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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.

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 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., 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 366-368, DOI: 10.11646/zootaxa.5258.4.1, http://zenodo.org/record/7784322, {"references":["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.","Moler, P. E. (2019 b) Pseudobranchus axanthus Nettingi and Goin 1942, Southern Dwarf Siren. In: Krysko, K. L., Enge, K. M. & Moler, P. E. (Eds.), Amphibians and Reptiles of Florida. University of Florida Press, Gainesville, Florida, pp. 124 - 126.","Enge, K. M. (2005) Herpetofaunal drift-fence surveys of steephead ravines in the Florida panhandle. Southeastern Naturalist, 4 (4), 657 - 678. https: // doi. org / 10.1656 / 1528 - 7092 (2005) 004 [0657: HDSOSR] 2.0. CO; 2","Strahler, A. N. (1964) Section 4 - II. Geology. Part II. Quantitative geomorphology of drainage basins and channel networks. In: Te Chow, V. (Ed.), Handbook of Applied Hydrology. McGraw-Hill, New York, New York, pp. 39 - 76.","Means, D. B. (1981) Steepheads: Florida's little-known canyon lands. ENFO (Florida Conservation Foundation), 81 (6), 1 - 4.","Means, D. B. (2000) Southeastern U. S. Coastal Plain habitats of the Plethodontidae: the importance of relief, ravines, and seepage. In: Bruce, R. C., Jaeger, R. J. & Houck, L. D. (Eds.), The Biology of Plethodontid Salamanders. Plenum, New York, New York, pp. 287 - 302. https: // doi. org / 10.1007 / 978 - 1 - 4615 - 4255 - 1 _ 14","Moler, P. E. (1985) A new species of frog (Ranidae: Rana) from northwestern Florida. Copeia, 1985 (2), 379 - 383. https: // doi. org / 10.2307 / 1444847","Wray, K. P., Means, D. B. & Steppan, S. J. (2017) Revision of the Eurycea quadridigitata (Holbrook 1842) complex of dwarf salamanders (Caudata: Plethodontidae: Hemidactyliinae) with a description of two new species. Herpetological Monographs, 31 (1), 18 - 46. https: // doi. org / 10.1655 / HERPMONOGRAPHS-D- 16 - 00011"]}

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."]}

Calamaria dominici Ziegler, Tran & Nguyen sp. nov. Figs1-4 Holotype: IEBR A.2018.1, an adult female collected on 28 May 2017 at 11:30 on a forest path by Anh Vu Tran in evergreen mixed forest of broadleaf and conifer trees within Ta Dung Nature Reserve, Dak Nong Province, Central Highlands, Vietnam, at an elevation of 1240 m asl. Diagnosis: A species of the genus Calamaria cha- racterized by the combination of the following characters: (1) rostral wider than high; (2) paraparietal surrounded by six shields and scales; (3) eye diameter larger than eye-mouth distance; (4) preocular present; (5) supralabials 5/4, 3-4/2-3 entering orbit; (6) maxillary teeth nine, modified; (7) infralabials 5/4, first three touching anterior chin shields; (8) mental touching tip of right anterior chin shield; (9) ventrals 1 + 174; subcaudal scales 18/17, divided; (10) precloacal plate single; (11) tail relatively short (6.2% of the total length), nearly as thick as body, slightly tapering, and ending in obtuse point; (12) dorsal scales reducing to six rows at position above 4th subcaudal, and to four rows above 13th subcaudal on tail; (13) dorsum dark with irregular yellow blotches; and (14) ventral side dark with few yellow blotches and bands. Description of holotype: Habitus vermiform; head indistinct from neck; pupil round; tail relatively short (6.2% of the total length), nearly as thick as body, slightly tapering, ending in obtuse point. Size. SVL: 395 mm; TaL: 26 mm; TL: 421 mm; ratio TaL/TL: 0.06. Dentition. Right upper maxilla with 9 modified maxillary teeth. Body scalation. Dorsal scale rows 13���13���13, smooth. Dorsal scales reducing to six rows at position above fourth subcaudal, to five rows above 12th subcaudal, and to four rows above 13th subcaudal on tail. 174 ventrals (+ 1 preventral); 18/17 subcaudals, all paired, first pair not in contact, followed by tail tip; precloacal single. Head scalation. Rostral wider than high, portion visible from above shorter than prefrontal suture. Prefrontal shorter than frontal, not entering orbit, and touching first two supralabials on right side and second and third supralabial on left side. Frontal hexagonal, nearly two times maximum width of supraocular. Paraparietal surrounded by six shields and scales. Length of parietal shorter than distance from posterior tip of frontal to posterior tip of rostral. Supraocular 1/1. Loreal 1/1. Preocular 1/1, distinctly higher than wide. Postocular 1/1, higher than wide, not as high as eye diameter. Eye diameter larger than eye-mouth distance. 