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Hynobius akiensis sensu lato has recently been split into three species based on short sequence analyses of cyt-b gene of mtDNA and without data of nuclear DNA, and strange sympatric distribution in some areas has been indicated in two species. We analyzed nuclear DNA marker (SNPs) and complete sequence of cyt-b in H. akiensis sensu lato to reassess species delimitation and genetic introgression among species. As a result, we found two lineages with discordant mitochondrial and nuclear DNA in some areas. Of H. akiensis sensu lato, each of the two contains the type locality of two species recently reported (H. sumidai and H. geiyoensis), and the use of these names has been previously advocated. However, their sympatric distribution was rejected based on nuclear DNA data, which we consider is more reliable than mtDNA. We thus clarify geographic boundary of these two species and revise the species delimitations.

A new salamander species of the genus Bolitoglossa is here described from the cloud forests of the western slopes of the Cordillera Oriental of Colombia, in the Cundinamarca department. The most salient characters of this new species are its numerous maxillary and vomerine teeth, its moderate webbing on hands and feet, its short and robust tail, and its chromatic variation. Based on molecular analyses this new species is assigned to the adspersa species group and its status established as the sister species of B. adspersa, with which it was previously confused. Lastly, the distribution, natural history, and conservation status of the new species are discussed.

For approximately four decades, scientists have known of the existence of several undescribed species of Siren in the southeastern United States Coastal Plain. One of these species, S. reticulata, was recently described, but a small, seepage-dwelling species has remained undescribed until now. To resolve outstanding questions concerning the phylogeny of Siren, we collected sequence and morphometric data from specimens across the range of Siren. We found S. lacertina and S. reticulata to represent strongly supported monophyletic groups, with S. reticulata having a sister relationship to all other Siren. Additionally, we found five distinct mtDNA lineages within what has been recognized as S. intermedia. Siren lacertina and type-locality S. intermedia (lineage A) are sister mtDNA lineages, whereas S. intermedia lineages B and C show a high level of mitogenomic divergence from type-locality S. intermedia. Analyses of two scnDNA loci revealed that S. lacertina is monophyletic but nested with low positional support in a clade including the three S. intermedia mtDNA lineages. Further study is needed to determine whether S. intermedia lineages A, B, and C represent distinct species or incompletely sorted lineages. We restrict the range of S. intermedia to the region from the Escambia and Perdido river drainages of Florida and Alabama eastward through Virginia (the combined ranges of lineages A, B, and C). We also elevate S. i. nettingi (lineage E) to species status and include the larger S. i. texana form in that taxon, generating a species that occurs from the Mobile Bay drainages westward through the Mississippi Basin and southwest into northeastern Mexico. Lastly, we describe a new miniature species, S. sphagnicola, that ranges from the Florida Parishes of Louisiana eastward to the westernmost tributary creeks of Choctawhatchee Bay in the western Florida panhandle.

We describe a new species of plethodontid salamander of the genus Pseudoeurycea from the Sierra de Zongolica, Veracruz, Mexico. The new species is distinguished from all other species in the genus by morphological and genetic features and by coloration. Based on a mtDNA phylogeny, the new species belongs to the Pseudoeurycea juarezi group and is most closely related to P. ruficauda from the Sierra Mazateca in northern Oaxaca. The newly described salamander increases the number of species of plethodontid salamanders from Veracruz to 43 and those recognized from Mexico to 140.

A crocodile newt, Echinotriton raffaellii, recently described from the Amami islands in the Ryukyu archipelago, western Japan has been regarded as Amami lineage of E. andersoni from the Okinawa islands, to which it is morphologically very similar, but genetically divergent. In the original description of E. raffaellii, only two specimens, including one born in captivity, were examined and only limited morphological comparisons with E. andersoni were made. Here we examined the morphological characteristics of E. raffaellii with an ample set of specimens, compared them with those of E. andersoni, and clarified their independent specific statuses.

The tiny moss salamanders of the genus Nototriton within Costa Rica have long been poorly known and understood. Due to their diminutive size and specialized ecology, this taxon is very rarely studied or even represented in collections, thus resulting in much confusion surrounding the actual taxonomic and distributional status of the species within the country. As a result of hundreds of hours of fieldwork, lab work, and morphological analysis and measurement, herein we provide an updated taxonomic revision of the Costa Rican members of the genus Nototriton, with the proposal of a new subgenus and four new species.

A new species of the genus Tylototriton is described from Ngoc Linh Mountain, Kon Tum Province, in the Central Highlands of Vietnam based on integrative taxonomy, namely by combining molecular and morphological evidence. Tylototriton ngoclinhensis sp. nov. differs from all other congeners based on morphological data, allopatric distribution, and molecular divergence. In terms of genetic divergence, Tylototriton ngoclinhensis sp. nov. distinctly differs from the sister species T. panhai (6.77%) and from T. ngarsuensis (12.36%) based on the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene. Tylototriton ngoclinhensis sp. nov. is a moderate sized and robust salamander species with large cephalic edges, parotoids, and vertebral ridge orange in coloration. The new taxon differs from its congeners by a combination of the following morphological characteristics: size medium (SVL 60.8–66.5 mm, TL 57.6–61.8 mm in males, and SVL 72.5–75.6 mm, TL 62.9–67.9 mm in females); head longer than wide; parotoids very prominent and enlarged, projecting backwards; tail length shorter than snout-vent length; vertebral ridge large, high and glandular in appearance; 14 large and distinct dorsolateral glandular warts; gular fold present; tips of fore and hind limbs overlapping when adpressed along the body; tips of fingers reaching between eye and nostril when foreleg is laid forward; dorsal surface and lateral sides of the head, upper and lower lips, dorsolateral glandular warts, vertebral ridge, the peripheral area of the cloaca and the ventral edge of the tail orange in coloration; the presence of a distinct black line extending from the posterior end of the eye towards the shoulder. Tylototriton ngoclinhensis sp. nov. is restricted to evergreen montane forests near water bodies on Ngoc Linh Mountain. We suggest that the new species should be classified as Endangered (EN) in the IUCN Red List. This new important discovery represents the eighth Tylototriton taxon described from Vietnam, and at the same time constitutes the southernmost distributional record for the whole genus in Asia.

Elmberg, Johan (2023): Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia). Zootaxa 5301 (3): 301-335, DOI: 10.11646/zootaxa.5301.3.1, URL: http://dx.doi.org/10.11646/zootaxa.5301.3.1

Smooth Newt Lissotriton vulgaris (Linneaus 1758) Distribution (Figure 2). Included records from Artportalen: all reports (N=670), as there is no confusion species except the much rarer Triturus cristatus. Widely distributed in the Southern Boreal; in a regular sampling grid covering the entire provinces of Gästrikland and Hälsingland, Sterner (2005) found the species in 42% of 111 squares in which randomly selected wetlands (et al. (2008) recorded the species in 23% of 155 randomly selected small wetlands. In the southern Middle Boreal this species is seemingly more patchily distributed, especially in the northern parts of the range. There are only two known extant occurrences in the Northern Boreal: Akkan and Abborrberg (4 kms apart in Stensele parish) in Lycksele lappmark (Persbo et al. 2006, Anders Forsgren pers. comm.). Interestingly, three of the northernmost occurrences are also the highest known from North Sweden (and in Sweden overall): Akkan and Aborrberg are both 520 m above sea level, and Jägarliden at 340 m (all in Lycksele lappmark; Persbo et al. 2006). Offshore occurrence is known from two islands in Medelpad (Brämön and Alnön; Elmberg & Ericsson 1983) and one in Hälsingland (Jättholmarna), indicating a fair dispersal capacity over brackish water. In coastal areas where the species occurs, many mainland records are very close to the sea (Elmberg 1995; Sterner 2005; Persbo et al. 2006; Olofsson et al. 2008). This pattern indicates a good general dispersal capacity, since such wetlands created by land uplift, are often less than 200 years old. There are not any data to gauge large-scale changes in distribution during the last 50 years, but local extinctions due to fish introduction have been documented (Dolmen 1978; Elmberg & Ericsson 1983; Persbo et al. 2006). One of these populations comprised neotenic individuals only (Långselberget, 460 m altitude, Stensele, Lycksele lappmark; Gislén & Kauri 1959; Dolmen 1978; Elmberg & Ericsson 1983). The northern distribution limit presented here runs farther north than in Gislén & Kauri (1959). This discrepancy is, however, not due to a range expansion but rather because the species has been overlooked before. Habitat and movements. Breeding habitats include tarns and small lakes in forested areas, as well as more or less permanent man-made wetlands (e.g., ponds, pasture pools, gravel pits). Tarns and lakes are usually oligoto mesotrophic, bordered either by floating Sphagnum mats or by moderately dense stands of aquatic grasses (Phragmites australis, Phalaris arundinacea) or sedges (Carex spp.) (Figure 11). The majority of known breeding wetlands are fishless (cf. Elmberg & Ericsson 1983; Sterner 2005; Persbo et al. 2006) and lack connection to other wetlands by streams. Several of the occurrences away from the more continuous range are isolated, either high on forested hills or in large peat bog complexes. Breeding sites are usually permanent wetlands, but the species has been recorded in ephemeral rock pools on the Baltic coast (Elmberg & Ericsson 1983; Aronsson et al. 2005; Figure 12). Summer habits and habitats are little known, as very few observations are made away from the breeding wetlands. Adults can be seen in the latter long into June, sometimes to mid-July. It is thus possible that many remain aquatic for a large part of the summer. During field work at Galtström (Medelpad) August 1 st we found active adults on land as well as in a nearby breeding pond. Terrestrial habits in North Sweden are little known; most records on land have been made under logs or woody debris in damp spruce-dominated forest close to breeding sites. Terrestrial summer records have also been made under flat sheltering objects near man-made wetlands. Hibernation habitats and habits in North Sweden have not been documented; it is not even known whether terrestrial or aquatic hibernation is the rule. However, aquatic hibernation was the only option in the neotenic population at Stensele in Lycksele lappmark, now extinct (Dolmen 1978). Very little is known about seasonal movements. The almost total lack of observations of adults moving to or from breeding wetlands strongly suggests they spend much of their life in the latter or their close proximity. Abundance estimates and trends. There are very little data on local abundance, and no indication that any breeding wetland holds more than 200 reproducing adults (e.g., Dolmen 1978; Elmberg & Ericsson 1983; Sterner 2005; Persbo et al. 2006; Olofsson et al. 2008). Given the patchy distribution, it is unwise to suggest abundance estimates for larger areas. However, the surveys by Sterner (2005) and Olofsson et al. (2008), which covered a mere fraction of the suitable wetlands in the Southern Boreal, suggest there must be thousands of breeding wetlands in this region. There are no indications of changes in abundance over the last 50 years, apart from local extinctions., Published as part of Elmberg, Johan, 2023, Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia), pp. 301-335 in Zootaxa 5301 (3) on pages 306-308, DOI: 10.11646/zootaxa.5301.3.1, http://zenodo.org/record/8030434, {"references":["Sterner, M. (2005) Inventering av Vattensalamandrar i Gavleborgs lan 2005. Rapport, Naturenheten Lansstyrelsen i Gavleborgs lan, Gavle, 2005, 19. [unknown pagination, in Swedish]","Olofsson, F., Persbo, F. & Ohmark, S. (2008) Inventering av storre vattensalamander Triturus cristatus i Vasternorrlands lan 2006. Lansstyrelsen Vasternorrland Miljoavdelningen Rapport Harnosand, 2008, 9. [in Swedish]","Persbo, F., Reinoja, J. & Ohmark, S. (2006) Inventering av mindre vattensalamander, Triturus vulgaris i Vasterbottens lan 2005. Project thesis, Department of Ecology and Environmental Science, Umea University. [unknown pagination, in Swedish]","Elmberg, J. & Ericsson, S. (1983) Skrattabbarn - om mindre vattenodlan Triturus vulgaris (L.) i Vasterbottens lan. Natur i Norr, 2 (1), 1 - 22. [in Swedish]","Elmberg, J. (1995) Groddjurens och kraldjurens utbredning i Norrland. Natur i Norr, 15 (2), 57 - 82. [in Swedish]","Dolmen, D. (1978) De neotene salamanderne (\" skrattabborrene \") ved Stensele. Fauna och Flora, 73 (4), 171 - 177. [in Norwegian]","Gislen T. & Kauri H. (1959) Zoogeography of the Swedish amphibians and reptiles, with notes on their growth and ecology. Acta Vertebratica, 1 (3), 197 - 397.","Aronsson, A., Karlberg, A., Persson, B., Ohrling, H. & Gothe, L. (2005) Oversiktlig kustinventering 2002 - 2004. Lansstyrelsen Vasternorrland, avdelningen for Kultur och natur. Rapport, 2005, 3. [in Swedish]"]}

