Tadpole of the Vietnamese jelly-nest tree frog, Feihyla palpebralis (Anura: Rhacophoridae): external morphology and chondrocranium.

The genus Feihyla Frost, Grant, Faivovich, Bain, Haas, Haddad, de S, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green & Wheeler was established relatively recently (Frost et al. 2006), but to date its composition has changed several times due to the extremely confusing and labile systematics of small rhacophorids (e.g., Grosjean et al. 2008; Hertwig et al. 2013; Biju et al. 2020). It embeds rather miniature, delicate tree frogs and currently comprises six species (Frost 2023). The type species, Feihyla palpebralis (Smith 1924), was described from Mount Langbian in the southern part of the Truong Son Range (Annamite Mountains) and was long considered endemic to Vietnam; recent reports that it is also distributed in China can be based on misidentification and need reliable confirmation (Biju et al. 2020).

Feeding habits and multifunctional classification of soil-associated consumers from protists to vertebrates.

Soil organisms drive major ecosystem functions by mineralising carbon and releasing nutrients during decomposition processes, which supports plant growth, aboveground biodiversity and, ultimately, human nutrition. Soil ecologists often operate with functional groups to infer the effects of individual taxa on ecosystem functions and services. Simultaneous assessment of the functional roles of multiple taxa is possible using food-web reconstructions, but our knowledge of the feeding habits of many taxa is insufficient and often based on limited evidence. Over the last two decades, molecular, biochemical and isotopic tools have improved our understanding of the feeding habits of various soil organisms, yet this knowledge is still to be synthesised into a common functional framework. Here, we provide a comprehensive review of the feeding habits of consumers in soil, including protists, micro-, meso- and macrofauna (invertebrates), and soil-associated vertebrates. We have integrated existing functional group classifications with findings gained with novel methods and compiled an overarching classification across taxa focusing on key universal traits such as food resource preferences, body masses, microhabitat specialisation, protection and hunting mechanisms. Our summary highlights various strands of evidence that many functional groups commonly used in soil ecology and food-web models are feeding on multiple types of food resources. In many cases, omnivory is observed down to the species level of taxonomic resolution, challenging realism of traditional soil food-web models based on distinct resource-based energy channels. Novel methods, such as stable isotope, fatty acid and DNA gut content analyses, have revealed previously hidden facets of trophic relationships of soil consumers, such as food assimilation, multichannel feeding across trophic levels, hidden trophic niche differentiation and the importance of alternative food/prey, as well as energy transfers across ecosystem compartments. Wider adoption of such tools and the development of open interoperable platforms that assemble morphological, ecological and trophic data as traits of soil taxa will enable the refinement and expansion of the multifunctional classification of consumers in soil. The compiled multifunctional classification of soil-associated consumers will serve as a reference for ecologists working with biodiversity changes and biodiversity-ecosystem functioning relationships, making soil food-web research more accessible and reproducible.

First description of the tadpole of Kurixalus baliogaster (Inger, Orlov, and Darevsky, 1999) (Anura: Rhacophoridae) from Vietnam, with comments on reproductive biology.

To date, 20 species of Kurixalus Ye, Fei, and Dubois have been described, and all of these species are distributed throughout South and Southeast Asia, from eastern India, throughout Myanmar and the mountainous regions of southern China, to Indochina, western and northern peninsular Thailand, Malaysia, Sumatra, Borneo, and the Philippines (Frost 2021). Descriptions of the tadpoles of only 6 species have been published: K. berylliniris and K. wangi Wu, Huang, Tsai, Li, Jhang, Wu (Wu et al. 2016); K. eiffingeri (Boettger) (Kuramoto Wang 1987); K. idiootocus (Kuramoto Wang) (Kuramoto Wang 1987); K. cf. verrucosus (Boulenger) (Ziegler Vences 2002), and Kurixalus yangi Yu, Hui, Rao, Yang (Humtsoe et al. 2020). A description of the tadpoles of K. baliogaster (Inger, Orlov, Darevsky) is also given in the species description (Inger et al. 1999), but described larvae are assigned tentatively to this species in the published text. Additional studies on the identification of the conspecificity of the described tadpoles with K. baliogaster have not been conducted. Based on the much larger size of the tadpole body (TL up to 40.3 mm), as well as the labial tooth row formula 6(26)/5(1) given by Inger et al. (1999), we concluded that these described tadpoles cannot be larval K. baliogaster and most likely belong to some other species of rhacophorid frogs.

