Research Article |
Corresponding author: Evan S. H. Quah ( evanquah@yahoo.com ) Academic editor: Raffael Ernst
© 2021 Evan S. H. Quah, L. Lee Grismer, Perry L. Wood Jr., Kelvin K. P. Lim, Paul Y. Imbun, M. S. Shahrul Anuar.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Quah ESH, Grismer LL, Wood Jr. PL, Lim KKP, Imbun PY, Anuar MSS (2021) An investigation into the taxonomy of Abavorana luctuosa (Peters, 1871) (Anura, Ranidae) and the resurrection of Rana decorata Mocquard, 1890 from Borneo. Vertebrate Zoology 71: 75-99. https://doi.org/10.3897/vz.71.e60921
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The taxonomic status of the ranid frog Abavorana luctuosa (Peters, 1871) was investigated using a combination of molecular and morphological data. The analyses revealed that A. luctuosa sensu lato is composed of two species in Borneo. One of these species agrees with the description of Rana decorata Mocquard, 1890 which is resurrected in the combination Abavorana decorata comb. nov. (Mocquard, 1890). Abavorana decorata is recovered as the sister lineage to the remainder of Abavorana and differs by a 16.0–17.0 % uncorrected pairwise sequence divergence from its congeners A. nazgul and A. luctuosa, respectively. It is distinguishable morphologically from A. luctuosa and A. nazgul by its ventral pattern (bold, black and white reticulations on its venter along with bold banding on the underside of hind limbs vs. generally immaculate and spotted in the latter two species), and a prominent white streak beneath the eye and/or tympanum extending to the corner of the jaw. Abavorana decorata further differs from A. luctuosa by having a significantly wider head and snout, larger interorbital and tympanum diameters, longer femur in both sexes, and various combinations of other mensural characters. Both species are sympatric in Borneo and this discovery adds to a growing number of widespread Sundaic species shown to be species complexes with distinct forms in Borneo.
amphibia, biodiversity, conservation, endemic, herpetofauna, phylogeny, Sundaland, systematics
The Mahogany frog (Abavorana luctuosa) is a colourful riparian species heard at night calling from tangled thickets along the edges of small streams and rivers as well as standing bodies of water such as ponds, pig wallows and pools in intermittent streams on the Thai-Malay Peninsula and Borneo. Abavorana luctuosa was described by Peters in 1871 from an unspecified location in Sarawak, East Malaysian Borneo (
Building upon these preliminary findings, we hypothesize that these populations are distinct evolutionary lineages and test this hypothesis with increased sample sizes from Borneo, Thailand, Peninsular Malaysia and detailed morphological, colour pattern, and genetic analyses based on the mitochondrial genes 16S, ND1 and flanking tRNAs (tRNA-Leu, tRNA-Ile, and tRNA-Gln). Given the results from the preponderance of data analyses herein, we rescind Boulenger’s synonymization of Rana decorata Mocquard, 1890 with Abavorana luctuosa and resurrect the species in the combination Abavorana decorata comb. nov. (Mocquard, 1890).
Nine additional specimens of Abavorana luctuosa sensu lato from Borneo were sequenced (Table
Taxon | Locality | Voucher | GenBank no. | Reference |
---|---|---|---|---|
Outgroups | ||||
Meristogenys stenocephalus | Borneo, Sabah, Crocker Range National Park | UMS:BORNEENSIS 8684 | AB526612.1 |
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Pulchrana sundabarat | West Malaysia, Kelantan, Pergau |
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KY982540 |
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Ingroup | ||||
Abavorana decorata | Borneo, Sabah, Lahad Datu district, Danum Valley |
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KY982535 |
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Abavorana nazgul | West Malaysia, Kedah, Gunung Jerai |
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KY982537 |
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Abavorana nazgul | West Malaysia, Kedah, Gunung Jerai |
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KY982538 |
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Abavorana nazgul | West Malaysia, Kedah, Gunung Jerai |
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KY982539 |
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Abavorana luctuosa | Borneo, Sabah, Sipitang district, Mendolong |
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KY982536 |
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Abavorana luctuosa | Borneo, Sabah, Sipitang district, Mendolong |
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MW478376 | This study |
Abavorana luctuosa | Borneo, Sabah, Sipitang district, Mendolong |
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MW478379 | This study |
Abavorana luctuosa | Borneo, Sarawak, Bintulu division, Tubau Camp |
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MW478378 | This study |
Abavorana luctuosa | Borneo, Sarawak, Bintulu division, Tubau Camp |
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MW478383 | This study |
Abavorana luctuosa | Borneo, Sarawak, Bintulu division, Tubau Camp |
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MW478380 | This study |
Abavorana luctuosa | Borneo, Sarawak, Bintulu division, Bukit Kana |
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MW478384 | This study |
Abavorana luctuosa | Borneo, Sarawak, Bintulu division, Bukit Kana |
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MW478377 | This study |
Abavorana luctuosa | Borneo, Sarawak, Bintulu division, Bukit Kana |
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MW478381 | This study |
Abavorana luctuosa | Borneo, Sarawak, Bintulu division, Samarakan |
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MW478382 | This study |
Abavorana luctuosa | West Malaysia, Kelantan, Gunung Stong |
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KY982519 |
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Abavorana luctuosa | West Malaysia, Kedah, Sungai Sedim |
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KY982520 |
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Abavorana luctuosa | West Malaysia, Penang, Penang Hill |
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KY982521 |
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Abavorana luctuosa | West Malaysia, Penang, Penang Hill |
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KY982522 |
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Abavorana luctuosa | West Malaysia, Penang, Penang Hill |
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KY982523 |
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Abavorana luctuosa | West Malaysia, Penang, Penang Hill |
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KY982524 |
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Abavorana luctuosa | West Malaysia, Selangor, Gombak |
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KY982525 |
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Abavorana luctuosa | West Malaysia, Selangor, Gombak |
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KY982526 |
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Abavorana luctuosa | West Malaysia, Pahang, Fraser’s Hill |
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KY982527 |
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Abavorana luctuosa | West Malaysia, Pahang, Fraser’s Hill |
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KY982528 |
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Abavorana luctuosa | West Malaysia, Pahang, Fraser’s Hill |
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KY982529 |
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Abavorana luctuosa | West Malaysia, Pahang, Genting Highlands |
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KY982530 |
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Abavorana luctuosa | West Malaysia, Pahang, Genting Highlands |
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KY982531 |
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Abavorana luctuosa | West Malaysia, Pahang, Genting Highlands |
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KY982532 |
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Abavorana luctuosa | West Malaysia, Pahang, Genting Highlands |
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KY982533 |
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Abavorana luctuosa | West Malaysia, Pahang, Genting Highlands |
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KY982534 |
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The new sequences were added to the molecular phylogenetic dataset of
Maximum likelihood (ML) and Bayesian Inference (BI) were used to generate phylogenetic trees. The data were partitioned into four partitions (ND1 by codon positions [1–3] and 16S+tRNAs) and best-fit models of evolution were determined in the IQ-TREE webserver version (
Uncorrected p-distances of the genus Abavorana. Mean (min–max) uncorrected p-distances (%) within examined populations of the genus Abavorana Oliver, Prendini, Kraus & Raxworthy 2015 computed in MEGA v6.06 (
n | A. decorata | A. nazgul | A. luctuosa (Pen. Malaysia) | A. luctuosa (Borneo) | |
A. decorata | 1 | — | — | — | — |
A. nazgul | 3 | 0.17 | 0.00 | — | — |
A. luctuosa (Pen. Malaysia) | 16 | 0.16 | 0.08 | 0.00 (0.00–0.02) | — |
A. luctuosa (Borneo) | 10 | 0.16 (0.16–0.17) | 0.08 (0.07–0.08) | 0.02 (0.02–0.03) | 0.01 (0.00–0.03) |
Morphological data from 62 adult specimens (34 males; 28 females) of Abavorana luctuosa sensu lato from 21 populations throughout the Thai-Malay Peninsula and Borneo (Appendix 1), and eight specimens of A. nazgul (seven males; one female) from the type locality in Peninsular Malaysia, were taken following the methodology of
Measurements of Abavorana decorata, A. nazgul and different populations of A. luctuosa. Abbreviations are listed in the Materials and methods.
