A Phylogenetic Analysis of Cranial Osteology in the Gerrhonotine Lizards Author(s): David A. Good Source: Journal of Herpetology, Vol.

21, No. 4, (Dec., 1987), pp. 285-297 Published by: Society for the Study of Amphibians and Reptiles Stable URL: http://www.jstor.org/stable/1563970 Accessed: 16/08/2008 13:44
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Journalof Herpetology, Vol. 21, No. 4, pp. 285-297, 1987 Copyright 1987 Society for the Study of Amphibians and Reptiles

A Phylogenetic Analysis of Cranial Osteology in the Gerrhonotine Lizards

DAVID A. GOOD
Museum of VertebrateZoology, University of California,Berkeley,California94720, USA
ABSTRACT.- The skulls of five genera and 13 species of gerrhonotine lizards were examined to accumulate characters useful in elucidating phylogenetic relationships within the group; 97 characters that are variable among the major species groups were found. Ancestral and derived states were estimated through outgroup comparison, using the other extant anguid subfamilies (the Anguinae and Diploglossinae) as outgroups. Among the hypotheses of relationships that were generated, one appears to be especially robust and it is

comparedwith those of previous workers.

Gerrhonotine lizards comprise a monophyletic (discussed by Gauthier, 1982) group of 38 currently recognized species distributed from southern British Columbia through the western United States, Mexico, and Central America as far south as western Panama. Despite numerous studies of the group (notably Smith, 1942; Tihen, 1949a, b;Stebbins, 1958; Criley, 1968; Waddick and Smith, 1974; Rieppel, 1980; and Gauthier, 1982), phylogenetic relationships among these species remain uncertain. Among past analyses, only those of Criley (1968), Rieppel (1980), Gauthier (1982), and to a limited extent Tihen (1949a) have involved discussions of osteology; all others have focused on external morphology. While Rieppel (1980) and Gauthier (1982) dealt extensively with skeletal material, their analyses were focused on a more inclusive group, the Anguimorpha; they did not analyze gerrhonotine relationships in depth. Tihen (1949a) briefly discussed some skeletal characters, but the material available to him was limited. Criley (1968) analyzed gerrhonotine cranial osteology with the view to discerning relationships within the subfamily, but he failed to find any systematically useful characters. I have reexamined skulls of all of the major groups of gerrhonotines and have found sufficient characters to permit construction of a robust hypothesis of relationships. In this paper I describe these characters and then discuss their phylogenetic implications in the light of previous phylogenetic hypotheses.

METHODS

Cranial material from the following five genera and 13 species was examined in order to find consistent qualitative differences among members of the subfamily (sample size in parentheses): Abroniadeppii (1), A. mixteca (1), A. oaxacae (1), A. taeniata (7), Barisiaimbricata(7), Elgaria coerulea (7), E. kingii(2), E. multicarinata(15), Gerrhonotus liocephalus(8), Mesaspis gadovii (3), M. monticola (3), M. moreleti (4), and M. viridiflava (3). Further data concerning these specimens are available on request from the author. Character states were scored for each of the species, and ancestral and derived conditions were determined by assuming that any condition characteristic of anguids outside of the Gerrhonotinae was ancestral for the subfamily. Because the relationships among the gerrhonotine subfamilies are unclear (as suggested by the conflicting hypotheses of McDowell and Bogert, 1954; Hoffstetter, 1962; Meszoely, 1970; Sullivan, 1979; Rieppel, 1980; Gauthier, 1982; and Good, 1987), a rigorous analysis of character state distributions among the outgroups, such as that discussed by Maddison et al. (1984), was impossible. A conservative approach was therefore taken and characters were ignored if more than one of the gerrhonotine states appeared outside of the subfamily or if none of them did. Many of the species included in this study are represented in collections by very little cranial material, often only single specimens. It is therefore impossible to de-

286

DAVIDA. GOOD

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and B. Elgaria 1. The skulls of: A. Gerrhonotus multicarinata. FIG. liocephalus,

termine the extent of intraspecific variation in these species. For this analysis, the character states seen in these species were assumed to be invariant if the states in species for which numerous specimens were available (see above) were invariant. Interspecific difference was considered to display no useful phylogenetic information if intraspecific variation in these relatively well represented species was high. Ontogenetic variation was largely ignored because juvenile specimens were lacking for most species. An analysis of ontogenetic variation and its relationship to the phylogenetic variation discussed here is presented elsewhere (Good, 1985).
CHARACTER ANALYSIS

Although Criley (1968) failed to find consistent variation in the skulls he examined, he admirably described the general condition of each element; I will therefore not describe them here. For ease of comparison with Criley's paper, the following character descriptions will be presented in the order in which he presented them.

The groups discussed below are essentially the genera as proposed by Tihen (1949a) except that his Barisia(sensu lato) is divided into two genera. Tihen's imbricata group is here placed in the genus Barisia (sensu stricto), and his gadovii and moreleti groups are combined into the genus Mesaspis(Good, 1985). Because osteological material from only one species was available from the gadovii group, it is referred to below as M. gadovii. The moreletigroup is referred to by that name. Grouped in this way, each of these taxa is monophyletic, although, on the basis of external characteristics (Good, 1985), the gadovii group of Tihen is paraphyletic when all of its species are included. The 97 characters used to diagnose these monophyletic units are summarized in Appendix 1. In the discussion below, the number in parentheses refers to the character number, and corresponds to the enumeration in the Appendix. Figs. 1-3 illustrate general skull morphology in the six major taxa included in the analysis. Premaxillae.--The ancestral position of

GERRHONOTINE OSTEOLOGY

287

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FIG.2.

