Brooding in Psolus patagonicus (Echinodermata: Holothuroidea) from Argentina, SW Atlantic Ocean

The mode, season, and time of brooding, egg diameter, egg number per brood, and the characteristics of newly released juveniles of Psolus patagonicus were investigated off Mar del Plata, Buenos Aires, Argentina, between October 1999 and February 2001. Individuals were attached to the Patagonian scallop, Zygochlamys patagonica. Spawning occurs between February and March. The mean egg diameter, 887 ± 26 μm, is the highest reported for the family Psolidae. Eggs are brooded under the mother’s sole until they develop into crawling juveniles within 7 months. The largest embryos reached a length of 1,941 ± 228 μm in September. During the brooding period (February–September) the number of brooded embryos decreased while their size increased. Our study confirms brooding behaviour in female P. patagonicus.     

Mass brooding of the blue octocoral,Heliopora coerulea on a sedimented equatorial reef

The octocoral Heliopora coerulea is a gonochoric surface brooder. Although the species is common on shallow Indo-Pacific coral reefs, information on its reproductive biology is limited and spawning timings have only been reported from four locations. We report the first observations of surface brooding in H. coerulea on a sedimented equatorial reef. In April 2014, 46 and 26 brooding colonies were recorded, respectively, from reefs fringing the islands of Kusu and Lazarus in Singapore. The brooded coral larvae were tightly bound to inflated coral polyps, forming a dense white mat which blanketed the brown coral tissue. The inflated polyps also protruded from layers of sediment, which accumulated in colony crevices, such that the brooded larvae were elevated above the settled sediment. Our observations supplement existing knowledge on the reproductive timing of H. coerulea and highlight the adaptations that improve survival of this species on sediment-impacted reefs.     

Exclusive Male Egg Care and Determinants of Brooding Success in a Millipede

The evolution of exclusive male parental care is a major and controversial issue in behavioural ecology. Although arthropods practicing paternal care are thought to be key taxa for investigating this issue, few studies have attempted to clarify the selection factors associated with male behaviour and fitness consequences in arthropods. In the millipede Brachycybe nodulosa, males curl their bodies around egg masses on the undersides of decaying logs. Male‐removal experiments in the laboratory strongly suggest that males defend the eggs against fungal infection. Orphaned eggs were soon covered by hyphae and no eggs hatched, whereas almost all eggs brooded by males successfully hatched. The egg‐brooding males showed no aggressive responses when disturbed. Only some mature males bred in the field. Furthermore, the number of eggs brooded varied greatly among the males. Selected generalized linear models revealed that males with a wide seventh body segment, which possesses gonopods (genital legs), tended to succeed in brooding; and males with a wider body also obtained more eggs. Colony attributes had no significant effects on male brooding. We discuss the possible sexual selection mechanisms that could accomplish this pattern of brooding success among male B. nodulosa.     

Maternal brooding in the children’s python <Emphasis Type="Italic">(Antaresia childreni)</Emphasis> promotes egg water balance

Parental care provides considerable benefits to offspring and is widespread among animals, yet it is relatively uncommon among squamate reptiles (e.g., lizards and snakes). However, all pythonine snakes show extended maternal egg brooding with some species being facultatively endothermic. While facultative endothermy provides thermal benefits, the presence of brooding in non-endothermic species suggests other potential benefits of brooding. In this study we experimentally tested the functional significance of maternal brooding relative to water balance in the children’s python, Antaresia childreni, a small species that does not exhibit facultative endothermy. Clutch evaporative water loss (EWL) was positively correlated with clutch mass and was much lower than expected values based on individual eggs. The conglomerate clutch behaved as a single unit with a decreasing surface area to volume ratio as clutch size increased. Maternal brooding had a dramatic impact on evaporation from eggs, reducing and possibly eliminating clutch EWL. In a separate experiment, we found that viability of unattended eggs is highly affected by humidity level, even in the narrow range from 75 to 100% relative humidity at 30.5°C (20–33 mg m⁻³ absolute humidity). However, the presence of the brooding female ameliorated this sensitivity, as viability of brooded clutches at 75% relative humidity was higher than that of non-brooded eggs at either the same absolute humidity or at near-saturated conditions. Overall, these results demonstrate that brooding behavior strongly promotes egg water balance (and thus egg viability) in children’s pythons.     

