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Data from: Causes and evolutionary consequences of population subdivision of an Iberian mountain lizard, Iberolacerta monticola
Aim: The study of the factors that influence population connectivity and spatial distribution of genetic variation is crucial for understanding speciation and for predicting the effects of landscape modification and habitat fragmentation, which are considered severe threats to global biodiversity. This dual perspective is obtained from analyses of subalpine mountain species, whose present distribution may have been shaped both by cyclical climate changes over ice ages and anthropogenic perturbations of their habitats. Here, we examine the phylogeography, population structure and genetic diversity of the lacertid lizard Iberolacerta monticola, an endemism considered to be facing a high risk of extinction in several populations. Location: Northwestern quadrant of the Iberian Peninsula. Methods: We analyzed the mtDNA variation at the control region (454 bp) and the cytochrome b (598 bp) loci, as well as at 10 nuclear microsatellite loci from 17 populations throughout the distribution range of the species. Results: According to nuclear markers, most sampling sites are defined as distinct, genetically differentiated populations, and many of them show traces of recent bottlenecks. Mitochondrial data identify a relatively old, geographically restricted lineage, and four to six younger geographically vicariant sister clades, whose origin may be traced back to the mid-Pleistocene revolution, with several subclades possibly associated to the mid-Bruhnes transition. Geographic range fragmentation of one of these clades, which includes lowland sites, is very recent, and most likely due to the accelerated loss of Atlantic forests by human intervention. Main Conclusions: Altogether, the data fit a "refugia within refugia" model, some lack of pattern uniformity notwithstanding, and suggest that these mountains might be the cradles of new species of Iberolacerta. However, the changes operated during the Holocene severely compromise the long-term survival of those genetic lineages more exposed to the anthropogenic perturbations of their habitats.
FIGURE 10 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 10. Photographs of some I. monticola specimens from several different localities across its distribution range: A, male (above) and female (below) from Llagu la Cueva, Somiedo (Asturias, Spain). These are perhaps the most common patterns in the main part of the species area. B, young male from the Picos de Europa (Asturias, Spain). Lizards from this area (males and females) take on a vivid green colour during the breeding period. C, young male from Cabeza Grande de Manzaneda (Orense, Spain). D, old female from Cabeza Grande de Manzaneda (Orense, Spain). E, wellgrown brown coloured male from Puerto de Vegarada (León, Spain). In some localities where the species inhabits earth taluses instead of rock outcrops, cryptic brown colours seem to be adopted. F, two hatchlings from Puerto de Vegarada (León, Spain). G, old female from Puerto de las Señales (León, Spain). H, adult male from A Torre, Serra da Estrela (Beira Alta, Portugal). Fullygrown specimens from this locality are very patterned, with a greyish, very light brown or more frequently green and occasionally bluish base colour.
FIGURE 9 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 9. Specimens of I. galani nov. and pictures of localites across its distribution range: A, hatchlings from Puerto del Morredero (Sierra del Teleno, León, Spain). B, hatchling from the Laguna de los Peces (Sierra Segundera, Zamora, Spain). C, female (paratype) melanistic from Puerto del Morredero (Sierra del Teleno, León, Spain). D, male (paratype) from the Lago de Truchillas (Sierra de la Cabrera Baja, León, Spain). Notice the malachite greenish tinge of the venter and the numerous ocelli on the shoulders. E, Puerto del Morredero (Sierra del Teleno, León, Spain). There is a small ski resort but I. galani nov. mainly inhabits the road and tracktaluses. F, Puerto de los Portillinos (Sierra de Teleno, León, Spain) is situated near the Puerto del Morredero. In this locality I. galani nov. inhabits not only the road taluses but also the extensive rocky (quarzites) outcrops at the higher parts and is syntopic with Podarcis bocagei. G, granitoid outcrops surrounded by heathland seen from the dam of the Laguna de los Peces (Sanabria). This is the typical habitat of I. galani nov. in Sanabria and is syntopic with Lacerta schreiberi. The lizard is especially abundant near the glacial lakes of this area. H, abandoned ski resort in the Peña Trevinca area. Iberolacerta galani nov. inhabits the slate slab accumulations by the road and track taluses such as the one seen in the centre of the picture (which constitutes the border between the provinces of León and Orense). It is syntopic with Podarcis bocagei in this area.
