Arnold, E.N. (1989) - Towards a phylogeny and biogeography of the Lacertidae: relationships within an Old-World family of lizards derived from morphology. - Bulletin of the British Museum (Natural History), Zoology, 55, 209-257. × Relationships of lacertid lizards were assessed on the basis of 84 primary and 112 binary characters drawn mainly from morphology, including features of the skeleton, external anatomy, various internal soft part systems and two aspects of behaviour. Among features not previously used, or not fully investigated before, are structure of the septomaxilla and nasal passages, arranged of the xiphisternal cartilages, mite pockets, kidney position, ulnar nerve arragement, thoracic fascia, aspects of the hemipenis and its associated muscles, female genitalia and jaw muscles. On the basis of parsimony analysis and compatibilty treatment of this character set, the Lacertidae fall into two main portions: A paraphyletic Palaearctic and Oriental group of primitive forms, from which is derived a holophyletic assemblage of Ethiopian and advanced Saharan and Eurasian taxa.
The former group ist not fully resolvable, but Psammodromus and Gallotia appear to be sister groups and are probably related to Lacerta parva and L. fraasi and then L. brandtii, Podarcis appears to be related successively to L. andreanszkyi, the sister species L. dugesii and L. perspicillata, and perhaps L. danfordi and L. laevis. This assemblage may be related to archaeolacertas and Algyroides. The separation of Lacerta lepida, L. pater and L. princeps from the agilis group, based on chemical evidence, is weakly contradicted by morphology. Takydromus may be most closely related to L. vivipara, and L. jayakari and L. cyanura constitute the most likely sister group of the Ethiopian and advanced Saharo-Eurasian assemblage.
Taxe in the Ethiopian and advanced SaharoEuroasian assemblage form a long essentially pectinate tree with relatively change between the side branches, except for a strong disjunction separating the more primitive from the more advanced taxa. Most of the former fall on two main branches, with ´Lacerta` australis and ´L.` rupicola possibly basal to them. 1. the Equatorial forest group containing Gastropholis, Bedriagaia, ´Lacerta` echinata, Adolfus, ´Lacerta` jacksoni and Holaspis. The first three of these constitute a holophyletic group and the same is probably true of the remainder. 2. Tropidosaura, Poromera and Nucras, the latter being the sister group of the more advanced forms. These include successively the Ethiopian Philochortus, Latastia, Ichnotropis and Heliobolus, Pseuderemias, Meroles and Aporosaura, and Pedioplanis, and then the Saharo-Eurasian Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita.
It seems probable that the ancestors of modern Lacertidae arose in western Eurasia, where the family is known since the Palaeocene and is still represented there largely by quite primitive forms (89 species and seven nominal genera). The family later invaded Africa, perhaps first in the early or middle Miocene. Relatively primitive lacertids spread widely in largely mesic situations in the Ethiopian region, radiating to some extent (six present genera and 16 species) and producing Nucras and the related series of advaned groups (eight genera and 54 species) whoich show increasing adaptation to xeric environments. These genera tend to have heir most primitive species in the northeast and north of the Ethiopian region. The most advaned gave rise to the Saharo-Eurasian clade, now made up to Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita. This invaded the arid areas of North Africa and Eurasia, where it is presently represented by 70 species. Many morphological changes in increasingly advanced lacertids may be functionally related to the problems of survival in arid, hot, open environments. Considerable ecological parallelism exists in lacertids, with members of separate stocks occupying similar niches in different geographical areas. Morphological adaptations associated with these niches contribute significantly to the high levels of character homoplasy found in the family. There is also some correlation between the degree of niche differentiation in various groups and the quality of the phylogenies that can be produced from their physical characters. A number of morphological parallels exist between advaned lacertids and New World macroteiids. In the skull at least, advaned lacertids show a complex mixture of paedomorphosis and acceleration.
