Reproduction,phenology and mortality sources of a species of Physalaemus (Anura: Leptodactylidae)

Physalaemus cf. fuscomaculatus is a foam-nesting frog of seasonally dry environments of south-eastern Brazil. We report on the reproduction and other natural history features of this species. Population parameters were based on trapped specimens. The advertisement call is described. The calling and spawning season is coincident with the rainy period. Calling and spawning were performed in ponds. Most of the adults were trapped in October and the juveniles during wetter months and the beginning of the dry season. At the end of the reproductive season, the median size of adults lowered. Egg clutches averaged 2350 eggs, each egg being about 1.4?mm in diameter. Males were more abundant than females in traps. Communal foam nests were frequent, involving up to 40 individual nests. Tadpoles entered the water within about 40?h. Maggots (Diptera) infested egg clutches; water bugs and mammals preyed upon adults. In this region, the dry/cold season may be so difficult for survival that early maturation at small size would be advantageous. The male-biased sample may be related to the greater motility of males. The rapid abandonment of the foam by the tadpoles and communal nesting may help to reduce predation by maggots. Skin toxins were not effective in deterring predation upon adults. The call of P. cf. fuscomaculatus differs from that of other species in the biligonigerus group and can help to define the specific status of the studied population.     

Reproductive aspects of Leptodactytus hylaedactylus (Anura: Leptodactylidae), a member of the Leptodactylus marmoratus species group,with a description of tadpoles and calls

In this paper we describe some aspects of the reproductive biology of Leptodactylus hylaedactylus from open areas of Central Amazonia, Brazil. We describe the calling site, reproductive season, daily pattern of calling activity, chamber structure, vocalizations and tadpole morphology. Males of Leptodactylus hylaedactylus called amidst grasses and bushes in open and urban areas, throughout the year, and a greater number of males were heard in the period from 16:00 to 20:00 h. The eggs are deposited in underground chambers, which are spherical or elliptical. Larval development occurs inside the chambers. Two types of vocalizations were detected, the advertisement and the territorial call. The advertisement call of the species studied herein differs from other calls described for L. hylaedactylus from other localities of Brazil, but is very similar to that described from Peru. Tadpole morphology is similar to that described from individuals in French Guiana.     

Reproductive features of three maternal-caring species of Leptodactylus (Anura: Leptodactylidae) with a report on alloparental care in frogs

We describe aspects of the reproduction of three shoaling/maternal-caring Leptodactylus species (Leptodactylus aff. latrans, Leptodactylus podicipinus and Leptodactylus aff. leptodactyloides), pointing to the relevance of the female to tadpoles and describe a case of alloparental care in frogs. Females of the three species connected water bodies by digging channels to their tadpoles. Leptodactylus aff. latrans females often expelled predatory snakes and conspecific males that approached shoals to prey upon tadpoles. In water bodies containing predatory teleosts, tadpoles of L. aff. latrans only reached metamorphosis in the presence of guardian females. Channel digging can provide tadpoles with access to new feeding grounds and prevent predation and desiccation. We found brood mixing and alloparental care between L. aff. latrans and L. podicipinus which, as in some Teleostei, may be regarded as the result of identification mistakes.     

Natural history of Phyllomedusa megacephala (Miranda-Ribeiro, 1926) (Anura: Hylidae) in southeastern Brazil,with descriptions of its breeding biology and male territorial behaviour

The breeding biology and male territorial behaviour of the large-headed leaf-frog (Phyllomedusa megacephala) were studied in three temporary altitudinal streams located in the Cerrado biome of southeastern Brazil, during three consecutive years (2007–2009). Phyllomedusa megacephala showed significant sexual dimorphism, with females being bigger and heavier than males. Acoustic and physical interactions between males defending or not defending territories including amplectant and non-amplectant males, patrolling behaviour by males without established territories, and triple amplexus were observed. Neither males nor females were observed using visual cues when searching for mates. We found that males that were present at breeding sites in previous years bred more successfully and maintained territory better.     

Vocalization and territorial behaviour of Phyllomedusa nordestina Caramaschi, 2006 (Anura: Hylidae) from southern Bahia,Brazil

Territoriality is common among amphibians with a prolonged breeding season and can be expressed through the defence of territories with vocalizations and aggressive interactions. The present study aims to describe the territorial behaviour of male Phyllomedusa nordestina, test whether the physical characteristics of males influence the results of disputes, and describe the vocalizations of the species. The study was conducted at temporary and permanent ponds located in the Ecological Reserve Michelin, Igrapiúna, Bahia, Brazil. Male P. nordestina defended territories and showed aggressive behaviour, consisting of vocal disputes, chasing and physical fighting. Winners were not bigger, heavier, or in better condition than the losers. The victory may be associated with the vocalization activity. Three types of vocalizations were recorded: the advertisement call, composed of one or two multipulsed notes, and two types of territorial calls. We also report on male patrolling behaviour and habitat use.     

