Sexual selection and mating system variation in anuran amphibians of the Arizona-Sonoran Desert

Mating system variation in anuran amphibians of the Arizona - Sonoran Desert was reviewed. Male density and breeding period duration were negatively correlated in seven bufonids and pelobatids. Variation in male mating behavior and ability of females to freely select their mates unhindered by active - searching males also was related directly to male density. These observations support hypotheses relating ecological factors to mating system organization. It is suggested that male calling behavior, and anuran lek mating systems in general, may be significantly influenced by predation on vocalizing males.           

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).     

XL.—A new Fossil form of Phasmatodea from Galkino (Turkestan), and on Mesozoic Phasmids in general

Insects of many groups form mating aggregations on hilltops. Males of some hilltopping species behave like certain classic lekking vertebrates, notably in their defense of territories that are devoid of resources useful to females but which are visited by females solely for mating. But even within a sample of 11 hilltopping insects found in the same habitat in central Arizona, there is considerable diversity with respect to resources transferred to mates by males, the degree to which males are aggregated and territorial, and the extent to which females can choose freely among hilltopping males. Thus ‘hilltopping’ is not a single mating system type; although some species conform closely to the traditional definition of a lek, others are very different. The ecological basis for the diversity in the behaviour of male hilltopping insects appears linked to differences in population density, which affect the costs of territoriality, and differences in the nature of female choice, which are little understood as yet.     

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.     

The function of the sphragis in Cressida cressida (Fab.) (Lepidoptera,Papilionidae): a visual deterrent to copulation attempts

During mating, males of Cressida cressida (Lepidoptera, Papilionidae) form a large external mating plug or sphragis that covers the female's copulatory opening and physically prevents remating. The sphragis has lateral and distal projections that make it an obvious structure. We tested experimentally the hypothesis that the sphragis acts as a signal of female mating status. Males pursue mated females with an intact sphragis only briefly and rarely make physical contact with them. When the sphragis of a mated female is removed or reduced in size, males are significantly more likely to physically contact the female and initiate the aerial takedown that preceeds a copulation attempt. These results suggest that the sphragis deters male sexual interest at a distance and thus functions as a signal of female mating status. The discussion focuses on the fitness consequences for females and their mates of the signal function of the sphragis.     

Breeding movements and reproductive activities of porcupine in the Great Basin Desert

I assessed movements of North American porcupines ( Erethizon dorsatum ) in the Great Basin of northwestern Nevada in relation to reproductive activities during the late summer and fall periods of 1991 and 1992. Porcupines exhibit a mate-defense polygynous mating system and I hypothesized that (1) competitively dominant males would have larger home ranges than both subordinate males and adult females, and (2) variation in home range size among adult male porcupines would be positively correlated with reproductive success. Results indicated that dominant male porcupines ranged over larger areas (average 95% minimum convex polygon home range = 20.7 ha) than subordinate males (average 95% MCP home range = 2.9 ha) and adult females (average 95% MCP home range = 8.2 ha). Analyses of movements in relation to body size and energetic requirements revealed that home ranges of dominant male porcupines were larger than predicted based on body size (approximately 10.2 ha). Breeding period home ranges of dominant male porcupines encompassed portions of the home ranges of 3 to 10 adult females, and indices of reproductive success based on observations of mate-guarding behaviors suggested a strong positive relationship between home range sizes of male porcupines and mating success. Together these data suggested that larger home ranges among dominant males were related to increased mating opportunities and not increased metabolic requirements associated with larger male body sizes. In the study area, however, female porcupines congregated around small, patchily distributed riparian areas, and dominant males with relatively small home ranges encompassing riparian areas may have gained mating access to multiple females. Finally, analyses of overlap among core home ranges (60% MCP) of adult male and adult female porcupines suggested that both sexes maintained relatively exclusive core home range areas, with males exhibiting significantly less range overlap with other males ( ￣x = 9.4%) than females with other females ( ￣x = 27.1%). It is possible that the small, patchily distributed riparian areas in this desertlike area were such a limited resource that females were unable to maintain exclusive use of their home range areas.     