Catalog no. | Species | SVL | HL | HW | SL | SW | IOD | IND | ED | TD | BL | FAL | ML | FL | CL | TL | PL | HG |
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A. decorata | 58.42 | 23.71 | 20.87 | 8.52 | 10.87 | 6.60 | 6.67 | 5.72 | 5.27 | 9.00 | 11.68 | 13.12 | 30.42 | 30.93 | 13.4 | 27.92 | ♀ |
SP02100 | A. decorata | 29.66 | 12.23 | 10.62 | 4.69 | 6.25 | 3.91 | 3.68 | 4.04 | 2.27 | 6.81 | 5.79 | 8.36 | 17.19 | 17.28 | 8.52 | 14.72 | ♀ |
SP02810 | A. decorata | 57.67 | 22.05 | 20.03 | 9.46 | 10.98 | 6.78 | 6.72 | 6.17 | 5.36 | 10.35 | 13.94 | 14.95 | 29.74 | 32.00 | 15.84 | 30.45 | ♀ |
SP20695 | A. decorata | 57.92 | 22.76 | 21.07 | 9.31 | 11.68 | 6.05 | 6.77 | 6.58 | 5.63 | 10.27 | 12.71 | 14.79 | 31.34 | 32.50 | 16.00 | 31.49 | ♀ |
SP26873 | A. decorata | 46.21 | 19.14 | 17.66 | 7.76 | 9.91 | 5.95 | 6.16 | 5.56 | 4.54 | 8.51 | 10.77 | 12.2 | 25.58 | 24.74 | 13.13 | 23.44 | 3.25 |
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A. decorata | 55.00 | 19.05 | 18.57 | 7.99 | 10.36 | 6.12 | 6.21 | 5.42 | 4.46 | 9.53 | 11.47 | 14.00 | 25.81 | 29.83 | 14.98 | 27.50 | ♀ |
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A. decorata | 56.00 | 19.05 | 19.53 | 8.22 | 10.71 | 5.39 | 6.09 | 5.93 | 4.6 | 10.06 | 11.18 | 15.00 | 26.1 | 29.33 | 14.98 | 28.00 | 3.89 |
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A. decorata | 33.69 | 13.9 | 11.92 | 6.42 | 7.07 | 4.29 | 4.48 | 3.94 | 2.78 | 6.89 | 6.59 | 9.33 | 17.25 | 17.74 | 9.19 | 17.12 | ♀ |
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A. nazgul | 43.99 | 17.14 | 15.83 | 7.44 | 8.70 | 4.48 | 5.54 | 3.86 | 3.80 | 8.87 | 8.13 | 11.00 | 23.11 | 24.01 | 11.84 | 24.08 | 2.51 |
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A. nazgul | 41.56 | 16.56 | 14.88 | 7.10 | 8.04 | 4.47 | 5.76 | 4.15 | 3.52 | 8.34 | 8.07 | 11.08 | 23.08 | 23.77 | 11.70 | 23.40 | 2.76 |
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A. nazgul | 44.10 | 16.68 | 14.72 | 7.16 | 8.02 | 3.99 | 5.45 | 4.06 | 3.61 | 8.76 | 8.29 | 11.05 | 22.56 | 23.85 | 11.30 | 22.76 | 2.41 |
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A. nazgul | 41.51 | 16.50 | 14.63 | 7.13 | 8.11 | 4.33 | 5.80 | 3.93 | 3.60 | 8.49 | 7.54 | 10.37 | 22.31 | 22.83 | 10.70 | 22.1 | 2.4 |
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A. nazgul | 42.70 | 16.50 | 15.03 | 7.25 | 8.39 | 4.12 | 5.73 | 4.24 | 3.58 | 8.48 | 7.84 | 10.90 | 22.20 | 23.52 | 11.14 | 23.18 | 2.47 |
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A. nazgul | 52.51 | 19.65 | 18.57 | 8.69 | 9.59 | 4.94 | 6.61 | 4.80 | 4.10 | 12.02 | 10.17 | 13.41 | 27.80 | 28.77 | 14.42 | 28.84 | ♀ |
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A. nazgul | 43.92 | 16.36 | 15.60 | 7.54 | 8.60 | 4.05 | 5.56 | 4.36 | 4.01 | 8.98 | 8.23 | 11.53 | 23.22 | 23.93 | 11.31 | 24.46 | 2.55 |
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A. nazgul | 41.61 | 16.52 | 15.41 | 7.57 | 7.98 | 3.94 | 5.67 | 4.56 | 3.91 | 8.32 | 8.39 | 11.25 | 22.43 | 23.26 | 10.95 | 23.22 | 2.98 |
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A. luctuosa Malay Peninsula | 44.39 | 15.20 | 13.83 | 6.48 | 8.75 | 4.81 | 6.41 | 5.05 | 3.24 | 9.49 | 9.23 | 11.24 | 21.92 | 24.69 | 12.39 | 22.52 | ♀ |
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A. luctuosa Malay Peninsula | 46.12 | 16.84 | 14.66 | 7.80 | 9.19 | 4.12 | 5.42 | 5.04 | 3.90 | 9.91 | 9.20 | 12.42 | 23.95 | 24.94 | 12.44 | 25.38 | ♀ |
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A. luctuosa Malay Peninsula | 44.17 | 16.69 | 15.58 | 7.56 | 8.62 | 4.59 | 5.44 | 4.60 | 3.32 | 7.99 | 7.90 | 10.58 | 19.82 | 21.59 | 11.61 | 21.72 | 3.40 |
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A. luctuosa Malay Peninsula | 41.27 | 16.37 | 14.50 | 6.99 | 8.52 | 4.33 | 5.27 | 4.10 | 3.54 | 7.80 | 7.66 | 10.25 | 21.27 | 22.03 | 11.02 | 22.03 | 3.36 |
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A. luctuosa Malay Peninsula | 37.20 | 14.70 | 13.39 | 6.75 | 8.21 | 4.03 | 5.08 | 3.80 | 3.41 | 8.03 | 8.14 | 10.46 | 19.80 | 22.14 | 10.92 | 20.50 | 3.89 |
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A. luctuosa Malay Peninsula | 48.7 | 19.59 | 15.98 | 8.29 | 9.38 | 4.93 | 6.72 | 4.75 | 3.82 | 9.11 | 9.64 | 13.72 | 25.14 | 27.29 | 14.30 | 27.32 | ♀ |