Great Crested Newt Triturus cristatus (Laurenti 1768) Distribution (Figure 3). Included records from Artportalen (N=320): due to risk of confusion with the more widespread Lissotriton vulgaris only records documented by a photograph or made by experienced observers have been included. Widely distributed in the Southern Boreal; in a regular sampling grid covering the entire provinces of Gästrikland and Hälsingland, Sterner (2005) found the species in 21% of 111 squares in which randomly selected wetlands (et al. (2008) recorded the species in 3% of 155 randomly selected small wetlands. In coastal southern Medelpad the occurrence is more or less continuous (Aronsson et al. 2005), as it is in coastal Hälsingland and Gästrikland. In the Middle Boreal there are scattered records north to 63 oN, most of which from east-central Jämtland. A recent photo-documented record in NE Ångermanland suggests that the species may range further north along the coast than currently known. The only record from the Northern Boreal during the study period is a possibly disjunct occurrence in western Jämtland (Figure 3). Although most known extant breeding sites are in coastal lowland, at least three are at 300–400 m altitude (Elmberg 1995; Olofsson et al. 2008). An extinct population at Långselberget (Lycksele lappmark (Middle Boreal)) was even higher, at 460 m. Offshore occurrence is known from only one site in North Sweden (Kråkön, Hälsingland). However, several breeding wetlands on the mainland are very near the sea (Sterner 2005; Olofsson et al. 2008). This indicates a fair general dispersal capacity, since such wetlands, created by land uplift, are often less than 200 years old. One local extinction has been documented in the Middle Boreal: the isolated occurrence in Lycksele lappmark, comprising only neotenic individuals, went extinct due to fish introduction in the 1960’s (Långselberget, Stensele; Gislén & Kauri 1959; Dolmen 1978; Elmberg & Ericsson 1983). There is not any indication that the general distribution has changed recently in North Sweden. Instead, the species has previously been much overlooked. It is more widespread, more numerous, and the range more extensive to the northwest than depicted in any previous source (cf. Gislén & Kauri 1959; Elmberg 1995). Habitat and movements. Most known breeding wetlands in North Sweden are small fishless tarns bordered by coniferous forest (e.g., Sterner 2005; Olofsson et al. 2008). These wetlands are generally oligotrophic and have floating mats of Sphagnum along the shore. However, in coastal areas a fair number of breeding sites of anthropogenic origin are known: farmland ponds, disused fishponds, peat quarries, gravel pits, and golf course ponds. Many of these are meso- to eutrophic and thus more resemble typical breeding habitats in South Sweden. Most known breeding wetlands are permanent, but some are more ephemeral, such as depressions in wet forest and rock pools near the Baltic (Olofsson et al. 2008). Observations of moving or migrating individuals are exceedingly few. Their scarcity and location strongly suggest very limited movements away from breeding wetlands and imply a largely aquatic lifestyle in summer, too. Summer habitat use is almost unknown; a few records of terrestrial adults have been made, mainly under woody debris or in brooks in old spruce-dominated forest near breeding wetlands. During field work at Galtström (Medelpad) August 1 st, we found active adults on land and in a nearby breeding pond, as well as hiding under woody debris in the upper part of the seashore littoral. Nothing is known about hibernation habits in North Sweden, that is, whether terrestrial or aquatic. However, aquatic hibernation must have been the rule in the extinct neotenic population at Stensele in Lycksele lappmark (Dolmen 1978). Abundance estimates and trends. There are not any data about local abundance. Given the patchy distribution, suggesting abundance estimates for larger areas is unwise. However, the surveys by Sterner (2005) and Olofsson et al. (2008), which covered a mere fraction of the suitable wetlands in the Southern Boreal suggest there must be several hundred breeding wetlands in this region. There are no indications of changes in abundance over the last 50 years, apart from local extinctions., Published as part of Elmberg, Johan, 2023, Amphibians and reptiles in North Sweden: distribution, habitat affinities, and abundance (Classes: Amphibia and Reptilia), pp. 301-335 in Zootaxa 5301 (3) on pages 308-309, DOI: 10.11646/zootaxa.5301.3.1, http://zenodo.org/record/8030434, {"references":["Sterner, M. (2005) Inventering av Vattensalamandrar i Gavleborgs lan 2005. Rapport, Naturenheten Lansstyrelsen i Gavleborgs lan, Gavle, 2005, 19. [unknown pagination, in Swedish]","Olofsson, F., Persbo, F. & Ohmark, S. (2008) Inventering av storre vattensalamander Triturus cristatus i Vasternorrlands lan 2006. Lansstyrelsen Vasternorrland Miljoavdelningen Rapport Harnosand, 2008, 9. [in Swedish]","Aronsson, A., Karlberg, A., Persson, B., Ohrling, H. & Gothe, L. (2005) Oversiktlig kustinventering 2002 - 2004. Lansstyrelsen Vasternorrland, avdelningen for Kultur och natur. Rapport, 2005, 3. [in Swedish]","Elmberg, J. (1995) Groddjurens och kraldjurens utbredning i Norrland. Natur i Norr, 15 (2), 57 - 82. [in Swedish]","Gislen T. & Kauri H. (1959) Zoogeography of the Swedish amphibians and reptiles, with notes on their growth and ecology. Acta Vertebratica, 1 (3), 197 - 397.","Dolmen, D. (1978) De neotene salamanderne (\" skrattabborrene \") ved Stensele. Fauna och Flora, 73 (4), 171 - 177. [in Norwegian]","Elmberg, J. & Ericsson, S. (1983) Skrattabbarn - om mindre vattenodlan Triturus vulgaris (L.) i Vasterbottens lan. Natur i Norr, 2 (1), 1 - 22. [in Swedish]"]}

Previous phylogenetic studies based on mitochondrial DNA data have consistently suggested that Hynobius tokyoensis consists of two major clades, clade A (northern clade) and clade B (southern clade). In this study, we newly estimated their population genetic structure and phylogenetic relationships by nuclear SNPs, and the results suggested heterospecific relationships of the two mitochondrial clades, without present hybridization in between. They were also recognized as morphologically different. The type locality of H. tokyoensis is in Tokyo Prefecture, and therefore clade B corresponds to H. tokyoensis sensu stricto, leaving clade A without available scientific name. We, thus, describe the clade A from northeastern Kanto to southern Tohoku as a new species Hynobius sengokui. The new species is distinguished from H. tokyoensis by its relatively longer axilla-groin distance, shorter trunk, and deeper vomerine teeth series, and is estimated to have diverged from it during the late Pliocene.

