Female mate choice,male-male competition and parental care in the river bullhead,Cottus gobio L. (Pisces,Cottidae)

Male body size was tested for its influence on female mate choice, male-male competition and ability to defend broods in the river bullhead, Cottus gobio L., a polygynous fish with paternal care. Females presented with two potential mates of different sizes significantly preferred to spawn with the larger male. Males smaller than, or 1·5 times longer than, the female were rarely selected as mates. Larger males were more successful in defending their brood from conspecifics, which may explain female preference for them. Unmated large males displaced smaller guarding males from their nests and retained the acquired egg masses. Competition between males for nest sites with eggs can be accounted for by the preference of females for males already guarding eggs: by seizing a nest containing egg masses, a male will increase his chance of being chosen.     

Spawning Behavior and Biparental Egg Care of the Crosshatch Triggerfish, Xanthichthys mento (Balistidae)

Reproductive ecology of the crosshatch triggerfish, Xanthichthys mento (Balistidae) was studied at Hachijojima, Izu Islands, Japan. Males established territories and repeatedly chased females passing nearby. There were 1–3 females in each male's territory before spawning and during egg care. This species spawned in pairs on the sandy bottom. Eggs were scattered and attached to sand particles. Females care for the eggs by blowing water on them and guarding them against intruders, while males helped in guarding. Thus, biparental egg care was observed for 2 days until hatching. Both the males and females disappeared from the territories after the egg care. The reproductive ecology of this species is compared with that of other balistids and the unique features of X. mento are described.     

Factors influencing egg parasitism in sub‐social insects: insights from the treehopper Alchisme grossa (Hemiptera,Auchenorrhyncha, Membracidae)

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Male mating success and female mate choice in the river sculpin,Cottus nozawae (Cottidae)

Synopsis Mating success of males and its correlates were investigated in a natural population of the polygynous fluvial sculpinCottus nozawae. Furthermore, the female mate preference of this species was examined experimentally under alternative conditions for mating in a stream. The mating success of individual males (the number of females with which a male mated) ranged between 0 and 8 with a mean of 2.41 in 1983 and 2.52 in 1989, in a population of which the sex ratio was about 1 : 2 in both years, skewed toward females. Mainly due to the excess of nests without egg masses and the few nests with one egg mass, the distribution of male mating success did not fit a Poisson distribution, indicating its non-randomness. Male mating success was not correlated either with the size of the nest rocks or with the male size, suggesting that these two variables are not determinants of mating success. The mate choice experiments demonstrated that females of this species more frequently chose smaller males as mates whose nests already contained eggs than large males without eggs. Additionally, an analysis of stomach contents of guarding males suggested that the parental males ate their own eggs during egg guarding (filial-cannibalism). Based on these results and on a comparison of reproductive characteristics with congeneric species, it is suggested that one of the most important determinants for female mate choice inCottus species may be whether or not parental males are filial egg cannibals.     

A new record of amphisexual care in an insect with exclusive paternal care: Rhynocoris tristis (Heteroptera: Reduviidae)

We describe the first record of amphisexual care in an insect with uniparental male care. Male Rhynocoris tristis are noted for aggressively guarding multiple egg masses. When a male is unable or unwilling to guard eggs, a female that has previously contributed eggs returns to the egg mass and defends it until the eggs hatch. We compared the frequency and intensity of parental aggression in both sexes and found no difference in either form of guarding behavior. Evidence of amphisexual care in R. tristis provides an opportunity to quantify the cost of care in both sexes within a single species.An erratum to this article can be found at     

Male body size and brooding success under natural conditions in the freshwater Japanese goby, Rhinogobius sp. OR (orange type)

Males of the Japanese goby Rhinogobius sp. OR (orange type) establish nests under stones in streams. They provide care to egg masses until hatching. We carried out field observations and a field experiment in a natural stream to examine the effect of male body size on brooding success. An experiment excluding guarding males indicated that their presence has a positive effect on nest defense. Field data showed that male body size was positively correlated with the size of the egg mass that was initially found at the nest. Furthermore, a sequential increase in egg mass size seen in the nests of larger males indicated that multiple females spawned during a single breeding cycle of larger males. To examine male brooding success, two dependent variables, egg mass persistence (days from initial egg mass to disappearance or hatching) and hatching success were analyzed. The results indicated that the variation found in both dependent variables was significantly explained by two independent variables, namely, initial egg-mass size and the number of matings. It is also suggested that male body size indirectly influenced brooding success through positive correlations with these independent variables.     