FIGURE 6 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 6. Morphology of the squamosal bone in Iberolacerta. Iberolacerta galani nov. (represented in black) has a fairly straight squamosal in comparison with the more gradually incurved ones of the other Iberolacerta spp. 1) I. monticola (male) from Sª Estrela; 2) I. monticola (female) from Sª Estrela; 3) I. monticola (female) from Sª Caurel; 4) I. monticola (female) from Pto. de las Señales; 5) I. monticola (male) from Somiedo; 6) I. monticola (male) from Somiedo; 7) I. monticola (female) from Cabeza de Manzaneda. 8) I. galani nov. (male) from Sanabria; 9) I. galani nov. (female) from Sanabria. 10) I. cyreni (male) from Pto. de Navacerrada; 11) I. cyreni (male) from Gredos; 12) I. cyreni (male) from Gredos; 13) I. martinezricai (female) from Las Batuecas.; 14) I. martinezricai (female) from las Batuecas; 15) I. aurelioi (female) from Coma Pedrosa (Andorra); 16) I. bonnali (male) from Bigorre; 17) I. aranica (female) from Mauberme massif; 18) I. horvathi (female) from Udine (Italy).
FIGURE 8 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 8. Specimens of I. galani nov. from several different localities across its distribution range: A–B, male (holotype) from the Laguna de los Peces (Sierra Segundera, Zamora, Spain) viewed from both sides. C, female (paratype) from Puerto del Morredero (Sierra del Teleno, León, Spain). D, female (ventral side) from the same locality as in C, ventral view. E, male (paratype) from Puerto de los Portillinos (Sierra del Teleno, León, Spain). F, same male as in E photographed under ultraviolet light. Notice the highly reflective blue ocelli on the shoulders. G, female (paratype) from the Trevinca Massif (Sierra del Eje, León, Spain). H, same female as in G, ventral view.
FIGURE 5 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 5. Karyograms of Iberian Iberolacerta species with 2n=36 chromosomes, showing distribution of heterochromatin (solid black blocks) and localization of NORs (black circles).
FIGURE 3. Maximumlikelihood tree using 1041 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 3. Maximumlikelihood tree using 1041 base pairs of the mitochondrial genes cytb and 12SrRNA and the nuclear gene cmos. Support values are presented above the branches (posterior probabilities, only asterisk if value is equal to or above 95%) and below the branches (left bootstrap value for ML and right bootstrap value for MP). Datations are presented for some nodes and are indicated by a dot and a value in My (millions of years).
FIGURE 2 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 2. Canonical Discriminant Analysis (CDA). Threedimensional representation of the three first canonical axes. OTUs (same abbreviations as in text) are represented by their centroid. The Minimumlength Spanning Tree (MST) connecting the closest samples is superimposed. A) Male analysis (76.9% of all explained variability). B) Female analysis (78.5% of all explained variability).
FIGURE 4 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 4. Metaphase plates (A, CF) and karyotype (B) of male (A) and female (BF) of I. galani from Sanabria, Giemsa stained (A and B), AgNOR banded (C), Cbanded (D) and sequentially stained with Cbanding+CMA3(E)+DAPI(F). Asterisks in A and C point to chromosomes bearing secondary constrictions and AgNOR positive spots, respectively; empty and filled arrows in DF point to W and Z chromosomes, respectively.
FIGURE 7 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 7. Lateral view of the head of the holotype of I. galani nov. from the Laguna de los Peces (Sanabria, Zamora), Spain.
