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Phylogenetic Relationships Within Squamata Richard Estes, Kevin De Queiroz, and Jacques Gauthier

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Phylogenetic Relationships Within Squamata Richard Estes, Kevin De Queiroz, and Jacques Gauthier Phylogenetic Relationships within Squamata Richard Estes, Kevin de Queiroz, and Jacques Gauthier INTRODUCTION Previous analyses of Squamata or "Lacertilia" were hampered by the lack of a well-corrobo- rated hypothesis of relationhips within Lepidosauria and their fossil relatives. The present attempt at réévaluation of squamate phylogeny was prompted by several factors. One impetus grew out of the compilation of the "lizard" fossil record given by Estes (1983a) in the Handbuch der Paläoherpetologie. The diagnoses of the groups prepared for the Handbuch provided an extensive initial list of osteological characters for both fossil and Recent "lizards." Camp (1923), who we honor with the papers in this volume, made extensive use of osteological characters in his Classi- fication of the Lizards. Supplementation and modification of his list of characters is therefore desir- able as one means of evaluating various aspects of Camp's hypotheses of squamate relationships. Availability of a phylogenetic systematic study of reptiles (Gauthier, 1984) and the related study on lepidosauromorph reptiles (Gauthier et al., 1988) was another factor important to the de- velopment of the present analysis. For the first time it is possible to analyze relationships within Squamata in light of a well corroborated hypothesis of relationships among squamates and their closest relatives. Our objectives in this paper are: first, to obtain a minimum-step computer-generated clado- gram of currently recognized "lizard" families based on a large number of osteological characters and characters from soft anatomy, including many of those used by Camp (1923); and second, to evaluate these results, identifying the principal problem areas, and offering a less highly resolved but better supported cladogram that can be used as an estimate of the actual phylogeny. METHODS CHARACTERS AND OUTGROUPS Ckiracters Osteological characters are the ones most applicable to the study of fossils, and it is our even- tual goal to use this data set to determine the relationships of fossil squamates to their living rela- tives. Most of the characters have been taken from the diagnoses of "lizard" families and suprafa- niilial taxa in Estes (1983a); others were offered by various colleagues or have come to our atten- tion during the preparation of this paper. Our present list includes 130 osteological characters and 18 from soft anatomy, a total of 148. Three kinds of characters have been omitted: (1) Characters whose derived states are restricted to a single family. These derived characters may serve to diag- nose that family, yet provide no information about interfamilial relationships. Although these characters are omitted from our character list, they are included in the diagnoses of our basic taxa. (2) Characters whose within-taxon variation is so great that it obscures the pattern of between- taxon variation. Such characters may be useful when working with less inclusive levels. (3) Characters that did not vary within Squamata or more inclusive groups. Such characters provide no information on relationships within Squamata; they are discussed where they exist as synapo- morphies of more inclusive taxa (Gauthier et al., 1988). The source of many of our osteological characters (Estes, 1983a) is itself largely a compila- tion and summary of information in numerous papers on both fossil and recent "lizards". Many of the characters used there to diagnose taxa were taken directly from the literature. Where detailed 120 Phylogenetic Relationships of the Lizard Families surveys or discussions of particular characters exist, we have cited these studies in our character de- scriptions, and in most cases have checked these characters on specimens. For new or undescribed characters, we have examined representatives of most of the "lizard" families personally as well as checking these on published figures. Another reason for emphasizing osteological characters in the present work is that they are readily determined in a wide variety of extant groups. Although Camp (1923) himself placed great importance on osteological characters, important aspects of his contribution lay in the analysis of characters of the soft anatomy, especially throat muscles. We have supplemented our list of osteo- logical features with a limited number of characters from soft anatomy. This list is limited be- cause it is more difficult to obtain a complete data set for characters of the soft anatomy than for osteological features; many workers in this area have studied only particular groups and preserved material is not always available to complete the data set. Variation in