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E. Phrynocephalus Size Diversity Phrynocephalus show a variety of snout vent lengths and tail lengths, (Table 6, Appendix 5). Body size per sample ranged from 106 mm for P. mystaceus to 35 mm for P. vindumi and P. interscapularis. Tail length per sample ranges from 133 mm for P. maculatus population 1 to 35 mm for P. luteoguttatus population 2. P. mystaceus and P. interscapularis are sympatric in the Caspian Basin, and habitat partition with P. mystaceus in large floating sand dunes, with P. interscapularis living on the edge of those large sand dunes, which may revolve around ecotype evolution in a predator-prey relationship. In northwestern Iran P. vindumi replaces the role of P. interscapularis, and is sympatric with P. mystaceus. Tail length appears to be the most sexually dimorphic characteristic. ......continued on the next page Snout vent lengths for clay ecotypes range from 62 mm for P. przewalskii population 1 to 49 mm for P. versicolor, with tail lengths ranging from 90 mm for P. przewalskii population 1 to 49 mm for P. persicus. The snout vent length for gravel ecotypes range from 64 mm for P. vlangalii population 1 to 40 mm for P. rossikowi, with tail lengths ranging from 90 mm for P. longicaudatus to 42 mm for P. theobaldi. Snout vent lengths for dry lakebed ecotypes range from 90 mm for P. maculatus population 1 to 60 mm for P. golubewii, with tail length ranging from 133 mm for P. maculatus population 1 to 78 mm for P. maculatus population 2. Snout vent lengths for small sand dune ecotypes range from 61 mm for P. vlangalii population 2 to 35 mm for both P. vindumi and P. interscapularis, with tail length ranging from 80 mm for P. przewalskii population 2 to 35 mm for P. luteoguttatus population 2. Snout vent lengths for large sand dune ecotypes range from 106 mm for P. mystaceus populations to 59 mm for P. salenskyi population 1, with tail lengths ranging from 110 mm for P. mystaceus population 1 to 70 mm for P. salenskyi population 1. The largest Phrynocephalus occupy large floating sand dunes, and the smallest Phrynocephalus species are found on the fringes of large floating sand dunes in the Caspian Basin and northern Iran. P. mystaceus occupying large sand dunes in these habitats coexists with the two smallest species that occupy the fringes of those sand dunes, P. interscapularis in the Caspian Basin, and P. vindumi in northwestern Iran. It has not escaped our attention that size and habitat partitioning may have occurred in these regions because it is likely that smaller species (P. interscapularis or P. vindumi) serve as a food source for larger species (P. mystaceus). Further work is needed for phylogenetic assessment of size evolution among Phrynocephalus species and populations considering the sexually dimorphic nature of this genus, both with body size and tail length.

A. Phrynocephalus Taxonomic Recommendations Based on phylogenetic analyses, a few taxonomic changes are proposed. The following taxonomic changes are based on the principle of monophyly as well as genetic and geographic differences. In each case, the reason a change is suggested is provided, whether data are genetic, distributional or phylogenetic (ie. non-monophyly). 1). Phrynocephalus maculatus Complex as Previously Recognized Based on complete regional sampling, this study recognizes all previous subspecies of P. maculatus as distinct species on the minim-basis of non-monophyly. Phrynocephalus maculatus was described from the Iranian Plateau (Anderson 1872) with the type locality of “Awada, Shiraz, Persia ”; corrected to “Abadeh [31o 10' N, 52o 37' E; Fars Province, Iran], north of Shiraz” by Blanford (1876). Phrynocephalus maculatus longicaudatus was described by Haas (1957), with a type locality of “Doha Dhalum, Saudi Arabia ”. The sole known population of P. maculatus from Turkmenistan was reported by Bogdanov et al. (1974), and subsequently described as a distinct species, Phrynocephalus golubewii with the type locality of “vicinity of the Bami rail-road station” [= 7 km N railway station Bamy], (Shenbrot & Semenov 1990). It is recommended to recognize P. golubewii, P. longicaudatus, and P. maculatus as distinct species based on phylogenetic results: a). While Phrynocephalus golubewii (Shenbrot & Semenov 1990) may look like P. maculatus living in saltbed dry lakes, it is directly-related to the clade containing P. turcomanus and P. helioscopus (mt-DNA, bootstrap 91%, decay index 6; combined data, bootstrap 87%, decay index 7; allozyme presence/absence, bootstrap 78%, decay index 1; allozyme step matrix, bootstrap 75%, decay index 2; non-monophyly in RAG- 1); in fact P. turcomanus occurs within meters of P. golubewii and may overlap in distribution. b). Phrynocephalus maculatus (Anderson 1872) is restricted to salt-bed dry lakes on the Iranian Plateau where its type locality is located, and adjacent Baluchistan Plateau including the base of the Sulaiman Range of southern Pakistan. P. maculatus is related, yet outside and sister to a clade containing P. arabicus and P. longicaudatus (see below) based on both mitochondrial and nuclear RAG-1 DNA evidence (mt-DNA, bootstrap 100%, decay index 32; RAG-1, bootstrap 100%, decay index 13; combined data, bootstrap 100%, decay index 46; all of these taxa were not sampled for allozyme data). c). Phrynocephalus longicaudatus (Haas 1957) New Status is restricted to the Arabian Peninsula. It is the sister taxon to P. arabicus which is also endemic to the Arabian Plate and supported by mitochondrial and nuclear RAG- 1 DNA evidence (mt-DNA, bootstrap 100%, decay index 22; RAG- 1, bootstrap 100%, decay index 7; combined data, bootstrap 100%, decay index 30; these taxa were not sampled for allozyme data). These taxa in any combination do not form a monophyletic group. 2). Small Sand Dune Species of Southwest Asia and Caspian Basin Phrynocephalus interscapularis interscapularis (Lichtenstien 1856; type locality “Bucharei” = Uzbekistan) and Phrynocephalus interscapularis sogdianus (Chernov 1948; type locality “ Tadjikistan, vicinity of the Pjandzh village”, approximately 37o 14' N, 69o 05' E) are recognized here as full species P. interscapularis and P. sogdianus based on allozyme and DNA differences. The mt-DNA pair-wise uncorrected sequence distance between P. interscapularis and P. sogdianus is 3.2% and these taxa show one fixed allozyme difference at the carboxylic ester hydrolase locus (EST; Table 2). Phrynocephalus ornatus vindumi [Golubev, 1998; type locality “Iran, Khorasan Prov., 35 km. N of Gonabad on road to Torbat-E. Heydariyeh (ca. 34o 49' N, 58o 47' E), 850 m. elevation”; sampling here within ten kilometers of the type locality, see appendix 1] is elevated to species status as Phrynocephalus vindumi New Status. Phrynocephalus ornatus (Boulenger 1887; type locality “between Nushki and Helmand, Baluchistan”) and P. vindumi appear in a non-monophyletic arrangement as P. vindumi occurring on the northeastern portion of the Iranian Plateau is separated from P. ornatus that occurs south of the Hindu Kush in Afghanistan and southwestern Pakistan. In fact, P. vindumi is related to small sand dune species in the Caspian Basin to the north: P. interscapularis and P. sogdianus. In the mt-DNA parsimony analysis, P. vindumi appears in a sister position to the clade containing P. interscapularis and P. sogdianus, with those three taxa grouping with high support (bootstrap 98%, decay index 13). Alternatively, P. ornatus is sister to P. clarkourum (bootstrap 99%, decay index 21), and those two taxa are included in a clade with the two P. luteogutattus samples that also receive high support (bootstrap 100%, decay index 26). Allozyme and RAG-1 DNA data did not sample appropriately to test this question. 3). Gravel Dwelling Phrynocephalus Species in the Caspian Basin Previously, P. reticulatus, P. bannikovi, and P. strauchi were all considered subspecies of P. reticulatus (Eichwald 1831): The nominal subspecies Phrynocephalus reticulatus reticulatus [Eichwald 1831; type locality "Hab. ad Oxum, pristinum amnem, in caspii maris litore orietali", corrected to the Amu-Darja River by Golubev (1991)] and two additional distinct subspecies, Phrynocephalus reticulatus strauchi [Nikolsky 1899; restricted type locality “Khodzhent, 40o 17' N, 69o 37' E, northern Tajikistan ” by lectotype designated by Dunayev (1995)] and Phrynocephalus reticulatus bannikovi (Darevsky, Rustamov & Shammakov 1976; type locality “Tuarkyr Mountains, northwestern Turkmenistan ”), (see Bannikov et al. 1977; although P. reticulatus and P. strauchi were originally described as species). Here these gravel ecotypes are recognized as distinct species based on genetic difference and lack of monophyly. In our sampling, P. bannikovi and P. strauchi do not appear monophyletic with P. strauchi and P. rossikowi grouping with light support (mt-DNA, bootstrap 55%, decay Index 2). P. reticulatus from central Uzbekistan was not sampled in our study or the study of Solovyeva et al. (2014). If P. bannikovi and P. rossikowi were removed from the parsimony analysis performed on the mt-DNA data, the topology of the trees in this study and that of Solovyeva et al. (2014) would be congruent. Based on incomplete sampling, it is recommended that members of the P. reticulatus complex be recognized as separate species Phrynocephalus bannikovi (Darevsky, Rustamov & Shammakov 1976) New Status, P. reticulatus, and P. strauchi. Solovyeva et al. (2014) appear to have not sampled any species from Turkmenistan, which includes P. raddei raddei and P. rossikowi, and in our phylogenetic analyses are related to the P. reticulatus complex. The inclusion of P. raddei raddei and P. rossikowi in the sampling presented here indicates the traditional P. reticulatus complex is not monophyletic, deserving species recognition for the three former subspecies. In common with this study, Solovyeva et al. (2014) only sampled P. strauchi from the P. reticulatus complex. In the sampling of this study, P. bannikovi and P. raddei are geographically oriented disjunctive-adjacent to each other in eastern and west-central Turkmenistan, respectively, and are sister taxa receiving high support from mt-DNA (bootstrap 100%, decay index 23), nuclear RAG-1 DNA (bootstrap 91%, decay index 3), and combined data analyses (bootstrap 100%, decay index 27). Solovyeva et al. (2014) did not sample P. bannikovi, P. rossikowi, or P. raddei raddei from Turkmenistan. However, Solovyeva et al. (2014) and this study did sample P. strauchi from the Fergan Valley of eastern Uzbekistan or northern Tadjikistan. Sampling between this study and Solovyeva et al. (2014) differs in the subspecies of P. raddei: This study uses P. raddei raddei, (Boettger 1888: type locality “Perewalnaja an der transcaspischen Bahn", = Perevalnaja railroad station, southwestern Turkmenistan; near where this study's sample originated from), whereas Solovyeva et al. (2014) uses P. raddei boettgeri (Nikolsky 1905; type locality “Schirabad, Buchara orient” = Sherabad, approximately 37o 42' N, 67o 04' E, southern Uzbekistan) with a vague locality provided as “Turan Depression” [= Caspian Basin]; but occurs in extreme southern Uzbekistan and southern Tajikistan along the Afghanistan border (Bannikov et al. 1977). Until a single study samples both subspecies of P. raddei, it will be unclear how they relate to P. bannikovi, P. reticulatus, P. strauchi, and P. rossikowi. In Solovyeva et al. (2014) no detailed locality information is available, nor is an appendix with this information provided. All GenBank entries do not include locality data, so this information is not archival with NIH GenBank. The Zoological Museum of Moscow University (ZMMU) (http://zmmu.msu.ru/en/about-muzeum/ fonds/amphibians-&-reptiles-collections) in which specimens reported in Solovyeva et al. (2014) are housed, does not offer an obvious means of searching by voucher number on their website. Additionally, this institution does not currently participate in global databases which compile data from institutions across the globe and allow free public access to museum collections involving voucher number relating to locality information. VertNet, a global organization whose participants include 64 institutions spanning 13 countries including Russia, is one prominent database to organize museum collections around the world. The Global Information Biodiversity Facility (GBIF) also offers a forum for researchers to access information on specimens collected in select countries on every continent, which is an added resource. The Zoological Museum of Moscow University is not currently participating in any of these databases, and localities in the manuscript of Solovyeva et al. (2014) are very vague, crossing large geographic regions, and not informative for the difficulties of dealing with Phrynocephalus taxonomy and sampling. Solovyeva et al. (2014) present GenBank sequences as common courtesy in the scientific community, but in those sequences, no locality information is provided so comparing Solovyeva et al. (2014) results with this study is difficult across all issues, starting with taxonomy. Published localities in Solovyeva et al. (2014) such as Central and Middle Asia are not informative, as in the Russian style geography, Middle Asia refers to the Caspian Basin, and Central Asia refers to Tibet, Mongolia and western China. Localities of such gross generality do not provide any information for a study of this detail. 4). Phrynocephalus helioscopus Complex in the Caspian Basin Phrynocephalus helioscopus turcomanus [Dunayev, Solovyeva & Poyarkov 2012 ( in Solovyeva et al. 2012); type locality 40° 01′ 00′′ N 52° 58′ 00′′ E, current name Turkmenbashi, Turkmenistan] is recognized as a distinct species Phrynocephalus turcomanus [Dunayev, Solovyeva & Poyarkov 2012 ( in Solovyeva et al. 2012)] New Status from P. helioscopus based on a north-south distributional disjunction in the Caspian Basin with significant mt-DNA differences. P. helioscopus [Pallas (1771) listed the type locality as "in deserti australioris collibus ardentissimis"; restricted (with no lectotype) to the "Inderskija Gory, Gebiet des unteren Uralflusses" by Mertens & Müller (1928), now Inderskije (Inder) Mountains, Atyrau Region, northwestern Kazakhstan] here is sampled from the northern Aral Sea region of Kazakhstan, and P. turcomanus from southwestern Turkmenistan (Appendix 1). While no fixed allozyme differences are detected between populations sampled, there is a high mt-DNA difference. The mt-DNA pairwise uncorrected sequence distance between P. helioscopus and P. turcomanus is 4.5%; therefore significantly diverged and in the range of sister species divergence for this segment of the mitochondrial genome among amphibians and reptiles (for summary see table 6 in Weisrock et al. 2001). 5). Use of Names in the Low Elevation Chinese Desert—Northern Caspian Basin Clade a). Phrynocephalus salenskyi The name P. salenskyi Bedriaga 1907 is applied here for populations sampled from the Junggur Depression of northwestern China, and the Zaysan Depression of northeastern Kazakhstan. P. salenskyi is the oldest name available by date and page number for Phrynocephalus populations of the low elevation Chinese desert – northern Caspian Basin clade in this geographic region, see below: Phrynocephalus Salenskyi Bedriaga 1907: 141, 213.—Restricted type locality: “ Urungu Fluss ” [= Ulungur River, Xinjiang Uygur Autonomous Region, China]. Phrynocephalus Isseli Bedriaga 1907: 229.— Type locality: Desert in vicinity of Mt. Ssalburty (in Saur Mountains of the Tarbagatay Range, Xinjiang Uygur Autonomous Region, China). Phrynocephalus Haeckeli Bedriaga 1907: 237.— Type locality: Desert in vicinity of Mt. Ssalburty (in Saur Mountains of the Tarbagatay Range), in the Tarbagataï (= Tacheng County) district, northern Xinjiang Uygur Autonomous Region, China. Phrynocephalus Arcellazzii Bedriaga 1907: 307.— Type locality: “ Gaschun, in der Nähe von Gutschen in der Dschungarei ” [= Qitai (44° 00' N, 89° 50' E), Junggar Depression, Xinjiang Uygur Autonomous Region, China]. Phrynocephalus Grum-Grzimailoi Bedriaga 1909: 420.—Restricted type locality: “Gutschen” [= Qitai (44° 00' N, 89° 50' E), Junggar Depression, Xinjiang Uygur Autonomous Region, China]. Phrynocephalus bedriaga Tsarevsky 1926: 214.—Restricted type locality: Zaissan Lake region, eastern Kazakhstan. Phrynocephalus albolineatus K. Zhao 1979: 113.— Type locality: "45 kilometers south of Tacheng County, Xinjiang Uygur Autonomous Region, China; 584 m elevation". The Irtysh River flows out of Zaysan Lake linking the Junggar Depression with the Zaysan Depression, which makes a compelling case for sampling presented in this study representing P. salenskyi populations. b). Phrynocephalus przewalskii Complex in the Gobi Desert Phrynocephalus Przewalskii Strauch 1876: 10.— Type locality: “deserto Alaschanico ” [= Tengger Desert, Nei Mongol Autonomous Region, China]; corrected to eastern Tengger Desert west from 106° E, southwest Nei Mongol Autonomous Region, China in Barabanov & Ananjeva (2007). This taxon is generally a sand dwelling ecotype. P. frontalis Strauch 1876: 15.—Type locality: locality “Ordos dicta” [= Ordos Desert, Nei Mongol Autonomous Region, China]. This taxon is generally thought of as a gravel dwelling ecotype, and here considered a synonym of P. przewalskii (see below). A lack of genetic diversity between populations from these taxa (two populations each), confirm that only the name P. przewalskii is valid, whereas P. frontalis is a synonym of P. przewalskii (following Gozdzik & Fu 2009). This is supported by nuclear encoded allozyme data, and mt-DNA data. Alternative names exist as reviewed in Barabanov & Ananjeva (2007), but without detailed sampling of type localities, the validity of alternate names cannot be addressed. Across the 25 allozyme loci sampled here, we detected no fixed differences among the four populations of P. przewalskii, indicating a fair amount of gene flow between the populations. The average mt-DNA pair-wise uncorrected sequence distances between population 2 and the clade containing populations 1, 3, and 4 is 3.64% (range 3.41–3.79%). The average pair-wise comparison within the clade containing populations 1, 3 and 4 is 0.57% (range 0.51–0.63%). These results match the phylogenetic topology discovered in the mt-DNA parsimony analysis, with population 2 placed in a sister position to the clade containing populations 1, 3 and 4 (bootstrap 100%, decay index 19). No resolution was discovered among the four P. przewalskii populations in the RAG- 1 DNA parsimony analysis. Urquhart et al. (2009) suggests a vicariant break among P. przewalskii populations roughly corresponding to the Helan Shan (mountains). Data reported here do not corroborate that hypothesis. Sampled are a single population east of the Yellow River and three populations west of the Yellow River, which flows north to south in this region. Population 2 is sampled on the east side of the river, and population 3 is sampled on the west side of the river, on banks directly across from each other in Shapotou, Ningxia Autonomous Region, China. Population 2, the only population on the east side of the Yellow River is the most divergent, with populations 1, 3 and 4 on the west side of the river nearly identical in our sampling. Population 1 (Wujiachuan, Gansu Province, China) is west of the Yellow River and further south than sampling conducted in Urquhart et al. (2009) and Wang & Fu (2004), yet nearly identical to other populations sampled west of the Yellow River. In our sampling, only population 4 lies west of the Helan Shan (Wuwei, Gansu Province, China) in the Tengger Desert, the type locality of P. przewalskii. Sampling in this study suggests the Yellow River could be responsible for a genetic break in P. przewalskii populations, and not the Helan Shan, though more detailed, fine-scale sampling and analysis will need to be done to confirm this hypothesis. In the Zhongwei region of Ningxia Autonomous Region, there is an indication that east and west haplotypes as described above occur in the same locality, which could indicate a point of hybridization, but further detailed work is needed (Wang & Fu 2004). Sampling conducted in similar areas in Wang & Fu (2004) and Urquhart et al. (2009) did not completely overlap, making conclusions between these studies and this study very difficult for comparative interpretation, further indicating additional work is needed. In sampling of the P. przewalskii complex in Jin & Brown (2013), populations to the east are grouped, and are referred to as P. frontalis, with populations to the west grouping and referred to as P. przewalskii, not conflicting with results reported here. 6). Use of Names in the Northern Tibetan Plateau