5/4 supralabials, on left side third and fourth entering orbit, fifth longest; on right side second and third entering orbit, fourth longest. Mental triangular, touching tip of right anterior chin shields. 5/4 infralabials, first three touching anterior chin shields. First pair of chin shields in contact mesially, second pair touching anteriorly and separated posteriorly. Coloration (in life). Eye black; tongue grey anteriorly, pinkish-grey posteriorly; dorsum of body and tail dark purplish-black, iridescent; head with irregular small and few medium-sized yellow blotches; body with mediumsized to large yellow blotches, irregularly arranged, in part forming transverse or oblique rows, sometimes in zig-zag pattern; hind part of body dorsum and dorsal surface of tail less intensely blotched; venter purplishblack with irregular yellow transverse bands or blotches; chin and throat region dark with yellow reticulation or blotches and bands; lower tail surface dark with few yellow blotches or bands. Coloration (in preservative). Ground colour purplishblack to brownish-black with whitish-cream pattern of irregularly arranged blotches. Comparisons: Comparisons of the new species with its congeners took place based on the following references (Inger & Marx, 1965; Grismer et al., 2004, Howard & Gillespie, 2007, Ziegler et al., 2008; Koch et al., 2009; Nguyen et al., 2009; Orlov, 2009; Orlov et al., 2010). In the following we first compare Calamaria dominici sp. nov. with the Calamaria species reported to occur in Vietnam: Calamaria dominici sp. nov. already differs by its color pattern from the species known from Vietnam: C. abramovi has a black dorsum without spots, the venter is covered with yellow-orange spots; C. buchi is blackish above with each dorsal scale having small light spots and its ventral scales having dark outermost corners; C. concolor has a uniform, patternless light brown body dorsum and a cream venter; C. gialaiensis has a light greyish brown dorsum with an indistinct dark neck collar and few dark blotches along posterior vertebral region, two pairs of light blotches on the tail, as well as a yellowish beige ventral side, with dark outermost corners of ventrals and anterior subcaudals; C. pavimentata usually has narrow, dark, longitudinal stripes, and a solid black collar behind the neck; C. sangi has a greyish brown dorsum with fine dark mottling, as well as a yellowish beige ventral side, with dark transverse bands and a dark longitudinal stripe below the tail; C. septentrionalis has dorsal scales with many small light dots forming a network; C. lovii ingermarxorum has an immaculate grey-bluish dorsum with light spots on each side of the neck covering four scales; and C. thanhi has distinct transverse light body bands. Calamaria dominici sp. nov. further differs from the species so far known from Vietnam in morphological characters: Calamaria dominici sp. nov. differs from C. abramovi Orlov, 2009 by its rostral being wider than high (versus its height equal to width), by the dorsal scales reducing to four rows above 13th subcaudal on tail (versus above 20th subcaudal in the female holotype of C. abramovi), by 18/17 versus 20 subcaudals in females, and by having 13-13-13 versus 12-13-13 dorsal scale rows. Calamaria dominici sp. nov. differs from C. buchi Marx & Inger, 1955 by having fewer ventral scales in females (1 + 174 versus 221-236 in C. buchi), by the rostral being wider than high (versus rostral higher than wide), by the dorsal scales reducing to four rows above 13th subcaudal on tail (versus above 3rd-4th subcaudal), by 18/17 versus 13-14 subcaudals in females, by having a ratio of tail length to total length of 6.2% (versus 3.9-4.1 in female C. buchi), and by the length of parietal being shorter than distance from posterior tip of frontal to posterior tip of rostral (versus length of parietal greater than distance from posterior tip of frontal to posterior tip of rostral). Calamaria dominici sp. nov. differs from C. concolor Orlov, Nguyen, Nguyen, Ananjeva & Ho, 2010 by having paraparietal surrounded by six shields and scales (versus by five shields and scales), and by the dorsal scales reducing to four rows above 13th subcaudal on tail (versus above 19th subcaudal in the male holotype of C. concolor). Calamaria dominici sp. nov. differs from C. gialaiensis Ziegler, Nguyen & Nguyen, 2009 by having paraparietal surrounded by six shields and scales (versus by five shields and scales), and by tail ending in obtuse point (versus with rounded end). Calamaria dominici sp. nov. differs from C. lovii Boulenger, 1887 by having a