Hynobius sumidai Sugawara, Naito, Iwata, and Nagano, 2022 Hynobius nebulosus (part, as Hiroshima morphotype) Okawa and Utsunomiya, 1989, 142. Hynobius nebulosus (part, as Aki morphotype) Okawa et al., 2005, 11. Hynobius akiensis (part) Matsui et al., 2019, 32. Holotype: HMNH-AM-102, from Tamaricho, Takehara-shi, Hiroshima Prefecture. Paratype: KPM-NFA 945, from the same locality of the holotype. Specimens referred in the present paper. Hiroshima Prefecture: Takehara-shi, KUHE 64274, from Nika-cho, collected by Y. Tomimori on 2 May 2022. KUHE 20220610 _1, 20220610_3, from Nika-cho, collected by K. Sakamoto on 22 April 2021. KUHE 61404-61406, from Tamari-cho, collected by Y. Tomimori on 4 Jury 2019. Higashihiroshima-shi, KUHE nb8-7, from the former Kurose-cho (Date of collection and collector unknown). KUHE 8682, 8683, 8685, from the former Kurose-cho, collected by M. Matsui on 10 March 1986. U 103–104, from former Kurose-cho, collected by T. Utsunomiya on 24 March 1986. KUHE OU 0297– 0299, from former Saijocho, collected by H. Okawa on 28 March 2007; KUHE OU 0300, from former Happonmatsu-cho, collected by H. Okawa on 28 March 2007; KUHE OU0318, 0319, from the former Akitsu-cho, collected by H. Okawa on 13 May 2007. KUHE OU 0340–0342, from former Happonmatsu-cho, collected by H. Okawa on 28 March 2008. Kure-shi U0842–U0845, from Gohara-cho, collected by T. Utsunomiya on 22 March 1990. Diagnosis. A small species of the genus Hynobius that breeds in hilly wetlands (adult male SVL 36.2–56.9 mm). Phylogenetically, H. sumidai forms a clade with H. geiyoensis in mitochondrial and nuclear DNA, but has a smaller SVL. than the latter. The dorsum of an adult dark brown and yellow stripes on upper and lower edges of tail often indistinct. Short limbs and tail; forelimb and hindlimb tips barely meeting (overlap of -2 to 0 costal folds in males). The fifth toe undeveloped. Egg sac short, loosely coiled into a string, with slightly fewer clutch size than in other lentic breeding species of the genus. Most similar to H. akiensis, but with shorter internarial distance (RIND 5.7% vs. 6.3%), interorbital distance (RIOD 6.0% vs. 6.4%), forelimb (RFLL 23.6% vs. 25.3%), and hindlimb (RHLL 29.5% vs. 30.8%). Color. In life, dorsum pale brown with minute black spots. Small white spots on ventral side, and yellowish brown stripe on dorsal edge of tail often indistinct, less distinct on ventral side. In preservative, ventral color tends to be faded to grayish-brown and white nuptial color on throat not clear. Variation. Morphological measurement data for males are summarized in Table 2. In 19 males examined, SVL 47.6 ± 5.2 (36.2–56.9) mm, tips of adpressed limbs separated or touching, the fifth toe generally weakly developed, with few individuals having only four toes (about 5.0% of the specimens examined). Body light brown with black spots, and some individuals with silvery white spots on dark brown background. Range. The species is known only in the low hills of southern area in the middle region of Hiroshima. Higashihiroshima-shi (including the former Saijo-cho, Kurose-cho, Fukutomi-cho, Toyosaka-cho, Akitsu-cho, and Happonmatsu-cho), Takehara-shi, and Kure-shi (Fig. 1). Altitude of the localities extends from 160.0– 455.0 m with the mean±SD of 296.3± 121.1 m (n=11). Natural history. Breeding usually occurs in hilly streams, wetlands, and paddy ditches from March to April. Egg sacs are string-like in shape and coiling. number of eggs is 26 ± 9.5 (3–67) (Average ± SD [range]) (Okawa et al. 2019). Morphological Comparisons. Hynobius sumidai has a yellowish-brown body color with small black dots on the dorsal side, a body color pattern that is often seen in lentic Hynobius (e.g., H. geiyoensis, H. akiensis, H. setouchi, H. utsunomiyaorum, H. abuensis, H. iwami, H. bakan, H. setoi, and H. kunibiki). However, H. iwami, H. bakan, H. setoi, and H. kunibiki can be distinguished from H. sumidai by distinct yellow stripes on dorsal and ventral sides of tail, while H. sumidai has an indistinct yellow stripe on dorsal side, if present. In addition, this species differs from H. iwami, which always lacks the fifth toe. Hynobius sumidai (mean male SVL 48.0 mm) is distinctly smaller than H. geiyoensis (60.4 mm) and H. abuensis (57.3 mm). Hynobius sumidai is distinguishable from the genetically related H. geiyoensis by the following ratio values: longer head, shorter interorbital, larger eyes, shorter trunk and tail, narrower medial tail, higher basal tail, shorter hindlimb and fifth toes, and wider vomerine teeth series. Hynobius sumidai significantly differs from H. akiensis in the following ratio values: shorter internarial, interorbital, forelimb, and hindlimb. Hynobius sumidai significantly differs from H. setouchi in the following ratio values: shorter snout, internarial, interorbital, tail, forelimb, hindlimb, third toe, and fifth toe. Hynobius sumidai is distinguishable from H. utsunomiyaorum by its significantly smaller SVL and following ratio values: larger head, longer lower jaw, larger eyes, narrower medial tail, higher basal tail, longer third finger and fifth toe, and larger vomerine teeth series. Protection: H. sumidai is in the Japanese Red List as Endangered (EN) under the name of H. akiensis (Ministry of the Environment Government of Japan 2020) and is listed as Vulnerable (VU) by the Red Data List Hiroshima (Okawa [2021] as H. akiensis). This species is presently protected by Higashihiroshima-shi, Hiroshima Prefecture., Published as part of Tomimori, Yusuke, Matsui, Masafumi, Okawa, Hiroshi, Nishikawa, Kanto, Tanabe, Shingo & Kamasaka, Ryo, 2023, Reassessment of species delimitation using nuclear markers in three lentic-breeding salamanders from the Chugoku District of Japan (Amphibia: Caudata: Hynobiidae), pp. 145-160 in Zootaxa 5293 (1) on pages 155-156, DOI: 10.11646/zootaxa.5293.1.6, http://zenodo.org/record/7959833, {"references":["Sugawara, H., Naito, J., Iwata, T. & Nagano, M. (2022) Molecular Phylogenetic and Morphological Problems of the Aki Salamander Hynobius akiensis: Description of Two New Species from Chugoku, Japan. Bulletin of The Kanagawa Prefectural Museum Natural Science, 51, 35 - 46. https: // doi. org / 10.32225 / bkpmnh. 2022.51 _ 35","Okawa, H. & Utsunomiya, T. (1989) Hynobius nebulosus from Hiroshima Prefecture. In: Matsui, M., Hikida, T. & Goris, R. C. (Eds.), Current Herpetology in East Asia. Herpetological Society of Japan, Kyoto, pp. 142 - 146.","Okawa, H., Okuno, T. & Utsunomiya, T. (2005) A group of Hynobius nebulosus distributed in Abu, Tsuwano and Yamaguchi region. Amphibian History, 14, 11 - 14.","Matsui, M., Okawa, H., Nishikawa, K., Aoki, G., Eto, K., Yoshikawa, N., Tanabe, S., Misawa, Y. & Tominaga, A. (2019) Systematics of the widely distributed Japanese clouded salamander, Hynobius nebulosus (Amphibia: Caudata: Hynobiidae), and its closest relatives. Current Herpetology, 38, 32 - 90. https: // doi. org / 10.5358 / hsj. 38.32","Okawa, H., Okuno, T. & Utsunomiya, T. (2019) Major groups of Hynobius nebulosus in Western Japan. Bulletin of the Herpetological Society of Japan, 2019, 9 - 21.","Okawa, H. (2021) H. akiensis. In: Rare Species Subcommittee (Ed.), Red Data Book Hiroshima, 2021, pp. 142."]}

Hynobius geiyoensis Sugawara, Naito, Iwata, and Nagano, 2022 Hynobius nebulosus (part, as Osakijima morphotype) Okawa and Utsunomiya, 1989, 142. Hynobius nebulosus (part, as Island morphotype) Okawa et al., 1999, 47. Hynobius nebulosus (part, as Shikoku morphotype) Okawa et al., 2007, 58. Hynobius akiensis (part) Matsui et al., 2019, 32. Holotype: HMNH (Hiwa Museum for Natural History) AM-101, from Osakikamijima-cho, Hiroshima Prefecture. Paratypes: KPM (Kanagawa Prefectural Museum of Natural History) NFA 943, KPM-NFA 944, from the same locality as the holotype. Specimens referred in the present paper. Hiroshima Prefecture: Osakikamijima Island, KUHE OU0957- 0961, from Nakano, collected by H. Okawa on 13 March 2019. KUHE 62645, 62660, 62666–62671, from Nakano, collected by Y. Tomimori on 17 February 2021. Ehime Prefecture: Imabari-shi, KUHE OU0343, 0344, 0347, 0348, from the former Namikata-cho, collected by H. Okawa on 5 April 2008. KUHE T 2839–2842, 2844, 2845, from the former Namikata-cho, collected by S. Tanabe and K. Okayama on 1 March 1997. KUHE T 3284, 3285, from the former Namikata-cho, collected by S. Tanabe and K. Okayama on 21 March 2005. Diagnosis. A medium-sized species (SVL 53.3–67.3 mm in males) of the lentic-breeding Hynobius; White dots are dense on the lateral and ventral sides, and black dots are visible on the dorsum. Tips of fore- and hindlimbs adpressed on body do not overlap in either male or female (overlap of −2.0 to 0 and -3.5 to -2.5 costal folds, respectively). Head, trunk, and tail clearly longer and the fifth toe more developed than in genetically close relative, H. akiensis and H. sumidai. Color. Body coloration greatly variable, with dorsum of body ranging from dark to yellowish brown, sometimes with black spots in life. Tail lacking yellowish-brown stripes on dorsal and ventral sides, although seldom with an indistinct stripe on dorsum. Ventral side of body grayish brown scattered with silvery white spots. White nuptial color on throat distinct in males. Variation. Morphological measurement data for males are summarized in Table 2. In 16 males examined, SVL 60.4 ±3.7 (53.3–67.3) mm, adpressed limbs always separated, and the fifth toe always present. In addition, all individuals with many fine white spots on ventral side of body, more pronounced in younger individuals. Range. Known only from two areas on Osaki-kamijima-cho, Toyota-gun, Hiroshima Prefecture and Imabari-shi, Ehime Prefecture (Fig. 1). Altitude of the localities extending from 9.6– 185.0 m a.s.l. with the mean±SD of 51.1± 37.4 m (n=25). Natural history. Breeding usually occurs from late January to early March in Hiroshima Prefecture, but from late December to early May in Ehime Prefecture. (Itano et al. 2016). Egg sacs are string-like in shape and coiling. Number of eggs is 62 ± 13.3 (34–93) (Tanabe and Okayama 2001). The minimum and maximum water temperatures at the time of breeding were 3.5–6.6°C and 8.0–13.6°C, respectively in Ehime Prefecture. (Itano et al. 2016). Morphological Comparisons. Hynobius geiyoensis has a dark brown body color with small white dots, a body color pattern that is often seen in lentic Hynobius (e.g., H. sumidai, H. akiensis, H. setouchi, H. utsunomiyaorum, H. iwami, H. abuensis, H. bakan, H. setoi, and H. kunibiki). However, H. iwami, H. bakan, H. setoi, and H. kunibiki can be distinguished from H. geiyoensis by distinct yellow stripes on dorsal and ventral sides of tail. Hynobius geiyoensis (mean male SVL 60.4 mm) is distinctly larger than H. sumidai (47.6 mm), H. akiensis (48.8 mm), and H. setouchi (51.5 mm). In addition, this species lacks striations on the envelope of egg sacs unlike H. abuensis. However, it may be confused with H. setouchi, which is known to be distributed in Shikoku and the islands of the Seto Inland Sea. Hynobius geiyoensis is distinguishable from the genetically close H. sumidai by its significantly larger SVL and by shorter head length, wider interorbital, smaller eyes, longer trunk and tail, wider medial tail lower basal tail, longer hindlimb and fifth toe, and narrower vomerine teeth series. Hynobius geiyoensis is distinguishable from H. akiensis by its significantly larger SVL and following ratio values: shorter head length, smaller eyes, longer trunk, tail and fifth toe. Hynobius geiyoensis is distinguishable from H. setouchi by its significantly larger SVL and following ratio values: shorter maximum head width, shorter lower jaw and snout, narrower upper eyelid, longer trunk, lower basal and medial tail, shorter third toe, and smaller vomerine teeth series. Hynobius geiyoensis is distinguishable from H. utsunomiyaorum by its significantly larger SVL and following ratio values: wider interorbital, longer trunk, tail, hindlimb, third finger, third toe, and fifth toe, and larger vomerine teeth series. Protection. H. geiyoensis is in the Japanese Red List as Endangered (EN) under the name of H. akiensis (Ministry of the Environment Government of Japan 2020) and is listed as Vulnerable (VU) by the Red Data List Ehime (Tanabe 2014). This species is presently protected by Ehime Prefecture, However, there is currently no effective protection law on Osakikamijima Island in Hiroshima Prefecture. Due to very limited distribution range, conservation status of this species needs to be immediately reassessed., Published as part of Tomimori, Yusuke, Matsui, Masafumi, Okawa, Hiroshi, Nishikawa, Kanto, Tanabe, Shingo & Kamasaka, Ryo, 2023, Reassessment of species delimitation using nuclear markers in three lentic-breeding salamanders from the Chugoku District of Japan (Amphibia: Caudata: Hynobiidae), pp. 145-160 in Zootaxa 5293 (1) on pages 153-154, DOI: 10.11646/zootaxa.5293.1.6, http://zenodo.org/record/7959833, {"references":["Sugawara, H., Naito, J., Iwata, T. & Nagano, M. (2022) Molecular Phylogenetic and Morphological Problems of the Aki Salamander Hynobius akiensis: Description of Two New Species from Chugoku, Japan. Bulletin of The Kanagawa Prefectural Museum Natural Science, 51, 35 - 46. https: // doi. org / 10.32225 / bkpmnh. 2022.51 _ 35","Okawa, H. & Utsunomiya, T. (1989) Hynobius nebulosus from Hiroshima Prefecture. In: Matsui, M., Hikida, T. & Goris, R. C. (Eds.), Current Herpetology in East Asia. Herpetological Society of Japan, Kyoto, pp. 142 - 146.","Okawa, H., Utsunomiya, T. & Okuno, T. (1999) Four toes population of Hynobius nebulosus. Hibakagaku (Journal of the Hiba Society of Natural History), 191, 4.","Okawa, H., Okuno, T. & Utsunomiya, T. (2007) Major groups of Hynobius nebulosus in Western Japan. Bulletin of the Herpetological Society of Japan, 2007, 58 - 59.","Matsui, M., Okawa, H., Nishikawa, K., Aoki, G., Eto, K., Yoshikawa, N., Tanabe, S., Misawa, Y. & Tominaga, A. (2019) Systematics of the widely distributed Japanese clouded salamander, Hynobius nebulosus (Amphibia: Caudata: Hynobiidae), and its closest relatives. Current Herpetology, 38, 32 - 90. https: // doi. org / 10.5358 / hsj. 38.32","Itano, K., Fujiwara, Y., Ikeuchi, K., Kobayashi, S. & Omori, K. (2016) Study on the breeding ecology of Clouded salamander (Hynobius nebulosus) in Imabari, Ehime Prefecture. Bulletin of the Ehime Pref. Museum, 21, 1 - 18.","Tanabe, S. & Okayama, K. (2001) About the lentic salamander discovered in Ehime Prefecture, Japan. Bulletin of the Ehime Prefecture Museum, 15, 23 - 27.","Tanabe, S. (2014) H. nebulosus. In: Ehime Red Data Book Revision Committee (Ed.), Red Data Book Ehime, 2014, pp. 97."]}

