taxonID	type	description	language	source
C30487CAFFF1FFB1838AED5BFE544E1D.taxon	discussion	Tang et al. (2023) described N. laojunshanensis from the Hengduan Mountains in Yunnan province, China, as the sister lineage of the lineage leading to Nanorana pleskei Günther, 1896 and Nanorana ventripunctata Fei & Huang, 1985. Morphologically, N. laojunshanensis exhibits a distinct combination of characteristics from known congeners, such as the presences of a tympanum, equal lengths of fingers I and II, a small body size, a yellow ventral surface of limbs, distinct vomerine teeth, indistinct subarticular tubercles, a head width greater than head length, slender supratympanic fold, the absence of the dorsolateral fold, the presence of nuptial spines on fingers I and II in adult males, the absence of a vocal sac, and paired brown spines on the chest. Within the nominal subgenus Nanorana, this new taxon is distinguished by indistinct subarticular tubercles and by “ lacking dark blotches on ventral surface and ventral surface of limbs yolk yellow ” (Tang et al. 2023). Coincidently, another species, Nanorana huangi Ji, Shi, Ma, Shen, Chang & Jiang, 2023, was described from the exact same area and published by a different team (Ji et al., 2023) during the same period (Autumn 2023). The type locality of N. huangi (26.874593 ° N, 99.544008 ° E; 3389 m a. s. l) is distanced by only about 30 km (by air line; Fig. 1 C) from the type locality of N. laojunshanensis (Mt. Laojun, Lijiang, Yunnan, China; 26 ° 37 ’ N, 99 ° 42 ’ E, 3982 m a. s. l.) and both populations are morphologically similar. In this respect, the morphological assessment of N. huangi is based on a greater sample size than N. laojunshanensis, and accordingly captures higher intraspecific variability (particularly for quantitative traits), which challenges the characteristics previously reported as diagnostic for N. laojunshanensis (e. g., body size, shape of supratympanic fold, coloration). Our re-analysis of 16 S and COI also reveals strong similarities, with both species sharing identical or closely related haplotypes (Fig. 1 A, B) with low level of genetic diversity (π = 0.2 %), and we therefore consider N. huangi and N. laojunshanensis as subjective synonyms (Fig. 1 A, B). Since the description of N. huangi was published on September 28, 2023, [vs. November 7, 2023, for N. laojunshanensis], N. huangi is treated as the oldest available name. The genetic distances highlighted by Tang et al. (2023) to justify the split of the new taxon from its closest relatives N. ventripunctata and N. pleskei, are relatively low, namely 1.6 % for 16 S and 7.4 % for COI, suggesting a young divergence (see also comment in Dufresnes and Litvinchuk 2022). These distances are typically lower than those presented for other species of the genus at the same loci (i. e., Liu et al. 2021). In addition, the study lacks a comparative analysis for a nuclear, biparentally inherited marker alone, although sequences of the nuclear gene RAG 1 were included. Given the relatively low observed genetic distances for the mtDNA markers, we expect the RAG 1 variability in the investigated sequences between taxa to be also low. Here, we found little differentiation in RAG 1 (for comparison with other species pairs, see Hofmann et al. 2023 a), namely 0.18 % between N. laojunshanensis and N. pleskei, 0.43 % between N. laojunshanensis and N. ventripunctata, and 0.18 % between N. pleskei and N. ventripunctata (all individual sequences were homozygous). Besides, the sequence KY 172605 specified by the authors in their RAG 1 dataset is a tyrosinase fragment. Drawing from these points, we advise caution towards the distinction of N. huangi as a separate species (see the topology in Fig. 1) without additional genetic (ideally genomic) evidence. This example is paradigmatic of hasty species descriptions and represents a certain trend observed in current taxonomic research that is not always beneficial for taxonomy / species conservation itself.	en	Daniel JABLONSKI, Sylvia HOFMANN (2024): Over-splitting and inconsistently applied criteria: a response to recent changes on the taxonomy of mountain spiny frogs (Dicroglossidae, Nanorana). Alytes 41 (1 - 4): 40-48, DOI: 10.5281/zenodo.16903363
C30487CAFFF7FFB083E4EA10FC174D1C.taxon	discussion	as the new subgenus Minipaa Tang, Liu & Yu, 2023 The up ranking of N. minica as its own subgenus, Minipaa, is inconsistent with the phylogenetic evidence. Depending on analysis, this taxon is alternatively placed within an unsupported clade that includes some but not all representatives of Chaparana, Paa Dubois, 1975, and Allopaa (16 S tree), or as the sister lineage of the clade regrouping Paa and Nanorana (the concatenated four genes phylogeny). For lumping Allopaa into Chaparana and erect N. minica as a separate subgenus, Tang et al. (2023) used published genetic data (see their Fig. 4) and mention two morphological information for N. minica taken from the Ohler & Dubois (2006), namely “ transculent or creamy nuptial spines and entirely whitish or creamy eggs, without colored animal pole. ”. However, only 15 of the 34 known species of the genus Nanorana (and only 9 of 13 species of the subgenus Paa) have been evaluated at that time by Ohler & Dubois (2006). Moreover, neither of those two features are included in the original description of N. minica. Whether these two characteristics are unique to N. minica thus remain to be demonstrated by investigating other spiny frogs. In addition, the molecular background of the new subgenus still requires confirmation because it remains unclear if the sequences attributed to N. minica in Tang et al. (2023) belong to this species. Thus, for reasons of taxonomic stability, N. minica should continue to be treated as member of the subgenus Paa (Ohler & Dubois 2006).	en	Daniel JABLONSKI, Sylvia HOFMANN (2024): Over-splitting and inconsistently applied criteria: a response to recent changes on the taxonomy of mountain spiny frogs (Dicroglossidae, Nanorana). Alytes 41 (1 - 4): 40-48, DOI: 10.5281/zenodo.16903363