Changes in male physiological condition during brooding activities in a natural population of a stream goby, <Emphasis Type="Italic">Rhinogobius</Emphasis> sp.

We investigated the change in physical condition of males during paternal care in a natural population of a stream goby Rhinogobius sp. LD. Hepatosomatic index (HSI) and Condition factor (K) of males guarding an egg mass were higher than those of non-breeding males. In the first brooding cycle, the HSI of males decreased with the progress of embryo development. Moreover, in the second brooding cycle, both the HSI and K decreased, suggesting that the energy reserves in the somatic tissue were used when the liver reserves were exhausted as a source of energy for paternal activities. After paternal care, males showed reduced K and were unable to recover HSI immediately. Comparisons of the physical condition between non-guarding males after the first brooding cycle and males guarding an early stage clutch in the second brooding cycle indicated that males which could recover HSI and K were able to have a second brooding cycle. This suggested that the physical condition determines participation in breeding activity and the number of brooding cycles, greatly influencing male reproductive success.     

Mate choice and cannibalism in a natural population of a stream goby,Rhinogobius sp.

Breeding ecology of the stream goby,Rhinogobius sp. LD (Large Dark), was investigated under natural conditions. Males selectively courted females of similar size to lead them to the nests, whereas females followed courting males preferentially when the relative male size was greater. Male-male competition for a female was relatively infrequent and not severe. Developmental stages of eggs and egg numbers in one nest indicated that males receive 1–3 clutches during one breeding cycle. Males guarding multiple clutches frequently ate some of the eggs, but those guarding single clutches rarely did so. Gravid females in the nest also frequently cannibalized eggs laid by a previous female, probably in order to extend the area available for egg deposition. Mate choice in this species is discussed in relation to paternal ability, limitation of available spawning area and the female-biased sex ratio.     

Parental care and social organization of the spiny eel, Aethiomastacembelus platysoma, in Lake Tanganyika

This is the first report demonstrating the occurrence of parental care in mastacembelids. Social organization and parental care of a spiny eel Aethiomastacembelus platysoma were studied in Lake Tanganyika. Both males and females maintained individual territories though the frequency of aggressive interaction was low. The male guarded offspring in a rock hole within its territory. The egg size was large (2.5–2.7 mm in diameter) and the brood size in a nest was 5.7 on average in spite of more oocytes in the ovary (65 large oocytes on average). The duration of guarding was around 30 days after hatching and the young became independent just after they began to feed. Guarding males seldom attacked fishes that approached the nest, and often went out of the nest to forage though the stomach contents of guarding males were less than those of non-guarding males. Compared with Tanganyikan cichlid fishes that show prolonged parental care at open sites, the post-hatching guarding interval is short and the egg size is large, which seem to be traits common to fishes that utilize closed spaces as guarding sites in the lake.     

Defensive capability of egg-guarding males of the goby,tridentiger kuroiwae brevispinis

The capability of egg-guarding maleTridentiger kuroiwae brevispinis to defend their own nests and egg masses, measured experimentally by matching them with conspecific male competitors of various body size, was almost total against competitors of body lengths ≦104% of defending male body length. Against competitors with body lengths >116% of defending male body length, nest and egg defence almost always failed, indicating an upper limit of competitor's relative body size (between 104% and 116%) against which successful nest and egg-guarding is possible. The specific growth rate of egg-guarding males was significantly lower than that of non-egg-guarding males, due to starvation and stress. The results suggested that male body size has a major influence on guarding males' reproductive success, through post-spawning male-male competition.     

