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Tropidonotus nicobarensis Sclater, 1891 is an enigmatic snake currently referred to the natricine genus Hebius Thompson, 1913. No specimens have been recorded since the original discovery of the name-bearing type in the late 19th century, which was allegedly collected on Kamorta Island in India’s Nicobar Archipelago. Recently, a second observation of this species was reported from Havelock Island (Swarajdweep) in the nearby Andaman Archipelago. However, the snake in question is clearly conspecific with another natricine, Thamnophis saurita (Linnaeus, 1766), native to eastern North America. This discovery prompted us to examine the type specimen of Tropidonotus nicobarensis, which revealed that it too is conspecific with Thamnophis saurita. The provenance of the Tropidonotus nicobarensis type specimen and the Havelock Island snake are discussed, with the latter likely representing an introduced animal.

Metal ions are ubiquitous in biology and aid in facilitating many enzymatic transformations. For instance, high–valent iron–oxido and –hydroxido species are proposed to be key intermediates in oxidative processes catalyzed by non-heme iron proteins. A subclass of these proteins contains bimetallic sites that house either FeFe or FeM cores. To investigate the properties of both mono- and bimetallic cores, the Borovik Lab has developed a symmetric tripodal ligand framework containing phosphinic amido groups ([poat]3–), which can stabilize high-valent metal centers, provide an auxiliary binding-site for a second metal ion, and contains a pendant arm that can provide hydrogen-bond stabilization as well as facilitate proton shuttling. The work presented in this dissertation describes my efforts to access high-valent homo-/heterobimetallic oxido and hydroxido species by a variety of synthetic routes.Chapter 1 provides an overview of bimetallic proteins, and efforts by chemists to synthetically model the structure and function of these active sites. Previous work performed in the Borovik Lab to approach these synthetic challenges is discussed.Chapter 2 explores the oxidation chemistry of the mononuclear complexes FeII– and FeIII–hydroxido complexes in the [poat]3– framework, which serve as important synthons in the stepwise assembly of subsequent bimetallic complexes. While previous members in the Borovik Lab have studied these complexes, the electrochemical properties of these compounds are reassessed in my effort to access high-valent Fe–hydroxido species.Chapters 3 and 4 introduce the synthesis and characterization of discrete heterobimetallic complexes, with an MII–OH–FeIII or MII–O–FeIV core (M = Zn, Cu, Ni). This stepwise assembly approach allows for the formation of dinuclear compounds in an unsymmetric framework. The Fe center is supported by the [poat]3– ligand, while MII is supported by an N-based macrocycle. These species are compared with bimetallic complexes in similar frameworks.Chapters 5 describes the preparation of a series of di-Fe compounds with an Fe–O–Fe or Fe–OH–Fe core. These complexes are housed within the same unsymmetric scaffold, and span across four oxidation levels, from FeIIFeII to FeIIIFeIV. This study illustrates the multifunctional nature of the [poat]3- framework, and provides useful insight into the thermodynamic parameters of proton and electron transfers in bimetallic systems. These findings have significant implications for di-Fe containing enzymes, such as ribonucleotide reductase and soluble methane monooxygenase.Using the synthetic strategies developed in the previous chapters, Chapters 6 and 7 describe the preparation and characterization of a library of FeMn and MnFe complexes to model their biological counterparts. Their structural, electrochemical, magnetic, and reactivity properties are discussed. An FeIII–O–MnIV species was trapped and studied spectroscopically, and its relevance to similar high valent FeMn active sites is biology is discussed.Chapter 8 discusses the observation of a rare high-spin CoIII center in a bimetallic system via different spectroscopic techniques. This is an unusual finding, as most six-coordinated CoIII complexes adopt a low-spin configuration due to significant ligand field stabilization. A series of CoFe and CoGa complexes were prepared to probe the factors for this phenomenon, including the P=O groups in [poat]3- being weak field ligands, and modulation of spin exchange interactions.

Redescription of Prymnomiodon chalceus A subjective junior synonym of Thamnophis saurita sackenii fide Cope (1892). The holotype (ANSP 5286) of Prymnomiodon chalceus is an adult male specimen in good condition (Figure 2), with a large ventrolateral incision on the posterior portion of the body and another incision below the vent. The tail is damaged and broken off. SVL 300 mm, TailL 38 mm (TotalL 338 mm), HeadL 14.0 mm, HeadW 7.2 mm, SnoutL 4.6 mm, EyeD 3.0 mm; TailL/ TotalL 0.112, HeadL/ W 1.95, EyeD/HeadL 0.22, EyeD/SnoutL 0.65. Head large and distinct from neck, longer than wide and wider than deep; eyes large with round pupils, smaller than half the length of the head and shorter than snout; nostrils round and oriented laterally. Rostral barely visible from above; internasals paired and wider than long, small, trapezoidal; prefrontals paired and wider than long, pentagonal, suture dividing the two scales less than half the length of the frontal; muzzle length/frontal length ratio 0.610; frontal large, longer than wide, pentagonal and shield shaped; anterior suture of frontal straight but posterior suture angled, border with supraoculars slightly convex; parietals paired and longer than wide, subhexagonal; length of each parietal longer than frontal. Loreal scale 1/1, moderately sized, slightly longer than wide and bottommost border longest; preoculars 1/1 and vertically elongate; postoculars 3/3, lowest postocular smallest in size; supralabials 8/7, with the 4 th and 5 th scales in contact with the orbit on the left side, and the 3 rd and 4 th scales in contact on the right side; 5 th or 6 th supralabial largest and rectangular shaped; temporal scale formula 1+2/1+2 on either side, with the anterior scale broad and subrectangular; infralabials 9/10, with first 1–4 scales in contact with the anterior chin shields on the left side, 1–5 scales in contact on the right side, and the 4 th scale being the largest. Dorsal scales strongly keeled, in 19–19–17 rows, ventrals 152; cloacal plate undivided; subcaudals paired, 24 on the incomplete tail. After more than 160 years of preservation, the dorsum is bluish gray with a pale vertebral stripe around 1.5–2.0 dorsal scales wide and edged with black, extending from the posterior parietal suture to the end of the tail; on each side a barely visible lateral stripe around 0.5 dorsal scales wide is present on the 3 rd and 4 th scale row, starting at the nape near the border of the last supralabial and ending at the terminus of the remaining tail; small black edges present across each stripe. Dorsal portion of head and nape blue green; snout and supralabial region pale beige; preocular with a distinct white spot bordered anteriorly by black; lower postocular also white, but without distinct dark edges; laterally, along the lower temporals, a small black streak extends posteriorly from the eye to the nape. Ventral surface beige, immaculate, much lighter than the dorsum, lighter than dorsal stripes. The original description by Cope (1860) indicates the specimen was originally more olive dorsally with the scales exhibiting a “metallic luster”, but overall the holotype is identical in general appearance and scale counts. Rossman (1961) also reported identical color characteristics, although he noted the bottommost postoculars had a more distinct bright spot that was not visible during our re-examination. Cope neglected to record the