The occurrence and function of the sphragis in the zerynthiine genera Zerynthia,Allancastria and Bhutanitis (Lepidoptera: Papilionoidea: Papilionidae)

In some butterfly species males attach a large external mating plug termed a sphragis to the female abdomen during mating. This is derived from male accessory secretions and covers the female ostium bursae and surrounding areas, thus preventing or delaying remating. Specimens of all 12 species of the genera Zerynthia, Allancastria and Bhutanitis (Lepidoptera: papilionidae), which form a natural clade within the Zerynthiini, were examined for presence or absence of a sphragis and their male and female genitalia were studied. In all three genera female genitalia lack a typical sinus vaginalis and the sterigma is modified to form an exposed, shiny, well-sclerotized genital plate, derived from the fusion and expansion of the lamellae ante- and postvaginales. The exposed ostium bursae is situated near the posterior end of the genital plate in Zerynthia, whereas in Allancastria and Bhutanitis it is near the anterior end. A crude irregularly formed sphragis was found at least facultatively in all species. The sphragides of Zerynthia and Bhutanitis were generally poorly developed, in most cases only partially covering the female genital plate. In Allancastria the sphragis mostly covered the genital plate entirely, and generally incorporated numerous long scales derived from the male’s 8th abdominal segment; scales were sometimes sparse or absent, probably due to depletion from repeated matings by males. In Zerynthia, males lacked the dense terminal abdominal tuft of elongated scales found in Allancastria, and their sphragis lacked scales. The sphragis of Bhutanitis thaidina incorporated scales from the male valves, whereas in the B. lidderdalii sphragis (and probably B. ludlowi) the scales derived from the male’s 8th abdominal segment. The role of the scales and possible reasons for the difference in the development of the sphragis among these genera are discussed.     

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.     

Occurrence of sexual morphs in Trama troglodytes von Heyden, 1837 (Hemiptera,Aphididae)

The root-feeding aphids of the tribe Tramini have been cited as a possible case of ancient apomixis, as there are no substantiated records in the literature of a functional bisexual generation, even in common and well-known European species. The karyotypic variation found in this group, involving considerable structural heterozygosity within species, also pointed to the decay of diploidy that would be an expected consequence of loss of sexuality and absence of meiosis. However, we have now found a small number of oviparae (mating females) and males in populations of the common species Trama troglodytes. Fertilized eggs were not obtained, but adult oviparae contained large, yolky eggs. The single adult male was apterous, small and blind, but had fully developed mouthparts and genitalia, and contained mature sperm. Both sexual morphs are described, and the karyotrypic variation is reassessed in the light of these findings. Occasional sexual reproduction is in concordance with recent molecular evidence that some recombination occurs in T. troglodytes and related species.     

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.     

Courtship,copulation and oviposition in the chalcedon checkerspot,Euphydryas chalcedona (Lepidoptera: Nymphalidae)

In this report we describe some aspects of the reproductive biology of a population of the chalcedon checkerspot, Euphydryas chalcedona, in central Arizona. Successful courtship in this species lasts about a minute and has a form similar to that of other nymphalids. During copulation the male forms a spermatophore within the female's bursa copulatrix that represents about 7% of the male's body weight. After copulation females may remate and older females may have mated as many as three times. The spermatophore deflates as time since deposition passes. Females oviposit every other day on average and lay their eggs in clusters. These data confirm previous brief reports of courtship structure and spermatophore size for E. chalcedona. Questions are raised about the control of receptivity in this species and the function of the mating plug.     

Palpal loss,single palp copulation and obligatory mate consumption in Tidarren cuneolatum (Tullgren, 1910) (Araneae,Theridiidae)

Males of Tidarren cuneolatum (Tullgren, 1910) amputate one of their palps a few hours after the penultimate moult, like T. sisyphoides (Walckenaer, 1841) from the USA (Branch, 1942). Hence adult males, which are minute, have only one palp, either left or right randomly. This palpal organ is not oversized, when compared with other small spiders. During courtship females are unusually active, signalling receptivity by continuous twanging with legs II. Males construct a mating thread. Copulation involves one insertion, which lasts ca 4 min. Thus, only one receptaculum is inseminated during copulation. With the advance of insertion the male's prosoma becomes shrunken. Copulation regularly ends in mate consumption. In copulation with a virgin female the palp was inserted contralaterally. Females taken in the field had both receptacula filled with sperm and therefore were polyandrous. Re-mating was also observed in the laboratory. Remarkably the second male performed an ipsilateral insertion, if it possessed the same palp as the previous male. Probably the virgin receptaculum was recognized. Postembryonic development is rapid in males, which moult three times and mature ca 41 days after hatching from the cocoon. Females need four or five moults and ca 69 days to reach maturity and then survive ca 2-4 months.     