The skulls of: A. Barisiaimbricata,and B. Mesaspis gadovii.

each medial ethmoidal foramen just inside the anterior end of the external naris is seen among gerrhonotines only in Elgaria. In Gerrhonotus,Barisia,and Mesaspis, an ossified bridge extends laterally from the nasal process of the premaxilla to isolate it from the naris (1). Abronia exhibits similarities to both of these conditions, possessing a lateral spur from the nasal process which, however, fails to completely isolate the foramen from the naris (2). This premaxillary bridge is broad in Barisia and much narrower in Mesaspis, being similar in width in the latter genus to the spur seen in Abronia. In Barisia,the premaxilla tends to extend less far onto the floor of the naris than it does in other forms (3). This floor is always comprised primarily of maxillary and septomaxillary elements, but the anterior part primitively tends to include a slight sloping shelf of premaxillary bone. Also seen only in Barisiaare two small foramina anterior and lateral to the medial ethmoidal foramen, one on the premaxilla and one on the premaxilla-maxilla suture (4). The width of the nasal process of the

premaxilla varies considerably among individuals of any given species, but there is a tendency toward its being broadened (5) in Barisia and narrowed (6) in Abronia. The process is parallel-sided between the nares in all forms except Barisia,in which it narrows posteriorly (7), and Gerrhonotus, in which it sometimes narrows anteriorly (8). The ancestral condition is parallel-sided. A derived contact of the nasal process with the frontal (excluding the nasals from contact with each other) (9) is seen in Barisia, in which contact is broad, and in M. gadovii and Abronia,in which it is narrow. Tihen (1949a) cited lack of contact in Abronia,but this is not corroborated by the present study. Lack of premaxillary-frontal contact in the moreleti group may be due to an anterior expansion of the nasals not seen in other gerrhonotines. In ventral view, there is some reduction of the size of the incisive process (10) in the moreleti group and a tendency toward convexity of this process (11) in Barisia, Mesaspis, and Abronia.A shelf on the posterior edge of the vomerine process running laterally from the incisive process

288

DAVIDA. GOOD

FIG.3. The skulls of: A. Mesaspis monticola,and B. Abroniamixteca.

creates a "channel" into the subnarial artery foramina in Mesaspis and Abronia(12). In all other gerrhonotines the ancestral condition is seen, with this shelf lacking and the slope into these foramina more gradual. Maxillae.-Contact with the premaxilla is broadened in all gerrhonotines, except and Elgaria,by the extension of Gerrhonotus the maxilla dorsomedially along the anterior edge of the naris (13). In M. gadovii, much of the bridge isolating the medial ethmoidal foramen from the naris is made up of a maxillary element rather than premaxillary (14). Widening of the premaxillary process appears to be concomitant with the deepening of the snout in Barisia(15). Three types of maxilla-prefrontal contact are seen in the Gerrhonotinae. That a straight suture line, seen in Gerrhonotus, is characteristic of the rest of the Anguidae. All other species except A. oaxacae,A. mixteca, and, to some extent, A. deppii,show a posteromedially pointed V-shaped pattern (16). The three Abroniaspecies show a lopsided W pattern. Contact with the frontal (17), a derived

and Abrocondition, is seen in Gerrhonotus nia. A supralabial groove usually runs anterodorsally from the jugal-maxillary suture (18) in Abronia,particularly A. deppii, A. oaxacae,and A. mixteca. The row of supralabial foramina is individually variable in foramen number; the only consistent trend is toward reduction in number in M. gadovii (19). Ventrally, the contact of the maxilla with the vomers between the premaxilla and the vomeronasal fenestra is narrowed in Barisia, Mesaspis, and Abronia (20). In addition, an ancestral, narrow, medially projecting process is present between the premaxilla and vomers in all gerrhonotines except Barisia(21). The palatal shelf is widened at the point at which it meets the maxillary process of the palatine in Barisia, Mesaspis,and Abronia (22). The anterior part of this shelf slopes strongly ventromedially in Mesaspis (23). It is ancestrally more-or-less horizontal. The maxilla extends posteriorly farther past the tooth row in Abroniathan in other genera (24). Vomers(Prevomersof Criley).-The pala-

GERRHONOTINE OSTEOLOGY tine processes of the vomers are elongate and, although the genus (25) in Gerrhonotus is rather variable, they are most divergent (26) in Abronia. The longitudinal ridges between the lateral and medial portions of the vomers slope more-or-less gradually in Mesaspis and Abronia (27), and are sharply defined in other gerrhonotines (the ancestral condition). The foramina (one on each vomer) for the medial palatine nerves penetrate straight through the bone in Mesaspis instead of showing the ancestral state by angling posteriorly (28). Palatines.-The palatines are widely divergent in A. deppii,A. oaxacae, and A. mixteca (29). Orientation of the palatines is variable in other forms but never reaches the condition seen in these species. Divergent palatines are found in most other anguids. Mesaspis is characterized by an anteromedial projection on the maxillary process of the palatine at its junction with the maxilla (30). The maxillary process curves outward to a greater degree in Abroniathan in any other gerrhonotine (31). It is more robust in Gerrhonotus, Elgaria,and Barisiathan in Mesaspis and Abronia(32). Although anguines have rather delicate processes, the condition in other anguids is robust and this is probably ancestral for the Gerrhonotinae. A pronounced dorsomedial flange, present on the dorsal surface of the vomerine process in Gerrhonotus,Elgaria, and Barisia, is absent or reduced in Mesaspisand Abronia (33); presence is ancestral. In ventral view, the lateral edge of the pterygoid process projects much farther posteriorly in all other genera than in Gerrhonotus(34). The reduced condition in this genus is derived. Contact of the palatines with the jugal on the anterior inside surface of the orbit has been excluded by the contact of the prefrontal with the maxilla in Gerrhonotus and Mesaspis (35). Ancestral contact is seen in most other gerrhonotines. The position of the infraorbital foramen is ancestrally near the ventral edge of the intraorbital surface of the palatine. This condition is seen in Gerrhonotus and Elgaria. The foramen is consistently more dorsally

