identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03E587C7FF9CCD0FFF73FA7FFC47F834.text	03E587C7FF9CCD0FFF73FA7FFC47F834.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. astreptophora (Lopez Seoane 1884)	<div><p>Natrix astreptophora astreptophora (López Seoane, 1884) – phenotypic variation</p><p>According to our analysis, Natrix astreptophora astreptophora from the Iberian Peninsula and France (Fig. 2)displays considerable variation in coloration and pattern. Three different non-melanistic phenotypes can be distinguished. Their occurrence is not mutually exclusive, but two phenotypes are geographically restricted. Kindler et al. (2018) had included samples from the entire European distribution range in their phylogeographic study, and since no obvious genetic differentiation was revealed, these distinct phenotypes merely represent local variation without taxonomic relevance.</p><p>Regular phenotype (n=490)</p><p>The most widespread phenotype occurs across the whole European range, and snakes representing this phenotype are always characterized by an orange to reddish iris. Hatchlings and small juveniles have a highly variable primary color, generally beige, grey, or greenish. Their dorsal body pattern consists of small black spots.Almost all hatchlings and small juveniles have a yellow collar, usually with black markings in the nuchal region; rarely the collar is whitish instead. Only 19 of the 47 studied hatchlings and small juveniles from Andalusia had a bicolored pileus. After the first year of life, the black color of the pileus, if present, fades. Later, the light collar increasingly fades and the primary body color becomes green or greenish; rarely adults may even show a turquoise body coloration. Typically, only aged females completely lose the black nuchal markings and the dark dorsal spots on the body, resembling then the pale northern phenotype (see below). In some populations (Puertollano, Girona, Alicante), also young adults lack dark dorsal markings, but differ from northern grass snakes in their primary coloration (beige instead of chestnut in northern conspecifics, see below). In Murcia, the Comunidad Valenciana, Catalonia, and France, adult specimens can show contrasting black nuchal and dorsal markings, similar to the western phenotype (see below), although the dorsal spots are typically smaller and the green color is more vivid in the regular phenotype. Other snakes in these areas resemble Moroccan grass snakes, but with more dorsal black markings and without the strong black color in the pileus in juveniles and young adults, typical of the Moroccan specimens.</p><p>The largest known N. astreptophora belongs to the regular phenotype. It was found in Toledo, Spain, and had a SVL of 112 cm and an estimated TL of 142 cm (García-Antón et al. 2017). For Andalusia, specimens with similar sizes are known (J.P. González de la Vega pers. comm.).</p><p>Northern phenotype (n=264)</p><p>López Seoane (1884) described Tropidonotus natrix var. astreptophorus from Galicia (Spain) based on the lack of a distinct light collar in adults and the plain chestnut brown body color, without any markings. This plain coloration develops during aging, when the light collar and the dark body markings increasingly fade. Such pale-colored adults occur across the whole European distribution range, but in northern and northwestern Iberia, all aged snakes are pale colored, i.e., in Galicia, Asturias, Cantabria, the Basque Country, the northern parts of Castilla León (north of Zamora, León, and Burgos), and the north of Portugal (Fig. 3). In other regions, pale-colored adults co-occur with patterned adults that retained at least some traces of the collar and body spots. Most pale individuals represent fully adult and aged snakes. An adult syntype of T. n. var. astreptophorus in the collection of the Muséum national d’Histoire naturelle, Paris (MNHN-RA-1889.582, Galicia, Spain), is a largely uniform pale specimen which has left only a few small dark spots on its back (see fig. 11 in Fritz &amp; Schmidtler 2020).</p><p>Hatchlings and small juveniles in the north are more variable patterned than elsewhere: They possess a brownred-grey primary color with many small dark markings, which can occur sometimes as wide bars or as a tessellate pattern. Also, plain-colored hatchlings and small juveniles, without any dark markings, or very dark individuals are frequent. Out of nine hatchlings from Asturias, seven were very dark both in the pileus and primary body color and had dorsally many dark spots. A hatchling syntype of T. n. var. astreptophorus in the collection of the Muséum national d’Histoire naturelle, Paris (MNHN-RA-1889.581, Cuntis, Pontevedra, Spain) shows many dark spots and a closed collar (see fig. 10 in Fritz &amp; Schmidtler 2020).</p><p>Approximately 80% of the hatchlings in these northern populations (n=25) had a reddish iris, while 20% had a yellow iris, resembling most populations of N. helvetica, or a maroon iris, suggestive of incipient melanism. This could be the reason why López Seoane (1884) did not mention a reddish iris in his astreptophora description – if he did not only refer to preserved material.</p><p>The distribution range of the northern phenotype is characterized by an oceanic climate, which is colder than the more southern and Mediterranean regions of Iberia, with cold night temperatures, frequent rainfall, and cloudy weather, even in summer (McKnight &amp; Darrel 2001). We suppose that these climatic conditions explain why we also recorded nearly all melanistic snakes in this region (see below the section on the melanistic phenotype). It seems likely that the plain colored and melanistic N. astreptophora reflect an adaptation to the relatively cool and humid oceanic climate. This adaptation seems to be so effective that N. astreptophora is one of the two most common snake species in Galicia, besides Coronella austriaca (see López Seoane 1884; Asensi Cabirta 2024). Likewise, N. astreptophora was recently identified as the second most frequent snake species in a Citizen Science Project (#Sugebizi of the Aranzadi Society, https://www.aranzadi.eus/sugebizi-proyecto-ciudadano) in the Basque Country, Spain.</p><p>Western phenotype (n=120)</p><p>Along the Atlantic coast of the Iberian Peninsula, from approximately the region of Sines-Botinhas (south of Lisboa, Portugal) to the north of La Coruña (Galicia, Spain), occur N. astreptophora that typically retain an intensely spotted body pattern in the adult stage, with many big dark dorsal markings. Each marking covers 3–6 scales. The dark elements can sometimes have a circular form or be bar-like and arranged like in N. helvetica . Pronounced large black nuchal markings are present, and the body has an orange-brown to dark green primary color; the iris is frequently yellow or maroon. Snakes of the western phenotype may resemble the coloration of N. maura (González de la Vega et al. 2021) . Hatchlings and small juveniles in the distribution range of the western phenotype may possess a rare and unique dorsal pattern with many big round dark markings.