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A. luctuosa Malay Peninsula | 49.75 | 19.35 | 16.84 | 8.07 | 8.37 | 5.20 | 6.04 | 5.21 | 3.96 | 7.67 | 9.38 | 12.50 | 24.52 | 26.48 | 14.56 | 26.30 | ♀ |
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A. luctuosa Malay Peninsula | 43.89 | 17.96 | 15.12 | 7.47 | 8.29 | 3.80 | 5.75 | 4.38 | 3.58 | 6.69 | 8.87 | 11.23 | 21.55 | 25.72 | 12.74 | 23.5 | ♀ |
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A. luctuosa Malay Peninsula | 51.06 | 19.15 | 16.17 | 8.24 | 9.25 | 5.49 | 6.53 | 5.19 | 4.24 | 9.37 | 11.39 | 12.55 | 26.37 | 28.75 | 14.52 | 27.86 | ♀ |
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A. luctuosa Malay Peninsula | 45.71 | 17.09 | 16.32 | 7.64 | 9.04 | 4.92 | 5.90 | 4.40 | 4.14 | 7.80 | 9.84 | 10.01 | 21.91 | 23.88 | 12.36 | 22.55 | ♀ |
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A. luctuosa Malay Peninsula | 43.06 | 15.85 | 14.58 | 6.95 | 8.18 | 5.14 | 5.70 | 4.20 | 3.66 | 7.08 | 8.55 | 9.83 | 20.78 | 23.49 | 12.26 | 22.55 | 4.41 |
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A. luctuosa Malay Peninsula | 49.50 | 18.05 | 16.76 | 8.30 | 8.97 | 5.64 | 5.87 | 4.97 | 3.93 | 8.48 | 11.07 | 11.60 | 23.72 | 26.39 | 13.14 | 25.46 | ♀ |
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A. luctuosa Malay Peninsula | 49.13 | 18.43 | 16.57 | 8.14 | 8.99 | 5.13 | 6.17 | 5.01 | 4.07 | 8.70 | 10.00 | 12.29 | 25.00 | 27.03 | 13.49 | 26.05 | ♀ |
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A. luctuosa Malay Peninsula | 46.08 | 16.08 | 14.74 | 7.37 | 8.21 | 5.15 | 5.73 | 4.65 | 3.79 | 8.57 | 8.96 | 10.92 | 23.67 | 24.95 | 12.25 | 25.15 | ♀ |
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A. luctuosa Malay Peninsula | 44.27 | 16.41 | 14.45 | 7.17 | 8.16 | 4.38 | 5.48 | 4.07 | 3.59 | 7.43 | 7.73 | 11.13 | 22.04 | 23.73 | 12.35 | 23.11 | 3.80 |
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A. luctuosa Malay Peninsula | 46.89 | 17.23 | 16.48 | 8.19 | 8.82 | 5.06 | 6.13 | 4.91 | 3.87 | 8.6 | 9.10 | 12.75 | 24.4 | 25.18 | 12.79 | 25.14 | ♀ |
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A. luctuosa Malay Peninsula | 40.56 | 16.26 | 14.34 | 7.43 | 8.12 | 4.59 | 5.14 | 4.70 | 3.41 | 7.37 | 8.81 | 10.69 | 21.91 | 22.36 | 11.30 | 22.00 | 2.58 |
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A. luctuosa Malay Peninsula | 40.17 | 14.45 | 13.78 | 6.62 | 7.58 | 3.25 | 5.30 | 4.15 | 2.62 | 8.85 | 8.27 | 10.34 | 18.62 | 21.43 | 10.72 | 20.83 | 2.81 |
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A. luctuosa Malay Peninsula | 38.09 | 14.96 | 14.51 | 6.43 | 8.04 | 4.11 | 5.07 | 4.31 | 3.24 | 8.84 | 8.30 | 10.54 | 20.54 | 21.28 | 10.66 | 19.62 | 3.50 |
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A. luctuosa Malay Peninsula | 40.77 | 15.75 | 13.94 | 6.70 | 8.26 | 4.21 | 5.88 | 4.38 | 3.12 | 8.14 | 8.01 | 9.65 | 19.58 | 20.75 | 10.00 | 20.58 | 3.09 |
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A. luctuosa Malay Peninsula | 37.83 | 14.94 | 13.35 | 6.20 | 7.11 | 4.89 | 4.93 | 4.17 | 3.05 | 8.34 | 7.55 | 9.68 | 18.38 | 21.03 | 10.31 | 20.59 | 3.09 |
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A. luctuosa Borneo | 37.78 | 15.92 | 13.49 | 6.33 | 7.52 | 3.80 | 4.66 | 4.09 | 3.00 | 5.78 | 7.98 | 8.69 | 17.82 | 19.70 | 10.69 | 18.40 | 3.87 |
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A. luctuosa Borneo | 43.58 | 17.60 | 14.96 | 7.30 | 7.71 | 4.70 | 5.10 | 4.33 | 3.77 | 7.38 | 9.55 | 10.33 | 20.10 | 23.72 | 12.82 | 21.97 | ♀ |
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A. luctuosa Borneo | 42.71 | 16.51 | 14.67 | 6.97 | 8.06 | 4.90 | 4.75 | 4.82 | 3.38 | 8.14 | 9.32 | 10.10 | 21.44 | 22.63 | 11.69 | 21.16 | ♀ |
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A. luctuosa Borneo | 37.43 | 15.75 | 13.69 | 6.54 | 7.83 | 4.56 | 5.03 | 4.00 | 2.81 | 6.93 | 7.83 | 8.69 | 18.99 | 20.83 | 11.05 | 19.96 | 4.05 |
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A. luctuosa Borneo | 45.19 | 16.26 | 15.74 | 6.99 | 9.02 | 5.60 | 5.99 | 4.79 | 3.63 | 8.47 | 9.89 | 11.29 | 23.09 | 24.98 | 13.48 | 23.73 | ♀ |
