Delineating ecological boundaries of Hanuman langur species complex in peninsular India using MaxEnt modeling approach.

Hanuman langur is one of the widely distributed and extensively studied non-human diurnal primates in India. Until recently it was believed to be a single species - Semnopithecus entellus. Recent molecular and morphological studies suggest that the Hanuman langurs consists of at least three species S. entellus, S. hypoleucos and S. priam. Furthermore, morphological studies suggested that both S. hypoleucos and S. priam have at least three subspecies in each. We explored the use of ecological niche modeling (ENM) to confirm the validity of these seven taxa and an additional taxon S. johnii belonging to the same genus. MaxEnt modeling tool was used with 19 bioclimatic, 12 vegetation and 6 hydrological environmental layers. We reduced total environmental variables to 14 layers after testing for collinearity and an independent test for model prediction was done using ENMTools. A total of 196 non-overlapping data points from primary and secondary sources were used as inputs for ENM. Results showed eight distinct ecological boundaries, corroborating the eight taxa mentioned above thereby confirming validity of these eight taxa. The study, for the first time provided ecological variables that determined the ecological requirements and distribution of members of the Hanuman langur species complex in the Indian peninsula

Philautus ochlandrae Gururaja, Palot & Radhakrishnan, 2007, sp. nov.

Philautus ochlandrae sp. nov. (Figure 1 a –h; Table 1) Holotype: Zoological Survey of India, WGFRS, Calicut (ZSI/ WGFRS /V/A/ 632), an adult male from Ochlandra setigera reed was collected at Kakkayam Reserve Forest, Calicut District, Kerala State, by DKP, MdJP and KVG on 23 rd April 2007 (11 ° 33 ’ 16 ” N, 75 ° 55 ’ 12 ” E, altitude ~ 745 m above mean sea level) between 17: 30 – 22:00 h. Paratypes: ZSI/ WGFRS /V/A/ 633, 634, two males and ZSI/ WGFRS /V/A/ 635, 636, a male and a female collected from the same locality on 24 th April 2007 at 08:00 h by DKP, MdJP and KVG, collection data same as holotype. ZSI/ WGFRS /V/A/ 637, a male collected by DKP and KVG on 27 th April 2007 at 21:00 h from the same locality. Diagnosis: A small-sized frog diagnosed as Philautus (Male: 22.1–25.6 mm; Female: 23.3 mm), having granular belly, all digits with well differentiated disks bearing circummarginal grooves, predominantly inhabiting shrubs (this species exclusively inhabits hollow tubular internodes of O. setigera reed brake) and having direct development as in Philautus cf. leucorhinus (Gururaja & Ramachandra, 2006). It is distinguished from all other congeners of Western Ghats by the following combination of characters: (i) body small, elongate (ii) habitus squat and flat; (iii) head arched, wider than long; (iv) snout short rounded, equal or sub equal to diameter of eye; (v) tympanum indistinct but visible; (vi) canthus rostralis rounded; (vii) eyes protruding, pupil with striking golden yellow dentition like marks; (viii) belly granular, under parts of forearm and thigh granular; (ix) vocal sac unpigmented and (x) fleshy brown to cream yellow dorsum with two distinct golden yellow lateral bands bordered by dark brown from upper eyelid to the posterior part of flanks. Description of the holotype: A small sized shrub frog (SVL = 25.6 mm), width of head broader than head length (HW = 8.6 mm; HL = 6.4 mm), arched, flat dorsally; snout rounded in total profile, slightly protruding beyond mouth. Snout length is equal to diameter of eye (SL = 2.9 mm, EL = 2.9 mm). Canthus rostralis rounded, loreal region slightly concave. Interorbital space (IUE = 2.8 mm) flat and broader than upper eyelid (UEW = 1.9 mm), wider than internarial distance (IN = 1.7 mm). Internarial distance between