FIGURE 1 in Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae)
FIGURE 1. Map of the Iberian Peninsula showing the current distribution ranges of all known species and subspecies of Iberolacerta (shaded areas). Numbers refer to sampling localities as in Table 1.
FIGURE 13 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 13. Comparison of the expected outcomes of four major geographic models of evolution. M = I. monticola s. str.; G = I. galani; NML = Iberolacerta monticola astur ssp. nov. from the Northern Montes de León. The three main cladogenetic events dated by mitochondrial markers are indicated by the width of the lines separating the populations. Block arrows indicate direction of gene flow after the initial separation of I. m. astur ssp. nov. Arrowheads within text boxes under each model mark the expected tendency (upward or downward) of the corresponding indicators of nuclear genetic divergence. Reference values of divergence, obtained from the observed data, are shown at the bottom of the figure.
FIGURE 9 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 9. Specimens of I. monticola astur ssp. nov. from several localities across its distribution range: A.—Adult male in breeding period, with the typical green coloration. Note a typical disposition of scales in front of the masseteric plate. Salientes (Palacios del Sil, León)[loc. 2 in Fig. 1A]. B.—Ventral view of the same specimen of Fig.9 A. Throat and belly show greenish tinges, being in this later in a gradation from darker toward the sides to clearer in the middle. C.—A young male specimen of the vermiculated morph in breeding period. Greenish to brownish gray above. Salientes (Palacios del Sil, León) [loc. 2 in Fig. 1A]. D.—Ventral view of the same specimen as in C. E.—HOLOTYPUS (MNCN 44652). A brown colored male with eclipsed coloration in summer (probably also brown during the breeding period). Salientes (Palacios del Sil, León) [loc. 2 in Fig. 1A]. F.—Male from Sierra de las Tiendas (Murias de Paredes, León) [loc. 10 in Fig. 1A]. Eclipsed coloration still showing green coloration. G.—Male from Sierra de Villabandin (Murias de Paredes, León) [loc. 14 in Fig. 1A]. Summer (eclipsed) coloration. Moreover the animal is close to the molt and shows a very dark appearance due to the old skin. In this locality, NML-II haplotypes have been identified. However, these animals fall perfectly inside the morphological variability of the NML-I haplotype populations. H.—Hatchlings from Salientes (Palacios del Sil, León) [parents from loc. 2 in Fig. 1A]. Tail tips cut for genetic analysis.
FIGURE 1. A in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 1. A) General situation and localities of Iberolacerta monticola in the study area. Iberolacerta monticola s. str. occupies the main axis of the Cantabrian Range, north to the Luna and Sil rivers (inverted triangles), and I. m. astur ssp. nov. (round symbols) the Northern Montes de León, south to the abovementioned rivers. Localities of I. m. astur ssp. nov. are: 1.—Pico Nevadín and Braña de Peña Vendimia. 2.—La Poza del Puerto, Puerto del Portillin (=Alto de Vivero). 3.—Vivero. 4.—El Molar de Montrando, El Tambarón. 5.—Braña La Forcada, Pico de la Robeza, Peña Carnicera, Cerneya and Peñona de Brañalibrán. 6.—Peña de Valdiglesia. 7.—Alto de los Grillos. 8.—Catoute. 9.—Collado de Ocidiello and Pico Arcos del Agua (=Fernán Pérez). 10.—Sierra de las Tiendas. 11.—El Suspirón and Mortera de la Vieja. 12.—Los Bayos. 13.—Valdeloso Valley. 14.—Villabandin (Sierra de Villabandin, Southern Slopes) and 15.—Riolago de Babia (Sierra de Villabandin, northern slopes).