SP02676 | A. luctuosa Borneo | 39.83 | 15.78 | 13.73 | 6.64 | 7.94 | 4.63 | 5.32 | 4.50 | 3.61 | 6.68 | 8.92 | 9.94 | 21.31 | 21.94 | 11.22 | 19.69 | 3.65 |
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A. luctuosa Borneo | 39.51 | 15.26 | 13.58 | 6.90 | 7.95 | 4.70 | 5.58 | 4.59 | 3.40 | 7.86 | 8.01 | 10.59 | 21.25 | 22.06 | 11.75 | 20.88 | 3.96 |
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A. luctuosa Borneo | 37.22 | 13.53 | 13.27 | 6.09 | 7.59 | 3.83 | 4.76 | 3.65 | 3.35 | 7.20 | 7.50 | 9.20 | 19.04 | 19.53 | 9.86 | 19.01 | 3.06 |
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A. luctuosa Borneo | 37.05 | 14.84 | 12.71 | 6.61 | 7.65 | 4.15 | 5.01 | 4.08 | 2.95 | 8.87 | 8.50 | 9.22 | 19.13 | 21.63 | 11.29 | 21.29 | ♀ |
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A. luctuosa Borneo | 38.86 | 15.15 | 13.64 | 6.67 | 7.69 | 3.78 | 5.26 | 4.33 | 3.11 | 8.56 | 7.22 | 10.10 | 19.81 | 20.72 | 10.28 | 20.32 | 3.92 |
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A. luctuosa Borneo | 36.30 | 14.16 | 13.17 | 5.66 | 7.33 | 4.13 | 4.89 | 3.98 | 2.50 | 7.37 | 7.25 | 9.49 | 18.31 | 20.11 | 10.19 | 19.53 | 3.12 |
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A. luctuosa Borneo | 35.41 | 14.72 | 12.92 | 6.35 | 7.77 | 4.22 | 5.04 | 4.18 | 2.75 | 7.38 | 7.77 | 8.73 | 19.13 | 20.29 | 10.71 | 19.21 | 3.95 |
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A. luctuosa Borneo | 35.05 | 14.25 | 12.28 | 6.39 | 7.47 | 4.10 | 5.41 | 4.20 | 3.26 | 8.65 | 7.36 | 8.79 | 18.53 | 20.82 | 10.39 | 20.55 | 2.50 |
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A. luctuosa Borneo | 36.78 | 14.54 | 12.25 | 6.26 | 7.57 | 4.10 | 5.01 | 4.30 | 3.05 | 8.92 | 7.23 | 9.03 | 17.81 | 21.22 | 10.84 | 20.09 | 2.22 |
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A. luctuosa Borneo | 38.19 | 14.44 | 13.39 | 6.01 | 7.57 | 4.00 | 4.90 | 4.40 | 3.34 | 8.53 | 7.51 | 9.36 | 19.43 | 20.91 | 11.27 | 20.50 | 3.85 |
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A. luctuosa Borneo | 38.22 | 15.27 | 12.90 | 6.50 | 7.65 | 4.59 | 5.05 | 4.12 | 3.07 | 8.96 | 7.13 | 9.47 | 19.32 | 20.93 | 10.86 | 21.28 | 3.79 |
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A. luctuosa Borneo | 39.74 | 15.98 | 13.93 | 6.75 | 7.84 | 4.62 | 5.44 | 4.54 | 3.04 | 9.06 | 8.13 | 9.55 | 20.63 | 21.39 | 11.36 | 20.87 | 3.58 |
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A. luctuosa Borneo | 37.27 | 15.51 | 13.25 | 6.96 | 7.90 | 4.53 | 5.50 | 3.79 | 3.06 | 8.42 | 7.42 | 9.13 | 19.86 | 21.5 | 11.55 | 20.93 | 3.22 |
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A. luctuosa Borneo | 38.37 | 15.04 | 12.80 | 6.62 | 8.22 | 4.41 | 5.28 | 4.05 | 3.00 | 8.94 | 7.27 | 9.43 | 20.12 | 21.23 | 11.27 | 21.82 | 4.16 |
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A. luctuosa Borneo | 39.01 | 15.74 | 13.65 | 6.63 | 8.02 | 4.19 | 5.38 | 3.95 | 3.16 | 9.05 | 7.49 | 9.03 | 20.98 | 21.36 | 10.97 | 20.96 | 3.33 |
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A. luctuosa Borneo | 45.01 | 17.49 | 15.52 | 7.40 | 8.79 | 4.86 | 5.80 | 5.22 | 3.67 | 10.2 | 9.92 | 10.61 | 23.68 | 24.27 | 13.11 | 22.82 | ♀ |
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A. luctuosa Borneo | 45.2 | 17.45 | 14.85 | 7.00 | 8.72 | 4.72 | 5.77 | 4.81 | 3.41 | 9.95 | 8.82 | 10.91 | 22.66 | 24.64 | 12.83 | 23.25 | ♀ |
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A. luctuosa Borneo | 43.75 | 17.06 | 14.51 | 7.34 | 8.71 | 4.65 | 5.74 | 4.54 | 3.47 | 10.13 | 9.01 | 11.08 | 22.17 | 22.99 | 12.71 | 23.01 | ♀ |
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A. luctuosa Borneo | 45.11 | 17.57 | 15.14 | 7.13 | 8.81 | 4.55 | 5.98 | 4.74 | 3.64 | 10.29 | 8.25 | 11.29 | 22.55 | 25.02 | 12.69 | 23.69 | ♀ |
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A. luctuosa Borneo | 42.18 | 17.72 | 13.92 | 6.93 | 7.75 | 4.58 | 5.31 | 4.75 | 3.18 | 8.11 | 9.11 | 10.65 | 21.05 | 19.83 | 11.71 | 22.00 | ♀ |
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A. luctuosa Borneo | 38.46 | 15.62 | 13.41 | 6.98 | 7.40 | 4.28 | 5.20 | 3.73 | 2.88 | 6.85 | 8.01 | 11.00 | 19.78 | 20.99 | 11.22 | 20.34 | 2.18 |