preocular scale (versus being absent in C. lovii), and by tail ending in obtuse point (versus with blunt end in the subspecies occurring in Vietnam, C. lovii ingermarxorum). Calamaria dominici sp. nov. differs from C. pavimentata Dum��ril, Bibron & Dum��ril, 1854 by the rostral being wider than high (versus rostral as broad as high or slightly higher than broad), and by the tail ending in obtuse point (tail tip with sharp point in C. pavimentata). Calamaria dominici sp. nov. differs from C. sangi Nguyen, Koch & Ziegler, 2009 by having fewer ventral scales (1 + 174 versus 2 + 190 in C. sangi), and by the dorsal scales reducing to six rows above 4th subcaudal on tail (versus above 8th subcaudal). Calamaria dominici sp. nov. differs from C. septentrionalis Boulenger 1890 by 18/17 versus 6-11 subcaudals in females, by having a ratio of tail length to total length of 6.2% (versus 2.6-4.3 in female C. septentrionalis), and by the tail ending in obtuse point (versus tail tip rounded in C. septentrionalis). Calamaria dominici sp. nov. differs from C. thanhi Ziegler & Le, 2005 by having a preocular scale (versus absent), by having fewer ventral scales (1 + 174 versus 198 in C. thanhi), and by the tail ending in obtuse point (tail tip with sharp point in C. thanhi). Calamaria dominici sp. nov. differs from C. yunnanensis Chernov, 1962, a species reported from southern China, which was judged as a doubtful form by Inger & Marx (1965), but subsequently listed as valid by Yang & Inger (1986) and Zhao & Adler (1993), by having a preocular (versus being absent), and by lacking narrow, dark, elongated stripes along the body. Calamaria dominici sp. nov. has a distinct preocular scale, which is lacking in the following species: C. alidae Boulenger, 1920, C. apraeocularis Smith, 1927, C. banggaiensis Koch, Arida, McGuire, Iskandar & B��hme, 2009, C. butonensis Howard & Gillespie, 2007, C. ceramensis De Rooij, 1913, C. gracillima (G��nther, 1872), C. javanica Boulenger, 1891, C. longirostris Howard & Gillespie, 2007, C. mecheli Schenkel, 1901, C. rebentischi Bleeker, 1860, and C. schmidti Marx & Inger, 1955. Calamaria dominici sp. nov. has modified maxillary teeth and thus differs from the following species, which have unmodified maxillary teeth: C. acutirostris Boulenger, 1896, C. curta Boulenger, 1896, C. lautensis De Rooij, 1917, C. leucogaster Bleeker, 1860, and C. ulmeri Sackett, 1940. Calamaria dominici sp. nov. has paraparietal surrounded by six shields and scales and thus differs from C. albiventer (Gray, 1835) (5), C. bicolor Dum��ril, Bibron & Dum��ril, 1854 (5), C. bitorques Peters, 1872 (5), C. brongersmai Inger & Marx, 1965 (5), C. everetti Boulenger, 1893 (5), C. griswoldi Loveridge, 1938 (5), C. hilleniusi Inger & Marx, 1965 (5), C. joloensis Taylor, 1922 (5), C. lateralis Mocquard, 1890 (5), C. lumbricoidea Boie, 1827 (4 or 5), C. lumholtzi Andersson, 1923 (5), C. muelleri Boulenger, 1896 (5), C. palavanensis Inger & Marx, 1965 (5), C. prakkei Lidth de Jeude, 1893 (5), and C. suluensis Taylor, 1922 (5). Calamaria dominici sp. nov. differs from the following species by a distinctly higher ventral scale count in the female sex: C. abstrusa Inger & Marx, 1965 (145-152), C. crassa Lidth de Jeude, 1922 (158-164), C. eiselti Inger & Marx, 1965 (151-153), C. linnaei Boie, 1827 (148- 166), and C. melanota Jan, 1862 (131-154). Calamaria dominici sp. nov. differs from the following species by a distinctly higher subcaudal scale count in the female sex: C. margaritophora Bleeker, 1860 (8- 11), C. nuchalis Boulenger, 1896 (9), and C. sumatrana Edeling, 1870 (10-14). Calamaria dominici sp. nov. differs from C. grabowskyi Fischer, 1885 by a distinctly lower subcaudal scale count in the female sex (20-28). Calamaria dominici sp. nov. has the first three infralabials touching the anterior chin shields versus only two pairs of infralabials touching anterior chin shields in C. borneensis Bleeker, 1860. In addition, Calamaria dominici sp. nov. differs from the remaining species at least by a distinct colour pattern: C. battersbyi Inger & Marx, 1965 (with narrow longitudinal stripes mid-dorsally), C. oesemani Inger & Marx, 1965 (with a continuous light stripe the entire length of the body), C. doederleini Gough, 1902 (with narrow dark brown crossbands on body and tail), C. forcarti Inger & Marx, 1965 (with narrow dark crossbands behind head, body without stripes, venter yellow), C. gervaisii Dum��ril, Bibron & Dum��ril, 1854 (usually with a dark-edged, interrupted, light stripe on first body scale row), C. ingeri Grismer, Kaiser & Yaakob, 2004 (with incomplete light transverse bands on body and