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.     

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.     

On the natural history of Leptodactylus syphax with comments on the evolution of reproductive features in the L. pentadactylus species group (Anura,Leptodactylidae)

We describe the reproduction of Leptodactylus syphax from Brazil and attempt to trace the evolution of reproductive traits within the L. pentadactylus species group. We quantified the proportion of eggs/embryos from fresh nests. We test whether L. labyrinthicus and L. syphax tadpoles can prey on eggs from heterospecific foam nests. Egg clutches were found outside the water at the edges of rocky/seasonal streams, usually within excavated basins. Males beat the foam with their legs moving from side to side. Clutches do not have trophic eggs, since most of their eggs develop into tadpoles. Tadpoles cling on rocks in current/shallow waters. Only L. labyrinthicus tadpoles could prey on eggs from heterospecific foam nests. If the L. pentadactylus group is monophyletic, excavated basins and foam‐beating in lateral motions may be possible synapomorphies of this group. However, large adults, carnivorous tadpoles and trophic eggs appear to be associated synapomorphies of a less inclusive group.     

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 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.     

Reproductive patterns of Trachycephalus venulosus (Laurenti, 1768) and Scinax fuscovarius (Lutz, 1925) from the Cerrado,Central Brazil

Animal mating systems are moulded by ecological factors, mainly the temporal patterns of reproduction. We report the reproductive period, use of microhabitats, clutch and egg sizes, reproductive efforts, occurrence of sexual size dimorphism, and the relation between body size and fecundity in the gravid females of Trachycephalus venulosus and Scinax fuscovarius at Serra da Bodoquena, Central Brazil. The reproduction of both species was correlated with the rainy season. Females of T. venulosus produced more eggs and their eggs were larger than those of S. fuscovarius. The females of both species were larger and heavier than males. The reproductive effort was measured as the ratio of gonad mass in relation to body mass. Females of both species presented similar reproductive efforts, but males of S. fuscovarius invested more in gonads compared to males of T. venulosus. Trachycephalus venulosus presented significant correlation between snout–vent length versus number of mature eggs and body mass versus number of mature eggs and ovarian mass. Scinax fuscovarius showed significant correlation between body mass versus ovarian mass. Trachycephalus venulosus is an explosive breeder, whereas S. fuscovarius has prolonged reproduction. These species are able to adapt to several types of habitat, due to behavioural and physiological plasticity.     

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.     

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.     

Ecological aspects of the pumpkin toadlet,Brachycephalus garbeanus Miranda-Ribeiro, 1920 (Anura: Neobatrachia: Brachycephalidae), in a highland forest of southeastern Brazil

Brachycephalus garbeanus is an aposematic toadlet, endemic to the Atlantic Forest in the mountains of the state of Rio de Janeiro, southeastern Brazil. Information regarding the ecology of species in this genus is scarce, with no information on most species, including B. garbeanus. We provide ecological information about B. garbeanus, in particular regarding microhabitat use, sexual dimorphism in body size, and feeding habits. Our study was conducted in a montane area of the Parque Estadual dos Três Picos, the largest State park in Rio de Janeiro. Most of the B. garbeanus in the study area were found on the forest floor leaf litter. Females were larger on average than males. Brachycephalus garbeanus consumed a wide array of prey, comprising exclusively invertebrates and predominantly arthropods. There were seasonal differences in the diet composition, but mites and ants were important in both wet and dry seasons.     

Reproductive ecology of the glass frog Espadarana prosoblepon (Anura: Centrolenidae) in an urban forest of the Central Andes of Colombia

