S.s.gallaica or S.s.bejarae?

Azhael

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Ok, i feel really dumb for having to ask this question but i´m genuinely confused. The S.salamandra populations of the central-north part of Spain, below the Cantabrian mountains have typically been called S.s.bejarae. I have seen them refered to as such in plenty of places and have always assumed this was the case. However, according to the distribution maps i´ve been able to find, S.s.bejarae doesn´t cross the northern plateau, and the distribution maps of S.s.gallaica sometimes include a narrow strip right below the Cantabrian mountains, extending under the bernardezi/fastuosa populations further north. The animals in this narrow strip show varying degrees of red pigmentation.
So it sounds like those animals are S.s.gallacia and not S.s.bejarae, but i haven´t been able to confirm this with any literature. The problem is, the head shape and the general coloration don´t seem to fit gallaica, but they don´t fit bejarae either, and this is further confused by the hybridization with fastuosa and possibly bernardezi.
If someone knows of a paper that addresses those populations, i´d be very grateful.
 

FrogEyes

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http://ib.berkeley.edu/labs/wake/2003_Evolution_Salamandra.pdf
http://www.uni-bielefeld.de/(de)/biologie/vhf/SF/documents/Phylogeny%20and%20systematics%20of%20salamanders/Veith%20et%20al.%201998.pdf
JSTOR: An Error Occurred Setting Your User Cookie [abstract only]
http://wwwedit.uni-bielefeld.de/bio...behavioral aspects/Steinfartz et al. 2000.pdf
From this one ^^^ I would conclude that morenica, crespoi, and longirostris comprise one or two separate species from all other "Salamandra salamandra", while bernardezi and gigliolii comprise one to three additional distinct species. The authors reach similar conclusions when they say "Thus, molecular and morphological data conform very well and one might, in the long term, consider assigning species status to the so far discussed subspecies." (p.406). All of the remainder [gallaica, fastuosa, bonalli, bejarae, hispanica, terrestris, some gigliolii, werneri, beschkovi, almanzoris, terrestris, salamandra, and implicity alfredschmidti] would amount to variously localized color variants of Salamandra salamandra, with no apparent subspecies. I say this because crespoi, morenica, and longirostris form distinct lineages separately and together; gigliolii and bernardezi form three distinct lineages separately and together; and ALL of the rest are indistinct,both separately and together. That is: 6 samples of gallaica were as different from one another as from 2 hispanica, 2 bejarae, 9 salamandra, and 10 terrestris (among others). So there is basically just as much variation between and WITHIN some populations as there is between subspecies, which makes most of those subspecies meaningless. That's not exactly a direct answer to your question, but it does coincide with my first instinct: some authors likely consider bejarae [and hispanica] to be populations of gallaica.

http://vipersgarden.at/PDF_files/PDF-081.pdf
Data from this study ^^^ would seem to confirm both evolutionary and biological species differences between almanzoris and bejarae (but NOT necessarily that each is valid - only that the two forms would seem to belong to separate species). It also suggests the possibility that bejarae alone could be composed of several distinctive groups. This all agrees with the study above [high diversity within most named subspecies], except their placement of almanzoris close to morenica and crespoi [closer to salamandra in the previous study]. This is likely at least partly due to the mode of inheritance studied, as this study is based on mtDNA, inherited via females only. The addition of one foreign female to a population can hypothetically [demonstrated in Plethodon shermani, Batrachoseps, and Batrachuperus] cause all or part of a species to appear closer to an unrelated species than to its closest kin [as demonstrated by nuclear DNA or morphology, both of which recombine and are subject to selective pressures].
http://www.biocongroup.eu/BioCon/R.Castilho_files/Reis_Genetica_2011.pdf
This is a good comparison ^^^. S.longirostris is clearly isolated from the rest, while morenica, crespoi, and almanzoris are grouped together as distinct entities. Bernardezi forms another distinct group made up of deep branches, but including one sample each of bejarae and gallaica. I would assume these to be animals which inherited genes of relatively recent hybridization with bernardezi. The fourth lineage contains gallaica plus two samples of bejarae and one each of bernardezi and salamandra. My conclusion would be that the bejarae are DCLs of gallaica [Deep Conspecific Lineages - ie, inheritors of the ancient genepool now known mostly as gallaica], while the other two represent hybrid introgression from secondary contact of the northern branch (salamandra and bernardezi) with the western Iberia branch (gallaica and bejarae). Based on this study, and in light of the others, I would treat S.morenica, S.crespoi, S.almanzoris, and S.longirostris as full species. I would treat S.s.bejarae as a synonym of S.s.gallaica, with the reservation that future studies could confirm species status of gallaica, with recognition of multiple subspecies or additional species. As I see it, there may be a LOT of distinct species of Salamandra, which differ in allozymes, mtDNA, environmental niche, behavior, reproduction, and morphology, but with only one or two species occupying vast regions. This would be comparable to the leopard frogs and tiger salamanders, in which one or two species has colonized vast regions post-glacially, but in tropical and subtropical regions is replaced by a wealth of species with small to tiny ranges.
http://www.herpetologica.org/nuevos_hallazgos/velo-anton_2007.pdf
A little off-topic, although the above paper ^^^ shows that viviparity is not an especially useful criterion to classify Salamandra, as it can result very quickly in isolated populations under selective environmental pressure. [live-bearing insular populations of S.s.gallaica].
http://vipersgarden.at/PDF_files/PDF-3815.pdf
THIS one is both new to me, and a lot to review. ^^^ Interesting in the inclusion of undescribed forms [in line with my own thoughts] and in the addition of ecological niche modeling [although see the previous paper, regarding rapid adaptation to viviparity within Salamandra lineages].
Heterochrony, cannibalism, and the evolution of viviparity in Salamandra salamandra [eScholarship]
Also off-topic. Interesting comparison of viviparity and ovoviviparity, with S.s.bejarae as one of the samples.
http://www.herpetologica.es/attachments/article/112/Nueva Lista Patrón 2011.pdf

In light of all this, and returning to your original questions...
Older maps and newer ones which treat bejarae as a synonym of gallaica, will map the bejarae range as gallaica. Since bejarae are nested within a more diverse gallaica, recognition of the former is questionable. In fact, recognition of the latter is questionable as well, although some data render the western Iberian Salamandra as distinctive relative to northern forms. Most recent authors seem to treat the ranges of bernardezi, fastuosa, salamandra, and bejarae as continuous and intermixing, to the exclusion of gallaica, although most genetic data merge gallaica and bejarae. Bejarae and almanzoris have intermeshing but distinct ranges in the Sistema Central, with limited hybridization. Morphologically, S.almanzoris and S.s.bejarae can usually be distinguished, but I don't think S.s.gallaica and S.s.bejarae can be, since they have overlapping variation. [I am ASSUMING the name "hispanica" to be younger than "bejarae", and the latter in turn younger than "gallaica". I haven't checked ASW].
 

Azhael

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Thank you! I´m going to need some coffee for these :)
 
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