Phylogeny and generic monophyly in the Testudinidae

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GeoTerraTestudo

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Fritz, U. and O.R.P. Bininda-Emonds. 2006. When genes meet nomenclature: Tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone. Zoology 110: 298–307.

Abstract
We used a five-gene data set (mtDNA: 12S rRNA, 16S rRNA, cyt-b; nDNA: Cmos, Rag2) comprising approximately two-thirds of all extant testudinid species and, for the first time, including all five Testudo species to investigate the question of whether all western Palaearctic testudinids are monophyletic. Further, we examined whether the recently suggested allocation of the African Geochelone pardalis in the otherwise exclusively South African genus Psammobates and of the Malagasy G. yniphora in the monotypic genus Angonoka is justified in the face of considerable morphological evidence against such placements. Our phylogenetic analyses do not support the paraphyly and generic break-up of Testudo, as suggested by previous papers using a smaller taxon sampling and mtDNA data only. We propose a continued usage of the generic name Testudo for all five western Palaearctic tortoise species. Within Testudo, two monophyletic subclades are present, one containing T. hermanni+T. horsfieldii, and the other comprising (T. kleinmanni+T. marginata)+T. graeca. Nomenclaturally, we demonstrate that Eurotestudo Lapparent de Broin et al., 2006, which was recently erected with the type species T. hermanni, is an objective junior synonym of Chersine Merrem, 1820 and Medaestia Wussow, 1916. Recognition of a monotypic genus Angonoka for G. yniphora is unwarranted according to both our re-analysis of sequence data and morphological data. Acknowledging the strong morphological similarity between G. yniphora and G. radiata, we suggest placing both species into the genus Astrochelys. Although sequence data for only one of the three Psammobates species was available for analysis, there is currently no cause to challenge the monophyly of this genus as established on the basis of morphological evidence. Thus, we hypothesize that G. pardalis is sister to a monophyletic Psammobates. In light of the clear morphological gap between G. pardalis and Psammobates species, the recognition of a distinct genus Stigmochelys for the former seems justified.

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bikerchicspain

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So let me get this right, The Monophyletic are considered as the only natural kind of group, so the group of Testudo are pure?

And that the clades are made up of 2 or more different species.?
 

HLogic

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It is interesting how quickly things change. Geochelone pardalis ssp. is now Stigmochelys pardalis (until such time as someone argues the distinction noted in the cited paper as sufficiently legitimate to resurrect the subspecific nomenclature).

An argument against combining genera based upon morphological differences is presented with respect to G. pardalis being placed in Psammobates but when presented with similar genetically based evidence regarding T. marginata, T. kleinmanni & T. graeca vs. T. horsfieldii and T. hermanni the author suggests retaining the monophyletic generic nomenclature of Testudo because of historical use, confusion and "practical" reasons. Those arguments didn't seem to apply in the Elaphe (European/N. American Ratsnakes) split into who-knows-how-many genera once the dust settles...

The utilization of a single specimen (i.e. Psammobates tentorius in this study and G. agassizii & I. forsteni in Le, 2006) as a basis for any argument sticks in my craw, too. Sample size counts!

Publish or die... I guess that is what it takes to get the grants but it seldom does anything to clarify the phylogeny of the specimens in question.

My guess is Stigmochelys p. pardalis & S. p. babcocki will be recognized in 50,000 years as separate subspecies once the subspeciation has had enough time to make them different enough for us to 'realize' it. In the same 50K years, Testudo will be split and combined at least half as many times as the number of years transpired. Last but not least, C. carbonaria (yes, I used Chelonoidis) will be split into at least two subspecies as soon as someone includes the phrase "...and the Effects of Global Warming on Population Stability" in their grant request.
 

GeoTerraTestudo

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Yes, "monophyletic" means a natural grouping, while "paraphyletic" means an artificial grouping (based on superficial similarities, rather than evolutionary relationship). There has been a lot of debate as to the validity of Testudo (see references in the paper), but according to this study, the genus reflects a real relationship among the five Testudo species (the Greek, Egyptian, marginated, Hermann, and Russian). Furthermore, there are two lineages within the genus: one is the Greek-Egyptian-marginated lineage, and the other is the Hermann-Russian lineage.

Another thing I really liked about this paper was that it showed how closely the African pancake tortoise (Malacochersus tornieri) is related to the Testudo tortoises, forming a clade, which is then in turn related to the Asian Indotestudo species (I. elongata, I. forstenii, and I. travancorica). I always thought the pancake tortoise and Indotestudo tortoises looked a lot like the Mediterranean tortoises, and this confirms that they are indeed closely related to each other.

Finally, the paper also showed that the African leopard tortoise is most closely related to the small Psammobates tortoises of southern Africa (P. geometricus, P. oculifer, and P. tentorius). This clade is in turn related to the other small African tortoises (Homopus, Chersina, and Kinyxis). Meanwhile, the closest relatives of the Sahel's big sulcata tortoise are Indian, not African (Geochelone elegans and G. platynota).

Overall, I think this is a really neat and useful tortoise study.
 

bikerchicspain

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Interesting, slightly confusing, had to read it 3 times before I got the hang of it,
A bit too scientific for me to take it, although when read in the context that it's meant to be it's actually straight forward,
You get to learn something new everyday on here.lol
Where do you find these things:D:D
 

HLogic

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Actually, paraphyletic is a legitimate phylogenetic relationship meaning a partial phlogeny (lineage). Monophyletic is the term used to denote all descendants of a common ancestor. Paraphyletic is used to denote a subset, sharing common ancestry, of a monophyletic lineage.

As a [very] loose analogy, siblings comprise a monophyletic group - all descended from a common 'ancestor', their parents. The female (or male) siblings considered as a group are paraphyletic - part of the complete monophyletic group. As an example from the diagrams in the article, Manouria and Gopherus taken together as a group are a paraphyletic clade of the family Testudinidae (the entirety of the chart). All of the other tortoises, as a group, are also a paraphyletic clade. The chart as a whole describes the monophyletic clade, Testudinidae (true tortoises), of the super family Testudinoidea (tortoise-oid, -like, -ish turtles). Another example of a paraphyletic group would be the group of all reptiles excluding birds (or snakes or lizards or...).

Not to add to the confusion, but there is another grouping known as polyphyletic - having multiple ancestors (e.g. warm-blooded animals comprised of mammals and birds).
 

GeoTerraTestudo

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In phylogenetics, paraphyly is to be avoided because it includes certain taxa within a clade, but selectively excludes others. Monophyly is preferred, because it shows real evolutionary relationships. Here is a nice illustration of that concept:

http://www.nicerweb.com/bio1152/Locked/media/ch26/clade-paraphyletic.html

In the context of this paper, the genus Testudo is monophyletic because those 5 species really are, by all indications, closely related. And within the genus, T. hermanni and T. horsfieldii form one clade, while T. graeca, T. kleinmanni, and T. marginata form a second clade. So what this means is, if you're going to stick the Russian tortoise in a separate genus (or subgenus) "Agrionemys," then you need to do the same with the Hermann tortoise. But because these 5 species are so closely related, there is no real justification for separating them at the generic level, although distinction at the subgeneric level may be warranted.
 

bikerchicspain

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this is getting way toooo complicated for some of us simpletons... :rolleyes:

My view is i have a tortoise, i know what species it is and i know how to look after properly, the rest i leave to vets and scientists. :D:D:D
 
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