Science Vision www.sciencevision.org Science Vision www.sciencevision.org Science Vision www.sciencevision.org Science Vision www.sciencevision.org

Science Vision 16 (4) October-December Original Research ISSN (print) 0975-6175 ISSN (online) 2229-6026

Observation on the breeding behaviour and analysis of the advertisement calls of maximus Günther, 1858 (Anura: )

Mercy H. Lalramdinfeli and H.T. Lalremsanga*

Department of Zoology, Mizoram University, Aizawl 796004,

Received 18 October 2016 | Revised 26 November 2016 | Accepted 8 December 2016

ABSTRACT

The breeding behaviour and the advertisement calls of the giant tree , Rhacophorus maximus was studied during the breeding season in Mizoram University Campus, Aizawl, Mizoram. At the first shower of the onset of monsoon, advertisement calls of male attract females to breeding areas and announce other males that a given territory is occupied. The aim of this study was to provide the detailed information on the breeding behaviour and analysis of advertisement calls of Rhacophorus maximus. Advertisement calls were emitted in series with variable call intervals. The morphometric measurement of the amplecting pairs were recorded and females are found to be larger than males.

Key words: Breeding behaviour, advertisement calls, Rhacophorus maximus, Mizoram.

INTRODUCTION Indian rhacophorids has been reported by many 5–7 workers. Anurans have a biphasic life cycle The diversity of reproductive modes in am- and they breed in a variety of water bodies such phibians is higher than in any other vertebrate as temporary rainfed ponds, ephemeral pools, group.1 One of the most interesting modes is that cemented tanks and permanent ponds. The of rhacophorids, where majority of them deposit breeding activity of rhacophorids like other an- their eggs in foam nests, while others exhibit urans is influenced by the abiotic factors like 8–10 direct development.2 The foam nests are thought temperature and rainfall. Certain reproduc- to protect the eggs and embryos from predation tive behaviour of many anuran like the and desiccation.3,4 Construction of foam nests by timing of their calling period are linked to cli- mate.11 Vocalization in is a common component of breeding behaviour as male an- Corresponding author: Lalremsanga urans call to advertise their breeding status, de- Phone: : +91-9862559482 fend territory and attract females.12 The social E-mail: [email protected]

CC BY-SA 4.0 International 2016 156 Observation on the breeding behaviour and analysis of the advertisement calls of Rhacophorus maximus behaviours of most anurans are associated with rus maximus commonly known as flying acoustic communication in the form of vocaliza- frog, Günther's tree frog or giant tree frog is tion.13 Advertisement calls are species specific. listed as Least Concern by the IUCN Red List The intra-specific diversity in call characteristics due to its extensive distribution and its tolerance allows females to discriminate among potential of a wide range of habitat types.21 It is a species conspecific mates based on some of the same of frog in the Rhacophoridae family found in acoustic parameters used for species identifica- southwestern (Yunnan, Tibet), northeast- tion.14 ern India, Nepal, western Thailand and north- Although most anurans are normally soli- ern Vietnam, and possibly in . Gupta tary, they often come together in large breeding et al.22 reported that the breeding biology of R. aggregations during the spring or rainy season. maximus have not been studied in the field. Re- Bioacoustic studies play fundamental roles in cently, Khongwir et al.23 reported the breeding understanding and resolving several issues re- and nesting behaviour of this species in Megha- lated to the study of anuran amphibians. Given laya, northeast India, but the call analysis was the importance of acoustic signaling in the not provided. Although it is known that like breeding ecology of most , detailed acousti- other rhacophorids, R. maximus construct foam cal and statistical descriptions of signals are an nests to protect the eggs from desiccation during important first step toward understanding the development there is scanty information on the reproductive and social behaviours of anurans. breeding biology and the advertisement call of Furthermore, basic knowledge of a species’ this tree frog. Therefore, the present investiga- acoustic behaviour has important implications tion was carried out for a better understanding for conservation in at least two respects. On the of its breeding and advertisement. one hand, bioacoustic data can be used as non- invasive tool for the purposes of population cen- MATERIALS AND METHODS sus and monitoring. The integration of bioacous- tic with other data sources can be important for Study sites for Breeding Behaviour effective conservation assessment, planning and management, especially for threatened and en- During the study period, it was observed that dangered species. Given the global decline in Rhacophorus maximus was a seasonal breeder and amphibians15 integrating bioacoustic with anu- its breeding activity coincides with the onset of ran conservation is an important goal. Quantita- monsoon i.e. late February to March in tive descriptions of vocalizations can Mizoram. The study was conducted at two inform an understanding of their evolutionary study sites, i.e., Site Ι: an artificial pond functions, the mechanisms for their utility in (circumference = 26.38 m) located near Lianch- , population monitoring, and conser- hiari road (with a GPS location of N 23° vation.16 44’15.7”: E 92°40’02.5” at an elevation of 824 m Reviews dealing with various aspects or vo- asl) and Site ΙΙ: an old water storage tank (3.54 calization in rhacophoridae have been published m x 2.32 m x 1.8 m) located near Lengteng Boys by few workers, such as the calls of Bornean Hostel (with a GPS location of N 23°44’18.0”: stream-breeding species of Rhacophorus E 92°39’43.2” at an elevation of 775 m asl) in- penanorum , R. kajau , R. belalongensis, R. anguli- side the campus of Mizoram University, Tan- rostris and R. gauni were reported.17-20 However, hril, Aizawl. in northeast India, reports on the breeding be- haviour and analysis of the advertisement call of any species under the , Rhacophorus is lim- Morphometric Measurement ited and there is no detailed study on the adver- tisement call of Rhacophorus maximus. Rhacopho- The morphometric measurement of the am-

