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- Australian blue tongue lizard ancestor was round-in-the-tooth
Thus, it seems likely this was the same specimen examined by Presch UUMZ 14 [ 21 ] prior to its desiccation, a conclusion previously confirmed by Avila-Pires [ 10 ]. No useful data could be obtained from this specimen. A Lacerta teguixin paralectotype NRM examined by us Fig 2 has five supraoculars, the first is the longest, the second is largest in area, and the fifth contacts two ciliaries. Three occipitals contact the interparietal scale and there are distinct, nearly round spots present on the dorsal surface of hind legs. The specimen has about rows of vertebrals. All traits suggest it is a member of our second clade.
However, the NRM lists four paralectotypes of Lacerta teguixin , 2 and scale counts made on other specimens by one of us AP suggests clade four may also be represented in this material. Given the above situation, we select NRM as described above and illustrated in Fig 2 as the neolectotype for Lacerta teguixin. Photo credit Sven O.
The name has long been considered a junior synonym of Tupinambis teguixin. The dorsal pattern is composed of wide dark bands separated by narrow light bands and four rows of white spots on the dorsum. It has five supraoculars, first is the longest, the second is the largest in area and the last one contacts two ciliaries.
Also, it has or vertebral rows. All are in agreement with our clade two. Here we retain this name as a junior synonym of Tupinambis teguixin based upon the data we have. Photo credit Aaron Bauer. Tupinambis nigropunctatus Spix was based upon five syntypes and all are extant. They note Spix was unsure of his own classification when it came to distinguishing it from T. Photos of ZMA a male illustrate morphology that agrees relatively well with our clade two Fig 4. Here we retain Tupinambis nigropunctatus Spix as a junior synonym of T.
Photo credit Michael Franzen. We gathered tissue samples from existing museum collections from 40 Tupinambis and Salvator , including 31 T. These data were combined with all available, vouchered individuals from GenBank for those genes for Crocodilurus , Dracaena , Tupinambis , and Salvator , representing the subfamily Tupinambinae, with Callopistes representing Callopistinae, following Harvey et al. We determined the optimal partitioning strategy for these loci using PartitionFinder [ 30 ] using the BIC criterion. We estimated phylogenies using MrBayes3.
We summarized the posterior distribution using a majority-rule consensus tree, with support estimated as the Posterior probability Pp for each node from the sampled trees. Specimen vouchers and GenBank accessions are given in [ S1 Table ]. The electronic edition of this article conforms to the requirements of the amended International Code of Zoological Nomenclature, and hence the new names contained herein are available under that Code from the electronic edition of this article. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN.
The LSID for this publication is: urn:lsid:zoobank. We reviewed the literature and examined illustrations and specimens said to be Tupinambis teguixin and T.
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We also consulted researchers with extensive taxonomic knowledge for their opinions on the status of some names. Taxonomic decisions are best made on the basis of recognizable morphological characters and concordant molecular evidence [ 33 ]. Thus, we reconcile geographic genetic variation with meristic and mensural characters from specimens to produce a robust taxonomic estimate with diagnostic evidence from both molecular and morphological data.
This integrates all available data, using the General Lineage Species Concept to delimit evolutionarily distinct clades as independent species [ 34 ]. For this work we examined extant museum specimens for morphological data [ S2 Table ]. Three previous works [ 8 , 10 , 18 ] provide detailed descriptions for Tupinambis.
However, some clarification as well as challenges regarding scale and scale arrangement terminology for Tupinambis are needed. While we use the terms and characters provided in these papers, we made some adjustments. Some scale counts and characters were found to contain information for distinguishing taxa, but most did not.
Some traditional characters were of limited use because the ranges overlapped extensively. These included vertebral row counts from the occiput to the row immediately posterior to the hind legs , transvers and longitudinal ventral scale rows counts, and lamellae on the fourth finger and fourth toe counted from the articulation points. Scales around mid-body were counted from one ventral around the mid-body, including scales of all sizes and shapes.
Characters that were more valuable for distinguishing taxa included the length and area of the supraoculars the longest vertebral axis and the largest area. Hoogmoed [ 19 ] described Tupinambis as having four supraoculars, and Avila-Pires [ 10 ] noted that a fifth scale is present that could be considered a supraocular; Harvey et al. Here we follow Avila-Pires [ 10 ] and consider the fifth and subsequent scales if present to be supraoculars, given their position above the orbit, and contact with ciliaries.
The number of occipital scales contacting the interparietal scale is relatively consistent within taxa; usually one or three occipital scales contact the interparietal, but occasionally the occipitals become fragmented or granulate in some specimens. Tupinambis has a gap between the pre-cloacal pores and the femoral pores. Pores are obvious in males while females tend to have pore-bearing scales with a small pore and a notch extending to the edge of the scale. Pore bearing scales were counted in both sexes. The number of enlarged supratemporal scales was somewhat useful, some taxa tend to have two enlarged supratemporals while others tend to have three.
