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1996 McGUIRE-SYSTEMATICS OF CROTAPHYTID LIZARDS 6 1 TRAN), 22.1 (posterior projections of the parabasisphenoid terminate at, or anterior to, the base of the sphenoccipital tubercles, fixed ACCTRAN, 23/ 24 of one step DELTRAN), 3 1.1 (number of premaxillary teeth, 0.38 of one step under both ACCT- RAN and DELTRAN), 40.1 (loss ofautotomic frac- ture planes of the caudal vertebrae, 20/24 of one step ACCTRAN, fixed DELTRAN), 42.1 (suprascapular notch present, 12/24 of one step ACCT- RAN, 13/24 of one step DELTRAN), 47.1 (medial and lateral plantar tubercles contact to form an arch, 21/24 of one step ACCTRAN, 19/24 of one step DELTRAN), 76.1 (dark nuchal spots present, 19/ 24 of one step under both ACCTRAN and DEL- TRAN), 90.6 (aGPD, 0.08 of one step under both ACCTRAN and DELTRAN), 9 1.2 or 9 1.6 (6-PGD, 0.12 of one step ACCTRAN, 0.05 of one step DEL- TRAN), 94.2 (GOTs, 0.69 of one step under both ACCTRAN and DELTRAN), 95.2 or 95.7 (Pro. 0.58 of one step ACCTRAN, 0.56 of one step DEL- TRAN), and 96.6 (Est I, 0.7 1 of one step under both ACCTRAN and DELTRAN). Four additional po- tential transformations at this node are discovered only under ACCTRAN optimization including: 28.1 (moderately developed ridge present on lateral surface of the surangular, fixed), 68.1 (white component of adult dorsal pattern composed of reticulations, fixed), 85.1 (dorsal coloration golden tan, fixed), and 89.2 (H-LDH, fixed). Finally, this stem may be supported by as many as 12 transformations that could not be polarized. It is equally parsimonious for each of these characters to support stem B (Gambelia) and a complete listing is given under the discussion of stem B. Stem F is supported by nine unambiguously placed transformations, four of which are fixed: 68.2 (white portion of dorsal pattern in the form of spots), 69.1 (sexual dichromatism of the dorsal color pattern), 85.4 (brown dorsal body coloration), 86.1 (saxicoly). The five unambiguous yet polymorphic apomor- phies include: 24.0 (posterior mylohyoid foramen equal with apex of coronoid, 1/24 of one step under both ACCTRAN and DELTRAN), 3 1.0 (number of premaxillary teeth, 0.01 of one step under both ACCTRAN and DELTRAN), 5 1.1 (terminal supradigital scales elevated from dorsal surface of claw, 4/24 of one step under both ACCTRAN and DEL- TRAN), 70.1 (paired melanic keels on ventral caudal extremity, 7/24 of one step under both ACCT- RAN and DELTRAN), and 77.1 (inguinal patches present, 7/24 of one step under both ACCTRAN and DELTRAN). This stem may also be supported by the following five ambiguously placed transformations, depending upon the optimization routine employed: 28.1 (ridge on lateral surface of the surangular, fixed DELTRAN), 43.1 (posterior coracoid fenestrae present, 8/24 of one step DELTRAN), 45.0 (clavicular fenestrations lost, 16/24 of one step ACCTRAN), 58.1 (postfemoral mite pockets present, fixed DELTRAN), and 89.2 (H-LDH, fixed DELTRAN). Stem G is weakly supported by eight unambiguously placed transformations, none of which are fixed. The unambiguous, yet polymorphic apomor- phies include 17.0 (palatine foramen present, 4/24 of one step under both ACCTRAN and DEL- TRAN), 26.0 (no bony shelf present between medial process of the surangular and ramus of mandible, 1/24 of one step under both ACCTRAN and DEL- TRAN), 32.1 (palatine teeth present, 2/24 of one step ACCTRAN, 5/24 ofone step DELTRAN), 42.1 (suprascapular notch present, 2/24 of one step ACCTRAN, 5/24 of one step DELTRAN), 47.1 (arch formed by contact of medial and lateral plantar tubercles, 3/24 of one step under both ACCTRAN and DELTRAN), 5 1.1 (terminal supradigital scales elevated from dorsal surface of claws, 20/24 of one step under both ACCTRAN and DELTRAN), 70.1 (paired melanic keels present on ventral caudal extremity, 17/24 of one step under both ACCTRAN and DELTRAN), and 77.1 (inguinal patches present, 17/24 of one step under both ACCTRAN and DELTRAN). Four ambiguously placed transformations may also support this node: 22.1 (posterior projections of parabasisphenoid terminate at, or anterior to, the base of the sphenoccipital tubercles, 1/24 of one step DELTRAN), 24.0 (posterior mylohyoid foramen equal with apex of coronoid, 1/24 of one step ACCTRAN), 90.1 (aGPD, 0.08 of one step ACCTRAN), and 95.5 (Pro, 0.43 of one step ACCTRAN). Stem H is weakly supported by one unambiguously placed polymorphic transformation: 76.0 (absence of dark nuchal spots, 13/24 of one step under both ACCTRAN and DELTRAN). This node may also be supported by two ambiguously placed transformations: 2.0 (ventral suture between vomers and premaxilla does not form a vertical ridge, 1/24 of one step ACCTRAN) and 90.1 (aGPD, 0.08 of one step, DELTRAN). Stem I is supported by five unambiguously placed transformations, three of which are fixed: 39.1 (lat-