number of maxillary teeth but stated that the teeth were larger anteriorly and decreased in size posteriorly. We counted 31 maxillary teeth, and instead found that the teeth gradually increase in size posteriorly, resembling the typical dentition of Thamnophis saurita., Published as part of Lee, Justin L., Chandramouli, S. R. & Bauer, Aaron M., 2023, Tropidonotus nicobarensis Sclater, 1891 is a junior synonym of Thamnophis saurita (Linnaeus, 1766) (Squamata: Serpentes: Natricinae), pp. 133-148 in Zootaxa 5285 (1) on pages 137-139, DOI: 10.11646/zootaxa.5285.1.5, http://zenodo.org/record/7935864, {"references":["Cope, E. D. (1892) A critical review of the characters and variations of the snakes of North America. Proceedings of the United States National Museum, 14, 589 - 694. https: // doi. org / 10.5479 / si. 00963801.14 - 882.589","Cope, E. D. (1860) Catalogue of the Colubridae in the Museum of the Academy of Natural Sciences of Philadelphia. Part 3. Proceedings of the Academy of Natural Sciences of Philadelphia, 12, 553 - 566.","Rossman, D. A. (1961) Nomenclatural status of the neotropical subspecies of the colubrid snake, Thamnophis sauritus. Notulae Naturae of The Academy of Natural Sciences of Philadelphia, 340, 1 - 2."]}

Redescription of Tropidonotus nicobarensis The holotype (ZSI 8895) of Tropidonotus nicobarensis is an unsexed juvenile specimen in fair condition, with an incomplete tail and partially damaged upper jaws (Figure 1). The right maxilla had earlier been dissected out, the incision leaving a blemish in the skin around the temporal region. SVL 164 mm, TailL 61 mm (225 mm TotalL), HeadL 11.0 mm, HeadW 5.0 mm, SnoutL 2.6 mm, EyeD 2.4 mm; TailL/TotalL 0.407, HeadL/ W 2.20, EyeD/HeadL 0.43, EyeD/SnoutL 0.92. Head large and distinct from the neck, ovoid, longer than wide, wider than head depth; eyes large with round pupils, slightly smaller than half the length of the head and shorter than snout; nostrils round and oriented laterally. Rostral barely visible from above; internasals paired and wider than long, small and subtriangular, narrower anteriorly; prefrontals paired and wider than long, pentagonal, suture dividing the two scales less than half the length of the frontal; muzzle length/frontal length ratio 0.510; frontal large, longer than wide, hexagonal and shield-shaped; anterior and posterior sutures of frontal all angled, border with supraoculars convex; parietals paired, longer than wide and subhexagonal; length of each parietal twice as long as frontal. Loreal scale 1/1 and moderately sized, slightly longer than wide and subtrapezoidal; preoculars 1/1, vertically elongate; postoculars 3/3, lowest postocular smallest; supralabials 8/7, with the 4 th and 5 th scales in contact with the orbit on the left side, and the 3 rd and 4 th scales in contact on the right side; 5 th or 6 th supralabial largest and trapezoidal-shaped; temporal scale formula 1+2/1+2 on either side, with the anterior scale broad and subrectangular; infralabials 10/10, with first 1–5 scales in contact with the anterior chin shields and the 4 th scale being the largest. Dorsal scales strongly keeled, in 19–19–17 rows, ventrals 160 (vs. 162 fide Sclater, 1891a), extending onto flanks; cloacal plate undivided; subcaudals paired, 88 under the incomplete tail (vs. 119 fide Sclater, 1891a; 120 fide Smith 1943). After more than 130 years of preservation, the dorsum is brown overall with a distinct light brown vertebral stripe around 1.0–1.5 dorsal scales wide edged with dark brown, extending from the posterior suture of the parietals to the tail tip; on each flank a small light lateral stripe around 0.5 dorsal scales wide present on the 3 rd to 4 th scale row, starting at the nape near the border of the last supralabial and ending at the start of the tail. The coloration of the head is similar to the dorsum, but the preocular has a small tannish spot bordered by dark brown; supralabial region slightly lighter than temporal region and adjacent upper portions of head. Ventral surface creamy-brown, immaculate, much lighter than dorsum and slightly lighter than dorsal stripes. According to past authors, the dorsal and ventral coloration of the specimen was much more vibrant. Sclater (1891a) described the dorsum as “bluish olive” and the snout and lateral portions of the head “yellowish” with a “dark streak” behind the eye. Smith (1943) noted the dorsum was Greenish olive above, with 3 light, black edged stripes, but erroneously reported the lateral stripes on the second and third dorsal scale rows, whereas our examination confirms their presence on the third and fourth rows. Smith (1943) also denoted the presence of small black spots edged along the lateral stripes, and the presence of two white spots on either side of the parietal suture. Neither characteristic is visible in current photographs of the holotype. Discrepancies in the number of ventral scales are probably indicative of different counting procedures, as Dowling (1951) ’s method was not used during the 19 th and mid 20 th century. Furthermore, the number of subcaudals is lower than past reports, but because the tail appears to be broken, it is possible this could be the result of damage since the specimen was last examined. Contrasting the conditions reported by Sclater (1891a) and agreeing with subsequent revisers, we confirm that the cloacal plate is undivided. The right maxilla was probably dissected out by Annandale (1905) because in-text he mentioned that he removed the bone to examine the dentition. Annandale (1905) reported a maxillary tooth count of 24, whereas Wall (1923) counted 25 teeth. Both authors noted that the teeth are gradually enlarged posteriorly., Published as part of Lee, Justin L., Chandramouli, S. R. & Bauer, Aaron M., 2023, Tropidonotus nicobarensis Sclater, 1891 is a junior synonym of Thamnophis saurita (Linnaeus, 1766) (Squamata: Serpentes: Natricinae), pp. 133-148 in Zootaxa 5285 (1) on pages 135-137, DOI: 10.11646/zootaxa.5285.1.5, http://zenodo.org/record/7935864, {"references":["Sclater, W. L. (1891 a) Notes on a collection of snakes in the Indian Museum, with descriptions of several new species. Journal of the Asiatic Society of Bengal, 60, 230 - 250, pl. 6.","Smith, M. A. (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese Sub-region. Reptilia and Amphibia. Vol. III. Serpentes. The Secretary of State for India, London, xii + 583 pp., folding map.","Grant, C. H. B. (1945) William Lutley Sclater, 1864 - 1944. The Ibis, 87, 115 - 121, pl. 1. https: // doi. org / 10.1111 / j. 1474 - 919 X. 1945. tb 01364. x","Dowling, H. G. (1951) A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1, 97 - 99.","Annandale, N. (1905) Additions to the collections of Oriental snakes in the Indian Museum. Part 2. Specimens from the Andamans and Nicobars. Journal and Proceedings of the Asiatic Society of Bengal, New Series, 1, 173 - 176.","Wall, F. (1923) A hand-list of the snakes of the Indian Empire. Part II. Journal of the Bombay Natural History Society, 29, 598 - 632."]}

