Home range behaviour in male and female poison frogs in Amazonian Peru (Dendrobatidae: Ranitomeya reticulata)

The poison frog Ranitomeya reticulata was studied for 3 months while mark–recapture surveys were performed. Ranitomeya species deposit terrestrial clutches and carry tadpoles to phytotelmata with few taxa performing biparental brood care including larval feeding. Home range size and spatial affinity to phytotelmata in the genus are linked to mating systems. In R. reticulata, individual home range size and overlap were similar in both sexes, indicating equal levels of site fidelity. Although territory defence was never observed, strong intrasexual intolerance within individuals' core areas was found. The large intersexual home range overlap for breeding pairs indicated that mate fidelity occurs. Individuals' home ranges were not overlapping, suggesting that R. reticulata lacks pair-bonding or strongly cooperative behaviour in parental duties. The number of ground bromeliads containing phytotelmata and home range size of males were positively correlated. Our findings suggest that females do not perform egg-feeding and male-only parental care is likely.     

Reproductive behaviour of the glass frog Hyalinobatrachium cappellei (Anura: Centrolenidae) in the Southern Amazon

Frog species of the Centrolenidae family exhibit a high variation in reproductive behaviours. Herein we describe the reproductive, ecological, acoustic and behavioural features of Hyalinobatrachium cappellei in Southern Amazon. Two hypotheses were also evaluated: (1) are ecological characteristics of the reproductive site related to male size and quantity of eggs and clutches? And (2) do males with one or more existing clutches at reproductive sites accumulate more new clutches than those males initially without them? This study was conducted at two streams, with sampling at 1 km each in Southern Amazon. Species reproduction occurred during the wet season and was considered prolonged breeding. Although male vocalization occurred on both the upper and underside of leaves, clutches were most often deposited on the underside of leaves. Males emitted a courtship call upon female approach. The process of female approach until oviposition lasted approximately eight hours. After oviposition, females left the breeding site while males remained near the clutch, covering it with the belly in ventral brooding behaviour. Thus, egg attendance is reported for the first time for this species. No intraspecific or interspecific aggressive behaviour was recorded. There was no relationship between habitat characteristics of the reproductive sites and male size, number of eggs and clutches. However, after 10 days of monitoring, we found evidences that parental males had a higher number of clutches than those males initially without clutches. Similar to other glass frog species, the behavioural characteristics of H. cappellei make this species an excellent model to use in evaluating the cost-effectiveness of parental care and the importance of egg attendance for offspring survival. Therefore, these data contribute towards a better understanding of the complex phylogenetic and biogeographic relationships between centrolenid lineages.     

Reproduction,development and parental care in two direct‐developing flatworms (Platyhelminthes: Polycladida: Acotylea)

Variations in reproductive modes, egg production, and the effects of parental care on hatching success are compared between Pleioplana atomata and Imogine zebra. Pleioplana atomata transfers sperm via true copulation, whereas I. zebra dermally impregnates spermatophores onto the dorsal surface of partners. Pleioplana atomata lays up to 750 large eggs over a 6‐week period, and I. zebra individuals lay up to 1346 small eggs in 12 days. Female fecundity is positively correlated with body size in both species. Developmental time lines are temperature‐dependent, and juvenile worms hatch after 3 and 6 weeks for I. zebra and P. atomata, respectively. Covering of egg masses by the adult is observed for both species and although this parental care is not necessary for egg development or hatching, it plays a significant role in the hatching success of P. atomata. In I. zebra, parental covering of egg masses may play a role in eggshell development.     

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.     

Male pheasant‐tailed jacanas commit infanticides to avoid cuckoldry when paternity of eggs is doubtful

Many hypotheses have been proposed to explain infanticide, killing the young or destruction of eggs by conspecific adults. We report cases of infanticide in the polyandrous pheasant‐tailed jacana (Hydrophasianus chirurgus) of Taiwan. In 2004, three males and one female bred at our study site. The female mated with the three males sequentially and laid clutches for all of them. Infanticidal behaviour was observed in all males and in all but one clutch, the first clutch of the breeding season. The infanticidal males threw out the first one or two eggs of a clutch, but then accepted replacement eggs and incubated normally. Because the intervals between copulation with different males were short (1–10 days) and sperm can remain viable within the female reproductive tract for at least 4–5 days, sperm competition among the males may have been intense, and the paternity of the first few eggs in each clutch was doubtful. We suggest that male pheasant‐tailed jacana may use infanticide as a strategy to avoid cuckoldry when the paternity of eggs is in doubt.     