Hynobius akiensis Matsui, Okawa et Nishikawa, 2019 Hynobius abei (part) Sato, 1943, 163. Hynobius nebulosus (part, as Transitional morphotype) Okawa and Utsunomiya, 1992, 12. Hynobius nebulosus (part, as Miyoshi morphotype) Okawa and Utsunomiya, 1992, 12. Hynobius nebulosus (part, as Aki morphotype) Okawa et al., 2005, 11 Holotype KUHE 35925, from Haizuka, Mirasaka-cho, Miyoshi-shi, Hiroshima Prefecture. Paratype KUHE 35924, one male, data same as the holotype. (KUHE 8682–8687 from Kurose-cho, Higashihiroshima-shi, in the original description (Matsui et al. 2019 removed.) Specimens referred in the present paper. Hiroshima Prefecture: Akitakata-shi, KUHE OU 0313, 0314, from the former Mukaihara-cho, collected by H. Okawa on 15 April 2007. KUHE OU 0368, 0369, from Takamiya-cho, collected by H. Okawa on 26 April 2009. KUHE OU 0510, from Takamiya-cho, collected by H. Okawa on 2 May 2010. KUHE 58834, from the former Koda-cho, collected by Y. Misawa on 1 March 2017. Higashihiroshima-shi, KUHE OU 0309–0312 from the former Toyosaka-cho, collected by H. Okawa on 5 April 2007. KUHE OU 0905, 0907, from the former Fukutomi-cho, collecter by H. Okawa on 4April 2017. Miyoshi-shi, KUHE OU 0301–0303, from the former Miwa-cho, collected by H. Okawa on 28 March 2007. KUHE OU 0387–0389, from Miwaka-cho, collected by H. Okawa on 3 March 2010. KUHE OU 0390, from Shimoshiwachi-cho, collected by H. Okawa on 3 March 2010. KUHE 58887, 58888, from Mirasaka-cho, collected by Y. Misawa on 7 March 2018. KUHE OU 0970–0972, from Miwaka-cho, collected by H. Okawa on 8 April 2019. KUHE 61385, 61385 _1, 61385_2, from Miwaka-cho, collected by H. Okawa on 5 Jury 2019. KUHE 62661–62664, from Megurikami-cho, collected by Y. Tomimori on 16 February 2021. Kitahiroshima-cho, Yamagata-gun, KUHE 61220, 61220 _1, 61220_2, collected by Y. Tomimori on 19 Jury 2019. Sera-cho, Sera-gun, KUHE OU 0472, from Sera-cho, collected by H. Okawa on 13 March 2010. KUHE OU 0474, from Sera-cho, collected by H. Okawa on 19 March 2010. KUHE OU 0479, from the former Seranishi-cho, collected by H. Okawa on 19 March 2010. Diagnosis. A small species of the genus Hynobius that breeds in wetlands in hilly to mountainous areas about 1000 m. altitude (adult male SVL 40.8–57.5 mm). Phylogenetically, H. akiensis forms a clade with H. geiyoensis and H. sumidai in nuclear DNA, which relationships not supported by mtDNA. Dorsum dark brown, some almost dark blue. Yellow stripes on upper and lower edges of tail indistinct. Limbs and tail short, forelimb and hindlimb tips barely meeting (overlap of -2 to 0.5 costal folds in males). Fifth toe least developed in H. akiensis sensu lato (R5TL: H. akiensis 1.8%, H. sumidai 2.0%, and H. geiyoensis 2.7%). Egg sac short, loosely coiled into a string, with smaller clutch size than in other lentic breeding species of the genus. Most similar to H. sumidai but with wider internarial (RIND 6.3% vs. 5.7%), interorbital (RIOD 6.4% vs. 6.0%), and longer forelimbs (RFLL 25.3% vs. 23.6%) and hindlimb (RHLL 30.8% vs. 29.5%). Color. In life, dorsum yellowish brown with minute black spots. Dark blue or dark brown individuals without black spots also present. Small white spots on ventral side, and often indistinct yellowish-brown stripe on dorsal edge of tail, less distinct on ventral side. In preservative, ventral color tends to be faded to grayish brown and white nuptial color on throat not clear. Variation. Morphological measurement data for males are summarized in Table 2. In 27 males examined, SVL 48.8 ± 4.4 (40.8–57.5) mm, adpressed limbs always separated, fifth toe generally weakly developed, some individuals with only four toes (about 15% of the specimens examined). Body color light brown with black spots, some individuals with silvery white spots on dark brown and dark blue background. Range. Hiroshima Prefecture: Akitakata-shi (including the former Yoshida-cho, Mukaihara-cho, Koda-cho, Takamiya-cho, Yachiyo-cho and the former Midori-cho [Sugahara et al. 2022]); Higashihiroshima-shi (including the former Fukutomi-cho, Toyosaka-cho); Mihara-shi (including the former Daiwa-cho, and the former Kui-cho [Sugahara et al. 2022]); Onomichi-shi (including the former Mitsugi-cho); Miyoshi-shi (including the former Kisacho, Mirasaka-cho, Miwa-cho); Kitahiroshima-cho, Yamagata-gun (including the former Chiyoda-cho, Oasa-cho, and the former Toyohira-cho, and Geihoku-cho [Sugahara et al. 2022]); Sera- cho, Sera-gun (including the former Seranishi-cho); Hiroshima-shi (only former Asakita-ku [Sugahara et al. 2022]); Akiota-cho (only former Togochicho [Sugahara et al. 2022]). Shimane Prefecture: Onan-cho (only former Mizuho-cho and Hasumi- mura [Sugahara et al. 2022]). Altitude of the localities extends from 260.0– 930.0 m with the mean±SD of 447.9± 145.4 m (n=23). Natural history. Breeding usually occurs in hilly streams, wetlands, and paddy ditches from March to April. Egg sacs are string-like in shape and coiling. number of eggs is 24.5 ± 8.6 (2–58) (Okawa et al. 2019). The minimum and maximum water temperatures at the time of breeding were 2.5–5.0°C and 8.1–11.1°C, respectively in Haizukadam, Mirasaka-cho, Miyoshi-shi, Hiroshima Prefecture. (Utsunomiya et al. 1996). Morphological Comparisons. Hynobius akiensis has a dark brown body color with small black dots on dorsal side and small white spots on ventral side, a body color pattern that is often seen in lentic Hynobius (e.g., H. sumidai, H. akiensis, H. setouchi, H. utsunomiyaorum, H. abuensis, H. iwami, H. bakan, H. setoi, and H. kunibiki). However, H. iwami, H. bakan, H. setoi, and H. kunibiki can be distinguished from H. akiensis because they have a distinct yellow stripe on dorsal and ventral sides of tail, while H. akiensis has an indistinct yellow stripe on dorsal side if present. Hynobius akiensis and H. iwami may be confused because of their close distribution and tendency of lacking the fifth toe. Hynobius akiensis (mean male SVL 48.8 mm) H. abuensis (57.3 mm). Hynobius akiensis significantly differs from H. setouchi in the following ratio values: shorter maximum head width, tail length, medial tail height, third toe and fifth toe, and smaller vomerine teeth series. Hynobius akiensis is distinguishable from H. utsunomiyaorum by its significantly smaller SVL and following ratio values: larger head, longer lower jaw, larger eyes, shorter axilla-groin distance and medial tail width, higher basal tail, longer hindlimb, third finger and fifth toe, and larger vomerine teeth series. Hynobius akiensis significantly differs from H. iwami in the following ratio values: shorter upper eyelid length, higher basal tail, and longer fifth toe. Morphological differences from allied species H. sumidai and H. geiyoensis are noted above for each species. Protection. H.akiensis is in the Japanese Red List as Endangered (EN) (Ministry of the Environment Government of Japan 2020). This species is on the list of Hiroshima Prefecture as Vulnerable (VU) by the Red Data List Hiroshima (Okawa 2021), and on the list of Shimane Prefecture as Near Threatened (NT) by the Revised Shimane Red Data Book 2014 Animals (Akiyoshi and Matsuno 2014)., Published as part of Tomimori, Yusuke, Matsui, Masafumi, Okawa, Hiroshi, Nishikawa, Kanto, Tanabe, Shingo & Kamasaka, Ryo, 2023, Reassessment of species delimitation using nuclear markers in three lentic-breeding salamanders from the Chugoku District of Japan (Amphibia: Caudata: Hynobiidae), pp. 145-160 in Zootaxa 5293 (1) on pages 156-157, DOI: 10.11646/zootaxa.5293.1.6, http://zenodo.org/record/7959833, {"references":["Matsui, M., Okawa, H., Nishikawa, K., Aoki, G., Eto, K., Yoshikawa, N., Tanabe, S., Misawa, Y. & Tominaga, A. (2019) Systematics of the widely distributed Japanese clouded salamander, Hynobius nebulosus (Amphibia: Caudata: Hynobiidae), and its closest relatives. Current Herpetology, 38, 32 - 90. https: // doi. org / 10.5358 / hsj. 38.32","Sato, I. (1943) A Monograph of the Tailed Batrachians of Japan. Nippon Shuppan-sha, Osaka, 520 pp.","Okawa, H. & Utsunomiya, T. (1992) On the toe bones of Hynobius nebulosus. Japanese Journal of Herpetology, 14, 214.","Okawa, H., Okuno, T. & Utsunomiya, T. (2005) A group of Hynobius nebulosus distributed in Abu, Tsuwano and Yamaguchi region. Amphibian History, 14, 11 - 14.","Okawa, H., Okuno, T. & Utsunomiya, T. (2019) Major groups of Hynobius nebulosus in Western Japan. Bulletin of the Herpetological Society of Japan, 2019, 9 - 21.","Utsunomiya, T., Utsunomiya, Y., Okawa, H. & Naito, J. (1996) Amphibians from the area around Haizuka projected reservoir in Hiroshima Prefecture, Southwest Japan. R eprinted from Natural History of Haizuka, 1996, 177 - 215.","Okawa, H. (2021) H. akiensis. In: Rare Species Subcommittee (Ed.), Red Data Book Hiroshima, 2021, pp. 142.","Akiyoshi, H. & Matsuno, A. (2014) Hynobius nebulosus. In: Shimane Red Data Book Revision Committee (Ed.), Revised Shimane Red Data Book, 2014, Animals, pp. 73."]}

The golden Alpine salamander Salamandra atra aurorae Trevisan, 1982 is an endemic subspecies found in Sette Comuni and Vezzena plateau in Veneto, Italy. We describe an aggressive interaction between two males which fought for four minutes, trying to go on top of each other and rubbing their chin on the antagonist’s head. This is the first documented case of aggressive behaviour in Salamandra atra aurorae.

