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.     

Mating behaviour,nympho-imaginal development and description of a new Mesabolivar species (Araneae: Pholcidae) from the Brazilian dry forest

A new species of Mesabolivar is described from Brazilian forests: Mesabolivar delclaroi. The mating behaviour, postembryonic development, number of egg sacs and eggs, birth rate, number of instars, developmental time until adulthood, sex ratio and size of cephalothorax (per instar) were recorded. The sexual behaviour was described and categorized into four steps: courtship, pre-copulation, copulation and post-copulation. After hatching, individuals presented five instars until maturity. The mean number of eggs (42 ± 16.6) and live births (31.5 ± 3.4) of the first egg sac were significantly greater than that of a second one (23.8 ± 3.8, and 19.25 ± 3.9, respectively). The developmental time from birth to adulthood (130.8 ± 9.6 days) did not differ significantly between egg sacs produced (128.61 ± 11.1). The size of the cephalothorax did not differ among adults or between sexes. The sex ratio revealed a shift in favour of females (4: 3).     

Long-term increases in clutch size in common snapping turtles (Chelydra serpentina) and diamondback terrapins (Malaclemys terrapin)

Clutch size (CS) is a well-studied life history characteristic, and optimal egg size theory predicts that CS should correlate positively with reproductive investment. Turtles are good subjects for studies of reproductive strategies because few turtle species exhibit parental care; therefore quantifying their short-term reproductive investment is mostly limited to measuring egg size and number. Clutch size is usually reported as an average value for each turtle population, and where CS variation is noted, it is usually attributable to variation in adult body size. In two long-term studies of ecologically dissimilar species we detected a dramatic increase in CS in a common snapping turtle (Chelydra serpentina) population and in a diamondback terrapin (Malaclemys terrapin) population. It is unknown whether these changes are due to either variation in adult body size or resource availability; but the temporal patterns we observed have apparently never been reported previously. These trends remain unexplained and should be explored further.     

Activity and reproduction in Megalobulimus paranaguensis (Gastropoda,Eupulmonata): implications for conservation in captivity for a South American land snail

ABSTRACT

In South America, Megalobulimus includes a number of threatened species, as well the largest land snails on the continent. The activity patterns and reproductive aspects of this group have not been documented. This work describes the daily and seasonal activity patterns and reproduction of M. paranaguensis. We maintained specimens in the laboratory for one year, and we quantified their behaviour for one hour at four different times of the day (0 h, 6 h, 12 h and 18 h) during three days in four months (August, September, April and May). The number of postures, hatching rate, time of hatching since oviposition and mortality rate among juveniles for each month were also quantified. Megalobulimus paranaguensis was more active in August, and had a egg laying peak one month after. Fifty-one eggs were laid by 12 captive individuals throughout the year, with a mean value of 4.25 eggs per individual. The hatching rate was 80.39%, and the time of hatching since oviposition was 56.7 ± 4.3 days. In two eggs, we observed the presence of twins. The mortality rate among juveniles was low (9.30%) indicating that rearing land snails in captivity has the potential to be an important and viable tool for the management and conservation of these organisms.     

Genetic differentiation and gene flow of Porcellionides pruinosus (Crustacea: Isopoda: Oniscidea) in Tunisia

Iso-enzymatic polymorphism analysis was carried out on Tunisian populations of Porcellionides pruinosus to clarify their confused taxonomic status, previously revealed by reproductive and morphological approaches. Seventeen allozyme loci were studied, using horizontal starch gel electrophoresis, in 230 adult specimens belonging to nine populations of P. pruinosus located in north, central and south Tunisia, to assess the levels of their intra-genetic and inter-genetic diversity, and to analyse their genetic structure. Our results indicate a low genetic variability with a polymorphism ratio ranging from 29.4% in Tabarka to 47.1% in the Garat Nâam and Raccada populations, and the mean number of alleles per locus was estimated at 1.51 ± 0.15. The direct-count of heterozygosity ranged between 8.1% and 15%. Substantial estimates of demic structuring were found at the total population (F_ST = 0.198), with a generalized and significant heterozygote deficiency detected in every population (with a mean value for all populations of F_IS = 0.372). Genetic distances between the studied populations were low, indicating that they belong to the same species. Nevertheless, the genetic analyses showed the divergence of the population of Tabarka from all the other studied populations, with the lowest mean gene flow, 0.58, and the highest mean genetic distance, (0.182) obtained also by the morphological and reproductive approaches.     

