Journal of Threatened Taxa | www.threatenedtaxa.org | 26 July 2025 | 17(7): 27171–27194

 

ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) 

https://doi.org/10.11609/jott.9461.17.7.27171-27194

#9461 | Received 23 October 2024 | Final received 12 June 2025 | Finally accepted 09 July 2025

 

 

Two new species of bush frogs (Anura: Rhacophoridae: Raorchestes) from Meghalaya, northeastern India

 

Holiness Warjri 1, Jayaditya Purkayastha 2, Hmar Tlawmte Lalremsanga 3  & Madhurima Das 4        

 

1,4 Department of Zoology, Assam Don Bosco University, Sonapur, Assam 782402, India.

2,4 Help Earth, 16, Raghunath Choudhury Path, Lachitnagar, Guwahati, Assam 781007, India.

3 Developmental Biology and Herpetology Laboratory, Department of Zoology, Mizoram University, Tanhril, Mizoram 796004, India.

1 holiness12345@gmail.com, 2 mail.jayaditya@gmail.com (corresponding author), 3 htlrsa@yahoo.co.in

4 madhurima.das@dbuniversity.ac.in (corresponding author)

 

 

 

 

ZooBank: urn:lsid:zoobank.org:pub:086F7F0F-6CBF-4339-BC55-89939B9C5203

Editor: S.R. Ganesh, Kalinga Foundation, Agumbe, India.         Date of publication: 26 July 2025 (online & print)

 

Citation: Warjri, H., J. Purkayastha, H.T. Lalremsanga & M. Das (2025). Two new species of bush frogs (Anura: Rhacophoridae: Raorchestes) from Meghalaya, northeastern India. Journal of Threatened Taxa 17(7): 27171–27194. https://doi.org/10.11609/jott.9461.17.7.27171-27194

  

Copyright: © Warjri et al. 2025. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article in any medium by providing adequate credit to the author(s) and the source of publication.

 

Funding: This research was funded by the Joint CSIR-UGC Junior Research Fellowship awarded to Holiness Warjri [CSIR-HRDG Ref. No.: Nov/06/2020(i)EU-V], provided by the Council of Scientific and Industrial Research  Human Resource Development Group (CSIR-HRDG), Government of India.

 

Competing interests: The authors declare no competing interests.

 

Author details: Holiness Warjri is a research scholar at the department of Zoology, Assam Don Bosco University. Jayaditya Purkayastha is a wildlife biologist at Help Earth NGO. Tlawmte Lalremsanga is a professor at the department of Zoology, Mizoram University. Madhurima Das is an assistant professor at the department of Zoology, Assam Don Bosco University.

 

Author contributions: HW—conceptualization, paper writing, collection of specimens, field study and data gathering; JP—paper writing and phylogenetic analysis; HTL—provided morphological data; MD—laboratory work and paper writing.

 

Acknowledgements: We extend our sincere gratitude to the principal chief conservator of forests, wildlife, and chief wildlife warden, Meghalaya (Memo. Nos. FWC/Research/115/2790−2796, FWC/Research/115/1653−1661, and FWC/Research/115/1134−1142) for permitting us to conduct this study. Financial support for this work was provided through the Joint CSIR-UGC fellowship [CSIR-HRDG Ref. No.: Nov/06/2020(i)EU-V], awarded to Holiness Warjri. We also thank the Department of Zoology, Assam Don Bosco University, for their support during the study. We deeply thank PI Gracy and Cynthia Mylliem Umlong for their assistance and support. We extend our heartfelt appreciation to Rijessing Warjri for his unwavering dedication and invaluable support throughout the study, whose expertise and commitment were instrumental in the success of this project. We also wish to express our deep appreciation to Nangiaisan Wahlang, Aldanes Syiemlieh, Joyforever Kharbani, Skhemjingmut Nongrum, Klasstarwell Warjri, Antarwell Warjri, Rises Warjri, and Ibankershisha Dkhar for their generous help and contributions in the fieldwork.

 

 

Abstract: The genus Raorchestes Biju et al., 2010 represents one of the most diverse lineages within the family Rhacophoridae, with its members characterized by small size, absence of vomerine teeth, direct development, and distinctive digital discs with circum-marginal grooves. Despite its location in the Indo-Burma biodiversity hotspot, the amphibian diversity of Meghalaya remains underexplored. In this study, we describe two new species, Raorchestes jakoid sp. nov. and Raorchestes jadoh sp. nov., from the Khasi Hills of Meghalaya, based on an integrative approach combining morphological, bioacoustic, and molecular data (16S rRNA). Additionally, we provide new records and supplementary descriptions for three species: R. kempiae, R. garo, and R. asakgrensis, thereby improving our understanding of their intraspecific variation and distribution. Phylogenetic analyses confirm the placement of the new species within the Raorchestes parvulus species complex. The new species are distinguished by a suite of morphological traits, unique call structures, and genetic divergence from congeners. Notably, both new species were discovered in secondary habitats near human settlements, suggesting ecological tolerance yet raising concerns about their long-term survival amid rapid habitat loss. The elevational distribution of the species studied ranges from 235 m to 1,655 m, with a concentration between 1,000–1,600 m, highlighting patterns of elevational partitioning and habitat specialization. Our findings add to the growing evidence of high cryptic diversity in the region and underscore the urgent need for targeted herpetofaunal surveys and conservation actions in Northeast India.

 

Keywords: 16SrRNA, acoustic, biodiversity, conservation, endemism, Indo-Burma biodiversity hotspot, morphology, parvulus complex, systematics, taxonomy.

 

 

INTRODUCTION

 

The genus Raorchestes Biju et al., 2010 is one of the most diverse in the family Rhacophoridae (Vijayakumar et al. 2014), currently comprising 80 recognized species (Biju & Bossuyt 2009; Frost 2025). The distribution of Raorchestes spans a wide geographical range, from southern and northeastern India to Nepal, extending through Myanmar, Thailand, Laos, and southern China, reaching Vietnam, Cambodia, and western Malaysia (Frost 2025). These small frogs are characterized by their unique morphological traits, including an adult snout-vent length between 15–45 mm and the absence of vomerine teeth. Notably, all documented species undergo direct development, with eggs hatching directly into froglets, bypassing the free-swimming tadpole stage. Additionally, the genus is distinguished by the expanded tips at its fingers, and toes, each ending in discs with circum-marginal grooves (Biju et al. 2010; Seshadri et al. 2012; Wu et al. 2021).

Meghalaya, a state in northeastern India, lies within the Indo-Burma biodiversity hotspot, a region renowned for its remarkable species richness and high levels of endemism (Myers et al. 2000). Despite its ecological significance, much of Meghalaya’s amphibian diversity remains underexplored, with many species still awaiting formal description. Recent studies describing new amphibian taxa points to the rich and underexplored amphibian diversity of Meghalaya (Mathew & Sen 2007, 2009, 2010a, 2010b; Das et al. 2010; Mahony et al. 2011, 2013, 2018; Purkayastha & Matsui 2012; Kamei et al. 2013; Saikia et al. 2023; Naveen et al. 2024, 2025).

Among the Rhacophoridae of Meghalaya, species within the genus Raorchestes are unique in exhibiting direct development. In contrast, all other rhacophorid genera recorded from the region (e.g., Rhacophorus, Polypedates, Kurixalus, & Chirixalus) undergo an indirect development cycle, with a prominent free-swimming aquatic tadpole stage following hatching. Additionally, members of the genus Raorchestes possess rudimentary toe webbing. In comparison, species of other rhacophorid genera exhibit moderate to extensive toe webbing. Based on morphology, bioacoustic and genetics (16S rRNA), herein, two new species of Raorchestes belonging to the Raorchestes parvulus species complex were described (Garg et al. 2021) along with additional data on the newly described R. asakgrensis Naveen, Chandramouli, Babu, Ryndongsngi, Karunakaran & Kumara 2024, and of redescribed R. garo (Boulenger, 1919), and R. kempiae (Boulenger, 1919) from the subtropical forests of Meghalaya. Naveen et al. (2025) synonymised two nominal taxa Philautus namdaphaensis Sarkar & Sanyal, 1985 and P. manipurensis Mathew & Sen, 2009 under R. kempiae. These newly described and redescribed species significantly enhance the known diversity within the genus, offering fresh insights into the biogeography of Raorchestes, and contributing to the broader understanding of amphibian diversity in this ecologically rich but understudied region.

 

 

MATERIALS AND METHODS

                                                                                                                

Field sampling

Fieldwork was carried out in different parts of Meghalaya where specimens were observed and collected (Figure 1), including Mawpat, South West Khasi Hills (25.359⁰ N, 91.255⁰ E, elevation 1,355 m) for ADBUHW0154, Mawsynram, East Khasi Hills (25.296⁰ N, 91.585⁰ E, elevation 1,445 m) for ADBUHW0141, Lawbah, East Khasi Hills (25.247⁰ N, 91.580⁰ E, elevation 815 m) for ADBUHW0143, ADBUHW0145, and ADBUHW0144, Langtor, Eastern West Khasi Hills (25.533⁰ N, 91.586⁰ E, elevation 1,655 m) for ADBUHW0124 and ADBUHW0123, Laittyra, East Khasi Hills (25.222⁰ N, 91.742⁰ E, elevation 755 m) for ADBUHW0052, Mawiong Nongkhlaw, Eastern West Khasi Hills (25.690⁰ N, 91.641⁰ E, elevation 875 m) for ADBUHW0049, Umdein, West Khasi Hills (25.637⁰ N, 91.047⁰ E, elevation 445 m) for ADBUHW0119, Lailad, Nongkhyllem, Ribhoi (25.897⁰ N, 91.775⁰ E, elevation 235 m) for ADBUHW0116, and Tura, West Garo Hills (25.519⁰ N, 90.210⁰ E, elevation 375 m) for ADBUHW0169, and ADBUHW0170 between 2022 and 2024.