posterior margins of the eyes 1.93 times that of anterior margins (IFE = 4.0, IBE = 7.7 mm). Nostrils oval, nearer to tip of snout (NS = 0.8 mm) and away from eye (EN = 1.9 mm). Pineal ocellus absent. Weak symphysial knob. Vomerine ridges absent. Eyes large, protruding, pupil horizontal, with golden yellow dentition like marks interspersed with black. Tympanum rather indistinct, rounded, barely visible behind the eye, 2.15 times in eye diameter (TYD = 1.4 mm). Tongue bifid, granular with a distinct retractile papilla. Supratympanic fold from behind eye to shoulder. Median subgular vocal sac with a pair of opening at the base of the lower jaw. Fore arm (FLL = 4.4 mm) less than hand (HAL = 7.0 mm). Relative length of fingers I<II<IV<III. Finger tips with well developed disks (fd 1 = 0.8 mm, fd 2 = 1.1 mm, fd 3 = 1.4 mm, fd 4 = 1.4 mm; fw 1 = 0.5 mm, fw 2 = 0.6 mm, fw 3 = 0.9 mm, fw 4 = 0.8 mm) with distinct circum–marginal grooves, fingers with dermal fringes on both sides. Webbing on palm absent, subarticular tubercles indistinct, rounded and pre–pollex tubercle oval, distinct. Supernumerary tubercles absent. Hind limb long, heels barely touch when folded at right angles to the body. Tibia 2.9 times longer than wide (TL = 10.1 mm, TW = 3.4 mm), subequal to femur (FL = 10.4 mm) and longer than foot (FOL = 9.5 mm). Heel to tip of fourth toe (TFOL = 14.9 mm) 2.76 times length of fourth toe (T 4 L = 5.4 mm). Relative toe length I<II<III<V<IV. Toe disk width and toe width are td 1 = 0.8 mm, td 2 = 1.0 mm, td 3 = 1.3 mm, td 4 = 1.5 mm, td 5 = 1.5 mm; tw 1 = 0.7 mm, tw 2 = 0.7 mm, tw 3 = 0.8 mm, tw 4 = 0.9 mm, tw 5 = 0.9 mm. Webbing moderate and distinct (MTTF = 5.5 mm, MTFF = 5.5 mm, TFTF = 4.4 mm, FFTF = 3.9 mm); web formula, I 2 – 2 II 1– 2 III 1–2 IV 2 – 1 V. Tibiotarsal articulation reaches tympanic region. First toe (T 1 L = 1.5 mm) nearly 1.9 times the length of inner metatarsal tubercle (IMT = 0.8 mm). Outer metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent. Skin: Snout, between eyes, sides of head, dorsum smooth. Dorsal part of forelimb, femur, tibia and tarsus smooth to very finely granular. Venter granular, larger granulation on belly extending up to throat, flanks (mid part of dorsolateral band) and thigh. Dorso–lateral folds and macroglands absent. Color in life: Dorsum fleshy brown with two lateral golden yellow bands bordered by dark brown from upper eyelid to the posterior part of the flanks (¾ of flank). Head, forelimbs and hind limbs brown dorsally. Hand ventrally cream, a few brown granulation along the dermal fringes of third and fourth finger. Upper lip and lower lip cream colored. Venter granular and uniform cream. Granulation on femur, tibia, tarsus and entire foot, brown ventrally. Webbing brown. Eyes with horizontal black pupil surrounded by golden yellow dentition like marks interspersed with black. Color in preservative: Dorsum and lateral parts of body light sandy brown with two lateral golden yellow bands, bordered by dark brown color from upper eyelid to the region of groin. Dorsal and lateral parts of limbs uniform without any cross bars. Ventral parts of head, body and hand cream white, belly translucent, granular. Granulation on foot, toes, disks and webbing, brown dorso–ventrally. Etymology: The species epithet refers to the generic name of Ochlandra setigera, the plant in which we observed this species (both male and female) as well as its eggs and advertisement calls. Sexual dimorphism: Nuptial spines absent in male, possess a median subgular vocal sac with a pair of openings at the base of the lower jaw. Female larger than the paired male (SVL: 23.3 mm and 22.1 mm respectively). Ovary large, with creamy white eggs. Female paler than males, dorsum golden yellow and two lateral stripes slightly thinner than males. Va r