Proximate Factors Regulating Maternal Options in the Eggplant Lace Bug, Gargaphia solani (Heteroptera: Tingidae)

The eggplant lace bug, Gargaphia solani, was used to investigate the proximate factors regulating maternal care and a noncaring, condition dependent strategy called egg dumping. We hypothesize that the act of delaying oviposition while searching for a dumping opportunity suppresses oogenesis and triggers guarding behavior. We examined several predictions of this hypothesis by measuring: (1) whether females do delay oviposition in the absence of dumping stimuli, (2) whether females in transition between egg dumping and egg guarding are capable of expressing either reproductive option, (3) the effect of nymphal interactions and antennal ablation on the duration of maternal care, and (4) oogenesis in guarding and dumping females. We found that females without a dumping opportunity wait, on average, 30 h longer to oviposit than females exposed to a dump mass. Females that had initiated their own egg mass could resume eggdumping if they had laid less than half of their eggs but were unlikely to abandon their eggs when most had been laid. Maternal care in G. solani can be prolonged if interactions with nymphs are artificially prolonged. Females require antennae to maintain maternalcare. Presumably antennae transduce cues from eggs and nymphs. Dissections of dumping and guarding females 72 h after their first oviposition demonstrated that dumpers continue to produce primary oocytes after a dumping event but guarders terminate oogenesis whilecaring for their first brood. We interpret all of these results within the context of the hypothesis that juvenile hormone titers regulate the expression of both egg dumping and egg guarding.     

Why egg-caring males of Isaza (<Emphasis Type="Italic">Gymnogobius isaza</Emphasis>, Gobiidae) refuse additional females: preliminary field observations

Like many other gobies, males of the Isaza goby, Gymnogobius isaza endemic to Lake Biwa, Japan, conduct parental care of eggs at nests, and females are likely to choose mates while visiting nests. The reproductive strategy should induce polygyny, but Isaza males never accept additional females in one breeding cycle. Sampling data of broods indicated that the egg mass areas were much smaller than the nest sizes, suggesting that nest size is not the limiting factor in obtaining further eggs from additional females. The brood size greatly decreased as the duration of care progressed. Few individuals including caring males ate eggs, and heterospecific egg predators were rarely observed. Sixty percent of egg masses at the middle stages of egg development were infected with aquatic fungi, some being covered with a fungus mat that drastically reduced survivorship. Infected egg masses contained more eggs than non-infected ones at the same stage, indicating that large egg masses are prone to be frequently destroyed by fungi. It is likely that the activity of parental males is lowered during the long care periods at low water temperature in early spring. Such lowered activity of caring males might be responsible for infections in large broods that would have needed more care. We propose the hypothesis that male rejection of additional females may be related to optimal brood size, which will be less susceptible to fungus infection and produce more hatching young than otherwise. This hypothesis will explain not only male avoidance of additional females but also some unique reproductive behaviors of this fish such as some females spawning of a portion of of the mature eggs in one nest.     

Does it Pay to Care?: Exploring the Costs and Benefits of Parental Care in the Hibiscus Harlequin Bug Tectocoris diophthalmus (Heteroptera: Scutelleridae)

Knowledge of the selective pressures favouring parental care can inform our understanding of the evolutionary origins and transitions of sociality in insects. Here, we report upon investigations designed to estimate the costs and benefits of parental care in the egg‐guarding hibiscus harlequin bug Tectocoris diophthalmus (Heteroptera: Scutelleridae). We found that the presence of a guarding female had no effect on hatching success under benign laboratory conditions. In the wild, however, guarding decreased the likelihood of total clutch failure, and produced a fourfold greater egg‐hatching rate relative to unguarded clutches. Females guarded against generalist invertebrate egg predators, including conspecific nymphs, but were ineffective against hymenopteran egg parasitoids. Females continued to feed during the guarding period and exhibited no signs of weight loss or increased mortality due to this behaviour. We did not observe the production of subsequent clutches in any experimental females; therefore, the lifetime fecundity costs of providing parental care in T. diophthalmus remain indeterminate. Overall, maternal egg guarding appears to function as a relatively low cost–low benefit strategy that increases hatching success by protecting against predation – but not parasitism.     

