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.     

Habitat,egg‐laying behaviour,eggs and tadpoles of four sympatric species of Pseudopaludicola (Anura,Leiuperidae)

In contrast to most leiuperid frogs, Pseudopaludicola falcipes does not lay eggs in foam nests; this could represent a reversion to the primitive state. We found that in four other Pseudopaludicola species, eggs were also not embedded in foam nests and had a well‐defined outermost jelly layer. The females also constantly moved or dived and males lacked vigorous “wiping/kicking” leg motions while eggs were being laid, features that make foam nesting difficult. The tadpoles of three species have two gaps in the marginal papillae along the lower labium and two posterior rows of labial teeth; the tadpoles of one species had three gaps and three rows, a pattern resembling that of some Physalaemus (Leiuperidae) species. Our data on tadpole morphology and reproductive behaviour do not allow us to rule out the paraphyletic nature of Physalaemus in relation to Pseudopaludicola but they provide some support against the “foam‐loss hypothesis” in Pseudopaludicola.     

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.     

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.     

Life History of Western Ghats endemic and threatened Anuran – Matheran leaping frog, (Indirana leithii) with notes on its feeding preferences

The species Indirana leithii (Anura: Ranixalidae) is endemic to the northern Western Ghats of India and is distributed between latitudes 16°N and 21°N in this region. The tadpoles of this species have been previously studied for their morphology and feeding behaviour; however, their natural history is still not completely understood. In the current study, we provide information about the mating behaviour, egg laying and metamorphosis of the species. We observed that the species possesses reproductive mode number 19 as these frogs lay eggs inside the natural cavities of vertical rock surfaces near streams and waterfalls and their tadpoles are adapted for adhering to the wet rock surface, where they feed on algae (including diatoms) growing in seepage on the rocks. Interestingly, there is no audible calling or territory guarding involved during the courtship behaviour. This has been observed for the first time in any Indian anuran. We also provide data on food preferences of adults and tadpoles.     

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.     

Comparative larval morphology of the European bone-skippers,Thyreophora cynophila (Panzer, 1798) and Centrophlebomyia furcata (Fabricius, 1794) (Diptera: Piophilidae), with notes on their coexistence and natural history

Considered at times as extinct, the European bone-skippers Thyreophora cynophila and Centrophlebomyia furcata (Diptera: Piophilidae) have been recently rediscovered in Spain, coexisting in some areas. Nevertheless, apart from their typical association with big mammal carcasses, little is known about them and the other species of subtribe Thyreophorina. In the present paper, the larval morphology of T. cynophila is described for the first time and compared with that of C. furcata. The morphology of cephalopharyngeal skeleton, the number and arrangement of the spines of ventral creeping welts, and the morphology and presence or absence of tubercles in the two last segments of the body, are the main characters allowing for larval identification of both species. Some preliminary field and laboratory observations on the behaviour and life cycle of the two species are also provided.     

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.     

Field and laboratory observations on reproductive aspects of Pseudopaludicola ameghini (Cope, 1887) (Leptodactylidae: Leiuperinae)

ABSTRACT

The genus Pseudopaludicola includes small-sized anurans, widely distributed throughout South America. Twenty-three out of the 25 species occur in Brazil. Although described more than a century ago, from mid-southern Mato Grosso state, central Brazil, Pseudopaludicola ameghini is poorly known. Here we describe the characteristics of reproductive site, amplexus, egg-laying behaviour, eggs and tadpoles of P. ameghini based on specimens and observations performed in the vicinities of the type locality. Pairs of P. ameghini reproduce in shallow and slow groundwater established on hydromorphic terrains, in open environments amidst the Brazilian Cerrado savanna. Reproductive activity occurred between 17:00 and 21:00 h at two sites studied from December 2013 to April 2014. Amplexus is axillary. Eggs are spherical, with a mean diameter of 5.1 mm (yolk plus jelly envelope). Oviposition occurs in shallow sites at a depth of nearly 2.0 cm. Females deposit their eggs one at a time, directly onto sediment at the bottom of waterbodies or attached to submerged vegetation. The tadpole body is oval in dorsal view and globular/depressed in lateral view; eyes are large and dorsally positioned. Nares are large, round, with a small apophysis on marginal rim, dorsally positioned, near the eyes. The spiracle is short, with posterodorsal opening. The oral disc is anteroventral, emarginated laterally, with one dorsal and two ventral gaps; the tooth row formula is 2(2)/2(1), and the upper jaw sheath is ‘arc’ shaped. Pseudopaludicola ameghini has a unique behaviour of oviposition among members of the genus. The differences in reproductive pattern and larval characteristics of P. mystacalis and P. ameghini reinforce the taxonomic validity of the latter, questioned until recently.     