Desmognathus aureatus (Martof, 1956) Leurognathus marmorata aureata Martof, 1956 Leurognathus marmorata roborata Martof, 1956, holotype UMMZ 111568 by original designation, collected 18 September 1954 by “the Bernard Martof family,” type locality “Reed Creek, along Burrell Ford Road, 0.5 of a mile from its junction with Glade School Road, about 3.5 miles northwest of Pine Mountain Community, Rabun County, Georgia; 2,350 feet elevation.” Paratypes UMMZ 111569 (lot of 26 specimens); same data. Informally synonymized with L. marmorata by Martof (1962) and resurrected by Dubois and Raffaëlli (2012) based on a suggestion by Jones et al. (2006). Here designated a junior subjective synonym of D. aureatus. Holotype: UMMZ 111566 by original designation, collected 11 September 1954 by “ Stephen and Andrew Martof,” type locality “ Jarrard’s Creek, 0.2 of a mile below its crossing of U.S. Route 19, about 9 air miles north-northeast of Dahlonega, Lumpkin County, Georgia; 1,550 feet elevation.” Paratypes: UMMZ 111567 (lot of 20 specimens); “data as for the holotype.” Description: After Martof (1956), adults of this species have no vomerine teeth (versus presence in Desmognathus intermedius and D. marmoratus [some females]); ventral color-pattern pale or mottled (versus dark or black in other Blue Ridge lineages); and dark brown to black background of dorsum with dorsal color-pattern consisting of paired, golden, marbled or lichenous blotches (albeit less distinct in northeastern populations previously assigned to L. m. roborata Martof, 1956). These characters should be re-evaluated for diagnostic consistency. In our sample, this is the smallest shovel-nosed species, with metamorphosed SVL = 34–73mm. Based on our size-corrected linear morphometric analyses, this species can be distinguished from D. intermedius by a shorter trunk (AG in original measurements = 17–40mm) and from D. marmoratus by a longer head (SG in original measurements = 8.1–17.1mm). From Martof (1962), females at sites in Georgia develop sexual maturity around 55–59mm SVL, typically laying 27– 54 eggs from May to July, with clutch size dependent on body size. Hatchlings are as small as 10–12mm SVL and metamorphose at 25–38mm (usually ~30) after 10–20 months, though up to three distinct size classes at some suggest possibly longer periods up to ~36 months. Larvae and adults are primarily insectivorous, likely preying opportunistically on available aquatic invertebrates (Martof and Scott 1957). Adults in some Georgia populations are occasionally bright yellow, a characteristic that apparently shifts rapidly in frequency within populations over short timescales (Martof and Walton 1965). Range: The Blue Ridge Mountains of northeastern Georgia, northwestern South Carolina, and small portions of the nearby border regions of North Carolina, in headwater streams of the Chattahoochee, Chattooga, and Tallulah River drainages (Fig. 1; Voss et al. 1995; Jones et al. 2006). Elevational range ~ 400–1100m. Habitat: As with all shovel-nosed salamanders, occurs almost exclusively in the rocky riffle zones of a fast-flowing, high-gradient mountain streams. Etymology: The specific epithet is a Latin singular adjective in the nominative case meaning “adorned with gold,” in reference to the dorsal coloration (Fig. 7). Standard English Names: For Desmognathus aureatus: Golden Shovel-nosed Salamander (Conant 1958); for Leurognathus marmoratus roboratus Martof, 1956: Husky Shovel-nosed Salamander (Conant 1958). We suggest that “Southern” Shovel-nosed Salamander is more appropriate. Conservation: This species is apparently widespread and abundant at numerous historical and recent sites across its range, a significant proportion of which is protected by state, federal, and private entities. Consequently, we suggest that it be considered “LC—Least Concern” based on available data (Maes et al. 2015). Notes: Implicitly synonymized with Leurognathus marmoratus (Moore, 1899) by Martof (1962). Indirectly suggested for resurrection by Jones et al. (2006), though this was essentially ignored. Later explicitly resurrected by Dubois and Raffaëlli (2012), though few subsequent authorities followed this arrangement. Part of the Nantahala clade (Jones and Weisrock 2018; Pyron et al. 2020, 2022c) and the sister lineage of the Dwarf Black-bellied Salamander Desmognathus folkertsi Camp, Tilley, Austin, & Marshall, 2002. The presence of the shovel-nosed phenotype in this lineage may have been transmitted to or from the Pisgah clade via an episode of ancient adaptive introgression (Pyron et al. 2020). Within the Pisgah clade, Beamer and Lamb (2020) and Pyron et al. (2020, 2022c) sampled specimens from the type localities of Leurognathus marmorata intermedia Pope, 1928 and L. m. melania Martof, 1956, which were highly similar genetically and belonged to the marmoratus C lineage. Therefore, we conclude that marmoratus C should be referred to as Desmognathus intermedius (Pope, 1928), including D. melanius (Martof 1956) in synonymy. Consequently, the species D. melanius recognized by Frost (2019) following Dubois and Raffaëlli (2012) and Raffaëlli (2013) is not valid but is instead preoccupied by, Published as part of Pyron, R. Alexander & Beamer, David A., 2023, A systematic revision of the Shovel-nosed Salamander (Plethodontidae: Desmognathus marmoratus), with re-description of the related D. aureatus and D. intermedius, pp. 262-280 in Zootaxa 5270 (2) on pages 273-274, DOI: 10.11646/zootaxa.5270.2.5, http://zenodo.org/record/7850637, {"references":["Martof, B. S. (1956) Three new subspecies of Leurognathus marmorata from the southern Appalachian Mountains. Occasional Papers of the Museum of Zoology, University of Michigan, 575, 1 - 14.","Martof, B. S. (1962) Some aspects of the life history and ecology of the salamander Leurognathus. American Midland Naturalist, 67, 1 - 35. https: // doi. org / 10.2307 / 2422814","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes, 28, 77 - 161.","Jones, M. T., Voss, S. R., Ptacek, M. B., Weisrock, D. W. & Tonkyn, D. W. (2006) River drainages and phylogeography: an evolutionary significant lineage of shovel-nosed salamander (Desmognathus marmoratus) in the southern Appalachians. Molecular Phylogenetics and Evolution, 38, 280 - 287. https: // doi. org / 10.1016 / j. ympev. 2005.05.007","Martof, B. S. & Scott, D. C. (1957) The food of the salamander Leurognathus. Ecology, 38, 494 - 501. https: // doi. org / 10.2307 / 1929894","Martof, B. S. & Walton, K. G. (1965) A study of the unusual pigmentation in the salamander Leurognathus marmoratus. Physiological Zoology, 38, 349 - 352. https: // doi. org / 10.1086 / physzool. 38.4.30152412","Voss, S. R., Smith, D. G., Beachy, C. K. & Heckel, D. G. (1995) Allozyme variation in neighboring isolated populations of the plethodontid salamander Leurognathus marmoratus. Journal of Herpetology, 29, 493 - 497. https: // doi. org / 10.2307 / 1565011","Conant, R. (1958) A field guide to reptiles and amphibians. Houghton Mifflin, Boston, 598 pp.","Maes, D., Isaac, N. J. B., Harrower, C. A., Collen, B., van Strien, A. J. & Roy, D. B. (2015) The use of opportunistic data for IUCN Red List assessments. Biological Journal of the Linnean Society, 115, 690 - 706. https: // doi. org / 10.1111 / bij. 12530","Moore, J. P. (1899) Leurognathus marmorata, a new genus and species of salamander of the family Desmognathidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 51, 316 - 323.","Jones, K. S. & Weisrock, D. W. (2018) Genomic data reject the hypothesis of sympatric ecological speciation in a clade of Desmognathus salamanders. Evolution, 72, 2378 - 2393. https: // doi. org / 10.1111 / evo. 13606","Beamer, D. A. & Lamb, T. (2020) Towards rectifying limitations on species delineation in dusky salamanders (Desmognathus: Plethodontidae): an ecoregion-drainage sampling grid reveals additional cryptic clades. Zootaxa, 4734 (1), 1 - 61. https: // doi. org / 10.11646 / zootaxa. 4734.1.1","Pope, C. H. (1928) Some plethodontid salamanders from North Carolina and Kentucky, with the description of a new race of Leurognathus. American Museum Novitates, 306, 1 - 19.","Frost, D. R. (2019) Amphibian Species of the World: an Online Reference. Version 6.1. American Museum of Natural History, New York, New York. Available from: https: // amphibiansoftheworld. amnh. org / (accessed 1 January 2023)","Raffaelli, J. (2013) Les urodeles du monde. Deuxieme Edition. Plumelec, Penclen, 480 pp."]}

