal; th, tympanohyal; tpas,
tympanic process of alisphenoid; zpsq, zygomatic process of squamosal.
near vertical lateral suture with the pars canalicularis described above and a sinuous foliate suture with the supraoccipital dorsally. Its medial and ventral edges are free; the
former forms the lateral margin of the foramen magnum
and the latter is the occipital condyle and ventral condyloid fossa. At the lateral suture with the pars canalicularis,
the exoccipital has a raised crest (“eoc” in Fig. 34), more
pronounced in the adults, that is continuous ventrally with
the faint paracondylar process and dorsally with a more
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pronounced crest on the supraoccipital described below.
Following Gaudin (1995), this is identified as an exoccipital crest.
Immediately dorsal to the occipital condyle is a shallow dorsal condyloid fossa. In the medial margin of the
dorsal condyloid fossa are openings into the condyloid canal that vary in size and number (“dcf” in Figs. 34, 35).
AMNH 185012 (right side), 212912, and CM 18069 (left
side) have a single large opening; AMNH 28271, 28272
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(left side), 185012 (left side), and CM 18069 (right side)
have large and small openings; and AMNH 28272 (right
side) has a large and two small openings. Two specimens,
AMNH 212912 and CM 18069, also have an additional
small opening to the condyloid canal just anteromedial to
the ventral condyloid fossa on the endocranial side of the
foramen magnum.
As McDowell (1958:145) noted, the solenodon occipital condyle has two facets joined by a narrow isthmus of
bone: “one ventral and anterior to the foramen magnum
and another lateral to the foramen magnum and facing
backward.” The larger posterodorsal facet (Figs. 34, 35)
is more uniform in shape across the examined specimens;
its main axis is oblique, from posteroventromedial to anterodorsolateral. In occipital view it is cylindrical and in
ventral view elliptical. The smaller anteromedial facet is
labial-shaped in the juveniles (Figs. 11, 18, 25, 26), but is
more quadrangular in the adults (Fig. 12).
Reflecting the external surfaces, the endocranial surface of the exoccipital in AMNH 28271 has horizontal and
vertical components contributing to the floor and posterior
wall of the caudal cranial fossa (Fig. 21C). At the lateral
margin of the horizontal component are the two hypoglossal foramina (Fig. 21C; three on the right side of AMNH
185012), visible in all specimens through the foramen
magnum (Fig. 35). At the anterior margin of the vertical
component in the terminus of the sigmoid sinus sulcus in
AMNH 28271 are several large openings into the condyloid canal (“coca” in Figs. 21C, 22), which runs posteriorly
through the exoccipital to the ventral and dorsal condyloid
foramina described above. The left side of AMNH 28271
has a large opening subdivided by a thread-like septum and
the right side has three large and two small openings. Sections preserving the condyloid canal are not preserved for
MPIH 6863. McDowell (1958) described the condyloid
canal in solenodon as venous, which is the case in the ox
(Sisson 1910) and dog (Evans 1993).
Supraoccipital (“so” in figures)
The supraoccipital forms the dorsal half of the occiput and
contributes to the posteriormost margin of the braincase
roof. The juvenile AMNH 28272 is the only skull fully
delimiting the borders of the supraoccipital (Figs. 5, 35);
the older juvenile AMNH 185012 preserves only the anterior sutures with the parietals and interparietal (described
above; Fig. 2).
On the occiput, the supraoccipital has an irregular ventral border and a longer curved dorsal border that is the
nuchal crest (“nc” in Fig. 35). The middle third of its ventral edge forms the dorsal margin of the foramen magnum
and the lateral thirds contact the exoccipitals (see above).
The adults AMNH 212912 and CM 18069 have a midline
external occipital crest that extends about halfway down
from the nuchal crest (the midline is not preserved in
AMNH 28271); in the juveniles this area is flat in AMNH
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28272 (Fig. 35) and slightly convex in AMNH 185012
(Fig. 34). At the lateral margin of the supraoccipital is a
triangular depression dorsal to the pars canalicularis delimited by medial and lateral ridges that unite dorsally, except
in AMNH 28272 where the bone is featureless. The medial ridge (“soc” in Fig. 34) is continuous ventrally with the
exoccipital crest (and the faint paracondylar process) and is
termed the supraoccipital crest here. The lateral ridge is on
the nuchal crest; the dorsal half of the lateral ridge is formed
entirely by supraoccipital, whereas the ventral half has a
contribution from the parietal, based on AMNH 185012.
All specimens have supraoccipital foramina (“socf” in
Figs. 34, 35) that connect internally to the occipital emissary vein described above (see Parietal), but the number,
size, and position vary both between and within specimens.
AMNH 28272 has a centrally positioned heart-shaped depression that contains a half dozen smaller openings (Fig.
35); AMNH 185012 has two larger openings per side, well
off the midline, and a number of smaller openings (Fig.
34); AMNH 212912 and CM 18069 have an ovoid depression just to the left of the external occipital crest that
contains several smaller openings as well as an additional
dozen small openings arrayed across the occiput.
On the braincase roof, the supraoccipital has a narrow
exposure with sutures anteriorly with the interparietal and
parietals described above (Figs. 2, 5). Where the occipital
and dorsal surfaces of the supraoccipital meet is the nuchal
crest. In dorsal view, the nuchal crest is evenly convex in
AMNH 28272 (Fig. 5), with a slight midline concavity in
AMNH 185012 (Figs. 2, 4), and with a more pronounced
midline concavity in AMNH 212912 and CM 18069 (Fig.
4). In lateral view, the dorsal and occipital surfaces meet at
roughly a right angle in AMNH 28272, but in the remaining skulls the nuchal crest overhangs the occiput (Fig. 8).
Sutures delimiting the supraoccipital within the caudal cranial fossa are visible through the foramen magnum
in the juvenile AMNH 28272 (see Fig. 21C). The middle
third of the supraoccipital’s contribution to the endocranial
roof reflects that bone’s external exposure on the braincase
roof, because of the edge-to-edge nature of the suture with
the interparietals. In contrast, the lateral thirds of the supraoccipital’s endocranial roof contribution are more substantial than the corresponding braincase roof contribution,
because the parietals broadly overlap the supraoccipital
externally. The middle third of the supraoccipital’s endocranial roof contribution has a depression for the vermis of the
cerebellum that extends anteriorly onto the interparietals
and posteroventrally onto the endocranial side of the supraoccipital’s occiput contribution. The lateral thirds have
an impression for the back of the cerebrum that extends
anteriorly but is much fainter on the interparietal. The lateral margin of the supraoccipital’s roof contribution abuts
the posterodorsal corner of the pars canalicularis of the
petrosal.
In the bisected skull AMNH 28271, the endocranial
surface of the supraoccipital differs in one major regard,
the presence of a wide sulcus and large foramen for the
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Fig. 36.—Solenodon paradoxus, AMNH 185012, drawings of right mandible in lateral view. Abbreviations: an, angular process; c, lower canine; coc,
coronoid crest; con, condylar process; cor, coronoid process; i1, lower first incisor; i2, lower second incisor; i3, lower third incisor; m1, lower first
molar; m2, lower second molar; m3, lower third molar; maf, masseteric fossa; mb, mandibular body; mf, mental foramina; mr, mandibular ramus; p1,
lower first premolar; p4, lower penultimate premolar; p5, lower ultimate premolar; pdm, process for digastric muscle; rms, retromolar space.
occipital emissary vein (“oev” in Fig. 21C), presumably a
difference of developmental stage. The sulcus arises at a
right angle from the posterodorsal margin of the sulcus for
the sigmoid sinus and is 4.0 mm long and 1.5 mm wide. It
ends dorsolaterally at an opening that ultimately leads to
the various supraoccipital foramina on the occiput (Figs.
34, 35). A part of the supraoccipital’s suture with the parietal is preserved at the anteroventral end of the sulcus for
the occipital emissary vein showing the origin to be on the
supraoccipital (Fig. 21C). In light of the position of the
supraoccipital and parietal in the juvenile AMNH 28272,
it seems likely that the bulk of the sulcus and foramen is
on the supraoccipital in AMNH 28271. Ventral to the large
foramen for the occipital emissary vein is a much smaller
one in AMNH 28271 that is interpreted to be part of the
same venous system.
Mandible (Figs. 36–42)
The paired mandible consists of the shallow, tooth-bearing
horizontal part (body) and behind that, the broad vertical part (ramus). Each mandible houses ten teeth: three
Wible.indd 375
incisors, canine, three premolars, and three molars. In
AMNH 28271, 28272, and 185012, the right and left mandibles are separated from each other, whereas in CM 18069
and AMNH 212912 they are held together by soft tissues
at the mandibular symphysis (Fig. 41). Allen (1908) described the teeth of the juvenile AMNH 28272, although
several typographic errors complicate identifications at
certain loci; McDowell (1958) redescribed the teeth and
made corrections to Allen (1908). The identification of
deciduous and permanent lower teeth followed here is in
accord with McDowell (1958); the only loci explicitly
known by two generations are the second incisor, and the
penultimate and ultimate premolars.
In lateral view, the body of the mandible (“mb” in Fig.
36) is fairly uniform in depth posterior to i3, with the deepest point below m3 and shallowest below m1. The body
tapers anteriorly and is very shallow below the tiny i1,
whose alveolus is anteroventral to the large i2. The m3 is
separated from the coronoid process on the mandibular ramus by a narrow retromolar space (“rms” in Fig. 36). In the
juveniles AMNH 28272 and 185012, the body is riddled
with many tiny foramina, especially below the canine and
premolars (Fig. 42), but the adults have far fewer such tiny
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Fig. 37.—Solenodon paradoxus, AMNH 185012, drawings of right mandible in medial view. Abbreviations: an, angular process; ars, articular surface; c,
lower canine; con, condylar process; cor, coronoid process; i2, lower second incisor; m1, lower first molar; manf, mandibular foramen; mas, mandibular
symphysis; mb, mandibular body; mhl, mylohyoid line; mr, mandibular ramus; p4, lower penultimate premolar; pdm, process for digastric muscle.
Fig. 38.—Solenodon paradoxus, AMNH 185012, drawings of right mandible in occlusal view. Abbreviations: an, angular process; ars, articular surface;
c lower canine; con, condylar process; cor, coronoid process; i2, lower second incisor; m1, lower first molar; mas, mandibular symphysis; p4, lower
penultimate premolar; pdm, process for digastric muscle; rms, retromolar space.
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foramina (Fig. 39). The five specimens have multiple mental foramina (four or more) in slightly different positions
(“mf” in Fig. 36). In AMNH 28272 (Fig. 42), there are four
large mental foramina (below the canine, the posterior root
of p1, p4, and the anterior root of m1) on the left side and
four (below the posterior root of p1, the anterior root of
dp5, the anterior root of m1, and the posterior root of m1)
on the right side (Fig. 39); there are extra small foramina
on the right side (below the canine, the middle of p1, and
the posterior root of m1). In AMNH 185012, there are four
large mental foramina on the left side (below the posterior
root of p1, the anterior root of p4, the anterior root of p5,
and the p5/m1 embrasure) and two large (p1/p4 embrasure
and posterior root of p4) and three small (canine, anterior
root of m1, and anterior root of m2) on the right side (Fig.
36). In CM 18069, there are four large (c/p1 embrasure, p4
anterior root, anterior root of p5, and anterior root of m1)
and a small one (middle of p4) on the right side (Fig. 39),
and three large (c/p1 embrasure, p4 anterior root, and posterior root of p5) and two small (middle of p4 and posterior
root of m1) on the left side. In AMNH 212912, there are
four large (c/p1 embrasure, anterior root of p4, posterior
root of p4, and anterior root of m1) on the right side (Fig.
39) and three large (posterior root of p1, posterior root of
p4, and anterior root of m1) and one small (anterior root
of p5) on the left. In AMNH 28271, there are three large
mental foramina (below the middle of p1 and p4, and the
anterior root of m1) and two small (below the canine and
anterior root of p5) on the right side (Fig. 39), and three
large (below the middle of p1, the posterior root of p5,
and the anterior root of m1) and two small (below the canine and anterior root of m2) on the left. In AMNH 28272,
185012, and 212912, the mental foramina are positioned
in roughly the same horizontal plane, but in AMNH 28271
and CM 18069 the anterior ones are higher than the posterior ones (Fig. 39).
In medial view, the most prominent feature on the body
is the cigar-shaped symphyseal surface, which extends posteriorly to beneath the anterior root of p5 in all specimens
(“mas” in Fig. 37). The posterior part of the symphyseal
surface is not flush with the surface of the body but is elevated. With the two mandibles in articulation, this elevated area produces a narrow shelf opposite p4 and p5 (Fig.
41). Ventral and posteroventral to m3 is a distinct crest,
the mylohyoid line (more rounded in the adults), which
continues posteriorly a short distance onto the mandibular
ramus (“mhl” in Fig. 37).
In lateral view, the dorsal aspect of the mandibular ramus (“mr” in Fig. 36) has the usual coronoid and condylar
processes, positioned anteriorly and posteriorly respectively (“cor” and “con” in Fig. 36), but the ventral aspect
appears to have two angular processes (“pdm” and “an”
in Fig. 36). The anteroventral one lies beneath the coronoid
process and is separated by a concavity from the posterodorsal one, which lies beneath the condylar process. The
anteroventral process is smaller than the posterodorsal one,
ends in a sharp point, and is slightly inflected medially. The
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377
Fig. 39.—Solenodon paradoxus, AMNH 185012, CM 18069, AMNH
28271, and AMNH 212912, photographs of right mandibles in lateral
view. Scale = 10 mm.
posterodorsal process is more extensive, positioned at the
posteroventral corner and extending up the posterior border of the ramus, rounded, and with a low, raised crest on
its outer contour in the adults. The author agrees with Allen (1910), who considered the posterodorsal process to be
the true angle, because in his dissections the masseter and
medial pterygoid muscles attach there, whereas the digastric muscle attaches to the unnamed anteroventral process.
The concavity separating the posterodorsal and anteroventral processes varies with the developmental stage, being
more pronounced in the juveniles (Figs. 39, 42).
The coronoid process is twice as high as the condylar
process measured from the retromolar space and the base
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Fig. 40.—Solenodon paradoxus, AMNH 185012 and AMNH 28271,
photographs of right mandibles in medial view. Scale = 10 mm.
of the coronoid lies at roughly a right angle to the occlusal
plane. The anterior and dorsal borders of the coronoid process have a thickened coronoid crest (“coc” in Fig. 36), especially in the adults (Fig. 39). At the base of the coronoid
process, the coronoid crest subtly turns posteriorly and then
posterodorsally toward the concavity between the coronoid
and condyle, demarcating the ventral and posterior limit of
the masseteric fossa (“maf” in Fig. 36). Immediately above
this subtle ridge is the deepest part of the masseteric fossa,
a circular depression in the juveniles AMNH 28272 and
185012 and an oval depression (longer than high) in the
adults (Fig. 39). Other than this depression, the surface of
the ramus is fairly flat. Little of the condylar process is visible in lateral view. It is positioned roughly a molar length
above the occlusal plane and is not delimited from the ramus by a penduncle or constricted neck in this view.
In the juveniles AMNH 28272 and 185012, the most
prominent feature on the ramus in medial view is a straight,
rounded prominence that extends from the mylohyoid line
on the body to the condyle dividing the ramus into dorsal and ventral parts (Figs. 37, 42). This prominence is not
horizontal but angled posterodorsally. Slightly posterior to
the midpoint of this prominence in its ventral aspect is the
opening into the mandibular canal, the mandibular foramen (“manf” in Fig. 37). The surface dorsal to this prominence on the medial aspect of the coronoid process is for
the temporalis muscle (Allen 1910). The adults differ from
the juveniles in that there is a stout, posteroventrally directed extension arising from this prominence anterior to
the mandibular foramen (Fig. 40); AMNH 185012 has the
ventral two-thirds of this extension in a weak form (Figs.
37, 40). It extends to the outer contour of the angle and
divides the medial aspect of the angle into subequal anteroventral and posterodorsal surfaces, the latter more deeply
Wible.indd 378
excavated than the former. Allen (1910:16) described the
medial pterygoid (his pterygoideus internus) as “inserting on the angle of the ramus” and the lateral pterygoid
(his pterygoideus externus) as inserting “on the lower jaw
inside the neck of the mandibular condyle forward to the
inferior dental foramen” (mandibular foramen). It is not
entirely clear how Allen’s descriptions relate to the morphology: either the lateral pterygoid occupies the entire
posterodorsal surface or it occupies some smaller subset
thereof with the medial pterygoid in the remainder.
The condylar process is most readily seen in occlusal
and posterior views (Figs. 38, 41). From above it is roughly triangular. The straight lateral side is the lateral surface
of the ramus; the convex posterior side holds the articular surface (“ars” in Fig. 38); and the anteromedial side
is slightly concave. The posterior and anteromedial sides
meet at a sharp, medially directed point. McDowell (1958)
described the articular surface in detail, noting the presence of two ovoid facets, corresponding to external and
entoglenoid [his postglenoid] facets on the squamosal.
The external facet is perpendicular to the plane of the ramus and faces dorsally and slightly posteriorly. The entoglenoid facet is only visible in posterior view because it
lies on the posterior surface of the medially directed point
mentioned above and is directed posteriorly and ventrally
(“ars” in Fig. 37). The posteromedial aspect of the external facet abuts the dorsolateral aspect of the entoglenoid
facet. The facets are present, but subtle in the juveniles
AMNH 28272 and 185012, and in the adults are subequal
Fig. 41.—Solenodon paradoxus, CM 18069 and AMNH 212912,
photographs of right and left mandibles in occlusal view. Scale = 10 mm.
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379
Fig. 42.—Solenodon paradoxus, AMNH 28272, photographs of left mandible in lateral (top) and medial (bottom) views. Third molar has fallen out and
is missing; it is figured in Allen (1908: pl. XXXIII, fig. 5). Scale = 10 mm. Abbreviations: c, lower canine; di2, lower deciduous second incisor; dp5,
lower deciduous third premolar; i1, lower first incisor; i2, crypt containing lower second incisor; i3, lower third incisor; m1, lower first molar; m2, lower
second molar; p1, lower first premolar; p4, lower penultimate premolar; p5, lower ultimate premolar.
in AMNH 28271 and CM 18069, but with the entoglenoid
facet much smaller than the external in AMNH 212912.
