Spring-Breeding and Reproductive Mode in Leptolalax khasiorum (Anura, Megophryidae) in North-Eastern India

Ronald K. L. Tron, Indraneil Das, Rupa N. K. Hooroo, Duwaki Rangad


We examined the breeding habitat, oviposition site and timing of breeding in Leptolalax khasiorum at its type locality at Mawphlang, Meghalaya State, north-eastern India. Surveys were conducted along a stream within a fragmented patch of forest for adults, tadpoles, eggs and nesting sites, with data collected monthly between January 2009 and December 2011. The species starts emerging from wintering microhabitats with the onset of the premonsoonal showers in February. Breeding activity (specifically, deposited eggs) was observed for a few weeks after emergence and lasted a few weeks between early March and April. Adults were found under rocks along the stream banks. Eggs are deposited on the underside of rocks that are flatly embedded on the gravel and over leafy bottom of the dry stream bed. The eggs of Leptolalax khasiorum are cream-colored, lacking an animal pole, and covered with a transparent jelly, showing adaptations for terrestrial development in sheltered microhabitats via avoidance of moisture loss and damage of eggs from solar radiation. In producing eggs out of water, showing free-swimming tadpoles, the reproductive mode can be classified as Mode 18, a first for a member of the genus Leptolalax. It is arguably the first report of terrestrial reproduction and parental care for a member of the family Megophryidae. Placement of eggs outside of water may coincide with low water levels of early spring, advantages of choice of timing being reduced competition for egg-laying sites and food for the larval and postlarval stages.


Leptolalax khasiorum; North-eastern India; Mawphlang; oviposition; reproductive mode

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Aichinger M. (1987), «Annual activity patterns of anurans in a seasonal neotropical environment», Oecologia, 71, 583 – 592.

Alford R. A. (1999), «Ecology. Resource use, competition, and predation», in: R. W. McDiarmid and R. Altig (eds.), Tadpoles. The Biology of Anuran Larvae, Univ. of Chicago Press, Chicago – London, pp. 240 – 278.

Altig R. and McDiarmid R. W. (2007), «Morphological diversity and evolution of egg and clutch structure in amphibians», Herpetol. Monogr., 21, 1 – 32.

Ancona N. and Capietti A. (1995), «Analysis of the breeding site characteristics of amphibians in a prealpine area (Triangolo Lariano)», in: G. A. Llorente, A. Montori, X. Santos, and M. A. Carretero (eds.), Scientia Herpetologica, Asociacion Herpetol. Esp., Barcelona, pp. 160 – 164.

Augert D. and Guyetant R. (1995), «Space occupation for egg deposition in amphibians living in plain woodland and pasture land (east of France)», in: G. A. Llorente, A. Montori, X. Santos, and M. A. Carretero (eds.), Scientia Herpetologica, Asociacion Herpetol. Esp., Barcelona, pp. 165 – 169.

Beebeee T. J. C. (1985), «Discriminant analysis of amphibian habitat determinants in southeast England», Amphibia–Reptilia, 6, 35 – 43.

Bertoluci J. and Rodrigues M. T. (2002), «Seasonal patterns of breeding activity of Atlantic rainforest anurans at Boracéia, southeastern Brazil», Amphibia–Reptilia, 23, 161 – 167.

Blaustein L. (1999), «Oviposition habitat selection in response to risk of predation: consequences for population dynamics and community structure», in: S. P. Wasser (ed.), Evolutionary Theory and Processes: Modern Perspectives, Kluwer Academic Publishers, Dordrecht, pp. 444 — 456.

Calef G. W. (1973), «Natural mortality of tadpoles in a population of Rana aurora», Ecology, 54, 741 – 758.

Cei J. M. (1980), «Amphibians of Argentina», Italian J. Zool. Monogr., 2, 1 – 609.

Crump M. L. (1991), «Choice of oviposition site and egg load assessment by a treefrog», Herpetologica, 47, 308 – 315.

Das I. (2007), Amphibians and reptiles of Brunei. A Pocket Guide, Natural History Publications (Borneo) Sdn Bhd, Kota Kinabalu.

Das I., Tron R. K. L., Rangad D., and Hooroo R. N. K. (2010), «A new species of Leptolalax (Anura: Megophryidae) from the sacred groves of Mawphlang, Meghalaya, north-eastern India», Zootaxa, 2339, 44 – 56.

