Mating behaviour and sexual selection in the African ball-rolling scarab Khepher platynotus (Bates) (Coleoptera: Scarabaeidae)

Mating behaviour of the ball-rolling dung beetle Kheper platynotus (Bates), adults of which are active in rainy seasons, was studied in southeastern Kenya. Male mating behaviour changed with the progression of the rainy season. In the early season when females made food balls, males attempted forced copulation with wandering and food-ball-making females on dung mounds. As the season progressed, females made brood balls, and males attempted to pair off with such females, as well as attempting forced copulation with wandering and food-ball-making females. A male who paired off with a brood-ball-making female cooperated with the female to make the ball. The male usually copulated after burying the ball (pre-copulatory mate guarding), but sometimes copulated during ball-making. In the latter case the male usually showed post-copulatory mate guarding until ball burial, but sometimes left soon after copulation. In the late season, when available females decreased because of maternal care in the underground nests, males started to make food balls to secure food resources for survival until the next rainy season. Male mate guarding involving ball-making, -rolling and -burying seemed to function to raise paternity confidence, but such assistance seemed to be less beneficial to females. Fights frequently occurred between guarding and intruding males on brood balls. Both residency and relative body sizes of fighters were important asymmetries influencing contest outcome. This seemed to result in not active but passive mate choice by brood-ball-making females.     

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.     

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.     

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 biology of Scinax fuscomarginatus (Anura,Hylidae) in south‐eastern Brazil

The present study describes the reproductive biology of Scinax fuscomarginatus in a remnant of Cerrado in south‐eastern Brazil. Observations were made between September 2002 and March 2004 at Estação Ecológica de Itirapina, State of São Paulo, south‐eastern Brazil. Breeding activities occurred in lentic and temporary bodies of water during the rainy season. Scinax fuscomarginatus exhibited a prolonged breeding pattern and a lek mating system. Males were smaller than females and defended individual calling areas through acoustic and physical interactions. Resident males consistently won encounters, but did not differ in size or mass from intruder males. Satellite behaviour was observed, but no female or amplected pair interception was registered. Scinax fuscomarginatus exhibited low operational sex ratios and the general reproductive mode in which eggs are laid in the water and tadpoles are aquatic. Amplexus was axillary and the eggs were deposited at the bottom of temporary ponds. Details on oocytes, egg masses, and eggs are included.     

The genus Grammostola Simon 1892 (Araneae: Theraphosidae): a new species from western Argentina,new synonymy and distributional data

Explosive breeding is a common reproductive strategy, especially in anurans. In some cases of explosive breeding in anurans, intense aggressive interactions occur among males attempting to gain access to females, and for terrestrial species that mate in water, females may die by drowning during long mating struggles. Such occurrences are obviously detrimental to females, and may also cause a decrease in the reproductive success of males through depletion of energy without obtaining access to a live female. However, the males of the small Amazonian frog Rhinella proboscidea can promote the ejection of oocytes from the abdominal cavities of dead females and fertilize them. This behaviour can minimize losses to both partners during explosive reproduction events. The existence of such a “functional necrophile strategy” shows that there may be possible selection in favour of stronger and more persistent males in explosive breeders.     

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.     

Reproductive ecology of a Tibetan frog Nanorana parkeri (Anura: Ranidae)

