Homatula guanheensis: A new species of Stone Loach from Henan Province, China

The Stone Loach genus Homatula includes a number of small, benthic Fish, which live in the drainage basins of the Yellow, Yangtze, Pearl, Lancang, Nujiang and Red rivers. They are sometimes considered to be a part of the genus Paracobitis, which is morphologically similar, but the two are generally considered separate, due to their different distributions, with Homatula known only from China, and Paracobitis found only in western Asia. Studies based on the morphology of Homatula have suggested that there is more diversity that the current number of species (19-20), but to what extent this reflects variation within species rather than the presence of undescribed species remained unclear, until scientists began to apply genetic identifications to the genus.

In a paper published in the Biodiversity Data Journal on 16 June 2021, Chuanjiang Zhou, Wenwen Ma, Xi Wang, Yongtao Tang, Xiaoling Meng, and Guoxing Nie of the Engineering Technology Research Center of Henan Province for Aquatic Animal Cultivation at Henan Normal University, describe a new species of Homatula from the Guanhe River, in the HanJiang River drainage (a tributary of the Yangtze River),  in Xixia County, Henan Province.

The new species is named Homatula guanheensis, where 'guanheensis' means 'coming from Guanhe', in reference to the Guanhe River where it was found living in cave environments. Like other Stone Loaches, the new species is an elongate, slender Fish. The known specimens range from 76.9 to 109.26 mm in length, and are cylindrical, and flattened towards the rear. The head is short and flattened, and lacks scales. The snout is short, with nostrils closer to the eye than the tip. The eyes are oval, and located towards the top of the head. The lips are thick, and cover the jaw; there are three pairs of barbels. 

 

Homatula guanheensis (holotype, HNU 010048, 99.6 mm SL). (A) Lateral view; (B) Dorsal view; (C) Ventral view; (D) Mouth characters; (E) Intestine form; (F) X–ray (lateral view). Zhou et al. (2021).

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Notopterus synurus: A cryptic species of Bronze Featherback from South Asia.

The complex geological history of the Oriental region caused a high degree of geographical genetic structure within freshwater organisms and species, once considered widely distributed in this region, are often formed by distinct genetic lineages. The Bronze Featherback Fish Notopterus notopterus, currently the only valid species of the genus Notopterus, is one of such widely-distributed species, occurring from the Indus basin (Pakistan and India) in the west, to the Mekong region – slightly extending east to the Annamite Range – (Cambodia, Laos, Thailand and Vietnam) in the east and to Java (Indonesia) in the south. Intriguingly, Notopterus notopterus has not yet been recorded in Borneo where it is most likely absent. Specimens of Notopterus notopterus are identifiable from all other Oriental Freshwater Fish based on (amongst other features) their distinct tapered tail and the corners of their mouth below eye (not behind as in the other Oriental Notopterid genus Chitala. It is reported that this species reaches up to 60 cm in standard length. A wealth of data is available for this species, documenting its reproductive behaviour and embryonic development, cytogenetics, and phylogeography. However, none of these studies simultaneously examined specimens sampled across the whole distributional range of Notopterus notopterus.

In 2009 a group of scientists led by Jun Inoue of the Ocean Research Institute at University of Tokyo published a paper in the journal Molecular Phylogenetics and Evolution, which reconstructed the molecular phylogeny of the family Notopteridae to discuss its evolution and biogeography. These authors sequenced the complete mitogenomes of two specimens of Notopterus notopterus, one from India and the other from Thailand. The comparison of these two mitogenomic sequences revealed that these two specimens diverged from each other about 25 million years ago. This result was unexpected because no consistent morphological variation was previously reported within this species. 

In a paper published in the journal Zoosystematics and Evolution on 1 July 2020, Sébastien Lavoué of the School of Biological Sciences at the Universiti Sains Malaysia, Siti Zafirah Ghazali, also of the School of Biological Sciences at the Universiti Sains Malaysia, and of the Institute of Marine Biotechnology at the Universiti Malaysia Terengganu, Jamsari Amirul Firdaus Jamaluddin, again of the School of Biological Sciences at the Universiti Sains Malaysia, Siti Azizah Mohd Nor, again of the Institute of Marine Biotechnology at the Universiti Malaysia Terengganu, and Khaironizam Zain, once again of the School of Biological Sciences at the Universiti Sains Malaysia, present the results of a study which investigated the genetic diversity within the genus Notopterus across its full range, for which, they analysed a dataset comprising 72 publicly available sequences of the standard barcoding fragment (655 base pairs) of the cytochrome oxidase I gene that were determined from specimens of Notopterus with precise information on their geographical collection plus two cytochrome oxidase I sequences extracted from two complete mitogenomes from two specimens of Notopterus without precise localities and four sequences Lavoué et al. newly determined from specimens collected in Peninsular Malaysia.