Hyoid Apparatus and Larynx
A broken piece of stylohyal (stylohyoid of NAV 1994) is
preserved in situ on the left side of AMNH 185012 and
212912 (“sh” in Figs. 25, 29), with the piece in the former
more substantial that the latter. In both, the posterolateral
end of the stylohyal articulates with the tympanohyal, and
medial to that the stylohyal abuts the posterior crus of the
ectotympanic and the caudal tympanic process of the
petrosal (Fig. 29B). In ventral view (Fig. 25), the broken
stylohyal is L-shaped with the union of the legs representing a small angle of the stylohyal.
J.A. Allen (1908: fig. 9) and G.M Allen (1910: pl. 7,
fig. 3) illustrated the hyoid bones and ossified larynx for
S. paradoxus, but only the latter included description of
these elements. From proximal to the skull base to distal,
Wible.indd 379
the hyoid apparatus includes the paired stylohyals, epihyals, ceratohyals, and the median basihyal, which is fused
to the paired thyrohyals. For completeness sake, the latter
Allen’s figure is reproduced here (Fig. 43) and his descriptions (Allen 1910: 42–43) follow below.
“It remains to describe briefly the laryngeal and hyoid
bones. These appear to be similar to those of S. cubanus as
figured by Peters (1864: pl. 2, fig. 11), and are well ossified. The thyroid is the largest, 13 mm in greatest length.
It is slightly more than half a complete ring and has at
the anterior dorsal margin on each side a process for the
articulation with the tips of the thyrohyals [thyrohyoids of
NAV 1994]. Posteriorly, the dorsal margin is similarly produced to form processes articulating with the posterolateral
margin of the cricoid. A low ridge arises about midway
of the straight dorsal border and curves ventrally to the
posterior edge. Above it on each side is a minute foramen
[Foramen thyroideum of NAV 1994] at the dorsal edge of
the bone. On the left side in our specimen there is in addition a minute foramen about 2 mm anterior to the first. The
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CONCLUSIONS
Fig. 43.—Solenodon paradoxus, line drawing of hyoid apparatus and ossified larynx in oblique left lateral view such that some paired elements
are visible bilaterally (redrawn and relabeled from Allen 1910: pl. 7, fig.
3). Median basihyal is fused to paired thyrohyals. Abbreviations: ashy,
angle of stylohyal; bhy, basihyal; chy, ceratohyal; crc, cricoid; ehy, epihyal; f, foramen; sh, stylohyal; thd, thyroid; thfc, tympanohyal facet on
stylohyal; thy, thyrohyal;
thyrohyal of each side has fused ventrally with the basihyal [basihyoid of NAV 1994] so that the three bones
thus form a half ring, bowed back at first, then forward
at the ventral side. The ceratohyals [ceratohyoids of NAV
1994] are appressed against the ventrolateral margin of
the ring. They are rather thick and about 4 mm in dorsoventral length. Their dorsal border, and the edge of the
thyrohyal adjacent, articulate on each side with the epihyal [epihyoid of NAV 1994], a broad but laterally flattened bone, that projects anteriorly from this articulation.
This in turn joins with the stylohyal, which is about 2 mm
longer, and much more rounded and slender. It joins the
skull by a very short bony process that projects at nearly
right angles from its proximal end. This process may be
fused to the tympanohyal [tympanohyoid of NAV 1994].
The cricoid at its anterior end is clasped by the converging
posterior processes of the thyroid and is a complete bony
ring, with a postero-dorsal extension. The vocal cords are
attached by cartilage, one at each side from the anterior
apex of this ring to its mid-ventral line, and pass forward
as a delicate strand to a median attachment just back of
the anterior edge of the thyroid. The first tracheal ring is
the broadest and fits into the posterior end of the cricoid,
to which it is bound by muscle fibers. Peters states that
in S. cubanus the first nine tracheal rings are complete
and that there are 21 in all. In S. paradoxus the number is
slightly more, 22 to 29, and all are incomplete dorsally.”
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The final arbiter of the significance of the detailed anatomy
presented above is phylogenetic analysis, which is well beyond the scope of this report. In the literature, a term frequently encountered in conjunction with the morphology
of the solenodon is “archaic.” For example, according to
Eisenberg (1981:114), solenodons “possess very archaic
morphological characters in conjunction with some apomorphic characters”, although he did not specify any of
either type. Possibly accounting for the purported archaic
(plesiomorphic) nature of the solenodon, molecular estimates place its divergence from other placentals at 76 million years ago (Roca et al. 2004). Does the solenodon skull
possess plesiomorphic characters lost in more derived
placentals?
Gregory’s (1910:253) monograph on the orders of
mammals provides the most detailed discussion of this
subject, albeit in a pre-cladistic paradigm. In his “phyletic
interpretation of the osteological characters of Solenodon
paradoxus,” he noted “primitive placental [eutherian]
characters,” “points of special resemblance to marsupials [metatherians],” and “primitive marsupio-placental
[metatherian-eutherian] characters” for the solenodon.
Skulls of Cretaceous therians were unknown in 1910, and
significant advances have occurred in that front in recent
years (Kielan-Jaworowska et al. 2004; Wible et al. 2005b).
Current status of the features named by Gregory is reviewed among basal eutherians and metatherians in order
to assess the supposed archaic nature of the solenodon.
The primitive eutherian characters highlighted by Gregory (1910) included: “Orbitosphenoid pierced by optic foramen and not depressed dorsoventrally in such a way that
the opposite sphenorbital fissures are confluent beneath it
(contrast Marsupials); alisphenoid relatively small and not
entering the glenoid fossa.” Both of these characters are
widely distributed among placentals and do not distinguish
the solenodon as plesiomorphic. An optic foramen with an
orbitosphenoid not depressed dorsoventrally occurs in the
few Cretaceous eutherians for which the orbitosphenoid is
well preserved (Wible et al. 2005b) and also in nearly all
extant placentals. In contrast, metatherians and the Early
Cretaceous prototribosphenidan Vincelestes Bonaparte,
1986, lack the optic foramen and have confluent sphenorbital fissures (Rougier 1993; Rougier et al. 1998); the only
exception is the Late Cretaceous metatherian Deltatheridium Gregory and Simpson, 1926, which has an optic
foramen (Wible et al. 2005b), although the presence of a
dorsoventrally depressed orbitosphenoid is unknown. Absence of a glenoid process of the alisphenoid is primitive
for eutherians, but is even more widely distributed, being
also absent in Vincelestes and Deltatheridium; the addition
of this process is a feature occurring within Metatheria
(Rougier et al. 1998).
The second category highlighted by Gregory (1910),
special resemblances (convergences) to metatherians, included: “Vomer with larger lateral posterior wings, joining
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Wible—On the Cranial Osteology of Solenodon paradoxus
ethmoid scroll. Large venous condylar foramina [condyloid canal]. A remnant of the tympanic wing [tympanic
process] of the alisphenoid. Posterior mental foramen beneath m1.” (1) The status of the vomer is known in so few
taxa that commentary is meaningless at this time. (2) The
solenodon does share a large venous condyloid canal with
marsupials, but it is not unique among placentals in that
regard (e.g., dog, Evans 1993; ox, Sisson 1910). Moreover,
a condyloid canal is absent in basal metatherians, such
as Pucadelphys Marshall and Muizon, 1988 (Marshall et
al. 1995) and Mayulestes Muizon, 1994 (Muizon 1998),
as well as in basal eutherians, such as Zalambdalestes
(Wible et al. 2004) and Maelestes (Wible et al. in press).
(3) The solenodon does have a small tympanic process of
the alisphenoid, which is widely distributed among marsupials (Horovitz and Sánchez-Villagra 2003), but it is not
unique among placentals in that regard (e.g., Erinaceus
europaeus Linnaeus, 1758, Potamogale velox (du Chaillu,
1860); Wible et al. 2007, in press). Moreover, a tympanic
process is absent in basal metatherians, such as Pucadelphys (Marshall et al. 1995) and Mayulestes (Muizon 1998),
and its addition is a synapomorphy of Marsupialia (Horovitz and Sánchez-Villagra 2003). (4) The solenodon does
share a posteriormost mental foramen beneath the lower
first molar with metatherians (Rougier et al. 1998; Wible
et al. 2007, in press), but this occurs in other eutherians
(e.g., Late Cretaceous Maelestes, Blarina brevicauda (Say,
1823), Potamogale velox; Wible et al. 2007, in press).
The third category highlighted by Gregory (1910),
primitive therian characters, was the longest by far and
included: “Interparietal paired; orbitosphenoid invaded anteriorly by ethmoid chamber; pituitary depression slight,
without posterior clinoid process; basioccipital very short.
Alisphenoids with prominent palatine flanges [entopterygoid processes]. Posterior border of palate ridged [postpalatine torus]. General arrangement of ethmoids and
maxillo-turbinals. Tympanic ring-shaped, oblique rather
than vertical. Arrangement of following nerve foramina:
olfactory, internal auditory [acoustic] meatus, facial canal,
fenestra ovalis [vestibuli], foramen lacerum anterius [sphenorbital fissure] and posterius [jugular foramen], sphenopalatine, post-palatine [minor palatine] stylomastoid, and
condylar [hypoglossal] foramina (the last occasionally
double). Venous foramina: transverse canal [foramen] in
basisphenoid (cf. Didelphis), ?sinus canal in temporal region [anterior opening, orbitotemporal canal], post-parietal
[foramen for ramus temporalis], post-squamosal [foramen
for ramus temporalis], post-glenoid [postglenoid], postmastoid [mastoid] foramina.”
Commentary on several features in this category is not
appropriate in light of our poor state of knowledge (i.e., orbitosphenoid invaded anteriorly by ethmoid chamber; general arrangement of ethmoids and maxillo-turbinals; olfactory foramina). The remainder can be divided into features
for which subsequent research has shown Gregory’s polarity to be incorrect, uncertain, or correct.
Five features thought by Gregory to be primitive for
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381
Theria are no longer so considered. (1) Rather than primitively paired in therians, the interparietal is wholly absent
in Cretaceous eutherians and basal metatherians (Rougier
et al. 1998; Wible et al. 2007, in press). Additionally, when
an interparietal occurs in placentals, it is usually a single
ossification and not paired (Novacek 1993); the paired interparietal of the solenodon is not unique (De Beer 1937),
but it is unusual. (2) The oval fenestra vestibuli of the
solenodon does not resemble the rounder version of this
aperture that occurs in basal metatherians and most basal
eutherians and is clearly the primitive condition for Theria (Rougier et al. 1998; Wible et al. 2007, in press). (3)
The solenodon sphenopalatine foramen with a contribution
from the ethmoid to its bony border does not resemble the
condition in basal eutherians and metatherians in which the
opening is either within the palatine or between the palatine
and maxilla (Wible et al. 2007, in press). (4) The solenodon stylomastoid notch with a contribution from the lateral
section of the caudal tympanic process of the petrosal does
not resemble the more open notch between only the tympanohyal and posterior continuation of the crista parotica
of basal eutherians and metatherians (Rougier et al. 1998;
Wible et al. 2001, 2007, in press). (5) The transverse canal
foramen in the basisphenoid is absent in Cretaceous eutherians (Wible et al. 2007, in press) and is a synapomorphy
within Metatheria (Horovitz and Sánchez-Villagra 2003).
The presence of a transverse canal in the solenodon is not
unique among placentals (e.g., Euphractus sexcinctus (Linnaeus, 1758), Wible and Gaudin 2004), but is unusual.
The shape and orientation of one element, the ectotympanic, is of uncertain polarity for Theria, because this element is poorly known in basal eutherians and metatherians.
Some forms (e.g., Maelestes) have a ring-shaped ectotympanic resembling that in the solenodon, whereas others
(e.g., Zalambdalestes) have a slightly expanded, fusiform
ectotympanic (Wible et al. 2007, in press). Even less is
known about the inclination of the ectotympanic in basal
therians than its form.
Gregory inferred the correct polarity for Theria for the
remaining features in the third category. However, there is
little here that distinguishes the solenodon as plesiomorphic when compared to other placentals. In fact, for five
features, the solenodon has some additional condition that
sets it apart from most placentals and/or basal eutherians.
(1) Little is known of the pituitary depression (hypophyseal fossa) in basal therians. Late Cretaceous Maelestes
and Ukhaatherium Novacek et al., 1997, have a shallow
fossa with a low dorsum sellae (Rougier et al. 1998; Wible
et al. 2007, in press) resembling that in the solenodon. Yet,
in both fossils and in placentals in general, the hypophyseal
fossa is within the basisphenoid. The solenodon is unusual
(perhaps even unique) in having a significant portion of
the hypophyseal fossa formed within the basioccipital. (2)
The solenodon has prominent entopterygoid processes as
in basal therians (Wible et al. 2005b, 2007, in press). However, it lacks the ectopterygoid processes that distinguish
the alisphenoids of basal eutherians and many placentals
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from those of metatherians (Wible et al. 2005b, 2007, in
press). (3) The solenodon minor palatine foramen with its
narrow bony bridge forming the posterior border resembles
the foramen that occurs widely in basal therians (Rougier
et al. 1998; Wible et al. 2007, in press). However, the solenodon aperture is variably subdivided into two ventral
apertures. (4) As in the solenodon, two or more hypoglossal foramina are present in Vincelestes (Rougier 1993),
basal metatherians and most basal eutherians (Wible et al.
2007, in press). However, the solenodon is distinguished
by having additional small foramina adjacent to the hypoglossal foramina that connect to the condyloid canal. (5)
The stapedial artery system of the solenodon includes a
well-developed anterior division of the ramus superior that
enters the orbit via the anterior opening of the orbitotemporal canal, as in basal eutherians, such as asioryctitheres and
zalambdalestids (Wible et al. 2004, 2007, in press).
However, the solenodon is distinguished by the close association of the anterior opening of the orbitotemporal
canal with the posterodorsal ethmoidal foramen and the
frontal diploic vein foramina, which is unknown in other
eutherians.
A recent phylogenetic analysis of morphological characters that includes the solenodon along with Cretaceous
eutherians by the author and colleagues (Wible et al. 2007,
in press) does not identify the solenodon as an archaic form
but nested well within Placentalia. Despite the general
conception of some, such as Eisenberg (1981), regarding
the archaic nature of the solenodon, its cranial osteology
presents few plesiomorphic features lost in more derived
placentals. The most striking of these, which is also previously unreported for the solenodon, is the presence of
a prootic canal for the lateral head vein. A prootic canal
is widely distributed among Mesozoic mammals, but only
among a few Cretaceous eutherians (Wible et al. 2001,
2007, in press; Ekdale et al. 2004) and no other placentals.
The unexpected discovery of this structure in the solenodon should alert others to be on the lookout for a prootic
canal in other placentals.
The overall form of the solenodon skull may appear to
be archaic with its elongated snout, small eyes, and tubular
braincase. However, rather than archaic, the cranial osteology described above paints a picture of an animal with
a highly specialized head with numerous unique adaptations. From the os proboscidis in front, to the numerous
venous foramina in the orbit (frontal diploic vein, supraoptic, and suboptic foramina), to the mallear articulation
on the petrosal in the ear region, to the numerous venous
foramina on the occiput, to the extra “angular” process
on the lower jaw, the solenodon is one peculiar beast.
ACKNOWLEDGMENTS
As a graduate student years ago, three outstanding monographs incorporating observations on the skull of the solenodon had a tremendous influence on me: Gregory (1910), McDowell (1958), and MacPhee (1981). In
the course of this project, I discovered several more amazing works on
Wible.indd 382
this unusual animal, in particular Brandt (1833), Peters (1864), and Allen
(1910). My first acknowledgment is to these researchers who motivated
me in this project. However, my greatest debt in this project is to Paul
Bowden, whose outstanding illustrations inspired me to get the anatomy
correct; Paul meticulously completed all the artwork, while I am responsible for the photographs. I am also extremely grateful to the curators in the
Department of Mammalogy of the American Museum of Natural History:
Ross MacPhee, Nancy Simmons, and Rob Voss. Without their willingness
to loan me the crucial skulls from their collection, this project would not
have been possible. For various favors aiding the completion of this project, I thank Pat Brunauer and Teresa Pacheco at the American Museum of
Natural History; Sue McLaren, Tim McCarthy, and Xianghua Sun at the
Carnegie Museum of Natural History; and Tim Smith at Slippery Rock
University. Thorough, insightful reviews by Guillermo Rougier and Norberto Giannini greatly improved the final product and helped me avoid
some embarrassments. This work was supported by Carnegie Museum of
Natural History and National Science Foundation ATOL grant 0629959.
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Wible, J.R., Y.-Q. Wang, C.-K. Li, and M.R. Dawson. 2005a. Cranial
anatomy and relationships of a new ctenodactyloid (Mammalia,
Rodentia) from the early Eocene of Hubei Province, China. Annals
of Carnegie Museum, 74:91–150.
Zeller, U. 1987. Morphogenesis of the mammal skull with special reference to Tupaia. In Morphogenesis of the Mammalian Skull (H.-J.
Kuhn and U. Zeller, eds.). Mammalia Depicta, 13:17–50.
———. 1989. Die Entwicklung und Morphologie des Schädels von
Ornithorhynchus anatinus (Mammalia: Prototheria: Monotremata).
Abhandlungen der Senckenbergischen Naturforschenden
Gesellschaft, 545:1–188.
Ziegler, A.C. 1971. A theory of the evolution of therian dental formulas and replacement patterns. Quarterly Review of Biology,
46:226–249.
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Glossary
In keeping with previous works by the author and collaborators (e.g., Wible 2003, 2007; Wible and Gaudin 2004;
Giannini et al. 2006), below in alphabetical order is a
treatment of 66 cranial structures (e.g., foramina, grooves,
canals) associated with soft tissues (e.g., nerves, arteries,
veins). The vast majority occurs in the solenodon, and the
few that do not are included for the sake of completeness.
As the point of reference, nearly every entry begins with
the condition in the dog, because an excellent, well-illustrated textbook of canine anatomy exists (Evans 1993) and
because the skull of the dog is more generalized than other
well-documented taxa (e.g., humans, horses, rats).