characters of the soft anato- my is less easily interpreted because of the incomplete data sets, and there is moreover a greater dif- ficulty in making polarity decisions owing to lack of knowledge of these characters in fossil out- groups. All of our characters are presented as linear and unidirectional transformation series. In other words, each numbered state for a given character is intended to imply direct transformation of the state designated by the next lower integer. We purposely described the characters in this way for two related reasons: (1) so that the character states of different characters would be direcdy compar- able as derived states (e. g., state 2 is always derived from state 1, not sometimes from state 0), and (2) so that the taxon-character matrix could be analyzed using a computer algorithm without reced- ing any of the characters. At the time of our study, options for treating multistate characters as unordered were not available; we suggest that this option be considered in the future for similar studies. The description of all characters as linear and unidirectional transformation series results in some unconventional characters, for example, characters 107 and 108. Because the transformation has gone in two different directions (eight cervical vertebrae is thought to be primitive; reduction and increase from this number have both occurred), the number of cervical vertebrae has been split into two characters. The result is that the plesiomorphic states of both characters contain what we believe to be derived morphologies, specifically those morphologies considered to be the derived state in the other member of the character pair. Because the character is thought to have trans- formed in two directions, we are simply focusing on one of those transformations while lumping all other morphologies as "not-such". In most cases (except reversals) this should create no prob- lems in phylogenetic analysis, because the heterogenous state is always the primitive one and we form groups only with derived states. ( Squamate Monophyly We have used outgroup comparison (Watrous and Wheeler, 1981; Farris, 1982; Maddison et al., 1984) for inferring character polarities. In order to use this method, it is desirable that the group under study (ingroup) be monophyletic; otherwise, groups used as outgroups that are actual- ly derived from within the ingroup can cause errors in polarity assessment. Gauthier et al. (1988) presented abundant evidence that Squamata, including "lizards," snakes, and amphisbaenians, is a monophyletic taxon. Within Squamata, however, the question arises as to whether snakes and am- phisbaenians are potential outgroups of "lizards" simply because they are placed in separate taxa, or whether they arose within "lizards." Rage (1982a) listed three presumed synapomorphies favoring a monophyletic Sauria (= Lacertilia of some authors) exclusive of amphisbaenians and snakes. All three of these synapomorphies are inadequate evidence for "lacertilian" monophyly. The first, "nerf J PHYLOGENY OF THE SQUAMATA - Estes et al. 121 glossopharyngien sortant du crâne par l'apertura lateralis recessus scalae tympani (état primitif chez les Lepidosauriens, le nerf glossopharyngien sort avec le nerf vague par le foramen jugulaire)," also occurs in most snakes (Rieppel, 1980b), and is variable in dibamids (Rieppel, 1984a; Greer, 1985), which are usually considered to be "lacertilians." Rage himself judged this character to be questionable because it has been checked only in a few taxa. The second presumed synapomorphy, "bord antérieur du scapulocoracoïde fenestré," is largely irrelevant to the question of "lacertilian" monophyly. The scapulocoracoid itself is absent in all snakes and is present in amphisbaenians only in Bipes, in which we are not convinced that fenestration (emargination) is actually lacking. Rage himself had doubts about this character as well; he indicated it with a question mark in his cladogram. Rage's third character, "pancreas rubanné," is also present in amphisbaenians. Rage suggested that it is convergent in the two groups, which is to some extent supported by the com- pact form of the pancreas in snakes, which Underwood (1967) refers to as the primitive condition (presumably for amniotes). Nevertheless, a ribbon-like pancreas could as well be a synapomorphy of squamates that has been further modified in snakes. Rage (1982) concluded that because of the questionable nature of these three presumed synapomorphies, a "definition" of "Lacertilia" is indeed fragile; he concluded that "Les Sauriens apparaissent surtout comme le regroupement des Squa- mates qui n'ont pas atteint le niveau évolutif de l'ensemble Amphisbénés-Serpents." We conclude that monophyly of "Lacertilia" is doubtful. If snakes and amphisbaenians are excluded from the "Lacertilia", we know of no synapomorphies that unite the members of this
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