A. Phrynocephalus Taxonomic Recommendations Based on phylogenetic analyses, a few taxonomic changes are proposed. The following taxonomic changes are based on the principle of monophyly as well as genetic and geographic differences. In each case, the reason a change is suggested is provided, whether data are genetic, distributional or phylogenetic (ie. non-monophyly). 1). Phrynocephalus maculatus Complex as Previously Recognized Based on complete regional sampling, this study recognizes all previous subspecies of P. maculatus as distinct species on the minim-basis of non-monophyly. Phrynocephalus maculatus was described from the Iranian Plateau (Anderson 1872) with the type locality of “Awada, Shiraz, Persia ”; corrected to “Abadeh [31o 10' N, 52o 37' E; Fars Province, Iran], north of Shiraz” by Blanford (1876). Phrynocephalus maculatus longicaudatus was described by Haas (1957), with a type locality of “Doha Dhalum, Saudi Arabia ”. The sole known population of P. maculatus from Turkmenistan was reported by Bogdanov et al. (1974), and subsequently described as a distinct species, Phrynocephalus golubewii with the type locality of “vicinity of the Bami rail-road station” [= 7 km N railway station Bamy], (Shenbrot & Semenov 1990). It is recommended to recognize P. golubewii, P. longicaudatus, and P. maculatus as distinct species based on phylogenetic results: a). While Phrynocephalus golubewii (Shenbrot & Semenov 1990) may look like P. maculatus living in saltbed dry lakes, it is directly-related to the clade containing P. turcomanus and P. helioscopus (mt-DNA, bootstrap 91%, decay index 6; combined data, bootstrap 87%, decay index 7; allozyme presence/absence, bootstrap 78%, decay index 1; allozyme step matrix, bootstrap 75%, decay index 2; non-monophyly in RAG- 1); in fact P. turcomanus occurs within meters of P. golubewii and may overlap in distribution. b). Phrynocephalus maculatus (Anderson 1872) is restricted to salt-bed dry lakes on the Iranian Plateau where its type locality is located, and adjacent Baluchistan Plateau including the base of the Sulaiman Range of southern Pakistan. P. maculatus is related, yet outside and sister to a clade containing P. arabicus and P. longicaudatus (see below) based on both mitochondrial and nuclear RAG-1 DNA evidence (mt-DNA, bootstrap 100%, decay index 32; RAG-1, bootstrap 100%, decay index 13; combined data, bootstrap 100%, decay index 46; all of these taxa were not sampled for allozyme data). c). Phrynocephalus longicaudatus (Haas 1957) New Status is restricted to the Arabian Peninsula. It is the sister taxon to P. arabicus which is also endemic to the Arabian Plate and supported by mitochondrial and nuclear RAG- 1 DNA evidence (mt-DNA, bootstrap 100%, decay index 22; RAG- 1, bootstrap 100%, decay index 7; combined data, bootstrap 100%, decay index 30; these taxa were not sampled for allozyme data). These taxa in any combination do not form a monophyletic group. 2). Small Sand Dune Species of Southwest Asia and Caspian Basin Phrynocephalus interscapularis interscapularis (Lichtenstien 1856; type locality “Bucharei” = Uzbekistan) and Phrynocephalus interscapularis sogdianus (Chernov 1948; type locality “ Tadjikistan, vicinity of the Pjandzh village”, approximately 37o 14' N, 69o 05' E) are recognized here as full species P. interscapularis and P. sogdianus based on allozyme and DNA differences. The mt-DNA pair-wise uncorrected sequence distance between P. interscapularis and P. sogdianus is 3.2% and these taxa show one fixed allozyme difference at the carboxylic ester hydrolase locus (EST; Table 2). Phrynocephalus ornatus vindumi [Golubev, 1998; type locality “Iran, Khorasan Prov., 35 km. N of Gonabad on road to Torbat-E. Heydariyeh (ca. 34o 49' N, 58o 47' E), 850 m. elevation”; sampling here within ten kilometers of the type locality, see appendix 1] is elevated to species status as Phrynocephalus vindumi New Status. Phrynocephalus ornatus (Boulenger 1887; type locality “between Nushki and Helmand, Baluchistan”) and P. vindumi appear in a non-monophyletic arrangement as P. vindumi occurring on the northeastern portion of the Iranian Plateau is separated from P. ornatus that occurs south of the Hindu Kush in Afghanistan and southwestern Pakistan. In fact, P. vindumi is related to small sand dune species in the Caspian Basin to the north: P. interscapularis and P. sogdianus. In the mt-DNA parsimony analysis, P. vindumi appears in a sister position to the clade containing P. interscapularis and P. sogdianus, with those three taxa grouping with high support (bootstrap 98%, decay index 13). Alternatively, P. ornatus is sister to P. clarkourum (bootstrap 99%, decay index 21), and those two taxa are included in a clade with the two P. luteogutattus samples that also receive high support (bootstrap 100%, decay index 26). Allozyme and RAG-1 DNA data did not sample appropriately to test this question. 3). Gravel Dwelling Phrynocephalus Species in the Caspian Basin Previously, P. reticulatus, P. bannikovi, and P. strauchi were all considered subspecies of P. reticulatus (Eichwald 1831): The nominal subspecies Phrynocephalus reticulatus reticulatus [Eichwald 1831; type locality "Hab. ad Oxum, pristinum amnem, in caspii maris litore orietali", corrected to the Amu-Darja River by Golubev (1991)] and two additional distinct subspecies, Phrynocephalus reticulatus strauchi [Nikolsky 1899; restricted type locality “Khodzhent, 40o 17' N, 69o 37' E, northern Tajikistan ” by lectotype designated by Dunayev (1995)] and Phrynocephalus reticulatus bannikovi (Darevsky, Rustamov & Shammakov 1976; type locality “Tuarkyr Mountains, northwestern Turkmenistan ”), (see Bannikov et al. 1977; although P. reticulatus and P. strauchi were originally described as species). Here these gravel ecotypes are recognized as distinct species based on genetic difference and lack of monophyly. In our sampling, P. bannikovi and P. strauchi do not appear monophyletic with P. strauchi and P. rossikowi grouping with light support (mt-DNA, bootstrap 55%, decay Index 2). P. reticulatus from central Uzbekistan was not sampled in our study or the study of Solovyeva et al. (2014). If P. bannikovi and P. rossikowi were removed from the parsimony analysis performed on the mt-DNA data, the topology of the trees in this study and that of Solovyeva et al. (2014) would be congruent. Based on incomplete sampling, it is recommended that members of the P. reticulatus complex be recognized as separate species Phrynocephalus bannikovi (Darevsky, Rustamov & Shammakov 1976) New Status, P. reticulatus, and P. strauchi. Solovyeva et al. (2014) appear to have not sampled any species from Turkmenistan, which includes P. raddei raddei and P. rossikowi, and in our phylogenetic analyses are related to the P. reticulatus complex. The inclusion of P. raddei raddei and P. rossikowi in the sampling presented here indicates the traditional P. reticulatus complex is not monophyletic, deserving species recognition for the three former subspecies. In common with this study, Solovyeva et al. (2014) only sampled P. strauchi from the P. reticulatus complex. In the sampling of this study, P. bannikovi and P. raddei are geographically oriented disjunctive-adjacent to each other in eastern and west-central Turkmenistan, respectively, and are sister taxa receiving high support from mt-DNA (bootstrap 100%, decay index 23), nuclear RAG-1 DNA (bootstrap 91%, decay index 3), and combined data analyses (bootstrap 100%, decay index 27). Solovyeva et al. (2014) did not sample P. bannikovi, P. rossikowi, or P. raddei raddei from Turkmenistan. However, Solovyeva et al. (2014) and this study did sample P. strauchi from the Fergan Valley of eastern Uzbekistan or northern Tadjikistan. Sampling between this study and Solovyeva et al. (2014) differs in the subspecies of P. raddei: This study uses P. raddei raddei, (Boettger 1888: type locality “Perewalnaja an der transcaspischen Bahn", = Perevalnaja railroad station, southwestern Turkmenistan; near where this study's sample originated from), whereas Solovyeva et al. (2014) uses P. raddei boettgeri (Nikolsky 1905; type locality “Schirabad, Buchara orient” = Sherabad, approximately 37o 42' N, 67o 04' E, southern Uzbekistan) with a vague locality provided as “Turan Depression” [= Caspian Basin]; but occurs in extreme southern Uzbekistan and southern Tajikistan along the Afghanistan border (Bannikov et al. 1977). Until a single study samples both subspecies of P. raddei, it will be unclear how they relate to P. bannikovi, P. reticulatus, P. strauchi, and P. rossikowi. In Solovyeva et al. (2014) no detailed locality information is available, nor is an appendix with this information provided. All GenBank entries do not include locality data, so this information is not archival with NIH GenBank. The Zoological Museum of Moscow University (ZMMU) (http://zmmu.msu.ru/en/about-muzeum/ fonds/amphibians-&-reptiles-collections) in which specimens reported in Solovyeva et al. (2014) are housed, does not offer an obvious means of searching by voucher number on their website. Additionally, this institution does not currently participate in global databases which compile data from institutions across the globe and allow free public access to museum collections involving voucher number relating to locality information. VertNet, a global organization whose participants include 64 institutions spanning 13 countries including Russia, is one prominent database to organize museum collections around the world. The Global Information Biodiversity Facility (GBIF) also offers a forum for researchers to access information on specimens collected in select countries on every continent, which is an added resource. The Zoological Museum of Moscow University is not currently participating in any of these databases, and localities in the manuscript of Solovyeva et al. (2014) are very vague, crossing large geographic regions, and not informative for the difficulties of dealing with Phrynocephalus taxonomy and sampling. Solovyeva et al. (2014) present GenBank sequences as common courtesy in the scientific community, but in those sequences, no locality information is provided so comparing Solovyeva et al. (2014) results with this study is difficult across all issues, starting with taxonomy. Published localities in Solovyeva et al. (2014) such as Central and Middle Asia are not informative, as in the Russian style geography, Middle Asia refers to the Caspian Basin, and Central Asia refers to Tibet, Mongolia and western China. Localities of such gross generality do not provide any information for a study of this detail. 4). Phrynocephalus helioscopus Complex in the Caspian Basin Phrynocephalus helioscopus turcomanus [Dunayev, Solovyeva & Poyarkov 2012 ( in Solovyeva et al. 2012); type locality 40° 01′ 00′′ N 52° 58′ 00′′ E, current name Turkmenbashi, Turkmenistan] is recognized as a distinct species Phrynocephalus turcomanus [Dunayev, Solovyeva & Poyarkov 2012 ( in Solovyeva et al. 2012)] New Status from P. helioscopus based on a north-south distributional disjunction in the Caspian Basin with significant mt-DNA differences. P. helioscopus [Pallas (1771) listed the type locality as "in deserti australioris collibus ardentissimis"; restricted (with no lectotype) to the "Inderskija Gory, Gebiet des unteren Uralflusses" by Mertens & Müller (1928), now Inderskije (Inder) Mountains, Atyrau Region, northwestern Kazakhstan] here is sampled from the northern Aral Sea region of Kazakhstan, and P. turcomanus from southwestern Turkmenistan (Appendix 1). While no fixed allozyme differences are detected between populations sampled, there is a high mt-DNA difference. The mt-DNA pairwise uncorrected sequence distance between P. helioscopus and P. turcomanus is 4.5%; therefore significantly diverged and in the range of sister species divergence for this segment of the mitochondrial genome among amphibians and reptiles (for summary see table 6 in Weisrock et al. 2001). 5). Use of Names in the Low Elevation Chinese Desert—Northern Caspian Basin Clade a). Phrynocephalus salenskyi The name P. salenskyi Bedriaga 1907 is applied here for populations sampled from the Junggur Depression of northwestern China, and the Zaysan Depression of northeastern Kazakhstan. P. salenskyi is the oldest name available by date and page number for Phrynocephalus populations of the low elevation Chinese desert – northern Caspian Basin clade in this geographic region, see below: Phrynocephalus Salenskyi Bedriaga 1907: 141, 213.—Restricted type locality: “ Urungu Fluss ” [= Ulungur River, Xinjiang Uygur Autonomous Region, China]. Phrynocephalus Isseli Bedriaga 1907: 229.— Type locality: Desert in vicinity of Mt. Ssalburty (in Saur Mountains of the Tarbagatay Range, Xinjiang Uygur Autonomous Region, China). Phrynocephalus Haeckeli Bedriaga 1907: 237.— Type locality: Desert in vicinity of Mt. Ssalburty (in Saur Mountains of the Tarbagatay Range), in the Tarbagataï (= Tacheng County) district, northern Xinjiang Uygur Autonomous Region, China. Phrynocephalus Arcellazzii Bedriaga 1907: 307.— Type locality: “ Gaschun, in der Nähe von Gutschen in der Dschungarei ” [= Qitai (44° 00' N, 89° 50' E), Junggar Depression, Xinjiang Uygur Autonomous Region, China]. Phrynocephalus Grum-Grzimailoi Bedriaga 1909: 420.—Restricted type locality: “Gutschen” [= Qitai (44° 00' N, 89° 50' E), Junggar Depression, Xinjiang Uygur Autonomous Region, China]. Phrynocephalus bedriaga Tsarevsky 1926: 214.—Restricted type locality: Zaissan Lake region, eastern Kazakhstan. Phrynocephalus albolineatus K. Zhao 1979: 113.— Type locality: "45 kilometers south of Tacheng County, Xinjiang Uygur Autonomous Region, China; 584 m elevation". The Irtysh River flows out of Zaysan Lake linking the Junggar Depression with the Zaysan Depression, which makes a compelling case for sampling presented in this study representing P. salenskyi populations. b). Phrynocephalus przewalskii Complex in the Gobi Desert Phrynocephalus Przewalskii Strauch 1876: 10.— Type locality: “deserto Alaschanico ” [= Tengger Desert, Nei Mongol Autonomous Region, China]; corrected to eastern Tengger Desert west from 106° E, southwest Nei Mongol Autonomous Region, China in Barabanov & Ananjeva (2007). This taxon is generally a sand dwelling ecotype. P. frontalis Strauch 1876: 15.—Type locality: locality “Ordos dicta” [= Ordos Desert, Nei Mongol Autonomous Region, China]. This taxon is generally thought of as a gravel dwelling ecotype, and here considered a synonym of P. przewalskii (see below). A lack of genetic diversity between populations from these taxa (two populations each), confirm that only the name P. przewalskii is valid, whereas P. frontalis is a synonym of P. przewalskii (following Gozdzik & Fu 2009). This is supported by nuclear encoded allozyme data, and mt-DNA data. Alternative names exist as reviewed in Barabanov & Ananjeva (2007), but without detailed sampling of type localities, the validity of alternate names cannot be addressed. Across the 25 allozyme loci sampled here, we detected no fixed differences among the four populations of P. przewalskii, indicating a fair amount of gene flow between the populations. The average mt-DNA pair-wise uncorrected sequence distances between population 2 and the clade containing populations 1, 3, and 4 is 3.64% (range 3.41–3.79%). The average pair-wise comparison within the clade containing populations 1, 3 and 4 is 0.57% (range 0.51–0.63%). These results match the phylogenetic topology discovered in the mt-DNA parsimony analysis, with population 2 placed in a sister position to the clade containing populations 1, 3 and 4 (bootstrap 100%, decay index 19). No resolution was discovered among the four P. przewalskii populations in the RAG- 1 DNA parsimony analysis. Urquhart et al. (2009) suggests a vicariant break among P. przewalskii populations roughly corresponding to the Helan Shan (mountains). Data reported here do not corroborate that hypothesis. Sampled are a single population east of the Yellow River and three populations west of the Yellow River, which flows north to south in this region. Population 2 is sampled on the east side of the river, and population 3 is sampled on the west side of the river, on banks directly across from each other in Shapotou, Ningxia Autonomous Region, China. Population 2, the only population on the east side of the Yellow River is the most divergent, with populations 1, 3 and 4 on the west side of the river nearly identical in our sampling. Population 1 (Wujiachuan, Gansu Province, China) is west of the Yellow River and further south than sampling conducted in Urquhart et al. (2009) and Wang & Fu (2004), yet nearly identical to other populations sampled west of the Yellow River. In our sampling, only population 4 lies west of the Helan Shan (Wuwei, Gansu Province, China) in the Tengger Desert, the type locality of P. przewalskii. Sampling in this study suggests the Yellow River could be responsible for a genetic break in P. przewalskii populations, and not the Helan Shan, though more detailed, fine-scale sampling and analysis will need to be done to confirm this hypothesis. In the Zhongwei region of Ningxia Autonomous Region, there is an indication that east and west haplotypes as described above occur in the same locality, which could indicate a point of hybridization, but further detailed work is needed (Wang & Fu 2004). Sampling conducted in similar areas in Wang & Fu (2004) and Urquhart et al. (2009) did not completely overlap, making conclusions between these studies and this study very difficult for comparative interpretation, further indicating additional work is needed. In sampling of the P. przewalskii complex in Jin & Brown (2013), populations to the east are grouped, and are referred to as P. frontalis, with populations to the west grouping and referred to as P. przewalskii, not conflicting with results reported here. 6). Use of Names in the Northern Tibetan Plateau, Published as part of Macey, Robert, Schulte, James A., Ananjeva, Natalia B., Van, Erik T., Wang, Yuezhao, Orlov, Nikolai, Shafiei, Soheila, Robinson, Michael D., Dujsebayeva, Tatjana, Freund, Gabriel S., Fischer, Clayton M., Liu, David & Papenfuss, Theodore J., 2018, A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics, pp. 1-81 in Zootaxa 4467 (1) on pages 34-38, DOI: 10.11646/zootaxa.4467.1.1, http://zenodo.org/record/1442134, {"references":["Anderson, J. (1872) On some Persian, Himalayan, and other Reptiles. Proceedings of the Zoological Society of London, 1872 (2), 371 - 404.","Blanford, W. T. (1876) List of Reptilia and Amphibia collected by the late Dr. Stoliczka in Kashmir, Ladak, Eastern Turkestan and Wakhan, with descriptions of new species. Journal of the Asiatic Society of Bengal, 44 (3), 191 - 196.","Haas, G. (1957) Some amphibians and reptiles from Arabia. Proceedings of the California Academy of Sciences, 29 (3), 47 - 86.","Bogdanov, O. P., Atayev, Ch. & Shammakov, S. (1974) The finding of Phrynocephalus maculatus in the USSR. Zoologicheskij Zhurnal, Moscow, 53 (2), 304 - 305. [in Russian]","Shenbrot, G. I. & Semenov (as Semyonov), D. V. (1990) New species of the genus Phrynocephalus (Reptilia, Agamidae) from Turkmenia. Zoologichesky Zhurnal, Moscow, 69 (9), 154 - 156. [in Russian with English summary]","Chernov, S. A. (1948) Reptiles - Reptilia. In: Pavlovsky, E. N. & Vinogradova, B. S. (Eds.), The Animals of the USSR. Fol. 2. The desert zone. USSR Academy of Sciences, Moscow-Leningrad, pp. 127 - 161. [in Russian]","Golubev, M. L. (1998) A new subspecies of Phrynocephalus ornatus Boulenger (Reptilia: Agamidae) from Eastern Iran, with a key to South-Western and Middle Asian microphrynocephalids. Hamadryad, 23 (2), 162 - 168.","Boulenger, G. A. (1887) Catalogue of the Lizards in the British Museum (Natural History), Second Edition. Fol. 3. Lacertidae, Gerrhosauridae, Scincidae, Anelytropidae, Dibamidae, Chamaeleontidae. Trustees of the British Museum, London, 575 pp.","Eichwald, E. (1831) Zoologia specialis quam expositis animalibus tum vivis, tum fossilibus potissimum Rossiae in universum, et Poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Filnensi habendarum. Pars posterior. Josephi Zawadzki, Vilnae, 396 pp.","Golubev, M. L. (1991) About the name Agama ocellata Lichtenstein in Eversmann, 1823 (Reptilia, Agamidae) with redescription of the types. Herpetological Researches, Leningrad, 1, 12 - 17. [in Russian with English summary]","Nikolsky, A. M. (1899) Reptiles and amphibians of the Turkestan general-governorship (Herpetologia Turanica). In: Fedschenko, A. P. (Ed.), Travel to the Turkestan of the member-founder of the A. P. Fedchenko Society, accomplished on behalf of the Imperial Society of Amateurs of natural history, anthropology, and ethnography. Zoogeographical Investigations. University Publishing, Moscow, Issue 23, Vol. 2, part 7, pp. 21 - 84. [in Russian]","Dunayev, E. A. (1995) Reviewed description of the types of Phrynocephalus strauchi Nikolsky, 1899 (Squamata, Agamidae) and materials on the history of its study, distribution, and variability. Russian Journal of Herpetology, 2 (2), 87 - 94.","Darevsky, I. S., Rustamov, A. K. & Shammakov, S. (1976) Status and distribution of Phrynocephalus reticulatus Eichwald (Sauria, Agamidae) in Central Asia. Teoreticheskije i prikladnyije aspekty okhrany prirody i okhotovedenija, Moscow, 84, 113 - 119. [in Russian]","Bannikov, A. G., Darevsky, I. S., Ischchenko, V. G., Rustamov, A. K. & Szczerbak, N. N. (1977) Opredelitel' Zemnovodnikh i Presmikayushchikhsya Fauni SSSR [Guide to the Reptile and Amphibian Fauna of the USSR]. Prosveshchenie, Moscow, 415 pp., figs. 1 - 108, 135 maps, 32 col. pls. [in Russian]","Solovyeva, E. N., Poyarkov, N. A., Dunayev, E. A., Nazarov, R. A., Lebedev, V. S. & Bannikova, A. A. (2014) Phylogenetic relationships and subgeneric taxonomy of Toad Headed Agamas Phrynocephalus (Reptilia, Squamata, Agamidae) as determined by mitochondrial DNA sequencing. Doklady Biological Sciences, 455, 119 - 124. https: // doi. org / 10.7868 / S 0869565214100260","Boettger, O. (1888) Uber die Reptilien und Batrachier Transcaspiens. Zoologischer Anzeiger, Leipzig, 11 (279), 259 - 263.","Nikolsky, A. M. (1905) Reptiles and amphibians of the Russian Empire (Herpetologica Rossica). Memoires de l'Academie Imperiale des Sciences de St. - Petersbourg, 17 (1), 518 pp. [in Russian]","Solovyeva, E. N., Dunayev, E. A. & Poyarkov, N. A. (2012) Interspecific taxonomy of Sunwatcher Toadhead Agama species complex (Phrynocephalus helioscopus, Squamata). Zoologicheski Zhurnal, Moscow, 91 (11), 1377 - 1396. [in Russian with English summary]","Pallas, P. S. (1771) Reise durch verschiedene Provinzen des Russischen Reichs. Bd. I. St. Petersburg: Kayserliche Acade-mie der Wissenschaften, 540 pp.","Mertens, R. & Muller, L. (1928) Liste der Amphibien und Reptilien Europas. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 41 (1), 1 - 62.","Weisrock, D. W., Macey, J. R., Ugurtas, I. H., Larson, A. & Papenfuss, T. J. (2001) Molecular phylogenetics and historical biogeography among salamandrids of the \" true \" salamander clade: Rapid branching of numerous highly divergent lineages with the rise of Anatolia in Mertensiella luschani. Molecular Phylogenetics and Evolution, 18 (3), 434 - 448. https: // doi. org / 10.1006 / mpev. 2000.0905","Bedriaga, J. (1907) Amphibien und Reptilien (Reptilia Przewalskiana). Wissenschaftliche Resultate der von N. M. Przewalski nach Central-Asien unternommenen Reisen. Zoologischer Theil, Bd. 3, Abth. 1, Lfrg. 2, St. Petersburg, pp. 71 - 278.","Bedriaga, J. (1909) Amphibien und Reptilien (Reptilia Przewalskiana). In: Wissenschaftliche Resultate der von N. M. Przewalski nach Central-Asien unternommenen Reisen. Zoologischer Theil, Bd. 3, Abth. 1, Lfrg. 3, St. Petersburg, pp. 279 - 502.","Tsarevsky (as Carevskij), S. (1926) Contributions sur la systematique et la distribution des lezards du genre Phrynocephalus (Reptilia). Comptes Rendus de l'Academie des Sciences de l'URSS, 1926, 211 - 214. [in Russian with French title]","Zhao, K. (1979) A survey of the classification and distribution of the toad-headed agamids (Phrynocephalus) in China. Acta Scientiarum Naturalium Universitatis Intramongolicae, Hohhot, 1979 (2), 111 - 121. [in Chinese with English summary]","Strauch, A. A. (1876) Part III. Reptilia and Amphibia. In: Imperial Russian Geographic Society (Ed.), Przevalsky N. M., Mongolia and the Tangut Lands, a Three-Year Journey in Eastern High Asia. Fol. II. Reptiles and Amphibians. V. S. Balashov printing house, St. Petersburg, pp. 1 - 55. [in Russian]","Barabanov, A. V. & Ananjeva, N. B. (2007) Catalogue of the available scientific species-group names for lizards of the genus Phrynocephalus Kaup, 1825 (Reptilia, Sauria, Agamidae). Zootaxa, 1399, 1 - 56. https: // doi. org / 10.11646 / zootaxa. 1399.1.1","Urquhart, J., Wang, Y. & Fu, J. (2009) Historical vicariance and male-mediated gene flow in the toad-headed lizards Phrynocephalus przewalskii. Molecular Ecology, 18 (17), 3714 - 3729. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2009.04310. x","Wang, Y. & Fu, J. (2004) Cladogenesis and vicariance patterns in the toad-headed lizard Phrynocephalus versicolor species complex. Copeia, 2004 (2), 199 - 206. https: // doi. org / 10.1643 / CG- 03 - 082 R 1","Jin, Y. - T. & Brown, R. P. (2013) Species history and divergence times of viviparous and oviparous Chinese toad-headed sand lizards (Phrynocephalus) on the Qinghai-Tibetan Plateau. Molecular Phylogenetics and Evolution, 68 (2), 259 - 268. https: // doi. org / 10.1016 / j. ympev. 2013.03.022","Bedriaga, J. (\" 1905 \" 1906) Verzeichnis der von der Central-Asiatischen Expedition unter Stabs-Kapitan W. Roborowski in den Jahren 1893 - 1895 gesammelten Reptilien. Annuaire du Musee Zoologique de l'Academie Imperiale des Sciences de St. - Petersbourg, 10 (3 - 4), 159 - 200.","Jiang, Y. - M., Huang, Q. - Y. & Zhao, E. - M. (1980) A new subspecies of Phrynocephalus vlangalii (Strauch) and preliminary observation on its ecology. Acta Zoologica Sinica, Beijing, 26 (2), 178 - 183. [in Chinese with English summary]","Jin, Y. - T., Brown, R. P. & Liu, N. - F. (2008) Cladogenesis and phylogeography of the lizard Phrynocephalus vlangalii (Agamidae) on the Tibetan plateau. Molecular Ecology, 17 (8), 1971 - 1982. https: // doi. org / 10.1111 / j. 1365 - 294 X. 2008.03721. x","Pang, J., Wang, Y., Zhong, Y., Hoelzel, A. R., Papenfuss, T. J., Zeng, X., Ananjeva, N. B. & Zhang, Y. - P. (2003) A phylogeny of Chinese species in the genus Phrynocephalus (Agamidae) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 27 (3), 398 - 409. https: // doi. org / 10.1016 / S 1055 - 7903 (03) 00019 - 8","Qi, Y., Yang, W., Lu, B. & Fu, J. (2013) Genetic evidence for male-biased dispersal in the Qinghai toad-headed agamid Phrynocephalus vlangalii and its potential link to individual social interactions. Ecology and Evolution, 3 (5), 1219 - 1230. https: // doi. org / 10.1002 / ece 3.532","Wu, Y., Fu, J., Yue, B. & Yin, Q. (2015) An atypical reproductive cycle in a common viviparous Asia Agamid Phrynocephalus vlangalii. Ecology and Evolution, 5 (21), 5138 - 5147. https: // doi. org / 10.1002 / ece 3.1783","Staub, N. L., Brown, C. W. & Wake, D. B. (1995) Patterns of growth and movements in a population of Ensatina eschscholtzii platensis (Caudata: Plethodontidae) in the Sierra Nevada, California. Journal of Herpetology, 29 (4), 593 - 599. https: // doi. org / 10.2307 / 1564743","Jackman, T. R. & Wake, D. B. (1994) Evolutionary and historical analysis of protein variation in the blotched forms of salamanders of the Ensatina complex (Amphibia: Plethodontidae). Evolution, 48 (3), 876 - 897. https: // doi. org / 10.1111 / j. 1558 - 5646.1994. tb 01369. x","Wake, D. B. & Schneider, C. J. (1998) Taxonomy of the plethodontid salamander genus Ensatina. Herpetologica, 54 (2), 279 - 298.","Noble, D. W. A., Qi, Y. & Fu, J. (2010) Species delineation using Bayesian model-based assignment tests: A case study using Chinese toad-headed agamas (genus Phrynocephalus). BMC Evolutionary Biology, 2010 (10), 197. https: // doi. org / 10.1186 / 1471 - 2148 - 10 - 197"]}

