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.     

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.     

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.     

In front of a mirror: visual displays may not be aggressive signals in nocturnal tree frogs

Some evidence indicates that in anuran amphibians, visual signalling can be important during social interactions such as territorial disputes among males, especially in diurnal species. The correct identification of a signal is not a trivial matter. A visual signal provides a visual cue during a social interaction, and to be effective it must elicit an immediate response in the receiver. We tested the hypothesis that visual displays in an agonistic context constitute aggressive signals, in three nocturnal species of Hylidae. We predicted that the production of visual displays would increase in the presence of a conspecific intruder male. Males of Hypsiboas raniceps, Dendropsophus nanus and Lysapsus limellum were submitted to two treatments: (1) Self Image, a reflection in a mirror, simulating the presence of an intruder; and (2) Control, a black rectangle covering the mirror. We observed three visual displays: vocal-sac display (inflate the vocal sac and maintain it inflated for some time), limb lifting (rapid up-and-down movements of one or more limbs), and toe/finger trembling (rapid up-and-down movements of one or more toes and/or fingers). This last display was observed only in H. raniceps males. Contrary to our hypothesis, the emission rates of all visual displays of the focal animals did not differ between treatments; and the behavioural response did not differ among species. Therefore, we suggest that these behaviours could not be used directly for communication in agonistic contexts, and may represent displacement activities (involuntary responses). Alternatively, an aggressive bimodal stimulus may be necessary to trigger a behavioural response by using visual signals during territory defence in these three species.     

Observations of unusual acoustic behaviour in two Australian leafhoppers (Hemiptera; Cicadellidae)

Acoustic substrate‐borne communication between hoppers has been well studied with certain patterns of acoustic behaviour considered typical. We report observations of acoustic behaviour in two Australian leafhoppers, Stenocotis depressa (Walker) and Austrolopa brunensis Evans, that are markedly atypical. Two types of unusual acoustic behaviour were observed: female–female interactions, and the transmission of an elaborate male call through direct physical contact with the female. We discuss the evolution of these acoustic displays based on our observations of these hoppers and their environments, including the potential roles of intra‐sexual competition and the carrying capacity of the plant substrate. The discovery of these unusual behaviour patterns emphasizes the need for broader surveys of the diversity of acoustic behaviour in the Hemiptera.     

The breeding system of an endangered pupfish ( Cyprinodon elegans )

We examined the breeding behavior of the endangered Comanche Springs pupfish ( Cyprinodon elegans ) during 3 separate breeding seasons in a pool of springwater originating in Phantom Cave, near Balmorhea State Park, Texas, in relation to behaviors reported for the species in swift-flowing canals (Itzkowitz 1969). In the quiet pool, unlike in swift water, the breeding system was characterized by 3 different male mating tactics: territorial defense, satellite positioning, and sneak spawning. Although the breeding strategies adopted by the males were conditional, mating tactics generally reflected male size. Territorial residents were the largest, satellites were medium-sized, and sneakers were the smallest adult males observed in the population. Territorial males traversed the least amount of area, defending compact territories, while satellites and sneakers covered more area in attempts to spawn. Although preferred territories in swift water centered on algal mats, territorial residents in quiet water seemed to prefer territories around large rocks. Males defending rocks had higher reproductive successes than males occupying other substrate types. The flexibility of the males' tactics was exhibited during one year of study when large males appeared to be absent from the population. During that year, medium-sized males switched from the satellite to territorial tactic, while small males remained sneakers.     

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.     

Spawning behaviour of Engystomops pustulatus (Anura: Leptodactylidae)

Foam nests have evolved independently in several amphibian groups as an adaptive response to prevent predation and desiccation in dry environments. Nests are normally laid on ponds, or in underground galleries, humid forest leaf litter or terrestrial bromeliads. They are built when males or females beat a foam precursor associated with the egg masses extruded by the female. The spawning process requires the synchronic actions of the mating pair to obtain a hemispheric nest that protects the offspring. Herein, we describe the spawning behaviour of Engystomops pustulatus based on videos from 13 nesting couples from the lowlands of western Ecuador. Three variables were measured as indicators of male effort: duration of mixing events, duration of resting periods, and number of kicks per mixing event. We consider that not only male physical effort but also female behaviour influences nest structure. We suggest that nest building requires prolonged and intense physical activity by the male as well as the female’s steady position during spawning and female’s oviposition site selection. Nest building has two phases. In the first phase, the duration of resting periods, the duration of mixing events, and the number of kicks increase and are highly variable. During the second phase the three variables stabilise until the end. The volume of the nest increased mainly during the second phase. In four nesting events we observed kicking movements by the female. To our knowledge, this is the first time that female kicking has been observed in leptodactylid frogs. The function of this behaviour is unknown but our observations suggest that it may be triggered by insufficient male effort. Traditionally, female mate choice in Engystomops has been explained under models of indirect benefits exclusively. We argue that the prolonged male activity during nesting could influence female fitness directly. This will allow the operation of sexual selection via direct benefits.     