Haplotypes of Notopterus segregate into two main groups having distinct distributions. The first group includes all haplotypes from India and Bangladesh (South Asia group). The second group includes all haplotypes from Peninsular Malaysia, Thailand (Mekong River), Indonesia (Sumatra and Java) and Myanmar (Lake Inle, Salween basin) (Southeast Asia group). These two main groups diverge by 7.5% p-genetic distance that represents, on average, 45 differences between any combination of two specimens sampled from different groups. In contrast, each of these two groups is genetically uniform with intra-group differentiation that does not exceed 1% (within the South Asia group, mean p-distance equals 0.4%; and within the Southeast Asia group, mean p-distance equals 1%). This represents six nucleotide differences on average within the Southeast Asian group and only two nucleotide differences on average within the South Asian group. Furthermore, the two specimens examined by Inoue et al. (2009) fell in their corresponding geographic origin groups.

The minimal genetic distance separating the genus Notopterus into two main groups is well above 3% (using cytochrome oxidase I marker) which is considered as a conservative threshold between population and species levels in Vertebrates. Furthermore, the existence of a so-called barcode gap between these two groups, along with the fixation of more than 40 diagnostic nucleotide changes in cytochrome oxidase I, strongly indicate that Notopterus is formed by two species. One species is distributed in South Asia (from the Indus basin to Ganga-Brahmaputra basin) and the other in Southeast Asia (from the Salween basin to Mekong basin plus Malay Peninsula, Sumatra and Java).

(A) Map covering the Oriental biogeographic region and showing the distribution of localities (black and red circles) of specimens of Notopterus examined. Black star indicates the estimated type locality of Notopterus notopterus in Java (i.e. Jakarta region) and red stars indicate the likely origin localities of the two syntypes of Notopterus synurus (i.e. coast of Malabar and Coromandel coast at Tharangambadi, formerly Tranquebar). Main Oriental river basins from West to East: (1) Indus basin; (2) Ganga-Brahmaputra river system; (3) Irrawaddy basin; (4) Salween basin, (5) Mekong basin. Insert shows the left lateral view of a specimen of Notopterus notopterus (Penang State, west Peninsular Malaysia; 20 cm in standard length, voucher specimen number USMFC (3) 00002, NO_1). (B) Unrooted network constructed with the software PopArt and a median-joining algorithm showing the cytochrome oxidase I haplotype relationships within the genus Notopterus. Branch lengths are not proportional to the number of changes. Red circles indicated haplotypes of Notopterus synurus (South Asia) and black circles indicated haplotypes of Notopterus notopterus (Southeast Asia). Lavoué et al. (2020).

A 1992 study by Tyson Roberts of the California Academy of Sciences examined the variation of several meristic characters of Notopterus notopterus throughout its entire range. He did not report any significant intraspecific variability that could be additional evidence for the recognition of more than one species in the genus Notopterus. The low amount of morphological variability within the genus Notopterus, however, is not surprising given that the family Notopteridae is known for its morphological stasis. Several valid species of Notopteridae are morphologically similar. For example, species of the genus Chitala are only distinguishable based on their colour pattern and more than one species is suspected to occur within Chitala lopis.

Lavoué et al.'s cytochrome oxidase I-based results strongly support the presence of two allopatric species of Notopterus which need names. There are several nominal species of Notopterus which have been described from Southeast Asia and South Asia and several of these names are available. To determine which name should be applied to each of our two species, Lavoué et al. examined synonym lists and checked the date of description and type locality of these synonyms.