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.     

The scramble competition mating system of the sphecid wasp Palmodes praestans (Kohl)

Males of the wasp Palmodes praestans engage in non‐territorial patrolling behaviour within a scramble competition mating system, showing great fidelity to their wide‐ranging patrolling routes without attempting to monopolize the areas covered. Scramble competition appears adaptive given that mating opportunities are not spatially aggregated in this species, whose females build isolated, single‐celled nests. Because of the female distribution pattern, a male's reproductive success in P. praestans probably depends upon the ability to find highly scattered, unmated females. The females of some close relatives of this sphecid wasp form fairly dense nesting aggregations. As mating system theory predicts, the males of some (but not all) of these species focus their searching at these concentrated nesting/emergence sites where potential mates are clustered spatially.     

Breeding behaviour and mating success of Phyllomedusa rohdei (Anura,Hylidae) in south‐eastern Brazil

Observations on the courtship behaviour, mating behaviour and mating success of a leaf‐frog, Phyllomedusa rohdei, population were conducted in a temporary flooded site of the Atlantic Forest, south‐eastern Brazil. We did not find any influence of size and body weight on male mating success across three scales: seasonal, nightly chorus and local groups (males found within 1?m of a mated pair). In addition, no evidence was found that female Phyllomedusa rohdei exhibit overt mate choice. The formation of mated pairs in P. rohdei occurred because males moved toward the females, rather than reverse. The only variable that was significantly related to male mating success was the number of nights of participation in chorus activity. Besides calling tactics, males showed active searching behaviour. There was no evidence of size‐ and mass‐assortative mating, and a high percentage of fertilization at all size and mass ratios of males and females was observed. Clutches in this species are produced in the same manner found in others members of Phyllomedusa; a purse‐like nest is made over water by folding a leaf around the egg mass (embryos and fluid‐filled capsules). Finally, we supply a list of several studies that examined the relationship between size or mass ratios of males and females in amplexus and its influence on fertilization rate, and show that size‐assortative mating in anurans is inconsistent (present in some species but not in other).     

Mating behaviour and maternal care in the tropical savanna funnel-web spider Aglaoctenus lagotis Holmberg (Araneae: Lycosidae)

Despite their abundance and diversity, Neotropical spiders have been the subject of relatively few studies of behavioural ecology. The sexual behaviour and maternal care of the spider Aglaoctenus lagotis (Lycosidae) is described here. We performed experimental manipulations to test the hypothesis that males are attracted by female cues in the web. Two predictions were tested: (a) webs used by a virgin female will attract more males than webs used by an inseminated female; (b) the main cue in the web is chemical, so webs recently used by virgin females will attract more males than those that are not recently used. Additionally, we tested the hypothesis that maternal care increases offspring survivorship. Results showed that males are attracted by recently used webs (z = 0.0015; p = 0.037; z-test) and webs of virgin females (z = 0.041; p = 0.036, z-test). Male sexual behaviour presented three different categories: court, pre-mate and mate. After the mating, the females built a cocoon that was transported adhered to the spinnerets and held by the last pair of legs. After the hatching of the eggs, the young migrated to the dorsal region of the mother's body for 5 days. During maternal care the mother eliminates a yellowish drop from its oral cavity, which is collected by the young. During the development of the young until adulthood, the offspring that have their mother present in the first stages of life have greater survival compared with those without maternal care. Our results make it possible to perform comparative analyses between a South American Lycosidae species and other species distributed worldwide.     

Extended parental care in two endobenthic amphipods

Extended parental care is described for two endobenthic amphipods which inhabit the estuarine soft-bottoms along the Gulf of Maine. The juveniles of both amphipod species, Leptocheirus pinguis and Casco bigelowi, remain in the burrow of their mother after they hatch from the brood pouch. Several consecutive clutches of L. pinguis can inhabit the female's burrow simultaneously, but in C. bigelowi there is always only one clutch of juveniles in the burrow of the female. Larger juveniles of L. pinguis start building their own small tubes at the bottom of the female's burrow. Casco bigelowi females increase the size of their burrows during the time when the juveniles grow. The juveniles of both species leave the female's burrow at about half adult size. The major tasks of female L. pinguis and C. bigelowi during the time of extended parental care is to irrigate and maintain the deep burrow. It is hypothesized that extended parental care in L. pinguis, C. bigelowi and other tube/burrow-living peracarids is a mechanism to protect small offspring from epibenthic predation.     