</p><p>Notably, we recorded the majority of grass snakes of the western phenotype along or near the coast (La Coruña, Pontevedra, Porto, Lisboa; n=41). However, other snakes of this phenotype were also found inland ( Santiago de Compostela, Braga, Coimbra, Peneda-Gerês, Serra de la Estrela; n=30). Representatives of the other two phenotypes also occurred together with the western phenotype, in particular in western Galicia.</p><p>Finally, it should be noted that there are adult specimens of the three European phenotypes in which the dorsal head coloration differs from that of the body. In the regular phenotype, a bluish head coloration may occur. In the northern phenotype, the head can be greenish, and in the western phenotype, grey. From northwestern Navarra, close to the Basque Country border, two adults of the regular phenotype had retained a yellow collar and had a complete black pileus. In close neighborhood, N. helvetica occurs and we cannot exclude that these unusually colored snakes are hybrids, as genetically confirmed for the Eastern Pyrenees (Asztalos et al. 2020). Figure 4 shows the typical body coloration of the three non-melanistic phenotypes from Europe compared to a Moroccan and a Tunisian grass snake.</p><p>Melanistic phenotype (n=46)</p><p>Completely black snakes are rare in Natrix astreptophora . Many melanistic individuals, even with jet black body and head, retain some lighter colored supralabials. Also, the collar and gular area are often lighter (Fig. 5). We never recorded black individuals with many small light speckles (the so-called ‘ picturata morphotype’) in N. astreptophora . Such snakes are known both from N. helvetica and N. natrix (Fritz &amp; Schmidtler 2020; Bruni et al. 2022; Fritz &amp; Ihlow 2022; Jablonski et al. 2023). The only known grass snake of the picturata morphotype from Spain, figured in Fernández Guiberteau et al. (2015), originates from the Val d’Aran, where N. helvetica occurs (Amat et al. 2024).</p><p>Melanistic N. astreptophora are unevenly distributed across the Iberian Peninsula. From Central Spain, there is only one record known (El Tiemblo, Ávila; J.C. Campos pers. comm.). The remaining 45 out of 46 melanistic snakes (98%) were from the northern and northwestern parts of the peninsula, regions where the northern and western phenotypes of N. a. astreptophora also occur (Fig. 3). The abundance of melanistic grass snakes has already been mentioned by previous studies for these regions (Albadalejo 2008; Fernández Guiberteau et al. 2015). Melanistic individuals of other reptile species ( Vipera seoanei, V. latastei, Coronella austriaca, Chalcides striatus) have also been reported from there (Martínez-Freiría et al. 2012; Benito et al. 2022). Several studies suggested that a dark or melanistic coloration in snakes and other reptiles is generally favorable, and thus under positive selection, in environments with reduced insolation, many cloudy days, or lower temperatures (e.g., Gibson &amp; Falls 1979; Andrén &amp; Nilson 1981; Meijide &amp; Pérez-Melero 1994; García Antón et al. 2023). This also explains the frequent occurrence of dark or melanistic grass snakes in Scandinavia, the northern distribution range in Russia ( N. natrix; Fritz &amp; Ihlow 2022), or at high altitudes in the Alps ( N. helvetica; Neumann et al. 2024).</p></div>	https://treatment.plazi.org/id/03E587C7FF9CCD0FFF73FA7FFC47F834	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
03E587C7FF99CD00FF73FF3CFD9CFBAC.text	03E587C7FF99CD00FF73FF3CFD9CFBAC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. algerica (Hecht 1930)	<div><p>Natrix astreptophora algerica (Hecht, 1930) (n=16)</p><p>Hecht (1930) described the taxon Tropidonotus natrix algericus based on two specimens from Algeria, one each from “Südalgier” (ZMB 19636, holotype) and “ Algier ” (ZMB 1935, paratype). Both are ethanol-preserved specimens in the collection of the Museum of Naturkunde, Berlin, and were examined by us. Fritz &amp; Schmidtler (2020) concluded that the type locality “Südalgier” (southern Algeria) must be corrected to an unidentified site in northern Algeria. In addition, they proposed to recognize Algerian and Tunisian populations, representing a unique genetic lineage of N. astreptophora, under the name Natrix astreptophora algerica (Hecht, 1930) as a distinct subspecies. We concur with this suggestion.</p><p>Grass snakes are only known from a few precise localities in the north of Algeria and Tunisia (Gervais 1848; Strauch 1862; Lallement 1866; Boulenger 1891; Olivier 1894; Doumergue 1901; Gadeau de Kerville 1908; Schleich et al. 1996; Brito et al. 2008; Escoriza &amp; Ben Hassine 2017; Rouag et al. 2024; R. Ouni pers. comm.; Appendix 2). One of them, “Tifret” (F. Lataste in Boulenger 1891), was traditionally identified with a village in northwestern Algeria called Tifrit, representing the only record in that part of the country (Sindaco et al. 2013; Trape 2023). However, according to the trips made by Lataste in Algeria, the correct locality should be Tifrit N’Aït el Hadj (P.- A. Crochet and M. Beddek pers. comm.), a site within the known distribution range of N. astreptophora . Thus, the actual distribution gap between the two North African lineages (Morocco and Algeria-Tunisia) is more than 650 km wide. Another putative locality, Gafsa, central Tunisia, with a physical voucher specimen in the collection of the Muséum national d’Histoire naturelle, Paris (MNHN-RA-1983.507, not studied by us, Appendix 2), is highly questionable because Gafsa is an oasis situated in desert climate.</p><p>Sochurek (1979) proposed that the subspecies “ Natrix natrix algirus ” [sic] is valid, but believed it is distributed from the Middle Atlas to northwestern Tunisia, while he tentatively assigned the grass snakes from the Rif Mountains to “ N. n. astreptophora ”. Sochurek (1979) characterized “ N. n. algirus ” by its closed light collar – this is a trait not present in grass snakes from Algeria and Tunisia. According to Sochurek (1979), the collar is still present in adults and is posteriorly bordered by longer dark nuchal markings compared to “ N. n. astreptophora ”; moreover, the body of “ N. n. algirus ” has frequently small dark spots. When it is considered that European N. a. astreptophora can become plain-colored early in ontogeny, this characterization is mostly correct. However, among 16 grass snakes from Algeria and Tunisia examined for the present study, there are also three large and aged specimens from Algeria (1) and Tunisia (2) without or with very faint collars, two of them with completely or nearly unicolored body (see Fig. 6C, D). The third specimen (BEV.8874) consists of the head only. The TL of the figured specimen from Algeria (Fig. 6D), most likely a female, exceeds 100 cm. Moreover, according to our data, the light collar of N. a. algerica is always interrupted by a narrow dark sagittal line, while Moroccan grass snakes have closed collars. This supports that Sochurek (1979) lumped the two North African lineages together, as is also evident from the distribution range assigned.