Offspring discrimination by female parents of the matriphagous spider Cheiracanthium japonicum (Araneae: Eutichuridae)

The ability of females of the matriphagous spider Cheiracanthium japonicum to discern their own offspring was examined based on a comparison of her maternal care for her own eggs and spiderlings with other conspecific females as well as those of other congeneric species. Cheiracanthium japonicum females did not discern their own nest from those of conspecific and congeneric individuals. In addition, the female did not distinguish her own egg sac from those of other conspecific or congeneric individuals. On the other hand, the female clearly discriminated between her own spiderlings and those of other congeneric species only if the congeneric spiderlings were introduced into the breeding nest. Consequently, the female will allow conspecific spiderlings to predate her (i.e. cannibalism). However, it is hardly expected that spiderlings of a certain female would eat another female in the field, as the reproductive period is quite short and synchronised in a population. Considering that the female can only recognise spiderlings of other species, the recognition ability of C. japonicum females will be evolved only to work as a protection against intruders of other species into the breeding nest.     

On the natural history of the South American pepper frog,Leptodactylus labyrinthicus (Spix, 1824) (Anura: Leptodactylidae)

Some species of Leptodactylus of the L. pentadactylus group lay their eggs outside water but the tadpoles need to reach water to complete the larval phase; other species complete development in terrestrial nests. Here we present details of the reproduction of L. labyrinthicus in south‐eastern Brazil. The proportion of tadpoles and trophic eggs in aged egg clutches was determined, as well as the growth of the tadpoles while in the nest. The gut contents of tadpoles that were in egg clutches of frogs were analysed. Adult males did not differ from females in size and had hypertrophied forearms and an enlarged spine on the thumb. Reproduction was initiated with the first rains of August/September and extended to mid‐January. Calling and spawning occurred at permanent or temporary water bodies. The foam nests were built in excavated basins outside of, but close to the water. The male determined the place of the basin construction; after amplexus, the female completed the excavation. The amplexus was axillary. One female spent the day after spawning in the foam. The eggs were pale grey, the yolk averaging 2.3?mm in diameter. The mean number of eggs was 2101 per egg clutch. The number of tadpoles in individual nests varied between 0.05% and 11.40% in relation to the total laid eggs. The tadpoles entered water when rains flooded the basin. The tadpoles grew to 12 times the weight of an individual egg while in the nest; no nesting tadpole was beyond stage 25. The longest time we followed tadpoles in a nest was 25 days. Tadpoles were found preying upon eggs of three other frog species and upon conspecific eggs. Males fought by grasping each other in a belly‐to‐belly position; the powerful arms and the thumb spines represent weapons. Even though males can reach maturity in the season following birth, small size would prevent them from establishing their own territory. All the species of the L. pentadactylus group may build their foam nests within excavated basins. The basins may protect the eggs and embryos from cannibalistic tadpoles and may have an anti‐desiccation effect. In order to produce trophic eggs, the female may delay laying additional unfertilized eggs until after the male has abandoned the foam nest. Anuran eggs represent an important food item for tadpoles after they leave the nest.     

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.     

A study of the anatomy of Pupa and Maxacteon (Acteonidae,Opisthobranchia), with an account of the breeding cycle of Pupa kirki

Summary

The external features, mantle cavity, reproductive system and nervous system of Pupa kirki (Hutton, 1873) and three species of Maxacteon Rudman, 1971 are described and compared with Acteon tornatilis (L). It is suggested that the reproductive system of Pupa and Maxacteon, with two accessory sacs, the gametolytic sac and an exogenous sperm sac, is more primitive than that of Acteon tornatilis.