Larval morphology of three syntopic species of emKaloula/em Gray (Anura: Microhylidae) from Vietnam.

The external larval morphology of three species of the microhylid genus Kaloula (K. indochinensis, K. mediolineata, and K. pulchra) inhabiting the southern and central regions of Vietnam is studied. The similarities and the distinctive features of their morphometric characters, the structure of their mouthparts and spiracle, and their coloration are analysed with consideration of the geographic variability. A description of the tadpole of K. indochinensis is provided for the first time. The interspecific comparison revealed the shape of the mouthparts and the spiracle as the most reliable diagnostic characters for the field identification of the coexistent Kaloula tadpoles. The first description of the larval chondrocranium and hyobranchial apparatus of K. indochinensis demonstrates a set of morphological traits characteristic of suspension-feeding microhylids. Some developmental parameters (egg number and size, duration of embryonic and larval development, larvae size and stage at hatching) are provided for K. indochinensis and K. pulchra.

Skull development in the Iberian newt, Pleurodeles waltl (Salamandridae: Caudata: Amphibia): timing, sequence, variations, and thyroid hormone mediation of bone appearance.

The cranial ossification sequence in Pleurodeles waltl is widely used in phylogenetic analyses of amphibian origin and evolution. However, the patterns published to date are far from completely resolved and contain certain discrepancies. Based on a large sample of P. waltl specimens ranging from early post-hatching larvae to post-metamorphic newts, we determined the most common cranial ossification sequence and revealed its intraspecific variations. Since thyroid hormones (THs) are involved in the mediation of skull development in salamanders, we studied the role of THs in the cranial development of P. waltl. The normal sequence and timing of bone appearance were compared with those in larvae reared under conditions of high (in 1 and 2 ng mL-1 triiodothyronine) and low [in 0.02% thiourea (TU), which inhibits thyroid gland activity] TH levels. Metamorphosis was greatly accelerated in the TH-treated larvae and was arrested in the TU-treated larvae, which retained the larval pattern of the palate and rudimentary external gills even after 2 years of the experiment. Early-appearing bones (the coronoid, vomer, palatine, dentary, squamosal, premaxilla, parasphenoid, pterygoid, prearticular, vomer, frontal, parietal, exoccipital, in this order) arise at the same stages and ages, and follow the same ossification sequence under different TH levels. The timing of the appearance of bones normally arising in the late larval and metamorphic periods (the quadratojugal, orbitosphenoid, prootic, maxilla, nasal, os thyroideum, prefrontal, quadrate, in this order) changes depending on the TH level. The maxilla and nasal display the most pronounced reaction to changes in the TH level: they appear precociously in TH-treated animals, while their appearance is postponed and they remain rudimentary in TU-treated animals. Because of different responses to THs, the order in which late-arising bones appear changes depending on the TH level. Although bones appearing early in larval ontogeny (e.g. the premaxilla, vomer, squamosal, palatine) display no TH-induced reaction when they start to develop, their further differentiation shows dependence on THs, and these bones become TH-inducible closer to metamorphosis. These findings indicate that TH involvement in the mediation of cranial development changes from minimal (if at all) in its early stages to maximal during metamorphosis. It is likely that the appearance of bones early in development is mediated by factors other than THs. Their further development is accompanied by changes in the mechanisms mediating their morphological differentiation. That is, likely non-hormonal mediation becomes replaced or/and complemented by hormonal mediation. The constituent parts of the same bone may exhibit differences in their reactions to changes in TH levels. Although in normal development, the overall cranial ossification sequence is constant, there was variation in the order in which late-appearing bones was recorded. These observations suggest that this variation results from individual variability in the internal TH level. Comparison with other salamanders suggests that (a) the pattern of TH mediation described in P. waltl is common for cranial development of metamorphosing urodeles and (b) the same bone may differ in its TH dependence in different salamanders, e.g. there are interspecific variations in the degree of TH dependence of individual cranial bones.

