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.     

Breeding biology of Phyllomedusa azurea Cope, 1862 and P. sauvagii Boulenger, 1882 (Anura) from the Cerrado,Central Brazil

Phyllomedusa azurea and P. sauvagii are treefrogs from seasonal dry environments of Central Brazil. We report on reproductive and other natural history features of these species. The study was carried out in Serra da Bodoquena, Mato Grosso do Sul, Brazil. The reproductive period was determined by the presence of vocalizing males, amplectant pair or presence of tadpoles. The reproductive effort was measured as the ratio of gonad mass in relation to body mass (BM). The reproduction of both species was correlated with the rainy season. Female P. sauvagii produced more eggs and their eggs were larger than those of P. azurea. Both species showed sexual dimorphism in relation to the snout–vent length (SVL) and BM. The mating behaviour of both species was similar. Males in both species showed similar reproductive efforts, but females of P. azurea invested more in gonads compared to the males of P. sauvagii. Phyllomedusa azurea showed a significant relation between SVL versus number of the mature eggs (NME), and P. sauvagii between SVL versus NME and BM versus ovarian mass.     

Breeding behaviour of Bokermannohyla nanuzae (Anura: Hylidae) at an Atlantic Forest site in southeastern Brazil

We studied the breeding activities of Bokermannohyla nanuzae at an Atlantic Forest site in southeastern Brazil from August 2010 to April 2012. We described courtship behaviour, acoustic parameters of calls, egg clutch features and the egg-laying site, and characterized the reproductive mode. We recorded calling males and gravid females almost year-round, except for a few months in the dry season. We observed the interaction of five male–female pairs that exhibited stereotyped behavioural sequences including visual and tactile signals. We identified three different types of calls that we considered as the advertisement call and two types of courtship calls. Egg clutches were placed in rocky crevices with water or in streamside puddles, among rocks, that would be likely to be flooded as stream water level raises, which characterizes a reproductive mode different from the one previously recognized for this species. The breeding biology of B. nanuzae is complex, with elaborate courtship behaviour and signalling.     

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.     

Reproductive ecology and behaviour of a species of Adenomera (Anura,Leptodactylinae) with endotrophic tadpoles: Systematic implications

The species of the Leptodactylus fuscus group and those of the genus Adenomera lay their eggs in underground chambers. According to current systematic arrangement of these genera, this behaviour is convergent. Here we describe reproduction, courtship, tadpole morphology, calls, chamber structure, and populational phenology of an Adenomera species, and compare some of these features with those of other species of the genus and with species of Leptodactylus of the fuscus and pentadactylus groups. We tested the tadpoles of the Adenomera sp. and those of Leptodactylus labyrinthicus (pentadactylus group) for their foam‐making abilities. There was sexual dimorphism in size; males averaged 22.0?mm and females 24.3?mm in snout–vent length. Males called from late September to late February; calling often began about 2–3?h before sunset. Males called exclusively in open (non‐forest) areas. Egg clutches were found around male calling sites. Males excavated spherical chambers which had a direct entrance. During mating, the male led the female towards a previously excavated chamber. Territorial interactions (aggressive calls and fights) occurred when an intruder male approached a calling male. Late‐stage tadpoles and newly metamorphosed juveniles were found within underground chambers. Clutch size averaged 10.6; eggs averaged 3.7?mm in diameter and were cream coloured. Tadpoles had horny beaks, but no labial tooth; spiracle was present. All tested groups of tadpoles generated foam within 10?h. As we report for Adenomera sp., the males of Leptodactylus species of the fuscus group lead the female to a previously excavated chamber. Tadpole foam‐making behaviour was reported in Leptodactylus of the fuscus group and was previously unknown for any species of Adenomera or for Leptodactylus of the pentadactylus group. The way in which the tadpoles of Adenomera sp., Leptodactylus spp. (of the fuscus group) and L. labyrinthicus made foam was quite similar. The foam‐making behaviour of the studied tadpoles may act to avoid compaction of the tadpoles at the bottom of the basin or underground chamber, avoiding overcrowding and increasing respiratory and excretory efficiency. As presently recognized, the phylogenetic position of Adenomera suggests that reproductive major behavioural features are convergent with some Leptodactylus species. Alternatively, our data point to close phylogenetic relationships between Adenomera and Leptodactylus of the fuscus and pentadactylus groups, reinforcing the paraphyletic nature of the genus Leptodactylus.     

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.     

Breeding biology of the rapids frog Limnomedusa macroglossa (Anura: Cycloramphidae) in southern Brazil

Limnomedusa is a monotypic genus that occurs in association with rocky outcrops and stream beds. Based on observational and mark–recapture fieldwork from 2005 to 2008, we report the breeding biology of Limnomedusa macroglossa in southern Brazil. The reproductive season lasted from late August to early February. Tadpoles were recorded from early September to late February. A clear pattern of emergence/recruitment was observed in juveniles. The occurrence of reproductive activities was clearly related to the longest photoperiods, when the highest temperatures occur. Males called from rocky or concrete substrates, mostly under rocks. Amplexus was axillary and the operational sex ratio was nearly even. Spawn occurred in lentic water bodies but tadpoles also completed their development in slow‐flowing water. Although using similar habitats for reproduction, L. macroglossa reached lower levels of specialization toward terrestriality than did the cycloramphids Cycloramphus and Thoropa. We classify L. macroglossa as a breeding habitat specialist that would be threatened by river damming.     