Population and reproductive behaviour aspects of the glass frog Espadarana prosoblepon are well known for populations from Central America, but similar information is lacking for populations from South America. We recorded the reproductive ecology of a population of the glass frog E. prosoblepon in the city of Armenia, Central Andes of Colombia. With weekly surveys with mark-recapture between December 2013 and November 2015, we monitored activity patterns and evaluated if the probability of males mating is associated to their body size or to chorus tenure (i.e. the number of nights a same individual is calling for attracting a mate). In addition, upon observing an amplexus, we recorded the behaviour of the individuals until oviposition and noted characteristics of egg clutch. We recorded 47 males and 32 females, nine amplexus events, eight ovipositions, and 21 egg clutches (35.4 ± 4.79 eggs per clutch, N = 29). Activity of males and females and the number of egg clutches were positively correlated to rainfall. Mating success of males was not associated to their body size, but it was positively associated to longer chorus tenure; clutches from large females had a higher number of eggs than clutches from small females. Oviposition occurred on average 3.76 ± 1.74 hours after we first observed the amplexus, occurring 93.1% of the time in fronds of Selaginella geniculata at an average height of 1.58 ± 0.44 m. After the oviposition, the male left the site, while the female remained between 30 and 45 min, partially covering the eggs. The reproductive behaviour of E. prosoblepon did not vary widely between a population in Costa Rica and our population in time to oviposition, and mating success of males associated to chorus tenure; however, in our study population we recorded a larger clutch size and the preference for S. geniculata fronds as oviposition substrate.     

Natural history of the lizard Enyalius perditus (Squamata: Leiosauridae) from an Atlantic forest remnant in southeastern Brazil

The natural history of the lizard Enyalius perditus was studied from August 2005 to July 2006, at Parque Estadual Nova Baden, in the municipality of Lambari, State of Minas Gerais, south-eastern Brazil. We found that E. perditus may be considered a semi-arboreal species, like other species in the genus, sleeping on vegetation (slim branches or large leaves) at night, possibly to minimize predation. Females with enlarged follicles or oviducal eggs were found from November to January and juveniles recruited at the end of the dry season and beginning of the rainy season, in October. The diet of this lizard is composed mainly of Lepidoptera larvae, Araneae and Formicidae. Females were larger than males, and juvenile coloration is similar to that of adult females.     

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.     

Natural history of the marsupial Thylamys macrurus (Mammalia,Didelphidae) in fragments of savannah in southwestern Brazil

The objective of this study was to furnish information on the natural history of the Neotropical marsupial Thylamys macrurus in the southern Cerrado of Brazil. A range expansion of the species was observed to the northeast of the Mato Grosso do Sul state, Brazil. Specimens were surveyed in fragments of savannah (40–600 ha) during dry and wet seasons in 2003 and 2004 with live‐trap grids and pitfall lines. With an effort of 9767 trap nights and 2200 pitfall nights, we obtained 102 captures and 49 recaptures of T. macrurus. The data analyzed (57 males and 40 females) showed that males are more abundant in the dry season and females in the wet season. Young were recorded mostly during the wet season. Thylamys macrurus was scansorial, using the floor and the understorey. Sexual dimorphism was recorded, with males being larger than females. Information on ectoparasites is supplied.     

Mating behaviour in the bees,Dieunomia heteropoda and Nomia tetrazonata,with a review of courtship in Nomiinae (Hymenoptera: Halictidae)

Males of many bees in the subfamily Nomiinae (Halictidae) have numerous secondary sexual characters, which provide much systematic information. Similarly, the use of these structures by males during sexual behaviour may provide useful characters, but the courtship and copulatory behaviour of most of these bees is not known. Structures and behaviour used for courtship and mating are described for two species of nomiine bees, Dieunomia heteropoda and Nomia tetrazonata, along with brief observations of mating in a parasitic bee, Triepeolus verbesinae (Apidae: Nomadinae). A review of mating behaviour within the Nomiinae shows no obvious phyletic patterns, based upon present limited knowledge. The species-specific nature of the male secondary sexual characters suggests they may have evolved in the context of sexual selection, but behavioural data from populations of additional species are needed to evaluate the hypothesis adequately.     