Bionomics,behaviour and immature stages of Pelecotoma fennica (Paykull) (Coleoptera: Rhipiphoridae)

Ptilinus fuscus (Anobiidae) was confirmed as the host species of Pelecotoma fennica (Rhipiphoridae, Pelecotominae). Females of Pelecotoma oviposit into the wood infested by the host larvae. First-instar larvae are elongate, unsclerotized, very different from the triungulinid larvae known in other rhipiphorids. They search actively for the host larvae in the wood; no phoresy is involved in the life cycle. The first-instar larvae temporarily paralyse the host larva and enter its body, overwintering (some perhaps more than once) as an endoparasite. In the spring of the year of emergence, the endoparasite engorges enormously (without moulting) and develops a special sclerotized caudal structure which is then used for perforation of the host's integument. The larva undergoes a further four ectoparasitic instars. The fifth (i.e. fourth ectoparasitic) instar differs considerably from the preceding three, and is capable of boring through the wood to prepare the emergence gallery for the adult. Pupation occurs in the wood. The rate of parasitization may locally far exceed 50%. Superparasitization by the first-instar larvae is possible, but usually only 1 larva survives to the ectoparasitic stage. Larval morphology of Pelecotoma is described and illustrated. Additional data on bionomics and larval morphology are also presented for the genus Metoecus (Rhipiphorinae). Comparing the biology and larval morphology of Pelecotoma with other Rhipiphoridae, it is assumed that ancestral rhipiphorids may have been xylophilous Tenebrionoidea with predaceous or omnivorous larvae, and that the ‘triungulinid’ larvae and phoretic habits may not belong to the groundplan of the family Rhipiphoridae. The widespread opinion considering Rhipiphoridae closely related to the family Mordellidae is questioned.     

Revision of the Australian endemic genus Hyptiogaster Kieffer (Hymenoptera: Gasteruptiidae), with descriptions of seven new species

The Australian hyptiogastrine wasp genus Hyptiogaster Kieffer is revised. Among other characters, species of Hyptiogaster can be readily distinguished from other Hyptiogastrinae by having an exserted ovipositor and the anterior face of the mesothorax truncate in lateral view. Ten species are recognised including seven which are newly described: H. arenicola Turner, H. humeralis (Schletterer), H. rufus (Westwood), H. cobarensis sp. nov., H. hulli sp. nov., H. kalbarrii sp. nov., H. kiefferi sp. nov., H. naumanni sp. nov., H. pinjarregaensis sp. nov., H. weowaniensis sp. nov. Two synonymies are proposed; H. flavosignata (Kieffer) with H. humeralis and H. crassiceps (Schletterer) with H. rufus. An illustrated key to species is presented and relationships among genera of Hyptiogastrinae and species of Hyptiogaster are discussed.     

The strangest tadpole: the oophagous,tree-hole dwelling tadpole of Rhacophorus vampyrus (Anura: Rhacophoridae) from Vietnam

The tadpole of the vampire flying frog Rhacophorus vampyrus from Vietnam is one of the strangest tadpoles known. These tadpoles develop from non-pigmented eggs suspended in a foam nest placed on the wall of a tree hole. The elongate, depressed body resembles that of some phytotelmon-breeding frogs, but the mouthparts bear little resemblance to any other tadpole: upper labium reduced to one large papilla-like structure on each side, upper jaw sheath with a few huge, widely spaced, hook-shaped serrations that face backwards into the buccal cavity, lower jaw sheath absent, sinistral spiracle visible only ventrally, and two large, forward facing, keratinized hooks accompanied laterally by two similar sized fleshy papillae on the margin of the reduced lower labium. All evidence suggests that the tadpoles are oophagous and that the mother returns to deposit trophic eggs.     