Brooding behavior in the shallow‐water brittle star Ophioderma wahlbergii

Brooding behavior and the brooded young of Ophioderma wahlbergii, a common, large, shallow‐water brittle star from South Africa, are described for the first time. Monthly samples of 20 specimens were collected from June 2013 to May 2014 (n=240 in total). The species was shown to be gonochoric. Females reached maturity at 17 mm disc diameter (dd), and their ovaries contained 5±3 yolky eggs of mean diameter 0.25±0.1 mm. No seasonal trends in reproduction were found, and >50% of females examined in each month were brooding. Brooding individuals contained an average of 7±6 and up to 33 developing young. Small stages were more frequent than larger young, suggesting that juveniles either emerge at different sizes, or more likely suffer mortality during development. Young within individual adults were of different clutches, but those within individual bursae were all of the same stage. No signs of larval stages were found, indicating that development is direct. Given the large size of emerging juveniles, the species is likely to be matrotrophic and hence truly viviparous. Brooding is a recent discovery in Ophioderma and, so far, the only known brooder has about 1000 small embryos. This study shows quite a different reproductive strategy within the genus, with few, large eggs developing into the largest brooded young known from any species of ophiuroid.     

Effects of seasonal variation in host quality and availability on parasitism by the egg parasitoid Telenomus coloradensis

Hemlock looper, Lambdina fiscellaria (Guenée) (Lepidoptera: Geometridae), is one of the most important defoliator in North American forests. The common egg parasitoid Telenomus coloradensis Crawford (Hymenoptera: Platygastridae) plays a significant role as a natural control agent, with parasitism levels in spring typically higher than in fall. The objectives of this study were to quantify changes in host acceptance and reproductive performance of the parasitoid in relation to (1) host egg fertilization in fall, (2) host diapause status, (3) host embryonic development in spring, and (4) host deprivation during summer. Our results indicate that T. coloradensis do not have the capacity to develop in unfertilized host eggs, whereas early‐diapausing eggs are more suitable for the parasitoid than post‐diapausing eggs. Furthermore, the host physiological suitability decreases with embryonic development in spring. Finally, a host deprivation period during the summer tends to negatively affect the parasitic activity of T. coloradensis. These laboratory results confirm previous hypotheses concerning T. coloradensis seasonal ecology and contribute to a better understanding of the effect of hemlock looper egg physiology and availability on the reproductive potential of T. coloradensis.     

Spawning strategy in Atlantic bobtail squid <Emphasis Type="Italic">Sepiola atlantica</Emphasis> (Cephalopoda: Sepiolidae)

This study aimed to determine the spawning strategy in the Atlantic bobtail squid Sepiola atlantica, in order to add new information to the knowledge of its reproductive strategy. A total of 12 females that spawned in aquaria were examined. Characteristics of the reproductive traits and egg clutches were similar to those of other known Sepiolidae. Clutch size varied from 31 up to 115 eggs. Females of this species had incorporated up to 1.58 times of their body weight into laid eggs. The size of laid eggs showed a positive correlation with maternal body size, supporting the idea that female size is a determinant of egg size. Our data suggest that S. atlantica is an intermittent terminal spawner, and that its spawning strategy comprises group-synchronous ovary maturation, multiple egg laying, and deposition of egg clutches in different locations. The obtained data provide insights for future comparative studies on reproductive allocation.     