Notopterus notopterus was described and illustrated by Peter Pallas in 1769 as Gymnotus notopterus from a specimen said to have been collected nearby Ambon (Ambon Island), in the Indian Ocean, a region where this species has never been recorded since. More recent studies have suggested that the type locality given in the description of Pallas is an error because this species does not occur on the Island of Ambon which lies east of the Wallace Line, in a different biogeographical region. Furthermore, the local vernacular name of this species 'Ikan Pangaio', reported by Pallas, is in the Malay language which was not used in Ambon at that time. For these reasons, ichthyologist Maurice Kottelat has suggested that the specimen used by Pallas  for the description of Notopterus notopterus should have been collected in Java where the Dutch established their main colony in Indonesia. Consequently, the name Notopterus notopterus should be retained for the species occurring in Southeast Asia with Java as its type locality. There is no type specimen known.

Marcus Bloch and Johann Schnieder described Clupea synura in 1801 from two syntypes, a name which was soon considered as a junior synonym of Notopterus notopterus. Bloch and Schneider first indicated that these specimens are from the coast of Malabar, India. However, in a following remark signed only by Johann Schneider, China and Tranquebar (now known as Tharangambadi, India) were listed as the localities of the syntypes. Notopterus, however, does not seem to occur in China, and zoologist Hans-Joachim Paepke found convincing explanations why Schneider could have mistaken China with the coast of Malabar, India. According to Paepke, the localities of the dry right skins of the two syntypes of Clupea synura that are housed in the Berlin Museum under catalogue numbers ZMB 8806 and ZMB 32057 are coast of Malabar and Tranquebar, respectively. Lavoué et al. suggest the revalidation of Clupea synura for the species of Notopterus occurring in South Asia which should be recognised as Notopterus synurus. 

Whereas the genetic evidence presented in Lavoué et al.'s work supports the recognition of two valid living species of Notopterus, a detailed morphological comparison of the two species is lacking. Such morphological study is needed to identify possible diagnostic characters (in addition to the molecular diagnostic characters presented by Lavoué et al.) and to document the early diversification of the genus Notopterus in the Orient. In this respect, a fossil of Notopterus, morphological similar to living species, indicates that this genus was already present in Sumatra at least 33 million years ago (the Eocene-Oligocene boundary). In addition, because of the cryptic diversity occurring in the genus Notopterus and the difficulty to identify the type localities of species of this genus, it will be important to designate a neotype for Notopterus notopterus and a lectotype for Notopterus synurus. Finally, the geographic coverage needs to be expanded with the study of specimens collected from the Mekong basin, Indus basin and, especially, from the region comprising the Irrawaddy basin situated between the Ganga-Brahmaputra river system and the Salween basin to determine the location of the exact distributional limit between these two species.

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Triplophysa weiheensis: A new species of Tibetan Loach from the Weihe River in Gansu Province, China.

Tibetan Loach of the genus Triplophysa are the most common members of the Stone Loach family, Nemacheilidae, found on the Qinghai-Tibet Plateau. These fish are found in almost all water bodies in the region, with new species of Triplophysa still being reported regularly. As a result, a total of 147 valid species of Triplophysa have been recorded to date, including Triplophysa zhaoi, which is found living in the swamps of the Lükqün Oasis in the Turpan Depression, at an altitude of 50 m below sealevel, making it the lowest living freshwater Fish in Asia, and Triplophysa stolickai which is found in hot springs near the Longmu Lake in western Tibet, at an altitude of 5200 m, making it the highest altitude Fish in Asia, and possibly the highest altitude Fish in the world. 

In a paper published in the journal Zoological Research in July 2020, Chen-Guang Feng of the Key Laboratory of Adaptation and Evolution of Plateau Biota of the Northwest Institute of Plateau Biology of the Chinese Academy of Sciences, and the School of Ecology and Environment at the Northwestern Polytechnical University, Yu Zhang, also of the Key Laboratory of Adaptation and Evolution of Plateau Biota, and of the University of the Chinese Academy of Sciences, Chao Tong, again of the Key Laboratory of Adaptation and Evolution of Plateau Biota, and of the Biology Department at the University of Pennsylvania, Bing-Zheng Zhou and Xiao-Hui Li, again of the Key Laboratory of Adaptation and Evolution of Plateau Biota, and of the University of the Chinese Academy of Sciences, Yong-Tao Tang, again of the Key Laboratory of Adaptation and Evolution of Plateau Biota, and of the College of Fisheries at Henan Normal University, and Wen-Zhu Song and Kai Zhao, once again of the Key Laboratory of Adaptation and Evolution of Plateau Biota, describe a new species of Tibetan Loach from the Weihe River in Gansu Province, China.