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A. luctuosa Borneo | 39.18 | 16.08 | 13.97 | 6.69 | 8.00 | 4.14 | 5.33 | 4.39 | 3.24 | 9.37 | 8.52 | 10.39 | 21.57 | 21.42 | 11.18 | 21.71 | 2.29 |
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A. luctuosa Borneo | 39.56 | 16.78 | 13.44 | 7.03 | 7.72 | 3.80 | 5.23 | 4.23 | 3.83 | 9.56 | 7.56 | 10.40 | 20.58 | 20.68 | 10.40 | 19.45 | 2.67 |
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A. luctuosa Borneo | 39.32 | 15.06 | 13.72 | 6.76 | 7.66 | 4.59 | 4.95 | 4.35 | 3.40 | 7.68 | 8.03 | 10.36 | 20.48 | 21.24 | 10.48 | 20.44 | 2.46 |
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A. luctuosa Borneo | 44.93 | 15.81 | 15.09 | 7.86 | 8.04 | 4.62 | 5.37 | 4.81 | 3.18 | 8.81 | 8.25 | 11.78 | 21.66 | 23.15 | 12.30 | 23.32 | ♀ |
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A. luctuosa Borneo | 40.87 | 17.33 | 14.49 | 7.36 | 8.41 | 4.31 | 5.70 | 4.67 | 3.59 | 10.00 | 8.23 | 11.36 | 21.55 | 22.37 | 11.21 | 22.53 | ♀ |
|
A. luctuosa Borneo | 37.25 | 14.80 | 13.60 | 6.62 | 7.53 | 4.25 | 5.10 | 4.30 | 3.19 | 7.97 | 7.45 | 10.22 | 19.28 | 19.5 | 10.17 | 19.49 | 3.14 |
|
A. luctuosa Borneo | 40.55 | 16.12 | 14.41 | 6.79 | 7.90 | 3.95 | 5.13 | 4.28 | 3.38 | 9.45 | 8.22 | 10.79 | 19.9 | 20.95 | 10.8 | 20.82 | 2.73 |
Museum abbreviations are:
All analyses were performed using the platform R version 3.2.1 (
A MANOVA analysis was performed to test for significantly different (p < 0.05) multivariate structure in the mensural data and subsequently viewed in a principal component analysis (PCA). PCA and discriminant analysis of principal components (DAPC) were performed using the ADEGENET package in R (
Independent of the MANOVA, an analysis of variance (ANOVA) was conducted on mensural characters. Since the species and population sample sizes were unequal, a Levene’s test for homogeneity of variances among the mensural characters was conducted prior to an analysis of variance (ANOVA). A Welch’s ANOVA was conducted on characters with unequal variances (p ≥ 0.05) and a standard ANOVA was conducted on characters with equal variances (p ≤ 0.05) to test for the presence of statistically significant mean differences (p ≤ 0.05) between all combinations of species or population pairs in the data set. Characters with equal variances and bearing statistical differences were subjected to a TukeyHSD test to ascertain which population pairs differed significantly from each other for those characters (Table
Pairwise matrix of statistically significant characters among species of the genus Abavorana. Pairwise matrix of statistically significant mean differences of characters among Abavorana decorata, A. nazgul, and A. luctuosa populations from the Malay Peninsula and Borneo. Abbreviations are in the Materials and methods.
A. decorata | A. nazgul | A. luctuosa Malay Peninsula | A. luctuosa Borneo | ||||||
♂ | ♀ | ♂ | ♀ | ♂ | ♀ | ♂ | ♀ | ||
A. decorata | ♂ | — | — | — | — | — | — | — | — |
♀ | — | — | — | — | — | — | — | — | |
A. nazgul | ♂ | HL, HW, SL, IOD, ED, TD, FAL, TL, PL, SW, IND, ML, CL, FL, HG | — | — | — | — | — | — | — |
♀ | — | HW, IOD, IND, FL, PL, SL, BL | — | — | — | — | — | — | |
A. luctuosa Malay Peninsula | ♂ | HL, HW, SL, IOD, ED, TD, FAL, TL, PL, SW, IND, ML, FL, CL | — | HL, HW, SL, TD, PL, IND, ML, FL, CL, HG | — | — | — | — | — |
♀ | — | HW, SW, IOD, TD, FL | — | HL, HW, SW, ML, FL, CL, PL, SL, BL, TL | — | — | — | — | |
A. luctuosa Borneo | ♂ | HL, HW, SL, IOD, ED, TD, TL, PL, SW, IND, FAL, ML, CL, FL | — | SVL, HL, HW, SL, TD, PL, SW, IND, ML, FL, CL, HG | — | SVL, HW, SL, PL, ML, CL | — | — | — |
♀ | — | HW, HL, SW, IOD, TD, FAL, ML, FL, CL, PL, ED | — | HL, HW, SW, IND, TD, FAL, ML, FL, CL, PL, SL, BL, TL | — | HW, SW, IND, TD, FAL, ML, FL, CL, PL, SL, TL | — | — |
Summary statistics and ventral colour pattern among members of the genus Abavorana. Summary statistics and ventral colour pattern for Abavorana decorata, A. nazgul, and A. luctuosa populations from the Malay Peninsula and Borneo, SD = standard deviation, N = sample size, / = no data. Humeral gland only present in males and all measurements in mm.