Thamnophis saurita (Linnaeus, 1766) Coluber saurita Linnaeus 1766: 385.— Neotype. FMNH 73119, adult female collected from 15 miles NNE of Charleston, Berkeley County, South Carolina, United States by John Quinby on 20 July 1953 [designation by Schmidt & Conant 1956, see discussion under Anonymous 1956 –1957]; a full chresonymy of this name and its other synonyms is too exhaustive to be discussed here, overviews are provided by Rossman 1970 and Ernst & Ernst 2003 Tropidonotus nicobarensis Sclater 1891a: 241–242 & pl. 6, fig. 5 (syn. nov.).— Holotype. ZSI 8895, immature specimen collected from “Camorta in the Nicobars [Kamorta Islands, Nicobar Islands, India] by Frederik Adolph de Röepstorff between 1868–1883; Sclater 1891b: vii & 42 (enforces original spelling as first reviser); Das et al. 1998: 151; Das 2003: 473; Chandramouli 2022: 31 Tropidonotus nicobaricus. — Sclater 1891a: 231 & 250 (incorrect spelling) Tropidonotus nicobariensis (incorrect subsequent spelling).— Boulenger 1893: 192; Annandale 1905: 174–175; Iskandar & Coljin 2001: 96; Vijayakumar 2005: 10 (misattributes spelling to Sclater 1891a); Rao et al. 2013: 81; Palot et al. 2013: 60 Natrix nicobariensis.— Wall 1923: 601–602 Natrix nicobarensis.— Smith 1943: 289; Das 2003: 473 Amphiesma nicobariensis.— Malnate 1960: 50; Das 1994: 31; Das et al. 1998: 151; Wrobel 2004: 23; Rao et al. 2013: 81 (misattributes authorship of name to Sclater 1891a) Xenochrophis nicobarensis.— Biswas & Sanyal 1980: 288 Paranatrix nicobariensis.— Mahendra 1984: 246 Amphiesma nicobarense.— David et al. 1998: 92; Das 2003: 473; Ziegler & Quyet 2006: 49; Vijayakumar & David 2006: 12; Palot et al. 2013: 60; Wallach et al. 2014: 31 Amphiesma nicobariense.— Vijayakumar 2005: 10; Harikrishnan et al. 2010: 395 Hebius nicobariense.— Guo et al. 2014: 438; Mohapatra & Ganesh 2021: 1 Hebius nicobarense.— Aengals et al. 2018: 21 Hebius nicobariensis.— Zhou et al. 2019: 78; Ziegler et al. 2020: 62; Das et al. 2020: 314; David et al. 2021: 54; Hauser et al. 2022: 26; Chandramouli 2022: 31; Li et al. 2022: 276; Uetz et al. 2023, Published as part of Lee, Justin L., Chandramouli, S. R. & Bauer, Aaron M., 2023, Tropidonotus nicobarensis Sclater, 1891 is a junior synonym of Thamnophis saurita (Linnaeus, 1766) (Squamata: Serpentes: Natricinae), pp. 133-148 in Zootaxa 5285 (1) on pages 141-142, DOI: 10.11646/zootaxa.5285.1.5, http://zenodo.org/record/7935864, {"references":["Linnaeus, C. V. (1766) Systema natur per regna tria natur, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio duodecima, reformata. Laurentii Salvii, Holmiae, 532 + 2 (errata) pp. https: // doi. org / 10.5962 / bhl. title. 68927","Schmidt, K. P. & Conant, R. (1956) Request that the International Commission on Zoological Nomenclature should direct that the nominal species Coluber sirtalis Linnaeus, 1758 and Coluber saurita Linnaeus, 1766, should be interpreted by the neotypes here designated by the present authors instead of, as hitherto proposed, by reference to specified previously published figures and descriptions. Bulletin of Zoological Nomenclature, 12, 194 - 199.","Anonymous [International Commission on Zoological Nomenclature] (1956 - 1957) Opinion 385. Determination under the Plenary Powers of the method to be adopted in interpreting the nominal species Coluber sirtalis Linnaeus, 1758, the common Garter Snake of the Eastern United States, and approval of the neotype designated for the foregoing species and of that designated for the nominal species Coluber saurita Linnaeus, 1766, the Eastern Ribbon Snake of North America. Bulletin of Zoological Nomenclature, 12, 191 - 230, pls. 1 - 2.","Rossman, D. A. (1970) Thamnophis sauritus. Catalogue of American Amphibians and Reptiles, 99, 1 - 2.","Ernst, C. H. & Ernst, E. M. (2003) Snakes of the United States and Canada. Smithsonian Books, Washington and London, xi + 668 pp.","Sclater, W. L. (1891 a) Notes on a collection of snakes in the Indian Museum, with descriptions of several new species. Journal of the Asiatic Society of Bengal, 60, 230 - 250, pl. 6.","Sclater, W. L. (1891 b) List of snakes in the Indian Museum. Trustees of the Indian Museum, Calcutta, x + 79 pp. https: // doi. org / 10.5962 / bhl. title. 8746","Das, I., Dattagupta, B. & Gayen, N. C. (1998) History and catalogue of reptile types in the collection of the Zoological Survey of India. Journal of South Asian Natural History, 3, 121 - 172.","Das, I. (2003) Growth of knowledge on the Reptiles of India, with an introduction to systematics, taxonomy and nomenclature. Journal of the Bombay Natural History Society, 100, 446 - 501.","Chandramouli, S. R. (2022) Snake fauna of the Andaman Islands, Bay of Bengal - A review of species richness, taxonomy, distribution, natural history and conservation status. Zootaxa, 5209 (3), 301 - 331. https: // doi. org / 10.11646 / zootaxa. 5209.3.1","Boulenger, G. A. (1893) Catalogue of the snakes in the British Museum (Natural History). Vol. I. Containing the families Typhlopid, Glauconiid, Boid, Ilysiid, Uropeltid, Xenopeltid, and Colubrid Aglyph, part. Trustees of the British Museum (Natural History), London, xiii + 448 pp., 28 pls.","Annandale, N. (1905) Additions to the collections of Oriental snakes in the Indian Museum. Part 2. Specimens from the Andamans and Nicobars. Journal and Proceedings of the Asiatic Society of Bengal, New Series, 1, 173 - 176.","Vijayakumar, S. P. (2005) Status and Distribution of Amphibians and Reptiles of the Nicobar Islands, India. Final Report. Rufford Foundation / Madras Crocodile Bank / Wildlife Institute of India, Uttarakhand, 44 pp.","Rao, D. V., Chandra, K. & Devi, K. (2013) Endemic Fauna of Andaman and Nicobar Islands Bay of Bengal. Zoological Survey of India, Kolkata, 292 pp.","Palot J. (2013) Reptiles. In: Venkataraman, K., Chattopadhyay, A. & Subramanian, K. A. (Eds.), Endemic Animals of India (Vertebrates). Zoological Survey of India, Kolkata, pp. 35 - 65, pls. 5 - 8.","Wall, F. (1923) A hand-list of the snakes of the Indian Empire. Part II. Journal of the Bombay Natural History Society, 29, 598 - 632.","Smith, M. A. (1943) The Fauna of British India, Ceylon and Burma, including the whole of the Indo-Chinese Sub-region. Reptilia and Amphibia. Vol. III. Serpentes. The Secretary of State for India, London, xii + 583 pp., folding map.","Malnate, E. V. (1960) Systematic division and evolution of the colubrid snake genus Natrix, with comments on the subfamily Natricinae. Proceedings of the Academy of Natural Sciences of Philadelphia, 112, 41 - 71.","Das, I. (1994) A check-list of the amphibians and reptiles of Andaman and Nicobar Islands. Journal of the Andaman Science Association, 10, 44 - 49.","Wrobel, M. (2004) Elsevier's Dictionary of Reptiles. 1 st Edition. Elsevier B. V., Amsterdam, ix + 761 pp.","Biswas, S. & Sanyal, D. P. (1980) A report on the Reptilia fauna of Andaman and Nicobars Islands in the collection of Zoological Survey of India. Records of the Zoological Survey of India, 77, 255 - 292. https: // doi. org / 10.26515 / rzsi / v 77 / i 1 - 4 / 1979 / 161856","Mahendra B. C. (1984) Handbook of the snakes of India, Ceylon, Burma, Bangladesh and Pakistan. Annals of Zoology, Agra, 22, i - xvi + 1 - 412.","David, P., Vogel, G. & Pauwels, O. S. G. (1998) Amphiesma optatum (Hu & Djao, 1966) (Serpentes, Colubridae): an addition to the snake fauna of Vietnam, with a list of the species of the genus Amphiesma and a note on its type. Journal of Taiwan Museum, 51 (2), 83 - 92.","Ziegler, T. & Quyet, L. K. (2006) A new natricine snake of the genus Amphiesma (Squamata: Colubridae: Natricinae) from the central Truong Son, Vietnam. Zootaxa, 1225 (1), 39 - 56. https: // doi. org / 10.11646 / zootaxa. 1225.1.4","Vijayakumar, S. P. & David, P. (2006) Taxonomy, natural history, and distribution of the snakes of the Nicobar Islands (India), based on new materials and with an emphasis on endemic species. Russian Journal of Herpetology, 13, 11 - 40. https: // doi. org / 10.30906 / 1026 - 2296 - 2006 - 13 - 1 - 11 - 40","Wallach, V., Williams, K. L. & Boundy, J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, Florida, xxvi + 1190 pp.","Guo, P., Liu, Q., Zhang, L., Li, J. X., Huang, Y. Y. & Pyron, R. A. (2014) A taxonomic revision of the Asian keelback snakes, genus Amphiesma (Serpentes: Colubridae: Natricinae), with description of a new species. Zootaxa, 3873 (4), 425 - 440. https: // doi. org / 10.11646 / zootaxa. 3873.4.5","Mohapatra, P. & Ganesh, S. R. (2021) Hebius nicobariense. The IUCN Red List of Threatened Species. Available from: https: // www. iucnredlist. org / species / 202837 / 2757236 (accessed 25 March 2023)","Aengals, R., Sathish Kumar, V. M., Palot, M. J. & Ganesh, S. R. (2018) A checklist of Reptiles of India. Zoological Survey of India, Kolkata, 35 pp.","Zhou, A. Z., Sun, Z., Qi, S., Lu, Y., Lyu, Z., Wang, Y., Li, P. & Ma, J. (2019) A new species of the genus Hebius (Squamata: Colubridae: Natricinae) from Hunan Province, China. Zootaxa, 4676 (1), 68 - 82. https: // doi. org / 10.11646 / zootaxa. 