</p><p>Actually, in all hatchlings, juveniles, and young adults from Algeria (n=5) and Tunisia (n=8) that we examined, the light neck collar is interrupted by a narrow black line (at least one scale wide). In contrast, all 11 individuals from Morocco in which the collar is visible have it continuous and closed. An interrupted collar may also occur in European grass snakes (Fig. 7). Elsewise, the head coloration of hatchlings and small juveniles of N. a. algerica resembled that of Moroccan populations. Hatchlings and small juveniles of the two North African lineages share a white-yellowish color of the collar that extends to the three postoculars, the preocular scales, and the first supralabial scales (with black lines separating the supralabial scales). The last supralabial scale has the same color as the collar, whereas the loreal scale is completely –or almost completely– black.</p><p>Hatchlings and small juveniles from Algeria and Tunisia have a brownish to greyish primary body color with or without small to medium-sized dorsal and lateral dark markings; if present, these are often less contrasting than in N. a. astreptophora and Moroccan grass snakes. Moreover, the shape of the dorsal markings can be different. Moroccan snakes tend to have a solid dark marking, while the dark markings may be divided in the Algerian-Tunisian specimens. Moreover, Algerian and Tunisian grass snakes seem to lose their hatchling spots very early and then become plain colored.</p><p>Based mainly on European material, Pokrant et al. (2016) supported their elevation of N. astreptophora as a distinct species by the lower number of ventral scales compared to other grass snake species (156–166 versus 162– 182). However, Hecht (1930) reported for the two types of Tropidonotus natrix algericus 163 and 171 ventral scales, respectively, suggesting that N. a. algerica might have a higher number of ventrals than European conspecifics. A Tunisian specimen in the collection of the Zoologisches Forschungsmuseum A. Koenig, used by Pokrant et al. (2016) and studied by us, has 163 ventrals (ZFMK 67196); two live snakes from Tunisia measured in May 2025 had 162 and 163 ventrals. In addition, the long tail of N. n. algerica is remarkable, representing 22.0–28.5% of the TL in the three studied complete museum specimens and the two Tunisian live snakes, corresponding to the longest tails known in N. astreptophora .</p></div>	https://treatment.plazi.org/id/03E587C7FF99CD00FF73FF3CFD9CFBAC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
03E587C7FF97CD07FF73F933FE7EF846.text	03E587C7FF97CD07FF73F933FE7EF846.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. soumiafahdae Mármol & Geniez & Neumann & Glaw & Fritz 2025	<div><p>Natrix astreptophora soumiafahdae subsp. nov.</p><p>http://zoobank.org/ urn:lsid:zoobank.org:act: 191629E9-980A-4B9D-AF3E-D17B576F5E38</p><p>Holotype. Naturhistorisches Museum Wien, NMW 32908 (Fig. 8), young adult (male?), Middle Atlas [Morocco], don. Erich Sochurek, 6 June 1982.</p><p>Paratypes. Collection BEV (housed in the CEFE-CNRS of Montpellier), BEV.6431 (adult female) and BEV.6432 (adult male), stream west of Oulmès, 33.4412, -6.0394, 1,069 m a.s.l. (Middle Atlas, Morocco), leg. Jean Garzoni 1964 (Fig. 9) .</p><p>Diagnosis. A medium-sized grass snake, with a known maximum TL of 133 cm in females and 95 cm in males; tail relatively long, representing 22–26% of TL. Iris red-colored. Differs from Natrix helvetica and N. natrix by the presence of a speckled dorsal pattern without pronounced side-bars, the bicolored pileus in hatchlings and small juveniles, and the red-colored eyes. Differs from N. a. algerica by the closed light neck collar and the presence of black dorsal and lateral markings in adults, except for very large aged females. Differs from most N. a. astreptophora by the presence of light spots on the parietal scales, the light color of the upper postocular scale, and the big black spot on the pileus that only disappears in aged individuals, whereas in N. a. astreptophora it is only present in some hatchlings.</p><p>Description of the alcohol-preserved holotype. SVL 49.4 cm, 15.4 cm TaL, corresponding to a TL of 64.8 cm; the tail represents approximately 23.8% of the TL. Body robust, with 19 longitudinal rows of keeled dorsal scales at midbody, 171 smooth ventral scales, 76 pairs of smooth subcaudal scales, anal scale smooth and divided. Body coloration greenish grey with two dorsal rows and two lateral rows of approximately 50 dark markings, those in the dorsal region smaller than the lateral ones. Venter: anterior half of the body yellow with large black quadrangular markings, posterior half of the body inclusive of tail almost completely black. Head well differentiated from body, with big eyes; pholidosis: on both sides, nasal scale divided, 1 loreal, 1 preocular, 3 postoculars, 1 anterior temporal, 2 posterior temporals. Dorsally closed light collar in the neck. Pileus bicolored; brown anteriorly and black posteriorly. In front of the white collar supralabial scales with broad black seams, on each side a large black blotch present, extending downwards from the black pileus spot. Two small round greenish spots on parietal scales.</p><p>Variation. The adult female paratype (BEV.6431) has a SVL of 106 cm; its estimated TL is 130 cm (tail incomplete); 19 scale rows at midbody and 168 ventrals are present, subcaudals were not counted; the body has a long slit ventrally (Fig. 9). The SVL of the adult male paratype (BEV.6432) is 75 cm, its TL is 98 cm; the TaL of 23 cm represents 23.4% of the TL. Nineteen scale rows are present at midbody, and there are 166 ventrals and 73 subcaudals. BEV.6432 is the largest known male of this subspecies; BEV.6431 is close to the largest known female (TL 133 cm; Fahd 2001). As in all grass snakes, the snout of N. a. soumiafahdae subsp. nov. is rounded, and there are 7 (rarely 8) supralabial scales on each side of the head, the 3 rd and 4 th supralabials are in contact with the eye. Generally, there are 1 (rarely 2) loreal scale, 1 preocular scale, 3 postocular scales, 1+2 temporal scales, and 19 rows of dorsal scales at midbody, which are keeled, the anal scale is divided. Aged females may have a very big head (Figs 9, 10). The contrasting hatchling coloration is present at least until an age of 3 years, i.e., the bicolored pileus as described above, the contrasting body pattern, and a well-pronounced light collar posteriorly bordered by two black blotches. In individuals exceeding 70 cm TL (i.e., older than 3 years) the collar fades and the head turns greyish to greenish, so that the pattern may disappear in older individuals.Among the ten adult specimens examined for coloration, the collar and the adjacent black blotches have completely faded out in three very large aged females, and only the lateral black body spots are still slightly visible in one specimen (Figs 9, 10).