The breeding cycle of Pupa kirki shows that the species has a basic annual cycle, a few specimens surviving for two years.     

Aspects of the life history and reproductive biology of the introduced terrestrial amphipod Arcitalitrus dorrieni (Hunt) at two sites in Co. Galway,Ireland

The reproductive biology of Arcitalitrus dorrieni (Hunt) was investigated over a period of 25 months at a deciduous and a coniferous forest site at Kylemore, Co. Galway, Ireland. The species has an annual, univoltine reproductive cycle with ovigerous females found throughout the year. Recruitment of juveniles begins in May–June with a peak of reproductive activity in August–September. The populations at each site show a bimodal size-distribution for most of the year, with a unimodal distribution in March, April and May. Mortality occurs exponentially, and the life-span is 20–22 months. Mean fecundity was 5·65 eggs per female and was dependent on female size, egg size and brood stage. Brood mortality was 37·5% at Addergoole and 29·8% at Tullywee Bridge.     

Biology and defensive secretion of myrmecophilous Thiasophila spp. (Coleoptera: Staphylinidae: Aleocharinae) associated with the Formica rufa species group

Various aspects of the biology of the myrmecophilous Thiasophila angulata and T. szujeckii (Staphylinidae; Aleocharinae), mainly associated with the Formica rufa species group, are studied. The composition of defensive gland secretions from adults of T. angulata has been analysed. The complete development (egg-L_1-3-pupa-adult form), including adult overwintering, of both species takes place solely in ant nests. Under laboratory conditions, this lasts for about 22 days in T. angulata, and the period elapsing between the old and new generations of insects is approximately two months. The mean fecundity of adults of this species (28 eggs) was determined, as well as the duration of its reproductive period (mean 28.5 days), and lifespan (mean 67 days). In natural conditions T. angulata, found within nests of Formica polyctena and F. rufa, produces three generations, and the sibling species T. szujeckii one generation per year. These two species differ in their phenologies and abundance dynamics of adults and larvae, which is linked to the size and temperature regime of the host nest. The results of this study uphold the recent separation of T. angulata and T. szujeckii based on morphological features of adults and selected molecular markers. Adults and larvae of T. angulata forage on both live and dead food items accumulated by the host, as well as on the host’s eggs and larvae. Both the beetle larvae and adults remain unmolested among the host workers. The adults use their defensive gland secretions, which contains substantial quantities of toxic quinones, when necessary. According to the current categories of myrmecophiles, T. angulata (and by analogy, T. szujeckii as well) should be classified as a species wholly integrated with the host.     

Studies on Alenophalera bronneomixia (Lepidoptera: Notodontidae)—a pest of the West African indigo plant,Lonchocarpus cyanescens (Papilionaceae)

Summary

This paper is concerned with the general biology of the notodontid moth, A. bronneomixia whose larvae are found to be serious defoliators of the West African indigo plant, Lonchocarpus cyanescens.

At temperatures fluctuating between 21°C and 29°C (mean 22·5°C) and humidities between 70 to 95 per cent eggs developed and hatched in a mean of 11·1 days from deposition, mean hatch being 93 per cent.

Under the said environmental conditions, the life cycle from egg to adult entailed six larval instars and a prepupal stage before the real pupa, all lasting 60 to 80 days. Larvae were strongly monophagous, fed communally and in nature pupated in soil/litter very close to the foot of their invaded food plant.

Adults were retiring and in the field deposited clusters of numerous eggs preferably on the abaxial surfaces of their food plants. In nature, this moth's population seemed strongly regulated by high egg and larval parasitization, larval predation and fungal attacks at the pupal stage.     

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.     

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.     

