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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 5 9 I G. wislizenii vestigium bicinctores reticulatus tt ebrius collaris dickersonae G. wislizenii C. vestigium C. bicincfores C. reticulatus C. nebrius C. collaris C. dickersonae Fig. 39.-(A) The single most parsimonious tree discovered in the reanalysis of the allozyme data set of Montanucci et al. (1 975) employing the approach in which frequency values are encoded into step matrices using Manhattan distances. (B) The 50 percent majority-rule consensus tree generated in the bootstrap analysis of this data set. stem J). Six additional unambiguous yet polymorphic apomorphies support this stem: 10.1 (supratemporal lies in a groove in ventral surface of supratemporal process of parietal, 23/24 of one step under ACCTRAN optimization, 2/24 of one step under DELTRAN optimization), 26.1 (bony shelf extends between medial process of surangular and ramus of mandible, 2/24 of one step under both ACCTRAN and DELTRAN optimization), 32.1 (palatine teeth present, 16/24 of one step ACCT- RAN, 13/24 of one step DELTRAN). 42.1 (scapular notch present, 10/24 of one step ACCTRAN, 5/24 of one step DELTRAN), 43.1 (posterior coracoid fenestrae present, fixed ACCTRAN, 16/24 of one step DELTRAN), and 45.1 (clavicular fenestrations present, fixed ACCTRAN, 8/24 of one step DEL- TRAN). Finally, Crotaphytidae may also be supported by two ambiguously placed synapomorphies: 40.1 (autotomic fracture planes of caudal vertebrae absent, 4/24 ofone step ACCTRAN) and 58.1 (postfemoral mite pockets present, fixed ACCTRAN). Stem B (Gambelia) is supported by 13 unambig- G. copei G. wislizenii G. corona$ G. silus C. reticulatus C. collaris C. nebrius C. antiquus C. dickersonae C. grismeri C. bicinctores C. vestigium Fig. 40.-The 50 percent majority-rule consensus tree generated in the bootstrap analysis of the complete (morphology + allo- zymes) data set. uous synapomorphies, six of which represent fixed character state changes: 12.1 (saddle-shaped suture between premaxilla and maxilla), 30.1 (slender, re- curved maxillary and dentary teeth), 44.1 (loss of the calcified cartilage border of the scapular fenes- tra), 46.1 (termini of the iliac blades round), 52.1 (femoral pore series extends beyond the inferior angle of the knee), and 62.1 (paired, paravenebrally arranged, blood-red spots present in juveniles of both sexes). This stem is also supported by seven unambiguous but polymorphic apomorphies: 2.1 (articulation between premaxilla and vomers in the form of a vertical ridge, 23/24 of one step ACCT- RAN, fixed DELTRAN), 15.1 (angle of jugal along anteroventral border of orbit approximately 90 degrees, 12/24 of one step under both ACCTRAN and DELTRAN), 17.1 (palatine foramen absent, 19/24 of one step ACCTRAN, 18/24 of one step DEL- TRAN), 24.1 (posterior mylohyoid foramen pos-
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 tenor to apex of coronoid, 20/24 of one step under both ACCTRAN and DELTRAN), 26.1 (bony shelf between the median process and ramus of the mandible, 22/24 of one step under both ACCTRAN and DELTRAN), 36.1 (second ceratobranchials widely separated, fixed ACCTRAN, 16/24 of one step DELTRAN), and 4 1.1 (one rather than two xiphisternal ribs, 20/24 of one step under both ACCT- RAN and DELTRAN). This stem also may be supported by five ambiguously placed transformations, the placements of which depend upon the optimization routine employed: 1.1 (nasal process of premaxilla narrow, 5/24 of one step ACCTRAN), 7.1 (tubercle present on anterolateral portion of post- orbital, 2/24 of one step ACCTRAN), 25.1 (poster- olingual process of coronoid angled posterolaterally at approximately 45 degrees, fixed ACCTRAN), 43.1 (posterior coracoid fenestrae absent, 8/24 of one step DELTRAN), and 58.1 (postfemoral mite pock- et present, fixed DELTRAN). Finally, this stem may be further supported by as many as 12 unpolarized characters: 8.0 (parietal roof not constricted poste- riorly), 