tail), C. modesta Dum��ril, Bibron & Dum��ril, 1854 (with ventrals having dark pigment at least laterally), C. schlegeli Dum��ril, Bibron & Dum��ril, 1854 (dark above, light below, head yellow above and below or black above and yellow below, or intermediate conditions), C. virgulata Boie, 1827 (dorsally dark brown, each scale with a light network, with or without longitudinal dark stripes). Etymology: Named dominici to honor Dominic T. Charles Scriven, founder of Wildlife at Risk (WAR), for his contribution towards wildlife conservation in Vietnam. Suggested common names: Dominic���s reed snake (English), Ran mai gam do-mi-nic (Vietnamese), Calamaire de Dominic (French), and Dominics Zwergschlange (German). Distribution: Calamaria dominici is currently known only from the type locality (Fig. 5). Natural history: The holotype was found in evergreen mixed forest of broadleaf and conifer trees. The snake was discovered on a forest path near a small creek, for about 50 m distance from a large creek (Figs 6-7). It was found, surface active, in a densely vegetated boggy area at 11:30. The surrounding habitat was primary forest consisting of dense understory punctuated with large boulders scattered over a ca. 20�� slope that descended to a large creek. Dissection of the female holotype revealed ovaries with some eggs enlarge to 4 mm., Published as part of Ziegler, Thomas, Tran, Vu A., Babb, Randall D., Jones, Thomas R., Moler, Paul E., Van Devender, Robert W. & Nguyen, Truong Q., 2019, A new species of reed snake, Calamaria Boie, 1827 from the Central Highlands of Vietnam (Squamata: Colubridae), pp. 17-26 in Revue suisse de Zoologie 126 (1) on pages 18-23, DOI: 10.5281/zenodo.2619512, {"references":["Inger R. F., Marx H. 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana: Zoology 49: 1 - 304.","Grismer L. L., Kaiser H., Yaakob N. S. 2004. A new species of reed snake of the genus Calamaria H. Boie, 1827, from Pulau Tioman, Pahang, West Malaysia. Hamadryad 28 (1 & 2): 1 - 6.","Howard S. D., Gillespie G. R. 2007. Two New Calamaria (Serpentes) Species from Sulawesi, Indonesia. Journal of Herpetology 41 (2): 237 - 242.","Ziegler T., Nguyen S. V., Nguyen T. Q. 2008. A new reed snake of the genus Calamaria Boie (Squamata: Colubridae) from Vietnam. Current Herpetology 27 (2): 71 - 80.","Koch A., Arida E., McGuire J. A., Iskandar D. T., Bohme W. 2009. A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia. Zootaxa 2196: 19 - 30","Nguyen T. Q., Koch A., Ziegler T. 2009. A new species of reed snake, Calamaria Boie, 1827 (Squamata: Colubridae), from central Vietnam. Hamadryad 34 (1): 1 - 8.","Orlov N. L. 2009. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from the central highlands (Ngoc Linh Nature Reserve, Ngoc Linh Mountain, Kon Tum Province), Vietnam. Russian Journal of Herpetology 16 (2): 146 - 154.","Orlov N. L., Nguyen T. Q., Nguyen T. T., Ananjeva N. B., Ho C. T. 2010. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from Thua Thien-Hue Province, Vietnam. Russian Journal of Herpetology 17 (3): 236 - 242.","Ziegler T., Le Q. K. 2005. A new species of reed snake, Calamaria (Squamata: Colubridae), from the Central Truong Son (Annamite mountain range), Vietnam. Zootaxa 1042: 27 - 38.","Yang D., Inger R. F. 1986. Key to the snakes and lizards of China. Smithsonian Herpetological Information Service 71: 1 - 21.","Zhao E., Adler K. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles, Contribution to Herpetology 10: 1 - 522."]}

Ziegler, Thomas, Tran, Vu A., Babb, Randall D., Jones, Thomas R., Moler, Paul E., Van Devender, Robert W., Nguyen, Truong Q. (2019): A new species of reed snake, Calamaria Boie, 1827 from the Central Highlands of Vietnam (Squamata: Colubridae). Revue suisse de Zoologie 126 (1): 17-26, DOI: 10.5281/zenodo.2619512, {"references":["Chen J.M., Poyarkov N.A., Suwannapoom C., Lathrop A., Wu Y.H., Zhou W.W., Yuan Z.Y., Jin J.Q., Chen H.M., Liu H.Q., Nguyen T.Q., Nguyen S.N., Duong T.V., Eto K., Nishikawa K., Matsui M., Orlov N.L., Stuart B.L., Brown R.M., Rowley J.J.L., Murphy R.W., Wang Y.Y, Che J. 2018. Large-scale phylogenetic analyses provide insights into unrecognized diversity and historical biogeography of Asian leaf-litter frogs, genus Leptolalax (Anura: Megophryidae). Molecular Phylogenetics and Evolution 124: 162-171.","Do T.V., Luu T.H., Wanke S., Neinhuis C. 2015. Three New Species and Three New Records of Aristolochia Subgenus Siphisia from Vietnam including a Key to the Asian Species. Systematic Botany 40(3): 671-691.","Dowling H.G. 1951. A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1(1): 97-99.","Grismer L.L., Kaiser H., Yaakob N.S. 2004. A new species of reed snake of the genus Calamaria H. Boie, 1827, from Pulau Tioman, Pahang, West Malaysia. Hamadryad 28 (1&2): 1-6.","Howard S.D., Gillespie G.R. 2007. Two New Calamaria (Serpentes) Species from Sulawesi, Indonesia. Journal of Herpetology 41(2): 237-242.","Inger R.F., Marx H. 