Factors influencing double brooding in Eurasian Hoopoes Upupa epops

Double brooding may be a good strategy for short‐lived species to maximize annual and lifetime reproductive success (ARS and LRS, respectively). Nevertheless, there is typically individual variation in the probability of producing a second clutch. Here we evaluate factors that influence the decision to double brood in the Eurasian Hoopoe Upupa epops. Analyses of an 11‐year dataset showed that 36% of the females and 21% of the males produced a second clutch after successfully raising a first clutch. Double‐brooded females had higher ARS (9.1 ± 1.9 fledglings; mean ± se) and LRS (0.93 ± 0.08 recruits) than single‐brooded females (ARS: 4.5 ± 2.1 fledglings; LRS: 0.36 ± 0.03 recruits). This suggests that double brooding is adaptive in Hoopoes, and raises the question of why most individuals only produce one clutch per season. The probability of double brooding varied only slightly between years, suggesting that it is influenced by individual characteristics rather than by external, population‐level environmental factors. In both sexes, the probability of double brooding increased with earlier timing of the first clutch, and the timing of reproduction was the most important factor influencing reproductive success. The latter is likely to be mediated by changes in resources during the season. The probability of double brooding also increased slightly with female age, due to differences in intrinsic quality among females rather than to a gain in experience. In contrast to many other studies, the probability of double brooding increased with an increasing number of fledglings from the first clutch, suggesting that it is a strategy of individuals of high quality. Taken together, we show that the individual quality of the breeder and the timing of their first clutches are key factors influencing the decision to double brood, and thereby that they are important determinants of reproductive performance in Eurasian Hoopoes.     

Suitability assessment of three Trichogramma species in the control of Mythimna separata (Lepidoptera: Noctuidae)

Mythimna separata (Walker) is a serious cosmopolitan pest. Trichogramma species have been used worldwide as biological control agents of lepidopteran pests. However, no data on the control efficiency of Trichogramma species on this pest have been reported. We evaluated the suitability of M. separata eggs for three Trichogramma species, Trichogramma dendrolimi Matsumura, T. ostriniae Pang et Chen and T. japonicum Ashmead, and compared the suitability with that of Corcyra cephalonica (Stainton) eggs. The parasitism rate, development time, emergence rate, female proportion, body size and egg load of female offspring indicated that the M. separata egg is a suitable host for the three Trichogramma species. On the other hand, the non‐emergence rate and the relationship between egg load and body size reflected that M. separata eggs are less suitable for the three Trichogramma species than C. cephalonica eggs. Egg resorption occurred in T. japonicum 4 days after emergence.     

Reproductive Ecology of Pterapogon kauderni, an Endemic Apogonid from Indonesia with Direct Development

Pterapogon kauderni does not have secondary sexual dimorphism, and has an equal sex ratio (50.8% males:49.2% females). Females first mature at a length of about 34mm. Data from both laboratory-bred and wild-collected eggs and embryos are not consistent with the presence of differential reproductive allocation in this species. Analysis from clutches brooded by wild-collected males shows a significant intra-clutch and among-clutch variation in egg size. Mean individual egg size (diameter, volume, or weight) is not correlated with the size (standard length, head length, or lower jaw length) of the males brooding those eggs. P. kauderni has a lunar reproductive cycle, with a major spawning peak during full moon and a smaller peak during the last quarter. Similarly, it has a major juvenile release (settlement) peak at full moon, with a minor peak at new moon. However, one population, which is located in a site with distinct environmental conditions and which was sampled a year earlier, shows a different reproductive pattern, last quarter being the phase where most of spawn and release events take place.     

Length rather than year‐round spawning,affects reproductive performance of RAS‐reared F‐generation pikeperch,Sander lucioperca (Linnaeus, 1758) – Insights from practice

The continuous production of large numbers of high quality gametes is essential for aquaculture, particularly in candidate species, such as pikeperch, Sander lucioperca (L.). The common practice of year‐round reproduction is under suspicion of inflicting adverse effects on the quality of the gametes through the disturbance of endogenous rhythms. We hypothesized that such perturbation does not affect RAS‐reared F‐generation broodstock. Reproductive performance (number of eggs) and gamete quality (fertilization and hatching rate) were assessed over the course of 3 years covering six independent, photothermal shifted spawning seasons in a commercial pikeperch hatchery (n = 31 egg batches of F‐generation fish in total). No substantial differences in fertilization or hatching rates could be detected between the individual spawning seasons. Fecundity varied, but there are indications for a size effect on female fecundity with intermediate sized females producing higher number of eggs (~65–70 cm). Low egg quality could be detected in batches of very large fish. In conclusion, size‐specific broodstock composition, but not year‐round reproduction of F‐generation pikeperch spawners affects the reproductive performance.     