9.0 (supratemporal processes tapered), 13.0 (maxilla-palatine articulation in the form of a low, rounded arch), 2 1.1 (posterior process of the ectop- terygoid crest absent), 23.0 (angular does not extend anteriorly beyond the fourth dentary tooth [counting forward] and rarely extends beyond the posterior- most tooth), 37.0 (skull nonrugose), 49.0 (supraor- bital semicircles absent), 50.0 (subocular scale series includes one very elongate scale), 55.1 (gular fold without closely approximating posteromedial folds), 56.0 (supra-auricular fold horizontal), 59.0 (hemi- penes large), and 82.1 (fully developed gular pattern in females). It is equally parsimonious for each of these characters to support stem E (Crotaphytus) depending upon their true polarity assignments. Stem C is supported by four unambiguous synapomorphies, two of which are fixed: 5.1 (loss of a vaulted cranium) and 20.0 (angle of the quadrate processes of the pterygoid approximately 18 degrees). The polymorphic apomorphies are characters 3 1.1 (number of premaxillary teeth, 0.35 of one step under both ACCTRAN and DELTRAN) and 32.1 (palatine teeth present, 6/24 of one step ACCT- RAN, 8/24 of one step DELTRAN). Nine more potential synapomorphies depend upon the partic- ular optimization routine employed. All but one of these (didovered during ACCTRAN optimization data runs) were coded as missing ("?') for G. coronat and thus may actually represent synapomorphies for stem D: 1 1.1 (septomaxillae slender and elongate, fixed), 15.1 (angle of jugal along anteroventral border of orbit approximately 90 degrees, 12/24 of one step), 24.1 (posterior mylohyoid foramen posterior to apex of coronoid, 2/24 of one step), 4 1.1 (one rather than two xiphisternal ribs, 3/24 of one step), 42.0 (scapular notch absent, 5/24 of one step), 53.1 (femoral pores of approximately equal size in males and females, fixed), 60.1 (females attain larger adult SVL than males, fixed), and 87.1 (territoriality absent, fixed). Finally, 25.2 (postero- lingual process of the coronoid angled posteroven- trally at approximately 45 degrees) may represent a fixed synapomorphy for this node (fixed DEL- TRAN). Stem D is supported by one unambiguous synapomorphy: 1.1 (nasal process of premaxilla long and slender, 19/24 of one step ACCTRAN, fixed DEL- TRAN). It may be further supported by as many as nine ambiguous (DELTRAN) characters including six of the characters (with the same frequency values) discussed under stem C (1 1, 1 5,24,4 1,53,60) for which G. coronat was coded as missing ("?'). The three remaining potential synapomorphies in- clude: 7.1 (tubercle on anterolateral border of post- orbital, 2/24 of one step), 10.1 (supratemporal lies in a groove along ventral border of supratemporal process, 21/24 of one step), and 40.1 (autotomic fracture planes absent, 4/24 of one step). Stem E (Crotaphytus) is supported by 24 unambiguous synapomorphies, 12 of which are fixed including: 19.1 (paraoccipital processes extend posterior to the occipital condyle), 27.1 (lateral process of the surangular present as a large protuberance), 33.1 (pterygoid tooth patch curls posterolaterally), 34.1 (ceratohyals with hook-like processes on prox- imal, medial edge), 35.1 (second ceratobranchials more than two-thirds the length of the ceratohyals), 54.1 (skin of the distal portion of tail weakly ad- herent to underlying musculature), 57.1 (presence of antehumeral mite pockets), 6 1.1 (subadult males acquire "gravid female" coloration), 66.1 (juvenile color pattern composed of white reticulations), 72.1 (acquisition of a black collar or collars outlined in white), 88.1 (lateral tail coiling behavior), and 98.3 (electromorph Tr). This stem is also supported by 12 unambiguously placed yet polymorphic transformations depending upon the optimization routine employed: 10.1 (supratemporal lies in a groove along ventral border of supratemporal process, 1/24 of one step ACCTRAN, 22/24 of one step DEL-
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