A. decorata | A. nazgul | A. luctuosa Malay Peninsula | A. luctuosa Borneo | |||||
---|---|---|---|---|---|---|---|---|
Sex | Male | Female | Male | Female | Male | Female | Male | Female |
N | 2 | 6 | 7 | 1 | 10 | 11 | 22 | 11 |
Snout-vent length (SVL) | ||||||||
mean (±SD) | 51.11 (±6.92) | 48.73 (±13.32) | 42.77 (±1.22) | 52.51 | 40.74 (±2.54) | 47.38 (±2.36) | 38.15 (±1.45) | 43.23 (±2.51) |
range | 46.21–56.00 | 29.66–58.42 | 41.51–44.10 | / | 37.20–44.27 | 43.89–51.06 | 35.05–40.55 | 37.05–45.20 |
Head length (HL) | ||||||||
mean (±SD) | 19.10 (±0.06) | 18.95 (±4.85) | 16.61 (±0.25) | 19.65 | 15.64 (±0.81) | 17.72 (±1.39) | 15.25 (±0.78) | 16.88 (±0.92) |
range | 19.05–19.14 | 12.23–23.71 | 16.36–17.14 | / | 14.45–16.69 | 15.20–19.59 | 13.53–16.78 | 14.84–17.72 |
Head width (HW) | ||||||||
mean (±SD) | 18.60 (±1.32) | 17.18 (±4.68) | 15.16 (±0.46) | 18.57 | 14.24 (±0.66) | 15.77 (±1.01) | 13.39 (±0.52) | 14.69 (±0.83) |
range | 17.66–19.53 | 10.62–21.07 | 14.63–15.83 | / | 13.35–15.58 | 13.83–16.84 | 12.25–14.41 | 12.71–15.74 |
Snout length (SL) | ||||||||
mean (±SD) | 7.99 (±0.33) | 7.73 (±1.85) | 7.31 (±0.20) | 8.69 | 6.88 (±0.43) | 7.82 (±0.56) | 6.55 (±0.34) | 7.17 (±0.33) |
range | 7.76–8.22 | 4.69–9.46 | 7.10–7.57 | / | 6.20–7.56 | 6.48–8.30 | 5.66–7.03 | 6.61–7.86 |
Snout width (SW) | ||||||||
mean (±SD) | 10.31 (±0.57) | 9.54 (±2.28) | 8.26 (±0.30) | 9.59 | 8.08 (±0.44) | 8.84 (±0.40) | 7.73 (±0.23) | 8.33 (±0.51) |
range | 9.91–10.71 | 6.25–11.68 | 7.98–8.70 | / | 7.11–8.62 | 8.21–9.38 | 7.33–8.22 | 7.65–9.02 |
Inter orbital distance (IOD) | ||||||||
mean (±SD) | 5.67 (±0.40) | 5.63 (±1.22) | 4.20 (±0.23) | 4.94 | 4.35 (±0.52) | 4.93 (±0.54) | 4.24 (±0.30) | 4.69 (±0.37) |
range | 5.59–5.95 | 3.91–6.78 | 3.94–4.48 | / | 3.25–5.14 | 3.80–5.64 | 3.78–4.70 | 4.15–5.60 |
Internarial distance (IND) | ||||||||
mean (±SD) | 6.13 (±0.05) | 5.76 (±1.34) | 5.64 (±0.13) | 6.61 | 5.33 (±0.30) | 6.06 (±0.38) | 5.16 (±0.24) | 5.50 (±0.42) |
range | 6.09–6.16 | 3.68–6.77 | 5.45–5.80 | / | 4.93–5.88 | 5.42–6.72 | 4.66–5.58 | 4.75–5.99 |
Eye diameter (ED) | ||||||||
mean (±SD) | 5.75 (±0.26) | 5.31 (±1.10) | 4.17 (±0.24) | 4.80 | 4.25 (±0.26) | 4.87 (±0.29) | 4.18 (±0.26) | 4.69 (±0.29) |
range | 5.56–5.93 | 3.94–6.58 | 3.86–4.56 | / | 3.80–4.70 | 4.38–5.21 | 3.65–4.59 | 4.08–5.22 |
Tympanum diameter (TD) | ||||||||
mean (±SD) | 4.57 (±0.04) | 4.30 (±1.43) | 3.72 (±0.19) | 4.10 | 3.30 (±0.31) | 3.87 (±0.27) | 3.16 (±0.29) | 3.44 (±0.25) |
range | 4.54–4.60 | 2.27–5.63 | 3.52–4.01 | / | 2.62–3.66 | 3.24–4.24 | 2.50–3.83 | 2.95–3.77 |
Brachium length (BL) | ||||||||
mean (±SD) | 9.29 (±1.10) | 8.81 (±1.60) | 8.61 (±0.26) | 12.02 | 7.99 (±0.59) | 8.58 (±0.92) | 8.14 (±1.04) | 9.12 (±1.03) |
range | 8.51–10.06 | 6.81–10.35 | 8.32–8.98 | / | 7.08–8.85 | 6.69–9.91 | 5.78–9.56 | 7.38–10.29 |
Forearm length (FAL) | ||||||||
mean (±SD) | 10.98 (±0.29) | 10.36 (±3.36) | 8.07 (±0.29) | 10.17 | 8.09 (±0.40) | 9.70 (±0.84) | 7.72 (±0.47) | 8.99 (±0.64) |
range | 10.77–11.18 | 5.79–13.94 | 7.54–8.39 | / | 7.55–8.81 | 8.87–11.39 | 7.13–8.92 | 8.23–9.92 |
Manus length (ML) | ||||||||
mean (±SD) | 13.60 (±1.98) | 12.43 (±2.86) | 11.03 (±0.35) | 13.41 | 10.32 (±0.48) | 11.93 (±1.04) | 9.67 (±0.72) | 10.78 (±0.71) |
range | 12.20–15.00 | 8.36–14.95 | 10.37–11.53 | / | 9.65–11.13 | 10.01–13.72 | 8.69–11.0 | 9.22–11.78 |
Femur length (FL) | ||||||||
mean (±SD) | 25.84 (±0.37) | 25.29 (±6.53) | 22.70 (±0.42) | 27.80 | 20.27 (±1.26) | 23.83 (±1.51) | 19.72 (±1.08) | 21.73 (±1.32) |
range | 25.58–26.10 | 17.19–31.34 | 22.2–23.22 | / | 18.38–22.04 | 21.55–26.37 | 17.81–21.57 | 19.13–23.68 |
Crus length (CL) | ||||||||
mean (±SD) | 27.04 (±3.25) | 26.71 (±7.19) | 23.60 (±0.42) | 28.77 | 21.98 (±0.99) | 25.94 (±1.41) | 20.88 (±0.70) | 23.20 (±1.58) |
range | 24.74–29.33 | 17.28–32.50 | 22.83–41.01 | / | 20.75–23.73 | 23.88–28.75 | 19.50–22.06 | 19.83–25.02 |
Tarsal length (TL) | ||||||||
mean (±SD) | 14.06 (±1.31) | 12.99 (±3.34) | 11.28 (±0.40) | 14.42 | 11.12 (±0.78) | 13.18 (±0.90) | 10.84 (±0.50) | 12.35 (±0.76) |
range | 13.13–14.98 | 8.52–16.00 | 10.70–11.84 | / | 10.00–12.35 | 12.25–14.56 | 9.86–11.75 | 11.21–13.48 |
Pes length (PL) | ||||||||
mean (±SD) | 25.72 (±3.22) | 24.87 (±7.13) | 23.31 (±0.79) | 28.84 | 21.35 (±1.09) | 25.20 (±1.75) | 20.28 (±0.88) | 22.62 (±0.90) |
range | 23.44–28.00 | 14.72–31.49 | 22.10–24.46 | / | 19.62–23.11 | 22.52–27.86 | 18.40–21.82 | 21.16–23.73 |
Humeral gland (HG) | ||||||||
mean (±SD) | 3.57 (±0.45) | / | 2.58 (±0.21) | / | 3.39 (±0.54) | / | 3.26 (±0.66) | / |
range | 3.25–3.89 | / | 2.40–2.98 | / | 2.58–4.41 | / | 2.18–4.16 | / |
Pattern of venter | Belly usually mottled with black and white reticulations along the flanks; limbs with bold black and white bands or reticulations | Spotted | Immaculate to very faint, sparse speckling | Immaculate to very faint, sparse speckling |
The BI and ML analyses recovered trees with identical topologies. Specimen
The MANOVA indicated there were significant (Wilks’ lamda, p=2.2-16) differences in mensural data between both sexes of the two pattern morphs of Abavorana luctuosa sensu lato and A. nazgul, as visualized in the PCA and DAPC (Fig.