Dehling J. M. (2012), «Redescription of Leptolalax gracilis (Günther, 1872) from Borneo and taxonomic status of two populations of Leptolalax (Anura: Megophryidae) from peninsular Malaysia», Zootaxa, 3328, 20 – 34.

Duellman W. E. and Trueb L. (1994), Biology of Amphibians, McGraw-Hill, New York.

Edgerly J. S., McFarland M., Morgan P., and Livdahl T. (1998), «A seasonal shift in egg-laying behaviour in response to cues of future competition in a treehole mosquito», J. Animal Ecol., 67, 805 – 818.

Ensabella F., Loriga S., Formichetti P., Isotti R., and Sorace A. (2003), «Breeding site selection of Bufo viridis in the city of Rome (Italy)», Amphibia–Reptilia, 24, 396 – 400.

Evans M., Yáber C., and Hero J. M. (1996), «Factors influencing choice of breeding site by Bufo marinus in its natural habitat», Copeia, 1996, 904 – 912.

Gallardo J. M. (1974), Anfibios de los alrededores de Buenos Aires, Editorial Universitaria de Buenos Aires, Buenos Aires.

Goldberg F. J., Quinzio S., and Vaira M. (2006), «Oviposition site selection of the toad, Melanophryniscus rubriventris in an unpredictable environment», Can. J. Zool., 84, 699 – 705.

Gururaja K. V. (2010), «Novel reproductive mode in a torrent frog Micrixalus saxicola (Jerdon) from the Western Ghats, India», Zootaxa, 2642, 45 – 52.

Haddad C. F. B., Cardoso A. J., and Castanho L. M. (1990), «Hibridação natural entre Bufo ictericus e Bufo crucifer (Amphibia: Anura)», Rev. Brasil Biol., 50, 739 – 744.

Haddad C. F. B. and Prado C. P. A. (2005), «Reproductive modes in frogs and their unexpected diversity in the Atlantic forest of Brazil», BioScience, 55, 207 – 217.

Haddad C. F. B. and Sazima I. (1992), «Anfíbios anuros da Serra do Japi», in: L. P. C. Morellato (ed.), História natural da Serra do Japi: ecologia e preservação de uma área florestal no sudeste do Brasil, Unicamp, Campinas, pp. 188 – 211.

Hazell D., Hero J.-M., Lindenmayer D., and Cunningham R. (2003), «A comparison of constructed and natural habitat for frog conservation in an Australian agricultural landscape», Biol. Conserv., 119, 61 – 71.

Hertwig S., Das I., Schweizer M., Brown R. M., and Haas A. (2011), «Phylogenetic relationships of the Rhacophorus everetti-group and implications for the evolution of reproductive modes in Philautus (Amphibia: Anura: Rhacophoridae)», Zool. Scripta, 41, 29 – 46.

Heyer W. R., McDiarmid R. W., and Weigmann D. L. (1975), «Tadpoles, predation and pond habitats in the tropics», Biotropica, 7, 100 – 111.

Inger R. F. (1985), «Tadpoles of the forested regions of Borneo», Fieldiana Zool. N. Ser., 26, 1 – 89.

Kats L. B., Petranka J. W., and Sih A. (1988), «Antipredator defenses and the persistence of amphibian larvae with fishes», Ecology, 69, 1865 – 1870.

Licht L. E. (1971), «Breeding habits and embryonic thermal requirements of the frogs, Rana aurora aurora and Rana pretiosa pretiosa, in the Pacific Northwest», Ecology, 52, 116 – 124.

Mahony S. (2008), «Redescription and generic reallocation of Rana mawphlangensis Pillai and Chanda, 1977 (Amphibia: Ranidae)», Hamadryad, 33, 1 – 12.

Mahony S., Teeling E. C., and Biju S. D. (2013), «Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region», Zootaxa, 3722, 143 – 169.

Malkmus R. and Kosuch J. (1999), «Beschrijving van het kikkervisje van Leptolalax arayai Matsui, 1997, eenkikkeruit de familie Megophryidae», Lacerta, 57(5), 161 – 165.

Murphy P. J. (2003), «Does reproductive site choice in a Neotropical frog mirror variable risks facing offspring?» Ecol. Monogr., 73, 45 – 67.

Newman R. A. (1988), «Adaptive plasticity in development of Scaphiopus couchii tadpoles in desert ponds», Evolution, 42, 774 – 783.