Two oft-cited monographs including observations on
the skull and basicranium of the solenodon respectively
are McDowell (1958) and MacPhee (1981). A common
denominator of these works is a glossary of cranial and
basicranial structures, which inspired the following.
Accessory Palatine Foramen.—In the dog (Evans 1993),
one or more accessory palatine foramina (following Wible
and Rougier 2000) (=minor palatine foramina of Evans
1993) in the horizontal process of the palatine transmit accessory palatine nerves to the mucosa of the hard palate.
These nerves arise from the major palatine nerve within the
palatine canal (Evans 1993). In the solenodon, numerous
openings occur in the palatomaxillary suture and the palatine bone on the back of the hard palate (Figs. 11–13). The
exact contents are not known, but these openings are either major or accessory palatine foramina. Until additional
study is done, those posterior to the anterior palatomaxillary suture are treated as accessory palatine foramina. Considerable variation occurs within and between specimens
in the number and location of accessory palatine foramina.
The right side of AMNH 28271 has the fewest with two
entirely within palatine, and the left side of AMNH 212912
and right side of AMNH 185012 the most with four in the
palatine and one in the palatomaxillary suture. Based on
the bisected skull AMNH 28271, the accessory palatine
nerves have a separate small foramen of exit from the nasopharyngeal meatus opposite the upper ultimate molar
(“apf” in Fig. 22).
Alisphenoid Canal.—In the dog (Evans 1993), the
alisphenoid canal (following Gregory 1910) (=alar canal
of Evans 1993) runs through the base of the alisphenoid
(=temporal wing of the sphenoid of Evans 1993); its smaller caudal opening transmits the maxillary artery and vein
and its larger rostral opening the maxillary artery, vein, and
nerve. The maxillary nerve enters the canal via the foramen rotundum in the canal’s roof. In the solenodon, the
alisphenoid canal lies anterior to the foramen ovale and
transverse canal foramen within the alisphenoid (“asc” in
Figs. 14–16) and is merely a foramen of entry into the skull
(=caudal alar foramen of the dog) rather than a canal. Gregory (1910:246) suggested that the solenodon alisphenoid
canal transmitted a branch of the external carotid artery
Wible.indd 386
(=ectocarotid artery of Gregory 1910); McDowell (1958),
citing Gregory (1920a), identified the ramus inferior of the
stapedial artery as the primary occupant. MPIH 6863 does
not preserve sections rostral to the foramen ovale, but the
major artery below that opening is the ramus infraorbitalis,
a primary branch of the ramus inferior of the stapedial artery (Wible 1987; Fig. 26C). Consequently, it seems more
likely that the ramus infraorbitalis is the primary occupant
of the alisphenoid canal.
Alveolar Foramina.—In the dog (Evans 1993:150), “leading from the infraorbital canal to the individual roots of the
premolar teeth (first four cheek teeth) are the alveolar canals
(canales alveolares), which open by numerous alveolar foramina (foramina alveolaria) in the apex of each alveolus.
The special incisivomaxillary canal (canalis maxilloincisivus) carries the nerves and blood vessels to the first three
premolar and the canine and incisor teeth. It leaves the medial wall of the infraorbital foramen, passes dorsal to the
apex of the canine alveolus with which it communicates,
and enters the incisive bone [=premaxilla].” In the solenodon, the usual pattern is two primary alveolar foramina: a
larger anterior one dorsomedial to the anterolabial root of
the upper ultimate premolar, anterior to the infraorbital foramen (“imc” in Fig. 6) and a smaller posterior one within
the infraorbital canal (“af” in Fig. 10). In AMNH 185012,
the former foramen can be traced through the thin bone
nearly to the premaxilla (Fig. 7) and, therefore, must be
equivalent to the incisivomaxillary canal of the dog.
Carotid Foramen and Sulcus.—In the dog (Evans 1993),
the internal carotid artery crosses the basicranium within
a perbullar canal (sensu Wible 1986) through the medial
wall of the auditory bulla. Three foramina are associated
with this course: one at the caudal entrance into the carotid
canal, a second at the rostral exit from the carotid canal,
and a third at the entrance into the cranial cavity between
the petrosal (=pars petrosa of the temporal bone of Evans
1993) and basisphenoid (for discussion of the terminology
of these foramina see Wible and Gaudin 2004:149). Within
the cranial cavity, the artery runs in a shallow, anteromedially directed carotid sulcus in the basisphenoid, posterolateral to the hypophyseal fossa. In the solenodon, the internal carotid artery (=entocarotid artery of Gregory 1910;
arteria promotorii of McDowell 1958) crosses the basicranium in a transpromontorial position (sensu Wible 1986)
in a sulcus on the pars cochlearis of the petrosal (“gica” in
Figs. 27A, 28; “ica” in Fig. 26C). It enters the cranial cavity either via the medial aspect of the piriform fenestra in
AMNH 28271, 28272 (Fig. 26C), 185012 (right side), and
CM 18069 (left side) or a separate carotid foramen in the
basisphenoid in the medial aspect of the piriform fenestra
in AMNH 185012 (left side, “cf” in Fig. 25), 212912, and
CM 18069 (right side). Within the cranial cavity is a faint,
anteromedially directed carotid sulcus on the basisphenoid,
anterolateral to the hypophyseal fossa (“gica” in Fig. 22).
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Cavum Epiptericum.—see Pterygoid Canal.
Caudal Palatine Foramen.—In the dog (Evans 1993),
the caudal palatine foramen for the major palatine nerve,
artery, and vein is separate from the sphenopalatine foramen, with both foramina in the perpendicular process of
the palatine. In the solenodon, a separate caudal palatine
foramen is lacking, and its contents pass through the sphenopalatine foramen (“spf” in Figs. 10, 15).
situated just rostral to the center of the basisphenoid interpreted as a craniopharyngeal canal (“cpc” in Figs. 25,
27A); in the adults this is reduced to a miniscule slit. A
craniopharyngeal canal does not occur in the available
sections of MPIH 6863, but only the posterior basisphenoid is represented. In AMNH 28272, Gregory (1910:245;
“f.ch.d.” in Fig. 1) identified a small midline hole between
the basi- and presphenoid as possibly lodging “a ventral
apophysis of the vestigial chorda dorsalis [=notochord]”,
but labeled in his figure is the open intersphenoidal synchondrosis.
Cochlear Canaliculus.—In the dog (Evans 1993), the
perilymphatic duct reaches the inner ear from the cranial
cavity via the external opening of the cochlear canaliculus
in the petrosal (=pars petrosa of the temporal bone of Evans 1993) in the rostral edge of the jugular foramen. The
jugular foramen is deeply recessed and partially obscured
by the auditory bulla, but the ventral edge of the aperture
can be seen from the ventral side (e.g., CM 30471). In the
solenodon, the external opening of the cochlear canaliculus (=aqueductus cochleae of McDowell 1958) is similarly
situated in the rostral edge of the jugular foramen with its
ventral edge visible from the ventral side (not visible in
the figures).
Cribroethmoidal Foramen.—In the dog (Evans 1993),
the ethmoidal nerve passes through an unnamed opening
in the cribriform plate to reach the nasal cavity. In humans
(Terry 1942), the anterior ethmoidal nerve enters the nasal
cavity via the ethmoidal fissure, a slit-like opening at the
side of the crista galli; the posterior ethmoidal nerve does
not enter the nasal cavity, but is distributed to the posterior
ethmoidal air cells and sphenoidal sinus. In the solenodon
(AMNH 28271), a large opening in the dorsomedial aspect of the cribriform plate (“cef” in Fig. 23) medial to
foramina to the nasoturbinate is interpreted here as for the
ethmoidal nerves. Following Moore (1981), this opening is
the cribroethmoidal foramen.
Condyloid Canal.—In the dog (Evans 1993:136), “the
rather large condyloid canal...runs through the medial part
of the lateral occipital bone [=exoccipital]. There is an intraosseous passage between the condyloid canal and the
hypoglossal canal. Usually there is also a small passage
between the condyloid canal and the petrobasilar fissure.”
The condyloid canal transmits the condyloid vein, which
connects the sigmoid sinus and the basilar sinus. In the
solenodon, the large condyloid canal (=venous condylar
foramen of Gregory 1910) differs in that its primary exit is
not endocranial. In the bisected skull, AMNH 28271, the
condyloid canal begins at several large openings at the terminus of the sigmoid sinus sulcus (“coca” in Fig. 21C) and
ends at the dorsal and ventral condyloid foramina (lacking in the dog) and hypoglossal foramina (“vcf” and “hf”
in Fig. 26B; “dcf” in Figs. 34, 35). In light of the size of
the condyloid canal, it represents the primary exit for the
sigmoid sinus; the contents of the canal are unknown for
MPIH 6863 because of lack of preservation. Two specimens, AMNH 212912 and CM 18069, also have a small
endocranial opening just internal to the foramen magnum
resembling the large caudal opening of the condyloid canal
in the dog.
Dorsal Condyloid Foramen.—In the solenodon, the dorsal condyloid fossa dorsal to the occipital condyle on the
exoccipital’s vertical process on the occiput has at least
one well-developed foramen in its medial margin that connects to the condyloid canal (“dcf” in Figs. 34, 35). AMNH
28271, 28272 (left side), 185012 (left side), and CM 18069
(right side) have an additional small opening; and AMNH
28272 (right side) has two small openings. In the dog (Evans 1993), dorsal condyloid foramina are not reported or
illustrated.
Cavum Supracochleare.—see Facial Canal and/or
Sulcus.
Craniopharyngeal Canal.—In the dog (Evans 1993:139),
in the basisphenoid “occasionally the small craniopharyngeal canal...persists in the adult, particularly in Bulldogs.
This midline canal is a remnant of the pharyngeal diverticulum to the hypophyseal fossa from which the pars glandularis of the hypophysis develops.” The juvenile solenodons
AMNH 28272 and 185012 have a visible midline foramen
Wible.indd 387
387
Ethmoidal Foramen.—In the dog (Evans 1993), two
ethmoidal foramina usually occur per side, a larger posterodorsal one in the orbital process of the frontal for
the external ethmoidal artery and vein, and a smaller anteroventral one in the suture between the frontal and orbitosphenoid for the ethmoidal nerve. Endocranially, these
foramina open posterolateral to the cribriform plate, with
the anteroventral one between the orbitosphenoid (=orbital
wing of the presphenoid of Evans 1993) and ethmoid; the
disposition of the posterodorsal foramen is not reported by
Evans (1993). In CM 25053, both ethmoidal foramina are
entirely within the frontal both extracranially and endocranially. The solenodon also has two ethmoidal foramina, a
smaller anteroventral one within the frontal and a larger
posterodorsal one between the frontal and orbitosphenoid,
based on AMNH 28272 (“aef” and “pef” in Figs. 16, 19).
The latter foramen is within a common depression with
the anterior opening of the orbitotemporal canal, except in
AMNH 212912. Endocranially, both foramina open into a
sulcus posterolateral to the cribriform plate of the ethmoid
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(“aef” and “pef” in Fig. 22); the bony disposition of these
foramina is uncertain.
Facial Canal and/or Sulcus.—In the dog (Evans 1993),
the course of the facial nerve through the middle ear is
entirely within a bony canal within the petrosal (=pars
petrosa of the temporal bone of Evans 1993) between the
cavum supracochleare (following Voit 1909) (=genu of the
facial canal of Evans 1993), the space housing the geniculate ganglion of the facial nerve, and the stylomastoid foramen. In the solenodon, based on MPIH 6863, the facial
nerve occupies a sulcus on the pars canalicularis of the petrosal (“fs” in Figs. 25, 26B, 28) between its exit from the
cavum supracochleare, the secondary facial foramen, and
the stylomastoid notch. In the segment of the facial sulcus
anterior to the fenestra vestibuli, the stapedial artery runs
ventral to the facial nerve. In the short segment posterior
to the fenestra vestibuli, the facial nerve runs medial to the
lateral head vein and then leaves the middle ear at the stylomastoid notch passing ventrolateral to the vein.
Foramen for Frontal Diploic Vein.—In the dog (Evans
1993), the frontal diploic vein, an emissary vein of the
diploë of the frontal bone to the dorsal external ophthalmic vein, leaves the skull via a small unnamed foramen in
the orbital surface of the postorbital process of the frontal (=zygomatic process of the frontal of Evans 1993).
Rather than a small foramen near the supraorbital rim as in
the dog, the solenodon has one or more, larger openings,
within the frontal in AMNH 28272, more ventrally placed
near the posterodorsal ethmoidal foramen and the anterior
opening of the orbitotemporal canal and/or the anteroventral ethmoidal foramen (“fdv” in Figs. 14, 16, 19). Based
on the bisected skull AMNH 28271, the frontal diploic
vein runs between the inner and outer table of the frontal
bone in the annular ridge, which demarcates the rostral and
middle cranial fossae, and communicates across the dorsal
midline (“fdv” in Fig. 21C). Gregory (1910) employed the
term supra-ethmoid foramen for the frontal diploic vein
foramen (Fig. 1).
Foramen for Inferior Petrosal Sinus.—In the dog (Evans 1993), the inferior petrosal sinus (=ventral petrosal sinus of Evans 1993) connects the cavernous sinus with the
internal jugular vein via the petrobasilar (petro-occipital)
canal, between the petrosal (=pars petrosa of the temporal
of Evans 1993) and the basioccipital (=basilar part of the
occipital of Evans 1993). Accompanying the inferior petrosal sinus is a condyloid artery off the occipital artery. The
foramen for inferior petrosal sinus (=posterior opening of
the petro-occipital canal of Evans 1993) lies anterior to the
jugular foramen. In the solenodon, based on MPIH 6863,
the inferior petrosal sinus occupies a groove on the medial
face of the pars cochlearis of the petrosal (“ips” in Fig.
25). Dorsal to the groove, the pars cochlearis overlaps the
basioccipital, forming a roof over the groove. There is also
a partial floor to the groove formed by a lamina on the pars
Wible.indd 388
cochlearis that varies in length; it runs the length between
the anterior pole and jugular foramen in AMNH 185012
(Fig. 25) but is confined to the rostral half of the pars
cochlearis in CM 18069 and AMNH 28271 and 212912.
The foramen for the inferior petrosal sinus occurs at the
posterior end of the lamina on the pars cochlearis.
Foramen for Ramus Superior.—In the solenodon (McDowell 1958), a separate foramen for the ramus superior
of the stapedial artery is lacking and the artery enters the
cranial cavity via the piriform fenestra (=pyriform fenestra of McDowell 1958) (“rs” in Fig. 26C), confirmed by
MPIH 6863 (see also MacPhee 1981). The dog lacks the
ramus superior of the stapedial artery and foramen.
Foramen Magnum.—In the dog (Evans 1993), the foramen magnum is enclosed between the basi-, ex-, and supraoccipital (=basilar, lateral, and squamous parts of the
occipital bone of Evans 1993), which are fused together in
the adult. The solenodon has the same arrangement. The
sectioned juvenile MPIH 6863 preserves the suture between the basi- and exoccipital and reveals that the basioccipital forms the odontoid notch and part of the adjacent
occipital condyles on the anterior rim of the foramen magnum, and AMNH 28272 preserves the suture between the
ex- and supraoccipital and reveals that the supraoccipital
forms the dorsal rim of the foramen magnum (“fm” in Fig.
35). The remaining specimens have a single occipital bone
rimming the foramen magnum. The foramen magnum is
elliptical, wider than high (Figs. 34, 35).
Foramen Ovale.—In the dog (Evans 1993), the foramen
ovale is in the base of the alisphenoid (=temporal wing of
the basisphenoid of Evans 1993) and transmits the mandibular nerve and a small emissary vein; a small notch or
even a separate foramen spinosum for the middle meningeal artery may be present in the posterolateral border of
the foramen ovale. In the solenodon, the foramen ovale is
in the alisphenoid immediately anterior to a low alisphenoid tympanic process (“fo” in Figs. 25, 27A). In MPIH
6863, the mandibular nerve is the primary occupant, but
also transmitted are small veins and a small meningeal artery off the ramus mandibularis (“mnb” in Fig. 26C); the
ramus inferior of the stapedial artery divides into the ramus infraorbitalis and ramus mandibularis posteroventral
to the foramen ovale (Fig. 26C).
Foramen Rotundum.—In the dog (Evans 1993), the maxillary nerve and a small emissary vein exit the cranial cavity via a foramen rotundum in the base of alisphenoid (=the
temporal wing of the basisphenoid of Evans 1993) and
then exit the skull via the rostral opening of alisphenoid
canal (=rostral alar foramen of Evans 1993). In the solenodon, a separate foramen rotundum for the maxillary nerve
is lacking and, according to Gregory (1910) the nerve exits
via the foramen lacerum anterius (=sphenorbital fissure)
(“sof” in Figs. 14, 16).
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Foramen for Ramus Temporalis.—In the solenodon, the
examined adult skulls have between three and six foramina
in the lateral braincase wall identified following MacPhee
(1994) as foramina for rami temporales (including the
post-parietal and post-squamosal foramina of Gregory
1910) for arterial and venous blood to the overlying temporalis muscle (“rt” in Figs. 2, 5, 6). The more posterodorsal of these foramina have well-developed grooves extending posterodorsally and dorsally into the temporal fossa.
Because the parietosquamous suture is fused in the adults
(Fig. 7), the exact bony location of these foramina is uncertain. In the juveniles, which retain the parietosquamous
suture, some foramina are in the parietosquamous suture,
others within the squamosal near the parietosquamous suture, and lastly one constant foramen within the squamosal
well away from the suture (Fig. 6). AMNH 28272 has two
foramina in the suture per side and two near the suture on
the left side only; AMNH 185012 has one in the left suture,
three in the right suture, and one near the suture on both
sides, with the one on the left minute. In contrast, the dog
has no foramina for rami temporales and the blood supply
to the temporalis muscle derives from the external carotid
system (Evans 1993).