157 Lalramdinfeli and Lalremsanga plecting pairs (males and females) were meas- (from the base of the first subarticular tubercle), ured using a dial caliper accurate to 0.01 mm, WFF: Webbing between fourth and fifth toe catalogued and preserved in 4% formalin in the (from the base of the first subarticular tubercle), Departmental Museum of Zoology, Mizoram T1 : From base of foot to tip of longest toe, T2: University (MZMU). From base of foot to tip of second toe, T3: From base of foot to tip of third toe , T4: From base of Abbreviations used foot to tip of fourth toe , T5: From base of foot to tip of fifth toe. SVL: Snout-vent length, HW: Head width, HL: Head length, MN: Distance from the back Acoustic analysis of mandible to the nostril, MFE: Distant from the back of the mandible to the front of the eye, Mating calls were recorded with the help of MBE: Distant from the back of the mandible to digital voice recorder Sony ICD-PX440 Proffe- the back of the eye, IFE: Distant between the sional compact voice recorder. The sampling front of the eye, IBE: Distant between the back used to convert the signals to digital format was of the eye, , IN: Inter nasal space, EN: Eye to 8 KHz with 16–bit precision. The oscillogram nostril (distance from the front of the eye to the was prepared and analysed with the help of a nostril),EL: Eye length, SL: Snout length software tool “SoundRuler Version 0.9.6.0 (distance from the front of the eye to the tip of (acoustic analysis)”. The notes are composed the snout) , SN: Snout to nostril (distance from of groups of pulses. Notes are measured the nostril to the tip of snout) ,TYD: Greatest from the beginning of the first pulse to the end of tympanum diameter, TYE: Distance from tym- the last pulse; intervals between two subsequent panum to the back of eye, IUE: Minimum dis- notes are measured from the end of the last pulse tant between upper eyelids, UEW: Maximum of the first note to the beginning of the first pulse width of inter upper eyelids, , FLL: Fore limb of the following note; note repetition rate is the length (from proximal end of arm to tip of long- number of notes per second; pulse repetition rate est finger), HAL: Hand length (from the base of is the number of pulses per second. outer palmar tubercle to tip of finger), TFL: Third finger length, PA: Width of pads of fin- RESULTS gers, WA: Width of fingers, FL: Femur length, TL: Tibia length, TFOL: Length of tarsus and Breeding season foot, FOL: Foot length, FTL: Fourth toe length, PP: Width of pads of toes, WP: Width of toes, In the present observation, the breeding activ- IMT: Length of inner metatarsal tubercle, ITL: ity of Rhacophorus maximus was stimulated by the Inner toe length, MTTF: Distance from the dis- first shower of monsoon rain in Mizoram. From tal edge of the metatarsal tubercle to the maxi- February to March, the frogs came out to mate mum incurvation of the web between third and in the pools. The atmospheric temperature re- fourth toe, TFTF: Distance from the maximum corded during the investigation period ranges incurvation of the web between third and fourth between 26 to 38ºC, water temperature between toe to the tip of fourth toe, MTFF: Distance 24 to 28ºC, pH between 5.54- 8.22 in both the from the distal edge of the metatarsal tubercle to study sites. the maximum incurvation of the web between fourth and fifth toe, FFTF: Distance from the Courtship and advertisement calls maximum incurvation of the web between fourth and fifth toe to the tip of fourth toe, During the study period, adult male frogs WTF: Webbing between third and fourth toe were observed to be the first to emerge from