PCA and DAPC for the genus Abavorana; PCAs (left) with convex hull polygons and DAPCs (right) showing the morphospatial relationships of male (top) and female (bottom) Abavorana decorata, A. nazgul, and A. luctuosa populations from the Malay Peninsula and Borneo. 1 =
The analyses also revealed that both venter phenotypes also differ in a number of statistically significant mensural metrics (Table
Significantly wider heads in both sexes, with HW of adult males 17.66–19.53 mm, mean 18.60 ± SD 1.32, n = 2 (vs. 12.25–14.41 mm, mean 13.39 ± SD 0.52, n = 22, P < 0.001, in Bornean Abavorana luctuosa, and 13.35–15.58 mm, mean 14.24 ± SD 0.66, n = 10, P < 0.001, in Malay Peninsula A. luctuosa) and adult females 10.62–21.07 mm, mean 17.18 ± SD 4.68, n = 6 (vs. 12.71–15.74 mm, mean 14.69 ± SD 0.83, n = 11, P < 0.001, in Bornean A. luctuosa, and 13.83–16.84 mm, mean 15.77 ± SD 1.01, n = 11, P < 0.01, in Malay Peninsula A. luctuosa); significantly wider snouts in both sexes, with SW of adult males 9.91–10.71 mm, mean 10.31 ± SD 0.57, n = 2 (vs. 7.33–8.22 mm, mean 7.73 ± SD 0.23, n = 22, P < 0.001, in Bornean A. luctuosa, and 7.11–8.62 mm, mean 8.08 ± SD 0.44, n = 10, P < 0.001, in Malay Peninsula A. luctuosa) and adult females 6.25–11.68 mm, mean 9.54 ± SD 2.28, n = 6 (vs. 7.65–9.02 mm, mean 8.33 ± SD 0.51, n = 11, P < 0.001, in Bornean A. luctuosa, and 8.21–9.38 mm, mean 8.84 ± SD 0.40, n = 11, P < 0.05, in Malay Peninsula A. luctuosa); significantly wider interorbital diameter in both sexes, with IOD of adult males 5.59–5.95 mm, mean 5.67 ± SD 0.40, n = 2 (vs. 3.78–4.70 mm, mean 4.24 ± SD 0.30, n = 22, P < 0.001, in Bornean A. luctuosa, and 3.25–5.14 mm, mean 4.35 ± SD 0.52, n = 10, P < 0.001, in Malay Peninsula A. luctuosa) and adult females 3.91–6.78 mm, mean 5.63 ± SD 1.22, n = 6 (vs. 4.15–5.60 mm, mean 4.69 ± SD 0.37, n = 11, P < 0.001, in Bornean A. luctuosa, and 3.80–5.64 mm, mean 4.93 ± SD 0.54, n = 11, P < 0.001, in Malay Peninsula A. luctuosa); significantly wider tympanum diameter in both sexes, with TD of adult males 4.54–4.60 mm, mean 4.57 ± SD 0.04, n = 2 (vs. 2.50–3.83 mm, mean 3.16 ± SD 0.29, n = 22, P < 0.001, in Bornean A. luctuosa, and 2.62–3.66 mm, mean 3.30 ± SD 0.31, n = 10, P < 0.001, in Malay Peninsula A. luctuosa) and adult females 2.27–5.63 mm, mean 4.30 ± SD 1.43, n = 6 (vs. 2.95–3.77 mm, mean 3.44 ± SD 0.25, n = 11, P < 0.001, in Bornean A. luctuosa, and 3.24–4.24 mm, mean 3.87 ± SD 0.27, n = 11, P < 0.05, in Malay Peninsula A. luctuosa); and significantly longer femur length in both sexes, FL of adult males 25.58–26.10 mm, mean 25.84 ± SD 0.37, n = 2 (vs. 17.81–21.57 mm, mean 19.72 ± SD 1.08, n = 22, P < 0.001, in Bornean A. luctuosa, and 18.38–22.04 mm, mean 20.27 ± SD 1.26, n = 10, P < 0.001, in Malay Peninsula A. luctuosa) and adult females 17.19–31.34 mm, mean 25.29 ± SD 6.53, n = 6 (vs. 19.13–23.68 mm, mean 21.73 ± SD 1.32, n = 11, P < 0.001, in Bornean A. luctuosa, and 21.55–26.37 mm, mean 23.83 ± SD 1.51, n = 11, P < 0.05, in Malay Peninsula A. luctuosa). In addition to these, the phenotypes differed significantly from each other and A. nazgul by a suite of other mensural characters (Table
The analyses demonstrate unequivocally that Abavorana luctuosa on Borneo are composed of two genetically and morphologically distinct forms. Abavorana luctuosa described by
Bornean Abavorana luctuosa; Dorsal (A & C) and ventral (B & D) pattern of Abavorana luctuosa from Borneo. A & B: Top row (from left to right);
Syntypes of Abavorana decorata; A & D: Dorsum and venter of
Examined Abavorana decorata specimens; Dorsal (A & C) and ventral (B & D) pattern of Abavorana decorata from Borneo. A & B (from left to right):
Limnodytes luctuosus:
Rana luctuosa
(in part):
Rana (Limnodytes) luctuosa:
Rana (Hylorana) luctuosa
(in part):
Rana (Hylarana) luctuosa
(in part):
Hylorana luctuosa
(in part):
Rana (Pulchrana) luctuosa
(in part):
Pulchrana luctuosa
(in part):
Hylarana luctuosa
(in part):
Abavorana luctuosa:
Hylarana (Abavorana) luctuosa
(in part):
Adult female (
Body robust, medium-sized; head moderate; snout short, rounded, canthus rostralis smoothly rounded; interorbital space broader than the upper eyelid; tympanum distinct, not quite two-thirds the size of the eye with no pale colouration on the margins of the tympanum; no vocal sacs in males; vomerine teeth in two small oblique patches on a level with the posterior edge of the choanae; length of 1st finger greater than 2nd finger; disc width to finger width ratios of finger 3 and toe 4 is 1–1.5; dorsolateral fold indistinct or absent; the humeral gland in males is prominent, raised and centrally positioned on the ventral surface of the upper arm; a weak or absent rictal ridge; outer metatarsal tubercle weak or absent; subarticular tubercles moderate; skin of dorsum smooth or