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.     

Natural history and behaviour of the primitively eusocial wasp Ropalidia marginata (Hymenoptera: Vespidae): a comparison of the two sexes

We report the natural history and behaviour of the primitively eusocial wasp Ropalidia marginata with a special reference to the males. We found that just as nests of this species are found throughout the year, so are the males. Females spend all their life in their nests but males stay in their natal nests only for 1–12 days and leave to lead a nomadic life. Males maintained in the laboratory can live for up to 140 days. Like all eusocial hymenopteran males, R. marginata males also do not perform any colony maintenance activities. We found that males did not forage or feed larvae. Compared with females, males showed fewer dominance and subordinate behaviours and being solicited behaviour and more feeding self and soliciting behaviours. By comparing males with young females, we found similar differences, except that the males showed similar rates of feeding self and higher rates of subordinate behaviour.     

‘On the rocks’: reproductive biology of the endemic toad Xanthophryne (Anura: Bufonidae) from the Western Ghats,India

Xanthophryne is a toad genus endemic to the northern Western Ghats of India, with two extant sister species – Xanthophryne koynayensis and Xanthophryne tigerina. Both species are local endemics and endangered. We studied reproductive biology of these toads and found that they are specialized to the lateritic rocky outcrops at mid-elevations in high rainfall areas. Xanthophryne toads have sporadic, multiple spawning bouts lasting 2–4 days during early monsoon. In this explosive breeding behaviour, we observed male toads to engage in ‘pelvic thrusts’, a unique and novel behaviour among anurans. Females oviposit in shallow pools in depressions of lateritic boulders where their tadpoles metamorphose. These ephemeral rocky pools have limited resources and they desiccate rapidly with a break in the rains. To mitigate the stochastic risk of desiccation and subsequent large-scale egg/tadpole mortality, females may disperse their reproductive investment spatially and temporally in multiple clutches, and tadpoles metamorphose rapidly. Here, we describe the amplexus, spawning and male advertisement call, and provide a comparative account of the life history traits of the two Xanthophryne species. These toads face numerous threats and are in need of urgent conservation action. These toads, seemingly well adapted to the isolated rocky outcrops, offer an excellent opportunity to understand endemism, mating systems, anuran ecology and behaviour.     

Expulsion and consumption of male ejaculates by promiscous female Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae)

While expulsion of male ejaculates by the females after copulation has been reported for various animal groups, expulsion followed by consumption of the expelled ejaculate is a rare behaviour outside spermatophylax-producing orthopterans. Among Diptera, this behaviour has been reported for a few species of Piophilidae, Empididae and Ulidiidae. Here we report on its occurrence among Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae). We also attempt to characterize the mating system of E. eluta in order to facilitate future hypothesis testing to understand the behavioural factors leading to the evolution of this peculiar behaviour. For this, courtship sequences, copulation duration, frequency of ejaculate expulsion and subsequent consumption, and latency to ejaculate expulsion for both E. eluta and E. mazorca were recorded. The time of sexual maturation, the time window of sexual receptivity during the day, and the mating frequency and variance in mating success for males and females (degree of polygamy) were determined for E. eluta. Both E. eluta and E. mazorca males engaged in elaborate courtship sequences involving visual and tactile displays before copulation. Females of both species almost invariably expelled and consumed ejaculates after copulation. Female E. eluta, required a 6- to 9-day period feeding on protein and sugar before becoming sexually receptive. Reproductive activity occurred continuously over the day with an early morning and late afternoon peak. Both males and females could mate multiply, with multiple partners over a 2-h observational period. Sexual network analysis revealed that some males and females had greater mating success than others. Ejaculate consumption appears to be widespread in the genus Euxesta. It is possible that females obtain nutrients from this behaviour while exerting some control over egg fertilization.     