The research was conducted under the permission of the Principal Chief Conservator of Forests, Wildlife and Chief Wildlife Warden, Meghalaya (Memo. No. FWC/Research/115/2790−2796, Memo. No. FWC/Research/115/1653−1661 & Memo. No. FWC/Research/115/1134−1142). The study used randomized walks (Lambert 1984) and visual encounter surveys (Crump & Scott 1994) on different sites. The frogs were located and recorded by paying attention to the male frogs’ calls. The frogs were photo-documented using Canon EOS M50 Mark II. Collected frogs were euthanized using lignocaine (2%), preserved in 10% formaldehyde solution, and deposited in the Assam Don Bosco University.

 

Morphological Study

The measurements were made using a vernier calliper to the nearest 0.01 mm. Different morphometric characters and descriptions were considered following Watters et al. (2016). The morphological characters include: head width (HW); snout-vent length (SVL); tibia length (TL); interorbital distance (IOD); head length (HL); eye diameter (ED); internarial distance (IND); eye−nostril distance (EN); snout length (SL); snout−nostril length (NS); foot length (FL); tympanum diameter (TD); thigh length (THL); hand length (HAL); forearm length (FLL); upper eyelid width (UEW); tarsus length (TSL); mandible to eye distance (MBE); upper arm length (UAL); horizontal tympanic annulus diameter (TAD); mandible−nostril distance (MN); hindlimb−length (HLL); lower arm length (LAL); body width (BW); snout−urostyle length (SUL); finger I length (Fin1L); finger II length (Fin2L); finger III length (Fin3L); finger IV length (Fin4L); finger I disk width (Fin1DW); finger II disk width (Fin2DW); finger III disk width (Fin3DW); finger IV disk width (Fin4DW); finger IV width (Fin4W); toe I length (Toe1L); toe II length (Toe2L); toe III length (Toe3L); toe IV length (Toe4L); toe V length (Toe5L); and toe IV disk width (Toe4DW). For morphological comparison, we used raw measurements. We compared these measurements with the members of Raorchestes parvulus complex (Garg et al. 2021; Naveen et al. 2024, 2025): Raorchestes kempiae, and Raorchestes garo.

 

Bioacoustics recording and analyses

The calls were captured in real−time using BOYA BY−DMR7 unidirectional handheld microphones (WAV format, 24−bit) and SONY ICD−PX470 stereo digital voice recorders (MP3 format, 256 kbps) between 1600 h and 2300 h. The distance between the recording equipment and the calling males was maintained at 30−150 cm, except few which are present at the top of big trees, with recording levels adjusted before each session (Prasad et al. 2020). The acoustic properties of the different species were analyzed using Raven Pro 1.6. For calls, we analyzed the temporal variables like the number of notes per call, note duration, duration of the silent interval between notes, call duration, duration of the silent intervals between the calls, and call repetition rate. The spectral variables include the dominant frequency. Descriptive statistics like mean, standard deviation, range, and coefficient of variance were computed using Microsoft Excel 2010.

Genetic study: Genomic DNA isolation from 10 samples of Raorchestes spp. (ADBUHW0049, ADBUHW0052, ADBUHW0116, ADBUHW0124, ADBUHW0141, ADBUHW0143, ADBUHW0145, ADBUHW0154, ADBUHW0169, ADBUHW0170) was done using Phenol: Chloroform: Isoamyl alcohol method, followed by 0.8% agarose gel electrophoresis, and visualized under UV light using Biorad Gel Imazer Gel Documentation Unit. The 16S rRNA gene were amplified using gene- primers pair AH-16S_S   5’- CGC CTG TTT ACC AAA AAC ATC GCC T-3’ and AH-16S_R 5’- TGC GCT GTT ATC CCY RGG GTA ACT-3’ following Caranza & Arnold (2006). Comparative genetic data from members of Raorchestes parvulus group were download from GenBank and were used in construction of phylogenetic tree (Table 2). Sequence alignment was done using MUSCLE (Edgar 2004) in MEGA7 (Tamura & Nei 1993; Kumar et al. 2016) with default parameter settings (max 2072 bp). Maximum likelihood (ML) phylogenetic tree was reconstructed using unpartitioned dataset in IQ-TREE (Nguyen et al. 2015) with the substitution model TIM2+F+R2 selected based on the BIC scores by Model Finder (Kalyaanamoorthy et al. 2017). The ML analysis was run with an ultrafast bootstrap option (Minh et al. 2013) for 1,000 iterations to assess clade support. The uncorrected pairwise p-distance was calculated in MEGA7 (Kumar et al. 2016) with pairwise deletions of missing data and gaps.

 

 

RESULTS

 

Based on 16S rRNA data generated, the species studied were seen to be members of the Raorchestes parvulus species group (Garg et al. 2021; Image 1; Supplement Table 1). The specimens ADBUHW0049 (PQ492285), ADBUHW0052 (PQ492286), and ADBUHW0145 (PQ492281) were seen to form a sister clade with R. kempiae differing by an uncorrected p-distance of 0.000−0.006 (0.6%). Specimens, ADBUHW0141 (PQ492279) and ADBUHW0154 (PQ492282) were seen to form a sister taxon to R. garo, differing by an uncorrected p-distance of 0.000.

ADBUHW0116 (PQ492287), ADBUHW0169 (PQ492283), and ADBUHW0170 (PQ492284) were seen to be sister taxa to R. asakgrensis, differing from each other with an uncorrected p-distance of 0.000−0.015 (0.15%).

Furthermore, specimen ADBUHW0124 (PQ492288) was seen to be sister taxon to R. shillongensis, differing by an uncorrected p-distance of 0.034 (3.4%), and ADBUHW0143 (PQ492280) was seen to be sister to R. rezakhani, differing by an uncorrected p-distance of 0.058 (5.8%) to 0.061 (6.1%).

 

Systematics

Raorchestes kempiae (Boulenger, 1919)

(Images 2 & 3; Tables 1 & 3)

 

Referred materials

ADBUHW0049, adult male collected from Mawiong Nongkhlaw, Eastern West Khasi Hills, Meghalaya, India (25.690⁰ N, 91.641⁰ E, elevation 875 m on 30 August 2023 at around 1845 h by Holiness Warjri and Rijessing Warjri.

ADBUHW0052 adult male collected from Laittyra, East Khasi Hills, Meghalaya, India 25.222⁰ N, 91.742⁰ E, elevation 755 m) on 7 August 2023 at around 1930 h by Holiness Warjri and Rijessing Warjri.

ADBUHW0145, adult male collected from Lawbah, East Khasi Hills, Meghalaya, India (25.247⁰ N, 91.580⁰ E, elevation 815 m) on 6 August 2024 at around 1750 h by Holiness Warjri and Rijessing Warjri.

 

Diagnostic characters

The species is allocated to the genus Raorchestes because of small size (adult SVL ranging from 15−45 mm) and absence of vomerine teeth; a large, transparent gular pouch visible during calls; in males, tips of all fingers and toes expanded into discs with circum-marginal grooves (Biju et al. 2010; Wu et al. 2021). Raorchestes kempiae is characterised by: webbing formula (I 1–2 II 2–1 III 1–3.5 IV 3–2 V); inner palmar tubercle absent and outer palmar tubercle with round shape; tibio-tarsal articulation reaching the eye when hindlimb is stretched alongside body (whereas Boulenger (1919) mentioned of tibio-tarsal articulation reaching the anterior border of an eye); nuptial pad present; inner metatarsal tubercle oval (IMT/Toe4L 0.09 mm long), outer metatarsal tubercle absent; upper part of body with small warts.

 

Color in life (Image 2)

Dorsal color beige, with or without an hour glass shaped marking; finger discs and toe discs greyish; dorsal surface of the hindlimb and forelimb with more or less dark crossbars; supratympanic fold dark brown; iris light golden; ventral surface creamy white with many white spots.

 

Advertisement calls: (Image 3; Table 3)

Ten advertisement calls were analyzed from two individuals having SVL of 23.6 and 24.4 mm. The calling position from the ground was observed to be 20−400 cm. When recording, calling males were sitting on trees and grasses near human settlements and calls were recorded between 1625 h and 1800 h on 7 August 2023. The ambient air temperature was 26⁰C and the relative humidity was 99%.

The advertisement call had a mean duration of 0.37 ± 0.05 seconds, with a mean inter−call interval of 1.32 ± 0.11 seconds, and an average call repetition rate of 2.73 ± 0.33 calls per minute. Each call contained an average of 2.0 ± 0.0 notes, with a note duration of 0.12 ± 0.008 seconds and a mean inter-note interval of 0.06 ± 0.007 seconds. The mean dominant frequency of the call was 2.73 ± 0.03 kHz, ranging 2.72−2.81 kHz, with a coefficient of variation of 1.09.

 

Raorchestes garo (Boulenger, 1919)

(Images 3 & 4; Tables 1 & 3)

 

Referred materials

ADBUHW0154 adult male collected from Mawpat, South West Khasi Hills, Meghalaya, India (25.3590 N, 91.2550 E, elevation 1,355 m) on 17 July 2024 at around 1830 h by Holiness Warjri and Rijessing Warjri.

ADBUHW0141 adult male collected from Mawsynram, East Khasi Hills, Meghalaya, India (25.2960 N, 91.5850 E, elevation 1,445 m) on 6 July 2024 at around 1910 h by Holiness Warjri and Rijessing Warjri.