Are there costs to extended larval transport in the Trinidadian stream frog,Mannophryne trinitatis (Dendrobatidae)?

Previous work has shown that male Mannophryne trinitatis (Dendrobatidae) carry their larvae on their backs for up to 4 days in search of a predator‐free pool in which to deposit them. The experiments reported here investigated whether costs to the larvae or to the adults limit transport duration. We simulated transport durations of 0, 4, 8, and 12 days for larvae, but found no deterioration in terms of ability to grow to metamorphosis; indeed, 12‐day larvae grew better than all the others. After 8 days of simulated transport, larvae had used up all their yolk reserves and begun to lose dry weight. Larvae on wet substrates gained wet weight and length but on drier substrates merely maintained weight, suggesting that dehydration could be a problem on the male's back. In a trial of locomotor performance (mean jump length; number of jumps to traverse a runway), females performed best with calling males not significantly different from transporting males, despite an average larval load equivalent to 15–20% of the frog's mass. Assessment of gut contents showed that females foraged more than males, but that transporting males foraged as much as did calling males. We found no differences between the three classes of adult frogs in fat body weight.     

The male of Culicoides reevesi Wirth, with a redescription of the female and new seasonal activity, distribution, and biting records (Diptera: Ceratopogonidae)

The previously unknown male of the biting midge, Culicoides reevesi Wirth, is described and illustrated; the female is also redescribed and this species is reassigned to the leoni group. Previously known from California, Arizona, and New Mexico, C. reevesi is recorded for the 1st time from Utah ( new record ). Females of this aggressive, hematophagous species were collected while biting humans during evening crepuscular periods in California. Females exhibited a strong attraction to CO 2 traps, and seasonal surveillance demonstrated that host-seeking occurred from late May until mid-October in both California and Utah. Small numbers of males were also collected in CO 2 traps; however, both sexes showed little attraction to ultraviolet and incandescent light traps.     

Do male tree frogs feed during the breeding season? Stomach flushing of five syntopic hylid species in Rio Grande do Sul,Brazil1

The diet of males of five syntopic hylid species from the Araucaria plateau of Rio Grande do Sul was studied by stomach flushing. Because the duration of their breeding seasons was rather different, the levels of food intake at the respective breeding sites were analysed. We found a positive correlation between the length of the male calling period and the ingestion of prey. The longer the species‐specific reproductive activity of the males, the more individuals fed during that time. The results are discussed under congruence aspects of feeding and mating microhabitats in these tree frogs. In addition, differences in niche breadth may influence the temporal and spatial partitioning between the species studied and result in quantitative as well as qualitative patterns of prey consumption, thereby ensuring a maximum of fitness gain.     

Prolonged parental behaviour by males of Limnonectes palavanensis (Boulenger 1894), a frog with possible sex-role reversal

ABSTRACT

In sex-role-reversed systems, males carry out all parental care duties, forgoing further mating opportunities. Cases of complete sex-role reversal have not been reported in anurans. We describe the parental care behaviour of the leaf-litter frog Limnonectes palavanensis, a Bornean endemic with egg-guarding and tadpole transport, possibly the first anuran with sex-role reversal. In this study, we tested one of the predictions of the sex-role reversal hypothesis: males invest more in parental care than females do, and this prevents them from quickly acquiring additional mates. We observed egg-guarding behaviour in captivity to determine the length and intensity of parental care. Also, we describe the tadpole climbing and tadpole transport behaviour. We found that males perform all of the parental duties, remaining with the clutch until hatching. They do not call during this time, nor do they care for more than one clutch at a time. Observations of tadpole transport in the field indicate that all tadpoles in the clutch are retrieved at once. The behaviour of males during tadpole retrieval suggests that they stimulate synchronous hatching. Together, these results indicate that the male parental expenditure in this species is high, resulting in lost mating opportunities. The prolonged male-only parental care behaviour of this species satisfies an important criterion for the occurrence of sex-role reversal.     