158 Observation on the breeding behaviour and analysis of the advertisement calls of Rhacophorus maximus their hiding places during the evening (ca. 3:35 male was noticed from evening to the next PM) and make advertisement calls onwards morning. During observation attracted by the (Fig. 1). It was also observed that multiple males calls, the female frog emerged and responded aggregate and produce advertisement call while towards the breeding ground. On seeing the fe- hiding behind the grasses, some damp places male, the male frog then emerged from its hiding near the breeding site, or while sub-merging on place and encircled the female. It then suddenly the water surface. Advertisement calls were au- grasped the female resulting in axillary amplexus dible to the human ear from a distance of about (Fig. 3). Amplexus was observed to take place 30-35 m away. The calling sound was usually both during the day and night-time. In the pre- heard during the evening and continued till early sent study, combating behavior of males was in the morning. It was observed that the call was observed, where one male frog tried to dislodge remarkably high in the evening after rainfall. another amplecting pair. Pairs in amplexus can Advertisement calls were emitted in series be seen in water submerging together usually at with variable call intervals. The call consisted of the corner of the pool. Amplecting was axillary a single note (Fig. 2) emitted at an interval of 0.8 and a single pair was monitored in the study site -0.9 s. The notes lasted 0.4 s and were composed Ι, whereas five pairs were also encountered in of a series of 2 pulses. The amplitude of the note the study site ΙΙ. Satellite males were also en- increased quickly in its second third and de- countered near the amplecting pair (Fig. 4). crease until the end. The frequency spectra had a They are found to breed in a group as sometimes dominant band at 1464.258 Hz and the band three pairs of amplexus were encountered in the width ranges from 367.1037–692.2985 Hz. same breeding area of study site II. Amplexus was observed to last several hours before the Mating and Spawning nest construction took place and the female de- posited its eggs in a large creamy white foam In this study, vocalization of unsuccessful nest. The nests were observed to be constructed

Figure 1. Emerging male (MZMU 600) start calling from the breeding ground at study site II.

159 Lalramdinfeli and Lalremsanga

Figure 2. Oscillogram, sonogram and frequency spectrum of an advertisement call of male Rhacophorus maximus (MZMU 600).

Figure 3. Axillary amplecting in Rhacophorus maximus (MZMU 1104 and 1105).

160 Observation on the breeding behaviour and analysis of the advertisement calls of Rhacophorus maximus

Figure 4. Amplecting pair (MZMU 1106 and 1107) and satellite male.

Figure 5. Foam nests of Rhacophorus maximus at study site I.