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.     

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.     

Mating behaviour in two closely related sympatric species of Philonthus (Coleoptera: Staphylinidae)

Summary

The intra-specific mating behaviour of two closely related sympatric staphylinid beetles, Philonthus cephalotes (Grav.) and P. sordidus (Grav.) is described in detail. Mating is essentially similar in the two species, although differing in the duration of copulation (1·7 min in cephalotes, 44·6 min in sordidus), and consists of four phases (A)-(D); (A) pre-copulatory phase, (1) examination of female abdominal apex by male, (2) examination of male abdominal apex by female, (3) prolonged examination of female by male, (4) mounting by male (5) extrusion of male genitalia; (B) copulatory phase; (C) terminal phase; and (D) post-copulatory phase.

An attempt to test the role of species specific patterns of peg setae, occurring on the genitalia of males, as a tactile signalling system was made by observation of inter-specific interactions between males and females of the two species. Under the trial conditions inter-specific intromission did not occur and behaviour was predominantly aggressive. Evidence suggests that signals during the early stages of an encounter between the sexes of different species are generally sufficient to deter further courtship. In two instances courtship reached a more advanced stage and here the patterns of peg setae on the male genitalia may have come into play, operating as a signalling device.     

Spawning behaviour and sexual dimorphism in the North American cyprinid fish Notropis leedsi,the bannerfin shiner

The spawning behaviour and sexual dimorphism of the little known North American cyprinid fish, Notropis leedsi, is described based on observations made in laboratory aquaria. Notropis leedsi is a crevice spawning species, a trait characteristic of members of the subgenus Cyprinella. Males establish breeding territories through ‘mock battles’ and defend them by chasing away intruding males or, at times, through threat displays or ‘mock battles’. Although wild-caught males are significantly larger than females, sexual dimorphism is more pronounced in the colour pattern and fin size, especially the dorsal and anal fins.

The modes of spawning behaviour in the genus Notropis are reviewed. The relationship between the promiscuous mating system (polybrachygamy), the crevice spawning mode, and sexual dimorphism in N. leedsi are discussed in relation to current notions regarding sexual selection.     

‘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.     

Post‐mounting courtship and the neutralizing of male competitors through “homosexual” mountings in the fly Hydromyza livens F. (Diptera: Scatophagidae)

Post‐mounting courtship behaviour of a 6–8 mm long scatophagid fly Hydromyza livens which lays its eggs in yellow water lily (Nuphar lutea) is described and analysed. Enigmatic male–male encounters are described, along with lengthy “homosexual” mounting behaviour. A possible interpretation of both types of behaviour is given, with reference to their adaptive significance and the special problems encountered by males in locating females.     

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.     

The vocal repertoire of Myrmeciza loricata (Lichtenstein, 1823) (Aves: Thamnophilidae)

The vocal repertoire is a tool that can be used to support systematics, conservation, ecology and behavioural studies. We characterize the acoustic parameters of the vocal repertoire of the white-bibbed antbird (Myrmeciza loricata) and describe the possible functions of each vocalization. The male’s song is related to territorial defence, and we found three calls in the repertoire. Call I is an alarm call, and call II is used for mutual recognition and remaining in contact with one another. Call III was emitted when the animals were foraging and when the individuals of the couple were very near to each other. Males and females sing as a duet, and their songs were individually distinct. The duet is possibly related to maintaining the pair bond, synchronizing breeding and warning potential intruders of the mated condition of the pair. This work is the first to describe the vocal repertoire of the white-bibbed antbird.     

The male of Lecane bulla (Gosse, 1851): new support for the synonymy of Lecane Nitzsch,Monostyla Ehrenberg and Hemimonostyla Bartos

The hitherto unknown male of Lecane bulla (Gosse, 1851) is described and figured. It is generally similar to the few known males of other Lecane species. However, whereas the degree of fusion of the toes accords between male and female in other species, this is not the case here: the male of L. bulla has two separate and movable toes, these are completely fused into a single toe in the female. This observation further questions the validity of the taxonomic separation of Lecane in three taxa of generic rank, based on the degree of fusion of the toes. The behaviour and reproduction of Lecane is commented upon.     

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.     

Intraspecific larval aggression in two species of Hyperini (Coleoptera: Curculionidae)

Two unusual types of behaviour (wandering and intraspecific aggressive behaviour) have been observed when rearing larvae of two species of the tribe Hyperini. Wandering of Hypera postica in search for food was observed in L1, L2 and L3 larvae, as well as in young L4 larvae. L1 larvae also disperse in response to crowding. Wandering of L2 to young L4 instars was a response to food shortage. Wandering of late L4 larva (‘prepupa’) occurs because of searching for a place to spin the cocoon and pupate. Encounters between the larvae may result in agonistic behaviour, and some larvae may die as a consequence of fighting. This aggression increases with food limitation. Agonistic intraspecific behaviour has not been described to date in weevil’s larvae and it probably may not occur under natural conditions when there is a plenty of food and larval densities are decreased by pathogens or parasitoids.