Glaserian Fissure.—According to Klaauw (1931:164),
during ontogeny, the Glaserian fissure forms in the anterior wall of the presumptive auditory bulla as an aperture
for Meckel’s cartilage, which subsequently disappears and
“later on we find the chorda tympani nerve in it and often
also the ramus inferior of the stapedial artery.” In the dog,
the ramus inferior is lacking (Tandler 1899; Bugge 1978)
and the chorda tympani nerve leaves the middle ear anteriorly “through a small canal in the rostrodorsal wall of
the tympanic [=ecto- + entotympanic] bulla, and emerges
through the petrotympanic fissure to join the lingual nerve
(Evans 1993:978).” This small unnamed canal is the Glaserian fissure. In the solenodon, the chorda tympani and ramus inferior do not leave the middle ear together in MPIH
6863 as noted by MacPhee (1981), and it is the course of
the former that marks the Glaserian fissure. Based on the
specimens with the ectotympanic in place (AMNH 28271
[left side], 185012 [right side], 212912, CM 18069 [right
side]), the chorda tympani nerve leaves the middle ear medial to the postglenoid foramen via a small opening between the rostral process of the malleus, anterior crus of the
ectotympanic, and entoglenoid process of the squamosal.
It then runs anteroventrally in a sulcus on the entoglenoid
process (“gct” in Figs. 25, 26B), the groove for the chorda
tympani of McDowell (1958).
Groove for Chorda Tympani Nerve.—see Glaserian
Fissure.
Hiatus Fallopii.—In the dog (Evans 1993), the greater
petrosal nerve (=major petrosal nerve of Evans 1993) runs
forward from the geniculate ganglion of the facial nerve in
the small petrosal canal in the petrosal (=pars petrosa of
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389
the temporal bone of Evans 1993) dorsal to the fossa for
the tensor tympani muscle. The hiatus Fallopii (=anterior
opening of the petrosal canal of Evans 1993) is in the petrosquamous suture. In the solenodon, the hiatus Fallopii is
also dorsal to the fossa for the tensor tympani muscle, but
anterior to the hiatus is the broad piriform fenestra. The
position of the opening in mammals has been characterized
(e.g., Sánchez-Villagra and Wible 2002) based on the relationship between its floor and roof (dorsal with floor longer than roof, intermediate with roof and floor subequal,
or ventral with roof longer than floor). All three conditions
are exhibited in the solenodon specimens examined (for
location see “hF” in Fig. 27A; nerve is reconstructed in
Fig. 27B).
Hypoglossal Foramen.—In the dog (Evans 1993), the hypoglossal foramen, the external opening of the hypoglossal
canal, is in the exoccipital (=lateral part of the occipital
of Evans 1993) posterolateral to the jugular foramen and
transmits the hypoglossal nerve and vein. In the solenodon, two hypoglossal foramina (=condylar foramina of
Gregory 1910) occur per side (“hf” in Figs. 25, 26B) except for the right side of AMNH 185012, which has three
(all three are not visible in direct ventral view in Fig. 25).
The exact contents of the hypoglossal foramina cannot be
verified, because MPIH 6863 does not preserve them. The
position and size of the foramina relative to each other varies among the examined skulls. In the adults, the foramina
open into a common depression, but this only occurs on
the right side of AMNH 185012 in the two juveniles (Fig.
25). All have small foramina adjacent to the hypoglossal
foramina that connect to the condyloid canal.
Incisive Foramen.—In the dog (Evans 1993), the beanshaped incisive foramen (=palatine fissure of Evans 1993)
is in the premaxilla (=incisive bone of Evans 1993) with
the maxilla forming the posterior border roughly medial
to the upper canine. It is longer than the alveolus for any
of the three upper incisors. The incisive foramen transmits
the nasopalatine duct connecting the oral and nasal cavities
with the vomeronasal organ, the rostral septal branch of the
major palatine artery, and the nasopalatine nerve (=septal
branch of the caudal nasal nerve of Evans 1993). In the solenodon, the oval incisive foramen (“inf” in Fig. 11) is almost completely in the premaxilla, with the maxilla forming the narrow posteriormost border, and is at the level of
the diastema between the first and second incisors, with
length approximating that of the upper third incisor alveolus, the smallest of the three upper incisors. Asher (2001)
reports the nasopalatine duct connecting the nasal and oral
cavities via the incisive foramen in a serially sectioned S.
paradoxus identified as neo/postnatal in age and also that
the vomeronasal duct opens directly into the nasal fossa
and not into the nasopalatine duct as in the dog.
Incisivomaxillary Canal.—see Alveolar Foramina.
Infraorbital Canal.—In the dog (Evans 1993), the
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infraorbital canal transmits the infraorbital nerve, artery,
and vein from the orbit to the snout and is roughly the
length of the enlarged upper carnassial tooth (P5), to which
it is dorsal; in CM 30471, the P5 is 30% longer than the
M1. In the solenodon, the infraorbital canal is relatively
shorter, slightly less than the length of the upper first molar
(M1), to which it is dorsal (Figs. 6, 7).
Infraorbital Foramen.—In the dog (Evans 1993), the elliptical infraorbital foramen, the anterior opening of the infraorbital canal, is within the maxilla on the snout dorsal to
the penultimate upper premolar (P4). In the solenodon, the
oval infraorbital foramen is dorsal to the anterolingual root
of the upper first molar (M1) (“iof” in Fig. 6).
Internal Acoustic Meatus.—In the dog (Evans 1993:143),
the internal acoustic meatus “is an irregularly elliptical
depression that is divided deeply by the transverse crest
(crista transversa). Dorsal to the crest is the opening of
the facial canal, which contains the facial nerve as well
as the cribriform dorsal [=superior] vestibular area (area
vestibularis utriculo-ampullaris) for the passage of nerve
bundles from the membranous labyrinth. Ventral to the
crest is the ventral [=inferior] vestibular area (area vestibularis saccularis), through which pass additional vestibular
nerve bundles that come from a deep, minute depression,
the foramen singulare. The spiral cribriform tract (tractus
spiralis foraminosus) is formed by the wall of the hollow
modiolus of the cochlea. The perforations are formed by
the fascicles of the cochlear nerve that arise from the spiral
ganglion on the outside of the modiolus.” The solenodon
AMNH 28271 (Figs. 20, 21A, C, 22) differs from the dog
(e.g., CM 106574) in the overall depth of the transverse
crest (“tc” in Fig. 22) and the foramen acusticum superius and inferius (the fossae on either side of the transverse
crest); in the former these are only slightly recessed from
the surrounding pars cochlearis and in the latter they are
very deeply recessed. The other major difference concerns
the number and size of the openings in the foramen acusticum inferius. In the dog (e.g., CM 106574) as in the newborn human (Terry 1942: fig. 137), the foramen centrale
cochleare in the spiral cribriform tract, which is the orifice
of the canal of the modiolus, is smaller than the inferior
vestibular area, whereas in the solenodon it is four times
larger. Additionally, in the dog (e.g., CM 106574), a foramen singulare separate from the inferior vestibular area is
not visible, whereas in the solenodon these two entities are
broadly separated (“fsi” and “iva” in Fig. 22).
Jugular Foramen.—In the dog (Evans 1993), the jugular foramen is between the petrosal (=pars petrosa of the
temporal bone of Evans 1993) and the occipital; based on
the disarticulated skull of a puppy (Evans 1993: fig. 4–45),
it appears to be largely or wholly the exoccipital (=lateral
part of the occipital of Evans 1993) that borders the foramen. Transmitted are the glossopharyngeal, vagus, and
accessory nerves and the sigmoid sinus, which then pass
through the petro-occipital and tympano-occipital fissures
Wible.indd 390
to reach the skull base. In the solenodon, the jugular foramen (=foramen lacerum posterius of Gregory 191; “jf”
in Figs. 25, 26B, 28) is largely between the petrosal and
exoccipital with only a minor contribution of the basioccipital to the anterior border based on MPIH 6863, the only
specimen to delimit the basi- and exoccipital. Transmitted
are the same nerves as in the dog (Gregory 1910); there is
some venous drainage based on MPIH 6863, but the primary conduit for the sigmoid sinus is the condyloid canal.
Lacrimal Fenestra.—In the dog (Evans 1993:1041), the
inferior oblique muscle (=musculus obliquus ventralis of
Evans 1993) “originates from a small depression in the
palatine bone near the junction of the palatomaxillary and
the palatolacrimal sutures. In prepared skulls this site may
appear as a foramen, since the attachment plate is thin and
easily lost.” Following Jollie (1968) and Gaudin and Wible
(2004), this depression is a lacrimal fenestra. In the solenodon juveniles, the lacrimal fenestra is a slit in the suture
between the orbital process of the lacrimal and maxilla
(“lacfe” in Figs. 6, 8, 10); in the adults, the lacrimal fenestra is replaced by a pit via the closure of the suture between
the lacrimal and maxilla.
Lacrimal Foramen.—In the dog (Evans 1993), the large
opening in the center of the orbital process of the lacrimal
is called the fossa for the lacrimal sac. The lacrimal sac,
formed by the union of the two lacrimal ducts, one from
each eyelid, leads into the nasolacrimal canal, by which
the nasolacrimal duct reaches the nasal vestibule. The solenodon has a similar arrangement with a large foramen
in the lacrimal immediately posteromedial to the orbital
rim that narrows into a canal anteriorly (“lacf” in Figs. 6,
8, 10). However, rather than calling this a fossa for the
lacrimal sac, the general term lacrimal foramen (=lacrymal
foramen of Gregory 1910) is employed.
Major Palatine Foramen.—In the dog (Evans 1993), the
major palatine nerve and vessels leave the pterygopalatine
fossa via the caudal palatine foramen, run through the palatine canal in the palatine bone, and reach the hard palate
via the major palatine foramen between the maxilla and
palatine opposite the distal part of the upper ultimate premolar. In the solenodon, there are a number of foramina
in the palatomaxillary suture and the palatine bone on the
back of the hard palate (Figs. 11–13). The exact contents
are not known, but these openings are either major or accessory palatine foramina. Until additional study is done,
those in the anterior palatomaxillary suture in the vicinity of the M1/M2 embrasure are treated as major palatine
foramina (“mapf” in Fig. 11). Variation occurs within and
between specimens in the number and position of major
palatine foramina. AMNH 185012 has one major palatine
foramen per side in the anterolateral palatomaxillary suture; AMNH 28271 (right side) and CM 18069 (right side)
have two in the suture, one anterolateral and the other anteromedial; AMNH 28271 (left side) and CM 18069 (left
side) have one anteromedial in the suture; and AMNH
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212912 has two per side, but the anteromedial one on the
right is within the maxilla and the anterolateral one on the
left is within the palatine. Based on the bisected solenodon
skull AMNH 28271, the major palatine nerve leaves the
pterygopalatine fossa via the sphenopalatine foramen, enters the nasopharyngeal meatus, and then exits the meatus
via a foramen opposite the embrasure between the upper
penultimate and ultimate molar (“mapf” in Fig. 21C).
Mandibular Foramen.—In the dog (Evans 1993), the
mandibular foramen, the caudal opening of the mandibular canal for the inferior alveolar nerve, artery, and vein
(=mandibular alveolar nerve, artery, and vein of Evans
1993), is on the medial side of the mandibular ramus, approximately in the antero-posterior center, ventral to the
alveolar margin. In the solenodon, the mandibular foramen
is slightly posterior to the antero-posterior midpoint of the
mandibular ramus, dorsal to the alveolar margin (“manf”
in Fig. 37).
Mastoid Canaliculus.—In the dog (Evans 1993), the auricular ramus of the vagus nerve in its course between the
jugular foramen and stylomastoid foramen runs in a small
canal in the petrosal (=pars petrosa of the temporal bone of
Evans 1993), the mastoid canaliculus of NAV (1994). In the
solenodon, the auricular ramus of the vagus is not enclosed
in a bony canal but passes through the back of the middle
ear, posterior to the fenestra cochleae, which it reaches by
penetrating broad medial and lateral gaps (“abX” in Fig.
27B). The medial gap lies lateral to the jugular foramen between the medial section of the caudal tympanic process,
the rostral tympanic process, and the pars canalicularis of
the petrosal (“mc” in Figs. 25, 27A, 29A); the lateral gap
lies medial to the stylomastoid foramen between the medial and lateral sections of the caudal tympanic process and
the pars canalicularis of the petrosal. In addition to transmitting the auricular ramus, this passageway also transmits
the lateral head vein (Fig. 27B).
Mastoid Foramen.—In the dog (Evans 1993), the mastoid foramen is on the occiput between the mastoid exposure of the petrosal (=mastoid process of Evans 1993)
and ex- and supraoccipital (=lateral and squamous parts
of the occipital of Evans 1993); it transmits the occipital
emissary vein, which joins the sigmoid sinus endocranially. Also reported by Evans (1993:608) is a caudal meningeal artery off the occipital artery that “goes through the
supramastoid foramen and ramifies in the dura [mater] of
the occipital cranial fossa”; the supramastoid foramen does
not appear elsewhere in the text and may be the mastoid
foramen. In the solenodon, McDowell (1958) reported the
mastoid foramen (his postmastoid foramen) to be absent,
but one or two small openings penetrate the mastoid exposure in all specimens (“msf” in Fig. 34) except the juvenile AMNH 28272 and the left side of AMNH 212912.
On the left side of AMNH 185012 and right side of AMNH
28271 is a dorsomedially placed foramen, near the suture
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391
with the parietal. On the right side of AMNH 185012 and
212912 and bilaterally in AMNH 28271 and CM 18069 is
a more ventrolaterally placed foramen, near the suture with
the squamosal. The latter openings bear a resemblance in
position to posterior openings into the posttemporal canal.
However, they do not lead into a canal between the pars
canalicularis and squamosal and, therefore, are not posttemporal foramina. MacPhee (1994: fig. 6 [p. 65]) illustrated a mastoid foramen for S. cubanus near the dorsomedial corner of the mastoid exposure’s suture with the fused
ex- and supraoccipital.
Maxillary Foramen.—In the dog (e.g., CM 30471), the
maxillary foramen, the posterior opening of the infraorbital canal, is in the anteroventral orbit dorsal to the posterior root of the upper ultimate premolar (P5) between the
maxilla, palatine, and jugal. In the solenodon, the maxillary foramen is entirely within the maxilla dorsal to the
embrasure between the upper first and second molars (M1
and M2) (“mxf” in Fig. 10).
Mental Foramen.—In the dog (Evans 1993), two or more
mental foramina in the mandibular body below the lower
anterior premolars transmit the mental nerves, arteries, and
veins from the mandibular canal. In the solenodon, the five
examined skulls have four or more mental foramina (“mf”
in Fig. 36) that vary in size and position both within and
between specimens (Figs. 39, 42). The anterior and posterior extremes are below the lower canine and below the
posterior root of the lower first molar (see Mandible for
details). The mental foramina are positioned in roughly
the same horizontal plane in AMNH 28272, 185012, and
212912, but the anterior ones are higher than the posterior
ones in AMNH 28271 and CM 18069.
Minor Palatine Foramen.—In the dog (Evans 1993), the
minor palatine nerve and artery arise in the pterygopalatine
fossa and run anteroventrally in a deep notch in the posterior edge of the palate between the palatine and maxilla
that rarely is closed to form a foramen, the minor palatine
foramen (following Wible and Rougier 2000). In the solenodon, McDowell (1958:146) noted the unusual nature
of the minor palatine foramen (his posterior palatine canal): “it is bored vertically, and forked, with a single dorsal
(orbital) opening, and two ventral (palatal) openings, one
anterior and one posterior to the postpalatine torus.” The
examined skulls show variability (see Fig. 13). The right
side of AMNH 212912 has the three openings reported by
McDowell (1958): one in the anteroventral pterygopalatine fossa, the dorsal minor palatine foramen; the second
leading anteriorly onto the hard palate, the anteroventral
minor palatine foramen; and the third leading posteriorly
onto the soft palate, the posteroventral minor palatine foramen. AMNH 185012 and 212912 (left side) have only two
foramina, because the anteroventral minor palatine foramen is a notch open ventrally, and CM 18069 has only
one foramen, because the dorsal and posteroventral minor
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palatine foramina are notches open posteriorly. McDowell
(1958) reported that in AMNH 28272 (currently with both
palatines missing) “the postpalatine torus is unossified,
and the posterior rim of the canal [minor palatine foramen]
is a simple notch in the rear border of the palatal surface
of the palatine.”
Occipital Emissary Vein Foramen.—In the dog (Evans
1993), the foramen of exit for the occipital emissary vein
from the caudal cranial fossa is the mastoid foramen between the mastoid exposure of the petrosal (=mastoid process of Evans 1993) and the ex- and supraoccipital (=lateral and squamous parts of the occipital of Evans 1993).
In the caudal cranial fossa of the bisected solenodon skull,
AMNH 28271, a large sulcus for the occipital emissary
vein runs perpendicular to the sulcus for the sigmoid sinus
dorsal to the petrosal (“oev” in Fig. 21C). The occipital
emissary vein sulcus ends posteriorly at a large foramen
that leads into the substance of the supraoccipital bone and
then to the supraoccipital foramina on the occiput (“socf”
in Figs. 34, 35). In light of the position of the occipital
emissary vein sulcus and foramen, it appears that both are
primarily on the supraoccipital bone.
Optic Foramen.—In the dog (Evans 1993), the optic foramen is centrally located in the orbitosphenoid (=orbital
wing of the presphenoid of Evans 1993) and transmits the
optic nerve, ophthalmic artery (=internal ophthalmic artery of Evans 1993), and the internal ophthalmic vein. In
the solenodon, the optic foramen is located near the ventral
limit of the orbitosphenoid (“of” in Figs. 15, 16). There
is left-right asymmetry in the anteroposterior position of
the optic foramen in three specimens (AMNH 185012 and
212912 and CM 18069). The solenodon optic foramen is
exceedingly small; its maximum diameter is approximately one fifth the size of the sphenorbital fissure, whereas in
the dog CM 30471 it is three quarters that of the orbital
fissure.