</p><p>Natrix astreptophora soumiafahdae subsp. nov. resembles N. a. algerica in that the dark part of the bicolored pileus persists much longer during ontogeny than in the nominotypical subspecies. As outlined above, in European hatchlings and small juveniles, the pileus can be even unicolored, a trait we did not observe in any North African specimens. The largest measured Moroccan grass snake with a bicolored pileus has a TL of 70 cm. As in N. a. algerica, Moroccan grass snakes always have the loreal scales black or almost entirely black, whereas in N. a. astreptophora individuals with light colored loreal occur. In all N. a. algerica hatchlings (n=4) and in all Moroccan hatchlings, juveniles, and young adults (n=8) the parietal scales bear light spots. Only 1.2% of the 920 studied N. a. astreptophora have such light spots. Furthermore, the upper postocular scale is always whitish or pure white, like the collar, in Moroccan and Algerian-Tunisian specimens. In contrast, in N. a. astreptophora the upper postocular has the same color as the supraocular scales. A unique trait of the Moroccan grass snakes is that the collar is always closed, without dorsal interruption as in N. a. astreptophora and N. a. algerica. This was already highlighted by Fahd (2001) and confirmed in the present study. In many N. a. astreptophora and in all known N. a. algerica, the collar is in its dorsal center interrupted by at least one dark scale.</p><p>Fahd (2001) reported for three males a ventral scale count of 167–173 and for two females, 166. We counted in the male and female paratypes from Oulmès 166 and 168 ventrals. A male from Moulay Abdeslam had 170 ventrals, a female from the same locality, 166, and the holotype from the Middle Atlas has 171 ventrals. This suggests that Moroccan N. astreptophora have more ventral scales than the nominotypical subspecies (maximum of 166; Pokrant et al. 2016). However, J.P. González de la Vega informed us of three Andalusian grass snakes with 170 and 175 ventral scales, and E. Fernández Meléndez found two specimens with 167 and 170 ventral scales in Andalusia. For a comparison of ventral counts of N. a. algerica, see above.</p><p>Etymology. We dedicate the new subspecies to the Moroccan herpetologist Soumia Fahd, who contributed significantly to the knowledge of Moroccan grass snakes in her doctoral thesis (Fahd 2001) and publications (Fahd &amp; Pleguezuelos 1992, 2001; Mediani et al. 2015).</p><p>Biogeography. Natrix astreptophora soumiafahdae subsp. nov. is only known from Morocco. Together with its counterpart N. a. algerica in Algeria and Tunisia, its distribution pattern matches a well-known phylogeographical paradigm for the Maghreb, with two deeply divergent lineages, (1) in Morocco west of the Moulouya River and (2) east of the Moulouya River to Algeria and Tunisia (Beddek et al. 2018; Kindler et al. 2018). This pattern has been described for many biota, among them scorpions ( Buthus: Sousa et al. 2012), salamanders ( Pleurodeles: Carranza &amp; Arnold 2004; Veith et al. 2004; Salamandra: Merabet et al. 2016), frogs ( Hyla: Dufresnes et al. 2019; Pelophylax: Nicolas et al. 2015; Dufresnes et al. 2024), freshwater turtles ( Emys: Stuckas et al. 2014), lizards ( Ptyodactylus: Metallinou et al. 2015; Timon: Paulo et al. 2008), snakes ( Daboia: Martínez-Freiría et al. 2017; Malpolon: Carranza et al. 2006; Natrix maura: Guicking et al. 2008), and rodents ( Meriones: Lalis et al. 2016).</p><p>Distribution and habitat. The geographical and altitudinal distribution of N. a. soumiafahdae subsp. nov. is determined by humidity and the presence of water bodies. The habitats can be streams, rivers, permanent ponds, and small lagoons or lakes (Fig. 11), including water reservoirs with dams. The distribution range is fragmented and resembles that of Emys orbicularis occidentalis in Morocco (Bons &amp; Geniez 1996; Velo-Antón et al. 2015). The range of N. a. soumiafahdae subsp. nov. includes disjunct occurrences in the Rif Mountains, the Middle Atlas, and a few records on the Atlantic plains (see Appendix 2 for a list of localities). The southernmost records reach the northern foothills of the High Atlas (Fig. 3). Most of the records are near rivers, streams, lagoons and other water bodies close to habitats with well-preserved old forests with Quercus pyrenaica, Cedrus atlantica, and Pinus pinaster as dominant tree species, but some sites are also near crop fields (Fahd 2001). Most records are in areas with high seasonal precipitation, and range from sea level to 1,960 m elevation at the Tanout ou Filal pass (Martínez del Mármol et al. 2019). In the Middle Atlas, most records are near Ifrane and Azrou; N. a. soumiafahdae subsp. nov. was also recorded near Oulmès. It has been recently recorded in the Jbel Tazzeka (Kindler et al. 2018). In the Rif Mountains, it was recorded at Tamorot (Beni Halet) by Galán (1931), and subsequent research increased the number of sightings for these mountains (Fahd &amp; Pleguezuelos 1992, 2001; Mediani et al. 2015), suggesting that the Rif Mountains are currently the stronghold for grass snakes in Morocco.</p><p>Food spectrum. Bons (1967) mentioned for the Middle Atlas the anurans Discoglossus scovazzi and Pelophylax saharicus, and for Oulmès, the skink “ Chalcides chalcides ” as prey for Moroccan grass snakes. We wonder whether the latter food item is due to a confusion with a Malpolon monspessulanus, a typical skink eater, whereas there are not any other records of grass snakes eating skinks in the literature (Braña 1998; Pleguezuelos 2018). Unicolored aged N. astreptophora are sometimes confused with M. monspessulanus, even by herpetologists or naturalists (López Seoane 1884), so that any non-vouchered identification should be treated with caution. Martínez del Mármol et al. (2019: fig. 714) figured an adult N. a. soumiafahdae subsp. nov. trying to swallow an adult Berber toad Sclerophrys mauritanica, suggesting that large individuals prey on big bufonids, as known for other grass snakes (Madsen 1983; Kabisch 1999).</p><p>Defensive behavior. When Natrix astreptophora soumiafahdae subsp. nov. cannot escape, they triangle the head and make puffing noises, or are feigning death, with open mouth, as known from other grass snake taxa (Kabisch 1999; Pokrant et al. 2017). A death feigning individual from Chefchaouen with open mouth is figured in Martínez del Mármol et al. (2019: fig. 715). Also, a bad-smelling liquid can be secreted from the cloaca. There are no reports of autohemorrhaging in this taxon, a behavior reported for N. natrix sensu stricto (Gregory et al. 2007; İğci et al. 2017) and N. helvetica (Amat et al. 2024), and very rarely for N. a. astreptophora (only reported in one specimen from Huesca, northeastern Spain; E. Ruiz Ara pers. comm.). Also, the ‘cobra stance’, known from European N. a. astreptophora (Pokrant et al. 2017), has not yet been described for Moroccan populations.</p><p>Reproduction. Fahd (2001) reported that a male with SVL of 38 cm had a great testicular volume, and two females of SVL 67 cm and 107 cm contained 11 and 28 eggs, respectively. This is in line with Bons (1967), who indicated for Moroccan populations clutch sizes of 10– 50 eggs.</p><p>Conservation status. In the European distribution range, Natrix astreptophora is widespread and in some regions abundant. However, in North Africa the species is generally rare, with a fragmented relict distribution (Bons &amp; Geniez 1996; Schleich et al. 1996; Martínez del Mármol et al. 2019; Fig. 3). The number of historical and recent observations is low (e.g., Bons &amp; Geniez 1996; Kalboussi &amp; Achour 2018). Starting with Gervais (1848) and the general mention of the species for Algeria by Guichenot (1850), we traced a total of only 27 records for Tunisia and Algeria (excluding the questionable localities Gafsa and “Südalgier”) and 55 records for Morocco (Appendix 2), confirming the rarity of this snake species in the Maghreb.