Structural comparisons of isomorphic breeding nests between closely allied spiders Cheiracanthium japonicum and Cheiracanthium lascivum (Araneae: Eutichuridae)

Closely allied spider species Cheiracanthium japonicum and Cheiracanthium lascivum make a closed breeding nest for egg laying and parental care. The nest provides the internal climatic stability required for suitable development of eggs and the physical durability required for protection against intruders. Although the breeding nests of these two spiders are quite similar in structure and appearance, their climatic stability and physical durability seem to be empirically different. Such physical features of the nests of these two spiders were compared based on a balance between the inner and outer air temperature and humidity of the nest as well as on the amount and size of spider silks lining the nest. In addition, the female’s relative energy allocation to egg production versus nest construction was examined based on the number or weight of eggs versus the climatic stability and physical durability of the nest. According to the results, the stability of temperature and humidity was maintained better in the breeding nest of C. japonicum than in that of C. lascivum. Furthermore, the nest of C. japonicum was more strongly constructed, with a greater volume and size of silks, than that of C. lascivum. On the other hand, the number or weight of eggs in relation to the female’s body weight in C. japonicum was smaller than that in C. lascivum. These results suggested that the reproductive effort towards nest construction for the purpose of egg and juvenile care in C. japonicum was larger than that in C. lascivum. In contrast, the effort towards egg production in C. japonicum was smaller than that in C. lascivum. Consequently, it is likely that the structural differences in breeding nests between these two spiders are responsible for the discrepancies in the female’s relative energy allocation to nest construction.     

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.     

Life history of Subulina octona (Brugüière) (Gastropoda: Pulmonata: Subulinidae) based on four-year laboratory observations and a comparative histological analysis of egg-retaining and ovoviviparous subulinids

In the present study, we aimed to characterize the life history of the land snail Subulina octona integrating information on life-history traits and morphology. We also compared the histology of the free-oviduct and spermoviduct of ovoviviparous and egg-retaining species of subulinids. We considered as ovoviviparous the species in which the complete embryonic development as well as egg hatching occurs inside the parent’s body and, at the end of this process, the parent releases juveniles instead of eggs. We considered as egg-retaining the species in which a great part of the embryonic development takes place inside the parent’s body and the eggs laid contain well-developed embryos. The results showed that the free-oviduct of both ovoviviparous and egg-retaining species shows a histological arrangement that confers greater strength to its walls and is probably related to egg retention. The wall of the spermoviduct is formed by pseudostratified columnar epithelium, with cell apical processes (probably cilia), and by underlying secretory cells. In gravid egg-retaining snails, the eggshells appear in close contact with the secretory cells of the spermoviduct. This fact suggests that these cells play a role in eggshell formation. The present study is the first account for histological features of S. octona, Allopeas gracile and Allopeas micra. The life history of S. octona is a combination of long lifespan, early sexual maturity, indeterminate growth and egg retention. Egg retention limits the number of young that can be produced in one reproductive event. However, the reproductive strategy adopted by S. octona, associating egg retention and K-strategism, compensates this cost of retaining eggs, because the reproductive success may be enhanced as a result of the higher survival of juveniles and the possibility of performing several reproductive events during the year.     

The biology and immature stages of Thrybius togashii Kusigemati (Hymenoptera: Ichneumonidae: Cryptinae), with a description of the male

Thrybius togashii Kusigemati is found to be an ectoparasitoid of larvae of a phytophagous eurytomid, Tetramesa sp. (Hymenoptera), growing gregariously in internodal cavities of reeds (Phragmites japonica Steud.), in Japan. The female of T. togashii oviposits into reed canes and its predaceous larva consumes almost all eurytomid larvae in an internodal cavity. This species is bivoltine. The mode of parasitism of this species is unique in some respects. First, when the female oviposits, no food resources are available in the reed. Second, the parasitoid allows part of the host larvae to grow and exploits increasing food resources. The egg, pupa and first to final instar larvae and adult male are described and figured for the first time. Colour dimorphism in both sexes and adult morphology adapted to habitat and ovipositional substrate are also discussed.     

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.     

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.