F. Phrynocephalus Species Diversity as Related to Geology, Habitat and Size Two regions show high species numbers in close proximity. Six species are found on the northern side of the Kopet-Dagh (mountains) along the southern margin of the Caspian Basin in Turkmenistan. Species that inhabit hard substrates are P. rossikowi, P. raddei, P. bannikovi, P. golubewii, and P. turcomanus, with P. interscapularis and P. mystaceus occurring in soft substrates (Table 5, Fig. 16). Among these species there are major size differences (Table 6). Six species are also found at the juncture of the Helmand Basin of southern Afghanistan with the Baluchistan Plateau including the base of the Sulaiman Range of southern Pakistan. Species that inhabit hard substrates are P. maculatus and P. scutellatus (Table 5, Fig. 16). Smaller species that inhabit soft substrates are P. clarkorum, P. ornatus, and P. luteoguttatus (Clade C in Figs. 4 and 8). A larger species inhabiting soft substrates in large floating sand dunes is P. euptilopus, which was not sampled in this study even though repeated efforts were made in both Afghanistan and Pakistan by the last author. Among these species there are major size differences (Table 6). Both of these examples share a plate tectonic past around the northern migration of the Arabian Plate and its impact on ancient tectonic plates to the north in Eurasia. The Kopet-Dagh (mountains), on the northern margin of the Iranian Plateau. descends into the Caspian Basin. This region of the Caspian Basin experienced the most dramatic changes via tectonic uplift, erosion, and sea floor drying of the Paratethys Sea creating unique and diversified habitats (see Figs. 3, 9–14; Table 4). Additionally, the Helmand Block of southern Afghanistan is being pushed south by impinging Arabia and further northern invasion of the Indian Plate (Fig. 9). This has created uplift in the Gulf of Oman creating the push-back of the Makron in southern Pakistan (Figs. 9–12). It is dynamic tectonics of this nature that created a series of habitats leading to speciation among Phrynocephalus. As adaptive niches were created, size diversity, morphological specializations, background matching, and defensive mimicry developed (Plates I – VIII). Future work on Phrynocephalus will need to take into account these aspects with particular attention to plate tectonic induced events in order to investigate phyletic patterns, taxonomy, and evolutionary diversity among one of the world’s most interesting and confusing lizard genera., Published as part of Macey, Robert, Schulte, James A., Ananjeva, Natalia B., Van, Erik T., Wang, Yuezhao, Orlov, Nikolai, Shafiei, Soheila, Robinson, Michael D., Dujsebayeva, Tatjana, Freund, Gabriel S., Fischer, Clayton M., Liu, David & Papenfuss, Theodore J., 2018, A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics, pp. 1-81 in Zootaxa 4467 (1) on page 51, DOI: 10.11646/zootaxa.4467.1.1, http://zenodo.org/record/1442134