</p><p>The new subspecies Natrix astreptophora soumiafahdae subsp. nov. is distributed allopatrically and endemic to northwestern Morocco (Fig. 3). It is largely confined to montane areas in temperate habitats characterized by high humidity and dense plant cover, usually near streams, ponds, and lakes (Fahd &amp; Pleguezuelos 1992, 2001; Martínez del Mármol et al. 2019; Fig. 11), with a few isolated records from the Atlantic plains (Fig. 3), remarkably in the same region from where also European pond turtles ( Emys orbicularis occidentalis), another taxon with a relict distribution in the Maghreb, have been recorded in the past (Bons &amp; Geniez 1996). Apart from the Rif Mountains, where several populations were found (Fahd &amp; Pleguezuelos 2001; Mediani et al. 2015), records from other regions are scattered and scarce, despite targeted searches by experienced herpetologists, including us. Fahd &amp; Pleguezuelos (2001) characterized Moroccan grass snakes to be very rare compared to the congeneric viperine snake N. maura (which might be a strong competitor) and as belonging to the rarest reptiles of Morocco.</p><p>Due to its strong genetic differentiation from both N. a. astreptophora and N. a. algerica, its relict distribution, and exceptional rarity N. a. soumiafahdae subsp. nov. should be considered as an evolutionarily significant conservation unit that deserves strict protection, continued research to improve the knowledge about its current distribution, and proactive conservation measures (especially prevention of habitat destruction and degradation) to ensure its long-term survival.</p></div>	https://treatment.plazi.org/id/03E587C7FF97CD07FF73F933FE7EF846	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
03E587C7FF8DCD1AFF73FCCCFC02F844.text	03E587C7FF8DCD1AFF73FCCCFC02F844.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. astreptophora (Lopez Seoane 1884)	<div><p>Natrix astreptophora astreptophora (López Seoane, 1884) (n=920)</p><p>FRANCE (n=16, photographic records):</p><p>Pyrénées Orientales (n=9): 3 hatchlings, 6 adults; Aude (n=4): 2 hatchlings, 1 juvenile, 1 adult; Hérault (n=3): 2 hatchlings, 1 adult.</p><p>PORTUGAL (n=146, photographic records):</p><p>Portugal (general, n=4): 1 hatchling, 3 adults; Alentejo (n=4): 1 juvenile, 3 adults ; Algarve (n=5): 1 hatchling, 4 adults ; Aveiro (n=2): 1 juvenile, 1 adult; Braga (n=12): 6 hatchlings, 1 juvenile, 5 adults; Castelo Branco (n=2): 1 hatchling, 1 adult; Coimbra (n=11): 3 hatchlings, 1 juvenile, 7 adults; Covilhã (n=1): 1 adult ; Évora (n=4): 1 hatchling, 1 juvenile, 2 adults; Guarda (n=1): 1 adult; Leiria (n=9): 3 hatchlings, 1 juvenile, 5 adults; Lisboa (n=24): 5 hatchlings, 19 adults; Peneda Gerês (n=14): 2 hatchlings, 12 adults ; Portalegre (n=4): 4 adults; Porto (n=39): 8 hatchlings, 4 juveniles, 27 adults; Salgueiro (n=1): 1 hatchling; Santa Maria da Feira (n=3): 2 hatchlings, 1 adult ; Santarém (n=1): 1 adult; Serra da Estrela (n=1): 1 adult ; Vila Real (n=1): 1 adult; Viana do Castelo (n=3): 2 hatchlings, 1 adult .</p><p>SPAIN (n=758, photographic records including 2 syntypes of Tropidonotus natrix var. astreptophorus López Seoane, 1884 in the Muséum national d’Histoire naturelle, Paris) :</p><p>Andalusia (n=199): 51 hatchlings, 14 juveniles, 134 adults; Aragón (n=26): 11 hatchlings, 1 juvenile, 14 adults; Asturias (n=94): 21 hatchlings, 2 juveniles, 71 adults; Basque Country (n=48): 19 hatchlings, 2 juveniles, 27 adults; Cantabria (n=46): 16 hatchlings, 4 juveniles, 26 adults; Castilla La Mancha (n=14): 2 hatchlings, 2 juveniles, 10 adults ; Castilla León (n=58): 12 hatchlings, 8 juveniles, 38 adults ; Catalonia (n=53): 20 hatchlings, 1 juvenile, 32 adults; Comunidad Valenciana (n=21): 7 hatchlings, 2 juveniles, 12 adults; Extremadura (n=14): 5 hatchlings, 2 juveniles, 7 adults; Galicia (n=156): 50 hatchlings, 18 juveniles, 88 adults; La Rioja (n=4): 4 adults; Madrid (n=16): 5 hatchlings, 11 adults; Murcia (n=3): 1 hatchling, 2 adults; Navarra (n=6): 2 hatchlings, 4 adults .</p><p>Main sources:</p><p>Unpublished photographs by the authors (Andalusia, Cantabria, Castilla León, Madrid, Navarra, and Portugal) and the following colleagues: José Agulló (Burgos), Javier Álvarez (La Rioja), Eduardo Fernández Meléndez (Málaga), Alberto González Estremera (Madrid, Porto), Ander Izagirre Egaña (Basque Country), Alex Mondéjar (Burgos), Jordi Ribo (Catalonia), and Rafael Vázquez Graña (Galicia, northern Portugal). Internet resources [retrieved by May 2025]:Asociación Herpetológica Timon (https://www.asociaciontimon.org), Atlas Herpetológico de Andalucía (http://anfibios-reptiles-andalucia.org), Biodiversidad Virtual (https://www.biodiversidadvirtual.org), Facebook group “Serpientes ibéricas”, iNaturalist (https://www.inaturalist.org).</p></div>	https://treatment.plazi.org/id/03E587C7FF8DCD1AFF73FCCCFC02F844	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
03E587C7FF8ACD1DFF73FF3CFCB2FB17.text	03E587C7FF8ACD1DFF73FF3CFCB2FB17.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. algerica (Hecht 1930)	<div><p>Natrix astreptophora algerica (Hecht, 1930) (n=16)</p><p>ALGERIA (n=6):</p><p>(1) BEV. T8085, hatchling, photographic record, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=6.4319&amp;materialsCitation.latitude=36.8746" title="Search Plazi for locations around (long 6.4319/lat 36.8746)">Aïn Kechra</a>, pond 5 km S Aïn Lemsid (36.8746, 6.4319), Olivier Peyre, Menad Beddek &amp; Boualem Dellaoui ;</p><p>(2) ZMB 1935, juvenile (possibly male), physical voucher specimen, paratype of Tropidonotus natrix algericus Hecht, 1930, “ Algier ” [=Algiers, 36.7538, 3.0588 or province of Algiers], leg. Wilhelm Martin Ernst Ludwig Graf von Schlieffen , c. 1855;</p><p>(3) hatchling, photographic record (Figs 6B, 7D–F), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=4.1258&amp;materialsCitation.latitude=36.4494" title="Search Plazi for locations around (long 4.1258/lat 36.4494)">Djurdjura mountains</a>, Tikjda station (36.4494, 4.1258), Olivier Peyre ;</p><p>(4) adult (probably female), photographic record (Fig. 6D), El Milia (36.6934, 6.3139), Krikra Oussama, Facebook group “Al-Zawahif Mag”;</p><p>(5) hatchling, photographic record, Lake Tonga, E El Kala (36.8814, 8.4981), https://www.inaturalist.org/ observations/265560908;</p><p>(6) ZMB 19636, juvenile (possibly male), physical voucher specimen (Fig. 8A), holotype of Tropidonotus natrix algericus Hecht, 1930, “Südalgier”, leg. Chatschatur Schaposchnikoff, May 1905 .