Orbitotemporal Canal, Anterior Opening.—In the solenodon, the orbitotemporal groove and canal in the middle
cranial fossa accommodates the anterior division of the ramus superior of the stapedial artery (see Wible 1987) and
accompanying veins based on MPIH 6863 (“otg” and “otc”
in Fig. 21C). The orbitotemporal groove is within the parietal, except for a small contribution from the squamosal at
its posterior extent based on AMNH 28272. Anteriorly, at
the posterior edge of the orbitosphenoid, the groove leads
into a short orbitotemporal canal, at least initially between
the parietal and orbitosphenoid. The canal ends at the anterior (orbital) opening between the frontal and orbitosphenoid in AMNH 28272 (“otc” in Figs. 16, 19), in a common
depression with the posterior frontal diploic vein foramen
(except the left side of AMNH 28272, which lacks the
venous foramen) and the posterodorsal ethmoidal foramen (except in AMNH 212912, in which the ethmoidal
foramen is anterior to the depression). The orbitotemporal
Wible.indd 392
canal and its orbital opening are lacking in the dog (Evans
1993) as is the anterior division of the ramus superior of
the stapedial artery (Bugge 1978; Wible 1984).
The history of usage for the solenodon is varied. Gregory (1910) called the orbitotemporal canal the sinus canal
following Parker (1886) and identified it as a venous structure; although the orbital aperture in AMNH 28272 was
labeled sinus canal (Gregory 1910: fig. 18A1; Fig. 1), supraorbital foramen was used in the text (p. 274). McDowell (1958) employed sinus canal foramen for the orbital
aperture and added the middle meningeal artery en route
to the ophthalmic arteries as an occupant. MacPhee (1994)
used cranio-orbital foramen and identified the occupants
as the sinus vein and ramus superior of the stapedial artery. The terminology followed here (orbitotemporal canal
and anterior opening of the orbitotemporal canal with occupants the anterior division of the ramus superior of the
stapedial artery and accompanying veins) is preferred as
stated by Wible and Gaudin (2004:162) “because it best
describes the position of this vascular canal and its orbital
egress, and it best reflects the broader homology of these
structures because it has already been applied to a broad
spectrum of cynodonts (see Rougier et al. 1992).”
Parietal Foramina.—In the dog (Evans 1993: fig. 12–22),
parietal foramina for emissary veins off the midline in the
parietal are not described but one with an emissary vein
is figured. Humans have small parietal foramina that are
variably present and differ in number, position, and symmetry (Boyd 1930). In the solenodon, the skulls examined
have a variable number of tiny, asymmetrically arranged
parietal foramina (Figs. 2, 5) except the juvenile AMNH
28272, which has none (Fig. 5).
Piriform Fenestra.—Following MacPhee (1981), the
piriform fenestra is the large gap in the skull base anterior
to the auditory capsule present in all fetal mammals and
some adults, including Solenodon. In the solenodon, the
piriform fenestra (“pf” in Figs. 18, 19, 25, 26B, 28, 29)
is bordered by the alisphenoid anteromedially, the squamosal anterolaterally, and the petrosal posteriorly. According to McDowell (1958), the tensor tympani muscle arises
in part from the membrane of the piriform fenestra (his
pyriform fenestra) and the internal carotid artery and ramus superior of the stapedial artery enter the cranial cavity
via this aperture (see Fig. 26C). The course of the internal
carotid artery is as noted by McDowell in AMNH 28272
and 185012 (and the left side of CM 18069), but not in
AMNH 28271 and 212912, where the artery is excluded
medially from the piriform fenestra by the basisphenoid in
a carotid foramen (“cf” in Fig. 25), a difference that may
result from developmental stage. The ramus superior of the
stapedial artery in MPIH 6863 dorsal to the piriform fenestra supplies three types of branches: the anterior division
traveling forward in the orbitotemporal groove, rami temporales, and the small posterior division traveling posteriorly in the posttemporal canal between the lateral surface
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of the pars canalicularis and squamosal. In addition, based
on MPIH 6863, the piriform fenestra transmits the internal
carotid nerve and veins connecting the cavernous sinus to
the pterygoid veins and inferior petrosal sinus. The piriform fenestra is lacking in the adult dog as the bones there
are in close contact (Evans 1993).
Postglenoid Foramen.—In the dog (Evans 1993), the
postglenoid foramen (=retroarticular foramen of Evans
1993) is in the squamosal (=pars squama of the temporal
bone of Evans 1993) posterior to the postglenoid process
(=retroarticular process of Evans 1993) and transmits the
postglenoid vein (=retroarticular vein of Evans 1993). In
the solenodon, the postglenoid foramen is in the squamosal posterior to the lateral end of the entoglenoid process (“pgf” in Figs. 25, 27A, 29A); a postglenoid process,
such as that in the dog, is absent. Although the foramen is
entirely within the squamosal, the anterior crus of the ectotympanic underlies the medial edge of the opening (Fig.
25).
Posttemporal Canal, Posterior Opening.—Among extant mammals, monotremes and some therians (e.g., some
didelphids, armadillos) have a posttemporal canal between
the lateral surface of the pars canalicularis of the petrosal
and the overlying squamosal, transmitting the posterior
division of the ramus superior of the stapedial artery, the
arteria diploëtica magna of Hyrtl (1853, 1854), and accompanying vein (Wible 1990; Wible and Hopson 1995; Wible
and Gaudin 2004). The posttemporal canal may open on
the occiput via the posterior opening of the posttemporal
canal or posttemporal foramen, which is near the lateral
margin of the occiput in or near the petrosquamous suture.
Neither the dog (Evans 1993) nor the examined solenodons have a posttemporal foramen (see Mastoid Foramen).
However, in the solenodon, MPIH 6863 has a small posterior division of the ramus superior that does not reach the
occiput running with significantly larger veins between the
pars canalicularis and squamosal, and AMNH 28272 has
a shallow posttemporal sulcus on the posterior part of the
lateral surface of the pars canalicularis, which represents
the medial part of a posttemporal canal (see Fig. 30).
Prootic Canal.—Among extant mammals, monotremes
and some marsupials have a prootic canal through the petrosal whereby the prootic sinus, the anterior distributary of
the transverse sinus, exits the cranial cavity to drain into
the lateral head vein in the middle ear (Wible 1990; Wible
and Hopson 1995; Sánchez-Villagra and Wible 2002;
Rougier and Wible 2006). A prootic canal is widely distributed among Mesozoic mammaliaforms including Cretaceous metatherians (Wible 1990; Rougier et al. 1998).
Within Eutheria, a prootic canal has only been described
in isolated petrosals referred to Early Cretaceous Prokennalestes (Wible et al. 2001) and Late Cretaceous zhelestids
(Ekdale et al. 2003), and in the skull of Late Cretaceous
Maelestes (Wible et al. 2007, in press). In Prokennalestes
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393
and non-therian mammaliaforms, the prootic canal is generally well developed and vertically oriented; in contrast,
in metatherians, zhelestids, and Maelestes, the prootic canal is reduced and horizontal.
The solenodon is the first placental, extant or extinct, for
which a prootic canal is reported. Its prootic canal resembles that in metatherians, zhelestids, and Maelestes, that is,
reduced and horizontal. The tympanic aperture (double in
AMNH 28272; single in AMNH 28271 and 212912) lies
on the pars canalicularis between the crista parotica and
facial sulcus (“prc” in Fig. 27A); the lateral aperture, visible only on the right side of the juvenile AMNH 28272 because of postmortem displacement of the squamosal, lies
on the lateral surface of the pars canalicularis in the ventral
end of the vertical sulcus for the prootic sinus (see Fig. 30).
Based on the bisected skull AMNH 28271, the adult differs from the juvenile in that a narrow sleeve of squamosal
with a small foramen in it intervenes between the lateral
aperture of the prootic canal and the sulcus for the prootic
sinus; the connection is maintained by passage from the
sulcus for the prootic sinus first into the small foramen in
the squamosal sleeve and then into the lateral aperture of
the prootic canal in the petrosal. MPIH 6863 confirms the
presence of a lateral head vein “lhv” in Fig. 27B), prootic
canal, and prootic sinus.
Prootic Sinus Sulcus.—In the dog (Evans 1993), the transverse sinus divides into anterior and posterior distributaries,
the temporal and sigmoid sinuses. The temporal sinus runs
in a canal between the pars petrosa and pars squamosa of
the temporal bone (=petrosal and squamosal) and exits the
skull at the retroarticular foramen (=postglenoid foramen).
In the bisected solenodon skull AMNH 28271, the anterior
distributary of the transverse sinus first runs anteriorly in a
canal between the parietal and the dorsalmost pars canalicularis of the petrosal (“ecps” in Fig. 21C) and then bends
anteroventrally and then medially into a deep sulcus in the
squamosal (“sps” in Fig. 21C). In the lateral wall of the canal is an opening (not visible in the figures) leading to the
posterodorsal foramen for ramus temporalis on the braincase wall and in the lateral wall at the bend is on opening
(“rt” in Fig. 21C) leading to the two anteroventral foramina
for rami temporales on the braincase wall. Additionally, at
the bend is the origination of the orbitotemporal groove.
Dorsal to the postglenoid foramen, the prootic canal enters
the posteromedial aspect of the sulcus (not visible in the
figures).
The homologies of the anterior distributary of the transverse sinus in the solenodon are at issue. Gelderen (1924)
showed that the vein exiting the postglenoid foramen has
a different developmental history in marsupials on the one
hand and placentals on the other. In marsupials, the primary ontogenetic anterior distributary of the transverse
sinus, the vena cerebralis media (=prootic sinus), and its
continuation as the vena capitis lateralis (=lateral head
vein) in the canalis prooticus (=prootic canal) are either
reduced or lost during later ontogeny (see also Wible 1990;
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Sánchez-Villagra and Wible 2002); a secondary ontogenetic anterior distributary of the transverse sinus forms,
Gelderen’s vena emissaria spheno-parietalis (=sphenoparietal emissary vein), so called because it leaves the chondrocranium via the sphenoparietal fenestra anterior to the
auditory capsule to reach the postglenoid foramen. In contrast, in placentals, the primary ontogenetic anterior distributary of the transverse sinus, the prootic sinus/lateral
head vein pathway, is lost and replaced by a secondary ontogenetic anterior distributary, Gelderen’s vena emissaria
capsulo-parietalis (=capsuloparietal emissary vein), so
called because it leaves the chondrocranium via the capsuloparietal foramen dorsal to the auditory capsule to reach
the postglenoid foramen. In recent years, several Cretaceous eutherians have been reported (Wible et al. 2001,
2007, in press; Ekdale et al. 2003) with an osteological
pattern closely resembling that of some marsupials (and
other metatherians), that is, a sulcus for the prootic sinus
on the lateral surface of the pars canalicularis of the petrosal confluent with a small prootic canal. Reported here is
the discovery of this osteological pattern in the solenodon.
The developmental history of the solenodon vessels is not
known and without that knowledge, the implications of the
discovery of this osteological pattern are uncertain. The author intends to pursue this topic, but of immediate concern
is a terminological issue regarding the anterior distributary
of the transverse sinus in the solenodon. For now, given
that a similar osteological pattern occurs in various Cretaceous and Paleocene metatherians (Wible 1990; Rougier
et al. 1998; Ladevèze 2004, 2007; Ladevèze and Muizon
2008), various extant marsupials (Wible 1990; SánchezVillagra and Wible 2002), various Cretaceous eutherians
(Wible et al. 2001, 2007, in press; Ekdale et al. 2003),
and the solenodon, it is judicious to apply the same terminology for the osteological correlates with the explicit
caveat that this is under review. Applying the metatherian/
Cretaceous eutherian terminology, AMNH 28271 has an
anteriorly directed endocranial canal for the prootic sinus
dorsally between the parietal and petrosal, then a ventrally
directed endocranial sulcus for the prootic sinus between
the squamosal and petrosal and ventrally just within the
squamosal. In the last segment is a foramen in the squamosal connecting to the prootic canal in the petrosal. Ventral to the prootic canal connection, the short segment of
the sulcus dorsal to the postglenoid foramen is for another
vein, which is called here the postglenoid vein to be congruent with the name of its foramen of exit.
Pterygoid Canal.—In the dog (Evans 1993), the caudal
opening into the small pterygoid canal is in the suture between the basisphenoid and posterodorsal margin of the
pterygoid bone on the skull base, and the rostral opening
is in the caudal pterygopalatine fossa anteroventral to the
sphenorbital fissure (orbital fissure of Evans 1993) between the pterygoid and the pterygoid process of the sphenoid (the ventral projection of the sphenoid that abuts the
pterygoid bone). It transmits the nerve of the pterygoid
Wible.indd 394
canal and occasionally a small artery off the maxillary
artery. In the solenodon, the caudal opening of the small
pterygoid canal (=vidian canal of MacPhee 1994) is between the caudal process of the pterygoid and basisphenoid at the anteromedial corner of the piriform fenestra,
based on AMNH 28272 and MPIH 6863 (“pc” in Figs. 19,
25, 27A). The rostral opening is deep within the cavum
epiptericum, the extradural space within the cranial cavity housing the trigeminal ganglion (Gaupp 1902, 1905),
at the level of the transverse canal foramen in the side of
the basisphenoid and is only visible through the sphenorbital fissure. A probe pushed into the caudal opening of
the pterygoid canal in the bisected skull AMNH 28271 appears in the transverse sinus canal within the basisphenoid
and then into the rear of the fossa for the trigeminal ganglion. In MPIH 6863, the pterygoid canal is separate from
(and passes ventral to) the transverse sinus canal. Based on
(unspecified) specimens with dried arteries in place, McDowell (1958:129) identified an artery of the pterygoid canal off the internal carotid artery (his vidian branch of the
arteria promontorii), which “passes through the basisphenoid or perhaps, between basisphenoid and alisphenoid) to
emerge in the cranial cavity, lateral to the pituitary fossa.
To this extent it is like the main carotid of marsupials and
therapsid reptiles, but the significance of this is uncertain.”
MacPhee (1981) noted that MPIH 6863 lacks such an artery and only vein accompanies nerve into the pterygoid
canal from behind.
Ramus Inferior Sulcus.—In the solenodon, based on
MPIH 6863, the ramus inferior of the stapedial artery (see
also McDowell 1958) and lesser petrosal nerve occupy a
sulcus on the posterior slope of the alisphenoid tympanic
process in their course between the piriform fenestra and
foramen ovale (“gri” in Figs. 25, 27A; “ri” in Fig. 26C).
The composition of this sulcus differs in the juveniles
AMNH 28272 and 185012, the only forms preserving a
suture here; in the former the alisphenoid and squamosal
contribute equally (Figs. 18, 27A), but in the latter it is
nearly entirely alisphenoid (Fig. 25). This difference may
result from the difference in developmental stages between
the two juveniles. In the dog, the ramus inferior is reported
to be absent (Wible 1984).
Secondary Facial Foramen.—see Facial Canal and/or
Sulcus.
Sigmoid Sinus Sulcus.—In the dog (Evans 1993:710),
the sigmoid sinus “is the roughly S-shaped caudoventral
continuation of the transverse sinus. It begins by forming
an arc around the proximal end of the petrous temporal
bone [=petrosal]. The first arc is continued by the second
arc, which lies medial to the petro-occipital synchondrosis.
The sinus terminates after traversing the jugular foramen
by continuing as the internal jugular vein.” In the bisected
solenodon skull, AMNH 28271, the sulcus for the sigmoid sinus lies dorsal to the subarcuate fossa in the caudal
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cranial fossa (“sss” in Fig. 21C). Anteriorly, the sulcus is
largely on the parietal with a minor contribution from the
pars canalicularis of the petrosal; posteriorly this relationship is reversed. At the posterodorsal corner of the petrosal
the supraoccipital makes a minor contribution to the dorsal
margin of the sulcus, which bends ventrally and ends at
the condyloid canal in the supraoccipital; the sulcus does
not extend to the jugular foramen. In light of the size of
the condyloid canal, it represents the primary exit for the
sigmoid sinus.
Sphenopalatine Foramen.—In the dog (Evans 1993), the
sphenopalatine foramen is within the perpendicular lamina of the palatine dorsal to the caudal palatine foramen.
It connects the pterygopalatine fossa and the nasal cavity,
and transmits the caudal nasal nerve and sphenopalatine
artery and vein. In the solenodon, a separate caudal palatine foramen is lacking and its contents, the major palatine
nerve, artery, and vein in the dog, are transmitted via the
sphenopalatine foramen. In the juvenile solenodon AMNH
28272, the only specimen preserving complete sutural information on the sphenopalatine foramen, this opening is
between the ethmoid, maxilla, and palatine (see “spf” in
Fig. 10); after a short rostrally directed canal it opens anteriorly within the nasopharyngeal meatus based on AMNH
28271 (“spf” in Fig. 22).
Sphenorbital Fissure.—In the dog (Evans 1993), the
sphenorbital fissure (=orbital fissure of Evans 1993) is located between the orbito- and alisphenoid (orbital and temporal wings of the sphenoid of Evans 1993), sandwiched
between the optic foramen and the rostral opening of alisphenoid canal (=rostral alar foramen of Evans 1993). It
transmits the oculomotor, trochlear, ophthalmic, and abducens nerves, the anastomotic artery connecting the maxillary and internal carotid arteries, and the ophthalmic venous
plexus. In the solenodon, the sphenorbital fissure is between
the orbito- and alisphenoid, somewhat separated from the
optic foramen and alisphenoid canal (“sof” in Figs. 14–16).
According to Gregory (1910), it transmits the oculomotor,
trochlear, ophthalmic, maxillary, and abducens nerves. It
most likely also transmits arteries and veins, the ramus infraorbitalis and ophthalmic veins.
One of the reviewers (G.W. Rougier) noted a problem
with the use of the term sphenorbital fissure as employed by
this author for the aperture both in marsupials (e.g., Monodelphis, Wible 2003) and in placentals (this report). In the
former, the sphenorbital fissure transmits the optic nerve, but
in latter it does not, because the optic nerve is enclosed in a
separate foramen. The developmental processes accounting
for this difference have been summarized by numerous authors (e.g., Kuhn 1971; Moore 1981; Kuhn and Zeller 1987;
Zeller 1989; Novacek 1993), but there appears to be no consensus on the naming of the resulting openings other than
that for the optic nerve. At least since Gregory (1910), the
tradition has been to employ sphenorbital fissure for the differing apertures in placentals and marsupials.