4674.1.3","Ziegler, T., Luu, V. Q., Nguyen, T. T., Van Ha, N., Ngo, H. T., Le, M. D. & Nguyen, T. Q. (2020) Rediscovery of Andrea's keelback, Hebius andreae (Ziegler & Le, 2006): First country record for Laos and phylogenetic placement. Revue suisse de Zoologie, 126, 61 - 71. https: // doi. org / 10.5281 / zenodo. 2619520","Das, A., Gower, D. J. & Deepak, V. (2020) Lost and found: rediscovery and systematics of the Northeast Indian snake Hebius pealii (Sclater, 1891). Vertebrate Zoology, 70, 305 - 318. https: // doi. org / 10.26049 / VZ 70 - 3 - 2020 - 04","David, P., Vogel, G., Nguyen, T. Q., Orlov, N. L., Pauwels, O. S. G., Teynie, A. & Ziegler, T. (2021) A revision of the dark-bellied, stream-dwelling snakes of the genus Hebius (Reptilia: Squamata: Natricidae) with the description of a new species from China, Vietnam and Thailand. Zootaxa, 4911 (1), 1 - 61. https: // doi. org / 10.11646 / zootaxa. 4911.1.1","Hauser, S., Smits, T. & David, P. (2022) On the distribution of the species of the genus Hebius Thompson, 1913 (Squamata: Natricidae) in northern Thailand, including the description of a new species and a discussion on snake diversity of this region. Zootaxa, 5116 (1), 1 - 39. https: // doi. org / 10.11646 / zootaxa. 5116.1.1","Li, M. - L., Ren, J. - L., Huang, J. - J., Lyu, Z. - T., Qi, S., Jiang, K., Wang, Y. - Y. & Li, J. - T. (2022). On the validity of Hebius sauteri maximus (Malnate, 1962) (Squamata, Natricidae), with the redescription of H. maximus comb. nov. and H. sauteri (Boulenger, 1909). Herpetozoa, 35, 265 - 282. https: // doi. org / 10.3897 / herpetozoa. 35. e 94920","Uetz, P., Freed, P, Aguilar, R., Reyes, F. & Hosek, J. (2023) The Reptile Database. Available from: http: // www. reptile-database. org (accessed 16 January 2023)"]}

Calamaria nebulosa sp. nov. Zoobank Registration: http://www.zoobank.org/ urn:lsid:zoobank.org:act: 13AC3177- FFBD-4C12-9132-A9BA5ED7BB48 Calamaria yunnanensis: Stuart and Heatwole (2008: 98), Quah et al. (2019: 3 in part) Calamaria cf. yunnanensis: Weinell et al. (2021: 3) Holotype. FMNH 258666 (field tag: HKV 64382), an adult female (Figures 3 and 6) collected dead on the road (DOR) on the mountainous road to Udomxai [= Oudomxai] about 25 km from Phongsaly City, Phongsaly District, Phongsaly Province, Laos (21.483333��N, 102.2��E; ~1000 metres above sea level). Collected by Mr Bounthavy Phommachanh on 6 October 1999. Diagnosis A species of Calamaria distinguished from all other members of the genus from mainland Southeast Asia by having the following combination of morphological characters: (1) rostral scale wider than high; (2) portion of rostral scale visible from above approximately half the length of the prefrontal suture; (3) six scales and shields surrounding the paraparietal scale; (4) absence of a preocular scale; (5) four supralabials, with the second and third touching the eye; (6) five infralabials, the first pair separating the mental scale from the anterior chin shields; (7) 179 ventrals and three gular scales; (8) 22 paired subcaudals; (9) dorsal colour pattern in preservative bluish-grey with five indistinct dark brown longitudinal stripes; (10) ventral surface of tail in preservative uniform yellow, immaculate; and (11) absence of a distinct white nuchal collar posterior to the head. A summary of diagnostic characters and comparisons is given below and in Tables 3���4. Description of the holotype Adult female in good condition, with some original layers of scales sloughed away on the head and midbody, small incision at base of tail and at midbody. SVL 326 mm, TailL 28 mm, TotalL 354 mm; HeadL 8.8 mm (7.3 mm measured from the parietal tip); HeadW 5.6 mm; SnL 2.8 mm; SnW 2.1 mm; EyeD 1.3 mm; NarEye 2.0 mm; IOD 3.5 mm. Body height at midbody 5.6 mm, at tail base 4.6 mm; eye-���mouth distance 1.1 mm; rostral scale width 2.0 mm, height 1.6 mm; prefrontal scale length 2.3 mm, suture 1.8 mm; frontal scale length 2.7 mm, width 1.9 mm; parietal scale length 4.0 mm, suture 2.5 mm. TailL/TotalL 0.079; HeadL/W 1.58; HeadL/SVL 2.25; SnL/HeadL 0.32; EyeD/SnL 0.46; NarEye/SnL 0.71; SnW/IOD 0.60; IOD/HW 0.63. Body elongate and vermiform, ellipsoid in cross section; tail thick, equal in diameter to rest of body anteriorly and medially; posterior portion of tail gradually tapering to the tip, which appears obtusely pointed in dorsal view, rounder in lateral view; head ovate in dorsal view, indistinct from neck; snout blunt, somewhat depressed posteriorly; nostrils small and elliptical; eyes small, round, diameter slightly larger than eye���mouth distance; pupil indistinct, round; rostral scale subtriangular in frontal view, wider than high, portion visible from above approximately 1/2 the length of the prefrontal suture, suture bordering prefrontal and rostral scale shallow-���V���-shaped; prefrontal scales subhexagonal, shorter than frontal; frontal scale shield-shaped, subhexagonal, 1.4�� longer than wide, around 1.1�� longer than parietal suture; posterior angle formed by the frontal/parietal sutures producing the posterior vertex of the frontal right (~90��); parietals subhexagonal, longer than wide, scale length longer than frontal; parietal suture 1.4�� longer than prefrontal suture; anterior parietal angle formed by the sutures between the parietal/frontal and the suture between the supraocular/parietal broad obtuse, pointing laterally (~130��); supraoculars subrectangular, width of frontal around 2.4�� maximum width of supraoculars; paraparietal shaped like a temporal scale with 6/6 scales and shields in contact; postocular rectangular, not as high as eye diameter; preocular absent; nasal scale small, barely surrounding nostrils; supralabials 4/4, 1st and 2nd smallest, 1st in contact with nasal, 1st and 2nd in contact with prefrontal, 2nd and 3rd in contact with eye, 4th largest and rectangular; infralabials 5/5, 1st smallest and blocking mental from contacting upper genials, 1st���3rd in contact with upper genials; 3rd���4th in contact with lower genials, 5th largest; mental triangular, unremarkable; upper genial scales paired and rectangular, completely in contact; lower genials in contact anteriorly and separated at midsection by the first gular scale, posterior portion truncate-tipped; three gular scales before first ventral scale, all rhomboid. (Continued) Table 3. (Continued). (Continued) Table 4. (Continued). Morphological character C. nebulosa sp. nov. C. abramovi C. albiventer C. andersoni C. buchi C. concolorC. dominici C. gialaiensis C. ingeri C. lovii gimletii Morphological character C. strigiventris C. lovii ingermarxorum C. lumbricoidea C. thanhi C. pavimentata C. yunnanensis C. sangi C. schlegeli schlegeli C. septentrionalisPortion of rostral scale visible from above greater than half with length of prefrontal suture (1) or less than or equal to to half its length (0)101100110Number of scales surrounding paraparietal64���55���65���65���6666���76Prefrontal: longer than frontal scale (1) or smaller than/equal to frontal scale (0)100000000Preocular: present (1) or absent (0)01110���11100Mental touching (1) or separated from (0) anterior genials110100000Posterior genials meeting in midline (1), diverging or meeting only in anteriorly (0)11100���11000End of tailBluntSharp pointSharp pointObtuse pointBluntRoundAbruptly to pointGradually to pointObtuse pointDorsal scales reducing to four rows above position of subcaudal anterior to tail tip4th���5th subcaudalLast���11th subcaudalLast���13th subcaudalLast���3rd subcaudal3rd���25th subcaudalDoes not decreaseLast���6th subcaudalDoes not decrease3rd-to-last subcaudalMaxillary teeth (modified = m)8 (m)9���11 (m)8���9 (m)9 (m)9���10 (m)8���9 (m)9���11 (m)9 (m)8���9 (m)Ventrals205 (M)137���229 (F) 144���196 (M)137���206 (F) 125���168 (M)2 + 190 (?)136���180 (F) 129���161 (M)168���188 (F) 148���166 (M)158���180 (F) 130���157 (M)198 (F) 184 (M)199 (F) 167���184 (M)Subcaudals23 (M)13���21 (F) 17���27 (M)8���20 (F) 13���33 (M)19 (?)19���37 (F) 25���44 (M)6���11 (F) 15���19 (M)20���30 (F) 29���33 (M)21 (F) 28 (M)19 (F) 15���20 (M)DorsumBluish-greyDark brown to blackBrownGreyish brownDark brown to blackDark brown to blackSlate grey to grey-brownDark blue to greyBlue-grey to olive-brownVenterDark greyYellowYellowYellowwhite or yellowYellowYellow and blackYellowYellow or redDorsal scales with light spots present (1) or absent (0)00���10001001Dorsum with light nuchal collar (1) or absent (0)100���1101000���1Tail with light rings: present (1) or absent (0)00���10���100���11010Underside of tail with dark midventral stripe present (1) or absent (0)000���1101100���1TailL/TotalL ratio (given as %)7.4 (M)3.9���8.3 (F) 6.3���11.4 (M)3.7���8.5 (F) 6.9���16.9 (M)6.2 (?)7.3���14.4 (F) 11.1���21.3 (M)2.6���4.3 (F) 6.3���8.6 (M)8.4���11.5 (F) 13.8���17.9 (M)6.8 (F) 9.9 (M)5.0 (F) 5.4���8.2 (M) Dorsal scale rows smooth, in 13���13���13 rows across entire body; dorsal scales reducing to six rows above the 12th subcaudal scale, reducing to four scales above the last subcaudal scale; ventrals 179; subcaudals 22, all paired; cloacal plate single; total body scales 202; subcaudal ratio 0.109; maxillary teeth 9/9, modified. Colouration of the holotype in preservative After formalin fixation and preservation in 70% ethanol for 21 years, the dorsum is bluish-grey, exposed areas of skin where layers of scales have been removed are dark brown; margins of dorsal scales dark grey with faint light yellow vermiculations on scales; there are five indistinct dark brown longitudinal stripes starting at the nape, fading posteriorly towards the tail; these stripes are one dorsal scale row wide, separated from each other by 1���1.5 dorsal scale rows; first three rows of nape pale yellow, immaculate; top of head same colour as dorsum, hued olive-brown on top of head shields; eye blue, pupil grey-blue; bottom half of supralabials and rostral pale yellow; underside of head and rest of venter also pale yellow, margins of ventral scales immaculate with yellow pigment extending onto the first three dorsal scale rows on nape, reducing to two rows across the rest of the body bordered by outermost row of longitudinal stripes; underside of tail the same as rest of venter. Colour in life not recorded. Comparisons Calamaria nebulosa sp. nov. was compared to all other members of the genus inhabiting mainland Southeast Asia. Since C. nebulosa sp. nov. is known only from a single female, morphological data from male Calamaria specimens were excluded from the comparisons. Calamaria nebulosa sp. nov. is morphologically assigned to Calamaria by having an elongate, cylindrical body, absence of a loreal scale, four supralabials, five infralabials and 13 rows of iridescent dorsal scales throughout the body. Additional morphological features that distinguish C. nebulosa sp. nov. from its congeners can be found in Table 4. The absence of a preocular scale in Calamaria nebulosa sp. nov. distinguishes it from all congeners inhabiting mainland Southeast Asia (i.e. Calamaria abramovi Orlov, 2009, Calamaria albiventer (Gray, 1834), Calamaria andersoni Yang and Zheng 2018, Calamaria buchi Marx and Inger, 1955, Calamaria concolor Orlov, Nguyen, Nguyen, Ananjeva and Ho, 2010, Calamaria gialaiensis Ziegler, Sang and Nguyen, 2009, Calamaria ingeri Grismer, Kaiser and Yaakob, 2004, C. lumbricoidea, C. pavimentata, Calamaria sangi Nguyen, Sang and Ziegler, 2009, C. septentrionalis and Calamaria strigiventris Poyarkov, Nguyen, Orlov and Vogel, 2019) except for Calamaria dominici Ziegler, Tran and Nguyen in Ziegler et al., 2019, Calamaria lovii gimletti Boulenger, 1905, C. lovii ingermarxorum Darevsky and Orlov, 1992, C. schlegeli schlegeli, C. thanhi Ziegler and Quyet, 2005 and C. yunnanensis, where the condition of the preocular is either absent or variable. Among these taxa, C. nebulosa sp. nov. is distinguished from C. schlegeli schlegeli (the only species of Calamaria in the region where the preocular can be present or absent) by having the end of the tail tapering to an obtuse point (vs blunt point), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs reducing above the 3rd���25th subcaudal), and by having a bluish-grey dorsum with longitudinal stripes (vs dorsum usually black with a reddish head or a yellow nuchal collar with no light vermiculations on dorsal scales); from C. dominici by having the mental separated from the anterior genials (vs mental in narrow contact with anterior genials), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs reducing above the 5th���6th subcaudal), 179 ventrals and 22 subcaudals in female (vs 174 ventrals and 17���18 subcaudals in females), tail length 7.9% of total length in female (vs 6.0% of total length in females) and a bluish-grey dorsum with longitudinal stripes (vs dark blue dorsum covered with bright yellow spots); from C. lovii gimletti by having the posterior genials meeting only anteriorly (vs meeting at midline), the end of the tail tapering to an obtuse point (vs tapering to a blunt point), 179 ventrals and 22 subcaudals in female (vs 215���249 ventrals and 10���12 subcaudals in females) and a bluish-grey dorsum with longitudinal stripes (vs dorsal colour variable, but usually with a light nuchal collar, small yellow spots on the nape and tail and no light vermiculations on dorsal scales); from C. lovii ingermarxorum by having the portion of the rostral scale visible from above roughly equal to half the length of the prefrontal suture (vs greater than half the length of the prefrontal suture), the length of the prefrontal smaller than the frontal scale (vs longer than frontal), mental separated from the anterior genials (vs in contact with anterior genials), posterior genials meeting only anteriorly (vs meeting at midline), the end of the tail tapering to an obtuse point (vs tapering to a blunt point), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs reducing above the 4th���5th subcaudal), 9 modified maxillary teeth (vs 8), and a bluish-grey dorsum with longitudinal stripes (vs dorsum blue-grey with a light nuchal collar and greyish venter); and from C. thanhi by having the end of tail tapering to an obtuse point (vs tapering gradually to a point), dorsal scales reducing to four rows above the last subcaudal anterior to the tail tip (vs not reducing to four rows), 179 ventrals and 22 subcaudals in female (vs 198 ventrals and 21 subcaudals in female) and by having a bluish-grey dorsum with longitudinal stripes (vs dorsum dark blue or grey with several light body and tail rings). Most specimens of C. yunnanensis can be distinguished from C. nebulosa sp. nov. by having a distinct nuchal colour posterior to the head and a dark brown midventral stripe under the surface of the tail (vs nuchal collar and midventral stripe absent) and by having dark brown margins on each of the ventral scales (vs ventral scales immaculate). However, some specimens of C. yunnanensis do not share these colour pattern traits. In specimens where the colouration is difficult to interpret in preservative or is variable, C. nebulosa sp. nov. can be differentiated from C. yunnanensis by having 179 ventrals in female (vs 199), subcaudal/total body scale ratio of 10.9% in female (vs 8.7%), tail length approximately 7.9% of total length in female (vs 5.0%), dorsal scales reducing to six rows above the 12th subcaudal (vs 3rd to 5th subcaudal) and reducing to four rows above the last subcaudal (vs third-to-last subcaudal). All specimens of C. yunnanensis have narrow dark brown longitudinal stripes