F. Phrynocephalus Species Diversity as Related to Geology, Habitat and Size Two regions show high species numbers in close proximity. Six species are found on the northern side of the Kopet-Dagh (mountains) along the southern margin of the Caspian Basin in Turkmenistan. Species that inhabit hard substrates are P. rossikowi, P. raddei, P. bannikovi, P. golubewii, and P. turcomanus, with P. interscapularis and P. mystaceus occurring in soft substrates (Table 5, Fig. 16). Among these species there are major size differences (Table 6). Six species are also found at the juncture of the Helmand Basin of southern Afghanistan with the Baluchistan Plateau including the base of the Sulaiman Range of southern Pakistan. Species that inhabit hard substrates are P. maculatus and P. scutellatus (Table 5, Fig. 16). Smaller species that inhabit soft substrates are P. clarkorum, P. ornatus, and P. luteoguttatus (Clade C in Figs. 4 and 8). A larger species inhabiting soft substrates in large floating sand dunes is P. euptilopus, which was not sampled in this study even though repeated efforts were made in both Afghanistan and Pakistan by the last author. Among these species there are major size differences (Table 6). Both of these examples share a plate tectonic past around the northern migration of the Arabian Plate and its impact on ancient tectonic plates to the north in Eurasia. The Kopet-Dagh (mountains), on the northern margin of the Iranian Plateau. descends into the Caspian Basin. This region of the Caspian Basin experienced the most dramatic changes via tectonic uplift, erosion, and sea floor drying of the Paratethys Sea creating unique and diversified habitats (see Figs. 3, 9–14; Table 4). Additionally, the Helmand Block of southern Afghanistan is being pushed south by impinging Arabia and further northern invasion of the Indian Plate (Fig. 9). This has created uplift in the Gulf of Oman creating the push-back of the Makron in southern Pakistan (Figs. 9–12). It is dynamic tectonics of this nature that created a series of habitats leading to speciation among Phrynocephalus. As adaptive niches were created, size diversity, morphological specializations, background matching, and defensive mimicry developed (Plates I – VIII). Future work on Phrynocephalus will need to take into account these aspects with particular attention to plate tectonic induced events in order to investigate phyletic patterns, taxonomy, and evolutionary diversity among one of the world’s most interesting and confusing lizard genera.