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana: Zoology 49: 1-304.","Koch A., Arida E., McGuire J.A., Iskandar D.T., Bohme W. 2009. A new species of Calamaria (Squamata: Colubridae) similar to C. ceramensis de Rooij, 1913, from the Banggai Islands, east of Sulawesi, Indonesia. Zootaxa 2196: 19-30","Nguyen T.Q., Koch A., Ziegler T. 2009. A new species of reed snake, Calamaria Boie, 1827 (Squamata: Colubridae), from central Vietnam. Hamadryad 34(1): 1-8.","Orlov N. L. 2009. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from the central highlands (Ngoc Linh Nature Reserve, Ngoc Linh Mountain, Kon Tum Province), Vietnam. Russian Journal of Herpetology 16(2): 146-154.","Orlov N.L., Nguyen T.Q., Nguyen T.T., Ananjeva N.B., Ho C.T. 2010. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from Thua Thien-Hue Province, Vietnam. Russian Journal of Herpetology 17(3): 236-242.","Rowley J.J.L., Tran D.T A., Le D.T.T., Dau V.Q., Peloso P.L.V., Nguyen T.Q., Hoang H.D., Nguyen T.T., Ziegler T. 2016. Five new, microendemic Asian Leaf-litter frogs (Leptolalax) from the southern Annamite mountains, Vietnam. Zootaxa 4085(1): 63-102.","Simmons J.E. 2002. Herpetological collecting and collections management. Revised edition. Society for the Study of Amphibians and Reptiles. Herpetological Circular, 31: 1-153.","Uetz P., Freed P., Hosek J. 2018. Eds. The Reptile Database, http://www.reptile-database.org, accessed 20.06.2018.","Wallach V.K., Williams L., Boundy J. 2014. Snakes of the World: A Catalogue of Living and Extinct Species. Taylor and Francis, CRC Press, 1237 pp.","Yang D., Inger R.F. 1986. Key to the snakes and lizards of China. Smithsonian Herpetological Information Service 71: 1-21.","Zhao E., Adler K. 1993. Herpetology of China. Society for the Study of Amphibians and Reptiles, Contribution to Herpetology 10: 1-522.","Ziegler T., Le Q. K. 2005. A new species of reed snake, Calamaria (Squamata: Colubridae), from the Central Truong Son (Annamite mountain range), Vietnam. Zootaxa 1042: 27-38.","Ziegler T., Nguyen S.V., Nguyen T.Q. 2008. A new reed snake of the genus Calamaria Boie (Squamata: Colubridae) from Vietnam. Current Herpetology 27(2): 71-80."]}

Macrochelys apalachicolae sp. nov. Common name. Apalachicola Alligator Snapping Turtle Holotype. UF 3998, partial skeleton from the Apalachicola River, Gadsden County, Florida, on 4 April 1953 by the Florida Museum of Natural History (see Figures 13, 14). (Central lineage; Figure 9). Paratypes. UF 52676, partial skeleton from Waddells Mill Creek, Jackson County, Florida, on 10 April 1978 by L. Richard Franz et al.; UF 152479 skull from Econfina Creek, Bay County, Florida (30.15274 o N, 85.55748 o W, elev. 2 m, 13.1 m depth), on 21 August 1982 by Joseph P. Ward and Joseph J. Ward. Diagnosis. Macrochelys apalachicolae is distinguished by the following: carapacial caudal notch narrow and triangular or narrow and U-shaped (Figure 13), relatively shallow, and reduced; posterior projection of the squamosal globular and obtusely angled in lateral aspect (Figure 5,14); pygal with two serrations, with medial suture; peripheral 11 with one serration; distal rib end of costal 1 enters posterior third of peripheral 3; pleural scute set 1 with slight to no overlap onto the nuchal; processus trochlearis oticum relatively straight with a single distal protuberance; posterior margin of squamosal-opisthotic contact relatively straight in dorsal aspect. Comments. Although there is a general pattern of small triangular pygal regions of the carapace, there is observable variation within the species. All cranial specimens are characterized by large, globular squamosal projections that are intermediate between those of M. suwannensis and M. temminckii. Although M. apalachicolae is genetically most similar to M. temminckii, in some ways it is morphologically more similar to M. suwanniensis; they share the unique synapomorphy of a sutured pygal. Macrochelys apalachicolae is somewhat morphologically intermediate between M. temminckii and M. suwannensis with regard to carapacial caudal notch proportions. The degree of overlap of pleural 1 onto the nuchal also suggests this (usually lying on or just anterior to the nuchalcostal 1 suture), as does a pygal that possesses two serrations (a western character) that is typically sutured medially (a