Callopistinae, Harvey & Ugueto & Gutberlet, 2012

Harvey, Michael B., Ugueto, Gabriel N. & Gutberlet, Ronald L., 2012, 3459, Zootaxa 3459, pp. 1-156 : 77

Callopistinae New Subfamily

Type Genus.— Callopistes by original designation.

Diagnosis.— Characters in the generic diagnosis of Callopistes distinguish the Callopistinae from the Teiinae and Tupinambinae . Unlike the Chamopsiinae , the Callopistinae has strong heterodonty and flat (non-concave) frontal and parietal bones. The Callopistinae lacks a parietal foramen and surangular fenestra ( Sullivan & Estes 1997).

Remarks.— In an unpublished doctoral dissertation, Teixeira (2003) assigned Callopistes to the Teiinae based on phylogenetic analysis of morphological data. Nonetheless as noted by Teixeira (2003) and Giugliano et al. (2007) this conclusion was only weakly supported. Teixeira’s data matrix and characters have several problems that raise questions regarding her results. For example, she incorporated data directly from several earlier studies ( Moro & Abdala 2000; Presch 1970; Vanzolini & Valencia 1965; Veronese & Krause 1997), in some cases perpetuating errors in those studies. She miscoded characters 43 and 44 (intertympanic and interangular sulci both coded as present in Tupinambis and absent in Crocodilurus based on Vanzolini & Valencia’s misunderstanding of these characters. Interangular sulci are absent in both genera, whereas intertympanic sulci are present in Tupinambis and Crocodilurus , but absent from Salvator ). As we point out elsewhere in this publication, some of these data matrices may include misidentified species and Teixeira apparently did not address this problem. Many of her new characters are not described except in brief one-line statements, and this problem is particularly acute in the hemipenial descriptions where variation appears to be over interpreted. Teixeira (2003) listed presence of teeth on the pterygoid (her character 6.1) as a synapomorphy of Teiinae + Callopistes , and she coded the various Teiinae genera as being fixed for this character (i.e., she assigned them state 1 rather than 0/1). This coding contradicts Presch’s (1974a, p. 347) observations that, “pterygoid teeth are present in K. intermedius but variably present in K. calcaratus and absent in all other species examined,” and “of the species of Ameiva that I have examined, only Ameiva bifrontata and Ameiva ameiva have pterygoid teeth.”

Our study did not consider most synapomorphies that Teixeira (2003) identified for Teiinae + Callopistes . Regarding 38.1 (supraoculars not in contact with supraciliaries), we suspect that this character is a synapomorphy of Teiidae rather than Teiinae + Callopistes ; a reversal is likely a shared character of the Tupinambis teguixin group and Crocodilurus . Scales between the supraoculars and supraciliaries rarely occur in gymnophthalmids ( Amapasaurus has them, for example). Teixeira (2003) examined specimens of Salvator merianae and S. duseni in her analysis. Both species invariably have her character 38.1, yet she coded Tupinambis as having 38.0. She coded Dracaena as polymorphic (0/1); all specimens we examined have 38.1.

Teixeira’s (2003) study remains unpublished and some of the problems might be resolved during any future review process. However, we think it inadvisable to uncritically accept her conclusions or incorporate her data into any type of combined analysis. For this reason, we do not have much confidence in the combined analysis of Giugliano et al. (2007).

Phylogenetic analysis of mitochondrial DNA supports assignment of Callopistes to a separate subfamily. Callopistes differs considerably from other Tupinambinae and is likely the sister genus of the remaining Teiidae or Tupinambinae . This conclusion is consistent with our survey of coloration, hemipenes, and external morphology. Furthermore, it is well supported by earlier morphological ( Presch 1974a; Sullivan & Estes 1997) studies. The parsimony analysis of 12S sequences by Giugliano et al. (2007) recovered Callopistes as the sister genus of all other Teiidae , whereas analysis of 16S and combined 12S and 16S sequences recovered Callopistes as the sister genus of the Tupinambinae . Finally, we note that Callopistes is an ancient lineage. Giugliano et al. (2007) concluded that Callopistes diverged from other teiids in the Paleocene, perhaps through vicariance related to formation of the Salamancan Sea.

In summary, a separate subfamily should be recognized for four reasons: (1) Callopistes is basal to one or both other extant subfamilies, (2) its exact relationship to the other subfamilies has yet to be resolved, (3) it likely represents a very old lineage distinguished from other teiids by a suite of novel characters, and (4) recognition of the subfamily does not render either the Teiinae or Tupinambinae paraphyletic, but failure to do so would produce a taxonomy inconsistent with teiid phylogeny.