E. Phrynocephalus Size Diversity Phrynocephalus show a variety of snout vent lengths and tail lengths, (Table 6, Appendix 5). Body size per sample ranged from 106 mm for P. mystaceus to 35 mm for P. vindumi and P. interscapularis. Tail length per sample ranges from 133 mm for P. maculatus population 1 to 35 mm for P. luteoguttatus population 2. P. mystaceus and P. interscapularis are sympatric in the Caspian Basin, and habitat partition with P. mystaceus in large floating sand dunes, with P. interscapularis living on the edge of those large sand dunes, which may revolve around ecotype evolution in a predator-prey relationship. In northwestern Iran P. vindumi replaces the role of P. interscapularis, and is sympatric with P. mystaceus. Tail length appears to be the most sexually dimorphic characteristic. ......continued on the next page Snout vent lengths for clay ecotypes range from 62 mm for P. przewalskii population 1 to 49 mm for P. versicolor, with tail lengths ranging from 90 mm for P. przewalskii population 1 to 49 mm for P. persicus. The snout vent length for gravel ecotypes range from 64 mm for P. vlangalii population 1 to 40 mm for P. rossikowi, with tail lengths ranging from 90 mm for P. longicaudatus to 42 mm for P. theobaldi. Snout vent lengths for dry lakebed ecotypes range from 90 mm for P. maculatus population 1 to 60 mm for P. golubewii, with tail length ranging from 133 mm for P. maculatus population 1 to 78 mm for P. maculatus population 2. Snout vent lengths for small sand dune ecotypes range from 61 mm for P. vlangalii population 2 to 35 mm for both P. vindumi and P. interscapularis, with tail length ranging from 80 mm for P. przewalskii population 2 to 35 mm for P. luteoguttatus population 2. Snout vent lengths for large sand dune ecotypes range from 106 mm for P. mystaceus populations to 59 mm for P. salenskyi population 1, with tail lengths ranging from 110 mm for P. mystaceus population 1 to 70 mm for P. salenskyi population 1. The largest Phrynocephalus occupy large floating sand dunes, and the smallest Phrynocephalus species are found on the fringes of large floating sand dunes in the Caspian Basin and northern Iran. P. mystaceus occupying large sand dunes in these habitats coexists with the two smallest species that occupy the fringes of those sand dunes, P. interscapularis in the Caspian Basin, and P. vindumi in northwestern Iran. It has not escaped our attention that size and habitat partitioning may have occurred in these regions because it is likely that smaller species (P. interscapularis or P. vindumi) serve as a food source for larger species (P. mystaceus). Further work is needed for phylogenetic assessment of size evolution among Phrynocephalus species and populations considering the sexually dimorphic nature of this genus, both with body size and tail length., Published as part of Macey, Robert, Schulte, James A., Ananjeva, Natalia B., Van, Erik T., Wang, Yuezhao, Orlov, Nikolai, Shafiei, Soheila, Robinson, Michael D., Dujsebayeva, Tatjana, Freund, Gabriel S., Fischer, Clayton M., Liu, David & Papenfuss, Theodore J., 2018, A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics, pp. 1-81 in Zootaxa 4467 (1) on pages 48-51, DOI: 10.11646/zootaxa.4467.1.1, http://zenodo.org/record/1442134

The cloud forests of Mesoamerica are notable for their high endemism, and plethodontid salamanders provide a striking example of divergence and microendemism across cloud forest blocks at a regional level. Salamanders that make use of arboreal bromeliad microhabitats in the cloud forest appear to be especially prone to divergence driven by natural habitat fragmentation, and are expected to show high endemism at small spatial scales. We use a multilocus dataset to investigate the biogeographic history and relationships among species of a small genus of salamander, Cryptotriton, restricted to the cloud forests of Nuclear Central America. We use a morphological data set along with a coalescent species delimitation method to reveal the presence of at least one undescribed species from an isolated cloud forest in eastern Guatemala. Biogeographic analyses show that Cryptotriton has a different biogeographic history than another clade of cloud forest-restricted salamanders in the same region, perhaps indicating that each genus restricted the spatial expansion and diversification of the other through preemptive occupancy. Our results suggest that isolation across relatively short geographic distances has led to range fragmentation and deep divergence between species. Exploration of remaining patches of cloud forest likely will continue to reveal undetected diversity.

Badiane, Arnaud, Garcia-Porta, Joan, Červenka, Jan, Kratochvíl, Lukáš, Sindaco, Roberto, Robinson, Michael D., Morales, Hernan, Mazuch, Tomáš, Price, Thomas, Amat, Fèlix, Shobrak, Mohammed Y., Wilms, Thomas, Simó-Riudalbas, Marc, Ahmadzadeh, Faraham, Papenfuss, Theodore J., Cluchier, Alexandre, Viglione, Julien, Carranza, Salvador (2014): Phylogenetic relationships of Semaphore geckos (Squamata: Sphaerodactylidae: Pristurus) with an assessment of the taxonomy of Pristurus rupestris. Zootaxa 3835 (1): 33-58, DOI: http://dx.doi.org/10.11646/zootaxa.3835.1.2

Weisrock, David W., Macey, J. Robert, Matsui, Masafumi, Mulcahy, Daniel G., Papenfuss, Theodore J. (2013): Molecular phylogenetic reconstruction of the endemic Asian salamander family Hynobiidae (Amphibia, Caudata). Zootaxa 3626 (1): 77-93, DOI: http://dx.doi.org/10.11646/zootaxa.3626.1.3

Sibon merendonensis new species (Figs. 2,3) Holotype. Museum of Vertebrate Zoology (MVZ) 263880, an adult female (Fig. 2), collected at Finca la Bendici��n, 4.0 km SE (by road) of soccer field at junction with road to Pinalito, then 7.6 km E (by road) of La Trementina, 14.93042��N, 89.4167��W (WGS 84 Datum), 1364 m, Departamento Zacapa, Guatemala, 16 January 2008, by Sean M. Rovito and Theodore J. Papenfuss. Original field number SMR 551. Diagnosis. Sibon merendonensis is a member of the S. annulatus group as defined by Savage (2002) in having the penultimate supralabial separated from the eye by the lower postocular (Fig. 3). It differs from all other members of the S. annulatus group in having a scale row formula of 15���17 ��� 17 rather than 13 ��� 13 ��� 13 in S. anthracops, 15���17 ��� 15 in S. perissostichon, and 15 ��� 15 ��� 15 in all other species in the group. Larger (TOL= 687 mm) than all other members of the S. annulatus group [female TOL range of 366 mm ��� 557 mm (McCranie 2007)], except S. dimidiatus (maximum female TOL of ca. 800 mm) and S. perissisticon (TOL= 668 mm). Dorsal scale rows retain normal orientation vs. changing orientation from sides of body to dorsal midline in S. perissostichon. Table 1 (modified from McCranie 2007) compares S. merendonensis with all other members of the S. annulatus group. Description of holotype. Adult female; large for a species of Sibon (SVL 498 mm; TOL 687 mm); tail short (TAL/TOL 0.275); head very distinct from neck (Fig. 2); snout short, blunt in dorsal outline; rostral not extending posteriorly between internasals, short, two-thirds length of prefrontal suture; prefrontals long, their median suture 87 % length of frontal; prefrontals bordering eye above loreal; frontal longer than wide, widest anteriorly, length two thirds that of parietals, in contact with prefrontals, supraoculars, and parietals; supraoculars longer than wide, equal in length to frontal; parietals long, wide, widest anteriorly, their median suture 20 % longer than frontal; parietals in contact with frontal, supraoculars, upper postoculars, anterior and posterior temporals, and seven nuchal scales; nasal entire, in contact with first three supralabials, loreal, prefrontal, internasal, and rostral; loreal single, longer than high, its length less than one-half length of snout (tip of snout to anterior edge of eye) and one-half length of eye; loreal bordering eye; preoculars absent; subpreoculars absent; postoculars 2, upper wider than lower; temporals 1 + 2, anterior one above 7 th supralabial, lower second on above 8 th supralabial; supralabials 8, with 5 th and 6 th bordering eye (enlarged penultimate supralabial separated from eye by lower postocular); pupil vertically elliptical; eye length two times length of loreal; infralabials 8 - 8, first pair separated behind mental by a single, large postmental, infralabials 1-2 in contact with postmental, infralabials 2-5 in contact with anterior chinshields; anterior chinshields paired, longer than wide, longer than posterior chinshields; posterior chinshields paired, in broad contact medially; mental groove present. Fig. 3 shows head scale characteristics discussed above. Ventrals 184; cloacal scute single; subcaudals 83; dorsal scales smooth, in 15 rows posterior to neck and 17 rows at midbody and anterior to cloaca, without apical pits or supracloacal ridges; vertebral row not enlarged anteriorly, barely enlarged posteriorly. Color in life (Fig. 2). Dorsal components of interspaces between bands very pale yellowish orange, with some cream-colored scales, brighter orange immediately behind head becoming paler posteriorly; dorsal components of bands medium brown. Lateral components of interspaces between bands mottled pale orange-brown; lateral components of bands cream-colored with brown mottling. Dorsal surface of head dark brown with orange mottling, orange patches condense into a stripe along midline posterior to parietals; supralabials brown with cream-colored mottling, sublabials mostly cream-colored with some brown. Iris golden-tan with darker brown flecks. Color in alcohol. Rostrum medium brown, middle of dorsal surface of head whitish with a slight orange tint. Interspaces of bands whitish with orange tint and some brown speckling, bands medium brown dorsally, lighter brown and more speckled on sides of body. Venter pale yellowish cream color with brown speckling from underside of head to about one-third down body, paler than rest of venter. Middle third of venter yellowish grey with more brown speckling, posterior third of venter and anterior half of tail very pale orange cream color with brown speckling. Irregularly spaced small dark brown patches throughout venter. Bands do not continue across venter. Ventral surface of posterior half of tail is a nearly continuous irregular band of medium to light brown with some lighter speckling, thinner where interspaces meet venter. Distribution and ecology. The holotype was found inside an arboreal bromeliad in broadleaf cloud forest with some pine trees. The type locality at La Monta��a de las Granadillas is part of a belt of mid-elevation cloud forest stretching along a ridge to the vicinity of La Uni��n, Zacapa. Although the eastern part of this forest, near La Uni��n, has been sampled fairly extensively without the discovery of any additional specimens, we expect that this species occurs in other portions of the forested ridge. The forest where La Monta��a de las Granadillas is located is separated from other areas of cloud forest by more arid lowland areas, including the arid Motagua Valley to the north. Given the isolated nature of this cloud forest, it seems unlikely that the species is found in other cloud forests in eastern Guatemala. Etymology. The specific epithet makes reference to the Sierra del Merend��n, the mountain range containing the type locality., Published as part of Rovito, Sean M., Papenfuss, Theodore J. & V��squez-Almaz��n, Carlos R., 2012, A new species of Sibon (Squamata: Colubridae) from the mountains of eastern Guatemala, pp. 62-68 in Zootaxa 3266 on pages 65-67, DOI: 10.5281/zenodo.280769, {"references":["Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica. A Herpetofauna between Two Continents, between Two Seas. University of Chicago Press, Chicago, Illinois, U. S. A. 934 pp.","McCranie, J. R. (2007) A second new species of Sibon (Squamata: Colubridae) from La Mosquitia, northeastern Honduras. Herpetologica, 63, 213 - 218."]}