289

located in Mesaspis and Abronia (36). The position in Barisiais variable. Ectopterygoids.-In ventral view, the suture joining the ectopterygoid with the pterygoid shows the ancestral U-shaped condition in Gerrhonotusand Elgaria,but is straight in Barisia, Mesaspis, and Abronia (37). The lateral spur of the ectopterygoid is usually elongate where it meets the maxilla in Elgaria (38). Pterygoids.-A prominent ridge on the posterodorsal edge of the transverse process is usually present in Barisia(39). Such a ridge is also often present in most other genera, but usually is not so prominently developed. The postepipterygoid groove on the dorsal surface of the pterygoid is usually reduced in Mesaspis and Abronia (40). Parietal.-There is a broadening of the parietal relative to its length in the moreleti group (41). When viewed from above, the parietal excludes the anterior end of the braincase from view in Abronia and most Gerrhonotus (42). It is visible ancestrally. The posterior edge of the parietal is rounded in dorsal or ventral view in A. deppii, A. oaxacae, and A. mixteca. The notched ancestral condition is present in all other gerrhonotines. The edges at the apex of this notch are twisted sharply downwardly in M. gadovii (43). Postorbitals. -The anterior end of the postorbital is expanded under the postfrontal in all forms except A. oaxacae and A. mixteca in which only a simple narrow slip of bone is present. A small process extending downward along the jugal lies anterior to it in many Elgaria rather than in the ancestral medial position (44). Posterior extension of the postorbital reaches the posterior end of the supratemporal fenestra and contacts the supratemporal bone in Mesaspis (45). This extension is usually broadened in M. gadovii (46). The posterior end of the supratemporal fenestra (but not the supratemporal bone) is also reached in anguines, but this seems to be the result of a reduction in fenestra size not seen in other anguids. -The postorbital bar (comSquamosals. of the posed squamosal and postorbital) lies on a plane with the parietal in Abronia, so that, in lateral view, the lateral edge of

290

DAVIDA. GOOD notines and this intermediate position is probably ancestral. Prefrontals.-In those groups with maxilla-frontal contact, the prefrontal is somewhat shortened. Contact with the maxilla is described above. In Barisia, an anteromedial arm extends anteriorly more than half way to the naris (55). This is a derived condition. The longitudinal supralacrimal ridge tends to be much more rounded (56) in Mesaspis and Abronia than in other gerrhonotines and the lacrimal foramen extends into the prefrontal to a greater degree (57) in Mesaspis.The tiny foramen on the dorsal surface of the prefrontal is closer to the lateral edge in Gerrhonotusand Elgaria than in other genera (58). This is ancestral. Palpebrals.--These elements were included by Criley (1968) in his discussion of the jugal. They are too often lost in preparation of skeletal material to be useful. Septomaxillae.--The septomaxilla, as viewed through the naris, is divided into anterior and posterior portions by an anteromedially-posterolaterally running ridge of individually variable distinctness. In all gerrhonotines except Mesaspis and Abronia,the ancestral condition is present, with the anterior portion much smaller than the posterior; in Mesaspisand Abronia, these portions are of approximately equal size (59). A spur, pointing anterolaterally and somewhat dorsally, is present on the lateral half of the dividing ridge in Gerrhonotus and Elgariaand may be the remnant of a prominent flange seen along the entire ridge in other anguids (60). The posterior edge of the posterior portion of the septomaxilla shows a bulge near its lateral end, making this end rather squared-off in appearance in A. oaxacaeand A. mixteca. The septomaxilla and maxilla form a relatively tight suture in the floor of the naris is all forms except Abronia, in which the septomaxilla overlaps the maxilla, especially at the lateral end of the dividing ridge (61). By examining intact skulls from the correct angle the posterior end of the septo-

the parietal is hidden (47). In A. oaxacae, the bar is bowed upward so that the parietal is visible below it. In all other gerrhonotines, the ancestral condition is seen, with the bar sloping downward laterally so that the edge of the parietal is visible, at least to some extent, above. The posterior end of the squamosal curls under itself in A. deppii,A. oaxacae,and A. mixteca. -Contact with the postSupratemporals. orbital is discussed above. There is considerable variation in the extent to which the supratemporal extends anteriorly along the medial edge of the supratemporal fossa and the degree to which it is hidden from above by the parietal; this seems to be variable among individuals of a species. The amount of parietal visible in lateral view ventral to the supratemporal is greater in Mesaspis than in other genera (48). Quadrates.-The cephalic condyle of the quadrate is expanded anteriorly in Barisia (49). It is either unexpanded or expanded laterally in other anguids. The constriction between the medial and lateral condyles of the articulating surface with the jaw is longer in M. gadovii than elsewhere (50). The quadrate is narrower in Abroniaand the moreletigroup than in other anguids. In Abroniathis is due to a narrowing of the lateral concave surface (51); in Mesaspisthe medial surface is reduced (52). Epipterygoids.-There is considerable variation in the degree of bowing of the epipterygoid, but this seems to be primarily either individual variation or the result of preparation. The epipterygoid is thinner in A. oaxacaeand A. mixteca than in other forms. Nasals.-Criley (1968) stated that the nasals are usually obscured by the cephalic osteoderms, particularly at their posterior end, but they are visible often enough to be useful. Contact of the nasals with each other in Gerrhonotus, Elgaria,and several Mesaspis is described above. The nasals fail to extend down the medial edge of the nares (53) in Barisiaand extend almost half way along it, thus causing loss of premaxilla-frontal contact (54), in the moreleti group. Extension is intermediate in all other gerrho-

