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FIGURE 5 in A subspecies of marbled newt (Triturus marmoratus) in the Iberian Peninsula newly resolved from congruent nuclear and mitochondrial DNA data

FIGURE 5 The Iberian Peninsula with the ranges of Triturus pygmaeus in light red and T. rudolfi in brown (Arntzen, 2023, 2024). Populations of the counterpart species T. marmoratus are coloured according to the dominant mitochondrial haplogroups with colours as in the legend and fig. 4. The solid or interrupted black line shows the northern range of T. m. harmannis ssp. nov., as determined by the green, yellow and blue sections in fig. 1. Note that haplogroups 1 and 2 are associated with T. m. marmoratus and that haplogroups 3, 4 and 5 are associated with T. m. harmannis ssp. nov. Haplogroup 6 is associated with T. pygmaeus and T. rudolfi (for details see table 2). Areas shown in white fall outside the documented range of the T. marmoratus species group and areas shown in grey are distant from a sampled locality.

FIGURE A1 in A subspecies of marbled newt (Triturus marmoratus) in the Iberian Peninsula newly resolved from congruent nuclear and mitochondrial DNA data

FIGURE A1 Holotype of Triturus marmoratus harmannis ssp. nov. at right and ventral view. Size bar is 1 cm. Stored at the Museo Nacional de Ciencias Naturales, Madrid, Spain under catalogue number 51792.

FIGURE 3 in A subspecies of marbled newt (Triturus marmoratus) in the Iberian Peninsula newly resolved from congruent nuclear and mitochondrial DNA data

FIGURE 3 Histogram of scores along the first axis for the discriminant analysis of size corrected morphometric data for Triturus marmoratus marmoratus (shaded bars) and T. m. harmannis ssp. nov. (open bars). This first discriminant axis is most strongly correlated with extremity lengths and not the other characters. Highest loadings on the second axis are for head length and head width (results not shown).

FIGURE 1 in A subspecies of marbled newt (Triturus marmoratus) in the Iberian Peninsula newly resolved from congruent nuclear and mitochondrial DNA data

FIGURE 1 Classification and geographical distribution of European marbled and pygmy newts from a panel of 32–33 nuclear genetic markers (data from Kazilas et al., 2024). (A) HIests plot with ancestry and heterozygosity for within marbled newts (top panel, with Triturus m. marmoratus left and T. m. harmannis ssp. nov. to the right) and for pygmy newts (bottom panel, with T. rudolfi to the left and T. pygmaeus to the right). (B) Investigated Iberian populations shown by black dots with surrounding areas coloured as in A. Areas shown in white fall outside the documented range of the T. marmoratus species group and areas in shown grey are distant from a sampled locality. The open square symbol in the Lisbon Peninsula corresponds to the open round symbol in A.

FIGURE 4 in Morphological and genetic diversification of pygmy and marbled newts, with the description of a new species from the wider Lisbon Peninsula (Triturus, Salamandridae)

FIGURE 4 Geographical clines observed for 'Triturus pygmaeus' in a longitudinal transect across central Portugal (see fig. 3), with T. rudolfi nov. sp. in the west and T. pygmaeus in the east. The horizontal axis is distance in km, measured from the nominal centre of the contact zone at Entroncamento railway station. The vertical axes are from top to bottom, left column – PCA2 for 54 SNP markers and the frequency of the eastern mtDNA haplotype, and in the right-hand column – the number of links and body size (lnSVl1) of adult males and females. Solid dots represent populations and the grey areas represent the 95% credibility intervals. Note that the colour bar is applied to fig. 3B. The formal cline descriptions are in supplementary table S3.

FIGURE 2 in Morphological and genetic diversification of Old-World marbled newts, with the description of a new and 'not-at-all-cryptic' subspecies from the Iberian Peninsula (Triturus, Salamandridae)

FIGURE 2 The distribution of two species of marbled newts over the western part of the Iberian Peninsula, as inferred from the morphological character Nlinks. Triturus marmoratus is shown in two shades of blue, T. pygmaeus in two shades of red and four localities with both species are shown in grey (see colour legend). The spatial extrapolation does not exceed ca. 50 km. Note that southern Iberian populations are all T. pygmaeus, irrespective of high (cluster C1), low (C2), or bimodal Nlink counts (Doñana National Park). Populations that were studied morphometrically are highlighted by a box. The type locality for T. pygmaeus lusitanicus ssp. nov. is Granja, Portugal and is marked by an asterisk.