The number of ciliaries in contact with the last supraocular was useful, as some taxa tend to have two ciliaries in contact with the last supraocular, while others tend to have three Fig 5. The shape and size of the largest scales on the anterior surface of the femur were of some use in distinguishing between taxa Fig 6A—6D.
The position of the anterior inside corner of the orbit defined as the posterior junction between the first subocular and the first ciliary over an upper labial was also useful. In some taxa it is over the third upper labial, in others it is over the fourth Fig 6E and 6F.
This is useful for identification since some taxa tend to have two ciliaries in contact with the last supraocular, while others tend to have three. The white markers denote the ciliaries in contact with the last supraocular. First, the shape and size of the scales on the anterior surface of the femur: A T. Second, the upper labial under the anterior corner of the orbit E, F. The inside corner of the orbit is over the third upper labial in Tupinambis teguixin , and the fourth upper labial in T.
The supratemporals are numbered. Tupinambis teguixin E usually has two supratemporals and T. Examination of photographs of type material including one of the paralectotypes of Lacerta teguixin Linnaeus NRM as well as Seps marmoratus Laurenti, and Tupinambis nigropunctatus Spix ZMA allowed for some comparison of the type material to the data collected from the specimens examined. Locality data was plotted using Arcview. Fig 7 illustrates localities sampled for DNA and morphology. Large circular markers denote the localities of specimens sampled for DNA. Smaller circular markers denote localities of specimens identified using morphology: Green is clade 1 T.
The two most northern red circles represent the islands of Tobago and Trinidad respectively.
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The other markers denote other species of Tupinambis not in the teguixin group. Measurements of body and tail lengths were taken to the nearest 1 mm using a ruler and tape measure. Dial calipers were used to measure scale lengths to the nearest 0. Univariate analyses of morphological data, Student t-tests, principal component analysis, and cluster analysis were applied when necessary. Because this project was started independently in the USA and Brazil not all data were collected for all specimens. The USA participants collected about 74 pieces of data on the specimens examined, the Brazilian contingent collected about 27 pieces of data on each specimen.
About 12 of these traits overlapped. Consequently, sample size in various analyses varies considerably. In this analysis, we used the following meristic counts: lower labials, upper labials, scales around midbody, vertebral rows, longitudinal ventral rows, transverse ventral rows, 4th finger lamellae and 4th toe lamellae.
We estimated prediction error based on replicates of fold cross-validation [ 39 ] of models with sequentially reduced number of predictors, ranked by importance. When building decision trees in random forests [ 40 ], regularization penalizes the selection of new features for splitting when the gain e. Several of the specimens used in the molecular analysis were also used in the morphological analysis for T.
The names established in this paper have been registered at ZooBank. Our results are similar to previous phylogenies of Tupinambinae [ 3 , 9 , 41 ]. The placement of the genera Dracaena and Crocodilurus is not strongly supported, likely due to the small amount of mitochondrial data available for those species.
We find weak support for a clade consisting of, respectively, Dracaena , Crocodilurus , and Tupinambis. We also find strong support for all sampled species, with possible paraphyly of S. Multi-locus nuclear datasets and deeper phylogeographic investigation will be needed to resolve deeper relationships in Tupinambinae and species limits in Salvator. Within Tupinambis , we find strong support for a clade of T. Interestingly, the T. Within the T. Some of these have been identified already by previous authors [ 3 , 42 ].
The first clade inhabits the Andean foothills and the western Amazon Basin. The second clade is widespread east of the Andes, in the Cerrado. The third clade appears restricted to the Maracaibo Basin in Venezuela and the fourth clade is primarily on the Guiana Shield and in the eastern Amazon basin. Each clade appears to correspond to a species-level taxon.
Note that we have not performed an explicit species-delimitation analysis, but these lineages have already been identified as distinct, putatively species-level taxa by previous authors, and are clearly diagnosable morphologically see below , while being relatively genetically and morphologically homogenous within each lineage. Their status as "cryptic" species is more a reflection of a lack of historical attention to their subtle morphological distinctiveness, resulting in a taxonomic burden of heritage [ 43 ]. Species illustrated: Top- Tupinambis cuzcoensis sp. Photo credit Mike Pingleton.
Second from top T. Photo credit Armida Madngisa. Bottom photo T. Photo credit JCM. With the exception of Tupinambis zuliensis , which was represented by only four individuals, these two variables permit a fairly good separation of the three other species Fig 9c. A Importance of meristic counts in predicting individual assignments to four species of Tupinambis lizards based on mean decrease in Gini accuracy as revealed by replicates of fold cross-validation of Guided Regularized Random Forests GRRF.
The higher the mean decrease in Gini accuracy, the higher the predictor importance. B Prediction error of GRRF models based on reducing number of predictors ranked by importance, as revealed by replicates of fold cross-validation. C Variation in vertebral rows and scales around midbody, the two best predictors of differences among four species of Tupinambis lizards. The results of the cluster analysis [ S2A Fig ] and PCA [ S2B Fig ] are in Based on the genetic and morphological analyses describe above, we split the species currently recognized as Tupinambis teguixin into four morphologically distinct species, three of which are new.