GERRHONOTINE OSTEOLOGY maxilla can be observed. In most gerrhonotines the ancestral, well developed, straight posterior process is present. This process is reduced (62) in M. gadovii and absent (63) from the moreleti group. In Elgaria it curves upward posteriorly (64). More of the septomaxilla is present in dorsal view (65) in Abronia than in other genera, in which the posterior edge of the naris obscures much of it from view (the ancestral condition). Frontal. Variation in contact of the frontal with the maxilla and the premaxilla is discussed above. Nothing other than individual variation was otherwise observed. -The posterior part of the lacLacrimals. rimal (and the anterior part of the jugal) tend to bend outward at the suture with the maxilla in Gerrhonotus,Elgaria,and Barisia(66). This condition is not seen in other anguids. The groove running posteriorly from the lacrimal foramen into the orbit is almost entirely on the lacrimal in Abronia(67). In other genera the ancestral presence of prefrontal bone on the medial edge of this groove is seen. Also seen only in Abronia is a reduction of a jugal spur ventral to the posterior end of the lacrimal on the medial surface of the zygomatic bar (68). This is especially true of A. deppii, A. oaxacae, and A. mixteca. At the anterior end of the orbit is a depression leading anteriorly onto the posterior dorsolateral part of the snout. This depression is lined almost entirely by the prefrontal in all forms except A. deppii, A. oaxacae, and A. mixteca, in which the anterior end of the lacrimal extends into it. Jugals.-There is some variation in characters such as the degree of contact with the postfrontals, the size of the jugal process, and the positions of foramina, but intraspecific variation is much greater than interspecific so little phylogenetic evidence is suggested. Postfrontals.-Contact with other elements is discussed above. When viewed from below, the posterior end of the postfrontal is rounded (69) in Mesaspis and more-or-less pointed in other genera (the ancestral condition). The foramen on the posterior arm is more posteriorly located

291

in A. deppii,A. oaxacae, and A. mixteca than elsewhere. Basisphenoid.-The basisphenoid is a single medial element making up the anterior floor of the braincase. It is fused with the more superficial parasphenoid ventrally, creating the sphenoid discussed by Criley (1968). The basipterygoid processes are long and slender in A. oaxacae and A. mixteca relative to other anguids. The alar processes have become divergent in Abronia and M. monticola (70). Parasphenoid.-This element was discussed by Criley (1968) in his section on the sphenoid. No significant variation was observed. Supraoccipital.-The supraoccipital tends to be wider and shorter in the moreletigroup than in other anguids (71). The medial ascendent process at its anterior end makes a sharper angle with the main body of the element in Gerrhonotus than in other forms (72). This is a derived feature. The ridges on the dorsal surfaces of the paraoccipital processes are in such a configuration that they create a noticeable depression in all gerrhonotines except Abronia,which shows a derived condition (73). Exoccipitals.-Exoccipitals are fused in the adult to the opisthotics to form the otoccipitals of Criley (1968). No significant variation was observed in these elements. Opisthotics.-Opisthotics were discussed by Criley (1968) in his section on the otoccipitals. The major variation observed in this element was the angle at which the paraoccipital processes diverge from the braincase (74). In Abronia and M. gadovii, these processes tilt upward at a greater angle than they do in other anguids, in which they are in a more-or-less horizontal plane. The reduced size of the moreleti group species results in a crowding of the braincase within the surrounding case of dermal bones, which may have a confounding effect on this divergence angle. There is a lateral projection from the posterior surface above the basioccipital process in all forms except Mesaspis (75). The projection is reduced in Abronia (76). (1968) reported Basioccipital.-Criley considerable variation in the size and shape

292

DAVIDA. GOOD angular (just above the condylar facet) in all gerrhonotines except Gerrhonotus (83). Coronoids. -In medial view, the angle of the posteroventral process of the coronoid is less oblique (i.e., closer to vertical) in Gerrhonotus than in other anguids (84). In M. gadovii, this process tends to be curved so that its end points almost directly posteriorly (85). In Gerrhonotus and Elgaria the coronoid projects anteriorly on the lingual surface of the jaw past the posteriormost tooth (86). A derived condition in which this is not the case is seen in Barisia, Mesaspis, or Abronia. Dentaries.--A derived sublabial longitudinal groove on the lateral surface of the dentary is present in Barisia,Mesaspis, and Abronia (87). The subdental shelf on the lingual surface is usually reduced in Mesaspis and Abronia(88). -The relative positions of the Splenials. anterior inferior alveolar foramen and the anterior mylohyoid foramen vary (89). In Elgaria, Barisia, Mesaspis, and Abronia, the anterior end of the mylohyoid foramen is approximately directly ventral to the posterior end of the alveolar foramen (or perhaps only slightly posterior to it). In Gerrhonotus, the foramina are much more condition widely spaced. The Gerrhonotus is ancestral. Less anterior projection of the splenial (90) is seen in Abroniathan in other genera and the posteriormost end of the bone tends to lie on the ventral edge of the jaw in A. deppii, A. oaxacae, and A. mixteca, while it extends somewhat onto the lingual surface in other forms. Angulars.-Little phylogenetic variation was observed in this element. Teeth. The retention of the ancestral and pterygoid teeth is seen in Gerrhonotus Elgaria (although they are somewhat reduced in E. coerulea).Reduction (91) is seen in Barisiaand complete loss (92) in Abronia and, usually, Mesaspis. Number and shape of maxillary, premaxillary, and dentary teeth are often mentioned as being of use in phylogenetic analysis in the Gerrhonotinae. For instance, Gauthier (1982) cited pointed, del-