A new species of Mantidactylus (subgenus Guibemantis) from Madagascar,with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae)

A new species of Mantidactylus (subgenus Guibemantis) from the rainforests of eastern Madagascar is described. The new species is sympatric with the closely related M. depressiceps (Boulenger, 1882) and M. tornieri (Ahl, 1929) but differs by larger size, a more strongly developed prepollex, and by advertisement calls. Mantidactylus acuticeps Ahl, 1929 and Rhacophorus mocquardii Boulenger, 1896 are confirmed to be junior synonyms of M. depressiceps. The presence of distinct femoral glands in the new species was noted; a feature so far not reliably recognized in other members of the subgenus Guibemantis. Femoral gland structure was investigated in representatives of each of the subgenera included in Mantidactylus by dissecting and reflecting the femoral skin. Four femoral gland types were identified. Type 1: many small, single granules in an ill-delimited patch, grouped in rosettes at the patch edges; type 2: a group of moderately sized granules in a well-delimited patch; type 3: comprising two different structures: distal granules enclosing externally a distinct central depression, and smaller proximal granules or groups of granules; type 4: as type 3, but lacking the proximal granules. Mantidactylus rivicola Vences, Glaw and Andreone, 1997, M. cf. malagasius (Methuen and Hewitt, 1913) and M. ventrimaculatus (Angel, 1935) differed from all other species examined; their femoral glands comprise a reduced number of large granules with an indistinct central depression in external view. The presence of a distal granule group with external central depression (types 3 and 4 above) provides a probable synapomorphy for the subgenera Brygoomantis, Ochthomantis, Hylobatrachus, Chonomantis and Mantidactylus. In females of these subgenera rudimentary femoral glands are present, but they lack completely in females of the other groups.     

Biology and immature stages of Panteles schnetzeanus (Hymenoptera: Ichneumonidae), a parasitoid of Lampronia fuscatella (Lepidoptera: Incurvariidae)

Panteles schnetzeanus (Roman), a parasitoid of the gall‐forming incurvariid moth Lampronia fuscatella (Tengström) and currently placed in the Stilbopinae, is shown to be a solitary koinobiont larval endoparasitoid. The large dark‐pigmented eggs possess a hook‐like tail which is embedded in host tissue, typically at the posterior or anterior end of the host. In cases of superparasitism, which represented 70% of cases, only a single egg hatches. The first instar, prepupa and final larval instar head capsules are described and illustrated. Many differences between the biology of Panteles and of Stilbops are discussed. Circumstantial evidence that suggests that Panteles may have an alternative host is also presented based on the emergence dates for it and the Lampronia together with the size, number and placement of the Panteles eggs. DNA sequence data additionally provide the first record of a eupelmid as attacking a gall‐forming incurvariid/parasitoid system, though it is not clear whether this was an hyperparasitoid on the Panteles or a primary on the incurvariid.     

Reproductive biology of two species of Hyalella Smith, 1874 (Crustacea: Amphipoda: Hyalellidae) from southern Brazil

Reproduction is a critical event in the life cycle of animals, and a high amount of energy is allocated to it. The aim of this study was to characterize the reproductive biology of two recently described species of Hyalella from the north-east region of state of Rio Grande do Sul in southern Brazil. The specimens were collected during 12 months from a stream source (Hyalella georginae) and from a water reservoir (H. gauchensis), in Palmeira das Missões municipality, state of Rio Grande do Sul. Upon sampling, ovigerous females and pairs in precopulatory behaviour were separated into microtubes. In the laboratory, the cephalothorax length (CL in mm) of all specimens was measured. Male size at onset of sexual maturity was larger than that of the females, in both species. There was a correlation between the size of paired males and females: larger males carried larger females, in both Hyalella species. Precopulatory pairs were found throughout the year; however, H. georginae had a higher frequency in spring, and H. gauchensis in autumn. There was a positive correlation between the CL of ovigerous females and the number of eggs/juveniles. A reduction in the number of eggs throughout the embryonic development was observed in H. georginae. The average fecundity of H. georginae was significantly higher than H. gauchensis. The differences between the reproductive biology of these species may be related to the physical and environmental conditions at which they are exposed in their habitats.     

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.