Diagnostic characters: Raorchestes garo is characterised by small body size (maximum SVL 22.7 mm), vomerine teeth absent, males possess a transparent gular pouch which is visible when calling, and tips of all fingers and toes expanded into discs with circum-marginal grooves. Tympanum may be distinct or hidden. Boulenger (1919) and Naveen et al. (2024) mentioned of distinct tympanum, whereas the specimens reported in this study (ADBUHW0141 and ADBUHW0154) have a indistinct tympanum; webbing between fingers absent; relative finger lengths: I < II < IV < III, relative toe lengths: I < II < III < V < IV; webbing formula for toe is: I 1−1 II 1−1 III 3−3 IV 3−2.5 V; inner palmar tubercle absent and outer palmar tubercle present with round shape; tibiotarsal articulation reaching the posterior border of an eye when hindlimb is stretched alongside body (whereas Boulenger (1919) mentioned tibio-tarsal articulation reaching the anterior border of an eye); nuptial pad present and whitish in color; inner metatarsal tubercle oval (IMT/Toe4L 0.07 mm long), outer metatarsal tubercle absent; dark brown interorbital triangle between eyes interorbital distance larger than eye horizontal diameter; upper part with small warts.

 

Color in life (Image 4)

Dorsal color beige, with a faint) (shaped marking; finger discs and toe discs greyish; dorsal surface of the hindlimb and forelimb with more or less dark crossbars; supratympanic fold dark brown; crotch with distinct black patches; iris golden brown; ventral surface creamy white with many white spots.

 

Advertisement calls: (Image 3; Table 3)

A total of 10 advertisement calls were analysed from 2 individuals having SVL of 22.5 and 22.8 mm. The calling position from the ground is observed to be between 50−500 cm. The calling males were observed and recorded between 1830 h and 2125 h on 17 July 2024 with most calls originating from bushes in dense forest areas and near human settlements. The ambient air temperature during the recordings was 24°C, with a relative humidity of 89 %. The advertisement calls analyzed had a mean duration of 0.03 ± 0.005 seconds, with a mean inter-call interval of 2.26 ± 0.18 seconds, and an average call repetition rate of 35.28 ± 6.15 calls per minute. Each call consisted of a single note, meaning the note duration was identical to the call duration, and the inter-note interval matched the inter-call interval. The mean dominant frequency of the call was 2.93 ± 0.04 kHz, with a frequency range of 2.91−3.00 kHz. The coefficient of variation was 1.35.

 

Raorchestes asakgrensis Naveen, Chandramouli & Babu, 2024

(Images 3 & 5; Tables 1 & 3)

 

Referred materials

ADBUHW0170 adult male collected from Tura, West Garo Hills, Meghalaya, India (25.5190 N, 90.2100 E, elevation 375 m) on 22 July 2024 at around 1830 h by Holiness Warjri and Rijessing Warjri.

Two adult males, ADBUHW0169 and ADBUHW0116 were collected. ADBUHW0169 was collected from the same locality and had the same information as the holotype, and ADBUHW0116 was collected from Lailad, Nongkhyllem, Ribhoi (25.8970 N, 91.7750 E, elevation 235 m) on 27 April 2024 at around 1800 h by Holiness Warjri and Ibankershisha Dkhar.

 

Diagnostic characters

Raorchestes asakgrensis is characterised by: very small body size with maximum SVL of 23.35 mm; head wider than long; tympanum may be distinct or hidden. Rudimentary webbing between toes present, webbing formulaI I 1−2 II 2−2 III 2−3.5 IV 3.3−2 V; relative finger lengths: I < II < IV < III, relative toe lengths: I < II < III < V < IV; inner palmar tubercle absent and outer palmar tubercle present with round shape; tibio-tarsal articulation reaches anterior border of an eye when hindlimb is stretched alongside of body; nuptial pad distinct; inner metatarsal tubercle oval (IMT/Toe4L 0.08 mm long), outer metatarsal tubercle absent; interorbital distance larger than eye horizontal diameter.

 

Color in life (Image 5)

Dorsal color light to dark brown, with no cross bars on the body; finger discs and toe discs light orange and greyish; dorsal surface of the hindlimb and forelimb with more or less dark crossbars; supratympanic fold black; iris light brown.

 

Advertisement calls: (Image 3; Table 3)

Fifteen advertisement calls were analyzed from three individuals having SVLs of 18.1, 21.6, and 23.4 mm. The calling position from the ground is observed to be 50−900 cm. The calling males were observed and recorded from trees and grasses near human settlements between 1745 h and 2100 h on 12 August 2023. The ambient air temperature was 28⁰C and the relative humidity was 99%.

The advertisement call had a mean duration of 0.24 ± 0.02 seconds, with a mean inter−call interval of 0.62 ± 0.09 seconds and an average call repetition rate of 4.17 ± 0.40 calls per minute. Each call contained an average of 4.0 ± 0.0 notes, with a mean note duration of 0.02 ± 0.003 seconds and a mean inter-note interval of 0.03 ± 0.003 seconds. The call’s mean dominant frequency was 3.39 ± 0.04 kHz, ranging 3.36–3.45 kHz, with a coefficient of variation of 0.

 

Raorchestes jakoid sp. nov.

(Image 3 & 6; Table 1, 3 & 4)

urn:lsid:zoobank.org:act:3240A3E3-FF1C-428C-86D0-E6712B4645CC

 

Holotype

ADBUHW0143 adult male collected from Lawbah, East Khasi Hills, Meghalaya, India (25.2470 N, 91.5800 E, elevation 815 m) on 6 July 2024 at around 1800 h by Holiness Warjri and Rijessing Warjri.

 

Paratype

ADBUHW0144 an adult male, other data same as holotype.

 

Etymology

The specific epithet ‘jakoid’ is derived from the Khasi word for “frog” (jakoid), used by the indigenous Khasi people of Meghalaya, northeastern India. The name is treated as a noun in apposition. This nomenclature honours the rich linguistic and cultural heritage of the Khasi community, and emphasizes the importance of local traditional knowledge in the discovery, and conservation of biodiversity in the Khasi Hills.

 

Diagnostic characters

While the developmental mode of the newly described species, Raorchestes jakoid sp. nov., remains uncertain, its placement within the genus Raorchestes is supported by the following combination of characteristics: small body size, vomerine teeth absent, male possesses transparent gular pouch which is visible when calling, and tips of all fingers & toes expanded into discs with circum-marginal grooves. The new species is distinguished from relevant congeners by phylogenetic position, by call analysis based on parameters like dominant frequency, call duration, inter-call interval, call repetition rate, number of note, note duration, and inter-note interval, and by the combination of the following morphological characteristics: small body size with SVL of 18.47−21.6 mm; head wider than long; tympanum indistinct, supratympanic fold distinct; tongue pyriform, and notched posteriorly; webbing between fingers absent; rudimentary webbing between toes present; relative finger lengths: I < II < IV < III, relative toe lengths: I < II < III < V < IV; inner palmar tubercle absent and outer palmar tubercle present with round shape; tibiotarsal articulation reaching posterior border of an eye when hindlimb is stretched alongside of body; nuptial pad distinctly whitish in colour; inner metatarsal tubercle oval (IMT/Toe4L 0.07 long), outer metatarsal tubercle absent; body surface slightly rough and has”)(“ shaped marking; interorbital distance larger than eye horizontal diameter; upper parts with very small warts.

 

Description of the holotype

(Measurement in mm, Table 1) ADBUHW0143 adult male. Body size is small (SVL 18.47). Head is wider than long (HW/HL 1.36); top of the head is relatively flat; snout is slightly rounded and longer than eye diameter (SL/ED 1.34); tympanum indistinct; supratympanic fold distinct; canthus rostralis rounded; internarial distance smaller than interorbital distance (IOD/IND 1.03); nostril is positioned slightly closer to the tip of the snout than to the front corner of the eyes (EN/NS 1.17); tongue pyriform and notched posteriorly; vomerine teeth absent; eye diameter small (ED 1.79) and a large external single subgular vocal sac present.

Forelimbs robust; lower arm is longer than the upper arm (LAL/UAL 3.71). Relative finger lengths: I < II < IV < III; tips of all four fingers expanded into discs in which the disc size is I<II<IV<III; webbing between fingers absent; circum-marginal grooves present; outer metacarpal tubercle distinct and rounded; inner metacarpal tubercle indistinct; no webbing between fingers; small nuptial pad present on the dorsal side of the first finger, whitish.

Hindlimb long (23.5); tibia length is longer than foot length (TL/FL 1.14); relative toe lengths: I < II < III < V < IV; tips of toes with discs and smaller than finger discs; rudimentary webbing between toes present (I 1−1 II 1−2.5 III 2−3 IV 3−1 V); inner metatarsal tubercle oval; outer metatarsal tubercle absent; tibio-tarsal articulation reaching the posterior border of an eye when hindlimb is stretched alongside body.

Dorsum is rough with small warts while the ventral regions are granular, the dorsal surface of the head, forelimb, and hindlimb are covered with very small granules. On the ventral side of digits rounded subarticular tubercles having different sizes are present. Relative size of 0.15 (third subarticular tubercles of Fin 3/Fin 3L) is recorded. Surfaces of hands and toes are granular.

 

Color in life (Image 6)

Dorsal colour light brown, with a black ”)(“ shaped marking; finger discs and toe discs light orange and greyish; dorsal surface of the hindlimb and forelimb with more or less dark crossbars; supratympanic fold black; iris light brown. The groin is dark and has less visible bands on the upper part of the thigh.

 

Advertisement calls (Image 3, Table 3)

The calls of two individuals having SVL of 18.5 and 21.6 mm were observed and recorded from small bushes at 1820−2040 h on 6 July 2024. The calling position from the ground is observed to be 100−350 cm. The ambient air temperature was 27⁰C and the relative humidity was 89%.

The advertisement call had a mean duration of 1.37 ± 0.93 seconds, with a mean inter-call interval of 1.17 ± 0.19 seconds, and an average call repetition rate of 1.07 ± 0.64 calls per minute. Each call contained an average of 17.0 ± 13.31 notes, with the mean note duration being 0.03 ± 0.003 seconds and the mean inter−note interval at 0.04 ± 0.005 seconds. The call’s mean dominant frequency was 3.19 ± 0.0 kHz, with a coefficient of variation of 0.