Wible.indd 395
395
Stapedial Artery Sulcus.—The stapedial artery is a branch
of the internal carotid artery that among extant adult mammals occurs in the platypus and various placentals (Tandler
1899, 1901; Bugge 1974; Wible 1987). In the placentals, the
stapedial artery often occupies a sulcus on the promontorium of the petrosal aimed at the fenestra vestibuli (Wible
1987). In the solenodon, based on MPIH 6863, the stapedial
artery sulcus has two separate parts: medial and anterolateral
to the fenestra vestibuli. The medial part arises about a millimeter medial to the fenestra vestibuli on the promontorium
from the (internal) carotid sulcus and crosses the posterior
half of that opening’s ventral rim (“gsa” in Fig. 26B). The
anterolateral part, also a millimeter long, extends between
the level of the secondary facial foramen and the back of the
piriform fenestra on the tegmen tympani (“gsa” in Fig. 27A).
Between these two sulci, the stapedial artery penetrates the
stapedial foramen in the stapes and runs ventral to the facial
nerve in the facial sulcus (“sa” in Fig. 26C). At is origin on
the promontorium, the stapedial artery sulcus is comparable
in width to the carotid sulcus. The stapedial artery (Tandler
1899; Bugge 1978) and sulcus (Evans 1993) are lacking in
the adult dog.
Stylomastoid Notch/Foramen.—In the dog (Evans 1993),
the stylomastoid foramen is deeply recessed ventral to the
mastoid exposure of the petrosal (=mastoid process of Evans 1993) and dorsal to the caudal part of the auditory bulla;
it transmits the facial nerve and the stylomastoid artery off
the caudal auricular artery, a branch of the external carotid.
In the solenodon, the stylomastoid foramen is a notch open
posteriorly and posteromedially; the notch is bordered laterally by the posterior continuation of the crista parotica, anteriorly by the tympanohyal, and anteromedially by the lateral
section of the caudal tympanic process (“smn” in Figs. 25,
27A). In MPIH 6863, the stylomastoid notch transmits the
facial nerve (“fn” in Fig. 27B), a branch of the ramus posterior of the stapedial artery (MacPhee 1981), and a small vein
off the lateral head vein.
Following Klaauw (1931), two types of stylomastoid foramina are often identified: the foramen stylomastoideum
primitivum and definitivum. The former occurs as a notch
in fetal mammals and some adults, including solenodon,
whereby the point of exit is between the tympanohyal and
pars canalicularis of the petrosal with contributions likely
also from the ectotympanic and stylohyal; the latter occurs
in mammals with inflated bulla (e.g., the dog), whereby the
point of exit is at the end of a canal in the bulla distal to the
fetal aperture.
Suboptic Foramen.—In the solenodon, according to Gregory (1910:248), “the suboptic foramen is located just below
the optic foramen; its canal runs downward and backward
to the transverse sinus in the presphenoid.” All solenodon
specimens have a suboptic foramen (“sbof” in Figs. 15, 19),
but the number and position varies considerably within and
between specimens with a maximum of six on the left side
of AMNH 28272 and a minimum of one on the left side
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of AMNH 28271. In AMNH 28272 and 185012, the only
specimens to preserve sutural information in the orbit, the
suboptic foramina are entirely within the orbitosphenoid
(Fig. 19). Continuity between the suboptic foramen and the
transverse sinus canal in the presphenoid was confirmed in
AMNH 28271. The dog (Evans 1993) does not have a suboptic foramen.
Supra-ethmoid Foramen.—see Foramen for Frontal
Diploic Vein.
Suprameatal Foramen.—A suprameatal foramen within
the squamosal dorsal to the external acoustic meatus and
ventral to the suprameatal crest (Novacek 1986a; Wible
2003; Wible et al. 2004) is lacking in the dog (Evans 1993).
In the solenodon, one or more small suprameatal foramina
(subsquamosal foramina of Gregory 1910; “f.sb.sq.” in Fig.
1) are found in the squamosal inferior to the suprameatal
crest in the vicinity of the roof of the external acoustic meatus and posttympanic crest (“smef” in Figs. 24, 27A). In
MPIH 6863, a tiny suprameatal foramen transmits a branch
of the ramus posterior of the stapedial artery into the substance of the squamosal (see Fig. 26C).
Supraoccipital Foramina.—In the dog (Evans 1993: figs.
12–21, 12–22), an unnamed foramen in the supraoccipital
(=squamous part of the occipital of Evans 1993) for a venous connection between the transverse sinus and occipital
emissary vein is illustrated, but not described in the text. In
the solenodon, the number, size, and position of foramina
in the supraoccipital bone varies (“socf” in Figs. 34, 35).
In AMNH 28272 a centrally positioned heart-shaped depression contains a half dozen smaller openings; in AMNH
185012 there are two larger openings per side, well off the
midline, and a number of smaller openings; and in AMNH
212912 and CM 18069, an ovoid depression just to the left
of the external occipital crest contains several smaller openings as well as an additional dozen small openings arrayed
across the occiput. All openings are presumably outlets for
the occipital emissary vein, but this cannot be verified in
MPIH 6863 because of lack of preservation.
Supra-optic Foramen.—In the solenodon, according to
Gregory (1910:248), the supra-optic foramen (above the
optic foramen) “runs downward and backward and joins
the suboptic” foramen, which in turn “runs downward and
backward to the transverse sinus in the presphenoid.” Although the opening that Gregory labeled as the supra-optic
foramen in AMNH 28272 is the posterodorsal ethmoidal
foramen of this report (cf. Figs. 1, 19), this specimen has
a small foramen within the frontal anteromedial to the posterodorsal ethmoidal foramen that likely is a supra-optic foramen based on the definition of Gregory (“s-of” in Fig. 19).
A probe passed into a similarly situated foramen on the right
side of AMNH 28271 reaches the transverse sinus canal in
the presphenoid. Sutures are lacking in AMNH 28271, but
it is doubtful that this foramen is entirely within the frontal and the orbitosphenoid likely contributes. A supra-optic
Wible.indd 396
foramen also occurs in CM 18069 (“4” in Fig. 15), but is
lacking in AMNH 185012 and 212912.
Transverse Canal Foramen.—In the solenodon, each side
of the bisected skull AMNH 28271 has an opening on the
cut edge of the basisphenoid for a vein crossing the midline based on MPIH 6863 (“tsc” in “bs” in Fig. 21C). The
principal exit of this vein is via the transverse canal foramen (transverse canal of Gregory 1910), posterior to but in
a common depression with the alisphenoid canal (“tcf” in
Figs. 15, 16, 19). Two others exits are evident: in the ventral
surface of the basisphenoid in or near the pterygoid suture
based on AMNH 28272 (Fig. 19) and in the dorsolateral
surface of the basisphenoid within the cranial cavity (lacking in AMNH 185012). Gregory (1910) reported a second
transverse sinus in the presphenoid, the opening of which
is preserved in the cut edges of that bone in AMNH 28271
(“tsc” in “ps” in Fig. 21C). According to Gregory (1910)
and confirmed here, two types of foramina connect to the
presphenoid transverse sinus: supra-optic and suboptic. A
transverse canal foramen and the presphenoid and basisphenoid transverse sinuses are absent in the dog (Evans 1993).
Tympanic Canaliculus.—In the dog (Evans 1993: fig. 19–
20), the course of the tympanic nerve is illustrated but not
described in the text. As figured, it arises from the glossopharyngeal nerve beneath the jugular foramen and runs laterally
into the middle ear between the pars cochlearis and auditory
bulla, through the tympanic canaliculus of NAV (1994). In
the solenodon, MacPhee (1981) reported a small tympanic
canaliculus in the caudal tympanic process in MPIH 6863,
but in the current report this process is identified as the rostral tympanic process because it arises from the pars cochlearis (“tca” in Figs. 25, 27A).
Ventral Condyloid Foramen.—In the solenodon, the ventral condyloid fossa between the occipital condyle and faint
paracondylar process on the exoccipital’s horizontal process
has two or more small foramina that connect to the condyloid canal (“vcf” in Fig. 26B). One specimen, AMNH
28271, has an additional large opening bilaterally. In the dog
(Evans 1993), ventral condyloid foramina are not reported
or illustrated.
Vestibular Aqueduct.—In the dog (Evans 1993:143), the
endolymphatic duct enters into the petrosal (=pars petrosa
of the temporal bone of Evans 1993) via the opening of
the vestibular aqueduct, which “is located caudodorsal to
the opening of the cochlear canaliculus in a small but deep
cleft.” In the solenodon AMNH 28271, the vestibular aqueduct is similarly situated (“av” in Figs. 21C, 22). However,
whereas the vestibular aqueduct is smaller than the cochlear
canaliculus in the dog (Evans 1993), the reverse is the case
in AMNH 28271.
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Wible—On the Cranial Osteology of Solenodon paradoxus
397
Appendix 1. List of Anatomical Terms. The terms used here are to the left with references; synonyms and/or Nomina Anatomica Veterinaria (NAV)
equivalents to the right.The * indicates structures discussed in the text but absent in Solenodon paradoxus (e.g., caudal palatine foramen).
Abducens Nerve (= Cranial Nerve VI)—Nervus abducens (NAV)
Accessory Nerve (= Cranial Nerve XI)—Nervus accessorius (NAV)
Accessory Palatine Foramen—(Wible and Rougier 2000); Middle
Palatine Foramen (Novacek 1986); Minor Palatine Foramen (Evans
1993)
Accessory Palatine Nerve—Nevus palatinus accessorius (NAV)
Ala temporalis—(De Beer 1937)
Alae of Vomer—Alae vomeris (NAV)
Alicochlear Commissure—(De Beer 1937; MacPhee 1981)
Alisphenoid—Os basisphenoidale, Ala temporalis (NAV)
Alisphenoid Canal—(Gregory 1910); Canalis alaris (NAV)
Alveolar Canal—Canalis alveolaris (NAV)
Alveolar Foramina—Foramina alveolaria (NAV)
Alveolar Margin—Margo alveolaris (NAV)
Alveolar Process of Maxilla—Os maxillare, Processus alveolaris (NAV)
Alveolar Process of Premaxilla—Os incisivum, Processus alveolaris
(NAV)
Alveoli—Alveoli dentales (NAV)
Ampulla of Lateral Semicircular Canal—Ampulla ossea lateralis (NAV)
Ampulla of Posterior Semicircular Canal—Ampulla ossea posterior
(NAV)
Anastomotic Artery*—Arteria anastomotica (Wible 1987; Evans 1993)
Angle of Mandible—Mandibula, Angulus mandibulae (NAV)
Angle of Stylohyal—Angulus stylohyoideus (NAV)
Angular Process—Processus angularis (NAV)
Annular Ridge of Frontal—(Rowe et al. 2005)
Anterior Crus of Ectotympanic—Annulus tympani, Crus anterior (NAV)
Anterior Crus of Stapes—Stapes, Crus rostrale (NAV)
Anterior Division of Ramus Superior of Stapedial Artery—(Wible and
Gaudin 2004)
Anterior Opening, Orbitotemporal Canal—(Rougier et al. 1992);
Supraorbital Foramen (Gregory 1910); Sinus Canal Foramen
(McDowell 1958); Foramen for Ramus Supraorbitalis (Wible 1987);
Cranio-Orbital Foramen (MacPhee 1994)
Anterior Pole of Promontorium—(Wible et al. 2001)
Anterior Semicircular Canal—Canalis semicircularis anterior (NAV)
Anteroventral Ethmoidal Foramen—this study
Antorbital Fossa—(Novacek 1986)
Aperture of Cochlear Fossula—(MacPhee 1981)
Aqueductus Vestibuli—(NAV)
Arteria diploëtica magna—(Hyrtl 1853, 1854; Wible 1987)
Artery of Pterygoid Canal*—(Evans 1993); Vidian Artery (McDowell
1958)
Auditory Bulla—Bulla tympanica (NAV)
Auditory Capsule—Capsula otica (NAV)
Auditory Ossicles—Ossicula auditus (NAV)
Auditory Tube—Tuba auditiva (NAV); Eustachian Tube (Gregory 1910)
Auricular Ramus of Vagus Nerve—Nervus vagus, Ramus auricularis
(NAV)
Axis—(NAV)
Basal Lamina of Ethmoid—Os ethmoidale, Lamina basalis (NAV)
Basal Lamina of Ventral Nasal Concha—(Evans 1993)
Basicranium—Basis cranii interna et externa (NAV)
Basihyal—Basihyoideum (NAV)
Basioccipital—Os occipitale, Pars basilaris (NAV)
Basipharyngeal Canal—(Evans 1993)
Basisphenoid—Os basisphenoidale, Corpus (NAV)
Body of Incus—Corpus incudis (NAV)
Body of Mandible—Corpus mandibulae (NAV)
Brain—Encephalon (NAV)
Braincase—Calvaria (NAV)
Canines—Dentes canini (NAV)
Canal for Modiolus—(Terry 1942)
Wible.indd 397
Capitular Crest of Malleus—(Henson 1961)
Capitular Spine of Malleus—(Henson 1961)
Capsuloparietal Emissary Vein*—(Gelderen 1924); Sinus temporalis
(NAV)
Capsuloparietal Foramen—(Gelderen 1924); Fissura occipitocapsularis
superior (De Beer 1937)
Carnassial Tooth*—(Evans 1993); Dens sectorius (NAV)
Carotid Canal*—(Evans 1993); Canalis caroticus (NAV)
Carotid Foramen—(Wible and Gaudin 2004); Canalis caroticus (NAV); Foramen lacerum medium (Gregory 1910)
Carotid Sulcus—Sulcus caroticus (NAV)
Cartilaginous Nasal Septum—Cartilago septi nasi (NAV)
Caudal Auricular Artery—Arteria auricularis caudalis (NAV)
Caudal Cranial Fossa—Fossa cranii caudalis (NAV); Occipital Cranial
Fossa (Evans 1993)
Caudal Meningeal Artery—Arteria meningea caudalis (NAV)
Caudal Nasal Nerve—Nervus nasalis caudalis (NAV)
Caudal Opening of Pterygoid Canal—(Wible and Gaudin 2004); Vidian
Foramen (McDowell 1958)
Caudal Palatine Foramen*—(Evans 1993); Foramen palatinum caudale
(NAV)
Caudal Process of Pterygoid—(Giannini et al. 2006)
Caudal Process of Squamosal—this study
Caudal Tympanic Process of Petrosal—(MacPhee 1981)
Cavernous Sinus—Sinus cavernosus (NAV)
Cavum Epiptericum—(Gaupp 1902, 1905; De Beer 1937)
Cavum Supracochleare—(Voit 1909; De Beer 1937); Geniculum canalis
facialis (NAV)
Central Buttress of Malleus—(Henson 1961)
Ceratohyal—Ceratohyoideum (NAV)
Cerebellum—(NAV)
Cerebral Hemisphere—Cerebrum (NAV)
Choanae—(NAV)
Chorda Tympani Nerve—Chorda tympani (NAV)
Circular Fissure—(Rowe et al. 2005)
Cleidomastoid Muscle—(Allen 1910); Musculus cleidomastoideus
(Evans 1993)
Cochlear Canaliculus—(MacPhee 1981); Apertura externa canaliculus
cochleae (NAV); Aqueductus cochleae (Evans 1993)
Cochlear Duct—Ductus cochlearis (NAV)
Cochlear Fossula—(MacPhee 1981)
Cochlear Nerve—Nervus cochlearis (NAV)
Common Nasal Meatus—Meatus nasi communis (NAV)
Conchal Crest—Crista conchalis (NAV)
Condylar Process—Processus condylaris (NAV)
Condyloid Artery—(Evans 1993); Arteria condylaris (NAV)
Condyloid Canal—(Evans 1993); Canalis condylaris (NAV); Venous
Condylar Foramen (Gregory 1910)
Condyloid Vein—(Evans, 1993)
Coronoid Crest—Crista coronoidea (Evans 1993)
Coronoid Process—Processus coronoideus (NAV)
Craniopharyngeal Canal—(Evans 1993)
Cranium—NAV