present on the margins of each dorsal scale row, occasionally restricted to the flanks in some specimens (vs five, wider dark brown longitudinal stripes, not present on every margin of each dorsal scale row), but these characters can be difficult to interpret on some specimens and should be treated with caution. Distribution and natural history The only known specimen was collected DOR on a mountainous road to the city of Udomaxi, Phongsaly Province, northern Laos (Figure 7). The type locality is approximately 1000 metres above sea level, where the surrounding habitat is dominated by dry evergreen forests and tropical montane deciduous forests (MacKinnon and MacKinnon 1986). The climate surrounding Phongsaly Province is monsoon influenced and humid subtropical (Cwa in the K��ppen climate classification), with a rainy season starting in the spring around May and ending in the fall around October (Peel et al. 2007; Geiser and Nagel 2013). Nothing else is known about the natural history of the new species. It is presumed to resemble other Calamaria in being an oviparous species that spends most of its life underground or below the forest floor, feeding on small soft-bodied invertebrates such as earthworms (Inger and Marx 1965). Conservation status Calamaria nebulosa sp. nov. is known from only a single specimen and has not been recorded since its discovery in 1999. The full extent of its distribution is unknown but the habitat around the type locality is relatively continuous across most of northern Laos, as well as adjacent Vietnam, northern Thailand, eastern Myanmar and southern China. It is likely that C. nebulosa sp. nov. also occurs in these countries, but until additional specimens are found, I recommend that this new species be listed as ��� Data Deficient��� following the International Union for Conservation of Nature (IUCN) Red List categories (IUCN 2019). Etymology The species epithet ��� nebulosa ��� is the nominative form of the word ��� nebulous ���, meaning ���misty, foggy or cloudy��� in Latin, and is given in the feminine form to match the female genitive declension of the genus name Calamaria. The species name is an allusion to both the type locality of this snake, situated in the mountainous regions of Laos, and the generally clouded appearance of its dorsal ground colour in preservative. I recommend the English common name ���Clouded reed snake���., Published as part of Lee, Justin L., 2021, Description of a new species of Southeast Asian reed snake from northern Laos (Squamata: Colubridae: Genus Calamaria F. Boie, 1827) with a revised diagnosis of Calamaria yunnanensis Chernov, 1962, pp. 531-560 in Journal of Natural History 55 (9 - 10) on pages 539-547, DOI: 10.1080/00222933.2021.1909165, http://zenodo.org/record/5476206, {"references":["Stuart BL, Heatwole H. 2008. Country records of snakes from Laos. Hamadryad. 33 (1): 97 - 106.","Quah ESH, Anuar S, Grismer LL, Wood PL Jr, Mohd Nor SA. 2019. Systematics and natural history of mountain reed snakes (genus Macrocalamus; Calamariinae). Zool J Linn Soc. 188 (4): 1236 - 1276. doi: 10.1093 / zoolinnean / zlz 092","Weinell J, Leviton AE, Brown RM. 2021. A new species of reed snake, genus Calamaria (Colubridae: Calamariinae), from Mindoro Island, Philippines. Philipp J Syst Biol. 14 (2): 1 - 14. doi: 10.26757 / pjsb 2020 b 14006","Zhao EM, Huang MH, Zong Y, Zheng J, Huang ZJ, Yang D, Li DJ, Eds. 1998. Fauna Sinica. Reptilia Vol. 3. Squamata Serpentes. Beijing: Science Press; p. i - xvii + 1 - 522, Pls. I - VIII, Col. Pls. I - IV. [in Chinese].","Yang D, Rao DQ. 2008. Amphibians and reptiles of Yunnan. Kunming: Yunnan Science and Technology Press; 409 pp. [in Chinese].","Chernov SA. 1962. On one yet unknown snake of the genus Calamaria from Yunnan (Scientific results of the Chinese-Soviet expeditions in 1955 - 1957 to South-Western China). Proc Zool Inst Russ Acad Sci USSR, Leningrad. 30: 382 - 384. [in Russian].","Orlov NL. 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. Russ J Herpeto. 16 (2): 146 - 154. doi: 10.30906 / 1026 - 2296 - 2009 - 16 - 2 - 146 - 154","Gray J. 1834. Illustrations of Indian zoology; chiefly selected from the collection of Major-General Hardwicke, F. R. S., L. S., M. R. A. S., M. R. I. A., & C. Volume II (parts XI - XX). London: Adolphus Richter and Co & Parbury, Allen, and Co. (3 unnumbered pages), Pls. 1 - 102.","Yang JH, Zheng X. 2018. A new species of the genus Calamaria (Squamata: Colubridae) from Yunnan Province, China. Copeia. 106 (3): 485 - 491. doi: 10.1643 / CH- 17 - 663","Marx H, Inger RF. 1955. Notes on snakes of the genus Calamaria. Fieldiana. 37: 167 - 209.","Orlov NL, Nguyen TQ, Nguyen TT, Ananjeva NB, Ho CT. 2010. A new species of the genus Calamaria (Squamata: Ophidia: Colubridae) from Thua Thien-Hue Province, Vietnam. Russ J Herpeto. 17 (3): 236 - 242. doi: 10.30906 / 1026 - 2296 - 2010 - 17 - 3 - 236 - 242","Nguyen QT, Koch A, Ziegler T. 2009. A new species of reed snake, Calamaria Boie, 1827 (Squamata: Colubridae), from central Vietnam. Hamadryad. 34 (1): 1 - 8. doi: 10.30906 / 1026 - 2296 - 2009 - 16 - 2 - 146 - 154","Grismer LL, Kaiser H, Yaakob NS. 2004. A new species of reed snake of the genus Calamaria H. Boie, 1827, from Pulau Tioman, Pahang, West Malaysia. Hamadryad. 28: 1 - 6.","Poyarkov NA Jr, Van Nguyen T, Orlov NL, Vogel G. 2019. A new species of the genus Calamaria Boie, 1827 from the highlands of the Langbian Plateau, Southern Vietnam (Squamata: Colubridae). Russ J Herpeto. 26 (6): 335 - 348. doi: 10.30906 / 1026 - 2296 - 2019 - 26 - 6 - 335 - 348","Ziegler T, Tran VA, Babb RD, Jones TR, Moler PE, Van Devender RW, Nguyen TQ. 2019. A new species of reed snake, Calamaria Boie, 1827 from the Central Highlands of Vietnam (Squamata: Colubridae). Rev Suisse Zool. 126 (1): 17 - 26. doi: 10.5281 / zenodo. 2619512","Boulenger GA. 1905. Descriptions of new Snakes in the collection of the British Museum. Ann Mag Nat Hist. Ser. 7. 15 (85): 453 - 456.","Darevsky IS, Orlov NL. 1992. A new subspecies of the dwarf snake Calamaria lowi ingermarxi ssp. nov. (Serpentes, Colubridae) from southern Vietnam. Asiatic Herpetol Res. 4: 13 - 17.","Ziegler T, Quyet LK. 2005. A new species of reed snake, Calamaria (Squamata: Colubridae), from the Central Truong Son (Annamite mountain range), Vietnam. Zootaxa. 1042 (1): 27 - 38. doi: 10.11646 / zootaxa. 1042.1.2","MacKinnon J, MacKinnon K. 1986. Review of the protected areas system in the Indo-Malayan realm. Cambridge (UK): International Union for the Conservation of Natural Resources; xii + 284 pp. + [separate] maps 3.1 - 3.6.","Peel MC, Finlayson BL, McMahon TA. 2007. Updated world map of the Koppen-Geiger climate classification. Hydrol Earth Syst Sci. 11 (5): 1633 - 1644. doi: 10.5194 / hess- 11 - 1633 - 2007","Geiser M, Nagel P. 2013. Coleopterology in Laos-an introduction to the nature of the country and its coleopterological exploration. Entomol Basiliensia Collectionis Frey. 34: 11 - 46.","Inger RF, Marx H. 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana Zool. 49: 1 - 304.","IUCN. 2019. IUCN International Union for Conservation of Nature. [accessed 2020 Apr 3]. https: // www. iucnredlist. org /."]}