 

Comparison (see Table 4)

Raorchestes jakoid sp. nov. differs from R. andersoni (Ahl, 1927) by 1) tympanum indistinct vs. tympanum distinct in R. andersoni; 2) rudimentary webbing between the toes vs. feebly webbed in R. andersoni; 3) inner palmar tubercle absent vs. present in R. andersoni; 4) outer palmar tubercle present vs. absent in R. andersoni.

Raorchestes jakoid sp. nov. differs from R. annandalii (Boulenger, 1906) by 1) tympanum indistinct vs. tympanum just distinguishable in R. annandalii; 2) rudimentary webbing between the toes vs. webbed at the base of the toe in R. annandalii; 3) tibiotarsal articulation reaches the posterior border of the eye vs. the eye in R. annandalii; 4) dorsum with small warts vs. smooth in R. annandalii; 5) inner palmar tubercle absent vs. present R. annandalii; 6) outer palmar tubercle present vs. absent R. annandalii.

Raorchestes jakoid sp. nov. differs from R. dulongensis Wu, Liu, Gao, Wang, Li, Zhou, Yuan & Che 2021 by 1) SVL of 18.47−21.6 mm vs. 15.0−19.0 mm in R. dulongensis; 2) tympanum indistinct vs. tympanum distinct in R. dulongensis; 3) nuptial pad present vs. absent in R. dulongensis; 4) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. dulongensis; 5) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. dulongensis; 6) inner palmar tubercle absent vs. present in R. dulongensis.

Raorchestes jakoid sp. nov. differs from R. hekouensis Du, Xu, Liu & Yu, 2024 by 1) SVL of 18.47−21.6 mm vs. 16.1–17.5 mm in R. hekouensis; 2) tympanum indistinct vs. distinct in R. hekouensis; 3) inner palmar tubercle absent vs. present in R. hekouensis; 4) outer palmar tubercle present vs. absent in R. hekouensis; 5) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. hekouensis.

Raorchestes jakoid sp. nov. differs from R. hillisi Jiang Ren, Guo, Wang & Li 2020 by 1) SVL of 18.47−21.6 mm vs. 15.9−17.7 mm in R. hillisi; 2) tympanum indistinct vs. tympanum distinct in R. hillisi; 2) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 1−1 II 1−2.5 III 1−2.5 IV 2.5−1 V of R. hillisi; 3) tibiotarsal articulation reaches posterior border of the eye vs. the eye in R. hillisi; 4) outer palmar present vs. absent in R. hillisi.

Raorchestes jakoid sp. nov. differs from R. huanglianshan Jiang, Wang, Ren, and Li 2020 by 1) SVL of 18.47−21.6 mm vs. 17.0−19.6 mm in R. huanglianshan; 2) tympanum indistinct vs. distinct in R. huanglianshan; 3) tibiotarsal articulation reaches posterior border of the eye vs. the eye in R. huanglianshan; 4) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 1−1 II 1−2 III 1−2 IV 2−1 V of R. huanglianshan.

Raorchestes jakoid sp. nov. differs from R. longchuanensis Yang & Li 1978 by 1) SVL of 18.47−21.6 mm vs. 21.4−23.9 mm in R. longchuanensis; 2) tympanum indistinct vs. tympanum distinct in R. longchuanensis; 3) inner palmar tubercle absent vs. present in R. longchuanensis; 4) relative toe length, I<II<III<V<IV vs. I<II<III ≈ V<IV in R. longchuanensis.

Raorchestes jakoid sp. nov. differs from R. malipoensis Huang, Liu, Du, Bernstein, Liu, Yang, Yu & Wu 2023 by 1) SVL of 18.47−21.6 mm vs. 14.6−17.7mm in R. malipoensis; 2) tympanum indistinct vs. distinct and small in R. malipoensis; 3) outer palmar tubercle present vs. absent in R. malipoensis; 4) inner metatarsal tubercle oval vs. round in R. malipoensis; 5) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. malipoensis; 6) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. malipoensis; 7) nuptial pad present vs. absent in R. malipoensis; 8) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 2−2 II 2−2 III 2−3 IV 3−2 V of R. malipoensis.

Raorchestes jakoid sp. nov. differs from R. meglaensis (Kou, 1990) by 1) SVL of 18.47−21.6 mm vs. 16.6−21.6 mm in R. meglaensis; 2) inner palmar tubercle absent vs. present in R. meglaensis; 3) outer metatarsal tubercle absent vs. present in R. meglaensis; 4) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. meglaensis; 5) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. meglaensis; 6) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 1−1 II 1−2 III 1−2.5 IV 2.5−1 V of R. meglaensis.

Raorchestes jakoid sp. nov. differs from R. parvulus (Boulenger, 1893) by 1) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 2−2 II 1.5−3.25 III 2−3.5 IV 3.25−2 V of R. parvulus; 2) inner metatarsal tubercle oval vs. small in R. parvulus; 3) nuptial pad present vs. absent in R. parvulus; 5) inner palmar tubercle absent vs. present in R. parvulus.

Raorchestes jakoid sp. nov. differs from R. rezakhani Al-Razi, Maria & Muzaffar, 2020 by 1) SVL of 18.47−21.6 mm vs. 18.85−20.90 mm in R. rezakhani; 2) supratympanic fold distinct vs. weakly distinct in R. rezakhani; 3) nuptial pad present vs. absent in R. rezakhani; 4) outer palmar tubercle present vs. absent in R. rezakhani; 5) inner metatarsal tubercle present vs. absent in R. rezakhani; 5) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. rezakhani; 6) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 2−2 II 1.75−2 III 1.5−3 IV 2.75−2 V of R. rezakhani.

Raorchestes jakoid sp. nov. differs from R. yadongensis Zhang, Shu, Liu, Dong & Guo 2022 by 1) SVL of 18.47−21.6 mm vs. 17.8–24.1 mm in R. yadongensis; 2) tympanum indistinct vs. distinct in R. yadongensis; 3) tibiotarsal articulation reaches posterior border of the eye vs. the snout in R. yadongensis.

Raorchestes jakoid sp. nov. differs from R. mindat Köhler, Dost, Than, Ohler, Thammachoti Charunrochana, Chuaynkern, Chuaynkern, and Geiss, 2025 by 1) SVL of 18.47−21.6 mm vs. 16.75−18.36 mm in R. mindat; 2) inner palmar tubercle absent vs. present in R. mindat; 3) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. II 2−3.5 III 2+−3.5 IV 3.25−2 V of R. mindat.

Raorchestes jakoid sp. nov. differs from R. leiktho Köhler, Dost, Than, Ohler, Thammachoti Charunrochana, Chuaynkern, Chuaynkern, and Geiss, 2025 by 1) SVL of 18.47−21.6 mm vs. 15.72−15.80 mm in R. leiktho; 2) supratympanic fold distinct vs. indistinct in R. leiktho; 3) inner palmar tubercle absent vs. present in R. leiktho; 4) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. III 2−3.5 IV 3.33−2 V of R. leiktho.

Furthermore, the new species differs from members of R. parvulus species complex, and other bush frogs of Meghalaya as follows:

Raorchestes jakoid sp. nov. differs from R. garo by 1) tympanum indistinct vs. tympanum distinct in R. garo; 2) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 1−1 II 1−1 III 3−3 IV 3−2.5 V of R. garo; 3) dorsum with small warts vs. smooth in R. garo.

Raorchestes jakoid sp. nov. differs from R. kempiae by 1) SVL of 18.47−21.6 mm vs. 22.96–24.4 mm in R. kempiae; 2) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 1−2 II 2−1 III 1.1−3.5 IV 3−2 V in R. kempiae.

Raorchestes jakoid sp. nov. differs from R. shillongensis (Pillai and Chanda, 1973) by 1) SVL of 18.47−21.6 mm vs. SVL of 10−20 mm of R. shillongensis; 2) inner metatarsal tubercle present and oval vs. indistinct in R. shillongensis; 3) tibiotarsal articulation reaches posterior border of the eye vs. the tympanic region R. shillongensis.

Raorchestes jakoid sp. nov. differs from R. asakgrensis by 1) SVL of 18.47−21.6 mm vs. SVL of 18.0–23.35 mm of R. asakgrensis; 2) tympanum indistinct vs. distinct in R. asakgrensis; 3) webbing formula of I 1−1 II 1−2.5 III 2−3 IV 3−1 V vs. I 1−2 II 2−2 III 2−3.5 IV 3.33−2 V in R. asakgrensis

 

Raorchestes jadoh sp. nov.

(Image 3 & 7; Table 1, 3 & 4)

urn:lsid:zoobank.org:act:FD80F608-0A64-4FDA-B377-EB6882554A5F

 

Holotype

ADBUHW0124 adult male collected from Langtor, eastern West Khasi Hills, Meghalaya, India (25.533o N, 91.586o E, elevation 1655 m) on 10 May 2024 at around 19:00 h by Holiness Warjri and Rijessing Warjri.

 

Paratype

ADBUHW0123 an adult male, other data same as holotype.

 

Etymology

The specific epithet ‘jadoh’ is derived from the name of a traditional rice and meat dish that is integral to the cuisine and cultural identity of the Khasi people of Meghalaya, northeastern India. The name is used here as a noun in apposition. This naming celebrates Khasi heritage and underscores the connection between local biodiversity and indigenous cultural practices.

 

Diagnostic characters

While the developmental mode of the newly described species, Raorchestes jadoh sp. nov., remains uncertain, its placement within the genus Raorchestes is supported by the following combination of characteristics: small body size, vomerine teeth absent, male possess transparent gular pouch which is visible when calling, and tips of all fingers and toes expanded into discs with circum-marginal grooves. The new species is distinguished from relevant congeners by phylogenetic position, by call analysis based on the parameters dominant frequency, call duration, inter-call interval, call repetition rate, number of note, note duration, and inter-note interval, and by the combination of the following morphological characters: very small body size with SVL of 13.68−14.01 mm; head wider than long; tympanum indistinct, supratympanic fold distinct; tongue pyriform and notched posteriorly, webbing between fingers absent; rudimentary webbing between toes present; relative finger lengths: I < II < IV < III, relative toe lengths: I < II < III < V < IV; inner palmar tubercle absent and outer palmar tubercle present with round shape; tibiotarsal articulation reaching posterior border of an eye when hindlimb is stretched alongside of body; nuptial pad present; inner metatarsal tubercle round (IMT/Toe4L 0.08 mm long), outer metatarsal tubercle absent; interorbital distance larger than eye horizontal diameter; upper part with small warts and six fairly distinct warts on the body towards the posterior of the eyes.