C30487CAFFF7FFB1833BEDC7FC26494C.taxon	discussion	Reassigning this species to a different genus than Nanorana (where it was placed by Qi et al. 2019) was recently discussed by Hofmann et al. (2023 a), based on its phylogenetic position in a 16 S mitochondrial tree and its high sequence differentiation (> 14 %) from any other Nanorana members. However, the type locality was not genetically assessed, and whether the analyzed vouchers correspond to this taxon deserves confirmation. Therefore, enforcing this species in Nanorana as Tang et al. (2023) suggested appears premature, and because its phylogenetic, and therefore taxonomic placement is unsettled, it would be correct and more parsimonious to classify this taxon as “ insertae sedis ”, and continue to refer to it as Odorrana arunachalensis Saikia, Sinha & Kharkongor, 2017 (for more details see the discussion in Hofmann et al. 2023 a). The case of Allopaa hazarensis (Dubois & Khan, 1979) in subgenus Chaparana Bourret, 1939 In the 16 S Bayesian phylogeny of Tang et al. (2023; Fig. 3), this taxon, endemic to northern Pakistan, branches (without support) as the sister lineage of Nanorana unculuanus (Liu, Hu, Yang, 1960) (subgenus Chaparana), noting that none of the known Nanorana subgenera form a monophyletic clade in this analysis. In their concatenated four genes phylogeny, A. hazarensis is robustly retrieved as the sister lineage of Chaparana. Based on these conflicting observations, Tang et al. (2023) present Allopaa as being lumped with Chaparana. However, the authors do not discuss this new arrangement, which otherwise appears superficial. According to Hofmann et al. (2021 a, 2023 a), the extremely disjunct distribution range (Allopaa is the westernmost Himalayan taxon among Nanorana, while Chaparana members occur at the opposite side of the Himalaya-Tibet orogen), as well as the morphological data of Ohler & Dubois (2006) on Allopaa, this taxon should continue to be treated as a separate subgenus of Nanorana (Hofmann et al. 2024).	en	Daniel JABLONSKI, Sylvia HOFMANN (2024): Over-splitting and inconsistently applied criteria: a response to recent changes on the taxonomy of mountain spiny frogs (Dicroglossidae, Nanorana). Alytes 41 (1 - 4): 40-48, DOI: 10.5281/zenodo.16903363
C30487CAFFF6FFB08327EEC0FA9848BE.taxon	discussion	In addition to the above-mentioned four cases, our re-analysis of the 16 S data available on GenBank also offer the opportunity to discuss the status of the recently described N. bangdaensis Rao, Hui, Zhu & Ma, 2022, which is nested within Nanorana parkeri (Stejneger, 1927) (Fig. 1 A, D). Since the sequences of the latter originated from very different localities across the Tibetan Plateau (Fig. 1 E), we suspect that N. bangdaensis might be conspecific with N. parkeri. This alpine species has been intensively studied by population genomics and standard single barcoding-gene approaches using sampling that covers its entire distribution range (Zhou et al. 2014, Wang et al. 2018). These studies revealed major populations [(E) ast and (W) est], defining them as evolutionarily significant units, and up to four subpopulations (E 1 - E 4) with only low whole-genomic differentiation among them. A similar east-west population structure has been previously identified in the endemic Tibetan snake Thermophis baileyi (Wall, 1907) (Hofmann et al. 2014), consistent with a scenario of range expansion from different refugia during interglacial and post-glacial times. Since one of the eastern N. parkeri subpopulation (E 4; KJ 811207 and KJ 811261) corresponds to the type locality of N. bangdaensis (Bangda, [Baxoi County], Qamdo Prefecture, Tibet Autonomous Region, China), we used the COI-sequence data of N. bangdaensis and of the N. parkeri (sub) populations W and E 1 ― E 4, comprehended by N. bangdaensis / N. parkeri sequence data of Ji et al. (2023) and Tang et al. (2023), to assess their respective sequence divergence (Fig. 1 D, E). As suspected, the COI sequences of N. bangdaensis are identical to sequences representatives of the East lineage of N. parkeri suggesting that N. bangdaensis and N. parkeri are weakly differentiated and likely belong to the same species. The lack of clear morphological diagnostic features further supports this assessment (https: // www. amphibiachina. org as cited from Rao et al. 2020; original in Chinese: “ The length of the head is approximately equal to the width of the head; the tympanic membrane is not obvious; the skin on the back is smooth, with a few short warts or skin folds on the back; the back is gray-green in life, with no obvious dark spots; the sides of the body are light in color, mixed with irregular spots. ”). While examples of “ super-cryptic species ” (i. e., cryptic species that experienced a mitochondrial capture, thus necessitating nuclear data for their detection; Dufresnes et al. 2019) do exist, and notwithstanding the possibility for an unusually young speciation event, it appears more parsimonious to assume that N. bangdaensis is part of the high intraspecific variability of N. parkeri, and might either be synonymized or be distinguished as a subspecies (see genetic distances, Fig. 1 D).	en	Daniel JABLONSKI, Sylvia HOFMANN (2024): Over-splitting and inconsistently applied criteria: a response to recent changes on the taxonomy of mountain spiny frogs (Dicroglossidae, Nanorana). Alytes 41 (1 - 4): 40-48, DOI: 10.5281/zenodo.16903363