Sibon merendonensis new species (Figs. 2,3) Holotype. Museum of Vertebrate Zoology (MVZ) 263880, an adult female (Fig. 2), collected at Finca la Bendición, 4.0 km SE (by road) of soccer field at junction with road to Pinalito, then 7.6 km E (by road) of La Trementina, 14.93042°N, 89.4167°W (WGS 84 Datum), 1364 m, Departamento Zacapa, Guatemala, 16 January 2008, by Sean M. Rovito and Theodore J. Papenfuss. Original field number SMR 551. Diagnosis. Sibon merendonensis is a member of the S. annulatus group as defined by Savage (2002) in having the penultimate supralabial separated from the eye by the lower postocular (Fig. 3). It differs from all other members of the S. annulatus group in having a scale row formula of 15–17 – 17 rather than 13 – 13 – 13 in S. anthracops, 15–17 – 15 in S. perissostichon, and 15 – 15 – 15 in all other species in the group. Larger (TOL= 687 mm) than all other members of the S. annulatus group [female TOL range of 366 mm – 557 mm (McCranie 2007)], except S. dimidiatus (maximum female TOL of ca. 800 mm) and S. perissisticon (TOL= 668 mm). Dorsal scale rows retain normal orientation vs. changing orientation from sides of body to dorsal midline in S. perissostichon. Table 1 (modified from McCranie 2007) compares S. merendonensis with all other members of the S. annulatus group. Description of holotype. Adult female; large for a species of Sibon (SVL 498 mm; TOL 687 mm); tail short (TAL/TOL 0.275); head very distinct from neck (Fig. 2); snout short, blunt in dorsal outline; rostral not extending posteriorly between internasals, short, two-thirds length of prefrontal suture; prefrontals long, their median suture 87 % length of frontal; prefrontals bordering eye above loreal; frontal longer than wide, widest anteriorly, length two thirds that of parietals, in contact with prefrontals, supraoculars, and parietals; supraoculars longer than wide, equal in length to frontal; parietals long, wide, widest anteriorly, their median suture 20 % longer than frontal; parietals in contact with frontal, supraoculars, upper postoculars, anterior and posterior temporals, and seven nuchal scales; nasal entire, in contact with first three supralabials, loreal, prefrontal, internasal, and rostral; loreal single, longer than high, its length less than one-half length of snout (tip of snout to anterior edge of eye) and one-half length of eye; loreal bordering eye; preoculars absent; subpreoculars absent; postoculars 2, upper wider than lower; temporals 1 + 2, anterior one above 7 th supralabial, lower second on above 8 th supralabial; supralabials 8, with 5 th and 6 th bordering eye (enlarged penultimate supralabial separated from eye by lower postocular); pupil vertically elliptical; eye length two times length of loreal; infralabials 8 - 8, first pair separated behind mental by a single, large postmental, infralabials 1-2 in contact with postmental, infralabials 2-5 in contact with anterior chinshields; anterior chinshields paired, longer than wide, longer than posterior chinshields; posterior chinshields paired, in broad contact medially; mental groove present. Fig. 3 shows head scale characteristics discussed above. Ventrals 184; cloacal scute single; subcaudals 83; dorsal scales smooth, in 15 rows posterior to neck and 17 rows at midbody and anterior to cloaca, without apical pits or supracloacal ridges; vertebral row not enlarged anteriorly, barely enlarged posteriorly. Color in life (Fig. 2). Dorsal components of interspaces between bands very pale yellowish orange, with some cream-colored scales, brighter orange immediately behind head becoming paler posteriorly; dorsal components of bands medium brown. Lateral components of interspaces between bands mottled pale orange-brown; lateral components of bands cream-colored with brown mottling. Dorsal surface of head dark brown with orange mottling, orange patches condense into a stripe along midline posterior to parietals; supralabials brown with cream-colored mottling, sublabials mostly cream-colored with some brown. Iris golden-tan with darker brown flecks. Color in alcohol. Rostrum medium brown, middle of dorsal surface of head whitish with a slight orange tint. Interspaces of bands whitish with orange tint and some brown speckling, bands medium brown dorsally, lighter brown and more speckled on sides of body. Venter pale yellowish cream color with brown speckling from underside of head to about one-third down body, paler than rest of venter. Middle third of venter yellowish grey with more brown speckling, posterior third of venter and anterior half of tail very pale orange cream color with brown speckling. Irregularly spaced small dark brown patches throughout venter. Bands do not continue across venter. Ventral surface of posterior half of tail is a nearly continuous irregular band of medium to light brown with some lighter speckling, thinner where interspaces meet venter. Distribution and ecology. The holotype was found inside an arboreal bromeliad in broadleaf cloud forest with some pine trees. The type locality at La Montaña de las Granadillas is part of a belt of mid-elevation cloud forest stretching along a ridge to the vicinity of La Unión, Zacapa. Although the eastern part of this forest, near La Unión, has been sampled fairly extensively without the discovery of any additional specimens, we expect that this species occurs in other portions of the forested ridge. The forest where La Montaña de las Granadillas is located is separated from other areas of cloud forest by more arid lowland areas, including the arid Motagua Valley to the north. Given the isolated nature of this cloud forest, it seems unlikely that the species is found in other cloud forests in eastern Guatemala. Etymology. The specific epithet makes reference to the Sierra del Merendón, the mountain range containing the type locality.

A new species of snake of the genus Sibon (Squamata: Colubridae) is described from La Montaña de las Granadillas, partof the Sierra del Merendón in eastern Guatemala. Sibon merendonensis is part of the S. annulatus group, and differs fromother members of the group in its scale row formula, head scalation and body size. The new species is known only from the holotype and was collected from an isolated area of cloud forest near Zacapa, Department of Zacapa, Guatemala.

Lavin, Brian R., Papenfuss, Theodore J. (2012): The phylogenetic position of Chalcides ocellatus (Squamata: Scincidae) from Yemen and Somalia. Zootaxa 3221 (1): 26-36, DOI: 10.11646/zootaxa.3221.1.2, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.3221.1.2

Oedipina nica sp. nov. Nicaraguan Highland Worm Salamander Figure 1 Oedipina cyclocauda (in part). K��hler 1999, 2001. Holotype. MVZ 263774, an adult male, from El Gobiado, Reserva Natural Cerro Datanl��-El Diablo, 13 ��09���N, 85 �� 52 ���W, 1420 m above sea level (a.s.l.), Dept. Jinotega, Nicaragua, collected 30 August 2009 by S. M. Rovito, T. J. Papenfuss, J. Sunyer, and L. A. Obando (original field number SMR 842). Paratypes (5). UF 156445, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156446 ��� 47, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156453 ��� 54, from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Referred specimens (12). SMF 78736, from Camp El Hielo, Reserva Natural Cerro Kilamb��, 1490 m a.s.l., Dept. Jinotega, Nicaragua; SMF 78737, 78739��� 40, from Camp 2, Reserva Natural Cerro Kilamb��, 13 �� 35.25 'N, 85 �� 41.50 'W, 1360 m a.s.l., Dept. Jinotega, Nicaragua; UF 156443 ��� 44, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156448 ��� 50, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156451 ��� 52, two tails dropped by escaped adults from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1600 m a.s.l., Dept. Jinotega, Nicaragua; UF 156455 from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Diagnosis. A slender, small-sized (largest specimen 48.5 mm SL) species assigned to the genus Oedipina based on the presence of a high number (19���20) of costal grooves between the short limbs, very narrow manus and pes, long tail (tail length ca 2 x SL), overall morphological similarity to other Oedipina species, and molecular phylogenetic evidence. This species is a member of the subgenus Oeditriton (based on molecular data; Fig. 2) and is distinguished from the only other members of that clade (O. kasios and O. quadra) by its more slender habitus and fewer vomerine teeth. Further distinguished from other Nicaraguan members of the genus Oedipina as follows: from O. collaris by being much smaller and more slender, having short limbs and narrow manus and pes, and having a short, rather bluntly tipped snout; from O. cyclocauda by being smaller and more slender without nearly pad-like feet; from O. pseudouniformis by being more slender with shorter, less robust limbs and narrower manus and pes, and by having almost uniformly dark brownishblack coloration with no distinct light pigmentation on the upper parts of the proximal segment of the limbs. Description of the holotype. The male holotype is 39.2 mm SL and is judged to be an adult by the presence of small papillae in the anterior margins of the vent and a small mental gland behind the anterior tip of the mandible. The small head is rounded in dorsal profile, and somewhat flattened in lateral profile with a relatively blunt snout and distinct nasolabial protuberances. Snout to posterior angle of vent 9.8 times head width and 6.5 times head length. Nostrils are small but conspicuous under magnification. The small eyes are inconspicuous and do not extend beyond the lateral margins of the head. The suborbital groove does not intercept the lip line. The skin of the dorsal surface of the head behind the eyes is depressed in a pattern that suggests the presence of a frontoparietal fenestra. There is a single slightly enlarged premaxillary tooth that is located distinctly anterior to the maxillary teeth. Maxillary teeth total 41. The small vomerine teeth are in a short, fairly straight series and total 11. There are 20 costal grooves between the small limbs with a limb interval of 13 intercostal folds. Hands and feet are tiny, narrow, and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with only a part of a phalangeal segment of digit III free between digits II���III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs; both outer digits are extremely short on the hind limbs. The triangular tip of digit III of the hind limb is distinct from the tiny rounded tips of digits II and IV, but less than a full phalangeal segment of that digit is free of the others. Subdigital pads are nearly imperceptible. Digits on forelimbs in order of decreasing length are III���II���IV���I; digits on hind limbs are III���II���IV���(V���I). The tail is round in cross section and tapers gradually to a narrow tip; the tail is almost twice SL. Measurements (in millimeters) of the holotype. Head width 4.0; snout to gular fold (head length) 6.0; head depth at posterior angle of jaw 1.8; eyelid width 1.1; eyelid length 1.5; eye to nostril 0.6; anterior rim of eye to snout 1.2; horizontal orbital diameter 1.0; interorbital distance 1.7; distance separating eyelids 0.7; nostril diameter 0.2; snout projecting beyond mandible 0.4; distance from eye to distal end of postorbital groove 1.9; snout to posterior angle of vent (SL) 39.2; snout to anterior angle of vent 37.0; snout to forelimb 9.4; axilla to groin 26.0; limb interval 13; shoulder width 2.7; tail length 77.0; tail width at base 2.4; tail depth at base 2.4; forelimb length (to tip of longest digit) 3.7; hind limb length (to tip of longest digit) 4.5; forelimb foot width 0.6; hind limb foot width 0.9; free length of longest digit 0.2. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are black. Costal grooves on the body and tail are the same color as the body or slightly lighter dark gray, but at the deepest part of the groove pigment is lacking so the grooves are prominent. Ventral surfaces of the head, body, and tail are slightly paler black than those dorsal surfaces. Dorsal and ventral surfaces of the limbs are the same as for the body. Tiny white speckles are most visible behind the eye and over the shoulder, then in an irregular, narrow line along the dorsolateral part of the body to the hind limbs, but they are few in number and obscure. White spots are a little larger in the gular and chest areas ventrally. The nasolabial protuberances are unpigmented and contrast with the surrounding tissue. Variation. Variation based on study of the holotype and three male (UF 156445, 156447, 156453), one female (UF 156446), and one juvenile (UF 156454) paratypes: the four males, judged to be mature based on having inconspicuous mental glands and anterior vent margins bearing fine pinnae, range from 38.5���44.5 SL; the female is 48.5 SL. Tails are present in the holotype, three male paratypes, and the juvenile and all taper gradually to slender tips. SL is from 0.86 (adult) to 0.52 (holotype) times tail length. SL in males is 9.8���11.8 times head width and 9.9 in the single female. Limbs are relatively short; number of costal folds between adpressed limbs is 13���14 in males and 13 in the female. There are 19���20 costal grooves between the limbs in the males and 20 in the female. Feet are narrow; 0.8���1.1. The digits are joined to their neighbors, with only the tips of the longest toes free. Males have 1���2 premaxillary teeth, the female has 4. Maxillary teeth range from 41���47 in males; 48 in the female. Males have 9���16 vomerine teeth; the female 18. If one adds the referred specimens, the size range of the adult males increases to 35.3���48.5 SL, and that of females from 37.5���48.5 SL. These specimens are consistent in measurements, tooth and costal groove counts, and coloration with the type series. SMF 78738, from Reserva Natural Cerro Kilamb�� (44.0 SL), fits within the range of the new species in all measurements (see discussion). Variation of coloration based on color photographs taken in life during daytime. Coloration is in general uniformly dark brownish-black with no distinguishing features. The upper side of the proximal segment of the limbs is the same color. The costal grooves are very evident because of the lack of pigment at their centers. Details for particular individuals follow. Adult male paratype (UF 156447; Fig. 1 b): Dorsum of body and tail blackish brown becoming a faded lighter brown on the dorsal and lateral surfaces of head extending to the chin. A lighter brown speckling is faintly noticeable dorsally from the head becoming imperceptible towards the base of the tail. Ventral surfaces of the body are blackish brown, similar in coloration to the tail. Limbs faded brown similar to color of head. Skin creases such as costal grooves, gular fold, and those around the paratoid region lack pigmentation and appear light grey. Iris dark rust brown. Adult male (UF 156450): Dorsum of body and tail blackish brown becoming faded lighter brown on the anterior and lateral surfaces of head. Gold speckling mixed with some white is located on the dorsal surface of the head and body. Speckling is most prominent on the top of the head becoming sparser posteriad, not extending beyond base of tail. Dorsolaterally above the anterior limbs a slight concentration of tiny white speckles forms a short, irregular line. Below the extent of the speckling ventrolaterally coloration is same as that of tail. Limb coloration is same as that of anterior and lateral surface of head. Iris dark rust brown. Juvenile paratype (UF 156454; Fig. 1 d): Dorsal and lateral surfaces of body, head, tail, and limbs are a uniform jet black. There is a faint overlay of tiny dirty white flecks along the dorsum of head, body, and tail giving a somewhat ashy appearance. This flecking is most concentrated dorsolaterally giving the impression of an irregular line extending from behind the jaw to the hind limb. Flecking is also present around the mouth and chest area. Larger white speckles are mixed sparsely and intermittently along the head, body, tail, and limbs. Habitat and distribution. This species is known to occur from 1360���1660 m elevation at three isolated localities in the Lower Montane Moist Forest formation (Holdridge 1967) in north-central Nicaragua (Fig. 3). The holotype was collected in a small patch of secondary cloud forest (Fig. 4 a���b) mostly surrounded by cattle pastures and agricultural land. It was found while raking during the daytime inside a moderately large rotten log. The paratypes and UF referred specimens from Reserva Natural Cerro Kilamb�� were collected in undisturbed, primary cloud forest (Fig. 4 c) during both diurnal and nocturnal surveys. All were found underneath logs, rocks, and other debris in areas along the mountain ridge where the substrate comprised a spongy mass of plant roots and mosses. The SMF referred specimens from Reserva Natural Cerro Kilamb�� were also collected in undisturbed, primary cloud forest during diurnal surveys. Most of them were found inside dead basal ramifications of giant ferns. The paratypes from Reserva Natural Macizos de Pe��as Blancas were collected in undisturbed, primary cloud forest (Fig. 4 d). One was uncovered during the daytime beneath a fallen log, and two others were found active on the forest floor around the campsite at 2130 h. Those found active within the camp were likely dislodged or disturbed by raking and uprooting logs that occurred during set-up of the campsite earlier the same day. Phylogenetic relationships. Phylogenetic analysis of sequence data from two mitochondrial genes (16 S and cyt b) confirms the monophyly of this taxon (ML bootstrap support = 99, Bayesian posterior probability = 1.0) and its position as sister species to Oedipina kasios in the Oeditriton clade (Fig. 2). A sister-species relationship between O. kasio s and O. nica is strongly supported (ML bootstrap support = 93, Bayesian posterior probability = 0.99) as is the monophyly of an Oeditriton clade containing O. kasio s, O. nica, and O. quadra (ML bootstrap support = 96, Bayesian posterior probability = 1.0). Oeditriton is the sister clade to the remaining Oedipina, which are divided into the subgenera Oedipina and Oedopinola (Fig. 2), which supports previous phylogenetic hypotheses for Oedipina (Garc��a-Par��s & Wake 2000; McCranie et al. 2008). All wellsupported phylogenetic relationships are congruent between the ML and Bayesian results. Etymology. The specific name nica is a short name for Nicaraguan (Nicarag��ense in Spanish) and is a colloquial word used in Central America to denote Nicaraguan people. The name alludes to the country of origin of the type specimens. Conservation status. Using the criteria established by IUCN for evaluating threatened species, Oedipina nica should be classified as Endangered (EN B 2 ab[iii]), due to its limited distribution (known only from three isolated highland forest areas with a total extent of less than 500 km 2) and the continued loss of habitat at these localities., Published as part of Sunyer, Javier, Wake, David B., Townsend, Josiah H., Travers, Scott L., Rovito, Sean M., Papenfuss, Theodore J., Obando, Lenin A. & K��hler, Gunther, 2010, A new species of worm salamander (Caudata: Plethodontidae: Oedipina) in the subgenus Oeditriton from the highlands of northern Nicaragua, pp. 29-39 in Zootaxa 2613 on pages 32-35, DOI: 10.5281/zenodo.197858, {"references":["Kohler, G. (1999) The amphibians and reptiles of Nicaragua - a distributional checklist with keys. Courier Forschungsinstut Senckenberg, 213, 1 - 212.","Kohler, G. (2001) Anfibios y Reptiles de Nicaragua. Herpeton. Offenbach, Germany.","Holdridge, L. R. (1967) Life Zone Ecology. Tropical Science Center. San Jose, Costa Rica.","McCranie, J. R., Vieites, D. R. & Wake, D. B. (2008) Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae). Zootaxa, 1930, 1 - 17."]}