</p><p>TUNISIA (n=10):</p><p>(7, 8) Juvenile and adult female, photographic records with measurements, Beni M’Tir (36.7306, 8.7052), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=8.7052&amp;materialsCitation.latitude=36.7306" title="Search Plazi for locations around (long 8.7052/lat 36.7306)">Gabriel Martínez del Mármol</a>, 26 May 2025 ;</p><p>(9) ZFMK 67196, hatchling, physical voucher specimen, Cap Serrat [37.2100, 9.2200], leg. Hemmo Nickel, 16 October 1997;</p><p>(10) adult, photographic record (Fig. 6C), Cap Serrat (37.2189, 9.2226), Martina Kiselová;</p><p>(11) hatchling, photographic record, Ghorra [36.5900, 8.3700], Mohsen Kalboussi, https://reptile-database. reptarium.cz;</p><p>(12) BEV.8874, adult, physical voucher specimen (head only), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=9.65&amp;materialsCitation.latitude=37.14" title="Search Plazi for locations around (long 9.65/lat 37.14)">Lake Ichkeul</a> [37.1400, 9.6500], unknown collector, 12 October 1979 ;</p><p>(13, 14) hatchling and young adult, photographic records, Nefza [36.9700, 9.0700], Ridha Ouni;</p><p>(15) hatchling, photographic record (Fig. 6A), Tabarka [36.9400, 8.7400], Mohsen Kalboussi, https://reptile-database.reptarium.cz;</p><p>(16) juvenile, photographic record in Brito et al. (2008: fig. 4), referring to one of the three northern Tunisian records in this publication (see Appendix 2), Tony Herrera.</p></div>	https://treatment.plazi.org/id/03E587C7FF8ACD1DFF73FF3CFCB2FB17	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
03E587C7FF8ACD1CFF73FB58FD34FE00.text	03E587C7FF8ACD1CFF73FB58FD34FE00.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. soumiafahdae Mármol & Geniez & Neumann & Glaw & Fritz 2025	<div><p>Natrix astreptophora soumiafahdae subsp. nov. (n=16)</p><p>MOROCCO (n=16):</p><p>Rif Mountains (n=12):</p><p>(1) Adult eating a Sclerophrys mauritanica, photographic record without exact locality, head of the snake not visible, Ahmed Idrissi (Martínez del Mármol et al. 2019: fig. 714; http://www.moroccoherps.com/ficha/natrix_ astreptophora/);</p><p>(2) adult, photographic record, Akchour [35.2200, -5.1900], “Khalid Chino” (photo received from Abdellah Bouazza);</p><p>(3) young adult, photographic record (Fig. 10A), Chefchaouen (35.1441, -5.2627), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.2627&amp;materialsCitation.latitude=35.1441" title="Search Plazi for locations around (long -5.2627/lat 35.1441)">Gabriel Martínez del Mármol</a> (Martínez del Mármol et al. 2019: fig. 715) ;</p><p>(4) young adult, photographic record, Chefchaouen [35.1600, -5.2600], <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.26&amp;materialsCitation.latitude=35.16" title="Search Plazi for locations around (long -5.26/lat 35.16)">Daniel Escoriza</a> (Martínez del Mármol et al. 2019: fig. 709) ;</p><p>(5, 6) hatchling and adult, photographic records, “Jebala” or “Djebala”, Daniel Escoriza (Martínez del Mármol et al. 2019: fig. 712; http://www.moroccoherps.com/ficha/natrix_astreptophora/);</p><p>(7) collection Salvador Carranza, juvenile female, physical voucher specimen, Moulay Abdeslam (35.2701, - 5.4881), Salvador Carranza (Kindler et al. 2018: fig. 1 and figure in Supplementary Information);</p><p>(8–10) hatchling, juvenile (head not visible), young adult male, photographic records, Moulay Abdeslam (35.2701, -5.4881), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.4881&amp;materialsCitation.latitude=35.2701" title="Search Plazi for locations around (long -5.4881/lat 35.2701)">Gabriel Martínez del Mármol</a> (hatchling: Fig. 7G–I) ;</p><p>(11) juvenile, photographic record (Fig. 1), between Moulay Abdeslam and Bouhachem National Park (35.2665, -5.4315), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.4315&amp;materialsCitation.latitude=35.2665" title="Search Plazi for locations around (long -5.4315/lat 35.2665)">Gabriel Martínez del Mármol</a> (Martínez del Mármol et al. 2019: fig. 710) ;</p><p>(12) adult female, photographic record (Fig. 10B, C), Souk El Arbaa [35.3100, -5.3500], <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.35&amp;materialsCitation.latitude=35.31" title="Search Plazi for locations around (long -5.35/lat 35.31)">Mario Schweiger</a> (Martínez del Mármol et al. 2019: figs 711, 713; http://www.moroccoherps.com/ficha/natrix_astreptophora/).</p><p>Middle Atlas (n=4):</p><p>(13) NMW 32908, young adult (male?), physical voucher specimen (Fig. 8B), holotype of Natrix astreptophora soumiafahdae subsp. nov., don. Erich Sochurek, 6 June 1982 ;</p><p>(14) adult (probably female), photographic record, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.21&amp;materialsCitation.latitude=33.43" title="Search Plazi for locations around (long -5.21/lat 33.43)">Azrou forest</a> [33.4300, -5.2100], Mohamed Amine Lyamani (Facebook group “Al-Zawahif Mag”) ;</p><p>(15, 16) BEV.6431, adult female, physical voucher specimen (Fig. 9B), BEV.6432, adult male, physical voucher specimen (Fig. 9A), paratypes of Natrix astreptophora soumiafahdae subsp. nov., stream W Oulmès (33.4412, - 6.0394), 1,069 m a.s.l., leg. Jean Garzoni 1964.</p></div>	https://treatment.plazi.org/id/03E587C7FF8ACD1CFF73FB58FD34FE00	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
03E587C7FF8BCD1FFF73FC93FC72FBE4.text	03E587C7FF8BCD1FFF73FC93FC72FBE4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. algerica (Hecht 1930)	<div><p>Natrix astreptophora algerica (n=29)</p><p>ALGERIA (n=13):</p><p>(1) Anonymous locality (36.8500, 4.3300, coordinates most likely approximately), literature record (Escoriza &amp; Ben Hassine 2017: Supplementary Material);</p><p>(2) anonymous locality (36.8500, 8.5300, coordinates most likely approximately), literature record (Escoriza &amp; Ben Hassine 2017: Supplementary Material);</p><p>(3) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=6.4319&amp;materialsCitation.latitude=36.8746" title="Search Plazi for locations around (long 6.4319/lat 36.8746)">Aïn Kechra</a>, pond 5 km S Aïn Lemsid (36.8746, 6.4319), physical voucher specimen (BEV. T8085) , hatchling; referring to same specimen: photographic record, Olivier Peyre, Menad Beddek &amp; Boualem Dellaoui and genetic record (Kindler et al. 2018: S1, BEV. T8085);</p><p>(4) Akfadou [36.6304, 4.6245], genetic record (Kindler et al. 2018: S1, CIBIO 868);</p><p>(5) Algiers or “Algier” [36.7538, 3.0588 or province of Algiers], physical voucher specimen (ZMB 1935), paratype of Tropidonotus natrix algericus Hecht, 1930, juvenile (possibly male), leg. Wilhelm Martin Ernst Ludwig Graf von Schlieffen, c. 1855 and literature records (Gervais 1848; Strauch 1862; Lallement 1866; Boulenger 1891: F. Lataste, 1880–1884) ;</p><p>(6) “ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=7.7469&amp;materialsCitation.latitude=36.8925" title="Search Plazi for locations around (long 7.7469/lat 36.8925)">Bône</a> ” [= Annaba, 36.8925, 7.7469], literature record (Doumergue 1901) ;</p><p>(7) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=4.1258&amp;materialsCitation.latitude=36.4494" title="Search Plazi for locations around (long 4.1258/lat 36.4494)">Djurdjura mountains</a>, Tikjda station (36.4494, 4.1258), photographic record, hatchling, Olivier Peyre ;</p><p>(8) La Chiffa [=Chiffa, 36.3999, 2.7667], literature records (Boulenger 1891: F. Lataste, 1880–1884; Doumergue 1901);</p><p>(9) SE El Milia (36.6934, 6.3139), photographic record, adult, Krikra Oussama, Facebook group “Al-Zawahif Mag”;</p><p>(10) Lake Tonga, E El Kala (36.8814, 8.4981), photographic record, hatchling, Telailia Salah, 16 March 2025, https://www.inaturalist.org/observations/265560908;</p><p>(11) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=7.6392&amp;materialsCitation.latitude=36.8825" title="Search Plazi for locations around (long 7.6392/lat 36.8825)">Mt. Edough</a> [=Djebel Edough, 36.8825, 7.6392], near Bona [= Annaba], literature records (Boulenger 1891: Dr. Hagenmüller; Olivier 1894) ;</p><p>(12) Tifret [=Tifrit N’Aït El Had, 36.7960, 4.4212], literature record (Boulenger 1891: F. Lataste, 1880–1884);</p><p>(13) “Südalgier”, physical voucher specimen (ZMB 19636), holotype of Tropidonotus natrix algericus Hecht, 1930, juvenile (possibly male), leg. Chatschatur Schaposchnikoff, May 1905 .