161 Lalramdinfeli and Lalremsanga on diverse substrata. Some of the foam nests ing of amphibians as is a single day with heavy constructed were attached to the grasses and rainfall.23 stones on the side of the pond and floating on Construction of foam nest by R. maximus is the water surface (Fig. 5). Both male and female suggested that to protect the eggs from desicca- frogs left the nests after construction and no pa- tion due to changes in the external environ- rental care was observed. Fresh laid eggs were mental factors during development. It is found collected from the breeding sites and were moni- that adult females of R. maximus are larger (SVL tored in the laboratory. = 70.27-76.90 mm) than males (SVL = 47.88- The study sites are also utilized for breeding 70.14 mm). Sexual dimorphism is represented by other anurans like, Duttaphrynus melanostictus by their sizes. In anurans, spectral call proper- (Bufonidae), Fejervarya limnocharis ties, such as dominant or fundamental fre- (Dicroglossidae), Kaloula pulchra and Microhyla quency, are usually negatively correlated with ornate (Microhylidae), and teraiensis body size because of morphological constraints (Rhacophoridae) during their respective breed- on the sound producing apparatus.24 The adver- ing season. tisement calls of Rhacophorus maximus are Morphometric measurements of the number strongly pulsed, short, consisting of 2 pulses, of six (6) amplecting males and females are as and have frequency spectra with a dominant shown in the Table 1. band of 1464.258 Hz and the band width ranges from 367.1037 to 692.2985 Hz. The calls of an- DISCUSSION other breeding species of Rhacophurus like R. belalongensis are very similar. They are strongly It was observed that Rhacophorus maximus is pulsed consisting of only 1-4 pulses, and have an early breeder, breeding activity is initiated high frequencies with energy maxima between 25 with the first shower of the onset of monsoon, 3500 and 6000 Hz. In constrast, the advertise- i.e. late February to March which is more or less ment calls of similarly sized pond- breeding spe- similar with that of March to April at Cherra- cies like R. rupifes have comparatively much 20 punjee and Mawsynram in Meghalaya.23 The lower frequencies . Several factors have been observation of the present study agreed with invoked to explain geographic variation in frog Khongwir et al.,23 where the species breeds after calls including reinforcement changes in the rainfall which leave standing water for the depo- acoustic environment or a divergence associated sition of spawn. The breeding activity and move- with morphological changes over the geographic 26-29 ments of frogs to the temporary rain fed pools range of the species. Further research on and ponds for spawning are initiated with an rhacophorids of NE India is needed so that more increase in temperature. Rainfall influences the information about their evolution, ecology and reproductive phenology of many spe- reproductive strategies could be obtained. cies, particularly in tropical forests with seasonal precipitation. Therefore, dependency upon an ACKNOWLEDGEMENT aquatic environment for reproduction results in breeding migrations before and after spawning. MHL expresses her sincere thanks to the co- It was suggested that only the onset of monsoon ordinator, DBT State Biotech-Hub, Department stimulates the to emerge from their sub- of Biotechnology, Mizoram University for pro- terranean retreats and strong choruses of breed- viding the studentship for this research work. ing aggregations have been heard following pro- We also thank the Head, Department of Zool- longed non-violent rains which lasted several ogy, Mizoram University for providing the nec- days. Prolonged droughts may completely pre- essary facilities. vent breeding and several continuous days with small rainfall may be as important in the breed-

162 Observation on the breeding behaviour and analysis of the advertisement calls of Rhacophorus maximus

F 2.1 6.14 20.4 7.55 4.59 7.01 6.73 4.62 5.99 1.13 4.37 5.08 5.98 1.32 3.91 2.28 3.79 1108 70.27 23.23 22.79 18.81 11.93 18.74 16.99 19.87 15.89 17.23 13.44 MZMU

M 4.9 3.9 3.4 2.9 8.64 2.08 10.8 5.87 5.67 6.67 8.87 1.37 4.39 8.53 8.19 1.14 3.77 3.92 1.02 2.05 2.33 1107 47.88 15.24 15.11 11.27 15.86 12.24 10.07 MZMU

F 8.9 4.7 24.8 6.52 7.06 6.01 6.04 5.08 2.11 9.85 5.15 2.09 3.56 5.99 4.74 2.79 3.19 4.56 1106 73.92 27.15 21.48 14.84 14.88 20.46 11.24 17.66 18.14 10.47 MZMU

F 7.7 1.5 4.9 8.99 6.23 5.04 5.16 11.5 5.79 2.96 8.89 6.69 4.41 4.16 1.64 3.77 3.31 3.84 1105 66.48 20.67 26.85 21.23 14.36 13.34 19.53 16.04 15.73 13.01 MZMU

M 9.7 22.7 5.62 6.84 6.97 9.92 5.91 4.68 2.81 8.93 18.2 1.55 4.79 4.41 3.63 1.31 2.42 2.52 2.29 1104 70.14 23.55 19.01 13.77 13.67 20.53 11.13 17.92 12.36 MZMU