character found in M. suwannensis). Distribution. Restricted to river drainages bounded by the Choctawhatchee and Ochlockonee rivers in Florida, Georgia, and Alabama. Etymology. Specific epithet refers to the new Latin apalachicol��� (referring to the Apalachicola River) and the Latin ���ae (treating the name of the river as a Latin cognate in the First Declension, genitive case), combined to form the composite noun apalachicolae. Fossil record. The earliest fossil representatives of Macrochelys in Florida are from the early Miocene, early Barstovian NALMA, ca. 15���16 Ma. These fossils are fragmentary and consist of a partial costal 8 (UF-Vertebrate Paleontology [VP] 259076) and partial hyo- and hypoplastron (UF-VP 259077). Although difficult to ascribe to the species level, they are contemporaneous with Macrochelys stricta (Matthew 1924) from the early Barstovian of Nebraska. Additional Macrochelys specimens are not observed in Florida until the late Miocene, early Hemphillian NALMA, ca. 8���9 Ma, with the occurrence of Macrochelys auffenbergi (Dobie 1968), which is represented by fairly complete material from the McGehee Farm locality in Alachua County. We reexamined these type specimens, as well as previously undescribed specimens of M. auffenbergi, to diagnose the species based on shell and skull characters and to distinguish it from extant Macrochelys. In M. auffenbergi, the nuchal and cervical are relatively narrow; pleural 1 does not contact the nuchal (shared with western M. temminckii and Chelydra); pygal is much longer than wide, with two serrations and a very narrow caudal notch; pygal lacks medial suturing and is keeled along vertebral 5; epiplastra are relatively wide and lobate (these are long and slender in extant Macrochelys); and sulcal impressions for scutes on the plastron are distinct and deeply incised (these scales are very thin and their impressions faint to lacking in extant Macrochelys). In extant Macrochelys, the nuchal and cervical are wide; Pleural 1 does (in both M. apalachicolae and M. suwanniensis) and does not (in M. temminckii) contact the nuchal; the pygal is much wider than long (in both M. apalachicolae and M. suwanniensis); the epiplastra are very narrow; and the plastron lacks well-defined scute sulcal impressions. The skull of M. auffenbergi, although relatively large, does not exhibit the extraordinary megacephaly expressed in extant Macrochelys. The relative head size is much smaller in M. auffenbergi than in modern Macrochelys, and in that way the fossil taxon is plesiomorphic. The triturating surfaces of the skull and mandible are slender and not as expanded as seen in extant Macrochelys, perhaps an indication that the fossil taxon was less durophagous (eating fewer hard-shelled organisms) than the extant Macrochelys. Increase in head size through time appears to correlate with a decrease in plastral forelobe width within Macrochelys. ,locality collection associated their summarizing specimens fossil Macrochelys unpublished previously and published. timeline of characteristics Paleontological identifying and, 15. age FIGURE geological Macrochelys next appears in Florida from Polk County, from the Widden Creek and Palmetto faunas of the Bone Valley Formation, late Hemphillian, latest Miocene-earliest Pliocene (Meylan 1995). These fossils are fragmentary and occur well south of the current range of Macrochelys. The isolated fossil elements are difficult to identify at the species level; however, the Bone Valley taxon appears larger than M. auffenbergi and more comparable in size to extant specimens. A few features indicate that the fossil taxon is somewhat intermediate in morphology between M. temminckii and M. auffenbergi. These features include having a pygal that is only slightly longer than wide or almost as wide as long (in M. auffenbergi, the pygal is much longer than wide, and in the extant clade, it is much wider than long), and relatively long slender dentaries without overlying expanded triturating surfaces (also seen in M. temminckii and M. auffenbergi). Macrochelys fossils are relatively common in late Blancan to Recent (from ca. 2.5 Ma) fluvial and estuarine deposits in Florida. Most of these fossils are fragmentary. Records include: 1) Late Blancan (ca. 2.5 Ma) US 19 bridge site from the Suwannee River, Gilchrist County. A pygal (UF-VP 247166) from this locality is wider than long with two serrations and unsutured medially. The dentaries (UF-VP 247163���247165) are generally slender as in M. auffenbergi and M. temminckii. 2) Late Blancan (ca. 2.5 Ma) Haile 15 A locality, Alachua County. A nuchal (UF-VP 259613) possesses no Pleural 1 sulci dorsally, the same condition as in M. auffenbergi and M. temminckii. 