</p><p>TUNISIA (n=16):</p><p>(1) Anonymous locality (37.1100, 9.7000, coordinates most likely approximately), literature record (Escoriza &amp; Ben Hassine 2017: Supplementary Material);</p><p>(2) anonymous locality (36.7400, 8.7100, coordinates most likely approximately), literature record (Escoriza &amp; Ben Hassine 2017: Supplementary Material);</p><p>(3) anonymous locality (36.9700, 8.8400, coordinates most likely approximately), literature record (Escoriza &amp; Ben Hassine 2017: Supplementary Material);</p><p>(4) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=8.7052&amp;materialsCitation.latitude=36.7306" title="Search Plazi for locations around (long 8.7052/lat 36.7306)">Beni M’Tir</a> (36.7306, 8.7052), photographic records, juvenile and adult female, Gabriel Martínez del Mármol, 26 May 2025 ;</p><p>(5) 1 km E from crossing to <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=8.6929&amp;materialsCitation.latitude=36.723" title="Search Plazi for locations around (long 8.6929/lat 36.723)">Beni M’Tir</a> (36.7230, 8.6929, 656 m a.s.l.), literature record (Brito et al. 2008), probably identical with genetic record of Kindler et al. (2018: S1, CIBIO 1561; 5 km W Beni M’Tir) ;</p><p>(6) Cap Serrat (37.2189, 9.2226), adult, photographic record, Martina Kiselová, same individual as iNaturalist record from Sejenane, Václav John, 2 July 2017, https://www.inaturalist.org/observations/144506762;</p><p>(7) Cap Serrat [37.2100, 9.2200], physical voucher specimen (ZFMK 67196), hatchling, leg. Hemmo Nickel, 16 October 1997, identical with genetic record of Kindler et al. (2018: S1, ZFMK 67196);</p><p>(8) 9 km S <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=9.0524&amp;materialsCitation.latitude=37.0497" title="Search Plazi for locations around (long 9.0524/lat 37.0497)">Cape Negro</a> (37.0497, 9.0524, 60 m a.s.l.), literature record (Brito et al. 2008), probably identical with genetic record of Kindler et al. (2018: S1, CIBIO 1582 for the same locality) ;</p><p>(9) Gafsa “ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=8.7757&amp;materialsCitation.latitude=34.4311" title="Search Plazi for locations around (long 8.7757/lat 34.4311)">O. Lella</a> (Gafsa)” [34.4311, 8.7757], physical voucher specimen (MNHN-RA-1983.507), collector unknown, 1982; this doubtful record is outside the known distribution range and therefore not included in Figure 3 (see above) ;</p><p>(10) Ghorra [36.5900, 8.3700], photographic record, hatchling, Mohsen Kalboussi, https://reptile-database. reptarium.cz;</p><p>(11) Kroumirie [= Khroumirie, Khroumérie, 36.5833, 8.4167], physical voucher specimen (MNHN-RA-1907.182), collector unknown (H. Gadeau de Kerville?), 1907, also literature records (Gadeau de Kerville 1908; Schleich et al. 1996: 512); the continued presence in the Kroumirie region was recently confirmed (Kalboussi &amp; Achour 2018);</p><p>(12) Lake Ichkeul [37.1400, 9.6500], unknown collector, physical voucher specimen (BEV. 8874, head only), adult, collector unknown, 12 October 1979;</p><p>(13) “Mogods” [37.1333, 9.5000], literature record (Schleich et al. 1996: 512);</p><p>(14) Nefza [36.9700, 9.0700], photographic records, young adult and hatchling, Ridha Ouni;</p><p>(15) Tabarka [36.9400, 8.7400], photographic record, hatchling, Mohsen Kalboussi, https://reptile-database. reptarium.cz;</p><p>(16) 10 km E <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=8.9353&amp;materialsCitation.latitude=36.9575" title="Search Plazi for locations around (long 8.9353/lat 36.9575)">Tabarka</a> (36.9575, 8.9353, 100 m a.s.l.), literature record (Brito et al. 2008, probably identical with genetic record of Kindler et al. (2018: S1, CIBIO 153) for the same locality.</p></div>	https://treatment.plazi.org/id/03E587C7FF8BCD1FFF73FC93FC72FBE4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
03E587C7FF88CD11FF73FBC8FE28FE90.text	03E587C7FF88CD11FF73FBC8FE28FE90.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Natrix astreptophora subsp. soumiafahdae Mármol & Geniez & Neumann & Glaw & Fritz 2025	<div><p>Natrix astreptophora soumiafahdae subsp. nov.</p><p>MOROCCO (n=55):</p><p>Atlantic plains (n=6):</p><p>(1) Anonymous locality [34.2200, -6.6800], literature record (Mediani et al. 2015), possibly identical with record 4 below;</p><p>(2) anonymous locality [35.1800, -6.1700], literature record (Mediani et al. 2015), possibly identical with record 5 below;</p><p>(3) El Khemis du Sahel (Khemis Sahel) [35.2530, -6.0830], physical voucher specimen (lost, originally in <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-6.083&amp;materialsCitation.latitude=35.253" title="Search Plazi for locations around (long -6.083/lat 35.253)">Estación Biológica de Doñana</a>, 20 July 1954) ;</p><p>(4) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-6.66&amp;materialsCitation.latitude=34.25" title="Search Plazi for locations around (long -6.66/lat 34.25)">Lake Sidi Boughaba</a>, Kenitra [34.2500, -6.6600, 45 m a.s.l.], literature records (Escoriza et al. 2011: September 2010; Escoriza &amp; Ben Hassine 2017: Supplementary Material), possibly identical with record 1 above ;</p><p>(5) mouth of Oued Loukos [35.2023, -6.1454], literature record (Mateo et al. 2003: 232), possibly identical with record 2 above;</p><p>(6) Sidi Sibara [33.4093, -6.7439], two physical voucher specimens (USNM 196481, 524232, see http://www. vertnet.org), collector unknown, 1969 .</p><p>Rif Mountains (n=39):</p><p>(7) Anonymous locality [35.3500, -5.5600], literature record (Escoriza &amp; Ben Hassine 2017: Supplementary Material), possibly identical with record 37 below;</p><p>(8) anonymous locality [35.3400, -5.5100], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(9) anonymous locality [35.2600, -5.5100], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(10) anonymous locality [35.0800, -5.1600], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(11) anonymous locality [35.0000, -4.9400], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(12) anonymous locality [35.0000, -4.7200], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(13) anonymous locality [34.9900, -4.6100], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(14) anonymous locality [34.9100, -4.6100], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(15) anonymous locality [35.6400, -5.4600], literature record (Mediani et al. 2015);</p><p>(16) anonymous locality [35.2900, -5.4900], literature record (Mediani et al. 2015);</p><p>(17) anonymous locality [35.2300, -5.4400], literature record (Mediani et al. 2015);</p><p>(18) anonymous locality [35.2000, -5.1000], literature