The reproductive ecology of Leptodactylus fuscus (Anura,Leptodactylidae): new data from natural temporary ponds in the Brazilian Cerrado and a review throughout its distribution

Recent studies indicate that populations historically called Leptodactylus fuscus (Schneider, 1799) comprise at least three well‐differentiated lineages. Herein, we describe the reproductive ecology of a southeastern clade population of L. fuscus, and review the characters of the reproductive ecology for several populations of the northern and southeastern clades. Most reproductive activity occurred in December and January, which coincided with the highest rainfall period. Males had an aggregated spatial distribution in the central area of the ponds. Three courtship interactions were observed. A female was observed closing the entrance to a burrow with moist sand after oviposition. This behaviour is described for the first time in L. fuscus. The review of the reproductive biology of the northern and southeastern clades indicates a relatively high plasticity in L. fuscus. Furthermore, the results corroborate the suggestion, based on molecular data, that the northern and southeastern clades of L. fuscus represent distinct evolutionary units.     

Development and demography of Phasmahyla jandaia (Bokermann and Sazima, 1978) (Anura,Hylidae) tadpoles in an Atlantic Forest site,southeastern Brazil

Mortality rates are often high during the anuran tadpole stage, making demographic parameters in this phase of their life cycle important to the understanding of population demography. Nevertheless, information on mortality rates or population size is lacking for most Brazilian anurans. Herein we report on the demographic parameters of Phasmahyla jandaia (Hylidae) during the larval stage, including number of individuals through time, mortality rates and time of recruitment. Tadpole recruitment occurred year‐round, although froglet recruitment to the terrestrial habitat was restricted to the warm, rainy season. This trend seems to result in high plasticity of the tadpole developmental period. Mortality rates decreased with advancing tadpole stage, totalling 77% during the whole aquatic phase. Survivorship can be regarded as intermediate compared with other anurans for which data are available, and is in accordance with a high investment in egg size and the low number of eggs laid by P. jandaia.     

On the natural history of the South American pepper frog,Leptodactylus labyrinthicus (Spix, 1824) (Anura: Leptodactylidae)

Some species of Leptodactylus of the L. pentadactylus group lay their eggs outside water but the tadpoles need to reach water to complete the larval phase; other species complete development in terrestrial nests. Here we present details of the reproduction of L. labyrinthicus in south‐eastern Brazil. The proportion of tadpoles and trophic eggs in aged egg clutches was determined, as well as the growth of the tadpoles while in the nest. The gut contents of tadpoles that were in egg clutches of frogs were analysed. Adult males did not differ from females in size and had hypertrophied forearms and an enlarged spine on the thumb. Reproduction was initiated with the first rains of August/September and extended to mid‐January. Calling and spawning occurred at permanent or temporary water bodies. The foam nests were built in excavated basins outside of, but close to the water. The male determined the place of the basin construction; after amplexus, the female completed the excavation. The amplexus was axillary. One female spent the day after spawning in the foam. The eggs were pale grey, the yolk averaging 2.3?mm in diameter. The mean number of eggs was 2101 per egg clutch. The number of tadpoles in individual nests varied between 0.05% and 11.40% in relation to the total laid eggs. The tadpoles entered water when rains flooded the basin. The tadpoles grew to 12 times the weight of an individual egg while in the nest; no nesting tadpole was beyond stage 25. The longest time we followed tadpoles in a nest was 25 days. Tadpoles were found preying upon eggs of three other frog species and upon conspecific eggs. Males fought by grasping each other in a belly‐to‐belly position; the powerful arms and the thumb spines represent weapons. Even though males can reach maturity in the season following birth, small size would prevent them from establishing their own territory. All the species of the L. pentadactylus group may build their foam nests within excavated basins. The basins may protect the eggs and embryos from cannibalistic tadpoles and may have an anti‐desiccation effect. In order to produce trophic eggs, the female may delay laying additional unfertilized eggs until after the male has abandoned the foam nest. Anuran eggs represent an important food item for tadpoles after they leave the nest.     