of the basioccipital processes, and found it to be dependent on the size of the skull. The probable ancestral condition of the occipital condyle, large and bulbous with its widest part being separated from the braincase by a slight constriction, is seen in Gerrhonotus,Elgaria, and Barisia. In all other forms, it is more delicately constructed and its widest point is where it joins the braincase (77). -There is much variation in the Prootics. size and shape of the alar processes and in the sutures of the ventral processes with the basisphenoid, but there does not seem to be a phylogenetic component to this variation: all conditions exist in all taxa. -The orbitosphenoids Orbitosphenoids. are small crescent-shaped ossifications on the posterior surface of the orbit. Since they do not contact any of the other bony elements of the skull, they are usually lost in preparation and so are of little use to this analysis. They were not discussed by Criley (1968). Prearticulars. These elements were incorrectly referred to by Criley (1968) as the articulars. The retroarticular process tends to be elongate (78) in Abronia and has a characteristic triangular shape in A. oaxacae and A. mixteca. The ventral point of this process is often somewhat hooked in M. tends to exhibit a gadovii (79). Gerrhonotus derived ventral bulging of the middle portion of the prearticular, anterior to the retroarticularprocess, giving the posterior end of the jaw a more robust appearance (80). Articulars.-Although the term "articular" is usually used in the literature to refer to the fused articular (as here used) and prearticular, the articular proper is a small element in the immediate vicinity of the condylar surface for articulation with the skull; the larger element including the retroarticular process is the prearticular. A derived shifting of the surangular-articular suture into the articular facet is seen in Barisia,Mesaspis, and Abronia(81). Surangulars.--The surangular is broader than in any othand stouter in Gerrhonotus er gerrhonotine and constitutes a derived state (82). There is an expansion of the dorsal edge at the posterior end of the sur-

OSTEOLOGY GERRHONOTINE icately constructed, recurved, widely spaced teeth as a synapomorphy of Barisia (sensu Tihen, 1949a, b) and Abronia. Unfortunately, tooth shape is much more variable intraspecifically than has been suggested. Some tendencies are observable, however. Long, sharp, strongly recurved teeth (93) are seen in Gerrhonotusand Abronia and, despite Gauthier's (1982) suggestion to the contrary, Barisia (sensu stricto) has short, rather blunt teeth (94). All other gerrhonotines tend to have intermediate conditions. The ancestral condition is unfortunately impossible to discern as all of these tooth shapes are seen elsewhere in the Anguidae, even within the single subfamily Anguinae. The bicuspid tooth form discussed by Gauthier (1982) as characteristic of the Gerrhonotinae is virtually lost in A. deppii, A. oaxacae, and A. mixteca. In tooth number, the only clearly derived state seen was a general increase in Gerrhonotus (95). Maxillary/premaxillary teeth number 14-18 in all forms except this genus, which has 21-24. Dentary teeth ancestrally number 18-23 in all other genera; in Gerrhonotusthey number 27-30. Teeth are somewhat more widely spaced in some Abronia(96). Osteoderms.-Heavy, deeply sculptured cephalic osteoderms are characteristic of Barisia,Mesaspis, and Abronia(97). This is a derived condition. ANALYSIS PHYLOGENETIC Hypotheses of ancestral versus derived states in characters variable within the Gerrhonotinae are provided in the preceding section. With these hypotheses, it is possible to estimate phylogenetic relationships among the gerrhonotines based on the realization that only shared derived character states (synapomorphies) provide any useful information about the genealogies of the organisms that possess them; shared primitive states (symplesiomorphies) do not (cf. Hennig, 1966). The monophyly of each of the major groups of gerrhonotines is supported by this analysis, although it must be remembered that, due to lack of material, many

293

species were not included. Gerrhonotusis uniquely diagnosed by eight derived character states, Elgaria by three, Barisiaby 11, M. gadovii by seven, the moreleti group by five, and Abronia by 17 (Appendix 1). Relationships among these groups are also suggested by unique synapomorphies (Appendix 1). Mesaspis (the moreletigroup and M. gadovii) possesses ten synapomorphies and the position of Abronia as its sister group is suggested by 11. Barisia, Mesaspis, and Abronia are allied by 13 synapomorphies. Elgaria is linked with them by three. It is likely that some of the characters used in this analysis are not independent. For instance, the ventral bulging of the prearticular (character 80), the broad, stout nature of the surangular (character 82), and perhaps even the vertical position of the posterior arm of the coronoid (character 84) and the increase in tooth number on the dentary (character 95) in Gerrhonotus are all manifestations of an overall increase in robustness of the jaw which might be better considered a single autapomorphy, rather than four autapomorphies, of the genus. However, all of the phylogenetic relationships discussed above are suggested by a variety of characters as independent as is possible in such a cohesive structure as a lizard skull, and combining such potentially non-independent characters would have no effect on the results of the analysis. Because of this, and because I cannot demonstrate that these possibly nonindependent characters are certainly nonindependent, I here discuss them separately. The hypothesis of relationships among the major groups of gerrhonotines analyzed is illustrated in Fig. 4. Only eight characters conflict with it. One of these, the outward bend of the lacrimal and jugal at the suture with the maxilla in Gerrhonotus, Elgaria, and Barisia (character 66), is best explained by the failure of outgroup comparison to suggest the true ancestral condition for the subfamily. If this outward bend is considered ancestral even though it is not seen in other anguids, the character ceases to conflict with Fig. 4. This reversal of polarity also best explains the

