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].