Reproductive ecology of a Tibetan frog Nanorana parkeri (Anura: Ranidae)

Endemic to the Tibetan Plateau, Nanorana parkeri figures among the highest altitude amphibians in the world. The present work, which was conducted in an alpine marsh at 4300 m altitude, is the first report of the species’ reproductive ecology. The breeding season, as indicated by the presence of amplectant pairs and fresh eggs in ponds, lasted from early Mary to early July. The ratios of adult male to female were 1.3 in hibernation ponds and 2.3 in spawning ones. Males were smaller than females. Amplexus was typically axillary (81.7%), with a few exceptional amplexus in which a male clasped another male or object rather than gravid females. Scramble competition occurred in 6.8% of the normal amplexus located. The body sizes of amplexed couples correlated positively. Deposited eggs were attached to submerged plants solitarily (60.5%) or in small clusters (39.5%), and the scattered arrangement of individual eggs could be a strategy to ensure embryonic development in oxygen-poor environments. Clutch size averaged 189 [standard deviation (SD) = 64], and correlated positively with female body size; egg size averaged 2.18 mm (SD = 0.10), independent of female body size; both traits within a clutch were negatively related with each other even controlling for female body size. Relatively few and large eggs laid by the Tibetan frogs, compared with those by lowland aquatic-spawning anuran species, should be adaptive to the harsh high-altitude conditions. Larvae hatched from eggs after 16.4 days on average (SD = 5.8). Froglets emerged between late July and early August, but some tadpoles were present in winter. These findings may help to understand the evolution of amphibians in extreme environments and to protect these remote species.     

Aspects of the natural history of Leptodactylus knudseni Heyer, 1972 (Anura: Leptodactylidae) in a pristine forest in Central Amazonia,Brazil, with comments on ontogenetic variation of its tadpoles

The Leptodactylus pentadactylus species group is comprised of medium to large species of Neotropical frogs. Leptodactylus knudseni, a member of this species group, has a wide distribution throughout the Amazon Basin. Herein we describe aspects of the reproductive biology of L. knudseni and provide notes about the ontogenetic variation of its tadpoles based on a population in a non-flooded forest near Manaus, Amazonian Brazil. Amplectant pairs of L. knudseni lay foam nests in excavated basins on the edge of temporary ponds located on clay soil and at least 50 m from a stream. The tadpole development happens initially in the foam nests with access to the pond after the rain flooding the basins. Studied clutches lacked trophic eggs and tadpoles did not produce foam. Ontogenetic variations in L. knudseni tadpoles are related to size, teeth formulae and body colour. The use of excavated basins for the deposition of foam nests has been reported in several species of the L. pentadactylus group. The absence of trophic eggs and production of foam by the tadpoles differ from other species of the L. pentadactylus group. The tadpole morphology is similar to that described for other species of the group.     

Mating behaviour,territoriality and natural history notes of Phyllomedusa ayeaye Lutz, 1966 (Hylidae: Phyllomedusinae) in south-eastern Brazil

The mating (male courtship, amplexus and oviposition) and male territorial behaviours of Phyllomedusa ayeaye are described from a high-altitude site in the state of Minas Gerais, south-eastern Brazil. Information related to male breeding success and territoriality, as well as egg clutch parameters, is provided, together with qualitative information regarding tadpole phenology and juvenile recruitment. In addition, estimations of sexual dimorphism and numbers of marked individuals are available through capture-mark-recapture sampling. Females bred with males within spatially clustered oviposition sites (broadleaf plants). In some cases, the amplectant couple actively searched for the oviposition site. Males defended territories from other males, employing both acoustic and physical interactions. Some males successfully maintained their calling sites over successive nights, and others seemed to switch among nearby sites during successive nights and tried to disrupt ongoing ovipositions. No significant relationship was found between physical attributes (snout vent length or body mass) of males with breeding success. Also, no influence of the number of nights a male was active in chorus and its breeding success was detected. Therefore, it is proposed that the mating system in P. ayeaye may be opportunistic. Additional information related to reproduction (egg clutch parameters and breeding behaviours) is also discussed for other species of the Phyllomedusa hypochondrialis group and compared with the results of the present study.     

Tadpole deposition behaviour in male stream frogs Mannophryne trinitatis (Anura: Dendrobatidae)

Male Mannophryne trinitatis transport their larvae on their backs to predator‐free pools and deposit them there. The experiments reported here investigated M. trinitatis male deposition behaviour in containers placed near a heavily fish‐populated stream at Mount Saint Benedict, northern Trinidad. Choice of deposition site was not related to height above or distance from the stream. The low mean number of tadpoles deposited in each container in the field and in a laboratory experiment indicated that most males tended not to deposit complete clutches in single pools. Deposition experiments in the laboratory with a choice of different‐sized containers showed that deposition was pool size dependent, with more tadpoles deposited in larger containers. Deposition behaviour when given a choice of containers with leaf litter (a source of food and shelter) and with no leaf litter showed no significant preference. Tadpoles grown in different water volumes with different food sources revealed that tadpole growth was best in larger water volumes containing leaf litter. Regular tadpole inspections at a pool in the field established that most tadpoles present were the result of downstream movement.     