294

DAVIDA. GOOD
Gerrhonotus

aria / vElgaria

spur/bridge is not polarizable, so it is impossible to determine whether the broad Barisiacondition or the narrow Mesaspis/
Abronia condition is derived. 2) Frontal-

maxilla contact (character 17) arose sepaar \Barisia

a/iM.

gadovii

moreleti

group

A bronia

of palatine-jugal contact inside the orbit (character 35) has occurred in Gerrhonotus and Mesaspis. 4) Gerrhonotusand Abronia share the loss of the anterior end of the braincase from dorsal view (character 42). 5) Divergent basisphenoid alar processes and M. monticola(character occur in Abronia
70) 1958; Waddick and 1980; and and Gauthier, Gauthier, Smith, 1974; Rieppel, 1980; 1949a, b; Stebbins,

rately in Gerrhonotus and Abronia. 3) Loss

the major groupsof gerrhonotinelizardsas suggested by analysisof cranialosteology.Rieppel,

4. Proposed phylogenetic relationships among FIG.

Seven major studies (Smith, 1942; Tihen,

apparently ancestral wide divergence of the palatines in A. deppii, A. oaxacae, and A. mixteca (character 29). Two characters show conditions in some forms which might explain their failure to agree with the hypothesis. Expansion of the nasals in the moreletigroup (character 54) is likely responsible for the failure of the premaxilla to contact the frontal. If this lack of contact is therefore considered secondary, contact can be suggested as a further synapomorphy of the group containing Barisia, Mesaspis, and Abronia. The crowding of the braincase within its shell of dermal bone might explain the failure in the moreleti group of the paraoccipital processes to show the upward angle seen in Abronia and other Mesaspis(character 74). This angle may therefore be another synapomorphy of these two genera. The patterns seen in the remaining five characters can best be explained as the result of homoplasy; using the criterion of parsimony, none of them suggests a likely hypothesis of relationships in view of the weight of opposing evidence provided by the characters discussed above. These five characters are: 1) Isolation of the medial ethmoidal foramen from the external naris (character 1). This apparently arose twice, once in Gerrhonotus and once in the ancestor of Barisia, Mesaspis, and Abronia. This assumes that the premaxillary spur seen in Abronia developed from the bony bridge seen in the other genera. The width of the

1982) have been conducted which include discussions of higher level intra-gerrhonotine relationships. None of these workers except Gauthier (1982) recognized the paraphyly of Barisia(sensu lato, including both Barisiaand Mesaspis as circumscribed here). Gauthier recognized that Abronia might be more closely allied with some of these "Barisia"than with others, and this is substantiated by the present analysis. All other workers placed Abroniaoutside of all other genera (except sometimes Coloptychon,a very rare form unrepresented in the present analysis). The species here referred to Gerrhonotus, Elgaria, Barisia, and Mesaspis almost universally have been considered to form a monophyletic group (except, again, by Gauthier, 1982). Hypotheses of relationships among these species have varied. Smith (1942) saw his antaugesgroup (=Mesaspis) as the most primitive of the group. Tihen (1949a) and Rieppel (1980) suggested that instead Gerrhonotusis the sister group to all others and that Tihen's Barisia (=Barisia and Mesaspis) is a monophyletic assemblage, with (according to Tihen, 1949b)the gadoviiand imbricata groups being sister taxa. Stebbins (1958) suggested that two clades are present, one containing Eland the other Barisia garia and Gerrhonotus (sensu lato) and E. coerulea. Waddick and Smith (1974) essentially agreed with this view except that they replaced E. coerulea in Elgaria. Tihen's (1949a) placement of Abronia

OSTEOLOGY GERRHONOTINE outside of all other gerrhonotines was based on the presence in that genus of a weak lateral fold, which he considered primitive (although it probably is not; Good, 1985), and osteoderms similar to the fossil Peltosaurus, a glyptosaurine whose close ties to the Gerrhonotinae are by no means clear. Smith (1942) had given no reason for his placement of Abroniain this position and most workers subsequent to Tihen essentially accepted it without question. Gauthier (1982), therefore, presented the first useful phylogenetic information for the placement of Abronia. Aside from Smith's (1942) placement of Mesaspis (his "antauges group") outside of the remaining genera, an hypothesis for which he gave no explanation, two basic hypotheses have been suggested for the relationships of Gerrhonotus, Elgaria,Barisia, and Mesaspis.The view that Gerrhonotus and form a natural 1958; Elgaria group (Stebbins, Waddick and Smith, 1974) is based on the sharing of an elongate body form, oviparity, and preference for relatively warm, dry habitat (Stebbins, 1958), and lack of anterior internasal contact (Waddick and Smith, 1974). All of Stebbins' characters are probably plesiomorphic and that of Waddick and Smith is not a synapomorphy since anterior internasal contact is lost in different ways in the various genera (Good, 1985). Hence no evidence exists to refute the alternate view (Gauthier, 1982), corroborated by this analysis, that Elgaria, Barisia, Mesaspis, and Abroniaform a natural group relative to Gerrhonotus. This analysis strongly suggests a single hypothesis for the relationships among the major groups of gerrhonotine lizards (Fig. 4). Although no really major variations were observed, the "highly surprising lack of consistent or characteristic osteological differences" cited by Criley (1968) was certainly not seen. Ample character variation was found. This situation points up the fact that care should be taken not to accept higher level groupings without real evidence for them. Criley's (1968) acceptance of the genera as proposed by Tihen (1949a) was in part responsible for his inability to find diagnostic features in the subfamily. He looked,