Expected future reproductive success and paternal behaviour in the sand goby, Pomatoschistus minutus (Pisces, Gobiidae)

We studied egg care and guarding behaviour in the paternal sand goby, Pomatoschislus minutus , in relation to egg number and age. A male's expected reproductive success is assumed to increase with larger egg masses and older eggs. Non-shareable care, i.e. fanning, was higher in larger and older egg masses. Egg age explained 65% and egg mass size 22% of the total variation in the amount of fanning performed by a male. Contrary to expectation, egg removal did not affect a male's fanning behaviour, Males that were approached stayed longer in the nest when they guarded larger egg masses. Males with older eggs were also more reluctant to leave the nest. The time since a male had been chased away from his nest until he returned decreased as the season progressed. The results are discussed in relation to tradeoffs between present and future reproductive events.     

Reproductive behavior of largemouth triplefin Ucla xenogrammus (Tripterygiidae) on reefs of Kuchierabu-jima Island, southern Japan

Reproductive behaviors of the largemouth triplefin Ucla xenogrammus were observed on reefs of Kuchierabu-jima Island, southern Japan. Males exclusively maintained home ranges including spawning sites on vertical walls of overhanging reefs where filamentous algae were densely distributed. Females visited a male's home range to release adhesive eggs into the dense algae. Males guarded multiple egg clutches, but rarely showed fanning behaviors or mouth cleaning to the eggs. Aquarium experiments showed that guarding males had no significant effect on egg development. Because of the favorable water exchange conditions around their spawning sites, male U. xenogrammus could avoid the energy costs for egg care.     

Facultative Second Oviposition as an Adaptation to Egg Loss in a Semelparous Crab Spider

Semelparity is prevalent in arthropod species that exhibit maternal care. Previous hypotheses postulated that long‐term maternal care constrains future reproduction in females, leading to the evolution of semelparity. Nevertheless, females may occasionally lose all or part of their offspring because of predation or other causes. Where females lose the first egg mass for any reason, the potential for females to produce an additional egg mass could be adaptive. This potential may be found widely among semelparous arthropods as a conditional strategy. We tested this hypothesis using the crab spider Lysiteles coronatus whose females guard their egg mass against predators. L. coronatus females did not consume food during the 40‐d guarding period; this resulted in a 30.2% loss in their weight. When the females were separated from their eggs immediately after oviposition and were provided with food, they resumed feeding and their ovaries redeveloped. Dissection of guarding females indicated that their ovaries developed temporarily during egg guarding and that the developed ovaries were subsequently reabsorbed. These results suggest that the females maintain the potential to produce a second egg mass in case of egg loss, but that this potential declines towards the end of the guarding period. Field observations showed that a small fraction of the females oviposited in late July, when most females had completed egg guarding. The size of the late broods was similar to the oocyte numbers that we found in the females fed in the laboratory. This result suggests that a few females produced a second egg mass after they had lost the first one. Thus, we suggest that facultative second oviposition in L. coronatus females has evolved as an adaptation to egg loss, and that the development of ovaries during the guarding period is intrinsically programmed for compensatory oviposition.     

Male counterstrategy against infanticide of the female giant water bugLethocerus deyrollei (Hemiptera: Belostomatidae)

The male giant water bug'sLethocerus deyrollei counterstrategy against egg mass destruction by the female was investigated in the laboratory. When mature females encounter a brooding male, they destroy his egg mass and thereby take over the mate and gain a nurse for their own egg mass. However, when the male stays on egg masses laid above the water surface, females cannot detect the male nor the egg mass. Eggs of this species fail to hatch without being supplied with water by males. In the observation, brooding males frequently ascend the stick to the egg mass and stay there for long periods, although most of the water attached to their body surface flowed down in 90 s. These long stays on egg masses are regarded as countestrategies against females.     