3) Latest Blancan (ca. 2 Ma) De Soto Shell Pit locality (pits 1 and 3 A), De Soto County. Records from this locality are farther south than the current range of the genus. A pygal (UF-VP 240915) from De Soto 3 A is wider than long with two serrations and unsutured medially. 4) Early Irvingtonian (ca. 1.6 -1.0 Ma) (Morgan & Hulbert 1995; Meylan 1995) Leisey Shell Pits (sites 1, 1B, and 3 B), Hillsborough County. Both Macrochelys and Chelydra occur at this locality, and we feel that there is some confusion with regard to Meylan���s (1995) Macrochelys vouchers. Some of the vouchers represent other taxa (e.g., UF-VP 84005 from pit 1 A is half of an emydid bridge peripheral), including a giant Chelydra species (e.g., a partial peripheral UF-VP 81198 from pit 1 A and a partial shell UF-VP 125099 from pit 2). Meylan (1995: 285) regarded some of his chelydrid identifications as tentative, being aware of an unnamed contemporaneous giant Chelydra (see below for discussion of Chelydra species). Also, the Macrochelys left dentary (UF-VP 116093) reported from pit 3 A is actually from pit 3 B. The Macrochelys fossils occur south of the extant range and are significant in that the dentaries are very robust with expansive triturating surfaces like those in modern M. suwanniensis. 5) Latest Irvingtonian to earliest Rancholabrean (ca. 0.3 Ma) Oldsmar locality, Pinellas County (Meylan 1995). UF-VP 135629 represents a partial posterior carapace with the pygal region well preserved. As with the Leisey specimens, this specimen most closely resembles M. apalachicolae and M. suwanniensis. The pygal is much wider than long, is sutured medially, and possesses two serrations, the condition most typically seen in the central assemblage. 6) Numerous late Pleistocene Rancholabrean NALMA records exist from Florida, including: Jug Springs, Ichetucknee River, Columbia County (Auffenberg 1957); Suwannee River sites; Hornsby Springs, Santa Fe River, Alachua County; and Aucilla River 1 A, Taylor County; as well as extralimital occurences from Wekiva Spring, Levy County; Rock Springs, Orange County; Oklawaha 1, Oklawaha River, Marion County; and Buzzard Island, Putnam County (Meylan 1995). Most of the Rancholabrean fossils are difficult to assess at the species level due to their incompleteness. Specimens from the Ichetucknee River (Suwannee River drainage), including some previously discussed by Auffenberg (1957), consist of nearly complete shells and skulls (e.g., UF-VP 259848, UF- VP 259849, and UF-VP 259842). However, the squamosal and pygal regions are not preserved in these fossils. In UF-VP 259848 and UF-VP 259849, peripheral 11 possesses only one serration, indicating the pygal was also serrated., Published as part of Thomas, Travis M., Granatosky, Michael C., Bourque, Jason R., Krysko, Kenneth L., Moler, Paul E., Gamble, Tony, Suarez, Eric, Leone, Erin, Enge, Kevin M. & Roman, Joe, 2014, Taxonomic assessment of Alligator Snapping Turtles (Chelydridae: Macrochelys), with the description of two new species from the southeastern United States, pp. 141-165 in Zootaxa 3786 (2) on pages 151-160, DOI: 10.11646/zootaxa.3786.2.4, http://zenodo.org/record/253006, {"references":["Matthews, W. D. (1924) Third contribution to the Snake Creek fauna. Bulletin of American Natural Hist ory, 50, 59 - 210.","Dobie, J. L. (1968) A new turtle species of the genus Macrochelys (Chelydridae) from the Florida Pliocene. Tulane Studies in Zoology, 15, 59 - 63.","Meylan, P. A. (1995) Pleistocene amphibians and reptiles from the Leisey Shell Pit, Hillsborough, County, Florida. Bulletin of the Florida Museum of Natural History, 37, 273 - 297.","Morgan, G. S. & Hulbert Jr., R. C. (1995) Overview of the geology and vertebrate biochronology of the Leisey Shell Pit local fauna, Hillsborough County, Florida. Bulletin of the Florida Museum of Natural History, 37, 1 - 92.","Auffenberg, W. (1957) The status of the turtle Macroclemys floridana Hay. Herpetologica, 13, 123 - 126."]}