finely shagreened; throat, abdomen, and flanks smooth; posterior section of venter and back of the thigh rugose; dorsum reddish-orange to chocolate-brown, encircled by a white or cream coloured dorsolateral line that encircles the snout, canthus rostralis, outer edge of the upper eyelids, and dorsum along the dorsolateral fold to the vent; lower flanks dark-brown or black below the dorsolateral line grading into a paler venter; dorsal colouration of the limbs same as the flanks with whitish or light-grey speckles or stripes. Abavorana luctuosa can be easily differentiated from its congeners on the basis of its ventral colour pattern which is usually immaculate or with only very faint, sparse, light speckling (Figs
Its distribution spans southern Peninsular Thailand, Peninsular Malaysia, Borneo, and Sumatra (Manthey & Grossmann 1997;
Rana decorata:
Rana luctuosa
(in part):
Rana (Hylorana) luctuosa
(in part):
Rana (Hylarana) luctuosa
(in part):
Hylorana luctuosa
(in part):
Rana (Pulchrana) luctuosa
(in part):
Pulchrana luctuosa
(in part):
Hylarana luctuosa
(in part):
Abavorana luctuosa:
Hylarana (Abavorana) luctuosa
(in part):
MNHNP 1889.226–28 from “Bornéo Kina Balu”, Sabah (Borneo), Malaysia. (
In addition to its phylogenetic placement (Fig.
Distribution (Fig.
Living specimens of Abavorana decorata and Bornean A. luctuosa; A: Abavorana decorata from Tawau Hills, Sabah, East Malaysia. B: A. decorata from Maliau Basin, Sabah, East Malaysia. C: A. decorata from Kubah National Park, Sarawak, East Malaysia. D: A. decorata from Melawi Regency, West Kalimantan Province, Indonesia. E & F: A. luctuosa from Kubah National Park, Sarawak, East Malaysia. G: A. luctuosa (
Rana luctuosa Sukumaran, J. 2005: 38
An adult male (
Abavorana nazgul can be differentiated from its congeners by the following combination of characters: adult males 41.51–44.1 mm SVL and single adult female 52.51 mm; adult males with SW 7.98–8.70 mm, single adult female SW 9.59 mm; adult males with IOD 3.94–4.48 mm, single adult female IOD 4.94 mm; adult males with TD 3.52–4.01 mm, single adult female TD 4.10 mm; adult males with FL 22.20–23.22 mm, single adult female FL 27.80 mm; nuptial pads absent in males; humeral glands in males small (2.4–2.98 mm) (Table
Abavorana nazgul is thus far confirmed from the upper elevations of Gunung Jerai, Kedah, Peninsular Malaysia (
Peninsular Malaysian, Thai and Sumatran Abavorana luctuosa, and A. nazgul; A: A. luctuosa from Sungai Durian Forest Reserve, Kelantan, Peninsular Malaysia. B: A. luctuosa from Janda Baik, Pahang, Peninsular Malaysia. C: A. luctuosa from Semenyih, Selangor, Peninsular Malaysia. D: A. luctuosa from Fraser’s Hill, Pahang, Peninsular Malaysia. E: A. luctuosa from Hala-Bala Wildlife Sanctuary, Thailand. F: A. luctuosa from Sumatra, Indonesia. G: A. nazgul (
Colour pattern comparison among species of the genus Abavorana; Dorsum (A,C & E) and venter (B,D & F) in life of A. luctuosa (
Prior to this study and that of
Borneo has long been a focal point of herpetological research in Southeast Asia (
1 | Mid-dorsal region of dorsum orange or reddish-brown | 2 |
– | Mid-dorsal region of dorsum generally black, sometimes with faint orange or yellow speckles, venter with light spots | A. nazgul |
2 | Venter immaculate or with only very fine, light speckling on flanks and underside of legs, and absence or only faint white streak under the eye and/or tympanum to the corner of the jaw | A. luctuosa |
– | Venter with light and dark spots or reticulations especially along the flanks, and underside of legs boldly marked with black and white reticulations or barring, and a prominent white streak under the eye and/or tympanum to the corner of the jaw | A. decorata |
We thank the Department of Wildlife and National Parks, Peninsular Malaysia for issuing a research permit (JPHL&TN (IP): 100-34/1.24 Jld 14 (71)). We are also indebted to Alan Resetar and Joshua Mata of
Appendix 1
Data type: .docx
Explanation note: Specimens of Abavorana that were examined for morphology in this study. *metamorph specimen that was not measured for comparison.
Appendix 2
Data type: .docx
Explanation note: Summary statistics and principal component analysis scores among male Abavorana decorata, A. nazgul, and A. luctuosa populations from the Malay Peninsula and Borneo. Abbreviations are in the Materials and methods.
Appendix 3
Data type: .docx
Explanation note: Principal component analysis scores among female Abavorana decorata, A. nazgul, and A. luctuosa populations from the Malay Peninsula and Borneo. Abbreviations are in the Materials and methods.
Appendix 4
Data type: .docx
Explanation note: P-values of the ANOVA tests on male and female morphometric datasets. Asterisks (*) denote characters that were found to be significantly different based on α=0.05.