Pombal J. P., Jr. (1997), «Distribuição espacial e temporal de anuros (Amphibia) em uma poça permanente na Serra de Paranapiacaba, sudeste do Brasil», Rev. Brasil Biol., 57, 583 – 594.

Purakayastha J. and Matsui M. (2010), «A new species of Fejervarya (Anura: Dicroglossidae) from Mawphlang, northeastern India», Asian Herpetol. Res., 3, 31 – 37.

Reading C. J. (1998), «The effect of winter temperatures on the timing of breeding activity in the common toad Bufo bufo», Oecologia, 117, 469 – 475.

Resitarits W. J., Jr. and Wilbur H. M. (1989), «Choice of oviposition site by Hyla chrysoscelis: role of predators and competitors», Ecology, 70, 220 – 228.

Richter-Boix A., Llorente G. A., and Montori A. (2006), «Breeding phenology of an amphibian community in a Mediterranean area», Amphibia–Reptilia, 27, 549 – 559.

Rowley J. and Trung C. T. (2009), «A new species of Leptolalax (Anura: Megophryidae) from central Vietnam», Zootaxa, 2198, 51 – 60.

Rowley J., Stuart B. L., Richards S. J., Phimmachak S., and Sivongxay N. (2010), «A new species of Leptolalax (Anura: Megophryidae) from Laos», Zootaxa, 2681, 35 – 46.

Rudolf V. H. W. and Rödel M. O. (2005), «Oviposition site selection in a complex and variable environment: the role of habitat quality and conspecific cues», Oecologia, 142, 316 – 325.

Salthe S. N. (1969), «Reproductive modes and the number and sizes of ova in the urodela», Am. Midl. Nat., 81, 467 – 490.

Salthe S. N. and Duellman W. E. (1973), «Quantitative constraints with reproductive mode in anurans», in: J. L. Vial (Ed.), Evolutionary Biology of the Anurans, Univ. of Missouri Press, Columbia, pp. 229 – 249.

Sengupta S., Sailo S., Lalremsanga H. T., Das A., and Das I. (2010), «A new species of Leptolalax (Anura: Megophryidae) from Mizoram, north-eastern India», Zootaxa, 2406, 57 – 68.

Skelly D. K. (2001), «Distributions of pond-breeding anurans: an overview of mechanisms», Israel J. Zool., 47, 313 – 332.

Smith D. C. (1983), «Factors controlling tadpole populations of chorus frog (Pseudacris triseriata) on Isle Royale, Michigan», Ecology, 64, 501 – 510.

Spieler M. and Linsenmeir K. E. (1997), «Choice of optimal oviposition sites by Hoplobatrachus occipitalis (Anura: Ranidae) in an unpredictable and patchy environment», Oecologia, 109, 184 – 199.

Summers K., McKeon C. S., Heying H., Hall J., and Patrick W. (2007), «Social and environmental influences on egg size evolution in frogs», J. Zool. London, 271, 225 – 232.

Telford S. R. and Dyson M. L. (1990), «The effect of rainfall on interclutch interval in Painted Reed Frog (Hyperolius marmoratus)», Copeia, 1990, 644 – 648.

Touchon J. C. and Warkentin K. M. (2008), «Reproductive mode plasticity: aquatic and terrestrial oviposition in a treefrog», Proc. Natl. Acad. Sci. USA, 105, 7495 – 7499.

Van Gomez-Mestre I., Pyron R. A., and Wiens J. J. (2012), «Phylogenetic analyses reveal unexpected patterns in the evolution of reproductive modes in frogs», Evolution, 66, 3687 – 3700.

Watling J. I. and Donnelly M. A. (2002), «Seasonal patterns of reproduction and abundance of leaf litter frogs in a Central American rainforest», J. Zool. London, 258, 269 – 276.

Wells K. D. (1977), «The social behaviour of anuran amphibians», Anim. Behav., 25, 666 – 693.

Wiest J. A. (1982), «Anuran succession at temporary ponds in a pot oak-savanna region of Texas», Herpetological Communities, 13, 39 – 47.

Wright A. H. (1914), North American Anura. Life-Histories of the Anura of Ithaca, Carnegie Institution of Washington, New York – Washington.

Wright A. H. and Wright A. A. (1949), Handbook of Frogs and Toads, Comstock, Ithaca, NY.

DOI: https://doi.org/10.30906/1026-2296-2015-22-3-224-232


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