Weihe River is a tributary of the Yellow River and originates from the southern part of Gansu Province. Previous studies have reported that southern Gansu is a hotspot area for Triplophysa, with an extensive distribution of species across the river systems. Until now, 17 species of Triplophysa have been reported from the Yellow River system, five of which have been recorded from Weihe River. Following an investigation of Triplophysa species from Weihe River, 15 specimens superficially resembling Triplophysa stoliczkae were collected and are described as a new species based on morphological and molecular analyses.

Map showing sampling sites of Triplophysa weiheensis  (star symbol, solid star for type locality). (1) Zhang River, at Chenjiamo Village, Zhangxian County; (2) Niutou River, at Hongbao Town, Qingshui County; (3) Niutou River, at Maiji District, Tianshui City. Feng et al. (2020).

Previous research has reported that Triplophysa stoliczkae is a striking case of morphological convergence and consists of distinct lineages that are not close relatives. As there has been no formal taxonomic revision for Triplophysa stoliczkae, Feng et al. treated it as a morphological species in this study but considered its different genetic lineages in  phylogeny. These lineages, which were initially mistaken as Triplophysa stoliczkae, exhibit very similar morphology. Thus, they represent a known morphological unit in the genus Triplophysa. Feng et al. specifically measured 61 Triplophysa stoliczkae specimens collected from various water systems and used principal component analysis to visualize morphological differences between Triplophysa stoliczkae and the new species.

The new species is named Triplophysa weiheensis, where 'weiheensis' means from the Weihe River. The body of Triplophysa weiheensis is thick and cylindrical. The dorsal profile of the body is arch-like. Maximum depth of body slightly greater than maximum width, occurring between pectoral and dorsal fins. Caudal peduncle laterally compressed, depth nearly uniform toward caudal-fin base, length longer than head length. Head width greater than depth. Cheeks slightly inflated, V-shaped outline in ventral view. Snout obtuse, sloping downward anterior to anterior nostril. Snout length shorter than postorbital length. Anterior and posterior nostrils close together. Valves around anterior nostrils, but not around posterior. Eyes small, dorsolaterally in head. Interorbital space wide. Mouth inferior. Lips thick and well-developed with furrows; lower lip continuous with shallow median incision. Lower jaw crescentic, uncovered by lower lip. Three pairs of barbels thick, moderately short; inner rostral barbels almost reaching corner of mouth; outer rostral barbels horizontally reaching posterior nostril; maxillary barbels horizontally reaching mid-point of eyes.

Lateral view, dorsal view, and ventral view of Triplophysa weiheensis, NWIPB 1505189, holotype, 65.8 mm. Feng et al. (2020).

Fins short, dorsal fin rounded distally, originating anterior to pelvic-fin origin; dorsal-fin origin near midway between tip of snout and caudal-fin base or slightly nearer to caudal-fin base. Pectoral fin short, not reaching halfway point to pelvic-fin base. Pelvic fin reaching past anus, distal fin tip attaining anal-fin origin when adpressed. Anal fin rounded distally, just posterior to anus. Caudal fin slightly emarginate.

Lateral view of Triplophysa weiheensis, NWIPB 1505189, holotype, 65.8 mm (a) and Triplophysa stoliczkae NWIPB 1707001, 85.1 mm, from the Yellow River system (b). Feng et al. (2020).

Skin scaleless. Lateral line tapering, interrupted on posterior trunk at pelvic-fin distal extremity; few lateral line pores close to caudal-fin base. Stomach U-shaped, expanded. Intestine long, with 6–7 loops, one hidden loop surrounded by 5–6 loops). Bony capsule of air bladder small and thin, closed laterally; posterior chamber of air bladder absent.

Micro computerised tomograph of skeletal system of Triplophysa weiheensis NWIPB 1505183, paratype, 64.1 mm. Feng et al. (2020).

In mature males, unbranched and four outer branched pectoral-fin rays thickened, covered with breeding tubercles on dorsal surface. Small breeding tubercles also present on both sides of head in liber apophyses extending from anterior lower margin of orbit to base of outer rostral barbel. These characters do not occur in females.

Sexual dimorphism of Triplophysa weiheensis. Dimorphic characters are highlighted in red circles. Feng et al. (2020).