Desmognathus marmoratus (Moore, 1899) Leurognathus marmorata Moore, 1899 Syntypes and Type Locality: Moore (1899) ’s description was explicitly based on both a male and female syntype along with an additional female specimen which was skeletonized for illustration. These were taken from “a large clear rocky pool beneath a waterfall of a stream on the south flank of Grandfather Mt., N. C., and at an elevation of about 3,500 feet.” A site matching this description is found at Green Mountain Creek Falls (36.114308, -81.778430; 1182m) on U.S. Hwy 221 near Blowing Rock (NC: Watauga), which we suggest is the type locality. The male syntype was deposited by Moore in 1916 at the Academy of Natural Sciences of Philadelphia as ANSP 19610. The two females (the alcoholic syntype and the skeleton) were said by Dunn (1917) to have been examined by himself at the University of Pennsylvania, presumably still in the collection of Moore, who was still a professor there at the time. Fowler and Dunn (1917) listed ANSP 19610 as a paratype, while Malnate (1971) listed it as the holotype. It is neither, but rather a syntype as per the original publication. We here designate it as the lectotype of Leurognathus marmorata Moore, 1899 to remove any ambiguity. This renders the second, female syntype a paralectotype. This specimen is of unknown disposition; it was originally in the P.J. Moore collection at the University of Pennsylvania, but inquiries to the university and the ANSP yielded no information. The female skeleton is also not known to exist but is not part of the type series and carries no name-bearing function. Description: After Moore (1899) and Martof (1956), a large (SVL = ~ 30–75mm), highly aquatic, fin-tailed salamander with slit-like internal nares; an indistinct dorsal color pattern consisting of a dark-brown background color and numerous mottled markings lacking light-colored centers; darker or grey venter; adult males lacking vomerine teeth, females with very few; large, prominent eyes; and snout usually not darker than the remainder of body (Fig. 5, 6). Can putatively be differentiated from the other two described shovel-nosed species based on the original characters given in the other accounts below, though these have not been re-assessed based on modern understandings of species limits and are of unknown diagnostic utility. Based on our size-corrected linear morphometric analyses, this species can be distinguished from Desmognathus aureatus and D. intermedius by a shorter head (SG in original measurements = 4.3–16.4mm) and from D. intermedius by a shorter trunk (AG in original measurements = 9.3–41.7mm). Of the 10 specimens observed in West Virginia, 8 were adults and the largest (a gravid female) was only ~ 60mm SVL, suggesting that this population may represent a dwarf race since the median SVL of 45 metamorphosed individuals sampled from southern populations is ~ 62mm. Ironically, little is known about the natural history of this species despite it being the earliest-described species as most studies have focused on D. aureatus and D. intermedius. Pope (1924) found a clutch of 28 eggs on 6 August 1924, and Bishop (1924) sampled females in October with 26–34 ovarian eggs. Previous reports suggest that larvae typically metamorphose between 26–33mm SVL (see Camp and Tilley in Lannoo 2005 and references therein). Range: In high-gradient mountain streams of the Appalachians, on the Blue Ridge northeast of the Pigeon River to southwestern Virginia. Populations are known from the South Fork Holston, Watauga, and Nolichucky (Upper Tennessee) and Upper Broad and Upper Catawba (Santee) River drainages. Also known from a handful of riffle zones along a 2–3km stretch of Glade Creek near Beckley, Raleigh County, West Virginia in the Lower New (Kanawha) River drainage. We are confident that this species is absent from several other similar creeks in the New River Gorge in which we exerted similar sampling effort (Fig. 2). In contrast, other nearby streams with apparently similar geological and hydrological characteristics may harbor populations of this species. This population is widely (~120 airline km) disjunct from the nearest known populations in the Upper Tennessee drainage in southwestern Virginia (Smyth and Washington Counties). Elevational range ~ 450–1600m (see Fig. 1 for localities). Habitat: Occurs almost exclusively in the rocky riffle zones of a fast-flowing, high-gradient mountain stream of the “typical” character associated with other Shovel-nosed Salamanders farther south in the Appalachian Mountains (Martof 1962). Etymology: The specific epithet is a Latin singular adjective in the nominative case meaning “marbled,” in reference to the dorsal color-pattern. Standard English Names: Moore’s Triton (Brimley 1907), Moore’s Salamander (Bishop 1943), Northern Shovel-nosed Salamander (Schmidt 1953), Shovelnose Salamander (Collins et al. 1978). We suggest that “Northern Shovel-nosed Salamander” is most appropriate. Conservation: The re-described concept of this species occupies a smaller range than the previous, more inclusive definition but is nevertheless widespread and abundant at numerous historical localities. Consequently, we suggest that it retain its “LC—Least Concern” status on the IUCN Red List based on the known information (see Maes et al. 2015). Special attention should be paid to the isolated population in the New River Gorge of West Virginia. Notes: First assigned to Desmognathus by Titus and Larson (1996) after the unpublished results of Titus (1992) and the informal suggestion of Bernardo (1994). Genetic variation in the widespread former concept of this species was first reported by Voss et al. (1995) and expanded by Jackson (2005), Kozak et al. (2005), and Jones et al. (2006), who confirmed the existence of multiple species-level taxa based on phylogenetic analysis of mitochondrial and nuclear DNA. This echoed earlier morphological work (Pope 1928; Martof 1956), though Martof (1962) had later downplayed the significance of this phenotypic variation. Accordingly, we offer revised accounts of the other known species of shovel-nosed salamander. These occur in two distantly related clades, Nantahala (Desmognathus aureatus) and Pisgah (D. intermedius). Beamer and Lamb (2020) and Pyron et al. (2020, 2022c) sampled the type locality of Leurognathus m. aureata Martof, 1956 and populations geographically proximate (5.5 airline km) to the type locality of L. m. roborata Martof, 1956, confirming their inclusion in marmoratus B. Acting as the First Reviser (Article 24.2.1), Raffaëlli (2013) had previously selected the former as the senior subjective synonym when the two taxa are considered conspecific. Consequently, this lineage should be referred to as Desmognathus aureatus (Martof, 1956) and include D. roboratus (Martof, 1956) in synonymy., Published as part of Pyron, R. Alexander & Beamer, David A., 2023, A systematic revision of the Shovel-nosed Salamander (Plethodontidae: Desmognathus marmoratus), with re-description of the related D. aureatus and D. intermedius, pp. 262-280 in Zootaxa 5270 (2) on pages 270-273, DOI: 10.11646/zootaxa.5270.2.5, http://zenodo.org/record/7850637, {"references":["Moore, J. P. (1899) Leurognathus marmorata, a new genus and species of salamander of the family Desmognathidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 51, 316 - 323.","Dunn, E. R. (1917) The salamanders of the genera Desmognathus and Leurognathus. Proceedings of the United States National Museum, 53, 393 - 433. https: // doi. org / 10.5479 / si. 00963801.53 - 2211.393","Fowler, H. W. & Dunn, E. R. (1917) Notes on salamanders. Proceedings of the Academy of Natural Sciences of Philadelphia, 69, 7 - 28.","Malnate, E. V. (1971) A catalog of primary types in the herpetological collections of the Academy of Natural Sciences, Philadelphia (ANSP). Proceedings of the Academy of Natural Sciences of Philadelphia, 123, 345 - 375.","Martof, B. S. (1956) Three new subspecies of Leurognathus marmorata from the southern Appalachian Mountains. Occasional Papers of the Museum of Zoology, University of Michigan, 575, 1 - 14.","Pope, C. H. (1924) Notes on North Carolina salamanders with especial reference to the egg-laying habits of Leurognathus and Desmognathus. American Museum Novitates, 153, 1 - 16.","Bishop, S. C. (1924) Notes on salamanders. New York State Museum Bulletin, 253, 87 - 102.","Martof, B. S. (1962) Some aspects of the life history and ecology of the salamander Leurognathus. American Midland Naturalist, 67, 1 - 35. https: // doi. org / 10.2307 / 2422814","Brimley, C. S. (1907) The salamanders of North Carolina. Journal of the Elisha Mitchell Scientific Society, 23, 150 - 156.","Bishop, S. C. (1943) Handbook of Salamanders. Comstock, Ithaca, New York, 555 pp.","Schmidt, K. P. (1953) A Check List of North American Amphibians and Reptiles. 6 th Edition. University of Chicago Press, Chicago, Illinois, 280 pp.","Collins, J. T., Huheey, J. E., Knight, J. L. & Smith, H. M. (1978) Standard common and current scientific names for North American amphibians and reptiles. SSAR Herpetological Circular, 7, 1 - 36.","Maes, D., Isaac, N. J. B., Harrower, C. A., Collen, B., van Strien, A. J. & Roy, D. B. (2015) The use of opportunistic data for IUCN Red List assessments. Biological Journal of the Linnean Society, 115, 690 - 706. https: // doi. org / 10.1111 / bij. 12530","Titus, T. A. & Larson, A. (1996) Molecular phylogenetics of desmognathine salamanders (Caudata: Plethodontidae): a reevaluation of evolution in ecology, life history, and morphology. Systematic Biology, 45, 451 - 472. https: // doi. org / 10.1093 / sysbio / 45.4.451","Titus, T. A. (1992) A phylogenetic analysis of the Desmognathinae (Caudata: Plethodontidae): evolutionary patterns inferred from mitochondrial DNA sequences. PhD Thesis, University of Kansas, Lawrence, Kansas, 119 pp.","Bernardo, J. (1994) Experimental analysis of allocation in two divergent, natural salamander populations. The American Naturalist, 143, 14 - 38. https: // doi. org / 10.1086 / 285594","Voss, S. R., Smith, D. G., Beachy, C. K. & Heckel, D. G. (1995) Allozyme variation in neighboring isolated populations of the plethodontid salamander Leurognathus marmoratus. Journal of Herpetology, 29, 493 - 497. https: // doi. org / 10.2307 / 1565011","Jackson, N. D. (2005) Phylogenetic history, morphological parallelism, and speciation in a complex of Appalachian salamanders (genus: Desmognathus). M. S. Thesis, Brigham Young University, Provo, Utah, 64 pp.","Kozak, K. H., Larson, A., Bonett, R. M. & Harmon, L. J. (2005) Phylogenetic analysis of ecomorphological divergence, community structure, and diversification rates in dusky salamanders (Plethodontidae: Desmognathus). Evolution, 59, 2000 - 2016. https: // doi. org / 10.1111 / j. 0014 - 3820.2005. tb 01069. x","Jones, M. T., Voss, S. R., Ptacek, M. B., Weisrock, D. W. & Tonkyn, D. W. (2006) River drainages and phylogeography: an evolutionary significant lineage of shovel-nosed salamander (Desmognathus marmoratus) in the southern Appalachians. Molecular Phylogenetics and Evolution, 38, 280 - 287. https: // doi. org / 10.1016 / j. ympev. 2005.05.007","Pope, C. H. (1928) Some plethodontid salamanders from North Carolina and Kentucky, with the description of a new race of Leurognathus. American Museum Novitates, 306, 1 - 19.","Pyron, R. A., O'Connell, K. A., Lemmon, E. M., Lemmon, A. R. & Beamer, D. A. (2022 c) Candidate-species delimitation in Desmognathus salamanders reveals gene flow across lineage boundaries, confounding phylogenetic estimation and clarifying hybrid zones. Ecology and Evolution, 12, e 8574. https: // doi. org / 10.1002 / ece 3.8574","Beamer, D. A. & Lamb, T. (2020) Towards rectifying limitations on species delineation in dusky salamanders (Desmognathus: Plethodontidae): an ecoregion-drainage sampling grid reveals additional cryptic clades. Zootaxa, 4734 (1), 1 - 61. https: // doi. org / 10.11646 / zootaxa. 4734.1.1","Raffaelli, J. (2013) Les urodeles du monde. Deuxieme Edition. Plumelec, Penclen, 480 pp."]}

Pyron, R. Alexander, Beamer, David A. (2023): A systematic revision of the Shovel-nosed Salamander (Plethodontidae: Desmognathus marmoratus), with re-description of the related D. aureatus and D. intermedius. Zootaxa 5270 (2): 262-280, DOI: 10.11646/zootaxa.5270.2.5, URL: http://dx.doi.org/10.11646/zootaxa.5270.2.5