Reproductive site selection in the leaf-frog Phyllomedusa azurea Cope, 1862 (Anura: Hylidae) in altered areas of the Brazilian Cerrado

The selection of reproductive sites by anurans may be associated with each species’ behavioural restrictions, reproductive modes, morphology and physiology. We investigated the use of reproductive sites by Phyllomedusa azurea in an area of Cerrado, Central-Western Brazil, which has been modified by agricultural and cattle ranching. Reproductive activity was studied during the peak of the rainy season in two consecutive years. No correlation was found between male body size and position of the calling site, and calling sites did not vary among ponds. Conversely, the height and distance from the water margin where clutches were found varied among ponds. Phyllomedusa azurea used mainly Brachiaria sp. and Brachiaria humidicola grasses as calling sites and the grass Brachiaria sp. and the shrub Melochia villosa (Malvaceae) as oviposition sites. Calling males used the vegetation according to its availability; however, individuals seemed to exhibit significant preferences regarding the type of vegetation used for oviposition.     

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.     

Breeding biology of Ameerega braccata (Steindachner, 1864) (Anura: Dendrobatidae) in the Cerrado of Brazil

Reproductive aspects of Ameerega braccata, a Cerrado endemic species of Dendrobatidae, were studied in the surroundings of the type locality of the species, Chapada dos Guimarães, mid-southern Mato Grosso State, Brazil. We found that males call mainly early in the morning and in the late afternoon. During courtship, the female approaches a calling male and tactile and visual communication occurs. Males call continuously while guiding females to the oviposition site, which may be situated as far as 24 m (in a straight line) from the initial encounter site. Spawn are deposited over fallen leaves on the soil surface or in burrows. Female snout–vent length and body mass did not explain the variation in ovary mass, or the variation in the number and size of post-vitellogenic oocytes. The results reported here for A. braccata might help to understand trends in the evolution of dendrobatids in open Cerrado environments.     

Spatial niche variation in two sympatric species of Bokermannohyla (Anura: Hylidae) in southeastern Brazil

We studied patterns of microhabitat use by adults of two sympatric Bokermannohyla species at the Reserva Particular do Patrimônio Natural Santuário do Caraça, southeastern Brazil. We selected three streams, one where both species occurred in syntopy and the other two where each one occurred alone. We sampled 150-m transects in each stream throughout 1 year, recording microhabitat features for each frog located (substrate type, height and distance from water). Microhabitat availability varied between dry and wet seasons in all streams, and overall microhabitat diversity changed in two streams. Bokermannohyla nanuzae seemed to have a niche contraction in the presence of Bokermannohyla martinsi, but this only happened during the dry season. Microhabitat requirements during the wet season may be closely linked to similar reproductive needs that probably represent a strong selective pressure, forcing niche overlap.     

Gastrointestinal helminths from fifteen species of Litoria and Nyctimystes (Anura: Hylidae) from Papua New Guinea

Three hundred and ninety‐nine individuals representing 10 species of Litoria and five species of Nyctimystes (Hylidae), L. arfakiana, L. bibonius, L. eucnemis, L. genimaculata, L. impura, L. infrafrenata, L. modica, L. nasuta, L. thesaurensis, L. wollastoni, N. gularis, N. kubori, N. papua, N. pulcher and N. semipalmatus collected in Papua New Guinea were examined for helminths. Gravid individuals representing one species of Monogenea (Parapolystoma bulliense), one species of Digenea (Mesocoelium monas), 18 species of Nematoda (Aplectana krausi, Aplectana macintoshi, Aplectana zweifeli, Cosmocerca novaeguineae, Falcaustra papuensis, Icosiella papuensis, Maxvachonia adamsoni, Maxvachonia ewersi, Meteterakis crombiei, Moaciria moraveci, Ochoterenella papuensis, Oswaldocruzia bakeri, Parapharyngdon maplestoni, Parathelandros allisoni, Parathelandros andersoni, Physalopteroides milnensis, Rhabdias australiensis, Seuratascaris numidica) and one species of Acanthocephala (Acanthocephalus bufonis) were found. Also included were immature forms representing one species of Cestoda (as a metacestode), three species of Nematoda (Abbreviata sp., Rictulariidae gen. sp., Ascarididae gen. sp. in cysts), one species of Acanthocephala (unidentified cystacanth), and one species of Pentastomida [Raillietiella sp. (nymph)]. The mean helminth species richness for infected hosts was 1.3±0.6 SD (range one to three species). Sixty‐seven new host records and two new locality records are reported.     