Calamaria yunnanensis Chernov, 1962 Calamaria yunnanensis: Chernov 1962: 383, in: ���On one yet unknown snake in the genus Calamaria from Yunnan (Scientific results of the Chinese���Soviet expeditions in 1955���1957 to South-Western China)���. Proceedings of the Zoological Institute, Academy of Science of the USSR, Leningrad, 30, 382���384, Inger and Marx (1965: 238), Yang and Inger (1986: 10), Zhao and Adler (1993: 232), Ziegler and Quyet (2005: 36), Zhao (2006: 184), Stuart and Heatwole (2008: 98 in part), Yang and Rao (2008: 275), Ziegler et al. (2008: 78), Orlov (2009: 152), Nguyen et al. (2009: 3), Koch et al. (2009: 20), Zaher et al. (2009: 138), Orlov et al. (2010: 239), Vogel and Luo (2011: 36), Yang and Zheng (2018: 485 in part), Ziegler et al. (2019: 23), Poyarkov et al. (2019: 341 in part), Wallach et al. (2014: 142, in part), Weinell et al. (2021: 3 in part). Holotype. NZMC, original catalogue number ZISP 17073 (Figure 1 (b)), collected ��� in the vicinity of Tszindun, in the gorge of monsoon forest 1100���1200 meters��� [= in the vicinity of Jingdong, Jingdong Yi Autonomous County, southern Yunnan Province, China] a male specimen (maturity unknown) collected on 31 May 1956 by Soviet Researcher Alexei K. Zaguliayev, as part of the joint Soviet���Chinese field expeditions to Yunnan Province, China. A translation of the original description can be found in the Supplementary data. Topotypic specimens. (Figure 5). KIZ 054176 (alternate catalogue: 75II0198), an adult female collected from Jingdong Yi Autonomous County, southern Yunnan Province, China (same locality as the type specimen) on 11 May 1975 (collector unknown); KIZ 056010���011 (alternate catalogues: 75II0224���225, respectively), two males from the same locality collected on 10 June 1975 (collector unknown); and KIZ 056009, sex unknown, from the same locality and date as KIZ 056009���010. Referred specimen. ROM 41547 (Figures 4 and 6), a male specimen (maturity unknown) found 8.8 km west of Simao, Yunnan Province, China (22�� 46 �� 47.928 �� ��� N, 100�� 54 �� 24.0372 �� ��� E; ~1270 metres above sea level) collected by Robert W. Murphy and Nikolai L. Orlov on 14 June 2002. Diagnosis A species of Calamaria distinguished from all other members of the genus found in mainland Southeast Asia by having the following combination of morphological characters: (1) rostral scale wider than high; (2) portion of rostral scale visible from above less than half the length of the prefrontal suture; (3) six scales and shields surrounding the paraparietal scale; (4) absence of a preocular scale; (5) four supralabials, with the second and third touching the eye; (6) five or six infralabials, the first pair separating the mental scale from the anterior chin shields; (7) 167���184 ventrals in males, 199 in female, three gular scales; (8) 15���20 subcaudals in males, 19 in female, all paired; (9) dorsal colour pattern variable, usually bluish-grey or olive-brown with indistinct dark brown longitudinal stripes across each dorsal scale row forming a network-like pattern, sometimes restricted to the flanks; (10) ventral surface red or yellow usually with dark brown margins on the ventral scales and a black midventral stripe under the tail, completely immaculate in some specimens; and (11) a light yellow nuchal collar posterior to the head usually present, absent in some specimens. A summary of diagnostic characters and comparisons is given below and in Tables 3���4. Description of the referred specimen Male (maturity unknown) in good condition, with one incision at the tail base and another along the posterior portion of body. SVL 226 mm, TailL 20 mm, TotalL 246 mm, HeadL 7.4 mm (6.4 mm measured from the parietal tip), HeadW 4.0 mm, SnL 2.6 mm, SnW 2.0 mm, EyeD 1.0 mm, NarEye 1.5 mm, IOD 3.1 mm. Body height at midbody 3.5 mm, at tail base 3.3 mm; eye���mouth distance 0.8 mm; rostral scale width 0.9 mm, height 1.1 mm; prefrontal scale length 1.8 mm, suture 1.0 mm; frontal scale length 2.4 mm, width 1.9 mm; parietal scale length 3.3 mm, suture 1.8 mm. TailL/TotalL 0.081; HeadL/W 1.6; HeadL/SVL 2.83; SnL/HeadL 0.41; EyeD/SnL 0.38; NarEye/SnL 0.58; SnW/IOD 0.66; IOD/HW 0.76. Body elongate and vermiform, round in cross section; tail thick, roughly equal in diameter to rest of body; posterior portion of tail slightly tapering, then abruptly tapering at the tip, which appears obtusely pointed in dorsal view (weakly acute), round in lateral view; head sub-oblong in dorsal view, indistinct from neck; snout blunt, weakly depressed posteriorly; nostrils small and elliptical; eyes small, round, diameter larger than eye��� mouth distance; pupil indistinct, round; rostral scale triangular in frontal view, wider than high, portion visible from above roughly 4/5 the length of the prefrontal suture, suture bordering prefrontal and rostral scale deep-���V���-shaped; prefrontal scales subhexagonal, shorter than frontal; frontal scale shield-shaped, hexagonal, 1.3�� longer than wide, around 1.4�� longer than parietal suture; posterior angle formed by the frontal/ parietal sutures producing the posterior vertex of the frontal right (~90��); parietals subhexagonal, longer than wide, longer than frontal; parietal suture 1.4�� longer than prefrontal suture; anterior parietal angle formed by the sutures between the parietal/frontal and the suture between the supraocular/parietal broadly obtuse, pointing laterally (~127��); supraoculars subrectangular, width of frontal around 2.7�� maximum width of supraoculars; paraparietal shaped like a temporal scale with 6/6 scales and shields in contact; postocular scale rectangular, not as high as eye diameter; preocular absent; nasal scale small, subrectangular, barely surrounding nostrils; supralabials 4/4, 1st and 2nd smallest, 1st in contact with nasal, 1st and 2nd in contact with prefrontal, 2nd and 3rd in contact with eye, 4th largest and rectangular; infralabials 5/5, 1st smallest and blocking mental from contacting upper genials, 1st���3rd in contact with upper genials; 3rd���4th in contact with lower genials, 5th largest; mental triangular, unremarkable; upper genial scales paired and rectangular, completely in contact; lower genials in contact anteriorly and separated at midsection by the first gular scale, posterior portion round-pointed; three gular scales before first ventral scale, all rhomboid. Dorsal scale rows smooth, in 13���13���13 rows across entire body; dorsal scales reducing to six rows above the 3rd subcaudal scale, reducing to four scales above the 16th (third-tolast) subcaudal scale; ventrals 167; subcaudals 19, all paired; cloacal plate single; total body scales 187; subcaudal ratio 0.102; maxillary teeth 8/8, modified. Colouration of the referred specimen in preservative After formalin fixation and preservation in 70% ethanol for 18 years, the dorsum is greybrown and highly iridescent; margins of all dorsal scales dark brown with faint light brown vermiculations on the medial portion of most dorsal scales; dark margins of dorsal scales forming a weak network-like pattern; along flanks, three dark brown longitudinal stripes, lower two stripes 0.5���1.0 dorsal scale rows wide, separated from each other by 0.5 dorsal scale rows; first two stripes on flanks broader (one dorsal scales wide) than highest stripe (0.5 dorsal scale rows wide) along the anterior portion of body; highest stripe two dorsal scales above lower stripes, reduced to one dorsal scale row at midbody; medial portion between stripes filled with light white vermiculations; all longitudinal stripes fade posteriorly, except for one pair that continues onto the posterior portion of tail; dorsal portion of head dark brown with occasional light brown hueing along cephalic scales; nuchal collar whitish-tan, around one dorsal scale in width at vertebrum then widening in lateral view, originating at the third and fourth row of dorsal scales; supralabials and nasal tannish, fourth supralabial mostly white; flanks along nape also white, connecting to nuchal collar; eye blue, pupil greyish-blue; underside of head cream with grey-brown mottling scattered along mental, infralabials, upper genials and nuchal region of first dorsal scales; venter white, margins dark brown; underside of tail with small dark spotting along margins and dark brown midventral stripe starting from tail tip, breaking up near the start of the cloacal plate. Description of the topotypic specimens and variation Zhao et al. (1998) and Yang and Rao (2008) published descriptions of C. yunnanensis based on the same three topotypic specimens (2 males, 1 female). Although they