F 8.8 4.6 4.5 7.7 3.8 33.4 24.8 8.43 1.42 19.2 2.61 4.53 5.27 5.36 2.89 3.86 3.99 1103 76.26 26.97 27.79 22.99 16.14 16.01 11.88 13.54 10.27 19.26 15.31 MZMU

M 7.64 5.19 5.62 8.82 7.44 5.84 2.36 9.42 6.22 3.24 5.24 5.94 5.64 2.72 2.96 3.78 3.74 1102 64.80 22.80 23.24 21.42 16.42 14.55 21.70 11.74 16.34 21.34 13.64 MZMU

F 6.88 8.76 6.78 7.93 5.21 5.92 2.75 8.46 4.75 2.29 4.03 4.56 5.88 2.62 2.47 2.63 3.93 1101 75.02 27.69 23.52 19.66 13.37 14.02 20.53 10.49 18.43 12.78 1O.04 MZMU

* * * * * * * * * * * F 3.9 6.77 8.86 5.36 6.74 5.42 4.73 8.19 4.11 1100 76.90 26.17 20.69 15.53 11.54 14.93 20.59 11.09 MZMU

M 1.4 4.1 602 4.11 6.95 4.76 6.81 4.77 4.48 7.94 2.33 3.93 4.68 4.65 1.99 2.13 3.21 3.34 63.75 24.84 21.24 16.92 11.24 13.21 20.07 10.23 17.35 18.36 11.79 MZMU

M 5.4 6.5 601 6.57 7.17 4.49 3.89 1.01 8.35 5.31 1.98 3.04 4.32 3.44 1.91 2.83 2.04 3.17 64.94 22.46 20.46 17.27 11.63 12.22 17.76 10.05 17.61 16.93 12.75 MZMU

M 7.2 5.9 600 4.84 5.34 3.05 5.93 3.88 9.31 3.45 1.93 4.56 1.51 2.72 3.38 3.78 1.19 2.09 2.41 2.43 56.09 19.24 18.73 16.44 10.87 10.59 11.72 15.07 14.91 MZMU

PA PA II PA III PA IV I WA II WA III WA IV WA PA PA I SVL HW HL MN MFE MBE IFE IBE IN EN EL SN SL TYD TYE IUE UEW FLL HAL TFL No.

SEX Museum 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21.

Table 1: Measurements of the mating frogs (in mm) (Dial caliper accurate to 0.01 mm) 0.01 to (in accurate mm)frogs Measurements the of 1: (Dialmating Table caliper

163 Lalramdinfeli and Lalremsanga

2.9 2.46 3.92 3.63 3.02 2.63 2.41 3.88 3.13 3.89 3.78 8.51 8.65 8.35 19.8 36.33 35.89 47.56 28.79 20.49 10.86 22.57 24.06 11.64 10.82 16.73 18.71 30.87 27.68 34.05

1.2 1.09 2.79 2.01 3.72 3.73 1.56 2.68 2.06 2.98 1.96 5.23 6.58 18.8 6.29 7.77 8.18 5.23 11.1 5.94 23.59 23.78 32.43 19.18 14.01 15.03 18.08 22.89 20.76 23.12

2.2 3.0 2.2 2.4 37.5 3.26 3.56 2.57 1.47 2.53 3.03 3.24 8.54 6.52 28.8 9.97 8.43 36.65 47.82 31.31 18.38 23.08 10.45 13.24 13.29 16.83 23.74 29.13 27.89 37.14

2.36 3.99 3.01 3.48 3.95 2.77 2.05 3.83 3.01 3.81 2.51 9.38 9.52 29.3 26.8 8.41 32.31 33.31 47.07 27.74 17.95 22.25 22.11 11.19 12.87 10.43 10.95 15.09 23.42 35.24

1.3 2.07 2.61 3.06 3.21 3.85 2.79 2.07 2.28 2.93 4.02 9.85 8.44 9.77 6.75 34.25 33.56 42.41 29.98 17.57 20.79 22.95 10.39 10.87 10.08 14.58 23.06 30.99 23.68 33.32