 

Description of the holotype (Measurements in mm, Table 1)

ADBUHW0124 adult male. Body size is very small (SVL 13.68). ADBUHW0124 Head is wider than long (HW/HL 1.39); top of the head is relatively flat; snout is slightly rounded; snout is longer than eye diameter (SL/ED 1.70); tympanum indistinct; supratympanic fold distinct; canthus rostralis rounded; internarial distance smaller than interorbital distance (IOD/IND 1.14); nostril is positioned slightly closer to the tip of the snout than to the front corner of the eyes (EN/NS 1.90); tongue pyriform and notched posteriorly; vomerine teeth absent; eye diameter small (ED 1.69) and a large external single subgular vocal sac present.

Forelimbs robust; lower arm is longer than the upper arm (LAL/UAL 1.74). Relative finger lengths: I < II < IV < III; tips of all four fingers expanded into discs in which the disc size is I<II<IV<III; webbing between fingers absent; circum−marginal grooves present; outer metacarpal tubercle distinct and rounded; inner metacarpal tubercle indistinct; no webbing between fingers; small nuptial pad present on the dorsal side of the first finger, whitish.

Hindlimb long (19.02); tibia length is longer than foot length (TL/FL 1.18); relative toe lengths: I < II < III < V < IV; tips of toes with discs, smaller than finger discs; rudimentary webbing between toes present (I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V); inner metatarsal tubercle round; outer metatarsal tubercle absent; tibiotarsal articulation reaching the posterior border of an eye when hindlimb is stretched alongside body.

Dorsum has small warts and six fairly distinct warts on the body towards the posterior of the eyes. At the same time, the ventral regions are granular the dorsal surface of the head, forelimb, and hindlimb are covered with very small granules; under the digits rounded subarticular tubercles having different sizes are present. Relative size of 0.10 (Third subarticular tubercles of Fin 3/Fin 3L) is recorded; surfaces of palm and sole are granular.

 

Color in life (Image 7)

Dorsal color light brown, with a black ”)(“ shaped marking; finger discs and toe discs light orange and greyish; dorsal surface of the hindlimb and forelimb with more or less dark crossbars; supratympanic fold black; iris light brown. The groin is dark and the rear part of the thigh has dark bands.

 

Advertisement calls: (Image 3; Table 3)

10 advertisement calls were analysed from two individuals having SVL of 13.7 and 14.0 mm. The calling males were observed and recorded from trees & bushes at 1800–2100 h on 17 July 2024. The calling position from the ground is observed to be 20−150 cm. The ambient air temperature at the time of recording was 24°C, with a relative humidity of 89%.

The advertisement call exhibited a mean duration of 0.93 ± 0.22 seconds, with a mean inter-call interval of 6.17 ± 0.54 seconds and an average call repetition rate of 1.12 ± 0.27 calls per minute. Each call contained an average of 3.5 ± 0.58 notes, with the mean note duration being 0.02 ± 0.003 seconds and the mean inter−note interval of 0.30 ± 0.01 seconds. The mean dominant frequency of the call was 3.66 ± 1.32 kHz, ranging 3.56−3.84 kHz, with a coefficient of variation of 3.63.

 

Comparison: (see Table 4)

Raorchestes jadoh sp. nov. differs from R. andersoni by 1) tympanum indistinct vs. tympanum distinct in R. andersoni; 2) rudimentary webbing between the toes vs. feebly webbed in R. andersoni; 3) inner palmar tubercle absent vs. present in R. andersoni; 6) outer palmar tubercle present vs. absent in R. andersoni.

Raorchestes jadoh sp. nov. differs from R. annandalii by 1) tympanum indistinct vs. tympanum just distinguishable in R. annandalii; 2) rudimentary webbing between the toes vs. webbed at the base of the toe in R. annandalii; 3) tibio-tarsal articulation reaches the posterior border of the eye vs. the eye in R. annandalii; 4) dorsum with small warts vs. smooth in R. annandalii; 5) inner palmar tubercle absent vs. present in R. annandalii; 6) outer palmar tubercle present vs. absent in R. annandalii.

Raorchestes jadoh sp. nov. differs from R. dulongensis by 1) SVL of 13.68−14.01 mm vs. 15.0−19.0 mm in R. dulongensis; 2) tympanum indistinct vs. tympanum distinct in R. dulongensis; 3) nuptial pad present vs. absent in R. dulongensis; 4) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. dulongensis; 5) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. dulongensis; 6) inner palmar tubercle absent vs. present in in R. dulongensis.

Raorchestes jadoh sp. nov. differs from R. hekouensis by 1) SVL of 13.68−14.01 mm vs. 16.1–17.5 mm in R. hekouensis; 2) tympanum indistinct vs. distinct in R. hekouensis; 3) inner palmar absent vs. present in R. hekouensis; 4) outer palmar tubercle present vs. absent in R. hekouensis; 5) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. hekouensis

Raorchestes jadoh sp. nov. differs from R. hillisi by 1) SVL of 13.68−14.01 mm vs. 15.9−17.7 mm in R. hillisi; 2) tympanum indistinct vs. tympanum distinct in R. hillisi; 2) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 1−2 II 1−2.5 III 1−2.5 IV 2.5−1 V of R. hillisi; 3) tibiotarsal articulation reaches posterior border of the eye vs. the eye in R. hillisi; 4) outer metacarpal present vs. absent in R. hillisi.

Raorchestes jadoh sp. nov. differs from R. huanglianshan by 1) SVL of 13.68−14.01 mm vs. 17.0−19.6 mm in R. huanglianshan; 2) tympanum indistinct vs. distinct in R. huanglianshan; 3) tibiotarsal articulation reaches posterior border of the eye vs. the eye in R. huanglianshan; 4) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 1−1 II 1−2 III 1−2 IV 2−1 V of R. huanglianshan.

Raorchestes jadoh sp. nov. differs from R. longchuanensis by 1) SVL of 13.68−14.01 mm vs. 21.4−23.9 mm in R. longchuanensis; 2) tympanum indistinct vs. tympanum distinct in R. longchuanensis; 3) inner palmar tubercle absent vs. present in R. longchuanensis; 4) relative toe length, I<II<III<V<IV vs. I<II<III ≈ V<IV in R. longchuanensis.

Raorchestes jadoh sp. nov. differs from R. malipoensis by 1) SVL of 13.68−14.01 mm vs. 14.6−17.7mm in R. malipoensis; 2) tympanum indistinct vs. distinctively small in R. malipoensis; 3) outer palmar tubercle present vs. absent in R. malipoensis; 4) inner metatarsal tubercle round vs. round in R. malipoensis; 5) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. malipoensis; 6) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. malipoensis; 7) nuptial pad present vs. absent in R. malipoensis; 8) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 2−2 II 2−2 III 2−3 IV 3−2 V of R. malipoensis.

Raorchestes jadoh sp. nov. differs from R. meglaensis by 1) SVL of 13.68−14.01 mm vs. 16.6−21.6 mm in R. meglaensis; 2) inner palmar tubercle absent vs. present in R. meglaensis; 3) outer metatarsal tubercle absent vs. present in R. meglaensis; 4) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. meglaensis; 5) tibiotarsal articulation reaches posterior border of the eye vs. anterior of the eye in R. meglaensis; 6) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 1−2 II 1−2 III 1−2.5 IV 2.5−1 V of R. meglaensis.

Raorchestes jadoh sp. nov. differs from R. parvulus by 1) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 2−2 II 1.5−3.25 III 2−3.5 IV 3.25−2 V of R. parvulus; 2) inner metatarsal tubercle round vs. small in R. parvulus; 3) nuptial pad present vs. absent in R. parvulus; 5) inner palmar tubercle absent vs. present in R. parvulus.

Raorchestes jadoh sp. nov. differs from R. rezakhani by 1) SVL of 13.68−14.01 mm vs. 18.85−20.90 mm in R. rezakhani; 2) supratympanic fold distinct vs. weakly distinct in R. rezakhani; 3) nuptial pad present vs. absent in R. rezakhani; 4) outer palmar tubercle present vs. absent in R. rezakhani; 5) inner metatarsal tubercle present vs. absent in R. rezakhani; 5) relative toe length, I<II<III<V<IV vs. I<II<V<III<IV in R. rezakhani.

Raorchestes jadoh sp. nov. differs from R. yadongensis by 1) SVL of 13.68−14.01 mm vs. 17.8–24.1 mm in R. yadongensis; 2) tympanum indistinct vs. distinct in R. yadongensis; 3) tibiotarsal articulation reaches posterior border of the eye vs. the snout in R. yadongensis.

Raorchestes jadoh sp. nov. differs from R. mindat by 1) SVL of 13.68−14.01 mm vs. 16.75−18.36 mm in R. mindat; 2) inner palmar tubercle absent vs. present in R. mindat; 3) inner metatarsal tubercle oval vs. rounded in R. mindat; 4) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. II 2−3.5 III 2+−3.5 IV 3.25−2 V of R. mindat.

Raorchestes jadoh sp. nov. differs from R. leiktho by 1) SVL of 13.68−14.01 mm vs. 15.72−15.80 mm in R. leiktho; 2) supratympanic fold distinct vs. indistinct in R. leiktho; 3) inner palmar tubercle absent vs. present in R. leiktho; 4) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. III 2−3.5 IV 3.33−2 V of R. leiktho.