Macrochelys temminckii (Troost in Harlan 1835) Common name. Alligator Snapping Turtle Holotype. MNHN-AC A. 4540, ���collected near Memphis��� (skull also figured in Bour 1987). (Western lineage; Figure 9). Amended diagnosis. Carapacial caudal notch narrow and triangular or U-shaped, contained wholly on the pygal and not extending onto peripheral set 11, and pygal with two serrations and without medial suture; Peripheral 11 with 1 serration; pleural scute set 1 does not overlap onto the nuchal; distal rib end of costal 1 enters middle of peripheral 3; posterior projection of the squamosal globular and obtusely angled in lateral aspect, usually upwardly inflected; dermal scale on the frontals reduced in size; processus trochlearis oticum relatively straight with a single distal protuberance; posterior margin of squamosal-opisthotic contact relatively straight in dorsal aspect; mandible relatively narrow with slender triturating surfaces. Although generally the caudal notch is small and triangular, observable variation occurs within the species., Published as part of Thomas, Travis M., Granatosky, Michael C., Bourque, Jason R., Krysko, Kenneth L., Moler, Paul E., Gamble, Tony, Suarez, Eric, Leone, Erin, Enge, Kevin M. & Roman, Joe, 2014, Taxonomic assessment of Alligator Snapping Turtles (Chelydridae: Macrochelys), with the description of two new species from the southeastern United States, pp. 141-165 in Zootaxa 3786 (2) on page 149, DOI: 10.11646/zootaxa.3786.2.4, http://zenodo.org/record/253006, {"references":["Bour, R. (1987) Type-specimen of the alligator snapper, Macroclemys temminckii (Harlan, 1835). Journal of Herpetology, 21, 340 - 343. http: // dx. doi. org / 10.2307 / 1563980"]}

Thomas, Travis M., Granatosky, Michael C., Bourque, Jason R., Krysko, Kenneth L., Moler, Paul E., Gamble, Tony, Suarez, Eric, Leone, Erin, Enge, Kevin M., Roman, Joe (2014): Taxonomic assessment of Alligator Snapping Turtles (Chelydridae: Macrochelys), with the description of two new species from the southeastern United States. Zootaxa 3786 (2): 141-165, DOI: http://dx.doi.org/10.11646/zootaxa.3786.2.4

Macrochelys suwanniensis sp. nov. Common name. Suwannee Alligator Snapping Turtle Holotype. UF 166146, adult male skeleton from Santa Fe River and State Road 235, Alachua County, Florida (29.87872 o N, 82.33619 o W, datum WGS 84, elev. 23 m), found dead, apparently from gunshot wounds, in very low water in 2003 by Jason R. Bourque (see Figures 10, 11, 12). (Suwannee lineage; Figure 9). Paratypes. UF 22267, partial skeleton from Santa Fe River, near Town of Santa Fe, Alachua County, Florida, on 9 April 1962 by George R. Zug; UF 12694, partial skeleton from Fletcher Spring, Lafayette County, Florida (29.84672 o N, 82.89256 o W, elev. 9 m), on 19 November 1961 by B. Sites, D. Desautels, and D. Young. Diagnosis. Macrochelys suwanniensis is distinguished by the following: carapacial caudal notch very wide and lunate (Figure 10), usually comprising the pygal and peripheral set 11 (shared with Chelydra); pygal sutured medially (composed of two bones) often with no serrations; Peripheral 11 with 1���2 serrations; distal rib end of costal 1 enters posterior third of peripheral 3; pleural scute set 1 with broad overlap onto the nuchal; dermal scale on the frontals very wide; processus trochlearis oticum with developed proximal and distal protuberances; squamosal contacts opisthotic anteriorly when viewed in dorsal aspect; mandible broad with expanded triturating surfaces and developed labial rugosity just anterior to the coronoid; posterior projection of the squamosal acutely angled in lateral aspect, dorsally straight or downwardly directed, and posteriorly extensive past the plane of the quadrate (Figure 11). Comments. Most carapaces of Macrochelys suwanniensis exhibited a medially sutured pygal. This feature is significant when considering caudal notch width and is likely at least part of the reason this species possesses the widest caudal notch amongst congeners. The extra suture may allow the caudal notch to expand as the turtle grows larger. This is in contrast to M. temminckii, which possesses a single unsutured pygal bone and consequently the narrowest caudal notch of extant Macrochelys. Peripheral 11 is usually doubly serrated; i.e., the serrations that are typically contained on the pygal bone in the western and central species have migrated onto the 11 th peripheral set in M. suwanniensis. Distribution. Restricted to the Suwannee River drainage in Florida and Georgia. Etymology. Specific epithet refers to combination of the new Latin suwanni��� (referring to the Suwannee River) and the Latin ���ensis (belongs to the) to form the composite noun suwanniensis. Specimens examined. See Appendix., Published as part of Thomas, Travis M., Granatosky, Michael C., Bourque, Jason R., Krysko, Kenneth L., Moler, Paul E., Gamble, Tony, Suarez, Eric, Leone, Erin, Enge, Kevin M. & Roman, Joe, 2014, Taxonomic assessment of Alligator Snapping Turtles (Chelydridae: Macrochelys), with the description of two new species from the southeastern United States, pp. 141-165 in Zootaxa 3786 (2) on pages 150-151, DOI: 10.11646/zootaxa.3786.2.4, http://zenodo.org/record/253006