Reproductive behaviour and development dynamics of Odontophrynus cordobae (Anura,Cycloramphidae)

We provide detailed information on breeding sites, release call, satellite behaviour, courtship, and embryonic and larval development of Odontophrynus cordobae. Field observations were carried out during two breeding seasons (2004–2005 and 2008–2009). Clutches were raised in captivity (artificial ponds) to analyse the development of embryos and tadpoles. Breeding sites were represented by streams and their backwaters, as well as temporary and permanent ponds of varying sizes. Operational sex ratio was male-biased varying between 0.154 and 0.25. Acoustic activity started about 8–10 pm and ended between 2 and 4 am the next day. Distance among calling males was approximately three metres. Male satellite behaviour was observed. Amplexus attempts among males were common. Release calls were structurally similar although shorter than advertisement calls. Courtship and formation of amplexus were documented.     

Helminth biodiversity of Costa Rican Anurans (Amphibia: Anura)

One hundred and sixty-three frogs representing 21 species collected in Costa Rica were examined for helminths. Those found were: one species of Digenea (Mesocoelium monas), one species of Cestoda (Cylindrotaenia americana), 15 species of Nematoda, 11 species represented by gravid individuals (Aplectana hylambatis, Aplectana incerta, Aplectana itzocanensis, Capillaria recondita, Cosmocerca parva, Cosmocerca podicipinus, Falcaustra costaricae, Hedruris heyeri, Oswaldocruzia costaricensis, Oswaldocruzia nicaraguensis, Rhabdias savagei) and four species represented by larvae (Physaloptera sp., Porrocaecum sp., Spirocamallanus sp., Ascarididae gen. sp.) and three species of Acanthocephala, one species represented by gravid individuals (Anuracanthorhynchus lutzi) and two species by cystacanths (Centrorhynchidae gen. sp. and Oligacanthorhynchidae gen. sp.). Mean helminth diversity (number of helminth species) per frog species was 2.4 ± 0.3 SE, range: 1–6. Forty-nine new host records are reported. Costa Rican frogs are infected by generalist helminths, which infect other species of frogs from Central and South America.     

Oviposition site preference and reproductive ecology of Teratohyla midas (Anura: Centrolenidae) in the Colombian Amazon

ABSTRACT

The family Centrolenidae (Glass frogs) includes 157 species of anurans with a Neotropical distribution. Although glass frogs have been the subject of several studies encompassing various aspects of their biology, natural history data on their reproductive ecology is still scarce. One of the important aspects of reproductive ecology is the oviposition site since it can impact the reproductive success of a species. Herein, we provide detailed information on the reproductive ecology of a population of Teratohyla midas from the Colombian Amazon foothills and test whether this species shows a preference on substrate and height of oviposition site. We monitored a population using capture-mark-recapture of individuals identified by a photograph of their unique pattern of dorsal yellow spots. Wild-ID Software was used to corroborate if the dorsal pattern was different between individuals. Additionally, we monitored the presence of egg clutches, amplexus and oviposition behaviour. We observed 110 males and 5 females of T. midas, seen at an average height of 3.1 m ± 2.9 (0–17.8 m) from the ground. Likewise, we recorded the amplexus behaviour of two pairs for approximately 150 min. Once the oviposition happened, the male retreated, and the female covered the egg clutches for approximately 60 min. We observed 25 egg clutches with 60% being deposited in leaves of Selaginella sp and found at an average height of 2.9 ± 1.7 m (1.1–7.0 m). Thus, the selection of oviposition site was not random but mostly specific to one type of substrate. Furthermore, there was an overlap of the re-sampled median height distributions of active individuals and clutches, which suggests that oviposition does not occur at a preferred height within the vertical distribution of individuals. The information presented here is the first detailed record of reproductive ecology for this species and we hope to provide a basis to improve our knowledge on the biology of this species and other glass frogs.     

Reproduction,planula development,and substratum selection in three species of Stylactaria (Cnidaria: Hydrozoa) from Hokkaido,Japan

Modes of reproduction, morphology and behaviour of planulae, and substratum specificity were studied and compared in three species of the hydrozoan genus Stylactaria from Hokkaido, S. conchicola, S. uchidai, and S. multigranosi. Observed differences in these attributes provide additional characters for discrimination of the three species. Stylactaria conchicola is oviparous, its planulae are sedentary, and its hydroid is substratum specific (living only on shells of the gastropod Homalopoma amussitatum). Stylactaria uchidai differs in being ovoviviparous, in having crawling planulae, and in being a substratum generalist. Stylactaria multigranosi is parthenogenetic, its planulae are sedentary, and it is substratum specific (occurring exclusively on the gastropod Nassarius multigranosus). Only female colonies of S. multigranosi are known, and gonophores of this species differ from those of S. conchicola and S. uchidai in being eumedusoid rather than styloid. Correlations were noted between attributes of the planula larva and substratum specificity in the three species. The cnidome differences between the species suggest that penetrant nematocysts are particularly important for sedentary planulae colonizing moving substrata. These characters are also considered useful in the taxonomy of species of Stylactaria. The phylogenetic significance of different gonophore types in the genus is considered.