Furthermore, the new species differs from members of R. parvulus species complex and other bush frogs of Meghalaya as follows

Raorchestes jadoh sp. nov. differs from R. garo by 1) tympanum indistinct vs. tympanum distinct or hidden in R. garo; 2) inner metatarsal present vs. indistinct in R. garo; 3) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 1−1 II 1−1 III 3−3 IV 3−2.5 V in R. garo; 3) tibiotarsal articulation reaches the posterior border of the eye vs. the anterior of the eye to the snout in R. garo; 4) dorsum with small warts vs. smooth in R. garo.

Raorchestes jadoh sp. nov. differs from R. kempiae by 1) tibiotarsal articulation which reaches the posterior border of the eyes vs. the eye to the snout in R. kempiae; 2) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 1−2 II 2−1 III 1−3.5 IV 3−2 V in R. kempiae

Raorchestes jadoh sp. nov. differs from R. shillongensis by 1) SVL of 13.68−14.01 mm vs. SVL of 10−20 mm of R. shillongensis; 2) Inner metatarsal tubercle present and round vs. indistinct in R. shillongensis; 3) rudimentary webbing between the toe vs. indistinct rudimentary webbing present between the fourth and fifth toe in R. shillongensis; 4) tibiotarsal articulation reaches posterior border of the eye vs. the tympanic region R. shillongensis.

Raorchestes jadoh sp. nov. differs from R. asakgrensis by 1) SVL of 13.68−14.01 mm vs. SVL of 18.0–23.35 mm of R. asakgrensis; 2) tympanum indistinct vs. distinct in R. asakgrensis; 3) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 1−2 II 2−2 III 2−3.5 IV 3.33−2 V in R. asakgrensis.

Raorchestes jadoh sp. nov. differs from Raorchestes jakoid sp. nov. by 1) SVL of 13.68−14.01 mm vs. 18.47–21.6 mm of R. jakoid; 2) inner metatarsal tubercle round vs. oval of R. jakoid; 3) upper part with small warts and six fairly distinct warts on the body towards the posterior of the eyes vs. small warts randomly distributed in R. jakoid; 4) webbing formula of I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V vs. I 1−1 II 1−2.5 III 2−3 IV 3−1 V in R. jakoid; 5) call duration of 0.93 ± 0.22 seconds vs. 1.37 ± 0.93 seconds of R. jakoid; 6) a mean inter−call interval of 6.17 ± 0.54 seconds vs. 1.17 ± 0.19 seconds of R. jakoid; 7) an average call repetition rate of 1.12 ± 0.27 vs. 1.07 ± 0.64 calls per minute of R. jakoid; 8) 3.5 ± 0.58 notes vs. 17.0 ± 13.31 notes of R. jakoid; 9) note duration 0.02 ± 0.003 seconds vs. 0.03 ± 0.003 seconds of R. jakoid; 10) inter note interval of 0.30 ± 0.01 seconds vs. 0.04 ± 0.005 seconds of R. jakoid; 11) dominant frequency of 3.66 ± 1.32 kHz vs. 3.19 ± 0.0 kHz of R. jakoid; 12) coefficient of variation 3.63 vs. 0 of R. jakoid.

 

 

DISCUSSION

 

Prior to this study, the bush frog diversity in Meghalaya was represented by only five species: R. kempiae, R. garo, R. annandalii, R. shillongensis, and R. asakrensis (Frost 2025). The discovery of two new species from the R. parvulus species complex significantly expands the known diversity of bush frogs in this region, highlighting Meghalaya as a hotspot for amphibian speciation. These findings underscore the critical role that northeastern India plays in amphibian biodiversity within the Indo-Burma region, an area known for its complex biogeography and high endemism (Myers et al. 2000).

In this study, in addition to the new species, an additional description for Raorchestes kempiae, Raorchestes garo, and Raorchestes asakgrensis was given. In R. kempiae, the tibiotarsal articulation reaches the eye when the hindlimb is stretched alongside the body, whereas Boulenger (1919) previously described it as reaching the anterior border of the eye. In R. garo, tympanum visibility varies. While Boulenger (1919) and Naveen et al. (2024) reported a small and fairly distinct tympanum, the specimens examined in this study exhibit an indistinct tympanum. Additionally, the tibiotarsal articulation extends to the posterior border of the eye when the hindlimb is stretched alongside the body, differing from Boulenger’s (1919) observation that it reaches the anterior border of the eye. Similarly, in R. asakgrensis, Naveen et al. (2024) documented a fairly visible tympanum, whereas the specimens analyzed in this study exhibit an indistinct tympanum. These variations in morphological characteristics contribute to a deeper understanding of intraspecific variation and potential taxonomic divergence within the genus Raorchestes. A recent study on Raorchestes of this region also revealed taxonomic inflation as two nominal taxa Philautus namdaphaensis and P. manipurensis were synonymised under R. kempiae, highlighting at the importance of redefining earlier-existing nominal taxa, as a prerequisite to identify new taxa (Naveen et al. 2025).

The distribution of the Raorchestes species across a wide range of altitudes, from 200 m (R. asakgrensis) to 1800 m (R. jadoh sp. nov.), provides valuable insights into habitat specialization and elevational partitioning within the genus. Most species diversity was concentrated between an elevation of 1,000 m and 1,600 m, a pattern observed in other amphibian taxa, where mid-elevation zones provide optimal microhabitats for species diversification (Rahbek 1995; Wu et al. 2021). This concentration of diversity at intermediate elevations is likely driven by the interaction of favourable temperature, humidity, and vegetation cover, which create stable environmental conditions necessary for the persistence of species with specific ecological requirements.

Interestingly, all new species were found in disturbed or secondary habitats near human settlements, suggesting a degree of ecological tolerance. However, this apparent adaptability should not be interpreted as resilience to environmental changes, particularly in a region where habitat degradation from deforestation and agricultural expansion is accelerating. The persistence of these species in human-modified landscapes raises concerns about their long-term survival, given the growing anthropogenic pressures on Meghalaya’s forests, wherein over the last 15 years there has been a 6% loss of forest cover in the West Khasi Hills District (Lyngdoh & Lyngdoh 2023).

The broad distribution of R. asakgrensis across disparate localities, including localities such as Tura and Lailad, Nongkhyllem, suggests that this species has a wider ecological niche than its congeners. This is in contrast to the more restricted ranges of the other species described in this study, which were confined to higher elevations. The aerial distance between populations of R. asakgrensis (approximately 170 km) may indicate either high dispersal ability or the existence of suitable but underexplored habitats between these regions, such as the Garbhanga Reserve Forest and adjacent areas of Assam. Similar patterns of species dispersal across fragmented landscapes have been reported in other Raorchestes species complexes, where geographic isolation and habitat fragmentation contribute to both speciation and range limitation (Vijayakumar et al. 2014; Wu et al. 2019).

The onset of calling activity for all species during the late dry season and the early monsoon (April−August) reflects a reproductive strategy synchronized with the arrival of rainfall, a pattern consistent with other Raorchestes species across their range (Biju et al. 2010; Vijayakumar et al. 2016). The behaviour of calling males, typically occupying small shrubs and bushes between 1–3 m in height, is a well-documented trait in the genus. The calls of R. asakgrensis were notably recorded at higher canopies (up to 9 m), particularly in  Lailad, Nongkhyllem. This canopy-calling behaviour may represent a unique adaptation to its low-elevation, forested habitat, distinguishing it from its higher-elevation congeners.

Meghalaya is home to around 70 species of amphibians (including the present finding) of which around 20 (29%) species were discovered or recorded in the current millennium (since the year 2000), highlighting the importance of the study of amphibians in the landscape of Meghalaya in specific and northeastern India in general. Describing new taxa not only highlights diversity but also helps update conservation criteria. An example of the high rate of cryptic diversity is Cyrtodactylus khasiensis (Jerdon, 1870) which was thought to be a single species with a distribution of whole of northeastern India and adjacent Myanmar. Current studies (mostly since 2018) have pointed to the fact that Cyrtodactylus khasiensis is a species complex currently represented by 35 species of which 26 are endemic to India (Boruah et al. 2024). Most of these species are now found to be point endemic and loss or fragmentation of a small patch of habitat may exterminate the species as a whole (Purkayastha et al. 2020, 2021, 2022; Bohra et al. 2022; Lalremsanga et al. 2022, 2023) Thus, describing such cryptic species has a very important local as well as global conservation implications.

 

 

Table 1.  Morphometric data of Raorchestes garo, R. jakoid sp. nov., R. jadoh sp. nov., R. asakgrensis, and R. kempiae from Meghalaya, India.