Endemic to the Tibetan Plateau, Nanorana parkeri figures among the highest altitude amphibians in the world. The present work, which was conducted in an alpine marsh at 4300 m altitude, is the first report of the species’ reproductive ecology. The breeding season, as indicated by the presence of amplectant pairs and fresh eggs in ponds, lasted from early Mary to early July. The ratios of adult male to female were 1.3 in hibernation ponds and 2.3 in spawning ones. Males were smaller than females. Amplexus was typically axillary (81.7%), with a few exceptional amplexus in which a male clasped another male or object rather than gravid females. Scramble competition occurred in 6.8% of the normal amplexus located. The body sizes of amplexed couples correlated positively. Deposited eggs were attached to submerged plants solitarily (60.5%) or in small clusters (39.5%), and the scattered arrangement of individual eggs could be a strategy to ensure embryonic development in oxygen-poor environments. Clutch size averaged 189 [standard deviation (SD) = 64], and correlated positively with female body size; egg size averaged 2.18 mm (SD = 0.10), independent of female body size; both traits within a clutch were negatively related with each other even controlling for female body size. Relatively few and large eggs laid by the Tibetan frogs, compared with those by lowland aquatic-spawning anuran species, should be adaptive to the harsh high-altitude conditions. Larvae hatched from eggs after 16.4 days on average (SD = 5.8). Froglets emerged between late July and early August, but some tadpoles were present in winter. These findings may help to understand the evolution of amphibians in extreme environments and to protect these remote species.     

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.     

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.     

Reproductive traits in the northern scorpion ( Paruroctonus boreus )

Despite its large geographic range, little is known about reproductive traits in the northern scorpion ( Paruroctonus boreus ). We analyzed reproductive traits for 36 females from a population near Billings, Montana. All data were collected within a single year. Litter size, offspring mass, total litter mass (TLM), and relative litter mass (RLM) were within the ranges of values reported for other species in the Vaejovidae. Female size (length or mass) was not correlated with any reproductive trait. Litter size and offspring mass were each positively correlated with RLM, suggesting that females investing relatively larger amounts of energy in reproduction increase both size and number of offspring. Finally, the within-litter coefficient of variation in offspring mass was negatively correlated with RLM, TLM, and mean offspring mass, suggesting that females investing more energy in reproduction produce more-uniformly sized offspring, an observation that appears common in scorpions.     

Egg production of two sympatric species of Hyalella Smith, 1874 (Crustacea,Amphipoda, Dogielinotidae) in aquaculture ponds in southern Brazil

This study tests the hypothesis that sympatric species of Hyalella may show different reproductive strategies that permit their coexistence. Ovigerous females were collected monthly with nets during a 1-year period in two aquaculture trout ponds in Vale das Trutas, São José dos Ausentes County, southern Brazil. In both species there was a decrease in the number of embryos in the brood pouch during embryonic development, and an increase in egg volume. However, the two species differed in the body size of ovigerous females, number of eggs and juveniles per female, brood mortality rate, egg size and hatching size of juveniles. Egg production continued throughout the year, with the highest number of eggs produced during winter and spring by Hyalella pleoacuta, and winter, spring and autumn by Hyalella castroi. These differences in the reproductive traits of species of Hyalella may permit their coexistence in nature.     

Reproductive characteristics of to symtopic lizards species, Sceloporus gadoviae and Sceloporus jalapae (Squamata: Phyrnosomatidae), from Tehuacán Valley, Puebla, México

We studied the reproductive characteristics of 2 syntopic lizard species, Sceloporus gadoviae and Sceloporus jalapae (Phrynosomatidae). Specimens of S. gadoviae ( N = 105) and S. jalapae ( N = 41) were collected in a tropical arid forest from Tehuacán Valley, Puebla, México. Males of S. gadoviae reached sexual maturity at the same snoutvent length (SVL; 45.0 mm) as S. jalapae , and a similar pattern occurred in females of both species (SVL; 41.0 and 42.0 mm, respectively). Males of S. gadoviae exhibited reproductive activity throughout the year, with a longer activity during the dry (November to May) and part of the wet season (June to September). In contrast, reproductive activity in S. jalapae males occurred during the wet season (July to September). Females of S. gadoviae showed continuous reproduction, whereas females of S. jalapae exhibited seasonal reproduction. Mean SVL of sexually mature females was higher for S. gadoviae (￣ x ± s￣x = 50.4 ± 0.52) than for S. jalapae (46.0 ± 0.54, P S. gadoviae was lower (3.9 ± 0.14 eggs) than for S. jalapae (5.6 ± 0.43). There was no significant correlation between snout-vent length of females and clutch size of S. gadoviae ( r 2 = 0.22, P > 0.05) or S. jalapae ( r 2 = 0.48, P > 0.05). Our study suggests that although both species inhabit the same environment, they have different reproductive characteristics.     