originate from the type locality, the written descriptions of these specimens by these authors have slight differences from Chernov (1962) ���s original description and from ROM 41547. Zhao et al. (1998) indicated that the midventral stripe and nuchal collar in C. yunnanensis may be present or absent, while Yang and Rao (2008) did not note the presence of either feature. Photographs of these specimens from the KIZ collection allow me to clarify this issue (Figure 5; Shuo Liu, pers. comm.) The official museum catalogue numbers for these specimens are different than those given in Yang and Rao (2008), but they can be associated with their specimen numbers based on scale counts and body measurements. In the single female specimen (KIZ 054176), the distinct nuchal collar and midventral stripe on the tail is absent. The dorsum is brown, becoming darker posteriorly near the tail, with indistinct longitudinal stripes along the edges of the dorsal scales forming a ���net-like��� pattern. The venter is immaculate yellow, with the light pigment extending onto the first outer row of dorsal scales. Zhao et al. (1998) and Yang and Rao (2008) also provide conflicting data on the number of ventrals and subcaudals for this specimen, with the former reporting 198 ventrals and 19 subcaudals and the latter reporting 201 ventrals with 21 subcaudals. A re-count of the scales for this specimen indicates that it has 199 ventrals and 19 subcaudals, agreeing more with Zhao et al. (1998). This number is significantly higher than for the male specimens of C. yunnanensis (167���184), but sexual dimorphism in the number of scale counts is common in Calamaria, and most females in the genus have a higher number of ventrals than the males (Inger and Marx 1965). In the other specimens (KIZ 056009���010), both males, the midventral stripe is present on the tail and a nuchal collar is present in one of the specimens (KIZ 056010). The other specimen (KIZ 056009) is partially destroyed with only the midbody and tail remaining, so it is uncertain whether or not it had a nuchal collar. Both specimens also have some dark brown vermiculate-shaped spots present on the underside of the tail in addition to the midventral stripe (KIZ 056009���010). It is unknown whether or not the type specimen contained this characteristic, but a few small dark vermiculations are present on ROM 41547. Otherwise, the colouration of these three specimens agrees with Chernov (1962) ���s original description of C. yunnanensis and with ROM 41547, with the only differing characteristic being the presence of indistinct dark brown longitudinal stripes on the dorsum. Chernov (1962) did not note this feature in his description but stated that the margins of the dorsal scales were often dark brown and ���continuous���, suggesting that there were indistinct stripes as described in these other topotypic specimens. Indeed, the striped pattern is barely visible on some the other topotypic specimens in the KIZ collection due to their state of preservation, and this artefact could have caused Chernov to miss this character when he was writing his description. Another characteristic shared by these specimens is the presence of dark brown margins along the ventral scales, which is shared with ROM 41547. The scale counts for both specimens are identical to those given by Zhao et al. (1998) and Yang and Rao (2008), except KIZ 056009 has 20 subcaudal scales instead of 22 scales as reported by these authors. A fourth specimen (KIZ 056011), unreported in either study, was also found in the collection. It is an immature individual and its sex cannot be accurately determined, but it clearly possesses the same colour pattern characteristics as KIZ 056009���010 and ROM 41547. Overall, the colour polymorphism observed in these topotypic specimens agrees with the description provided by Zhao et al. (1998), while Yang and Rao (2008) ���s omission of these characteristics may have caused confusion in diagnosing the species. Photographs of a live C. yunnanensis shown in Yang and Zheng (2018, fig. 4) and in Figure 1 offer some insight into the live colouration of this species. The specimen agrees with the colour pattern observed in KIZ 054176. However, the dorsum is predominately olive-brown, becoming dark grey-brown posteriorly, with well-defined dark brown longitudinal stripes across the body forming a network-like pattern on the dorsum. The venter in this specimen is a bright coral red, extending onto the lower flanks of the first two dorsal scale rows and on the labial region of the head and nostrils. A streak of red pigment forms a small marking posterior to the parietals but is too indistinct to be considered a nuchal collar, as it is nowhere near as pronounced as the markings present in other specimens. Red colouration may have been present on the ventral surface of other topotypic C. yunnanensis specimens and ROM 41547, but this pigment tends to fade quickly after preservation. Distribution and natural history So far, C . yunnanensis has been collected from three localities in south-central Yunnan Province, China (Figure 7). The type locality, Jingdong Autonomous County, is situated 1100���1200 metres above sea level, while the locality of ROM 41547 was at 1270 metres above sea level. The type specimen was found underneath a rock in a forested montane valley. The photographed individual in Yang and Zheng (2018) was located approximately 60 km north-west of the type locality in Nanjian Yi Autonomous County, Dali, Yunnan Province, China (24��48 �� 09.3��� N, 100��32 �� 58.5��� E; ~1700 metres above sea level). It was found on 27 July 2011 at night between 17:00 and 18:00 UTC, attempting to climb across a rock along the road���s shoulder (Chung-Wei You, pers. comm). The surrounding habitats in these localities include a mix of tropical montane deciduous forests near Simao, and montane evergreen broadleaf forests near Nanjing Yi and Jingdong Autonomous Counties. The climate in these regions is monsoon influenced and considered humid subtropical (Cwa) or subtropical highland (Cwb) according to the K��ppen Classification (Peel et al. 2007). Besides this information, nothing else is known regarding its natural history, but given these observations, it is likely a nocturnal species. Similar to other species of Calamaria, it is presumed to be a fossorial species of snake that spends most of its life underground or below the forest floor, feeding on small soft-bodied invertebrates such as earthworms (Inger and Marx 1965). Conservation status Calamaria yunnanensis has been recorded from two localities in southern Yunnan Province China based on a total of six to seven known specimens. The full extent of this species distribution in Yunnan Province is unknown. I recommend that C. yunnanensis be listed as ��� Data Deficient��� following the IUCN���s Red List categories (IUCN 2019). Etymology The species epithet ��� yunnanensis ��� is a reference to the type locality situated in Yunnan Province, China. I recommend the English common name ��� Yunnan reed snake���., Published as part of Lee, Justin L., 2021, Description of a new species of Southeast Asian reed snake from northern Laos (Squamata: Colubridae: Genus Calamaria F. Boie, 1827) with a revised diagnosis of Calamaria yunnanensis Chernov, 1962, pp. 531-560 in Journal of Natural History 55 (9 - 10) on pages 548-555, DOI: 10.1080/00222933.2021.1909165, http://zenodo.org/record/5476206, {"references":["Chernov SA. 1962. On one yet unknown snake of the genus Calamaria from Yunnan (Scientific results of the Chinese-Soviet expeditions in 1955 - 1957 to South-Western China). Proc Zool Inst Russ Acad Sci USSR, Leningrad. 30: 382 - 384. [in Russian].","Inger RF, Marx H. 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana Zool. 49: 1 - 304.","Yang D, Inger RF. 1986. 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