FIGURE A1 in Morphological and genetic diversification of pygmy and marbled newts, with the description of a new species from the wider Lisbon Peninsula (Triturus, Salamandridae)

FIGURE A1 Holotype of Triturus rudolfi sp. nov. at right (top) and left lateral view (middle), and in ventral view (bottom). Size bar is 1 cm. Stored at the Museo Nacional de Ciencias Naturales, Madrid, Spain under catalogue number 51784. Downloaded from Brill.com 07/10/2024 02:01:22PM via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 license. https://creativecommons.org/licenses/by/4.0/

FIGURE 4 in Morphological and genetic diversification of Old-World marbled newts, with the description of a new and 'not-at-all-cryptic' subspecies from the Iberian Peninsula (Triturus, Salamandridae)

FIGURE 4 Morphological and genetic variation in Triturus pygmaeus from in and around Doñana National Park (DNP). A – histograms of Nlink counts for four population groups with, from top to bottom: southwestern cluster of T. pygmaeus (localities 65, 436, 438 and 1004 in the south of Portugal, together marked C1 in fig. 2), populations in the northern section of DNP (ER, locality 1005 and BS, locality 1006), population in the southern section of DNP (locality 271) and the Betic cluster of T. pygmaeus (localities 270, 463, 465 and 1003 in the very south of Spain, together marked C2 in fig. 1). Exceptionally, the northern Doñana museum material contains 15 juvenile specimens (open bars) along with three adults (shaded bars). B – geographical cline in microsatellite genetic variation. The horizontal axis shows 'Distance from the Guadalquivir River' in km and the vertical axis presents Structure Q-scores. The solid round symbols represent populations and the grey area shows the 95% credibility interval. The Structure Q-scores were extracted from a published figure (Albert & García-Navas, 2022: Figure 4) and are unavoidably imprecise. The position of the 'Torre Carbonero', a cultural landmark at the Doñana beach, is shown by an arrow (TC). Three populations with morphological data available, that can be associated to the transect are El Rocío (arrow marked ER) and the biological station (BS) in northern Doñana and Zacallón C. de los Junqueros in southern Doñana (arrow marked 271). For details, see table 1 and supplementary table S1.

FIGURE 5 in Morphological and genetic diversification of pygmy and marbled newts, with the description of a new species from the wider Lisbon Peninsula (Triturus, Salamandridae)

FIGURE 5 Histogram for the number of links (Nlinks) observed in Iberian large-bodied newts, with low values for Triturus marmoratus (top panel), intermediate values for T. rudolfi sp. nov. (middle panel) and low to high values for T. pygmaeus (bottom panel). A distinction is made between T. p. pygmaeus from the Betic region (grey bars) and T. p. lusitanicus from the remainder of the species range (open bars) (see Arntzen, 2024). The optimal separation of T. rudolfi nov. sp. versus T. p. lusitanicus is achieved at Nlinks = 6.8, as shown by an interrupted line. To the right examples are shown of individuals with low and high link counts. Animals are facing left, with T. marmoratus from Gerês, northern Portugal at the top and T. p. lusitanicus from Sagres, southern Portugal at the bottom. Links are counted over the left and right side of the newts' bodies, in between the insertion of the fore- and hind leg. THE IMAGERY IS REPRODUCED FROM ARNTZEN (2018)

FIGURE 3 in Morphological and genetic diversification of Old-World marbled newts, with the description of a new and 'not-at-all-cryptic' subspecies from the Iberian Peninsula (Triturus, Salamandridae)

FIGURE 3 Nine marbled newts from three (sub)species each in dorsal and ventral view. Link counts for the left and right side of the body are given in parentheses. Top row – Triturus marmoratus from Jublains, France; left male (3, 2) and right female (1, 1). Middle row – T. pygmaeus lusitanicus ssp. nov. from Cardeña, Spain; from left to right female (2, 4), male (4, 3) and male (4, 5). Bottom row – T. p. pygmaeus from Salinas, Spain; from left to right male (2, 1), male (1, 2) male (2,0) and female (3, 1). Note that the colour pattern in T. marmoratus and T. p. pygmaeus is horizontally banded whereas in T. p. lusitanicus ssp. nov. it is reticulated. PHOTOGRAPHY L. A. VAN DER LAAN

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Morphometric data to describe phenotypical variation across Pygmy and Marbled newts (genus Triturus)

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FIGURE 4 in A subspecies of marbled newt (Triturus marmoratus) in the Iberian Peninsula newly resolved from congruent nuclear and mitochondrial DNA data

FIGURE 4 Minimum spanning network for mitochondrial ND 4 sequences retrieved from pygmy newts (Triturus rudolfi and T. pygmaeus) with a single haplogroup in red (H6), and marbled newts (T. marmoratus) with five haplogroups shown as H1 – green, H2 – light green, H3 – light blue, H4 – blue and H5 – purple. Satellite haplotypes are allocated to the nearest main haplotype, based on minimum uncorrected p-distances (details see text). Size of the circles corresponds to sample size (see legend). Internal and external branches have lengths of one or two substitutions, except for those marked with an 'x' that have 5‒7 substitutions.

FIGURE 2 in A subspecies of marbled newt (Triturus marmoratus) in the Iberian Peninsula newly resolved from congruent nuclear and mitochondrial DNA data

FIGURE 2 Histograms of Nlink values observed for adults of Triturus m. marmoratus (N = 167, shaded bars) and T. m. harmannis ssp. nov. (N = 635, open bars), with (A) all data and (B) averages for populations with a sample size ≥ 10. (C) Juvenile T. marmoratus from the subspecies transition area (Poço do Inferno near Valongo, Portugal). The photo aims to illustrate the species, and to discuss the usefulness of the character Nlinks for subspecies identification (details see text).