Morphological variation of Eucyclops elegans (Herrick, 1884) (Copepoda: Cyclopoida) in the Americas and comments on records of Eucyclops conrowae Reid, 1992

Based on an examination of material from different areas of the American continent, we evaluate the morphological variations of Eucyclops elegans (Herrick, 1884) with an emphasis on characters used in the current taxonomy of the genus. Eucyclops elegans is clearly a member of the serrulatus group. Differences in both female and male specimens from North America and South America and the lack of records in southern Mexico and Central America suggest that it is possible that this nominal species contains at least two taxa. Characters such as the ornamentation of the antennal basis and coxa of P4 are described for the tropical species E. conrowae Reid, 1992. This species is clearly not a member of the serrulatus group and differences with respect to the type specimens indicate that records of E. conrowae in Mexico do not correspond to this taxon and should be reassigned to other species.     

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.     

Life history of Paraselenis (Spaethiechoma) dichroa (Germar, 1824) (Coleoptera: Chrysomelidae: Cassidinae) in natural conditions of Atlantic Forest from Brazil

Cassidinae Gyllenhaal? is the second largest subfamily of Chrysomelidae (Coleoptera), which presents characteristic morphology, biology and behaviour. The current study describes the life cycle of Paraselenis(Spaetiechoma) dichroa (Germar), ?determining biological and behavioural aspects as well as the action of natural enemies on populations, when the species has maternal care. The study was conducted between February and April 2015 in the National Forest of Passa Quatro, municipality of Passa Quatro, Minas Gerais state, Brazil. Daily observations were made (morning and afternoon) to evaluate the offspring of females on Ipomoea sp. and Merremia macrocalyx (O’Donell?) (Convolvulaceae), as well as the actions of natural enemies on the young. The eggs are deposited on the midrib and abaxial surface of leaves (n = 25). Egg clusters are pedunculated, arranged in groups and devoid of any cover. On average, each female produced 27.3 ± 5.6 (n = 16) eggs, with a mean incubation period of 12.7 ± 2.9 days (n = 10 females with their young). The larvae remained grouped in the leaves throughout development, except when they fed. They retained faeces and exuviae as a stacked faecal structure on their mobile urogomphi (caudal process) like a faecal shield. The mean number of larvae per female was 12.7 ± 10.1 (n = 19) and the larval period lasted 24.9 ± 4.5 days (n = 12). Pupation occurred on the stem of the plant (n = 16). On average there were 8.1 ± 8.5 individuals for pupae progeny (n = 14). The mean duration of the pupal stage was 10.4 ± 3.3 days (n = 5). Overall, 3% of adults emerged (n = 13 individuals), with a total cycle time of 41.3 ± 8.4 days (n = 6). Adults are sexually dimorphic in the elytral shape. Females remain with the offspring throughout development. The natural enemies of immatures were Emersonella pubipennis Hansson? (Hymenoptera: Eulophidae), Tachinidae (Diptera) and Stiretrus decastigmus (Herrich-Schaeffer?) (Heteroptera: Pentatomidae), Conura sp. Spinola? (Hymenoptera: Chalcididae) and Brachymeria sp. Westwood? (Hymenoptera: Chalcididae). The information presented here on the natural history of P. dichroa are important as they may serve as a starting point to understand evolutionary questions and multispecies interactions.     

Aspects of natural history in a sand boa,Eryx muelleri (Erycidae) from arid savannahs in Burkina Faso,Togo, and Nigeria (West Africa)

The ecology of West African arid savannah snakes is still poorly known, especially in regard to the fossorial species. Here, field data on distribution, morphometrics, habitat use and diet are reported for the sand boa Eryx muelleri (Erycidae) from three different countries, i.e. Burkina Faso, Togo and Nigeria. This snake was observed in 21 distinct localities, and a total of 39 individuals was recorded. The species is likely fairly common and locally abundant. Mean body length of snakes was similar among countries and between sexes, but males had proportionately longer tails than females. Body length was significantly positively correlated with both tail length and head length. Adult sex ratio was close to parity. Most of the specimens were found in well-vegetated spots with low percentage of bare soil. There were apparently both ontogenetic and intersexual dietary variations: (1) a single analysed juvenile fed upon a small-sized lizard, whereas adults fed considerably upon young rodents; and (2) females fed exclusively upon young rodents and males upon both lizards and young rodents. In conservation terms, the species is actively exported from Togo for the pet trade, and additional individuals may also be illegally collected in other countries.