41.2 48.2 29.0 3.89 3.42 4.12 5.88 4.11 2.91 2.64 3.37 3.47 3.96 3.19 7.39 8.51 9.22 15.9 24.2 9.68 41.21 13.64 10.55 26.21 10.35 27.19 10.59 33.88 29.63 41.03

4.1 3.64 4.22 4.64 4.72 4.98 2.66 2.84 3.32 3.42 3.00 8.00 6.52 5.55 35.32 35.06 47.58 31.22 22.08 11.62 17.28 11.92 26.26 11.90 10.93 15.77 22.62 30.72 27.13 34.08

12 3.9 2.2 3.6 2.02 4.52 5.27 5.88 3.52 2.66 2.42 7.09 14.8 9.42 36.87 34.74 48.16 30.94 21.36 11.32 12.54 26.84 11.56 11.64 12.24 12.08 22.56 26.30 28.34 36.52

2.5 9.4 2.27 3.91 4.56 3.77 1.58 2.71 2.01 2.15 2.01 2.47 6.79 6.72 32.8 37.57 36.09 48.19 33.13 18.88 25.56 27.13 12.97 10.25 11.68 11.12 19.17 25.27 32.54 29.99

36.5 1.62 2.71 2.87 3.36 2.95 1.36 2.57 2.73 2.58 2.32 2.67 7.31 8.61 7.85 7.55 9.14 9.23 33.98 44.71 30.17 17.75 20.49 22.13 11.23 14.19 20.95 27.61 24.24 34.31

27.4 2.21 2.49 2.75 3.53 3.64 1.71 1.82 1.05 2.36 2.23 2.69 7.29 7.91 8.83 7.45 33.4 32.18 33.17 41.27 15.81 10.81 20.44 21.51 10.95 10.73 13.89 20.79 24.12 22.67

2.7 1.5 6.8 28.1 1.56 2.15 2.97 2.85 0.87 1.84 1.93 1.85 2.28 6.05 8.35 8.15 7.41 7.25 22.5 7.22 30.98 35.36 23.53 15.69 16.59 17.77 12.75 18.39 24.85 30.63

PP II PP III PP IV PP V WP II WP III WP IV WP V WP FL TL TFOL FOL FTL PP I WP I IMT ITL MTTF TFTF MTFF FFTF WTF WFF T1 T2 T3 T4 T5 TBW TBL

22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. * = Amputated limbs fed on by carnivorous tadpoles. limbscarnivorous by on fed * = Amputated