295

for instance, for characters to diagnose the genus Barisia,which, as he defined it, contained both Barisiaand Mesaspis, and failed to find any; none were found in the present analysis either. This is, of course, because Abronia is phylogenetically closer to Mesaspis than is Barisia (sensu stricto) and all of the derived features of both of the latter two are shared by the former. Criley should have analyzed the gerrhonotines on a species by species level, without reference to preconceived notions of relationship. In this paper I have suggested an hypothesis of relationships among the alligator lizards which is strongly suggested by the data presented. Although much can be learned about many attributes of an organism such as an alligator lizard without reference to its phylogenetic position, without knowledge of phylogeny nothing can ever be known about the evolution of these attributes. This analysis represents a step toward providing a genealogical framework for future comparative work on this fascinating group of lizards which shows a wide range of distributional, ecoand logical, behavioral, physiological, attributes. of these will morphological Many be discussed at length in subsequent papers. Acknowledgments.-This work was conducted as part of a Ph.D. dissertation at the University of California, Berkeley, and I would like to thank both the Zoology Department and the Museum of Vertebrate Zoology there for support and encouragement throughout my career as a graduate student. The following curators of herpetological collections provided specimens: Pere Alberch, Museum of Comparative Zoology, Harvard University (MCZ); Arnold G. Kluge, University of Michigan Museum of Zoology (UMMZ); Charles W. Myers, American Museum of Natural History (AMNH); John W. Wright, Natural History Museum of Los Angeles County (LACM); and George R. Zug, United States National Museum (USNM). I would like to thank Thomas Duncan, Harry W. Greene, David B. Wake, Aaron M. Bauer, and Kevin de Queiroz for useful

296

DAVID A. GOOD
Characters diagnostic of Gerrhonotus: 8. Nasal process of the premaxilla narrowing anteriorly between the external nares. 25. Elongate palatine processes of the vomers. 34. Reduction in posterior projection of the palatines at the junction with the pterygoids. 72. Ascendent process of the supraoccipital angled sharply upward. 80. Middle part of prearticular bulging ventrally. 82. Surangular broad and stout. 84. Posterior arm of the coronoid more-or-less vertical. 95. Increase in tooth number. Characters diagnostic of the group containing Elgaria, Barisia,Mesaspis,and Abronia: 16. Loss of a straight maxilla-prefrontal suture. 83. Expansion of the dorsal edge of the posterior end of the surangular. 89. Narrowing of the separation of the anterior inferior alveolar foramen and anterior mylohyoid foramen. Characters diagnostic of Elgaria: 38. Elongate lateral spur of the ectopterygoid at the junction with the maxilla. 44. Anterior placement of the jugal process of the postorbital. 64. Upward curve of the posterior process of the septomaxilla. Characters diagnostic of the group containing Barisia, Mesaspis, and Abronia: 11. Convex incisive process of the premaxilla. 13. Extension of the maxilla along the anterior end of the naris. 20. Narrow maxilla-vomer contact. 22. Widened palatal shelf of the maxilla where it meets the palatine. 36. Dorsal shift in the position of the infraorbital foramen. 37. Straight pterygoid-ectopterygoid suture. 58. Medial placement of the foramen on the dorsal surface of the prefrontal. 60. Loss of the septomaxillary spur. 81. Placement of the surangular-articular suture on the condylar facet. 86. Loss of the projection of the coronoid anterior past the posterior teeth. 87. Sublabial groove on dentary. 91. Reduction of pterygoid teeth. 97. Heavy, deeply sculptured cephalic osteoderms. Characters diagnostic of Barisia: 3. Reduction of the amount of premaxillary bone on the floor of the naris. 4. Presence of two small foramina on the premaxilla anterior and lateral to the medial ethmoidal foramen. 5. Broad nasal process of the premaxilla. 7. Nasal process of the premaxilla narrows posteriorly between the external nares. 15. Broad premaxillary process of the maxilla.