Reproductive ecology of black-tailed prairie dogs in Montana

Reproductive ecology of black-tailed prairie dogs ( Cynomys ludovicianus ) was studied on the Charles M. Russell National Wildlife Refuge in northeastern Montana (1978–1980 and 1985). Breeding took place from early March through early April in most years. Persistent snow cover and below normal temperatures in February and March of 1978 delayed the start of breeding. Litter size averaged 4.4 but varied significantly among years. Average yearly litter size was correlated (r 2 = 0.986) with summer (June-September) precipitation prior to the breeding season. Largest average yearly litter size (5.0) followed record precipitation, while the smallest average yearly litter size (3.8) followed extreme drought. More than half the yearling females failed to breed while 88% of the females two years and older bred. Testes weights were greatest early in the breeding season and regressed rapidly during April.     

Heterospecific mating between distantly related species of stink bugs and its evolutionary implications

The copulatory process in animals is preceded by a series of integrated communication processes which are often under selection to guarantee species-specific courtship and copula. However, individuals of species supposedly in reproductive isolation frequently copulate in nature, which has sparked the attention of evolutionary biologists for decades. While studies on this topic have strongly focused on recently diverged species or populations, few instances of heterospecific copula between distantly related species are known. Here we report an intriguing case of interspecific copula between two distantly related species of stink bugs in nature. We replicated this observation in laboratory conditions and revealed an asymmetric relation: males of Chinavia obstinata copulated with females of C. erythrocnemis but the reverse combination did not occur. We propose a series of hypotheses to explain our observations in the light of the current knowledge on mate recognition and reproductive isolation in stink bugs, considering the potential pre- and postcopulatory barriers involved. Once these two species have probably evolved in allopatry and contacted secondarily, the absence of directional selection against mating signals differentiation is a probable process. We argue that similarities in genital shape and size, most likely a result of convergent evolution, may have played a fundamental role in guaranteeing that these species achieved proper coupling.     

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.     

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.     

Effective strategy of the overwintering of semiaquatic bugs: overwintering of Velia caprai (Heteroptera: Gerromorpha: Veliidae)

Overwintering of Velia caprai was studied during the winter months 2006–2007 in the area of South Bohemia (Czech Republic) by (1) field phenological observations and samples, (2) study of artificially established experimental cohorts in an unheated glasshouse, and (3) auxiliary experiments in a laboratory. Velia caprai has a unique way of overwintering within the gerromorphan bugs that is rare among insects generally; it can overwinter in both an adult and an egg stage. Most females reach sexual maturity before overwintering, mate with mature males, overwinter actively on the water surface and lay eggs during the winter. The dormant eggs can be induced to hatch by increasing temperatures. Winter oviposition can be interpreted as a sign of a risk‐spreading reproductive strategy. The overwintering of both adults and eggs provides a chance to adapt to changes in the local environmental conditions and could be an important determinant in broad variability of life‐history traits.     

Some relationships of black-tailed prairie dogs to livestock grazing

Reproductive ecology of black-tailed prairie dogs ( Cynomys ludovicianus ) was studied on the Charles M. Russell National Wildlife Refuge in northeastern Montana (1978–1980 and 1985). Breeding took place from early March through early April in most years. Persistent snow cover and below normal temperatures in February and March of 1978 delayed the start of breeding. Litter size averaged 4.4 but varied significantly among years. Average yearly litter size was correlated (r 2 = 0.986) with summer (June-September) precipitation prior to the breeding season. Largest average yearly litter size (5.0) followed record precipitation, while the smallest average yearly litter size (3.8) followed extreme drought. More than half the yearling females failed to breed while 88% of the females two years and older bred. Testes weights were greatest early in the breeding season and regressed rapidly during April.