164 Observation on the breeding behaviour and analysis of the advertisement calls of Rhacophorus maximus

REFERENCES AS, Fischman DL & Waller RW (2004). Status and trends of amphibian declines and extinctions worldwide. Science, 306, 1783–1786. 1. Caldwell JP (1992). Diversity of reprodutive modes in anurans: facultative nest construction in gladiator frogs. 16. Thomas A, Suyesh R, Biju SD & Bee MA (2014). Vocal In: Reproductive biology of South American vertebrates, (WC behavior of the elusive purple frog of India Hamlet, ed.), NY, Springer-Verlag, pp. 85–97. (Nasikabatrachus sahyadrensis), a fossorial species endemic to the . PLoS ONE, 9(2), e84809. 2. Duellman WE & Trueb L (1994). Biology of Amphibians, doi:10.1371/journal.pone.0084809. McGraw-Hill Publishing Company, New York, pp. 1–670. 17. Dehling JM (2008). A new tree frog (Anura: Rhacophori- 3. Heyer WR (1969). The adaptive ecology of the species dae: Rhacophorus) from Gunnung Mulu, Borneo. Salaman- group of the genus Leptodactylus (Amphibia, Leptodactyli- dra, 44, 193–205. dae). Evolution, 23, 421–428. 18. Dring J (1983). Frogs of the genus Leptobrachella 4. Downie JR (1993). Functions of the foam in foam-nesting (Pelobatidae). Amphibia-Reptilia, 4, 89–102. leptodactylids: the nest as a post-hatching refuge in Phy- salaemus pustulosus. Herpetol J, 3, 35–42. 19. Dehling JM, Yambun P & Lakim MB (2010). Advertise- ment calls of Rhacophorus angulirostris Ahl, 1927 and 5. Mohanty-Hejmadi P & Dutta SK (1988). Life history of Rhacophorus everetti macroscelis Boulenger, 1896 from Gun- the common Indian tree frog, (Gray, nung Kinabalu, Sabah, East Malaysia (Borneo). Herpetozoa, 1834) (Anura: Rhacophoridae). J Bom Nat Hist Soc, 85, 512 23, 51–57. –517. 20. Inger RF (1966). The systematics and zoogeography of 6. Kadadevaru GG & Kanamadi RD (2000). Courtship and the amphibia of Borneo. Fieldiana, 52, 1–402. nesting behavior of the Malabar gliding frog, Rhacophorus malabaricus (Jerdon, 1870). Curr Sci, 79, 377–380. 21. IUCN, Conservation International and NatureServe. ( 2 0 0 6 ) . Global Amphibian Assessment, 7. Dutta SK, Mishra S & Rath S (2001). Breeding and devel- www.globalamphibians.org. Accessed on 09 October opment of Polypedates maculatus (Anura: Rhacophoridae). 2007. Pranikee – J Zool Soc, spl. vol, 121–134 22. Gupta BK, Vasudevan K, Taply B & Goetz M (2015). Ex 8. Reading CJ (2003). The effects of variation in climatic situ Management of Amphibians. Assam State Zoo cum Bo- temperature (1980–2001) on breeding activity and tadpole tanical Garden, Guwahati, p. 72. stage duration in the commom toad, Bufo bufo. Sci Total Environ, 310, 231–236. 23. Khongwir S, Hooroo RNK & Dutta SK (2016). Breeding and nesting behaviour of Rhacophorus maximus (Anura: 9. Roy D, Choudhury A & Borah B (2004). Role of weather Rhacophoridae) in Meghalaya, North East India, Curr Sci, condition on the daily appearance and advertisement call 110, 1102–1105. initiation time of Polypedates leucomystax during the breeding season. Zoo’s Print J, 19, 1408–1410. 24. Berry PY (1964). The breeding patterns of seven species of Singapore Anura. J Anim Ecol, 33, 227–243. 10. Deuti K (2001). Breeding ecology of Annandale’s tree frog Chirixalus simus (Anura: Rhacophoridae) near Kol- 25. Dehling JM & Grafe TU (2008). A new treefrog of the kata, West Bengal. J Bom Nat Hist Soc, 98, 341–346. genus Rhacophorus (Anura: Rhacophoridae) from Brunei Darussalam (Borneo). Salamandra, 44, 101–112. 11. Gibbs JP & Breisch AR (2001). Climate warming and calling phenology of frogs near Ithaca, New York, 1990– 26. Martin WF (1972). Evolution of vocalization in the genus 1999. Conserv Biol, 15, 1175–1178. Bufo. In: Evolution in the Genus Bufo, (WF Blair, ed.), Univer- sity of Texas Press, Austin, pp. 279–309. 12. Mitchell NJ (2002). Nest-site selection in a terrestrially breeding frog with protracted development. Aust J Zool, 27. Butlin R (1987). Speciation by reinforcement. Trends Ecol 50, 225–235. Evol, 2, 8–13. 13. Krishna SN & Krishna SB (2005). Female courtship calls 28. Loftus-Hills JJ & Littlejohn MJ (1992). Reinforcement of the litter frog (Rana curtipes) in the tropical forests of and reproductive character displacement in Gastrophyne Western Ghats, South India. Amphibia-Reptilia, 26, 431- carolinensis and G. olivacea (Anura: Microhylidae): a re- 435. examination. Evolution, 46, 896–906. 14. McClelland BE, Wilczynski W & Ryan MJ (1996). Corre- 29. Howard DJ & Gregory PG (1993). Post-insemination lations between call characteristics and morphology in signaling systems and reinforcement. Phil Trans Royal Soc male cricket frogs (Acris crepitans). J Exp Biol, 199, 1907- London Biol Sci, 340, 231–136. 1919. 15. Stuart SN, Chanson JS, Cox NA, Young BE, Rodrigues

165