Oedipina nica sp. nov. Nicaraguan Highland Worm Salamander Figure 1 Oedipina cyclocauda (in part). Köhler 1999, 2001. Holotype. MVZ 263774, an adult male, from El Gobiado, Reserva Natural Cerro Datanlí-El Diablo, 13 °09’N, 85 ° 52 ’W, 1420 m above sea level (a.s.l.), Dept. Jinotega, Nicaragua, collected 30 August 2009 by S. M. Rovito, T. J. Papenfuss, J. Sunyer, and L. A. Obando (original field number SMR 842). Paratypes (5). UF 156445, from Reserva Natural Cerro Kilambé, 13 ° 34 ’N, 85 ° 42 ’W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156446 – 47, from Reserva Natural Cerro Kilambé, 13 ° 35 ’N, 85 ° 43 ’W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156453 – 54, from Reserva Natural Macizos de Peñas Blancas, 13 ° 17 ’N, 85 ° 43 ’W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Referred specimens (12). SMF 78736, from Camp El Hielo, Reserva Natural Cerro Kilambé, 1490 m a.s.l., Dept. Jinotega, Nicaragua; SMF 78737, 78739– 40, from Camp 2, Reserva Natural Cerro Kilambé, 13 ° 35.25 'N, 85 ° 41.50 'W, 1360 m a.s.l., Dept. Jinotega, Nicaragua; UF 156443 – 44, from Reserva Natural Cerro Kilambé, 13 ° 34 ’N, 85 ° 42 ’W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156448 – 50, from Reserva Natural Cerro Kilambé, 13 ° 35 ’N, 85 ° 43 ’W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156451 – 52, two tails dropped by escaped adults from Reserva Natural Cerro Kilambé, 13 ° 34 ’N, 85 ° 42 ’W, 1600 m a.s.l., Dept. Jinotega, Nicaragua; UF 156455 from Reserva Natural Macizos de Peñas Blancas, 13 ° 17 ’N, 85 ° 43 ’W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Diagnosis. A slender, small-sized (largest specimen 48.5 mm SL) species assigned to the genus Oedipina based on the presence of a high number (19–20) of costal grooves between the short limbs, very narrow manus and pes, long tail (tail length ca 2 x SL), overall morphological similarity to other Oedipina species, and molecular phylogenetic evidence. This species is a member of the subgenus Oeditriton (based on molecular data; Fig. 2) and is distinguished from the only other members of that clade (O. kasios and O. quadra) by its more slender habitus and fewer vomerine teeth. Further distinguished from other Nicaraguan members of the genus Oedipina as follows: from O. collaris by being much smaller and more slender, having short limbs and narrow manus and pes, and having a short, rather bluntly tipped snout; from O. cyclocauda by being smaller and more slender without nearly pad-like feet; from O. pseudouniformis by being more slender with shorter, less robust limbs and narrower manus and pes, and by having almost uniformly dark brownishblack coloration with no distinct light pigmentation on the upper parts of the proximal segment of the limbs. Description of the holotype. The male holotype is 39.2 mm SL and is judged to be an adult by the presence of small papillae in the anterior margins of the vent and a small mental gland behind the anterior tip of the mandible. The small head is rounded in dorsal profile, and somewhat flattened in lateral profile with a relatively blunt snout and distinct nasolabial protuberances. Snout to posterior angle of vent 9.8 times head width and 6.5 times head length. Nostrils are small but conspicuous under magnification. The small eyes are inconspicuous and do not extend beyond the lateral margins of the head. The suborbital groove does not intercept the lip line. The skin of the dorsal surface of the head behind the eyes is depressed in a pattern that suggests the presence of a frontoparietal fenestra. There is a single slightly enlarged premaxillary tooth that is located distinctly anterior to the maxillary teeth. Maxillary teeth total 41. The small vomerine teeth are in a short, fairly straight series and total 11. There are 20 costal grooves between the small limbs with a limb interval of 13 intercostal folds. Hands and feet are tiny, narrow, and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with only a part of a phalangeal segment of digit III free between digits II–III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs; both outer digits are extremely short on the hind limbs. The triangular tip of digit III of the hind limb is distinct from the tiny rounded tips of digits II and IV, but less than a full phalangeal segment of that digit is free of the others. Subdigital pads are nearly imperceptible. Digits on forelimbs in order of decreasing length are III–II–IV–I; digits on hind limbs are III–II–IV–(V–I). The tail is round in cross section and tapers gradually to a narrow tip; the tail is almost twice SL. Measurements (in millimeters) of the holotype. Head width 4.0; snout to gular fold (head length) 6.0; head depth at posterior angle of jaw 1.8; eyelid width 1.1; eyelid length 1.5; eye to nostril 0.6; anterior rim of eye to snout 1.2; horizontal orbital diameter 1.0; interorbital distance 1.7; distance separating eyelids 0.7; nostril diameter 0.2; snout projecting beyond mandible 0.4; distance from eye to distal end of postorbital groove 1.9; snout to posterior angle of vent (SL) 39.2; snout to anterior angle of vent 37.0; snout to forelimb 9.4; axilla to groin 26.0; limb interval 13; shoulder width 2.7; tail length 77.0; tail width at base 2.4; tail depth at base 2.4; forelimb length (to tip of longest digit) 3.7; hind limb length (to tip of longest digit) 4.5; forelimb foot width 0.6; hind limb foot width 0.9; free length of longest digit 0.2. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are black. Costal grooves on the body and tail are the same color as the body or slightly lighter dark gray, but at the deepest part of the groove pigment is lacking so the grooves are prominent. Ventral surfaces of the head, body, and tail are slightly paler black than those dorsal surfaces. Dorsal and ventral surfaces of the limbs are the same as for the body. Tiny white speckles are most visible behind the eye and over the shoulder, then in an irregular, narrow line along the dorsolateral part of the body to the hind limbs, but they are few in number and obscure. White spots are a little larger in the gular and chest areas ventrally. The nasolabial protuberances are unpigmented and contrast with the surrounding tissue. Variation. Variation based on study of the holotype and three male (UF 156445, 156447, 156453), one female (UF 156446), and one juvenile (UF 156454) paratypes: the four males, judged to be mature based on having inconspicuous mental glands and anterior vent margins bearing fine pinnae, range from 38.5–44.5 SL; the female is 48.5 SL. Tails are present in the holotype, three male paratypes, and the juvenile and all taper gradually to slender tips. SL is from 0.86 (adult) to 0.52 (holotype) times tail length. SL in males is 9.8–11.8 times head width and 9.9 in the single female. Limbs are relatively short; number of costal folds between adpressed limbs is 13–14 in males and 13 in the female. There are 19–20 costal grooves between the limbs in the males and 20 in the female. Feet are narrow; 0.8–1.1. The digits are joined to their neighbors, with only the tips of the longest toes free. Males have 1–2 premaxillary teeth, the female has 4. Maxillary teeth range from 41–47 in males; 48 in the female. Males have 9–16 vomerine teeth; the female 18. If one adds the referred specimens, the size range of the adult males increases to 35.3–48.5 SL, and that of females from 37.5–48.5 SL. These specimens are consistent in measurements, tooth and costal groove counts, and coloration with the type series. SMF 78738, from Reserva Natural Cerro Kilambé (44.0 SL), fits within the range of the new species in all measurements (see discussion). Variation of coloration based on color photographs taken in life during daytime. Coloration is in general uniformly dark brownish-black with no distinguishing features. The upper side of the proximal segment of the limbs is the same color. The costal grooves are very evident because of the lack of pigment at their centers. Details for particular individuals follow. Adult male paratype (UF 156447; Fig. 1 b): Dorsum of body and tail blackish brown becoming a faded lighter brown on the dorsal and lateral surfaces of head extending to the chin. A lighter brown speckling is faintly noticeable dorsally from the head becoming imperceptible towards the base of the tail. Ventral surfaces of the body are blackish brown, similar in coloration to the tail. Limbs faded brown similar to color of head. Skin creases such as costal grooves, gular fold, and those around the paratoid region lack pigmentation and appear light grey. Iris dark rust brown. Adult male (UF 156450): Dorsum of body and tail blackish brown becoming faded lighter brown on the anterior and lateral surfaces of head. Gold speckling mixed with some white is located on the dorsal surface of the head and body. Speckling is most prominent on the top of the head becoming sparser posteriad, not extending beyond base of tail. Dorsolaterally above the anterior limbs a slight concentration of tiny white speckles forms a short, irregular line. Below the extent of the speckling ventrolaterally coloration is same as that of tail. Limb coloration is same as that of anterior and lateral surface of head. Iris dark rust brown. Juvenile paratype (UF 156454; Fig. 1 d): Dorsal and lateral surfaces of body, head, tail, and limbs are a uniform jet black. There is a faint overlay of tiny dirty white flecks along the dorsum of head, body, and tail giving a somewhat ashy appearance. This flecking is most concentrated dorsolaterally giving the impression of an irregular line extending from behind the jaw to the hind limb. Flecking is also present around the mouth and chest area. Larger white speckles are mixed sparsely and intermittently along the head, body, tail, and limbs. Habitat and distribution. This species is known to occur from 1360–1660 m elevation at three isolated localities in the Lower Montane Moist Forest formation (Holdridge 1967) in north-central Nicaragua (Fig. 3). The holotype was collected in a small patch of secondary cloud forest (Fig. 4 a–b) mostly surrounded by cattle pastures and agricultural land. It was found while raking during the daytime inside a moderately large rotten log. The paratypes and UF referred specimens from Reserva Natural Cerro Kilambé were collected in undisturbed, primary cloud forest (Fig. 4 c) during both diurnal and nocturnal surveys. All were found underneath logs, rocks, and other debris in areas along the mountain ridge where the substrate comprised a spongy mass of plant roots and mosses. The SMF referred specimens from Reserva Natural Cerro Kilambé were also collected in undisturbed, primary cloud forest during diurnal surveys. Most of them were found inside dead basal ramifications of giant ferns. The paratypes from Reserva Natural Macizos de Peñas Blancas were collected in undisturbed, primary cloud forest (Fig. 4 d). One was uncovered during the daytime beneath a fallen log, and two others were found active on the forest floor around the campsite at 2130 h. Those found active within the camp were likely dislodged or disturbed by raking and uprooting logs that occurred during set-up of the campsite earlier the same day. Phylogenetic relationships. Phylogenetic analysis of sequence data from two mitochondrial genes (16 S and cyt b) confirms the monophyly of this taxon (ML bootstrap support = 99, Bayesian posterior probability = 1.0) and its position as sister species to Oedipina kasios in the Oeditriton clade (Fig. 2). A sister-species relationship between O. kasio s and O. nica is strongly supported (ML bootstrap support = 93, Bayesian posterior probability = 0.99) as is the monophyly of an Oeditriton clade containing O. kasio s, O. nica, and O. quadra (ML bootstrap support = 96, Bayesian posterior probability = 1.0). Oeditriton is the sister clade to the remaining Oedipina, which are divided into the subgenera Oedipina and Oedopinola (Fig. 2), which supports previous phylogenetic hypotheses for Oedipina (García-París & Wake 2000; McCranie et al. 2008). All wellsupported phylogenetic relationships are congruent between the ML and Bayesian results. Etymology. The specific name nica is a short name for Nicaraguan (Nicaragüense in Spanish) and is a colloquial word used in Central America to denote Nicaraguan people. The name alludes to the country of origin of the type specimens. Conservation status. Using the criteria established by IUCN for evaluating threatened species, Oedipina nica should be classified as Endangered (EN B 2 ab[iii]), due to its limited distribution (known only from three isolated highland forest areas with a total extent of less than 500 km 2) and the continued loss of habitat at these localities.