Studies on the life cycle of the cicadellid bug Empoasca dolichi Paoli,in Southern Nigeria

Summary

Studies carried out on the biology of E. dolichi Paoli showed that its eggs are palegreen to creamy white, elongate and often curved, and have a mean length and width of 0·84 mm and 0·16 mm respectively. The egg-incubation period averaged 8·3 days under uncontrolled insectary conditions of 21–32°C and 45–95% relative humidity, while the life-cycle from egg to adult averaged 17·5 days. In the insectary, mating in E. dolichi usually started between the third and sixth day after adult emergence. Over 56% of observed individuals mated on the 4th day. The observed diurnal mating period was mainly within 05·30–09·00 hours and 16·00–20·00 hours local time, with an average copulation period of 65·7 min. The mean pre-oviposition period of 3·0 days for 23 mated females was significantly shorter than 4·0 days for 21 unmated females. An average of 116 eggs and 96 eggs were laid by mated and unmated females respectively, but the fecundity rate (3 eggs/day of total life-span) of mated females was significantly higher than that (2 eggs/day) of unmated females. Unmated females lived longer (mean=49·7 days) than mated females with a mean of 38·3 days. The average life-span for mated and unmated males was 31·0 and 32·4 days respectively. Insectary and field observations revealed that females of E. dolichi preferentially lay their eggs into the stems, leaf-veins and leaf petioles of cowpea plants. Only mated females of E. dolichi laid viable eggs, and the observed male: female sex-ratio of their progeny in the insectary was 1:1·04.     

Assortative mating in soldier beetles (Cantharidae: Chaulignathus ): test of the mate-choice hypothesis

Soldier beetles of 2 species, Chauliognathus basalis and C. deceptus , were examined to test the Crespi hypothesis that positive assortative mating by size is caused by mate choices. Specifically, we tested the prediction that if mate choice involves choosing the largest mate available, then mating individuals will be larger than nonmating individuals. Four samples were taken, at different times during the mating season, from each of 2 sites. Each sample consisted of mating pairs, nonmanting males, and nonmating females. Some of the samples contained beetles of both species; others contained beetles of a single species. For each gender elytron lengths of mating individuals were compared with elytron lengths of nonmating individuals. We found no effect of mating status (mating vs. nonmating) on elytron lengths in samples that exhibited assertive mating (which occurs where 2 species coexist). Surprisingly, we found a consistent effect of mating status on elytron lengths in samples that did not exhibit assortative mating (which occurs where only 1 species exists). Our results do not support the mate-choice hypothesis. Instead, mate choice and assortative mating appear to be alternative mating patterns in which mate choice occurs where a single species exists and assortative mating occurs where 2 species coexist.     

An imperfect imago? Post-mating loss of iridescent scales in Cheimas butterflies may change female from attractive to cryptic (Lepidoptera: Nymphalidae: Satyrinae)

Adults of both sexes of Cheimas opalinus (Staudinger), a Satyrinae butterfly occurring in the cloud forests of the Venezuelan Cordillera de Mérida, have simple wing colour patterns, dark brown dorsa marked by one conspicuous element, large greenish-blue patches in the discal area of the hind wing. It was observed that in young, freshly emerged females these patches are frequently absent. They fade away and eventually disappear during mating and nuptial flights when brushed out by lateral movements of the male’s hind wing put between folded female hind wings. The falling off of the scales is made easy because they do not adhere to the wing as firmly as brown background scales due to their particular folded extremities. We speculate that this process is related to the fitness of the females. Prior to mating, blue-green patches are advantageous because they attract the attention of the opposite sex and enhance the chances of successful mating. After mating they lose their sexual role. Females without the blue-green patches become cryptic. Thus, they are less apparent not only for the males, which helps them avoid sexual harassment, but also for potential bird predators. Additionally, the loss of blue reflecting scales may speed up the warming up of the abdomen and egg maturation. In the males, positive role of the patches (signalling), and negative (bird attraction), is balanced during their entire life span, and they fade gradually due to regular usage of wings scaling in flight. Our hypothesis is supported by morphological, optical, experimental and statistical analysis in which we used 509 individuals of both sexes. A similar ratio of young (with undamaged wing) females with or without blue patch was recorded, whereas in the males no individuals without any trace of blue patch were reported, and a correlation of age (wing damage) and gradual fading off of the patch was demonstrated. Such an adaptation involving an active change of the appearance of adults, i.e. colour patterns related to sexual selection, has not been reported previously in Lepidoptera.     

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.