Specimens were collected from flowing streams with gravel or sandy substrates at 1360–1768 m above sealevel. River water became muddy at about 0.5 m deep. Periphytic Algae, sand, and Insect larvae (e.g. Chironomids) were found in the stomach of specimens. Other species collected with Triplophysa weiheensis included Triplophysa dalaica, Triplophysa minxianensis, Pseudorasbora parva, Gobio huanghensis.

Habitat of Triplophysa weiheensis. (a) Type locality: Zhang River, at Chenjiamo Village, Zhangxian County; (b) Niutou River, at Hongbao Town, Qingshui County. Kai Zhao in Feng et al. (2020).

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Trematocranus pachychilus: A new species of Cichlid Fish from Lake Malaŵi.

The Cichlids are an extremely successful group of freshwater Perciform Fish (Perches), found in North, Central and South America, as well as the Caribbean, the Middle East, South Asia and, most notably Africa, where the group reaches its highest diversity. Over 800 species have been described from Lake Malaŵi alone, which is more species of freshwater fish than have been described from the entire Northern Hemisphere and slightly over 1% of every vertebrate species ever described. The genus Trematocranus currently contains three species, all endemic to Lake Malaŵi. These are fairly large Fish, reaching about 250 mm in length, typically having a spotted pattern, with several rows of teeth on their pharyngeal plates.

In a paper published in the journal ZooKeys on 14 March 2018, Katrien Dierickx and Mark Hanssens of Ichthyology at the Royal Museum for Central Africa, Bosco Rusuwa of the Department of Biology at the University of Malawi, and Jos Snoeks, also of Ichthyology at the Royal Museum for Central Africa, and of the Laboratory of Biodiversity and evolutionary genomics at Leuven University, describe a new species of Trematocranus from Jafua Bay on the Mozambique shore of Lake Malaŵi.

The new species is named Trematocranus pachychilus, meaning 'thick-lip', in reference to the lips of this species, which are notably thicker than those of other members of the genus. The species is described from three specimens caught in 1998 and stored in alcohol at the Royal Museum for Central Africa. These range from 117.7 to 154.6 mm in length, and have lost their original colouration. They are deep bodied and laterally flattened, with teeth sharp and pointed at the front of the mouth and flat and rounded at the back; the rear teeth of the outer rows are bicusped, while the inner row are unicusped, though in larger specimens there are more bicusped teeth.

Trematocranus pachychilus, preserved specimen, adult male, 154.6 mm SL, Lake Malawi, Jafua Bay. Dierickx et al. (2018).

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Charax awa: A new species of Charicid Fish from Maranhão State in northestern Brazil.

Charicids are freshwater Boney Fish related to Piranhas and Tetras that are found across South and Central America and as far north as Mexico and parts of Texas. Members of the genus Charax are medium sized members of this group, reaching a maximum of about 130 mm in length, most abundant in the Andean regions, but also found in southern and western Brazil and parts of the Guyanan Shield. No described member of this genus has a distribution that includes the coastal basins of northeastern Brazil, but there are several records of Fish apparently belonging to this group being found there. 

In a paper published in the journal Zoosystematics and Evolution on 1 February 2018, Erick Cristofore Guimarães of the Programa de Pós-Graduação em Biodiversidade e Conservação, and the Laboratório de Sistemática e Ecologia de Organismos Aquáticos at the Universidade Federal do Maranhão, Pâmella Silva De Brito of the Programa de Pós-Graduação em Biodiversidade e Biotecnologia da Amazônia and the Laboratório de Sistemática e Ecologia de Organismos Aquáticos at the Universidade Federal do Maranhão, Beldo Rywllon Abreu Ferreira, also of the Programa de Pós-Graduação em Biodiversidade e Conservação at the Universidade Federal do Maranhão, and Felipe Polivanov Ottoni, of the Programa de Pós-Graduação em Biodiversidade e Conservação, the Programa de Pós-Graduação em Biodiversidade e Biotecnologia da Amazônia, the Laboratório de Sistemática e Ecologia de Organismos Aquáticos, and the Programa de Pós-Graduação em Oceanografia at the Universidade Federal do Maranhão, describe a new species of Charax from the Rio Mearim, Rio Munim and Rio Turiaçu basins, of Maranhão State in northestern Brazil.