Siren intermedia Barnes, 1826 (Figs. 1‒7, 9, 10, & 13) Common name. Intermediate Siren Neotype: UF Herp 190369 (Fig. 7A), adult female (gravid) from Wayne Co., Georgia, USA (31.50109°N, 81.91324°W, datum WGS84, elev. 13 m). Collected on 30 January 2020 by Dirk Stevenson, Arik Hartmann, and Matthew Fedler. Description of neotype: The neotype has 32 costal grooves and faint black spots on the dorsum extending from the head to the fourth costal groove behind the forelimbs. In life, it had a bluish gray venter and sides and a dark grayish brown dorsum. The slightly darker tail tip may indicate old regeneration. A broken chartreuse labial stripe runs from a few millimeters posterior of the nares to just anterior of the gills. Light yellow spots are present ventrolaterally on the head, often surrounding sensory pores. Sparse and more randomly placed small (Paratypes: UF Herp 188598, 189657 (Fig. 7B), 188604 (Fig. 7C), 188605 (Fig. 7D), 186989 (Fig. 7F), 190370 (Fig. 7E), 190371, 190374, 190375. Locality information for these specimens can be found via the Florida Museum of Natural History’s web database (http://specifyportal.flmnh.ufl.edu/herps/). Diagnosis. Siren intermedia has typical Siren characteristics: external gills with three fimbriate gill stalks, three associated gill slits, four toes on the forelimbs, lack of pelvic girdle and hindlimbs, and a thin, pigment-bearing mucus layer that overlies the keratinized skin. This species is best distinguished from other sympatric Siren lineages by costal groove count, which varies geographically. We restrict this species to the recovered S. intermedia lineages A, B and C. All S. intermedia lineages combined that we examined have 29‒33 costal grooves (Fig. 4). Lineage B has a more restricted costal groove range of 29 (n =12), 30 (n = 23), or 31 (n = 3) in areas of sympatry with S. sphagnicola., which has 31‒33 costal grooves in the East Bay, Yellow, Blackwater, and Escambia river drainages. Specimens of lineage B that are comparable in size to S. sphagnicola typically have yellow labial stripes, rostral patches, and ventrolateral flecking (Fig. 10). Siren intermedia lineages A (type clade) (Fig. 7) and C (Fig. 10) can be distinguished from sympatric Siren species by having fewer than 34 costal grooves. These two lineages exhibit hypervariability in pattern and coloration. Yellowish labial stripes and rostral patches may be present or absent. Numerous distinct black spots may be present, absent, or occur in patches on the dorsal and lateral surfaces of the head and body, and some individuals possess reticulated patterns (UF Herp 186989, 192033, and an unvouchered specimen from the same locality as 192033). These traits are also present in other Siren species and may be variable within populations of any of these species, emphasizing the need for accurate costal groove counts to distinguish from other species in the vicinity. Original description, first redescription, and comments. Le Conte (1828) provided the following description: “Its greatest length is twelve inches: colour uniform dusky, very slightly paler beneath, sometimes faintly speckled with darker above.” Barnes (1826) and Harlan (1827) announced that the species was going to be described and provided the same information when summarizing Le Conte’s forthcoming manuscript.According to Harlan (1827): “Colour resembling that of the Lacertina; branchiae resembling those of the Striata [i.e., Pseudobranchus striatus Le Conte]. Length about one foot.” We designated a neotype from near the type locality for the express purpose of clarifying the taxonomic status of a nominal taxon whose specimens (syntypes) were destroyed. Smith et al. (1975) clarified that the name should be attributed to Barnes (1826). Le Conte’s (1828) description provided additional traits that are generic among Siren and failed to identify any features that distinguish it from other Siren given our current understanding of the genus. Siren intermedia is extremely variable in both color and pattern, and many phenotypes overlap those of S. lacertina and S. nettingi. The original type locality was purportedly Le Conte’s Woodmanston Plantation, Riceboro, Georgia (Harper 1935). We chose the neotype locality, which is approximately 45 km SW of Woodmanston Plantation, because all sequenced specimens from the surrounding region and Altamaha River drainage belong to the same mtDNA lineage as our single Woodmanston Plantation specimen. The neotype locality is easily accessible and yielded many S. intermedia compared to other localities in the area. Additionally, the locality has stereotypical habitat for S. intermedia consisting of a cypress swamp with many small creeks and large accumulations of muck, leaf packs, and submerged Creeping Rush (Juncus repens [Michaux]). Noble and Marshall (1932) provided a redescription of S. intermedia but failed to state where the specimens they examined came from, and they likely included specimens from populations later split into the nettingi and texana subspecies of S. intermedia. This would explain their higher costal groove range for S. intermedia (up to 36 grooves). Additionally, they stated that the outer capsule of the egg was wider for S. lacertina than S. intermedia. This may ultimately be proven true by future studies, but their data are contradictory in light of the revised Siren taxonomy. Noble and Marshall (1932) compared specimens of what they considered to be S. intermedia and S. lacertina from the same locality (Maverick Co., Texas) to each other, as well as S. intermedia and S. lacertina from Oakley, Berkeley Co., South Carolina. A recent genetic study (LaFortune 2015) found that the Rio Grande drainage has only one genetic group of Siren; thus, Noble and Marshall (1932) and Flores-Villela and Brandon (1992) were comparing large and smaller individuals of the same species (now S. nettingi) using different names. Their findings indicate that different-sized individuals of the same species may have different-sized eggs, or they were comparing specimens with eggs at various stages of development. Size. The largest specimen we examined (UF Herp 186989) measured 191 mm SVL and 125 mm TaL (316 mm TL). Noble & Marshall (1932) reported a male S. intermedia (MCZ Herp A-140) from Georgetown, South Carolina, that measured 212 mm SVL and 347 mm TL. This animal is likely a member of S. intermedia lineage C. Goin (1957) provided 240 mm SVL and 381 mm TL for the maximum size of “ S. i. intermedia ” specimens he examined. However, he failed to provide the collecting locality or attribute the measurements to a vouchered specimen; thus, we are unable to assign it to a lineage. Because Goin assumed that only S. i. intermedia was present in the coastal drainages of Mississippi, Alabama, and the Florida Parishes of Louisiana, he may have attributed the size to a specimen that we now recognize as S. nettingi. Furthermore, his measurements closely match the size of the largest S. nettingi specimen we sequenced (LSUMZ 87289; 270 mm SVL, 408 mm TL) and the largest S. nettingi in the AUM collection (AUM 40435; 242 mm SVL, 365 mm TL). The smallest sexually mature female (UF Herp 190904) measured 75 mm SVL and 119 mm TL. Distribution. Our findings restrict S. intermedia to Atlantic and Gulf drainages from the vicinity of Chesapeake Bay south to central Florida and west throughout the Florida panhandle (Fig. 9). Specimens attributed to this species from the Mobile Bay drainage westward are likely either S. nettingi or S. sphagnicola. Common name. The first common name assigned to S. intermedia was Intermediate Siren (Gray 1831), and we suggest using this name instead of Lesser Siren because there are three larger sirens (S. lacertina, S. reticulata, S. nettingi) and three smaller sirens (S. sphagnicola, Pseudobranchus axanthus, P. striatus). This also aligns the epithet with the common name. Specimens examined. See Supplemental Table 1., Published as part of Fedler, Matthew T., Enge, Kevin M. & Moler, Paul E., 2023, Unraveling Siren (Caudata: Sirenidae) systematics and description of a small seepage specialist, pp. 351-378 in Zootaxa 5258 (4) on pages 369-374, DOI: 10.11646/zootaxa.5258.4.1, http://zenodo.org/record/7784322, {"references":["Barnes, D. H. (1826) An arrangement of the genera of batracian [sic] animals, with a description of the more remarkable species; including a monograph of the doubtful reptils [sic]. American Journal of Science and Arts, 11 (2), 268 - 297.","Le Conte, J. E. (1828) Description of a new species of Siren. Annals of the Lyceum of Natural History of New York, 2 (1), 133 - 134. https: // doi. org / 10.1111 / j. 1749 - 6632.1826. tb 00242. x","Harlan, R. (1827) Genera of North American Reptilia, and a synopsis of the species. Journal of the Academy of Natural Sciences of Philadelphia, 5, 317 - 372.","Smith, H. M., Smith, R. B. & Sawin, H. L. (1975) The authorship and date of publication of Siren intermedia (Amphibia: Caudata). Great Basin Naturalist, 35 (1), 100 - 102.","Harper, F. (1935) Records of amphibians in the southeastern states. The American Midland Naturalist, 16 (3), 275 - 310. https: // doi. org / 10.2307 / 2420030","Noble, G. K. & Marshall, B. C. (1932) The validity of Siren intermedia Le Conte, with observations on its life history. American Museum Novitates, 532, 1 - 17.","LaFortune, T. C. (2015) Species Identification and Habitat Assessment of the South Texas Siren. M. S. Thesis, University of Texas at Brownsville, Brownsville, Texas, 91 pp.","Flores-Villela, O. & Brandon, R. A. (1992) Siren lacertina (Amphibia: Caudata) in northeastern Mexico and southern Texas. Annals of the Carnegie Museum, 61, 289 - 291. https: // doi. org / 10.5962 / p. 226656","Goin, C. J. (1957) Description of a new salamander of the genus Siren from the Rio Grande. Herpetologica, 13 (1), 37 - 42. https: // doi. org / 10.2307 / 1439392","Gray, J. E. (1831) A synopsis of the species of the Class Reptilia. In: Griffith, E. & Pidgeon, E. (Ed.), The Animal Kingdom arranged in conformity with its organization, by the Baron Cuvier, member of the Institute of France, with additional descriptions of all the species hitherto named, and of many not before noticed. Volume the Ninth. The Class Reptilia arranged by the Baron Cuvier, with specific descriptions. Appendix. Whittaker Treacher, and Co. London, pp. 1 - 86 + 1 - 110 (Appendix), pls. 1 - 55."]}