ia ti o n: Table 1 details the morphometric and meristic variations observed in six individuals. Color on the dorsum varied from dark brown to brown interspersed with yellow blotches to broad yellow bands (this is apart from two lateral distinct golden yellow bands from upper eyelid to the posterior part of the flanks). In individuals with brown dorsum and yellow spots, the yellow spots were in four–six stripes, two stripes commencing at snout dividing in to four at interorbital space, and widening at mid dorsum to which two more stripes are added that coalesce near vent. In individuals with brown dorsum and yellow stripes, the yellow stripe starts singly at snout, bifurcating at dorsum and ending near vent. Dorsum of forelimb and hind limb brown with yellow blotches. A yellow stripe along the canthus rostralis was also noticed in an individual. Additional information from paratypes: Morphometric data are given in Table 1. All the paratypes are in good condition, except for ZSI/ WGFRS /V/A/ 636, which was incised ventrally to determine sex. TABLE 1 Morphometric (in mm) and meristic data for the type series of Philautus ochlandrae sp. nov. Prefix ZSI/ WGFRS /V/A/ for all Holotype and Paratypes. For abbreviations see text. fd 1 0.8 0.7 0.7 0.7 0.8 0.7 ± 0.07 (0.7–0.8) 0.9 fd 2 1.1 0.9 1.1 1.0 1.1 1.0± 0.10 (0.9–1.1) 1.1 fd 3 1.4 1.2 1.4 1.3 1.5 1.3 ± 0.10 (1.2–1.5) 1.4 fd 4 1.4 1.2 1.2 1.2 1.3 1.3 ± 0.11 (1.2–1.4) 1.3 fw 1 0.5 0.6 0.6 0.6 0.6 0.6 ± 0.04 (0.5–0.6) 0.6 fw 2 0.6 0.9 0.8 0.7 0.7 0.8 ± 0.11 (0.6–0.9) 0.9 fw 3 0.9 0.9 0.9 0.9 1.1 0.9 ± 0.07 (0.9–1.1) 1.0 fw 4 0.8 1.0 0.8 1.0 0.8 0.9 ± 0.12 (0.8 –1.0) 0.9 ......continued Ecology and natural history notes: This species was first noticed on a reed brake on 23 rd April 2007 calling at 19: 30 h while it was drizzling. Other species calling at that time were Nyctibatrachus cf. aliciae Inger, Shaffer, Koshy and Bakde, Sylvirana aurantiaca (Boulenger), Philautus cf. wynaadensis, and P. ponmudi. Two more individuals were collected during the same night from reed brakes. On 24 th April 2007 at 8:00 h, two individuals were collected, of which one was female. These frogs reside in the hollow tube of internodal region of O. setigera reed brake. On four occasions, calling males were noticed inside culms of O. setigera. Generally, this species remain within these culms throughout the day at a height of 2.42 ± 0.25 m (range: 2.25–2.7 m) above ground. These culms (girth: 81.5 ± 9.4 cm, range: 70–93 cm) had slit like openings (length: ~27.0 mm, width: ~5.0 mm) which the frogs used as portals. Males start calling early (around 16.00h) on rainy days, increase in number at dusk and subsequently last until around 22:00h. However, on two occasions, we heard them calling at 8:00 h in the morning. We observed that individuals reside only in live reeds. Specimens were found as pairs (a male and a female) in culms on two occasions, and solitarily in different culms and reed brakes at the type locality. In subsequent field visits (29 th May 2007) we observed an egg clutch with six developing embryos (Figure 1 b) (diameter of eggs with jelly cover: 4.94 ± 0.06 mm, range: 4.87–5.01 mm) ensconced in jelly cover, attached to inner walls of hollow reeds, ~ 12 cm above the opening. Embryos were cream-white in color and had pigmented eyes with visible heart-beats and movements; which would eventually hatch into froglets as in other Philautus of the region (Gururaja & Ramachandra, 2006). This is similar to development mode 20 (Duellman & Trueb, 1994). Both males and females were in the same hollow reed, indicating the possible provisioning of parental care. However, this requires further field investigations to substantiate. Advertisement call analysis: Calls were recorded using Olympus digital voice recorder (W– 10, Olympus) within 10–20 cm from calling males. The air temperature and relative humidity were 93.75 ± 3.3 % (range: 90–98 %) and 21.76 ± 1.45 °C (range: 20–23.6 °C) respectively. Calls were recorded from four individuals, on two days. Six calls were analyzed using SIGVIEW 32 Ver.1.9.3.2. Call terminology were based on Giacomo & Castellano (2001). Advertisement call had a short four pulse call (shriek ‘ shreaaw ’ note) and a long 27–73 pulse call (‘ tink tink tink tink ….’ notes). Average dominant frequency was 2796.82 ± 125.49 Hz (range: 2691.23–2978.56 Hz), Call duration was 5.42 ± 1.90 sec (range: 3.04–8.23 sec), short call pulse duration 0.22 ± 0.05 sec (range: 0.17–0.27 sec), short pulse rate 19.37 ± 4.37 sec– 1 (Range: 14.83–24.24 Sec– 1), long pulses were 48.33 ± 17.84 (range: 27–73), long call pulse duration 5.11 ± 1.92 sec (range: 2.74–7.94 sec), long pulse rate 9.47 ± 0.63 sec– 1 (range: 8.52–10.21 sec– 1). Figure 2 illustrates call spectrum and amplitude of a single advertisement call of 8.22 sec duration. Comparisons: We compared the new species with all 22 known species from south India listed in Appendix-I. Sri Lankan and South-east Asian Philautus are excluded from comparison as they form phylogenetically distinct clades (Bossuyt et al. 2004; Manamendra– Arachchi & Pethiyagoda, 2005). However, the phylogenetic status of P. cf. leucorhinus and P. cf. variabilis is not fully resolved, and they were therefore included in morphological comparison. Morphometric and meristic data of type specimens were taken from the original description of the respective species (Table 2), additional information is compiled from subsequent publications as cited in Appendix–I. Philautus ochlandrae sp.nov. differs from all other Indian species of Philautus in a number of characters. Appendix–I provides the opposing suites of characters of these congeners compared to P. ochlandrae sp.nov. Considering the morphological information of 17 male type specimens of Philautus, we performed a cluster analysis using STATISTICA software (version 5.5). This analysis was used to compare the new species with other known congeners, based on unweighted pair group averages and squared Euclidian distance measure of 19 morphometric and three meristic characters. We included only male type specimens for cluster analysis, hence P. tinniens and P. variabilis (with female type specimens) were excluded from the analysis. We also excluded P. beddomii, P. bombayensis, P. chalazodes, and P. travancoricus, for lack of data. From the dendrogram generated (Figure 3), despite a minor branching among the individuals of P. ochlandrae sp. nov., overall clustering clearly indicates the species to be new. There is an overlap of P. ochlandrae sp. nov. with P. griet, however, P. griet differs from P. ochlandrae sp.nov., in the following characters: dorsum brown with large spines; tongue without papilla; supernumerary tubercles on both fingers and toes; webbing on toes rudimentary; webbing transparent with black spots; vocal sac and throat light gray; thighs cross barred. Note: For abbreviations see text. Species 1. P. anili; 2. P. bobingeri; 3. P. charius; 4. P. dubois; 5. P. flaviventris; 6. P. glandulosus; 7. P. graminirupes; 8. P. griet; 9. P. cf. leucorhinus; 10. P. luteolus; 11. P. neelanethrus; 12. P. nerostagona; 13. P. ponmudi; 14. P. signatus; 15. P. tinniens; 16. P. tuberohumerus; 17. P. variabilis; 18. P. wynaadensis. STF: present = 1, absent = 0; LP: present = 1, absent = 0, WF: 0.2 = reduced, 0.3 = partial, 0.5 = medium, 0.8 = nearly full, 1.0 = full.Published as part of Gururaja, Kotambylu Vasudeva, Palot, Muhamed Jafer & Radhakrishnan, C, 2007, A new species of Philautus Gistel (Amphibia: Anura: Rhacophoridae) from southern Western Ghats, India, pp. 1-16 in Zootaxa 1621 on pages 2-8, DOI: 10.5281/zenodo.17921