Predatory risk increased due to egg‐brooding in Armadillidium vulgare (Isopoda: Oniscidea)

Cost of reproduction is associated with a reduction in subsequent survival or future breeding success. A decrease in survival rate of parents during or after reproduction reduces the probability of their future reproduction. However, few studies have demonstrated such survival costs to parents. Females of Armadillidium vulgare hold their eggs in a marsupium and brood these until the young hatch. Caring for eggs in a marsupium seems to place a large burden on brooding females, and it restricts their predator avoidance behaviour. As such, costs of care may increase the mortality rates of brooding females. To reveal the costs of parental care, we examined the effects of egg brooding on behaviour and predation risk. Egg‐brooding females decreased speed of locomotion and rolling duration, and were killed by predators at a higher rate. Our results indicate that egg brooding in A. vulgare has costs in the form of predation risk.     

The relationship between fecundity and fertility in the mouthbrooding cichlid fish Tilapia leucosticta

The development of fry in the mouth of the female mouthbrooder Tilapia leucosticta Trewavas is described. The egg production by fish of various lengths is plotted and a curvilinear regression for standard and total length fitted. The number of eggs produced is approximately equal tothe square of the total length in centimetres of the parent fish. The brood numbers of 460 femalesare plotted against length, and linear regressions for mean brood number and mean fry number fitted. The relationship between the expressions for numbers of fry brooded and egg production gives anestimate of brooding efficiency that decreases with increasing parent length. Individual female fish show signs of reripening and the timing of this process is estimated from gonad state and from the length of fry brooded in the mouth. The relationship between fecundity (egg production) and fertility (fry brooded) is considered and the effect of maturation size, growth rate and spawning frequency assessed.     

Reproduction of the wedge sole (<Emphasis Type="Italic">Dicologoglossa cuneata</Emphasis> Moreau) in captivity: spawning parameters and influence of the natural temperature

In this work, the wedge sole (Dicologoglossa cuneata) spawning under a natural thermocycle has been studied. The spawning parameters were similar to those of other flatfish in captivity, the relative fecundity being very high (1.7 ± 0.1 × 10⁶ eggs kg female⁻¹ per spawning season). The percentage of floating eggs decreased with time and was directly related to egg diameter, daily relative fecundity and fertilization rate. Nevertheless, the hatching rate variations were not correlated to any other spawning parameter. The temperature range during the spawning season was very wide (11–21°C) though the largest spawns occurred between 12 and 16°C. The natural temperature cycle influenced wedge sole egg production, the winter temperature minimum indicating the onset of spawning, but the variations in the reproductive season did not appear to affect the daily egg production. Future research should focus on the photoperiod and its interaction with temperature as key factor for spawning.     

The relationship between fecundity and fertility in the mouthbrooding cichlid fish Tilapia leucosticta

The development of fry in the mouth of the female mouthbrooder Tilapia leucosticta Trewavas is described. The egg production by fish of various lengths is plotted and a curvilinear regression for standard and total length fitted. The number of eggs produced is approximately equal to the square of the total length in centimetres of the parent fish. The brood numbers of 460 females are plotted against length, and linear regressions for mean brood number and mean fry number fitted. The relationship between the expressions for numbers of fry brooded and egg production gives an estimate of brooding efficiency that decreases with increasing parent length. Individual female fish show signs of reripening and the timing of this process is estimated from gonad state and from the length of fry brooded in the mouth. The relationship between fecundity (egg production) and fertility (fry brooded) is considered and the effect of maturation size, growth rate and spawning frequency assessed.     