Spawning and biparental egg-care in a temperate filefish,Paramonacanthus japonicus (Monacanthidae)

Synopsis Reproductive habits of a temperate filefish, Paramonacanthus japonicus, were studied on a rocky reef at Tsuyazaki, Fukuoka, Japan, from 1989 through 1990. Males had territories of 30–70m² and defended them from conspecific males and potential egg predators such as another filefish, Stephanolepis cirrhifer. Egg masses were found on the sandy bottom in male territories. Individual discrimination of males and females occurring in three male territories revealed that males and females stayed in stable pairs during one month of observation in 1989. In these stable pairs, males fed only within their territories, but females occasionally foraged outside. The occurrence of egg masses within male territories and biparental egg care showed that fish were reproducing as monogamous pairs. Contrary to this, males tagged in 1990 changed their territories after the disappearance of females, and males and females mated polygamously. Spawning was observed only four times during the study period, between 1633 and 1754h. Prior to spawning, the female prepared a spawning bed on the sandy bottom. The male nuzzled the female and the pair spawned, touching their gonopores on the spawning bed. Spawning was very quick and took only 1–3 seconds. The adhesive eggs were spherical with a diameter of 0.56 mm. They were mixed with sand particles and formed a doughnut-shaped mass of about 4 cm in diameter. One egg mass contained 3300–3800 embryos of similar developmental stage, which hatched 2–3 days later. P. japonicus appears to be monogamous but may also practice polygamy when pair-bonds are unstable.     

Sex-specific defence behaviour against brood parasitism in a host with female-only incubation

Nest protection against intruders is an indispensable component of avian parental care. In species with biparental care, both mates should evolve nest defence behaviour to increase their reproductive success. In most host-parasite systems, host females are predicted to have more important roles in nest defence against brood parasites, because they typically are primarily responsible for clutch incubation. Male antiparasitic behaviour, on the other hand, is often underestimated or even not considered at all. Here we investigated sex-specific roles in four aspects of great reed warbler (Acrocephalus arundinaceus) nest defence against a brood parasite—the cuckoo (Cuculus canorus), namely (1) mobbing, (2) nest attendance/guarding, (3) nest checking and (4) egg ejection. Using dummy experiments, simulating brood parasitism and by video-monitoring of host nests we found that males took the key roles in cuckoo mobbing and nest guarding, while females were responsible for nest checking and egg ejection behaviours. Such partitioning of parental roles may provide a comprehensive clutch protection against brood parasitism.     

Paternal care behaviour of the giant water bug Kirkaldyia deyrolli (Heteroptera: Belostomatidae) against ants

1. Although potentially vulnerable to predators, the offspring of subsocial insects are effectively protected by their parent(s). The female giant water bug Kirkaldyia deyrolli lays its egg masses on the vegetation above the water surface in aquatic environments and the males supply the eggs with water and guard them against cannibalistic females until hatchling dispersal. Field observations showed that egg masses are attacked by ants if the attending males are not present. 2. Laboratory experiments were conducted to evaluate the effects of paternal care by K. deyrolli against the ant Tetramorium tsushimae by means of four treatments: attending male with ant approach (WM‐WA); no attending male with ant approach (NoM‐WA); attending male without ant approach (WM‐NoA); and no attending male without ant approach (NoM‐NoA). 3. The rate of offspring survival was lower in the NoM‐WA group (45.3%) than in any other group, which showed similar offspring survival (WM‐WA = 80.4%, WM‐NoA = 75.1%, NoM‐NoA = 80.3%). Moreover, there were a total of 44 interactions between the attending male and ants in WM‐WA, and of these, a chemical compound was released by the attending male four times; this probably deterred ants from attacking because the ants went back to their colony. 4. In conclusion, the attending male can protect its eggs from ant predators and its care has an important role. To the best of the authors' knowledge, this is the first report of Lethocerinae males protecting their egg masses from ants by means of physical and chemical defence.