Paramesotriton ermizhaoi species nov. (Fig. 3 ABC) Holotype: CIB 88141, an adult male collected in Mt. Dayao (24 ��07��N, 110 �� 13��E, 881m elevation), Jinxiu Yao Autonomous County, Guangxi Zhuang Autonomous Region, P. R. China; collected by Jiatang Li on July 7, 2006. Paratypes: Same locality as holotype: CIB 88140, CIB 95998 -96000, collected with the holotype on the same date; MVZ 230616 - 230621, collected by E. Zhao in March 1999. Diagnosis: Paramesotriton ermizhaoi is assigned to the genus Paramesotriton because of its slender and nearly straight epibranchial bones, maxillary bones oriented angular to the body axis, and laterally compressed tail. This species can be diagnosed from other congeners by the following combination of characters: skin relatively smooth; granular warts absent on head and body; vertebral ridge not prominent; head depressed and nearly flat in profile; habitus slender and depressed; limbs short���when adpressed against flank, fingers and toes hardly meet; dorsum olive brown with irregular black flecks; ventral orange-red blotches irregular in shape. Description of the holotype: A slender and flat newt, preserved with mouth open. Head oval in shape, depressed and nearly flat in profile. Snout truncate and slightly projecting beyond lower mandible. Nostrils close to snout tip. Labial fold evident throughout posterior half of upper jaw. Skull relatively narrow, with maxillary bones oriented angular to body axis. Maxillary tips lie anterior and lateral to pterygoid bones; no contact between maxillary and pterygoid. Fronto-squamosal arch complete but not very robust. Epibranchial bones slender, nearly straight, only slightly flared dorsally. Vomerine tooth patch ��-shaped, tooth rows converging anteriorly and exceeding the anterior limit of choanae. Tongue adhering to mouth floor with free lateral margins. Parotoid region evident, but not enlarged. Gill filaments absent. Gular fold present. Skin relatively smooth on head and body, with small transverse wrinkles. Vertebral ridge thin, slightly elevated. Lateral dorsal ridge absent. Thirteen trunk vertebrae. Four fingers, five toes, without webbing. Tail laterally compressed; posterior dorsal caudal fin evident, ventral caudal fin indistinct; tail extremity round in profile. Cloaca swollen, with a few papillae near posterior limit. Color of holotype in life: Dorsum olive brown with irregular black flecks. Vertebral ridge with orange tint. Venter paler brown. Irregular orange-red blotches on venter and anterior portion of cloaca with indistinct black margin; some blotches connected into short irregular stripes. Orange-red on underside of tail. In preservative, dorsum and venter black. Vertebral ridge inconspicuous. Ventral bright orange fading to yellow to milky white; black margins invisible. Variation: Morphology of paratypes resembles that of holotype except that females have smaller and nonswollen cloaca without papillae. Linear measurements are summarized in Table 2. Color of ventral blotches ranges from orange to red, spreading to chin and underside of axillae, varying in shape and arrangement. Black marks border those blotches or intersperse on venter. Holotype Females (N = 6) Males (N = 5) Measurements (CIB 88141) Range Mean �� SE Range Mean �� SE SVL 63.5 46.6���66.8 56.3 �� 2.7 46.4���63.5 56.1 �� 3.1 TTL 126.0 94.0��� 137.5 112.8 �� 5.9 92.2���127.6 111.5 �� 6.8 TAL 57.7 44.7���65.5 53.4 �� 3.0 43.3���60.2 51.1 �� 3.3 TAD 7.5 5.5���8.1 7.0 �� 0.4 5.5���8.5 7.3 �� 0.5 HL 17.7 14.4���17.4 15.9 �� 0.5 14.8���17.8 16.1 �� 0.7 HW 12.1 8.8���12.6 10.7 �� 0.5 9.1���12.3 10.9 �� 0.6 IO 7.1 5.3���6.9 6.0 �� 0.2 5.4���7.1 6.4 �� 0.3 EN 4.9 3.2���4.2 3.8 �� 0.2 3.1���4.9 4.0 �� 0.3 IN 3.7 2.7���3.6 3.1 �� 0.1 2.4���3.7 3.1 �� 0.2 AX 31.3 20.2���36.7 27.7 �� 2.2 21.4���31.3 27.6 �� 1.7 AL 15.6 12.6���15.4 13.8 �� 0.4 12.5���17.4 14.6 �� 0.9 PL 16.0 13.1���15.6 14.2 �� 0.4 13.5���17.4 15.2 �� 0.7 Etymology: The new species is named after Ermi Zhao, a prominent Chinese herpetologist and educator, for his great contribution to the development of herpetological study and the training of a new generation of scientists in China. Habitat: Stream in broadleaf forest with herbaceous plants and vines. Stream is 3-4 meters wide and shallow, flowing slowly in a valley (Fig. 3 C). Substrates include gravels, scattered small rocks, and semisubmerged larger rocks. Along the stream are pools with a very slow current. Newts are found at the stream bottom, usually under rocks or between crevices during daytime. Fishes and small aquatic invertebrates coexist with Paramesotriton ermizhaoi, and Pachytriton labiatus is found in the same stream drainage at a higher elevation. Discussion: Paramesotriton ermizhaoi is unusual insofar as a large portion of individuals possess 13 trunk vertebrae; other congeneric species normally have 12 trunk vertebrae (Chan et al. 2001). Although variation has been observed in other Paramesotriton, it is rare. However, six of eleven P. ermizhaoi (including the holotype) have 13 trunk vertebrae (Fig. 4); the other five specimens have 12. The number of trunk vertebrae does not appear to be correlated with gender. A similar situation occurs in the European salamandrid Ommatotriton ophryticus, in which the modal number of trunk vertebrae varies from 12 to 13 in different geographic populations (Litvinchuk et al. 2005). Orska and Imiolek (1962) reported the correspondence of vertebral number to developmental temperature in salamanders. Future research is necessary to evaluate the correlation between environmental temperature and embryonic development in P. ermizhaoi. Paramesotriton ermizhaoi may have been long misidentified as another Asian salamandrid species. Scholz (1998) reported a potentially new species of Pachytriton from the pet trade, named Pachytriton C. In fact, most descriptions of morphology, coloration, and behavior of Pachytriton C match with P. e r m i z h a o i. Due to lack of molecular data and known locality, Scholz (1998) did not make a definitive taxonomic statement on Pachytriton C. We suspect that the two names refer to the same species. The only known locality of P. ermizhaoi, Mt. Dayao, is located within the putative Guangxi population of P. c h i n e n s i s (Fei et al. 1999; Zhang & Wen 2000; Fei et al. 2006), which was first described in rivers at inland from Ningbo, Zhejiang Province (Gray 1859). However, P. ermizhaoi has been identified mistakenly as P. chinensis despite their different morphologies. For instance, the paratypes of P. e r m i z h a o i from MVZ were originally catalogued as P. c h i n e n s i s. We also note that the P. chinensis used by Lu et al. (2004) from Mt. Dayao actually are P. ermizhaoi, based on mitochondrial sequences. To our knowledge, no true P. c h i n e n s i s has been collected from Mt. Dayao. It is possible that claims of a disjunct Guangxi population of P. chinensis are incorrect due to misidentification. However, Zhang and Wen (2000) provide an account on Guangxi P. chinensis that matches the diagnostic characters of this species. More fieldwork is needed to determine if P. chinensis occurs in Guangxi Zhuang Autonomous Region. Comparative material examined: P. caudopunctatus (MVZ 236250-236254 from Guizhou, China), P. chinensis (MVZ 230360, CIB 95899, CIB 95907 -95911 from Zhejiang, China), P. fuzhongensis (MVZ 230622 - 230625 from Guangxi Zhuang Autonomous Region, China), P. hongkongensis (MVZ 230365 - 230370 from Hong Kong, China), P. deloustali (MVZ 222122-222123, MVZ 223627-223629 from Tam Dao, Vietnam), P. laoensis (FMNH 255450, FMNH 257850, FMNH 257852, FMNH 257853 from Xiang Khouang, Laos)., Published as part of Wu, Yunke, Rovito, Sean M., Papenfuss, Theodore J. & Hanken, James, 2009, A new species of the genus Paramesotriton (Caudata: Salamandridae) from Guangxi Zhuang Autonomous Region, southern China, pp. 59-68 in Zootaxa 2060 on pages 64-66, DOI: 10.5281/zenodo.186817, {"references":["Chan, L. M., Zamudio, K. R. & Wake, D. B. (2001) Relationships of the salamandrid genera Paramesotriton, Pachytriton, and Cynops based on mitochondrial DNA sequences. Copeia, 4, 997 - 1009.","Litvinchuk, S. N., Zuiderwijk, A., Borkin, L. J. & Rosanov, J. M. (2005) Taxonomic status of Triturus vittatus (Amphibia: Salamandridae) in western Turkey: trunk vertebrae count, genome size and allozyme data. Amphibia - Reptilia, 26, 305 - 323.","Orska, J. & Imiolek, Z. (1962) Preliminary studies on the effect of temperature on the development of meristic characters in the Urodela. Acta Universitatis Wratislaviensis, 3, 134 - 154.","Scholz, K. P. (1998) Uber eine rauhhautige Pachytriton-Art. Salamandra, 34, 375 - 380.","Fei, L., Ye, C., Huang, Y. & Liu, M. (1999) Atlas of Amphibians of China. Henan Science and Technical Press. Zhengzhou, 432 pp (in Chinese).","Zhang, Y. & Wen, Y. (2000) Amphibians in Guangxi. Guangxi Normal University Press, Guilin, 183 pp (in Chinese).","Fei, L., Hu, S., Ye, C. & Huang Y. (2006) Fauna Sinica, Amphibia Vo l. 1. Beijing: Science Press. 470 pp (in Chinese).","Gray, J. E. (1859) Description of new species of salamanders from China and Siam. Proceedings of the Zoological Society of London, 1859: 229 - 230.","Lu, S. Q., Yuan, Z. G., Pang, J. F., Yang, D. T., Yu, F. H., McGuire, P., Xie, F. & Zhang, Y. P. (2004) Molecular phylogeny of the genus Paramesotriton (Caudata: Salamandridae). Biochemical Genetics, 42, 139 - 148."]}