record (Mediani et al. 2015);</p><p>(19) anonymous locality [35.1500, -4.9900], literature record (Mediani et al. 2015);</p><p>(20) anonymous locality [35.0100, -5.2100], literature record (Mediani et al. 2015);</p><p>(21) anonymous locality [35.1500, -4.9400], literature record (Mediani et al. 2015);</p><p>(22) anonymous locality [35.0700, -4.8300], literature record (Mediani et al. 2015);</p><p>(23) anonymous locality [35.0300, -4.7200], literature record (Mediani et al. 2015);</p><p>(24) anonymous locality [35.0700, -4.6600], literature record (Mediani et al. 2015);</p><p>(25) anonymous locality [34.8900, -4.6000], literature record (Mediani et al. 2015);</p><p>(26) anonymous locality [34.9800, -4.3900], literature record (Mediani et al. 2015);</p><p>(27) anonymous locality, fountain in western Rif Mountains [35.3200, -5.5300, 1,175 m a.s.l.], literature record (Escoriza et al. 2011: November 2008), possibly identical with genetic record 33 below;</p><p>(28) anonymous locality, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.35&amp;materialsCitation.latitude=35.73" title="Search Plazi for locations around (long -5.35/lat 35.73)">Rif Mountains</a> / Mediterranean coast [35.7300, -5.3500], literature record (Mediani et al. 2015) ;</p><p>(29) Akchour [35.2200, -5.1900], photographic record, adult, “Khalid Chino” (photo received from Abdellah Bouazza);</p><p>(30) S Bab <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.26&amp;materialsCitation.latitude=35.16" title="Search Plazi for locations around (long -5.26/lat 35.16)">Taza</a> (35.0225, -5.2060, 1,200 m a.s.l.), observation Soumia Fahd (pers. comm. May 2024);</p><p>(31) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.26&amp;materialsCitation.latitude=35.16" title="Search Plazi for locations around (long -5.26/lat 35.16)">Chefchaouen</a> (35.1441, -5.2627), photographic record, young adult, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.26&amp;materialsCitation.latitude=35.16" title="Search Plazi for locations around (long -5.26/lat 35.16)">Gabriel Martínez del Mármol</a>;</p><p>(32) Chefchaouen [35.1600, -5.2600], photographic record, young adult, Daniel Escoriza;</p><p>(33) 30 km NW of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.5247&amp;materialsCitation.latitude=35.356" title="Search Plazi for locations around (long -5.5247/lat 35.356)">Chefchaouen</a> [35.3560, -5.5247], genetic record (Kindler et al. 2018: S1, ENAT1 M), possibly identical with literature record 27 above ;</p><p>(34) 1 km SW <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.2957&amp;materialsCitation.latitude=35.1064" title="Search Plazi for locations around (long -5.2957/lat 35.1064)">Derdara</a> [35.1064, -5.2957], genetic record (Kindler et al. 2018: S1, E21023 x8M) ;</p><p>(35) J[e]bel Bouhalla [35.1700, -5.1600], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(36) J[e]bel Tazaot [35.2600, -5.1700], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(37) Jem A Allah near Targuist [34.9900, -4.3900], literature record (Fahd &amp; Pleguezuelos 2001);</p><p>(38) 6 km SE <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.61&amp;materialsCitation.latitude=35.41" title="Search Plazi for locations around (long -5.61/lat 35.41)">Koudia El Mal</a> [35.4100, -5.6100], genetic record (Kindler et al. 2018: S1, e10127x1M), possibly identical with record 7 above ;</p><p>(39) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.4881&amp;materialsCitation.latitude=35.2701" title="Search Plazi for locations around (long -5.4881/lat 35.2701)">Moulay Abdeslam</a> (35.2701, -5.4881), several photographic records, Gabriel Martínez del Mármol and one physical voucher specimen, juvenile female, collection of Salvador Carranza ;</p><p>(40) between Moulay Abdeslam and Bouhachem National Park (35.2665, -5.4315), photographic record, juvenile, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.4315&amp;materialsCitation.latitude=35.2665" title="Search Plazi for locations around (long -5.4315/lat 35.2665)">Gabriel Martínez del Mármol</a>;</p><p>(41) Souk El Arbaa [35.3100, -5.3500], photographic record, adult female, Mario Schweiger;</p><p>(42) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.51&amp;materialsCitation.latitude=35.69" title="Search Plazi for locations around (long -5.51/lat 35.69)">Souk Khemiss Anjra</a> (= Souk Khémis des Anjra, Soq Khmis Anjra) [35.6900, -5.5100], literature record (Fahd &amp; Pleguezuelos 2001) ;</p><p>(43) Tamorot (Beni Halet) [34.9376, -4.7793], literature records (Galán 1931: one individual, 15 June 1930; Fahd &amp; Pleguezuelos 2001);</p><p>(44) northern slope of Yebel Haus (=Jebel Haouz) [35.6995, -5.4586], literature record (Mateo et al. 2003: 232, two individuals);</p><p>(45) Yebel Tazaot (=Jebel Tasaot) [35.2236, -5.0669], literature record (Mateo et al. 2003: 232, one individual).</p><p>Middle Atlas (n=10):</p><p>(46) <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-5.21&amp;materialsCitation.latitude=33.43" title="Search Plazi for locations around (long -5.21/lat 33.43)">Azrou forest</a> [33.4300, -5.2100], adult (probably female), Mohamed Amine Lyamani, Facebook group “Al-Zawahif Mag” ;</p><p>(47) Dayet Aoua [33.6545, -5.0338], observation Uwe Sattler (pers. comm. early 1990s);</p><p>(48) Ifrane [33.5228, -5.1110], physical voucher specimens (MNHN-RA-1961.318, 1961.319), J. Bons &amp; H. Saint Girons, 1958;</p><p>(49) J[e]bel Tazekka [34.0903, -4.1836], genetic record (Kindler et al. 2018: S1, CIBIO 10510);</p><p>(50) Naour [32.4803, -5.9086], literature record (Schleich et al. 1996: 512);</p><p>(51) stream W Oulmès (33.4412, -6.0394), 1,069 m a.s.l., physical voucher specimens (BEV. 6431, 6432), paratypes of Natrix astreptophora soumiafahdae subsp. nov., adult female and adult male, leg. Jean Garzoni 1964;</p><p>(52) Oulmès (<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-6.1499&amp;materialsCitation.latitude=32.5101" title="Search Plazi for locations around (long -6.1499/lat 32.5101)">Tarmilète</a> = Oulmès, Tarmilate) [33.4171, -6.0511], literature record (Schleich et al. 1996: 512); (53) Rhorm el Alem [32.5101, -6.1499], literature record (Schleich et al. 1996: 512);</p><p>(54) Tanout Ou Filal pass (Tizi Tanout Ou Filal) (32.6767, -5.4550), 1,960 m a.s.l., historical record by Jörg Wittenberg, from database of Bons &amp; Geniez (1996);</p><p>(55) 18 km S <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-3.9911&amp;materialsCitation.latitude=34.0461" title="Search Plazi for locations around (long -3.9911/lat 34.0461)">Taza</a> [34.0461, -3.9911], physical voucher specimen (USNM 196480, see http://www.vertnet.org), collector unknown, 1970 .</p></div>	https://treatment.plazi.org/id/03E587C7FF88CD11FF73FBC8FE28FE90	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Mármol, Gabriel Martínez Del;Geniez, Philippe;Neumann, Adrian;Glaw, Frank;Fritz, Uwe	Mármol, Gabriel Martínez Del, Geniez, Philippe, Neumann, Adrian, Glaw, Frank, Fritz, Uwe (2025): Phenotypic variability in the western grass snake Natrix astreptophora, with description of a new subspecies from Morocco. Zootaxa 5693 (1): 41-65, DOI: 10.11646/zootaxa.5693.1.3, URL: https://doi.org/10.11646/zootaxa.5693.1.3