Considering the morphological data collected for this study, it is clear why these lizards have been confused for more than two centuries. Differences are subtle, the coloration and pattern are variable, complex and have an ontogenetic component. Table 1 summarizes the morphology for the four species of the Tupinambis teguixin group discussed here, and Table 2 compares the known species in the genus Tupinambis. Data for longilineus , palustris , quadrilineatus were taken from the literature and on-line photographs. Here, we provide a taxonomic revision to bring taxonomy into concordance with the molecular and morphological results for the Tupinambis teguixin complex.
First, we provide a re-description of:. Figs 1a and 1c , 3a and 7 ,. In the molecular analysis this is clade 2. The largest Tupinambis teguixin measured was a male, mm SVL with a mm tail. The smallest was a neonate 84 mm SVL and a mm tail. Supraoculars five, six or seven are not common , the first is the longest; last supraocular usually in contact with two ciliaries; all specimens had three occipitals contacting the interparietal, except one specimen which had two; and all had an incomplete interangular fold; suboculars usually six, one specimen had seven; upper labials 8—10, third or fourth the longest; lower labials 7—8; sublabials 3—5, usually four; chin shields in four pairs, rarely five pairs; lamellae on fourth finger 14—16; lamella on fourth toe 31— Tupinambis teguixin is distinguished from the sympatric Tupinambis cryptus sp.
Tupinambis teguixin differs from T. Tupinambis teguixin differs from Tupinambis zuliensis sp. Natural History. Because this species has been long confused with other species of the T. Considering that Tupinambis teguixin and T. This species corresponds to clade 4 in the molecular analysis. Holotype AMNH , male. Size SVL mm, tail broken. Collected 5 March by Charles J. Cole and Carol R. Townsend at the Dubulay Ranch on the Berbice River, ft asl, 5. Description of holotype. Color in alcohol.
Crown has mottled plates which are mostly dark with light areas; face is brown-black with light spots on each scale; chin is olive green; throat is mostly olive green with some yellow; neck red brown with gray vermiculated pattern; trunk is a mosaic of light and dark indistinct bands; dorsal surface of legs vermiculated with red brown and yellow-gray; ventral surface is yellow with some black intruding laterally, and black along the seams of the ventral plates; tail is uniform red brown above and laterally, yellow ventrally, distally striped with wide black and slightly narrower yellow bands.
The cloacal plate has seven or eight rows of scales from the level of the femoral pores to the free edge of the plate, five of these rows are plate-like scales. In alcohol the head is olive green with some dark spotting on the crown scales. The lower jaw is yellow and heavily mottled with black pigment. The dorsum of the neck and body is mostly uniform dark brown to black with about 12 rows of indistinct spots. The ventral surface is mostly yellow with dark pigment intruding from the side along the seams of the ventrals, and some patches of dark pigment scattered.
This same coloration occurs on the underside of the legs. Proximally, the tail is a solid dark brown-black above with very narrow yellow rings. Coloration of juveniles and adults in life can be seen in Fig 1.
There is considerable pattern variation in Tupinambis cryptus. None of the adults from Trinidad and Tobago have this pattern, although we have seen a few individuals in the field with indistinct bands.
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The largest Tupinambis cryptus sp. The smallest was a neonate that was 85 mm SVL and a 42 mm tail. Smallest individual measured 85 mm SVL. Tupinambis cryptus sp. It can be distinguished from T. Tupinambis cryptus is known from Trinidad and Tobago, Venezuela, the Guianas, and as far south as the confluence of the Rio Negro and Rio Branco in Brazil, but its range may be more extensive. It ranges as far west as Falcon, Venezuela, and appears to range into the Andes in the vicinity of Bucaramango, Colombia.
The co-occurrence of this species with T. However, T. Our observations of this lizard suggest they use secondary forest, savannas, and human modified habitats. We have not observed them in primary forests proper, but at the forest edge. It may avoid dense forest because of the reduced number of basking sites.
Australian blue tongue lizard ancestor was round-in-the-tooth
Like other species of Tupinambis , T. We have observed this lizard investigating caiman nests, foraging along streams on the floor of secondary forests and in mangroves. Usually their tongue is flicking and they are probing the leaf litter with their head. Tupinambis cryptus is most readily observed foraging under the bird feeders at the Asa Wright Nature Center Trinidad were they scavenge pieces of fruit.
The ecology of the Trinidad population was examined by Everard and Boos [ 44 ]. They trapped Tupinambis cryptus sp n. Traps were baited with chicken remains. At the Waller Field study site 56 T.
Flinders University. Australian blue tongue lizard ancestor was round-in-the-tooth. Retrieved July 1, from www. But new research reveals a major problem with this Their study shows not only that the animal is a close relative of the ancestor Ancient fossilized scrapes recently Below are relevant articles that may interest you. ScienceDaily shares links with scholarly publications in the TrendMD network and earns revenue from third-party advertisers, where indicated.
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