Nomenclatorial changes are as follows: Cabrita is synonymised with Ophisops, necessitating a new name, Ophisops nictans, for Cabrita jerdonii. Aporosaura is synonymised with Meroles, Platyplacopus with Takydromus, and Bedriagaia with Gastropholis. ´Lacerta` (or Centromastyx) echinata is also transferred to the latter genus and Lacerta jacksoni to Adolfus. ´Lacerta` australis and ´L.` rupicola are put in a new genus, Australolacerta. It is recommended that Lacerta dugesii and L. perspicillata should not be placed in the otherwise very uniform genus Podarcis. Although clearly paraphyletic, Lacerta s. lat. Should be retained at least for the present and, if necessary putative relationships within it indicated by informal groups or subgenera. Edwards, S. & Branch, W.R. & Vanhooydonck, B. & Herrel, A. & Measey, G.J. & Tolley, K.A. (2013) - Taxonomic adjustments in the systematics of the southern African lacertid lizards (Sauria: Lacertidae) - Zootaxa, 3669 (2): 101–114. × Molecular phylogenetic analyses of southern African lacertid lizards (Eremiadini) using mitochondrial and nuclear
markers revealed two examples of generic assignments incompatible with monophyletic clades. Australolacerta
Arnold 1989, a genus endemic to South Africa and to which two isolated species have been referred, is
paraphyletic at the generic level. In addition, the species Ichnotropis squamulosa Peters 1854 was found to be
embedded within the genus Meroles. To resolve the paraphyly in Australolacerta we erect a new genus,
Vhembelacerta Edwards, Branch, Herrel, Vanhooydonck, Measey, & Tolley, gen. nov., to accommodate Lacerta
rupicola FitzSimons 1933. To maintain a monophyletic Ichnotropis Peters 1854, Ichnotropis squamulosa Peters
1854 is transferred to Meroles Gray 1838, now named Meroles squamulosus comb. nov. Where necessary the
genera affected by these actions are re-characterized. Engleder, A. & Haring, E. & Kirchhof, S. & Mayer, W. (2013) - Multiple nuclear and mitochondrial DNA sequences provide new insights into the phylogeny of South African Lacertids (Lacertidae, Eremiadinae). - Journal of Zoological Systematics and Evolutionary Research, Berlin, 51 (2): 132-143. × Eremiadinae, one of three subfamilies of Lacertidae, are distributed throughout Asia and Africa. Previous phylogenetic studies suggested that one of
the main groups of Eremiadinae (the Ethiopian clade) consist of two clades with predominately East-African and South-African distribution. Yet, especially
the latter one, which includes the genera Pedioplanis, Meroles, Ichnotropis, Tropidosaura and Australolacerta, was not well supported in the
molecular phylogenetic analysis. In this study, we analysed the phylogenetic relationships among the genera of the ‘South African clade’ to assess
whether this group actually forms a highly supported clade and to address questions concerning the monophyly of the genera. We sequenced sections
of the widely used mitochondrial genes coding for 16S rRNA, 12S rRNA and cytochrome b (altogether 2045 bp) as well as the nuclear genes c-mos,
RAG-1, PRLR, KIF24, EXPH5 and RAG-2 (altogether 4473 bp). The combined data set increased the support values for several nodes considerably.
Yet, the relationships among five major lineages within the ‘South African clade’ are not clearly resolved even with this large data set. We interpret
this as a ‘hard polytomy’ due to fast radiation within the South African lacertids. The combined tree based on nine marker genes provides strong support
for the ‘South African Clade’ and its sister group relationship with the ‘East African Clade’. Our results confirm the genus Tropidosaura as a
monophylum, while Ichnotropis is paraphyletic in our trees: Ichnotropis squamulosa appears more closely related to Meroles than to Ichnotropis capensis.
Furthermore, the monophyly of Meroles is questionable as well. Based on our results, I. squamulosa should be transferred from Ichnotropis into
the genus Meroles. Also, the two species of Australolacerta (A. australis and A. rupicola) are very distantly related and the genus is perhaps paraphyletic,
too. Finally we propose a phylogeographical scenario in the context of palaeoclimatic data and compare it with a previously postulated hypothesis.
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