comments and discussions during the preparation of this paper. The critical comments of Paul E. Hertz, associate editor of the Journal of Herpetology, and two anonymous reviewers are also appreciated.
LITERATURE CITED B. B. 1968. The cranial osteology of gerCRILEY, rhonotiform lizards. Am. Midi. Nat. 80:199-219. GAUTHIER, J. A. 1982. Fossil xenosaurid and anguid lizards from the early Eocene Wasatch Formation, southeast Wyoming, and a revision of the Anguioidea. Contr. Geol., Univ. Wyoming 21:7-54. GOOD,D. A. 1985. Studies of interspecific and intraspecific variation in the alligator lizards (Lacertilia: Anguidae: Gerrhonotinae). Unpubl. Ph.D. dissertation, Univ. California, Berkeley. 622 pp. . 1987. An allozyme analysis of anguid subfamilial relationships. Copeia 1987:696-701. W. 1966. Phylogenetic systematics. Univ. HENNIG, Illinois Press, Urbana. 263 pp. R. 1962. Observations sur les osteoHOFFSTETTER, derms et la classification des anguides actuels et fossiles (reptiles, sauriens). Mus. Natl. d'Hist. Nat., Paris, ser. 2, 34:149-157. MADDISON, W. P., M. J. DONOGHUE,AND D. R. MADDISON. 1984. Outgroup analysis and parsimony. Syst. Zool. 33:83-103. S. B., ANDC. M. BOGERT. 1954. The sysMCDOWELL, tematic position of Lanthanotusand the affinities of the anguinomorphan lizards. Bull. Amer. Mus. Nat. Hist. 105:1-142. C. A. M. 1970. North American fossil MESZOELY, anguid lizards. Bull. Mus. Comp. Zool. 139:87-150. 0. 1980. The phylogeny of anguinomorph RIEPPEL, lizards. Birkhauser Verlag, Basel. 86 pp. H. M. 1942. Mexican herpetological miscelSMITH, lany. 3. A tentative arrangement and key to Mexican Gerrhonotus,with the description of a new race. Proc. U.S. Nat. Mus. 92:363-369. R. C. 1958. A new alligator lizard from the STEBBINS, Panamint Mountains, Inyo County, California. Am. Mus. Novitates 1883:1-27. R. M. 1979. Revision of the Paleogene SULLIVAN, genus Glyptosaurus (Reptilia, Anguidae). Am. Mus. Nat. Hist. Bull. 163:1-72. TIHEN, J. A. 1949a. The genera of gerrhonotine lizards. Am. Midl. Nat. 41:580-601. . 1949b. A review of the lizard genus Barisia. Univ. Kansas Sci. Bull. 33:217-256. WADDICK, J. W., AND H. M. SMITH. 1974. The significance of scale characters in evaluation of the lizard genera Gerrhonotus, Elgaria,and Barisia.Great Basin Nat. 34:257-266. Accepted: 8 October 1986. 1 APPENDIX Derived cranial character states diagnostic of monophyletic groups within the Gerrhonotinae. The character numbers correspond to those in the text.

GERRHONOTINE OSTEOLOGY
21. Loss of the medial process of the maxilla between the premaxilla and the vomers. 39. Prominent dorsal ridge on the ectopterygoid. 49. Anterior expansion of the cephalic condyle of the quadrate. 53. Failure of the nasals to extend down the medial edge of the external nares. 55. Extension of the prefrontals more than half way to the nares. 94. Short, blunt teeth. Characters diagnostic of the group containing Mesaspis and Abronia: 12. Pronounced channel into the subnarial artery foramina on the palatal process of the premaxilla. 27. Gradual slope from ventral surface into medial depression of the vomers. 32. Delicate maxillary process of the palatines. 33. Reduction of the dorsal flange on the vomerine process of the palatines. 40. Reduction of the postepipterygoid groove on the pterygoid. 56. Rounding of the supralacrimal ridge. 59. Approximately equal size of the anterior and posterior portions of the septomaxilla. 76. Reduction of the lateral projection from the posterior surface of the otoccipital. 77. Reduction in robustness of the occipital condyle. 88. Reduction of the subdental shelf of the dentary. 92. Loss of pterygoid teeth. Characters diagnostic of Mesaspis: 23. Ventromedial slope of the palatal shelf of the maxilla. 28. Medial palatine nerve foramina on the vomers penetrating straight through the bone. 30. Anteromedial projection on the maxillary process of the palatine at its junction with the maxilla. 45. Extension of the postorbital to the posterior end of the supratemporal fenestra. 48. Increase in the amount of parietal ventral to the supratemporal in lateral view. 52. Narrowing to the medial concave surface of the quadrate. 57. Enhanced extension of the lacrimal foramen onto the prefrontal. 62. Reduction in the posterior process of the septomaxilla. 69. Rounded posterior end of the postfrontal. 75. Loss of the lateral projection from the posterior surface of the opisthotic. Characters diagnostic of Mesaspis gadovii:

297

14. Maxillary rather than premaxillary bridge between the medial ethmoidal foramen and the naris. 19. Reduction in number of supralabial foramina. 43. Downward twist on the edges of the posterior notch in the parietal. 46. Broadened postorbital. 50. Elongate articular facet of quadrate. 79. Hooked ventral point of the retroarticular process. 85. Horizontal direction of the posterior arm of the coronoid. Characters diagnostic of the moreleti group of Mesaspis: 10. Reduction in size of the incisive process of the premaxilla. 41. Broadened parietal. 54. Extensive expansion of the nasals anteriorly along the medial edge of the external nares. 63. Loss of the posterior process of the septomaxilla. 71. Widened, shortened supraoccipital. Characters diagnostic of Abronia: 2. Premaxillary spur between the medial ethmoidal foramen and the naris. 6. Narrow nasal process of the premaxilla. 18. Supralabial groove on the maxilla. 24. Enhanced posterior extension of the maxilla posterior to the teeth. 26. Divergent palatine processes of the vomers. 31. Outward curve of the maxillary process of the palatine. 47. Upward bowing of the postorbital bar. 51. Narrowing of the lateral concave surface of the quadrate. 61. Loss of tight contact of the septomaxilla and maxilla. 65. Increase in amount of septomaxilla visible in dorsal view through external naris. 67. Groove running posteriorly from the lacrimal foramen entirely on the lacrimal. 68. Loss of jugal spur ventral to the lacrimal on the inside of the zygomatic bar. 70. Divergent basisphenoid alar processes. 73. Loss of the depression on the dorsal surface of the paraoccipital processes. 78. Elongate retroarticular processes. 90. Decrease in anterior projection of the splenial. 96. Widely spaced teeth.