 

ADBUHW0154

ADBUHW0141

ADBUHW0143

ADBUHW0144

ADBUHW0123

ADBUHW0124

ADBUHW0169

ADBUHW0170

ADBUHW0116

ADBUHW0049

ADBUHW0052

ADBUHW0145

Species

R. garo

R. jakoid sp. nov.

R. jadoh sp. nov.

R. asakgrensis

R. kempiae

SVL

22.47

22.75

18.47

21.6

14.01

13.68

21.57

23.35

18.08

22.96

24.4

23.62

SUL

21.41

18.11

16.5

20.13

13.24

13

20.52

22.93

16.41

21.64

23.07

22.1

BW

7.21

7.11

7.4

7.51

7.2

7.33

8.53

10.38

7.35

8.59

9.1

9.7

HW

5.79

7.67

7.05

7.52

6.02

5.43

8.35

9.21

6.42

8.87

8.56

8.8

HL

3.95

5.34

5.2

6.11

4.01

3.91

6.54

7.68

5.26

7.13

8.97

7.22

IOD

2.18

2.68

2.19

1.7

2.12

2.1

2.21

2.05

2.13

2.61

2.33

2.79

IND

2.8

1.52

2.12

2.1

1.97

1.85

1.84

2.01

1.98

2.67

3.15

1.89

UEW

1.51

1.81

1.63

1.75

1.51

1.46

1.94

2

1.52

2.06

1.74

1.78

ED

1.68

2.51

1.79

1.8

1.71

1.69

2.24

2.6

1.82

3.21

2.77

2.64

EN

1.71

2.61

1.7

1.89

1.71

1.69

2.24

2.94

1.6

2.48

2

2.53

SL

2.9

3.41

2.4

2.45

2.9

2.87

3.24

4.32

2.78

3.64

4

3.41

NS

0.91

1.41

1.45

1.28

0.9

0.89

1.99

1.71

2.6

1.86

2.01

1.61

TD

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

TAD

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

MBE

1.31

1.31

1.07

1.45

1.21

1.25

1.74

2.17

1.04

2.42

2.04

1.41

MN

3.06

3.58

4.41

5.45

2.91

3.01

4.63

4.88

4.27

5.69

5.05

6.01

FLL

3.49

4.79

4.22

4.29

3.42

3.41

3.72

4.92

4.12

5.33

5.15

5.73

UAL

4.12

4.41

3.09

3.33

4.15

4.11

3.8

3.73

3.08

3.96

3.49

3.25

LAL

7.11

9.11

11.45

12

7.11

7.15

14.33

4.66

11.29

11.04

10.87

10.01

HAL

4.45

5.65

4.85

4.91

4.61

4.5

5.68

6.09

4.78

7.37

7.87

7.19

HLL

19.05

24.01

23.5

23.16

18.61

19.02

32.03

34.97

23.5

33.77

36.47

31.21

THL

6.75

9.19

7.75

8.16

6.75

6.7

11.51

10.8

8.07

12.11

10.71

11.1

TL

7.16

10.11

8.4

10.62

8.11

7.1

11.49

12.91

8.43

11.58

12.23

14.23

TSL

4.08

6.61

3.69

5.49

4.09

4.02

7.49

6.73

3.88

6.53

6.52

5.98

FL

6.04

8.11

7.35

8.11

6.05

6.03

8.58

9.47

7.36

9.38

8.91

9.53

Fin1L

1.61

4.25

2.93

3.18

1.43

1.66

2.45

2.1

2.25

3.12

4.33

3.01

Fin2L

2.1

5.35

3.55

4.11

2

2.11

3.19

3.92

3.65

4.69

5.23

4.15

Fin3L

4.14

6.41

5.35

6.45

4.12

4.15

5.87

6.1

5.3

7.51

7.4

6.94

Fin4L

3.61

5.99

4.28

4.5

4.62

3.59

4.63

5.2

4.22

5.48

6.45

5.73

Fin1DW

0.33

0.46

0.32

0.44

0.33

0.39

0.36

0.44

0.49

0.53

0.61

0.45

Fin2DW

0.61

0.86

0.86

0.89

0.62

0.65

0.81

0.82

0.85

0.93

1.12

0.91

Fin3DW

1.19

0.97

1.2

1.61

1.36

1.29

1.25

1.25

1.14

1.42

1.48

1.45

Fin4DW

0.81

0.88

0.92

0.99

0.99

0.68

0.87

0.84

0.9

1.44

1.48

1.16

Fin4W

0.25

0.56

0.59

0.91

0.21

0.25

0.72

0.87

0.58

0.6

1

0.58

Toe1L

1.31

2.99

2.4

2.91

1.21

1.26

2.3

3.65

2.41

3.29

2.85

3.59

Toe2L

3.39

2.11

3.45

2.48

3.18

2.42

3.1

2.97

2.4

5.55

4.97

4.22

Toe3L

5.19

5.2

5.6

6.78

6

6.11

6.33

6.7

5.49

7.27

6.68

5.99

Toe4L

6.19

7.11

7.5

7.69

6.01

6.45

8.85

9.16

7.41

9.19

9.52

9.52

Toe5L

6.19

6.99

6.7

6.9

5

5.12

7.54

7.36

6.55

8.03

7.32

7.81

Toe4DW

0.87

1.2

0.98

1.03

0.83

0.86

1.12

1.4

0.94

1.41

1.4

1.29

 

 

Table 2. GenBank accession of 16s rRNA gene for species used to construct maximum likelihood phylogenetic tree.

Species

Locality

Voucher specimen

GenBank accession No

Raorchestes andersoni

Medog, Tibet, China

KIZ 014104

MW023610

Raorchestes andersoni

Medog, Tibet, China

KIZ YPX16167

MW023609

Raorchestes annandalii

Nepal

CDZMTU419

MT983169

Raorchestes asakgrensis

Tura, West Garo Hills, Meghalaya

ADBUHW0169

PQ492283

Raorchestes asakgrensis

Tura, West Garo Hills, Meghalaya

ADBUHW0170

PQ492284

Raorchestes asakgrensis

Lailad, Nongkhyllem, Ribhoi, Meghalaya

ADBUHW0116

PQ492287

Raorchestes asakgrensis

Nongkhyllem, Meghalaya

ZSIS-M7

MN524577

Raorchestes asakgrensis.

NA

CESF420

JX092712.1

Raorchestes asakgrensis.

Asakgre Community Reserve, Garo Hills, Meghalaya, India

SACON VA 805

PQ512828

Raorchestes cf. parvulus

Gunung Jerai, Kedah, Malaysia

LSUHC 10473

MH590203

Raorchestes cf. parvulus

Pulau Langkawi, Kedah, Malaysia

LSUHC 7596

MH590202

Raorchestes cf.  parvulus

Gunung Stong, Kelantan, Malaysia

LSUHC 11118

MH590201

Raorchestes dulongensis

 Qinlangdang Village, Yunnan, China

KIZ 035125

MW537815

Raorchestes dulongensis

Qinlangdang Village, Yunnan, China

KIZ 035082

MW537814

Raorchestes dulongensis

Qinlangdang Village, Yunnan, China

KIZ 035126

MW537816

Raorchestes garo

Mawpat, South West Khasi Hills, Meghalaya

ADBUHW0154

PQ492282

Raorchestes garo

Mawsynram, East Khasi Hills, Meghalaya

ADBUHW0141

PQ492279

Raorchestes garo

Daribokgre Community Reserve, East Garo Hills, Meghalaya, India

SACON VA 809

PQ585812

Raorchestes hekouensis

Hekou, Yunnan, China

GXNU YU000536

OQ859106

Raorchestes hekouensis

Hekou, Yunnan, China

GXNU YU000537

OQ859107

Raorchestes hekouensis

Hekou, Yunnan, China

GXNU YU000160

ON986422

Raorchestes hillisi

Xiding, Yunnan, China

MT488411

MT488411

Raorchestes hillisi

Xiding, Yunnan, China

CIB 116329

MT488412

Raorchestes hillisi

Xiding, Yunnan, China

CIB 116330

MT488413

Raorchestes huanglianshan

Mt. Huanglian, Lvchun, Yunnan, China

CIB 116365

MT488414

Raorchestes huanglianshan

Mt. Huanglian, Lvchun, Yunnan, China

CIB 116353

MT488415

Raorchestes huanglianshan

Mt. Huanglian, Lvchun, Yunnan, China

CIB 116354

MT488417

Raorchestes jadoh sp. nov.

Langtor, Eastern West Khasi Hills, Meghalaya

ADBUHW0124

PQ492288

Raorchestes jakoid sp. nov.

Lawbah, East Khasi Hills, Meghalaya

ADBUHW0143

PQ492280

Raorchestes kempiae

Mikadogre Community Reserve

SACON VA 806

PQ512827 

Raorchestes kempiae

Mawiong Nongkhlaw, Eastern West Khasi Hills, Meghalaya

ADBUHW0049

PQ492285

Raorchestes kempiae

Laittyra, East Khasi Hills, Meghalaya

ADBUHW0052

PQ492286

Raorchestes kempiae

Lawbah, East Khasi Hills, Meghalaya

ADBUHW0145

PQ492281

Raorchestes kempiae

Cangyuan, Yunnan, China

KIZ 015855

MN475866

Raorchestes kempiae

Cangyuan, Yunnan, China

KIZ 015856

MN475867

Raorchestes kempiae

Cangyuan, Yunnan, China

KIZ 015857

MN475868

Raorchestes leiktho

Near Leiktho, Hp-an District, Kayin State, Myanmar

SMF 106284

PP694058

Raorchestes leiktho

Near Leiktho, Hp-an District, Kayin State, Myanmar

SMF 106234

PP694060

Raorchestes longchuanensis

Gongdong, Longchuan county, Yunnan, China

KIZ 048492

MN475871

Raorchestes longchuanensis

Gongdong, Longchuan county, Yunnan, China

KIZ 048468

MN475870

Raorchestes malipoensis

Malipo, Yunnan, China

SWFU 3110

ON128247

Raorchestes malipoensis

Malipo, Yunnan, China

SWFU 3111

ON128241

Raorchestes malipoensis

Pac Ban, Tuyen Quan, Vietnam

ROM30288

GQ285674

Raorchestes menglaensis

Zhushihe, Mengla, Yunnan, China

CIB 116338

MT488403

Raorchestes menglaensis

Zhushihe, Mengla, Yunnan, China

CIB 116340

MT488404

Raorchestes mindat

Ovatmataung National Park, Kanpetlet township, Mindat District, Chin State, Myanmar

CAS 234782

PP694093

Raorchestes mindat

Ovatmataung National Park, Kanpetlet township, Mindat District, Chin State, Myanmar

CAS 234783

PP694094

Raorchestes rezakhani

Maulovibazar, Bangladesh

JnUZool-A0319

MN072374

Raorchestes rezakhani

Maulovibazar, Bangladesh

JnUZool-A0419

MN072375

Raorchestes rezakhani

Maulovibazar, Bangladesh

JnUZool-A0619

MN615901

Raorchestes rezakhani

Bangladesh

A0619

MW165454

Raorchestes shillongensis

Shilong, meghalaya, India

ZSIS-M1

MN519707

Raorchestes shillongensis

Malki forest, Shilong, meghalaya, India

R2

MG980283

Raorchestes shillongensis

Risa forest, Shilong, meghalaya, India

R1

MG980282

Raorchestes sp.