Siren sphagnicola sp. nov. (Figs. 1‒6, 11–14) Common name. Seepage Siren Holotype. UF Herp 185209 (Fig. 11A), adult female from Junior Walton Pond in Okaloosa Co., Florida, USA (30.69270°N, 86.47250°W, datum WGS84, elev. 30 m) (Fig. 12 A & 12B). Collected on 18 January 2019 by Matthew Fedler, Paul Moler, and Pierson Hill. Paratypes. UF Herp 161516, 162498, 162568, 163271, 164240, 164241, 164242, 164243, 184285, 185195, 185200, 185201, 185208, 185209, 185214, 185215, 188766, 188767, 190036, 190037, 185205, 185216, 185197, 185198, 185210. Locality information for the various paratype localities is available via FLMNH’s UF Herpetology database (http://specifyportal.flmnh.ufl.edu/herps/). Description of holotype. The holotype has 32 costal grooves and faint black dorsal spots extending from the head to the forelimbs. It lacks approximately 3 mm of its tail tip. In life, it was mouse gray on its venter and sides with a grayish brown dorsum. Sensory pits on the head are well defined and beige in color. Measurements are 97 mm SVL, 49 mm TaL, 3.9 mm interorbital distance, 7.1 mm head width, 11.1 mm head length, 3.6 mm eye-snout distance, and 4.4 mm chest width. Diagnosis. Siren sphagnicola has typical Siren characteristics: external gills with three fimbriate gill stalks, three associated gill slits, four toes on the forelimbs, lack of pelvic girdle and hindlimbs, and a thin, pigment-bearing mucus layer that overlies the keratinized skin. A combination of traits distinguishes it from other members of the genus. It has 30‒33 costal grooves (Fig. 3) and a mouse gray base color with occasionally a light, grayish brown sheen on the dorsum (Fig. 11). Small juveniles in the post-macrocephalic larval stage, which is>2 months of age according to diagrams of growth/transition rates of S. nettingi provided by Noble & Marshall (1932), have the same gray coloration as adults (Fig. 11) and lack the orange, red, or yellow highlights present on other Siren juveniles of similar age (Fig. 13). A few adult specimens examined have small, well-defined black spotting on the head and occasionally on the dorsum (Fig. 11). Sensory pits on the head are more visible than those on heads of other Siren species and are typically ivory to beige colored, which may denote an absence of gray pigment rather than the presence of chromatophores (Fig. 11). This species lacks the yellow labial stripe present in young S. lacertina (Figs. 13B & 13F), S. intermedia (Figs. 13A, 13C, 13G, & 13I), and S. nettingi examined from the Mobile Bay drainage. Some juvenile S. intermedia in eastern populations also lack the light labial stripe. A few small juvenile S. sphagnicola have yellow spots or a short, broken stripe where a labial stripe is present in other species (Figs. 13E & 13H). Siren sphagnicola also lacks the post-cranial yellow or gold flecking found in many S. lacertina (Fig. 8), S. intermedia (Fig. 10), and S. nettingi. Gill stalk coloration is typically rosy pink to red in recently captured specimens but fades to grayish pink in captivity, likely due to changes in acidity or oxygenation of water. Intact tail tips are rounded, whereas partially regenerated tails (frequently observed) often taper to an abrupt point after the tail fin blade (Fig. 11). Regenerated portions of the tail seem to lack the density of gray pigment found in non-regenerated portions; thus, the regenerated portion is easily distinguished by its pinkish gray hue. Regenerated portions of the tail of other Siren species examined match the normal body coloration or have a brownish hue. Size. Siren sphagnicola is the smallest known species in the genus Siren. Additionally, all specimens examined are shorter than the maximum length given for both species of Pseudobranchus (Moler 2019b, c), making S. sphagnicola the smallest member of Sirenidae based on our current understanding. The largest specimen examined (AUM 27973) had an SVL of 126 mm, but it lacked a complete tail. The largest specimen with a complete tail (AUM 8960) had an SVL of 120 mm and a TaL of 76 mm (196 mm TL). We attributed these AUM specimens to S. sphagnicola based upon costal groove count, lack of labial striping and gold flecking found on sympatric S. nettingi and S. intermedia, and presence of beige-colored facial pores. Reproductive females have been found as small as 71 mm SVL (111 mm TL). When comparing measurement distributions of Siren lineages, S. sphagnicola was not distinct from any other single lineage in, at most, two of seven measurements (Table 4). Natural history and distribution. Based on our surveys in Florida, populations appear to be robust and widely distributed in suitable microhabitats in the Blackwater and Yellow river drainages and the western two-thirds of Eglin Air Force Base, including several streams that flow into the western side of Choctawhatchee Bay (Fig. 14). This suspected microhabitat specialist has been found in headwater seepage areas of steephead streams, mucky seeps farther downstream, muddy and/or densely vegetated seepage bogs, shallow-water depressions lined with dense sphagnum moss or filled with leaves along seepage-fed streamside terraces, and other types of shallow streams with mucky, detrital, or sandy bottoms (Enge 2005) (Fig. 12). In contrast, S. intermedia collected at localities near (S. sphagnicola were found in leaf packs not associated with seeps and adjacent to deeper water. Incised (gully-eroded) first- and second-order streams (Strahler 1964) lack the microhabitats used by S. sphagnicola (and many other salamander species), because accumulations of leaf litter and other detritus are constantly flushed from streams and scoured from surface pools by heavy rainfall events. Common, syntopic amphibian species are the Southern Cricket Frog (Acris gryllus [Le Conte]), Bronze Frog (Lithobates clamitans Latreille), Southern Two-lined Salamander (Eurycea cirrigera [Green]), and Southern Red Salamander (Pseudotriton ruber vioscai Bishop). Onetoed Amphiuma (Amphiuma pholeter Neill) and Two-toed Amphiuma (A. means Garden) may be present but are less abundant than the aforementioned species (Enge 2005). Deep, steephead ravine systems (Means 1981, 2000) and more shallow-gradient, seepage bogs in upland habitats near the Gulf of Mexico may have served as “evolutionary engines” during periods of elevated sea levels, producing the Florida Bog Frog (Lithobates okaloosae Moler, 1985), Bog Dwarf Salamander (Eurycea sphagnicola Wray, Means, & Steppan, 2017), and S. sphagnicola. A sea level rise of only 2‒5 m would have led to saltwater inundation of the mouths of these deep steephead valleys, thus isolating ancestral populations of freshwater species (Means 2000). We suspect the range of S. sphagnicola is similar to that of E. sphagnicola, which also inhabits the sphagnum-lined margins of streams and associated seepage habitats (Wray et al. 2017). Siren sphagnicola has a smaller geographic distribution than other Siren species. Most specimens have been found in the Blackwater,Yellow, and Escambia/Conecuh river drainages of Florida and Alabama (Fig. 14). Elsewhere, its range is poorly known, but we believe it is restricted to the environs of sandy, seepage-fed creeks in the lower Gulf Coastal Plain as far west as the Florida Parishes of Louisiana (Fig. 14). Locality information from outside Florida is entirely based on preserved AUM specimens and sequence data from a GenBank specimen that match both mtDNA and scnDNA markers. Few Siren museum vouchers with genetic material exist from Mississippi (42 total specimens via Vertnet search and only one with available tissue, which we sequenced) and the Florida Parishes of Louisiana (63 total via Vertnet search; one of two tissues requested yielded DNA), and we did not examine most museum specimens from this area that lacked tissue samples. Etymology and common name. The specific epithet is derived from Sphagnum, the generic name for sphagnum moss, and the Latin suffix -cola, meaning inhabitant or dweller. The species epithet is used as noun in apposition. This siren is frequently found in and under mats of Sphagnum in and along streams and the margins of other bodies of water. Because of its affinity for seepage-fed streams and wetlands, we suggest Seepage Siren as the common name. Specimens examined. See Supplemental Table 1.

Siren nettingi (Goin, 1942) (Figs. 1‒6 & 9) Common name. Western Siren Holotype. Carnegie Museum 7580. Adult female collected in May 1928 from Imboden, Lawrence Co., Arkansas, USA (Goin 1942). Paratypes. See Goin (1942). Diagnosis. Siren nettingi has typical Siren characteristics: external gills with three fimbriate gill stalks, three associated gill slits, four toes on the forelimbs, lack of pelvic girdle and hindlimbs, and a thin, pigment-bearing mucus layer that overlies the keratinized skin. Goin (1942) distinguished S. nettingi from S. intermedia by the presence of well-defined light spots on the sides and venter and also specifying that these were not the bar-like streaks found in S. lacertina. Additionally, Goin (1942, p. 212) stated that S. nettingi has “about two more costal grooves (usually 33 in intermedia, 35 in nettingi),” a value slightly overlapping but less than the average for S. lacertina. See comments below for possible reasons for variation in data. In areas of sympatry, the costal groove range of S. nettingi overlaps that of S. sphagnicola. Siren nettingi has 32‒34 costal grooves, with two outliers of 31 and 36 (Fig. 4). Goin (1942) gave a slightly higher range of 33‒37 costal grooves, but we infer that he included the axial groove or terminated counting grooves at a point posterior to our stopping point. Siren nettingi specimens similar in size to S. sphagnicola can be distinguished by the presence of a solid yellow labial stripe and rostral patch, whereas S. sphagnicola has, at most, several beige spots where the labial stripe occurs in other species. In addition, bodies of small S. nettingi often have a green or yellowish hue and gold flecking, both of with are lacking in S. sphagnicola. Comments. This species occupies a large geographic area, and we analyzed relatively few specimens, mainly from Alabama and Louisiana (Fig. 9). Our information may not represent the full extent of variation found in this species. Reinhard et al. (2013) provided photographs of larvae and mature adults of this species. After examining the holotype, we concluded that the well-defined, light spots on the venter referenced by Goin (1942) represent sensory pores rather than chromatophores. Chromatophores on the holotype are neither defined nor extensive in coverage. Regardless of this distinction, neither the distinct, light-colored sensory pores nor the yellow to yellow-green flecking of chromatophores is unique to S. nettingi. These pores are frequently obfuscated by the slime layer but become more visible when this is removed. This slime layer frequently sloughs off of poorly preserved or frozen specimens. Validity of the Rio Grande Siren (S. intermedia texana Goin, 1957) as a taxonomic unit distinct from S. nettingi has been questioned in recent decades. Our analyses suggest that S. nettingi and “texana” are either closely related sister taxa or ecomorphs of the same species adapted to habitats present in different regions. Further investigation of this topic is needed, but we treated “texana” as a distinct unit for morphological analyses. Furthermore, S. i. “texana” is no longer recognized by Highton et al. (2017), who accepted the erroneous designation by Flores-Villela and Brandon (1992) that the large siren from southern Texas and Mexico is S. lacertina (LaFortune 2015). Size. Siren nettingi has a reported maximum TL of 686 mm for the larger “texana” form found in southern Texas and Mexico, whereas the smaller form found in the rest of its range has a reported maximum TL of 502 mm (Martof 1973a). Flores-Villela and Brandon (1992) cited Goin (1957) for total lengths given by Martof (1973a and 1973b) and incorrectly reported the TL as the SVL for both the “texana” form of S. nettingi (then S. intermedia texana) and S. lacertina. Distribution. Based on our examination of museum specimens and genetic analyses, S. nettingi occurs from the Mobile Bay drainage westward, and native populations do not occur in any Florida drainage (Fig. 9). A single specimen that sequenced as S. nettingi was collected from Tates’s Hell State Forest, Franklin Co., Florida, but we suspect this specimen was translocated as live fish bait. Based on phenotypes and costal groove counts, Goin (1942) suggested that Florida Parishes of Louisiana represented the break between the then-recognized subspecies intermedia and nettingi. This boundary was later considered to be a hybridization zone that stretched into Mississippi (Boyd & Vickers 1963) and then to the Mobile Bay drainage (Caldwell & Howell 1966). We suspect part of this confusion resulted from examining S. sphagnicola and attributing them to S. i. intermedia because of their resemblance to grayish S. intermedia that can be found farther east. Furthermore, S. nettingi is currently thought to be the only Siren species that occurs from the Mississippi River west and south to Veracruz, Mexico. Common Name. We suggest the common name Western Siren. Except for the Mobile Bay drainage system, where S. reticulata is present, S. nettingi represents the largest or only siren (from the Mississippi River westward) present throughout its range; thus, the “Lesser” moniker is misleading. Specimens examined. See Supplemental Table 1., Published as part of Fedler, Matthew T., Enge, Kevin M. & Moler, Paul E., 2023, Unraveling Siren (Caudata: Sirenidae) systematics and description of a small seepage specialist, pp. 351-378 in Zootaxa 5258 (4) on pages 368-369, DOI: 10.11646/zootaxa.5258.4.1, http://zenodo.org/record/7784322, {"references":["Goin, C. J. (1942) Description of a new race of Siren intermedia Le Conte. Annals of the Carnegie Museum, 29, 211 - 217. https: // doi. org / 10.5962 / p. 215158","Reinhard, S., Voitel, S. and Kupfer, A. (2013) External fertilisation and paternal care in the paedomorphic salamander Siren intermedia Barnes, 1826 (Urodela: Sirenidae). Zoologischer Anzeiger-A Journal of Comparative Zoology, 253 (1), 1 - 5. https: // doi. org / 10.1016 / j. jcz. 2013.06.002","Goin, C. J. (1957) Description of a new salamander of the genus Siren from the Rio Grande. Herpetologica, 13 (1), 37 - 42. https: // doi. org / 10.2307 / 1439392","Highton, R., Bonett, R. M. & Jockusch, E. L. (2017) Caudata salamanders. In: Crother, B. I. (Ed.), Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in our Understanding. Society for the Study of Amphibians and Reptiles Herpetological Circular No. 43. 8 th Edition. Society for the Study of Amphibians and Reptiles, s. n., pp. 18 - 30.","Flores-Villela, O. & Brandon, R. A. (1992) Siren lacertina (Amphibia: Caudata) in northeastern Mexico and southern Texas. Annals of the Carnegie Museum, 61, 289 - 291. https: // doi. org / 10.5962 / p. 226656","LaFortune, T. C. (2015) Species Identification and Habitat Assessment of the South Texas Siren. M. S. Thesis, University of Texas at Brownsville, Brownsville, Texas, 91 pp.","Martof, B. S. (1973 a) Siren intermedia Le Conte. Lesser Siren. Catalogue of American Amphibians and Reptiles, 127, 1 - 3.","Martof, B. S. (1973 b) Siren lacertina Linnaeus. Greater Siren. Catalogue of American Amphibians and Reptiles, 128, 1 - 2.","Boyd, C. & Vickers, D. H. (1963) Distribution of some Mississippi amphibians and reptiles. Herpetologica, 19 (3), 202 - 205.","Caldwell, R. D. & Howell, W. M. (1966) Siren intermedia nettingi from Alabama. Herpetologica 22 (4), 310 - 311."]}