Larval morphology of Hylarana lateralis (Boulenger) (Anura: Ranidae) from southern Vietnam.

The Kokarit Frog, recently referred to as Hylarana lateralis (Boulenger), is widely distributed throughout Indochina (Cambodia, Laos, Vietnam), Thailand, Myanmar and southern China. The position of this species in ranid systematics remains unclear; accordingly to the latest revision by Oliver et al. (2015), it was formally withdrawn from the genus Hylarana, but not assigned to any other hylaranine genus and currently is referred to as "Hylarana" lateralis (Frost 2019).

Tadpole of Microhyla picta Shenkel (Anura: Microhylidae), an endemic narrow-mouthed frog from Vietnam.

The Painted Rice Frog, Microhyla picta Schenkel, is one of the least studied species among narrow-mouthed frogs of the genus Microhyla. To date it is known only from Vietnam, where it is distributed mostly in eastern coastal areas in central and southern parts of the country (Nguyen et al. 2009; Nguyen Hoang 2013). The species was also found in Con Dao archipelago (Poyarkov Vassilieva 2011). No data are available on the ecology and reproductive biology of the species, and its larval morphology is also not described. Microhyla frogs are widespread and numerous in terrestrial ecosystems and microhylid tadpoles are known to play an important role in the aquatic communities of temporary waterbodies of monsoon ecosystems in Southeast Asia (Heyer 1973; Vassilieva et al. 2017). The ability to identify microhylid tadpoles in the field is pertinent for biodiversity and ecological studies. We provide a description of larvae of M. picta from various areas in southern Vietnam.

Molecular, morphological and acoustic assessment of the genus Ophryophryne (Anura, Megophryidae) from Langbian Plateau, southern Vietnam, with description of a new species.

Asian Mountain Toads (Ophryophryne) are a poorly known genus of mostly small-sized anurans from southeastern China and Indochina. To shed light on the systematics within this group, the most comprehensive mitochondrial DNA phylogeny for the genus to date is presented, and the taxonomy and biogeography of this group is discussed. Complimented with extensive morphological data (including associated statistical analyses), molecular data indicates that the Langbian Plateau, in the southern Annamite Mountains, Vietnam, is one of the diversity centres of this genus where three often sympatric species of Ophryophryne are found, O. gerti, O. synoria and an undescribed species. To help resolve outstanding taxonomic confusion evident in literature (reviewed herein), an expanded redescription of O. gerti is provided based on the examination of type material, and the distributions of both O. gerti and O. synoria are considerably revised based on new locality records. We provide the first descriptions of male mating calls for all three species, permitting a detailed bioacoustics comparison of the species. We describe the new species from highlands of the northern and eastern Langbian Plateau, and distinguish it from its congeners by a combination of morphological, molecular and acoustic characters. The new species represents one of the smallest known members of the genus Ophryophryne. At present, the new species is known from montane evergreen forest between 700-2200 m a.s.l. We suggest the species should be considered Data Deficient following IUCN's Red List categories.

The phytotelm tadpoles of Microhyla arboricola (Anura: Microhylidae) from Vietnam, with comments on reproductive biology and development.