Phoretic behaviour of Attacobius attarum (Roewer, 1935) (Araneae: Corinnidae: Corinninae) dispersion not associated with predation?

Attacobius attarum spiders exhibit a phoretic behaviour on the winged sexual of Atta leaf-cutting ants during their mating flight. However, it is unclear if this behaviour is for dispersion or to facilitate the predation of ants in the new colonies. A nest of Atta sexdens was monitored on the day of the mating flight, and the winged ants, as well as the spiders, were collected. The results obtained corroborate the hypothesis that phoretic behaviour is commonly used for dispersion of the spider A. attarum, predominantly females. Of these spiders, 64 individuals of A. attarium were collected, of which 62 were females (96.9%) and two were males (3.1%). Regarding the winged leaf-cutting ants sampled, 378 females and 361 males were collected, totaling 739 individuals. Of these, 64 individuals (8.7%) had a spider attached to its back for phoretic dispersal and none was observed on the queens after the nuptial flight. In our study, A. attarum females perform phoretic dispersal into the environment on winged leaf-cutting ants but do not settle in the new nests.     

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.     

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 biology of Limnoria chilensis: another boring peracarid species with extended parental care

Several boring peracarid species engage in extended parental care with parents tolerating small juveniles in their burrows, but only anecdotal observations have been reported from boring isopods of the genus Limnoria. The isopod Limnoria chilensis Menzies is frequently found in holdfasts of the macroalgae Macrocystis pyrifera (L.) C. Agardh and Durvillea antarctica (Chamisso) Hariot. In the present study, holdfasts of D. antarctica that harboured L. chilensis were carefully dissected in order to examine the association pattern of reproductive females and small juveniles. In most infested holdfasts, L. chilensis reached very high densities of 43.7 ± 3.9 (mean ± SE) individuals cm^?2 (range: 0–90 individuals cm^?2). The overall sex ratio (females:males) was close to unity (1.2). Male size varied between 1.5 and 2.6 mm body length (BL) and female size between 1.5 and 3.6mm BL. The percentage of reproductive females was low (19.7% of all females), and only the largest females (>2.6 mm BL) were reproductive. Embryo numbers ranged between six and 19 per female and a significant correlation between female BL and the number of embryos was found. Several females were found with small juveniles (0.8–1.2 mm BL) in the terminal end of their burrows. On several occasions, aggregations of similar-sized juveniles (1.0–1.5 mm BL) in their own burrows were found near a female burrow, indicating that these juveniles initiated their first individual burrows from within the maternal burrows. Some females with small juveniles in the terminal end of their burrows were either accompanied by males or they were ovigerous, indicating that they may have been in the process of producing a second brood. The fact that juveniles build their first burrows in the protection of female burrows suggests that such small juveniles have not yet developed full boring capacity. Extended parental care in this (and other boring) peracarid species represents a mechanism facilitating high juvenile survival rates. Given the highly local recruitment, it is suggested that the reproductive biology of this isopod has strong implications for Its population biology.     

Constraints on convergence: hydrophobic hind legs allow some male pollinator fig wasps early access to submerged females

Pollinator fig wasps (Hymenoptera: Agaonidae) display numerous adaptations linked to their obligate association with fig trees (Ficus). Ceratosolen fig wasps pollinate figs that often fill temporarily with liquid, and one clade has males with unusually long hind legs. We investigated their morphology and behaviour. Scanning electron microscopy (SEM) revealed that the cuticle of their hind legs is highly modified and covered with numerous hydrophobic setae and microtrichia that can prevent blockage of the wasps’ large propodeal spiracles by liquids. In deep liquid, the males floated on the surface, but when only a thin layer of liquid was present, the legs allowed males to access females without the risk of drowning. Access to females was facilitated by an air bubble that forms between the hind legs and maintains a column of air between the spiracles and the centre of the figs. Sexual selection should favour males that can gain earlier access to mates, and the modified legs represent an adaptation to achieve this. Convergent adaptations are known in some unrelated non-pollinating fig wasps that develop in similar liquid-filled figs, but these species have enlarged hydrophobic peritremata at the ends of their metasoma to protect the spiracles located there. Unlike non-pollinating fig wasps, pollinator males need to insert their metasoma deep into females’ galls during mating. This difference in mating behaviour has constrained the extent of convergence.     

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.