Cribroethmoidal Foramen—(Moore 1981); Ethmoidal Foramen (Sisson
1910); Ethmoidal Fissure (Terry 1942)
Cribriform Plate of Ethmoid—Os ethmoidale, Lamina cribrosa (NAV)
Cricoid Cartilage—Cartilago cricoidea (NAV)
Crista Frontalis—(NAV)
Crista Galli of Ethmoid*—Os ethmoidale, Crista galli (NAV)
Crista Interfenestralis—(Wible et al. 1995)
Crista Parotica—(De Beer 1937)
Crista Petrosa—(Wible 1990); Crista parties petrosae (NAV)
Crista Stapedis—(Henson 1961)
Crista Supramastoidea—(NAV)
Crista Tympanica—(MacPhee 1981)
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Appendix 1., continued
Crural Sulcus of Stapes—(Henson 1961)
Crus Breve of Incus—Incus, Crus breve (NAV)
Crus Commune of Semicircular Canals—(Wible 1990); Crus osseum
commune (NAV)
Crus Longum of Incus—Incus, Crus longum (NAV)
Deciduous Teeth—Dentes decidui (NAV)
Diastema—(NAV)
Digastric Muscle—Musculus digastricus (NAV)
Diploë—(NAV)
Dorsal Condyloid Foramina—(Wible and Gaudin 2004)
Dorsal Condyloid Fossa—Fossa condylaris dorsalis (NAV)
Dorsal Nasal Meatus—Meatus nasi dorsalis (NAV)
Dorsum Sellae—(NAV)
Dura Mater—Dura mater encephali (NAV)
Ectopterygoid Process*—(Giannini et al. 2006); Ectopterygoid Lamina
(McDowell 1958); Ectopterygoid Crest (Novacek 1986)
Ectoturbinates—(Smith and Rossie 2006); Ectoturbinalia (NAV)
Ectotympanic—(MacPhee 1981); Annulus tympanicus (NAV);
Tympanic (Gregory 1910)
Embrasure—Septa interalveolaria (NAV)
Endocranial Canal for Prootic Sinus—this study
Endocranium—Cavum cranii (NAV)
Entoglenoid Capitular Facet of Glenoid Fossa—Postglenoid Capitular
Facet of Glenoid Fossa (McDowell 1958)
Entoglenoid Facet of Mandibular Condyle—Postglenoid Facet of
Mandibular Condyle (McDowell 1958)
Entoglenoid Process of Squamosal—(McDowell 1958); Postglenoid
Process (Allen 1910); Post-Glenoid (Entoglenoid) Process (Gregory
1910); Modified Entoglenoid Process (McDowell 1958; MacPhee 1981),
Pseudopostglenoid (Entoglenoid) Process (Novacek 1986a);
Pseudoglenoid Process (MacPhee 1994)
Endolymphatic Duct—Ductus endolymphaticus (NAV)
Entopterygoid Process—(Giannini et al. 2006); (Internal) Pterygoid
Lamina (McDowell 1958); Entopterygoid Crest (Novacek 1986)
Entotympanic*—(Klaauw 1922); Os temporale, Pars endotympanica
(NAV)
Epihyal—Epihyoideum (NAV)
Epitympanic Recess—(Klaauw 1931); Recessus epitympanicus (NAV)
Epitympanic Wing of Petrosal—(MacPhee 1981)
Ethmoid—Os ethmoidale (NAV)
Ethmoidal Foramen—Foramen ethmoidale (NAV); Ethmoid Foramen
(Gregory 1910) Ethmoidal Fossae—Fossae ethmoidales (NAV)
Ethmoidal Nerve—Nervus ethmoidalis (NAV)
Ethmoturbinate I—(Smith and Rossie 2006); Endoturbinalia II (Evans
1993)
Ethmoturbinate II—(Smith and Rossie 2006); Endoturbinalia III (Evans
1993)
Ethmoturbinate III—(Smith and Rossie 2006); Endoturbinalia IV (Evans
1993)
Ethmoturbinate IV—(Smith and Rossie 2006)
Ethmoturbinates—(Smith and Rossie 2006); Ethmoturbinalia (NAV)
Exoccipital—Os occipitale, Pars lateralis (NAV)
Exoccipital Crest—(Gaudin 1995)
External Acoustic Meatus—Meatus acusticus externus (NAV); External
Auditory Meatus (Gregory 1910)
External Capitular Facet of Glenoid Fossa—(McDowell 1958)
External Carotid Artery—Arteria carotis externa (NAV); Ectocarotid
Artery (Gregory 1910)
External Ethmoidal Artery—Arteria ethmoidalis externa (NAV)
External Ethmoidal Vein—Vena ethmoidalis externa (NAV)
External Facet of Manidbular Condyle—(McDowell 1958)
External Lamina of Ethmoid—(Evans 1993); Os ethmoidale, Lamina
tectoria + Lamina orbitalis + Lamina basalis (NAV)
External Nasal Aperture—Apertura nasi ossea (NAV); External Nasal
Opening or Piriform Aperture (Evans 1993)
Wible.indd 398
External Occipital Crest—Crista occipitalis externa (NAV)
Eyelids—Palpebrae (NAV)
Facet for Ectotympanic on Squamosal—(Wible et al. 2004)
Facet for Malleus on Petrosal—this study
Facet for Tympanic Process of Petrosal on Ectotympanic—(McDowell
1958)
Facial Canal—(MacPhee 1981); Canalis facialis (NAV); Fallopian
Aqueduct (McDowell 1958)
Facial Nerve (= Cranial Nerve VII)—Nervus facialis (NAV)
Facial Process of Lacrimal—Os lacrimale, Facies facialis (NAV)
Facial Process of Premaxilla—Os incisivum, Facies labialis (NAV)
Facial Sulcus—(MacPhee, 1981)
Facial Surface of Maxilla—Os maxillare, Facies facialis (NAV)
Falx Cerebri—(NAV)
Fenestra Cochleae—(NAV); Fenestra Rotunda (McDowell 1958)
Fenestra Vestibuli—(NAV); Fenestra Ovalis (Gregory 1910)
Footplate (= Base) of Stapes—Stapes, Basis stapedis
Foramen Acusticum Inferius—(Sisson 1910); Ventral Vestibular Area
(Evans 1993)
Foramen Acusticum Superius—(Sisson 1910); Facial Canal + Dorsal
Vestibular Area (Evans 1993)
Foramen Centrale Cochleare—(Terry 1942)
Foramen for Chorda Tympani Nerve of Malleus—(Henson 1961)
Foramen for Inferior Petrosal Sinus—(Wible 1983)
Foramen for Occipital Emissary Vein—this study
Foramen for Ramus Temporalis—(Wible and Gaudin 2004); Post Parietal Foramen; Post-Squamosal Foramen (Gregory 1910);
Subsquamosal Foramen(Wible et al. 2004)
Foramen Magnum—(NAV)
Foramen Ovale—(NAV)
Foramen Rotundum*—(NAV)
Foramen Singulare—(NAV)
Foramen Spinosum*—(NAV)
Foramen Stylomastoideum Definitivum*—(McDowell 1958)
Foramen Stylomastoideum Primitivum—(McDowell 1958)
Fossa for Lacrimal Sac—Fossa sacci lacrimalis (NAV)
Fossa for Stapedius Muscle—(McDowell 1958); Fossa m. stapedius
(Evans, 1993)
Fossa for Tensor Tympani Muscle—(MacPhee, 1981); Fossa m. tensor
tympani (Evans, 1993)
Fossa Incudis—(MacPhee 1981)
Frontal—Os frontale (NAV)
Frontal Diploic Vein—Vena diploica frontalis (NAV)
Frontal Diploic Vein Foramen—(Thewissen 1989); Supra-Ethmoid
Foramen (Gregory 1910); Frontal Diploic Foramen (MacPhee 1994)
Frontal Sinus—Sinus frontalis (NAV)
Frontoethmoidal Suture—Sutura frontoethmoidalis (NAV)
Fundus of Nasal Fossa—(Evans 1993)
Geniculate Ganglion—Ganglion geniculi (NAV)
Glaserian Fissure—(MacPhee 1981); Fissura Glaseri (Klaauw 1931)
Glenoid Area of Squamosal—this study
Glenoid Fossa—Fossa mandibularis (NAV); Retroarticular Fossa (Evans
1993)
Glossopharyngeal Nerve (= Cranial Nerve IX)—Nervus glossopharyngeus
(NAV)
Greater Petrosal Nerve—Nervus petrosus major (NAV); Great Superficial
Petrosal Nerve (McDowell 1958)
Groove for Auditory Tube—(Wible 2003)
Groove for Chorda Tympani Nerve of Malleus—(Henson 1961)
Groove for Chorda Tympani Nerve of Squamosal—(McDowell 1958)
Groove for Internal Carotid Artery—(Wible 1986)
Groove for Postglenoid Vein—this study
Groove for Ramus Inferior of StapedialArtery—(Wible 1987)
Groove for Rostral Process of Malleus of Ectotympanic—Groove for
Folian Process of Malleus of Ectotympanic (McDowell 1958)
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Appendix 1., continued
Groove for Stapedial Artery—(Wible 1987)
Hard Palate—Palatum osseum (NAV)
Head of Malleus—Caput mallei (NAV)
Head of Stapes—Caput stapedis (NAV)
Hiatus Fallopii—(McDowell 1958); Petrosal Canal (Evans 1993)
Horizontal Part of Exoccipital—(Wible 2003)
Horizontal Part of Vomer— (Evans 1993)
Horizontal Process of Palatine—Os palatinum, Lamina horizontalis
(NAV)
Horizontal Sulcus of Cribriform Plate—this study
Hyoid Apparatus—Apparatus hyoideus (NAV)
Hypoglossal Foramen—Canalis n. hypoglossum (NAV); Condylar
Foramen (Gregory 1910); Foramen hypoglossi (Evans 1993)
Hypoglossal Nerve (= Cranial Nerve XII)—Nervus hypoglossus (NAV)
Hypoglossal Vein—Vena canalis hypoglossi (Evans 1993)
Hypophyseal Fossa—Fossa hypophysealis (NAV); Pituitary Fossa or
Depression (Gregory 1910)
Hypophysis—(NAV); Pituitary Body (Gregory 1910)
Incisive Foramen—Fissura palatina (NAV)
Incisive Incisure of Vomer— (Evans 1993)
Incisivomaxillary Canal—Canalis maxilloincisivus (Evans 1993)
Incisors—Dentes incisivi (NAV)
Incudomallear Joint—Articulation incudomallearis (NAV)
Incudostapedial Joint—Articulation incudostapedia (NAV)
Incus— (NAV)
Inferior Alveolar Artery—Arteria alveolaris inferior (NAV); Mandibular
Alveolar Artery (Evans 1993)
Inferior Alveolar Nerve—Nervus alveolaris inferior (NAV); Mandibular
Alveolar Nerve (Evans 1993)
Inferior Alveolar Vein—Vena alveolaris inferior (NAV); Mandibular
Alveolar Vein (Evans 1993)
Inferior Articular Facet of Malleus—(Henson 1961)
Inferior Articular Surface of Incus—(Henson 1961)
Inferior Oblique Muscle—Musculus obliquus ventralis (NAV)
Inferior Petrosal Sinus—Sinus petrosus ventralis (NAV)
Inferior Vestibular Area—(Terry 1942); Area vestibularis superior
(NAV)
Infraorbital Artery—Arteria infraorbitalis (NAV)
Infraorbital Canal—Canalis infraorbitalis (NAV)
Infraorbital Foramen—Foramen infraorbitale (NAV)
Infraorbital Nerve—Nervus infraorbitalis (NAV)
Infraorbital Vein—Vena infraorbitalis (NAV)
Inner Lamella of Malleus—(Henson 1961)
Interfrontal Suture—Sutura interfrontalis (NAV)
Intermaxillary Suture—Sutura intermaxillaris (Evans, 1993); Rostral Part
of Sutura palatina mediana (NAV)
Internal Acoustic Meatus—Meatus acusticus internus (NAV); Internal
Auditory Meatus (Gregory 1910)
Internal Carotid Artery—Arteria carotis interna (NAV); Entocarotid
Artery (Gregory 1910); Arteria promontorii (McDowell 1958)
Internal Carotid Nerve—Nervus caroticus interna (NAV)
Internal Jugular Vein—Vena jugularis interna (NAV)
Internasal Suture—Sutura internasalis (NAV)
Interpalatine Suture—Caudal Part of Sutura palatina mediana (NAV)
Interparietal—Os interparietale (NAV)
Intersphenoidal
Synchondrosis—Synchondrosis
intersphenoidalis
(NAV)
Jugal*—Os zygomaticum (NAV); Malar (Gregory 1910)
Jugular Foramen—Foramen jugulare (NAV)
Jugular Incisure—Incisura jugularis (NAV)
Jugular Process of Exoccipital—Processus jugularis (NAV)
Jugum Sphenoidale—(NAV)
Lacrimal—Os lacrimale (NAV); Lachrymal (Gregory 1910)
Wible.indd 399
Lacrimal Duct—Canaliculus lacrimalis (NAV)
Lacrimal Fenestra—(Wible and Gaudin 2004)
Lacrimal Foramen—Foramen lacrimale (NAV)
Lacrimal Sac—Saccus lacrimalis (NAV)
Lacrimal Tubercle*—(McDowell 1958)
Lateral Laminae of Vomer—Laminae lateralis of Vomer (Evans, 1993)
Larynx—(NAV)
Lateral Aperture of Prootic Canal—(Wible 2003)
Lateral Head Vein—(Wible 1990; Wible and Hopson 1995; Rougier and
Wible 2006)
Lateral Head Vein Sulcus—Groove for Lateral Head Vein (Wible and
Hopson 1995)
Lateral Laminae of Vomer­—(Evans 1993)
Lateral Mass of Ethmoid—(Evans 1993); Os ethmoidale, Labyrinthus
ethmoidalis (NAV)
Lateral Process of Malleus—Malleus, Processus lateralis (NAV)
Lateral Pterygoid Muscle—Musculus pterygoideus lateralis (NAV)
Lateral Section of Caudal Tympanic Process—(MacPhee 1981)
Lateral Semicircular Canal—Canalis semicircularis lateralis (NAV)
Lenticular Process of Incus—Incus, Processus lenticularis (NAV)
Lesser Petrosal Nerve—Nervus petrosus minor (NAV)
Levator Labii Superioris Proprius—(Whidden 2002); Musculus levator
labii superioris (NAV)
Lingual Nerve—Nervus lingualis (NAV)
Longus Capitis Muscle—Musculus longus capitis (NAV)
Major Palatine Artery—Arteria palatina major (NAV)
Major Palatine Foramen—Foramen palatinum majus (NAV); Anterior
Palatine Foramen (Gregory 1910)
Major Palatine Nerve—Nervus palatinus major (NAV)
Major Palatine Vein—Vena palatinus major (NAV)
Mallear Facet on Crista Parotica—this study
Malleus—(NAV)
Mandible—Mandibula (NAV)
Mandibular Canal—Canalis mandibulae (NAV)
Mandibular Foramen—Foramen mandibulae (NAV)
Mandibular Nerve—Nervus mandibularis (NAV)
Mandibular Symphysis— (Evans 1993)
Manubrium of Malleus—Manubrium mallei (NAV)
Masseter Muscle—Musculus massetericus (NAV)
Masseteric Fossa—Fossa masseterica (NAV)
Mastoid Canaliculus—(NAV); Passage for Auricular Nerve (McDowell
1958)
Mastoid Exposure of Petrosal—(Wible 2003); Processus mastoideus
(NAV); Mastoid Portion of Periotic (Gregory 1910)
Mastoid Foramen—Foramen mastoideum (NAV); Post-Mastoid
Foramen (Gregory 1910); Postmastoid Foramen (McDowell 1958)
Mastoideohyoideus Muscle—(Thiel 1955); Musculus mastoideostyloideus
(Saban 1968); Musculus jugulohyoideus (Evans 1993)
Maxilla—Os maxillare (NAV); Maxillary (Gregory 1910)
Maxillary Foramen—Foramen maxillare (NAV)
Maxillary Nerve—Nervus maxillaris (NAV)
Maxillary Process of Frontal—(Evans 1993)
Maxillary Recess—Recessus maxillaris (NAV)
Maxillary Vein—Vena maxillaris (NAV)
Maxilloturbinate (= Ventral Nasal Concha)—Os conchae nasalis
ventralis (NAV)
Meckel’s Cartilage—(De Beer 1937)
Medial Pterygoid Muscle—Musculus pterygoideus medialis (NAV)
Medial Section of Caudal Tympanic Process of Petrosal—(MacPhee
1981)
Mental Artery—Arteria mentalis (NAV)
Mental Foramen—Foramen mentale (NAV)
Mental Nerves—Nervi mentales (NAV)
Mental Vein—Vena mentalis (NAV)
Mesocranium—(Wible et al. 2004)
Middle Cranial Fossa—Fossa cranii media (NAV)
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Annals of Carnegie Museum
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Appendix 1., continued
Middle Nasal Meatus—Meatus nasi medius (NAV)
Middle Ear—Auris media (NAV)
Middle Ear Ossicles—Ossicula auditus (NAV)
Middle Meningeal Artery—Arteria meningea media (NAV)
Minor Palatine Artery—Arteria palatina minor (NAV)
Minor Palatine Foramen—(Wible and Rougier 2000); Foramen
palatinum caudale (NAV); Postero-External Palatine Foramen
(Gregory 1910); PosteriorPalatine Canal (McDowell 1958)
Minor Palatine Nerve—Nervus palatinus minor (NAV)
Molars—Dentes molares (NAV)
Muscular Process of Malleus—Malleus, Processus muscularis (NAV)
Muscular Process of Stapes—Processus muscularis stapedius (Henson
1961)
Muscular Tubercle—Os occipitale, Pars basilaris, Tuberculum musculare
(NAV)
Mylohyoid Line—Linea mylohyoideus (NAV)
Nasal—Os nasale (NAV)
Nasal Cavity (= Nasal Fossae)—Cavum nasi (NAV)
Nasal Fossa—(Evans 1993); Olfactory Recess (Moore 1981)
Nasal Septum—Septum nasi osseum (NAV)
Nasal Surface of Maxilla—Os maxillare, Facies nasalis (NAV)
Nasal Vestibule—Vestibulum nasi (NAV)
Nasolacrimal Canal—Canalis lacrimalis (NAV)
Nasolacrimal Duct—Ductus nasolacrimalis (NAV)
Nasopalatine Duct—(Asher 2001); Ductus incisivus (NAV)
Nasopharyngeal Meatus—Meatus nasopharyngeus (NAV)
Nasoturbinate, Ethmoid—(Smith and Rossie 2006); Concha nasalis
dorsalis (NAV); Endoturbinalia I (Evans 1993)
Nasoturbinate, Nasal—(Smith and Rossie 2006); Crista ethmoidalis
(NAV)
Neck of Malleus—Malleus, Collum mallei (NAV)
Nerve of Ampulla of Posterior Semicircular Canal—(Terry 1942);
Nervus ampullaris posterior (NAV)
Nerve of Pterygoid Canal—Nervus canalis pterygoidei (NAV); Vidian
Nerve (McDowell 1958)
Nerves of Saccule—(Terry 1942); Nervus saccularis (NAV)
Nerve of Utricle—(Terry 1942); Nervus utricularis (NAV)
Notochord—(Evans 1993)
Nuchal (= Lambdoid) Crest—Crista nuchae (NAV)
Occipital Artery—Arteria occipitalis (NAV)