FIGURE 2 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 2. Principal Components Analysis. Males. Bidimensional plot of the specimens scores. Triangle: BABIA (Northern slopes of Sierra de Villabandín; NML-II haplotype). Inverted triangles: SALIENTES (Nevadin-Tambarón-Catoute populations; NML-I haplotype). Crosses: TIENDAS & SUSPIRON (Sierra de las Tiendas and Suspiron peak; NML-II haplotype). Blades: VILLABANDIN (Sierra de Villabandín or La Cañada; NML-II haplotype). Despite to be significant, the two first axes had a very poor discriminative power, with only a 33.8 % of variance explained (see text for details) and show a considerable overlap of the NML-I specimens that are distributed overlapping with the main part of the other samples. The second axis seems to have a geo-climatic interpretation, separating in the positive part the most northern (oceanic influenced) samples (BABIA) and in the negative part the ones inhabiting the more continental and harsh climate areas facing to the Spanish Northern Plateau (Meseta) (VILLABANDIN -Southern slopes of the same Sierra de Villabandin-), BAYOS, TIENDAS and SUSPIRON. Note that BABIA (triangle symbol) specimens are geographical neighbors (in the northern slopes) of VILLABANDIN (blades symbol) (in the southern slopes), but appear totally opposite in the PCA, in accord with this bioclimatic influence in the scalation. These more continental climate populations show the greater values for most scalation characteristics and the smaller ones for Tympanic plate size.
FIGURE 8 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 8. Three-dimensional representation of a Factorial Correspondence Analysis based on microsatellite genotypes of Iberolacerta monticola astur ssp. nov. individuals from the Northern Montes de León. The figure includes individuals of I. galani and I. monticola s. str. to be used as reference.
FIGURE 7 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 7. Summary plot of estimated membership coefficients for each individual, in each cluster (Q). Each individual is represented by a single vertical line broken into k colored segments, with lengths proportional to each of the k inferred clusters. The numbers (1–4) correspond to the predefined populations.
FIGURE 3 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 3. Principal Components Analysis. Females. Symbols as in Fig. 2. As in males, it is significant but also has a poor discrimination of the specimens into differentiated samples. The two axes together reach 32.3 % of variability explained (Fist axis: Eigenvalue of 2.91 and 17.14 % variance; Second axis 2.57 (15.14 % of variability), which is a very poor discriminative result. As in the male analysis, SALIENTES samples overlap with all the other ones, not allowing discriminating among groups or any geographic or climatic interpretation of the axes.
FIGURE 5 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 5. Canonical Discriminant Analysis. Females. First axis has two thirds of the total variability (Eigenvalue=1.3796; 69 % variance; Canonical Correlation 0.76) whereas the second one had the remaining one third (eigenvalue 0.6169, 31 % variance; Canonical Correlation 0.61). The female analysis presents less overlap among samples than the male analysis. The first axis separates I. galani in its positive part from I. monticola s. str. and I.m. astur ssp. nov. in its negative part. The second axis separates I.m. astur ssp. nov. in its positive part from the other two taxa. Discriminant axes obtained an 80.7 % of correct reclassification. Contrary to the male analysis, the I. m. astur ssp. nov. centroid is closer to the centroid of I. monticola than to the centroid of I. galani.
FIGURE 4 in A new mountain lizard from Montes de León (NW Iberian Peninsula): Iberolacerta monticola astur ssp. nov. (Squamata: Lacertidae)
FIGURE 4. Canonical Discriminant Analysis. Males. Projection of the first two axes. First axis: Eigenvalue=0.958; 70.6 % variance explained; Canonical Correlation 0.70. Second axis: Eigenvalue=0.39; 29.4 % variance explained; Canonical Correlation 0.53. Both axes are significant and configure a bidimensional space with moderate overlap among the three taxa. The first axis separates I. monticola s. str. in its negative part from I. m. astur ssp. nov. and I. galani in the positive one. The second axis separates I. m. astur ssp. nov. from the other two taxa. These discriminant axes obtain a 78.3 % of correct reclassification. Iberolacerta galani and I. m. astur ssp. nov. centroids are the closest in the males bidimensional space, thus, their males are more similar in our analysis.
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