The new species is named Charax awa, where ‘awa’ is the name used by the Guajá people of Maranhão State to describe themselves. Adults of this species range from 40.2 to120.0 mm in length and are silvery in colour, darker on the upper surface and towards the front, and with blotchy markings on the fins. The upper part of the head is distinctly concave, and the eyes somewhat small. The upper and lower jaws both have enlarged canine teeth as well as rows of smaller, conical teeth. 

 Charax awa, live specimen from a stream at the Miranda do Norte, in the Mearim River Basin of Maranhão State, Brazil. Erick Guimarães in Guimarães et al. (2018).

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Pseudogobiopsis lumbantobing: A new species of freshwater Gobiid Fish from Java and Sumatra.

The freshwater Gobiid Fish genus Pseudogobiopsis is found from Thailand southeast to Papua New Guinea, though it is best known from the European aquarium trade, with collectors frequently sending professional ichthyologists photographs of unidentified species for identification.

In a paper published in the Raffles Bulletin of Zoology on 1 June 2017, Helen Larson of the Museum and Art Gallery of the Northern Territory and the Museum of Tropical Queensland, Renny Hadiaty of the Museum Zoologicum Bogoriense of the Indonesian Institute of Sciences and Nicolas Hubert of the Institut de Recherche pour le Développement, describe a new species of Pseudogobiopsis from the freshwater drainages of western Sumatra and western Java.

The new species is named Pseudogobiopsis lumbantobing, in honour of Daniel Lumbantobing of Jakarta, who collected the first specimens from the wild that were directly brought to the attention of scientists (although a number of aquarists had previously sent Helen Larson photographs of similar Fish of unknown provenance). The species is described from eighteen specimens, ranging from 18 to 33.5 mm in length, pearly grey in colour, with a with a white belly and a scattering of small orange spots. 

 

 Captive specimen of Pseudogobiopsis lumbantobing, imported via Singapore. Emma Turner in Larson et al. (2017).

This also potentially solves the mystery of a painting by Kuhl and van Hasselt, made between 1820 and 1823, of an unknown Gobiid Fish from Java. Unfortunately the original of this painting is in the collection of the Naturalis Library, which is currently undergoing renovation, so that the picture was in storage and unavailable for inspection prior to publication of the paper.

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Devario fangae & Devario myitkyinae: Two new species of Striped Devario from northern Myanmar.

Devarios, Devario spp., are small freshwater Cyprinid Fish found in hillstreams across South and Southeast Asia. There are currently about 38 described species from Bhutan, India, Myanmar, Nepal, Pakistan, Sri Lanka, southern China, Cambodia, Laos, Malaysia, Vietnam, and Thailand, which can be divided into two groups, Barred and Striped Devarios, based upon their markings, though this probably does not reflect the true diversity of the group. There are currently six species listed from Myanmar, though one of these, Devario aequipinnatus, is almost certainly a misidentification.

In a paper published in the journal Zootaxa on 3 February 2017, Sven Kullander of the Department of Zoology at the Swedish Museum of Natural History describes two new species of Striped Devario from the Ayeyarwaddy River Basin in northern Myanmar.

The first new species is named Devario fangae, in honour of the late ichthyologist Fang Fang Kullander (wife of Sven Kullander), who first identified the new species, but sadly passed away before she was able to formally describe it. The species was found in hillstreams around Putao in Kachin State in March 1998, and is 43-61 mm in length. The specimens from which the species is described are pale creamish brown in colour with darker stripes, though they have been kept in preservative for over a decade and their original colour is unclear. 

Devario fangae. (a) Male, 47.9 mm; (b) male 58.5 mm; (c) female, 49.2 mm. All from Kachin State, Putao, Ayeyarwaddy River drainage, Nan Hto Chaung. Kullander (2017). 

The second new species is named Devario myitkyinae, in reference to the city of Myitkyina in Kachin State, near which it was discovered close to. The species is described from specimens collected from Hpa Lap stream in the upper Ayeyarwaddy River Basin by Fang Fang Kullander in 1997. These are 42.8-68.7 mm in length, and pale brown with a yellowish white belly and darker stripes in preservative, the original colour is unknown. 

Devario myitkyinae. (a) Male , 65.8 mm; (b) male, 61.8 mm; (c) female, 56.7 mm. All from Myanmar: Kachin State, Ayeyarwaddy River drainage, Hpa Lap Chaung near Myitkyina. Kullander (2017).

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