62 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 32 erally compressed tail with dorsal and ventral fat bodies), 65.1 (off-white dorsal caudal stripe present), and 78.1 (enlarged inguinal patches in adult males). The two unambiguous yet polymorphic transformations are 3 1.1 (number ofpremaxillary teeth, 0.0 1 ofone step under both ACCTRAN and DELTRAN) and 76.0 (dark nuchal spots lost, 6/24 of one step under both ACCTRAN and DELTRAN). Three ambiguously placed transformations may also support this node: 24.0 (posterior mylohyoid foramen equal with the apex of coronoid, 1/24 of one step ACCTRAN), 47.1 (arch formed by contact of the medial and lateral plantar tubercles, 2/24 ofone step ACCTRAN), and 75.1 (ventral extent of posterior collar marking just reaches forelimb insertion, fixed ACCTRAN). Stem J is supported by six unambiguously placed transformations, two of which represent fixed changes: 7 1.0 (loss of black oral melanin) and 75.2 (posterior collar terminates within antehumeral fold). The unambiguous yet polymorphic transformations include: I. 1 (nasal process of premaxilla narrow, 1/24 of one step under both ACCTRAN and DEL- TRAN), 32.0 (palatine teeth lost, 8/24 of one step under both ACCTRAN and DELTRAN), 44.1 (cal- cified cartilage anterior border of scapular fenestra absent, 3/24 of one step under both ACCTRAN and DELTRAN), and 95.4 or 95.5 (Pro, 0.1 1 of one step under ACCTRAN, 0.41 under DELTRAN). This stem may be further supported by two ambiguously placed transformations, both of which were discovered under ACCTRAN optimization: 89.3 (H-LDH, fixed) and 94.3 (GOTs, fixed). Stem K is weakly supported by two unambiguously placed transformations, neither of which is fixed: 45.1 (clavicular fenestrations present, 4/24 of one step under both ACCTRAN and DELTRAN) and 83.1 (enlarged melanic axillary patches present, 18/24 of one step ACCTRAN, 15/24 of one step DELTRAN). The stem may be further supported by four ambiguously placed transformations: 70.0 (paired melanic keels absent from ventral caudal extremity, 13/24 of one step ACCTRAN), 89.3 (H-LDH, tixed DELTRAN), 94.3 (GOTs, fixed DELTRAN), and 95.4 (Pro, 0.11 of one step DEL- TRAN). Stem L is supported by seven unambiguously placed synapomorphies, three of which are fixed: 73.1 (posterior collars widely separated), 75.3 (posterior collar terminates ventrally before entering the antehumeral fold), and 84.1 (olive green ventrolat- era1 coloration present). Olive green ventrolateral coloration is not a fixed state in C. vestigium as northern populations are characterized by burnt- orange ventrolateral coloration. This was an artifact of the multistate character coding scheme employed in this analysis and resulted at least in pan because a satisfying estimate of the frequencies of the orange and green ventrolateral conditions in C. vestigium could not be obtained from preserved material. The four unambiguous but polymorphic transformations include: 16.1 (acquisition of extravomerine bones, 9/24 of one step under both ACCTRAN and DELTRAN), 26.1 (bony shelf between medial process of surangular and ramus of the mandible present, 5/24 of one step under both ACCTRAN and DELTRAN), 3 1.1 (number of premaxillary teeth, 0.14 of one step under both ACCTRAN and DEL- TRAN), and 45.0 (ciavicular fenestrations lost, 2/24 of one step under both ACCTRAN and DEL- TRAN). Finally, four ambiguously placed transformations may also support this node: 68.3 (presence of slender, transversely arranged, white dorsal stripes, fixed ACCTRAN), 9 1.1 (6-PGD, 0.1 2 of one step ACCTRAN), 95.3 (Pro, 0.29 of one step ACCT- RAN), and 96.1 (Est l,0.7 1 ofone step ACCTRAN). In an attempt to assess the amount of character support for each clade, bootstrap and decay index analyses were performed for the combined data set. From each of these analyses it is clear that a number of clades are rather unstable. For example, in the bootstrap analysis (Fig. 40), nodes G and H were supported in less than 50 percent of the bootstrap replications. Stems D and F also were found to be relatively weakly supported with bootstrap propor- tion values of 66 and 60, respectively. Despite its low bootstrap P value, stem F is supported by four fixed, unambiguous synapomorphies. The amount of support for stem D may be underestimated because a number of derived character states were coded as missing ("?') for the fossil taxon G. co- ronat, thus rendering these transformations ambig- uous when both ACCTRAN and DELTRAN optimization routines are considered. Notably, when G. corona* is excluded from the analysis, the bootstrap P value of stem D is 100. The remaining clades appear to be well supported, given that Hillis and Bull (1 993) found bootstrap proportions to be con- servative estimates of stem support with P values 2 70 corresponding to a probability r 0.95 that the corresponding clade is real. The results of the decay index analysis (Fig. 41)
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Cbrr- Branch acler Steps 31 38 33 2
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