The reproductive biology of Microhyla arboricola (Microhylidae) was studied in two regions of the southern Annamite Mountains in Vietnam. M. arboricola is an obligate phytotelm-breeder that reproduces in water-filled tree hollows in montane evergreen forests. Clutches are attached above the water level in the hollows and contain 5-37 pigmented, relatively large eggs. Larvae hatch at markedly advanced stages and develop in water until metamorphosis is completed. The developing tadpoles are obligately oophagous and feed on conspecific eggs and embryos. M. arboricola tadpoles differ from tadpoles of pond-dwelling Microhyla species in their external morphology (extremely long tails, dorsolateral position of the eyes, dark pigmentation), digestive tract morphology (large, extensible larval stomach and short intestine), and oral morphology. The larval chondrocranium and hyobranchiumof M. arboricola is described. M. arboricola shares its habitat with other hollow-breeding species of anurans. To date, M. arboricola is the only known arboreal species of the genus Microhyla that has a unique reproductive mode. The ecological niche of this species differs greatly from those occupied by other microhylids of Indochinese Peninsula.

Larval morphology and complex vocal repertoire of Rhacophorus helenae (Anura: Rhacophoridae), a rare flying frog from Vietnam.

We present new data on the distribution, reproduction, larval morphology and vocalization of Rhacophorus helenae (Rhacophoridae), a narrowly distributed frog from southern Vietnam. Two new populations of R. helenae were discovered during field surveys in the lowland monsoon forests in Dong Nai and Ba Ria-Vung Tau provinces in 2010-2013. Spawning was observed in May 2013. Egg clutches containing small (2.3±0.1 mm) unpigmented eggs were embedded in a foam nest and suspended high on trees above temporary ponds. The tadpoles of R. helenae have a morphology typical of pond-dwelling Rhacophorus larvae with a moderate tail length and a labial tooth row formula of 5(2-5)/3. Postmetamorphic juveniles differed from adult frogs in the features of their coloration and less developed webbing. The complex vocal repertoire of R. helenae included five types of tonal, wideband and pulsed calls and several transitional signal types differentiated by frequency and amplitude parameters. Calls were uttered as singular signals (pulsed calls) or within non-stereotyped series of variable duration (other call types). The complex structure of the advertisement call markedly distinguishes R. helenae from other members of the Rhacophorus reinwardtii species complex.

A new species of the genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from coastal southern Vietnam.

A new species of the genus Oligodon from the coastal area of Binh Chau-Phuoc Buu Nature Reserve, Ba Ria-Vung Tau Province, southern Vietnam, is described. Oligodon arenarius sp. nov. is distinguishable from all other species by the unique combination of the following characters: medium size; 17 dorsal scale rows; 6-8 maxillary teeth, the posterior three being enlarged; head scalation lacking a loreal but usually including a presubocular; divided nasal; two postoculars; 131-144 ventrals; 36-60 subcaudals; unforked hemipenis, without spines or obvious papillae; sexual dimorphism displayed in the number of subcaudals (36-40 in females, 58-60 in males) and a relative tail length, tails being quite long in males (TaL/TL = 0.26-0.28) and moderate in females (TaL/TL = 0.13-0.17); head coloration pattern including ocular band, temporal bands and chevron-shaped mark on nape; dorsal coloration without distinct pattern, uniform or with dark speckling; ventrals pinkish in life, immaculate. To date, this species appear to be endemic to Vietnam.

A new species of Leptolalax (Anura: Megophryidae) from the western Langbian Plateau, southern Vietnam.