Delineating Ecological Boundaries of Hanuman Langur Species Complex in Peninsular India Using MaxEnt Modeling Approach

Hanuman langur is one of the widely distributed and extensively studied non-human diurnal primates in India. Until recently it was believed to be a single species - Semnopithecus entellus. Recent molecular and morphological studies suggest that the Hanuman langurs consists of at least three species S. entellus, S. hypoleucos and S. priam. Furthermore, morphological studies suggested that both S. hypoleucos and S. priam have at least three subspecies in each. We explored the use of ecological niche modeling (ENM) to confirm the validity of these seven taxa and an additional taxon S. johnii belonging to the same genus. MaxEnt modeling tool was used with 19 bioclimatic, 12 vegetation and 6 hydrological environmental layers. We reduced total environmental variables to 14 layers after testing for collinearity and an independent test for model prediction was done using ENMTools. A total of 196 non-overlapping data points from primary and secondary sources were used as inputs for ENM. Results showed eight distinct ecological boundaries, corroborating the eight taxa mentioned above thereby confirming validity of these eight taxa. The study, for the first time provided ecological variables that determined the ecological requirements and distribution of members of the Hanuman langur species complex in the Indian peninsula

Mud-packing frog: A novel breeding behaviour and parental care in a stream dwelling new species of Nyctibatrachus (Amphibia, Anura, Nyctibatrachidae)

Gururaja, Kotambylu Vasudeva, Dinesh, K. P., Priti, H., Ravikanth, G. (2014): Mud-packing frog: A novel breeding behaviour and parental care in a stream dwelling new species of Nyctibatrachus (Amphibia, Anura, Nyctibatrachidae). Zootaxa 3796 (1): 33-61, DOI: 10.11646/zootaxa.3796.1.

Breeding in bamboo: a novel anuran reproductive strategy discovered in Rhacophorid frogs of the Western Ghats, India

Amphibians exhibit extraordinarily diverse sets of reproductive strategies among vertebrates. Understanding life history strategies in an evolutionary framework is lacking for many amphibian species in the tropics. Here, we report a novel reproductive mode where adult frogs enter hollow internodes of bamboo via a small opening, deposit direct developing eggs, and provide parental care. This behaviour is observed in two species of the frog genus Raorchestes. The first description of this unique life history and details of nest site characteristics and embryo development are provided along with ecological comparisons. Evolution of novel reproductive modes and parental care are discussed in context of natural selection. Dearth of natural history information on amphibians in the Western Ghats and much of the South-East Asian region is highlighted with suggestions for further studies.(c) 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114, 1-11

Predicted area of distribution for all the taxa.

<p>Predicted area of distribution for all the taxa.</p

Proportion of the predicted area of taxa A occupied by the predicted area of taxa B.

<p><b>For above the diagonal values, species in row-heads are taxa A and column-heads are taxa B. For below the diagonal values, taxa in row-heads are taxa B and column-heads are taxa A.</b></p

Maxent distribution modeling logistic output for <i>S. p. priam</i>, <i>S. entellus</i>, <i>S. h. iulus</i> and <i>S. johnii</i>.

<p>Maxent distribution modeling logistic output for <i>S. p. priam</i>, <i>S. entellus</i>, <i>S. h. iulus</i> and <i>S. johnii</i>.</p

Percentage Niche overlaps (%) between taxa pairs to their area of prediction.

<p>Percentage Niche overlaps (%) between taxa pairs to their area of prediction.</p

Non-overlapping occurrence data points of the various taxa used in the present study.

<p>Non-overlapping occurrence data points of the various taxa used in the present study.</p