Occipital Complex—(Wible 2003); Os occipitale (NAV)
Occipital Condyle—Condylus occipitalis (NAV)
Occipital Emissary Vein—Vena emissaria occipitalis (NAV)
Occiput—(NAV)
Oculomotor Nerve (= Cranial Nerve III)—Nervus oculomotorius (NAV)
Odontoid Notch—(Wible 2003); Incisura intercondyloidea (Evans
1993)
Olfactory Bulb—Bulbus olfactorius (NAV)
Olfactory Nerve (= Cranial Nerve I)—Nervi olfactorii (NAV)
Ophthalmic Artery—Arteria ophthalmica interna (NAV)
Ophthalmic Nerve (= Cranial Nerve V1)—Nervus ophthalmica (NAV)
Ophthalmic Vein—Vena ophthalmica interna (NAV)
Optic Foramen—Canalis opticus (NAV)
Optic Nerve (= Cranial Nerve II)—Nervus opticus (NAV)
Oral Cavity—Cavum oris (NAV)
Orbicular Apophysis of Malleus—(Henson 1961)
Orbit—Orbita (NAV)
Orbital Crest—Crista orbitalis (NAV)
Orbital Lamina of Ethmoid—Os ethmoidale, Lamina orbitalis (NAV)
Orbital Rim—Margo orbitale (NAV)
Orbital Surface of Lacrimal—Os lacrimale, Facies orbitalis (NAV)
Orbital Surface of Maxilla—Os maxillare, Facies orbitalis (NAV)
Orbitosphenoid—Os presphenoidale, Ala orbitalis (NAV)
Orbitotemporal Groove—(Rougier et al. 1992)
Orbitotemporal Canal—(Rougier et al. 1992); Sinus Canal (Gregory
Wible.indd 400
1910; McDowell 1958)
Orbitotemporal Crest—Crista orbitotemporalis (NAV)
Orbitotemporal Fossa—Orbita + Fossa temporalis (NAV)
Orifice of Canal for Modiolus—(Terry 1942)
Os Proboscidis—(Brandt 1833)
Osseous Lamina of Mallei—(Evans, 1993); Lamina (Henson 1961)
Osseous Nasal Septum—Septum nasi osseum (NAV)
Outer Lamella of Malleus—(Henson 1961)
Oval Window—Fenestra vestibuli (NAV)
Palatine—Os palatinum (NAV)
Palatine Canal—Canalis palatinus (NAV)
Palatine Process of Maxilla—Os maxillare, Processus palatinus (NAV)
Palatine Process of Premaxilla—Os incisivum, Processus palatinus
(NAV)
Palatine Surface of Maxilla—Os maxillare, Facies palatina (NAV)
Palatine Surface of Palatine—Os palatinum, Facies palatina (NAV)
Palatine Surface of Premaxilla—Os incisivum, Facies palatina (NAV)
Palatolacrimal suture—Sutura palatolacrimalis (NAV)
Palatomaxillary Suture—Sutura palatomaxillaris (Evans, 1993); Sutura
palatina transversa (NAV)
Paracondylar Process of Exoccipital—Processus paracondylaris (NAV);
Paroccipital Process of Exoccipital (Gregory 1910)
Paraflocculus of Cerebellum—Paraflocculus (NAV)
Parietal—Os parietale (NAV)
Parietal Foramen—(Boyd 1930, 1934)
Parietosquamosal Suture—Sutura squamosa (NAV)
Paroccipital Process of Petrosal—(Wible and Gaudin 2004); Mastoid
Process (Gregory 1910); Mastoid Eminence (MacPhee 1981)
Pars Canalicularis of Petrosal—(Wible 1990; Wible et al. 1995, 2001)
Pars Cochlearis of Petrosal—(Wible 1990; Wible et al. 1995, 2001)
Pars Glandularis of Hypophysis—(Evans 1993); Adenohypophysis
(NAV)
Pars Orbitalis of Frontal—Os frontale, Pars orbitalis (NAV)
Pars Processus Anterioris of Malleus*—(Henson 1961)
Pedicle of Incus—(Henson 1961)
Peduncle of Condylar Process of Mandible*—(Luo et al. 2002)
Perbullar Canal for Internal Carotid Artery*—(Wible, 1986)
Perilymphatic Duct—Ductus perilymphaticus (NAV)
Perilymphatic Vein—Vein of Cochlear Canaliculus (Patten 1942)
Permanent Teeth—Dentes permanentes (NAV)
Perpendicular Plate (Lamina) of Ethmoid—Os ethmoidale, Lamina
perpendicularis (NAV)
Perpendicular Process of Palatine—Os palatinum, Lamina perpendicularis
(NAV)
Petrobasilar Fissue—(Evans 1993); Fissura petrooccipitalis (NAV)
Petrosal (= Petrous Temporal)—Os temporale, Pars petrosa (NAV)
Petrosal Process of Rostral Process of Malleus—this study
Petrosquamous Suture—Sutura squamosomastoidea (NAV)
Petrotympanic Fissure*—(Evans 1993); Fissure petrotympanica (NAV)
Piriform Fenestra—(MacPhee 1981); Foramen Lacerum Medium
(Gregory 1910); Pyriform Fenestra (McDowell 1958)
Posterior Clinoid Process*—(Gregory 1910); Os basisphenoidalis,
Processus clinoideus caudalis (NAV)
Posterior Continuation of Crista Parotica—(MacPhee 1981); Lateral
Section of Caudal Tympanic Process of Petrosal (MacPhee 1981)
Posterior Crus of Ectotympanic—Annulus tympanicus, Crus posterior
(NAV)
Posterior Crus of Stapes—Stapes, Crus caudale (NAV)
Posterior Digastric Muscle—Musculus digastricus, Venter caudalis
(NAV)
Posterior Division of Ramus Superior of Stapedial Artery—(Wible and
Gaudin 2004)
Posterior Ligament of Incus—Ligamentus incudis posterius (NAV)
Posterior Nasal Spine—Spina nasalis caudalis (NAV)
Posterior Opening of Posttemporal Canal—(Wible and Gaudin 2004);
Posttemporal Foramen (Wible 2003)
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401
Appendix 1., continued
Posterior Semicircular Canal—Canalis semicircularis posterior (NAV)
Posterodorsal Ethmoidal Foramen—this study
Posterodorsal Process of Premaxilla—(Wible et al. in press)
Postglenoid Area of Squamosal—this study
Postglenoid Foramen—Foramen retroarticulare (NAV); Post-Glenoid
Foramen (Gregory 1910)
Postglenoid Process*—Processus retroarticulare (NAV)
Postglenoid Vein—Vena emissaria foraminis retroarticularis (NAV)
Postorbital Process*—Os frontale, Processus zygomaticus (NAV)
Postpalatine Torus—(McDowell 1958); Crista palatina (NAV)
Posttemporal Canal—(Rougier et al., 1992); Percranial Canal (MacPhee
1994)
Posttemporal Foramen (= Posterior Opening of Posttemporal Canal)
(Wible and Gaudin 2004)
Posttemporal Sulcus—Posttemporal Groove (Wible et al. 2001); Sulcus
for Diploic Vessels (Wible 1990)
Posttympanic Crest of Squamosal—(Wible et al. 2004)
Posttympanic Process of Squamosal—(Kielan-Jaworwoska 1981;
Novacek 1986); Processus retrotympanicus (NAV); Post-Tympanic
Process of Squamosal (Gregory 1910)
Prefacial (= Suprafacial) Commissure—(De Beer 1937)
Preglenoid Area of Squamosal—this study
Premaxilla—Os incisivum (NAV); Premaxillary (Gregory 1910)
Premolars—Dentes praemolares (NAV)
Presphenoid—Os presphenoidale, Corpus (NAV)
Presphenoid-Basisphenoid Synchondrosis—Synchondrosis intersphe noidalis (NAV)
Presphenoid Wing—this study
Primary Lamina of Ethmoturbinal I and II—(Smith and Rossie 2006)
Process for Digastric Muscle of Mandible—this study
Processus alaris, Ala temporalis—(De Beer 1937)
Promontorium of Petrosal—(Evans, 1993)
Prootic Canal—(Wible 1990; Wible and Hopson 1995; Rougier and
Wible 2006)
Prootic Sinus—(Wible 1990; Wible and Hopson 1995; Rougier and
Wible 2006)
Pterygoid Canal—Canalis pterygoideus (NAV)
Pterygoid Hamulus—Hamulus pterygoideus (NAV); Hamular Process
(McDowell 1958)
Pterygoid Veins—Venae pterygoideae (NAV)
Pterygopalatine Fossa—Fossa pterygopalatina (NAV)
Pterygopalatine Suture—Sutura pterygopalatina (NAV)
Ramus Inferior of Stapedial Artery—(Wible 1987)
Ramus Infraorbitalis—(Wible, 1987); Arteria maxillaries (NAV)
Ramus of Mandible—Ramus mandibulae (NAV)
Ramus Mandibularis—(Wible, 1987)
Ramus Posterior of Stapedial Artery—(MacPhee 1981)
Ramus Superior of Stapedial Artery—(Wible 1987)
Ramus Supraorbitalis—(Wible, 1987)
Ramus Temporalis of Stapedial Artery—(Wible, 1987)
Recessus Frontalis of Pars Intermedia of Nasal Capsule—(Smith and
Rossie 2006); Posterior Superior Recess (McDowell 1958)
Recessus Lateralis of Pars Intermedia of Nasal Capsule—(Smith and
Rossie 2006); includes Recessus Maxillaris and Recessus Frontalis
Recessus Maxillaris of Pars Intermedia of Nasal Capsule—(Smith and
Rossie 2006); (NAV)
Recessus Meatus—(Klaauw 1931; McDowell 1958)
Rectus Capitis Lateralis Muscle—Musculus rectus capitis lateralis
(NAV)
Rectus Capitis Ventralis Muscle—Musculus rectus capitis ventralis
(NAV)
Retromolar Space—Retromolar Fossa (Hiatt and Gartner 1987)
Rostral Alar Foramen*—Foramen alare rostrale (NAV)
Rostral Cranial Fossa—Fossa cranii rostralia (NAV)
Rostral Opening of Pterygoid Canal—(Wible and Gaudin 2004)
Rostral (= Anterior) Process of Malleus—Malleus, Processus rostralis
Wible.indd 401
(NAV); Folian Process of Malleus (McDowell 1958); Tympanic Plate
of Anterior Process of Malleus (Henson 1961)
Rostral Septal Branch of Major Palatine Artery—Rami septi rostrales
(Evans, 1993)
Rostral Tympanic Process of Petrosal—(MacPhee 1981)
Rostrum—(NAV)
Root of Teeth—Dentes, Radix dentis (NAV)
Round window—Fenestra cochleae (NAV)
Saccule—Sacculus (NAV)
Sagittal Crest—Crista sagittalis externa (NAV)
Sagittal Part of Vomer—(Evans 1993)
Secondary Facial Foramen—(Wible 1990; Wible and Hopson 1993)
Secondary Tympanic Membrane—Membrana tympani secundaria
(NAV)
Semicircular Canal—Canalis semicircularis (NAV)
Septal Branch of Caudal Nasal Nerve (= Nasopalatine Nerve)—(Evans,
1993); Nervus nasopalatinus (NAV)
Septal Process of Maxilla—(this study)
Septal Process of Nasal—Os nasale, Processus septalis (NAV)
Septal Process of Premaxilla—(this study)
Septum Frontomaxillare—(Smith and Rossie 2006)
Sigmoid Sinus—Sinus sigmoideus (NAV)
Shoulder of Anterior Crus of Stapes—(Henson 1961)
Shoulder of Posterior Crus of Stapes—(Henson 1961)
Sphenoethmoidal Suture—Sutura sphenoethmoidalis (NAV)
Sphenoid Complex—(Wible 2003)
Sphenoidal Fossa—(Evans 1993)
Spheno-occipital Synchondrosis—Synchondrosis spheno-occipitalis
(NAV)
Sphenopalatine Artery—Arteria sphenopalatina (NAV)
Sphenopalatine Foramen—Foramen sphenopalatinum (NAV)
Sphenopalatine Vein—Vena sphenopalatina (NAV)
Sphenoparietal Emissary Vein—Vena spheno-parietalis (Gelderen 1924)
Sphenorbital Fissure—(Gregory 1910; Novacek 1986); Fissura orbitalis +
Foramen Rotundum + Foramen alare rostrale + Foramen alare
parvum (NAV); Orbital Fissure (Evans 1993)
Spiral Cribriform Tract—Tractus spiralis foraminosus (NAV)
Spiral Ganglion—(Terry 1942); Ganglion spirale cochleae (NAV)
Squama frontalis—(NAV)
Squama of Squamosal—(Wible 2003)
Squamosal—Os temporale, pars squamosa (NAV)
Stapedial Artery—Arteria stapedia (Tandler 1899, 1902; Wible 1984,
1987)
Stapedial Foramen—Intercrural Foramen; Obturator Foramen (Henson
1961); Intracrural Foramen (Wible 2003)
Stapedius Fossa—(Wible 1990); Fossa for Musculus Stapedius (Evans
1993)
Stapedius Muscle—Musculus stapedius (NAV)
Stapes—(NAV)
Sternomastoid Muscle—(Allen 1910); Musculus sternomastoideus
(Evans 1993)
Styliform Process of Ectotympanic—(Klaauw 1931); Anteromesial Spine
of Ectotympanic (McDowell 1958); Anterior Process of Ectotympanic
(MacPhee 1981); Os temporale, Pars tympanica, Processus
muscularis (NAV)
Stylohyal—Stylohyoideum (NAV)
Stylohyal Facet on Tympanohyal—this study
Stylomastoid Artery—Arteria sylomastoidea (NAV)
Stylomastoid Foramen* —Foramen stylomastoideum (NAV)
Stylomastoid Notch—Foramen stylomastoideum (NAV)
Subarcuate Fossa—Fossa subarcuata (NAV); Floccular Fossa
(Gregory 1910)
Submandibular Gland—Glandula submandibularis (NAV); Submaxillary
Gland (Allen 1910)
Suboptic Foramen—(Gregory 1910)
Sulcus for Inferior Petrosal Sinus—Sulcus sinus petrosa ventralis (NAV)
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Appendix 1., continued
Sulcus for Occipital Emissary Vein—New term
Sulcus for Prootic Sinus—(Wible 1990)
Sulcus for Sigmoid Sinus—(Wible 1990)
Sulcus septi nasi—(Evans, 1993); Sulcus vomeris (septalis) (NAV)
Sulcus tympanicus—(NAV)
Superior Articular Facet of Malleus—(Henson 1961)
Superior Articular Surface of Incus—(Henson 1961)
Superior Vestibular Area—(Terry 1942); Area vestibularis superior
(NAV)
Suprameatal Crest—Dorsal Boundary of External Acoustic Meatus
(Evans 1993)
Suprameatal Foramen—(Novacek 1986); Subsquamosal Foramen
(Gregory 1910)
Supraoccipital—Os occipitalis, Squama occipitalis (NAV)
Supraoccipital Crest—this study
Supraoccipital Foramina—(Wible et al. 2004)
Supra-Optic Foramen—(Gregory 1910)
Symphyseal Surface of Mandible—(Evans 1993)
Tectorial Lamina of Ethmoid—Os ethmoidale, Lamina tectoria (NAV)
Tectum nasi—(Zeller 1987)
Tegmen Tympani—(De Beer 1937; Moore 1981); (NAV)
Temporal Fossa—Fossa temporalis (NAV)
Temporal Line—Linea temporalis (NAV)
Temporal Region—Regio temporalis (NAV)
Temporal Sinus*—(Evans 1993); Sinus temporalis (NAV); Capsuloparietal
Emissary Vein (Gelderen 1924)
Temporalis Muscle—M. temporalis (NAV)
Tensor Tympani Muscle—Musculus tensor tympani (NAV)
Tentorium Cerebelli—Tentorium cerebelli membranaceum (NAV)
Thyrohyal—Thyrohyoideum (NAV)
Thyroid Cartilage—Cartilago thryroidea (NAV)
Thyroid Foramen—Foramen thyroideum (NAV)
Tracheal Rings—Cartilagines tracheales (NAV)
Transverse Canal of Basisphenoid—(Gregory 1910)
Transverse Canal of Presphenoid—(Gregory 1910)
Transverse Canal Foramen—(Sánchez-Villagra and Wible 2002);
Transverse Canal (Gregory 1910); Foramen Vesalii (McDowell
1958)
Transverse Crest of Petrosal—Crista transversa (NAV)
Transverse Lamina—(Smith and Rossie 2006)
Transverse Sinus—Sinus transversus (NAV)
Transverse Sinus of Basisphenoid—(Gregory 1910)
Transverse Sinus of Presphenoid—(Gregory 1910)
Trigeminal Ganglion—Ganglion trigeminale (NAV); Gasserian or
Semilunar Ganglion (McDowell 1958)
Trigeminal Ganglion Fossa—Gasserian Fossa (McDowell 1958);
Trigeminal Fossa (Wible 1990); Semilunar Recess (Rougier et al. 1992)
Trigeminal Nerve (= Cranial Nerve V)—Nervus trigeminus (NAV)
Trochlear Nerve (= Cranial Nerve IV)—Nervus trochlearis (NAV)
Tympanic (= Middle Ear) Cavity—Cavum tympani (NAV)
Tympanic Canaliculus—Canaliculus tympanicus (NAV)
Tympanic Incisure—(Henson 1961)
Tympanic Nerve—Nervus tympanicus (NAV)
Tympanic Process of Alisphenoid—(MacPhee 1981); Tympanic Crest of
Alisphenoid (Gregory 1910); Preotic Crest of Alisphenoid
(McDowell 1958)
Tympanohyal—Tympanohyoideum (NAV)
Tympano-Occipital Fissure *—(Evans 1993); Fissura tympanooccipitalis
(NAV)
Tympanum—(NAV)
Vena diploëtica magna—(Hyrtl 1853, 1854)
Ventral Condyloid Foramina—(Wible and Gaudin 2004)
Ventral Condyloid Fossa—Fossa condylaris ventralis (NAV)
Ventral Nasal Concha (= Maxilloturbinate)—Os conchae nasalis
ventralis (NAV)
Ventral Nasal Meatus—Meatus nasi ventralis (NAV)
Vermis of Cerebellum—Vermis (NAV)
Vertical Part of Exoccipital—(Wible 2003)
Vestibular Aqueduct—Apertura externa aqueductus vestibuli (NAV);
Opening for Vestibular Aqueduct (Evans 1993)
Vestibular Fossula—(MacPhee 1981)
Vomer—(NAV)
Vomeronasal Organ—Organum vomeronasale (NAV)
Vomeropalatine Suture—Sutura vomeropalatina dorsalis (NAV)
Wings of Vomer—Alae vomeris (NAV)
Yoke—Jugum sphenoidale (NAV)
Zygoma*—Arcus zygomaticus (NAV)
Zygomatic Process of Lacrimal—(Wible 2003)
Zygomatic Process of Maxilla—Os maxillare, Processus zygomaticus
(NAV)
Zygomatic Process of Squamosal—Os temporale, Pars squama,
Processus zygomaticus (NAV)
Zygomaticus Minor— (Whidden 2002); Musculus zygomaticus (NAV)
Utricle—Utriculus (NAV)
Vagus Nerve (= Cranial Nerve X)—Nervus vagus (NAV)
Vascular Foramen of Lacrimal—(Giannini et al. 2006)
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