FIGURE 5 in Morphological and genetic diversification of Old-World marbled newts, with the description of a new and 'not-at-all-cryptic' subspecies from the Iberian Peninsula (Triturus, Salamandridae)

FIGURE 5 Results of a principal component analysis on size corrected morphometric data for Triturus pygmaeus pygmaeus (solid lines) and T. p. lusitanicus ssp. nov. (interrupted lines). The ellipses show the 95% confidence interval of the mean. The left panel is for males and the right panel is for females. Percentages of total variance explained are indicated along the axes. For details, see text.

FIGURE 3 in Morphological and genetic diversification of pygmy and marbled newts, with the description of a new species from the wider Lisbon Peninsula (Triturus, Salamandridae)

FIGURE 3 (A) Genetically investigated populations of large-bodied newts from the western part of the Iberian Peninsula plotted with the nearest point method, with a maximum spatial extrapolation of ca. 25 km. Colour codes are: blue – T. marmoratus, red – T. pygmaeus and ochre – T. rudolfi sp. nov. Inferred hybrid populations of T. marmoratus and 'T. pygmaeus' are shown in grey (as in fig. 2). Note that populations from densely sampled areas do not always stand out separately. Five populations from along the lower Tejo area added a posteriori are shown by black dots. The boxed area in between 39.3-39.7 N encompasses the transect studied for spatial variation in morphometrics (35 populations), Nlinks (35 populations), mitochondrial DNA (69 populations) and nuclear genetic composition (82 populations) with HZAR software (for results see fig. 4). The nominal centre of the T. pygmaeus – T. rudolfi sp. nov. hybrid zone is at Entroncamento railway station (E, coordinates -8.478 E, 39.456 N). (B) Dirichlet tessellation of populations studied for variation in nuclear genetic composition, with colours like above. The yellow to red colour bar corresponds to the one shown in fig. 4. Distances in km are relative to Entroncamento railway station. Downloaded from Brill.com 07/10/2024 02:01:22PM via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 license. https://creativecommons.org/licenses/by/4.0/

FIGURE A2 in Morphological and genetic diversification of Old-World marbled newts, with the description of a new and 'not-at-all-cryptic' subspecies from the Iberian Peninsula (Triturus, Salamandridae)

FIGURE A2 Holotype of Triturus pygmaeus lusitanicus ssp. nov. in live condition, at the left (top) and right dorso-lateral view (middle), and in ventral view (bottom). The scale bar equals 10 mm.

FIGURE 1 in Morphological and genetic diversification of Old-World marbled newts, with the description of a new and 'not-at-all-cryptic' subspecies from the Iberian Peninsula (Triturus, Salamandridae)

FIGURE 1 Distribution of marbled newts in the Iberian Peninsula with Triturus marmoratus in blue and T. pygmaeus in red (after Wielstra et al., 2014). The arrow indicates the position of a T. pygmaeus subspecies border inside Doñana National Park (present paper). Downloaded from Brill.com 07/10/2024 02:00:28PM via Open Access. This is an open access article distributed under the terms of the CC BY 4.0 license. https://creativecommons.org/licenses/by/4.0/

FIGURE A1 in Morphological and genetic diversification of Old-World marbled newts, with the description of a new and 'not-at-all-cryptic' subspecies from the Iberian Peninsula (Triturus, Salamandridae)

FIGURE A1 Variation in ventral colouration of Triturus pygmaeus. Top row – T. p. lusitanicus ssp. nov.: male (with dark tail underside) and two females from Villalba and male from Cardeña. Middle row – T. p. pygmaeus: male from Salinas, male and female from Los Barrios and male from Rio Alberite. Bottow row – T. p. pygmaeus: two males and two females from Rio Alberite. PHOTOGRAPHY L. A. VAN DER LAAN

FIGURE 1 in Morphological and genetic diversification of pygmy and marbled newts, with the description of a new species from the wider Lisbon Peninsula (Triturus, Salamandridae)

FIGURE 1 The Iberian Peninsula with the approximate distributions of four taxa of large-bodied newts as described in table 1. Colour codes are: blue – Triturus marmoratus, dark red – T. pygmaeus pygmaeus, light red – T. p. lusitanicus and brown – T. rudolfi nov. sp., i.e., the newly described species from the wider Lisbon Peninsula. Letters indicate the capital cities Lisbon, Portugal (L) and Madrid, Spain (M) as well as Peniche (P) at the Atlantic coast. Major rivers partially coinciding with (sub)species borders are the Guadalquivir, the Vouga and the Tejo. The new species' type locality Lagoa Seca near Valado dos Frades is indicated by a long arrow. The insert shows an adult male T. marmoratus. ANIMAL DRAWING BY BAS BLANKEVOORT