We describe a new species of megophryid frog from Loc Bac forest in the western part of the Langbian Plateau in the southern Annamite Mountains, Vietnam. Leptolalax pyrrhops sp. nov. is distinguished from its congeners by a combination of the following morphological attributes: (1) presence of distinct dark brown/black dorsolateral markings, including blackish spots on flanks and dark canthal and/or temporal streaks; (2) rudimentary webbing on toes; (3) tympanum externally distinct; (4) dorsal skin finely shagreened with numerous small tubercles and pustules; (5) medium size for the genus (30.3-33.9 mm in 2 adult males, 30.8-34.3 mm in 7 females); (6) grey-pinkish to dark brownish-violet chest and belly with numerous whitish speckles, also covering the lateral sides of body; (7) ventrolateral glands small, indistinct, do not form a distinct line; (8) pectoral glands comparatively small, comprising 1-3% of adult SVL; (10) iris bicolored, typically bright orange-red in upper two-thirds, fading to silvery green in lower third. The male advertisement call of the new species is also unique among those Leptolalax species for which calls are known, with a single long 'introductory' note, consisting of 5-12 pulses, followed by of 4-5 predominantly single-pulsed notes, and an average dominant frequency of 1.91-2.23 kHz. From the morphologically similar L. applebyi, L. melicus and L. bidoupensis, Leptolalax pyrrhops sp. nov. can be further distinguished by 13.5%, 13.7% and 10.3% sequence divergence at the 16S rRNA mtDNA gene. At present, the new species is known from montane evergreen forest between 800-1100 m elevation. We suggest the species should be considered as Data Deficient following IUCN's Red List categories. To date our finding represents the southernmost known record of the genus Leptolalax from Vietnam.

Two new species of Kalophrynus Tschudi, 1838 (Anura: Microhylidae) from the Annamite mountains in southern Vietnam.

We describe two new mountain-dwelling microhylid species of the genus Kalophrynus from the southern part of the Annamite Mountains in Vietnam. The two new species differ from all known congeners in morphological characters and mtDNA; phylogenetically, they form a sister clade to the large-bodied K. interlineatus (1009 bp, 16S rRNA gene, mtDNA). Both species share the following characteristics: snout pointed in dorsal and lateral views, slightly sloping in profile; tympanum distinct, smaller than eye in diameter; toe webbing moderate; outer metatarsal tubercle present; light dorsolateral line absent. Kalophrynus cryptophonus sp. nov. from Loc Bao, Lam Dong Province is a small-sized species distinguishing from its congeners by a combination of: SVL 27.9-30.4 mm in males, 23.4 mm in female; canthus rostralis indistinct; males with large sharp conical spines on the skin covering mandible margins and finely asperous nuptial pads on the dorsal surface of fingers I-III; dark ocelli in the inguinal region usually present, small, without a light border; anterior palatal dermal ridge short, restricted to medial part of palate. Kalophrynus honbaensis sp. nov. from Hon Ba, Khanh Hoa Province is a medium-sized Kalophrynus, distinguishing from its congeners by a combination of: SVL 26.7-36.8 mm in males; canthus rostralis distinct; males without distinguishable spines on the mandible margins nor the nuptial pads; dark ocelli in the inguinal region present, large, without a distinct light border, anterior palatal dermal ridge developed, parallel to posterior one. Kalophrynus cryptophonus sp. nov. reproduces in hollow bamboo stems; we describe larval morphology and bioacoustics of this species in relation to phytotelm breeding. A review of the distribution of the genus Kalophrynus in Indochina is provided.

A new species of Kukri Snake (Oligodon Fitzinger, 1826; Squamata: Colubridae) from the Cat Tien National Park, southern Vietnam.

We describe a new species of the genus Oligodon from the lowland forests of Cat Tien National Park, Dong Nai Province, in southern Vietnam. Oligodon cattienensis sp. nov. is distinguished from the remaining Southeast Asian kukri snakes by the combination of the following characters: medium-sized, deeply forked hemipenes without spines, 17-17-15 dorsal scale rows, nasal entire, 2 small postoculars, almost equal in size, 167-178 ventrals, 31-35 subcaudals, 24-35 + 5 large dark-edged vertebral blotches in combination with a yellow-orange or red vertebral stripe between blotches, head pattern including ocular band, temporal bands and elongated chevron, ventrals pink or whitish (reddish in juveniles) in life, some bearing a quadrangular dark blotch on each lateral side, or ventrals being entirely dark. Based on the hemipenial morphology the new species is assigned to the Oligodon cyclurus species group. A comparison table for all Indochinese Oligodon is provided.