Helminths of ten species of Litoria frogs (Anura: Hylidae) from Papua New Guinea

One hundred and ninety-one frogs collected in Papua New Guinea from 10 species of the genus Litoria, namely, L. angiana, L. arfakiana, L. caerulea, L. congenita, L. eurynastes, L. gasconi, L. infrafrenata, L. iris, L. kumae and L. modica, were examined for helminths. Eighteen species of helminths were found: one species of Monogenea, Parapolystoma bulliense; three species of Digenea, Diplodiscus amphichrus, Halipegus zweifeli and Mesocoelium monas; 13 species of Nematoda, gravid specimens of Aplectana macintoshii, Cosmocerca novaeguineae, Falcaustra papuensis, Maxvachonia adamsoni, Maxvachonia flindersi, Maxvachonia ingens, Meteterakis crombiei, Moaciria moraveci, Parathelandros mastigurus, Physalopteroides milnensis, Pseudorictularia dipsarilis, Seuratascaris numidica, and larvae (in cysts) of Abbreviata sp. and, one species of Acanthocephala, Pseudoacanthocephalus bufonis. Mean number of helminth species per host species was 3.0 ± 1.4 SD (range 1–5), mean number of helminth species per infected frog was 1.1 ± 0.4 SD (range 1–2), and mean number of helminths per infected host was 33.3 ± 5.8 SD (range 1–36). Twenty-eight new host records and two new locality records are reported. The biogeography of the recovered helminths is discussed.     

Spatial and temporal organization in three syntopic species of the Scinax ruber group (Anura: Hylidae) in the Atlantic rainforest,southeastern Brazil

Coexistence of anuran species might lead to acoustic interference among communications, especially where species are closely related. This problem might be minimized through resource partitioning. To test this idea, we analysed calling with respect to temporal and physical space usage in three syntopic, closely related species (Scinax cuspidatus, Scinax aff. hayii and Scinax aff. x-signatus) in a temporary pond in the Atlantic rainforest. The chorus attendance of each species resulted from a different combination of factors in the rainy season. Although there was some temporal overlap of daily activity among species, peak activity time differed, particularly between the two species whose calls were most similar. Although all three species tend to aggregate in open microhabitats, their coexistence is apparently facilitated by differential use of vertical strata and organization of calling sites near the sites of conspecifics. Together, these factors apparently reduce acoustic interference among heterospecific calls.     

Age,body size and growth pattern of Argenteohyla siemersi pederseni (Anura: Hylidae) in northeastern Argentina

In this study we analysed the body size, age structure, age at maturity, longevity and growth pattern of Argenteohyla siemersi pederseni in north-eastern Argentina using skeletochronological methods. Body size was sexually dimorphic; females were significantly larger and heavier than males, regardless of age. As A. s. pederseni is an explosive breeder with only one reproductive episode per year, we assumed that each Line of Arrested Growth (LAG) observed was equivalent to one period of decreasing growth, LAGs reflecting a direct estimation of individual age. The minimum number of LAGs counted was two for males and three for females, but there was no sexual difference in longevity. The von Bertalanffy's growth coefficients (K) were higher in males than in females for both size and body mass. Females had a lower growth rate than males, took longer to reach the minimum maturation size, and were older and larger at reproduction.     

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.     

Vocalizations of the Brazilian torrent frog Hylodes heyeri (Anura: Hylodidae): Repertoire and influence of air temperature on advertisement call variation

In this study we analysed the acoustic activity of Hylodes heyeri in areas of Atlantic Forest, in southern Brazil. Observations were made in November 2001 and from January to April 2002, at Estação II Instituto Agronômico do Paraná, municipality of Morretes, State of Paraná. Males displayed three different vocalizations: advertisement calls, territorial calls, and courtship calls. Temporal and spectral parameters of advertisement and territorial calls were analysed, along with call intensity. The duration of the advertisement call showed a negative correlation with air temperature, with males decreasing the duration of advertisement calls at higher temperatures. Male body size was not correlated with any of the acoustic parameters. Details on habitat use, distances to nearest vocalizing neighbour, and daily calling activity are also included.     

Geographic variation in body size and sexual size dimorphism in the giant spiny frog Paa spinosa (David, 1875) (Anura: Ranoidae)

We describe the spatial pattern of variation in body size and sexual size dimorphism in the giant spiny frog, Paa spinosa (David, 1875). The study also aims to identify potential factors responsible for geographic morphological variation in the frog. The results indicate that primary production, which positively correlates with precipitation and summer temperature in China, had an important effect on body size of P. spinosa, whereas Bergmann's rule was more efficient to explain the influence of average minimum temperature in January. The species was sexually dimorphic for body size and body shape: the body size, jaw length and forelimb length of males were significantly larger than those of females. Furthermore, the degree of sexual size dimorphism varied geographically, but the relationship between the geographic variation of sexual size dimorphism and the climate was complex. Specifically, absolute sexual size dimorphism was positively correlated with the increased ambient temperature and decreased precipitation.