 Tam Dao, Vinh Phu, Vietnam

ROM30298

MN475869

Raorchestes yadongensis

Yadong, Xizang, China

YBU 21222

OP345440

Raorchestes yadongensis

Yadong, Xizang, China

YBU 21223

OP345441

Nasutixalus jerdonii

Meriema, Nagaland, India

SDBDU 2007.060

KU170003

 

 

Table 3. Acoustic data of studied species of Raorchestes from Meghalaya. India.

Species

Calling position from the ground (cm)

Call Properties

Call Duration (s) ±SD (min–max) CV

Inter-call interval (s) ±SD (min–max) CV

Number of notes per call ±SD (min–max) CV

Note duration(s) ±SD (min–max) CV

Call Repetition Rate(calls/min) ±SD (min–max) CV

Inter-note interval(s) ±SD (min–max) CV

 

Dominant Frequency(kHz)±SD (min–max) CV

Raorchestes garo

50–500

0.03±0.005 (0.02–0.04) 18.2

2.26±0.18 (1.86–2.49) 7.8

1.0±0.0 (1.0–1.0) 0

0.03±0.005 (0.02–0.04) 18.2

35.28±6.15 (25.32–45.66) 17.4

2.26±0.18 (1.86–2.49) 7.8

2.93±0.04 (2.91–3) 1.35

Raorchestes asakgrensis

50–900

0.24±0.02 (0.21–0.28) 10.0

0.62±0.09 (0.5–0.8) 15.3

4.0±0.0 (4.0–4.0) 0

0.02±0.003 (0.02–0.03) 16.7

4.17±0.40 (3.6–4.9) 9.60

0.03±0.003 (0.03–0.04) 10.3

3.39±0.04(3.36–3.45) 0

Raorchestes kempiae

20–400

0.37±0.05 (0.30–0.43) 12.4

1.32±0.11 (1.16–1.46) 8.5

2.0±0.0 (2.0–2.0) 0

0.12±0.008 (0.11–0.13) 6.9

2.73±0.33 (2.32–3.31) 12.3

0.06±0.007 (0.04–0.07) 11.8

2.73±0.03(2.72–2.81) 1.09

Raorchestes jakoid sp. nov.

100–350

1.37±0.93 (0.55–2.81) 68.0

1.17±0.19 (0.94–1.51) 15.8

17.0±3.31 (5.0–35.0) 78.3

0.03±0.003 (0.02–0.03) 10.6

1.07±0.64 (0.36–1.81) 59.5

0.04±0.005 (0.03–0.04) 12.4

3.19±0.0

Raorchestes jadoh sp. nov.

20–150 -

0.93±0.22 (0.74–1.13) 23.6

6.17±0.54 (5.36–6.49) 8.8

3.5±0.58 (3.0–4.0) 16.5

0.02±0.003 (0.02–0.03) 12.9

1.12±0.27 (0.89–1.35) 23.6

0.30±0.01 (0.29–0.31) 13.3

3.66±1.32(3.56–3.84) 3.63

 

 

Table 4. A comparative table of member species of Raorchestes parvulus species group and species of bush frogs found in northeastern India.

Character

R. dulongensis

R. meglaensis

R. longchuanensis

R. parvulus

R. malipoensis

R. rezakhani

R. hillisi

R. huanglianshan

R. shillongensis

R. annandali

R. andersoni

R. garo

SVL of male (mm)

15.0–19.0

16.6–21.6

21.4–23.9

NA

14.6-17.7

18.85–20.90

15.9–17.7

17.0–19.6

10–20

16.0

24.0

22.47–23.8

Tympanum

Distinct

Indistinct

Distinct

Hidden

Distinct

Indistinct

Distinct

Distinct

Indistinct

Just distinguish-able

Distinct

Distinct or hidden

Supratympanic fold

Distinct

NA

NA

NA

Distinct

Weakly distinct

Distinct

Distinct

Distinct

NA

NA

Distinct

Nuptial pad

Absent

Present, white

Present

Absent

Present

Absent

Present

Present

NA

NA

NA

Present

Inner palmer tubercle

Present

Present

Present

Present

Absent

Absent

Indistinct

NA

NA

Present

Present

Absent

Outer palmar tubercle

Present

Present

Present

Present

Absent

Absent

Indistinct

NA

NA

Absent

Absent

Present

Inner metatarsal tubercle

Present

Present

Present

Present

Present, round

Absent

Present, Oval

 

Absent 

Absent

Present

Present

Outer metatarsal tubercle

Absent

Present

Absent

Absent

Absent

Absent

Absent

NA

Absent 

Absent

Absent

Absent

Toe web

Rudimentary webbing between toes

Toes are partially webbed (II 1 – 2 III 1 – 21/2 IV

21/2 – 1 V)

Toes are partially webbed

Webbed at the base

Rudimentary (I 2 – 2 II 2 – 2 III 2 – 3 IV 3 – 2 V)

Moderate webbing (I2 – 2 II 1¾ – 2 III 1½

– 3 IV 2¾ – 2 V)

No web between
toe I and toe II, web rudimen-tary between other toes (II 1 – 2 III 1–21/2 IV

21/2 – 1 V)

Rudimentary web on toes (II 1 – 2 III 1 – 2 – IV

2 – 1 V)

Indistinct rudiment of web present between the fourth and fifth toe

Webbed at the base

Feebly webbed

Webbed at the base (I 1−1 II 1−1 III 3−3 IV 3−2.5 V)

Relative toe length

I < II < V < III < IV

I<II<III ≈ V<IV, or I<II<III<V<IV

I<II<III ≈ V<IV

I<II< III <V<IV

I<II<V<III<IV

I < II < V < III < IV

I < II < III < V < IV

I < II < III < V < IV

I < II < III < V < IV

I < II < III < V < IV

I < II < III = V < IV

I < II < III < V < IV

Tibiotarsal articulation

Reaches anterior of the eye.

NA

NA

Reaches posterior border of the eye.

Reaches anterior border of the eye.

NA

Reaches the eye

Reaches the eye

Reaches the tympanic region

Reaches the eye

Reaches the eye

Reaches the posterior and above the eye

Dorsal surface

Distinct “) (”-shaped dark marking

Rough with small warts, dark butterfly-shaped spot on the back

Scattered small warts and "X" shaped markings

Curved dark band on each side of the back

X-shaped marking on the dorsal surface

“)-(“ or “)(“ shaped blackish mark on the dorsal surface

An indistinct “) (” shaped dark brown blotch on fore part of back.

An indistinct “X” shaped brown blotch on back.

"V" shaped broad light band on the back

Dorsal surface is smooth

Scattered tubercles on the dorsal surface

Dorsal surface is covered with hourglass-shaped
blotch covering the head from between the eyes and the back

 

Table 4 cont. A comparative table of member species of Raorchestes parvulus species group and species of bush frogs found in northeastern India.

Character

P. kempiae

R. hekouensis

R. yadongensis

R. asakgrensis

R. mindat

R. leiktho

R. jakoid sp. nov.

R. jadoh sp. nov.

SVL of male (mm)

22.96 –24.4

16.1–17.5

17.8–24.1

18.0–23.35

16.75–18.36

15.72–15.80

18.47–21.6

13.68– 14.01

Tympanum

Hidden

Distinct

Distinct

Indistinct

Indistinct

Indistinct

Indistinct

Indistinct

Supratympanic fold

Distinct

NA

NA

Distinct

Distinct

Indistinct

Distinct

Distinct

Nuptial pad

Present

Present

Present

Present

Present

Present

Present, whitish

Present

Inner palmer tubercle

Absent

Present

NA

Absent

Present

Present

Absent

Absent

Outer palmar tubercle

Present

Indistinct

NA

Present

Present

Present

Present

Present

Inner metatarsal tubercle

Present

Present, oval

Present

Present, oval

Present, rounded

Present, oval

Present, Oval

Present, round

Outer metatarsal tubercle

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Absent

Toe web

Webbed at the base ( I 1-2 II 2-1 III 1-3.51 IV 31- 2 V)

Rudimentary webbing on toes

Rudimentary webbing

Rudimentary webbing (I 1−2 II 2−2 III 2−3.5 IV 3.33−2 V)

Webbing between toes 1 and 2 vestigial, other rudimentary (II 2−3.5 III 2+−3.5 IV 3.25−2 V)

Webbing between toes 1 and 2 absent, other rudimentary (III 2−3.5 IV 3.33−2 V)

Rudimentary webbing (I 1−1 II 1−2.5 III 2−23 IV 23−1 V)

Rudimentary webbing (I 1−1 II 1−1.5 III 1−3.5 IV 3−1 V)

Relative toe length

I < II < III < V < IV

I < II < III < V < IV

I< II < III < V < IV

I < II < III < V < IV

I < II < III < V < IV

I < II < III < V < IV

I < II < III < V < IV

I < II < III < V < IV

Tibiotarsal articulation

Reaches the eye or the snout

Reaches the anterior  of an eye

Reaches the tip of snout.

Reaches anterior border of an eye

Reaches level of posterior margin of the eye

Reaches level of mid eye

Reaches posterior border of the eye.

Reaches posterior border of the eye.

Dorsal surface

Dorsal surface rough with granules with or without hour glass shaped marking

Distinct X-shaped dark brown marking on back

 

Upper parts with small wartsand with or without ”)(“ shaped marking. 

Body surface with X-shaped markings

 

Body surface slightly rough with very small warts and has”)(“ shaped marking

Upper part with small warts and six fairly distinct warts on the body towards the posterior of the eyes. Body also has ”)(“ shaped marking

 

 

Supplement Table 1. Uncorrected pdistance (16s rRNA) amongst the members of Raorchestes parvulus species group (Preceding the species name is the GenBank accession number).

Click here

 

FOR FIGURES & HTML - - CLICK HERE FOR FULL PDF

 

 

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