Maternal investment increases with altitude in a frog on the Tibetan Plateau

Reproducing females can allocate energy between the production of eggs or offspring of different size or number, both of which can strongly influence fitness. The physical capacity to store developing offspring imposes constraints on maximum clutch volume, but individual females and populations can trade off whether more or fewer eggs or offspring are produced, and their relative sizes. Harsh environments are likely to select for larger egg or offspring size, and many vertebrate populations compensate for this reproductive investment through an increase in female body size. We report a different trade‐off in a frog endemic to the Tibetan Plateau, Rana kukunoris. Females living at higher altitudes (n = 11 populations, 2000–3500 m) produce larger eggs, but without a concomitant increase in female body size or clutch size. The reduced diel and seasonal activity at high altitudes may impose constraints on the maximum body size of adult frogs, by limiting the opportunity for energy accumulation. Simultaneously, producing larger eggs likely helps to increase the rate of embryonic development, causing tadpoles to hatch earlier. The gelatinous matrix surrounding eggs, more of which is produced by large females, may help buffer developing embryos from temperature fluctuations or offer protection from ultraviolet radiation. High‐altitude frogs on the Tibetan Plateau employ a reproductive strategy that favours large egg size independent of body size, which is unusual in amphibians. The harsh and unpredictable environmental conditions at high altitudes can thus impose strong and opposing selection pressures on adult and embryonic life stages, both of which can simultaneously influence fitness.     

Seasonality and 3D‐visualization of brooding in the hermaphroditic ophiuroid Amphiura capensis

This article describes the reproductive biology and developing young of Amphiura capensis, a small brooding brittle star found in the intertidal zones of Namibia and South Africa. Each month from October 2014 to September 2015, 20 specimens were collected from Mouille Point, Cape Town, South Africa, and dissected for internal examination of brooding characteristics. Disc diameters of 238 analyzed adults ranged 2.5–8.7 mm, with a mean of 6.15 mm (SD = 1.10 mm). Brooding was exhibited in 60.5% of all sampled individuals, of which 30.6% contained young of only one of six subjectively chosen size classes, 31.9% contained two size classes, and the rest (37.5%) contained more than two size classes at the same time. Young within the same bursa were commonly of the same size class, which suggests sequential brooding. However, multiple size classes of young were often present within different bursae of the same parent, which thus exhibited characteristics of both sequential and simultaneous brooding. Of 584 brooded young retrieved from dissections, an average of 2.5 (SD = 3.2) and a maximum of 14 young were recorded within a single parent. As no larval stages were observed, development was assumed to be direct. Three‐dimensional visualizations of μCT scans revealed the positions of the brooded young to be quite different from those in other species, facing with their mouths downwards instead of upwards, and not all pressed against the adult's bursal wall. Brooding occurred throughout the year, but numbers of young peaked in austral winter, coinciding with warmer water temperatures at this site. The duration of brooded development was estimated to be ~6 months, comparable with other species with similar reproductive biology.     

The reproductive capability of Ooencyrtus kuvanae reared on eggs of the factitious host Antheraea pernyi

Ooencyrtus kuvanae is a key egg parasitoid of Lymantria dispar (L.), an important defoliating pest in China and North America. We have developed mass rearing techniques for O. kuvanae and in this study compared the reproductive capacity of O. kuvanae when it was previously reared on the natural host, L. dispar or the factitious host, Antheraea pernyi Guerin‐Meneville. There was no significant difference in the oviposition period or total number of eggs laid between L. dispar‐reared and A. pernyi‐reared females. However, the mean number of offspring successfully emerging from those eggs was significantly larger from the A. pernyi‐reared females compared with L. dispar‐reared females. From this, we can conclude that, with respect to reproductive capability, eggs of A. pernyi are suitable factitious hosts for mass rearing of O. kuvanae.