Brood defence and parental roles in a biparental cichlid fishLamprologus toae in Lake Tanganyika

Brood defence of a cichlid fish,Lamprologus toae, was investigated in its natural habitat in Lake Tanganyika. Both parents guarding a brood attacked both conspecific and heterospecific intruder fishes at different locations. The heterospecific intruder fishes could be classified into three groups on the basis of the locations at which the attacks against each species took place. The distinction of groups by the parents seemed to be primarily based on food habits and feeding behaviour of the intruder fishes. The piscivorous species which were more dangerous for the brood were attacked by both parents at more distant locations from the brood. Parental defence of breeding territory changed with the development of the young. The frequency of attacks against each group decreased after the young reached the size too large for the fishes of the group to prey on. Division of labour in the territorial defence was recognized between male and female. The male parent mainly defended the peripheral region of the territory and the female parent defended the inner region. Significance of the selective attack against intruders and the division of labour between the two sexes in brood defence is discussed.     

Maternal investment in a bee species with facultative nest guarding and males heavier than females

1. Maternal investment can be influenced by several factors, especially maternal quality and possibilities for future reproduction. Mass provisioning Hymenoptera are an excellent group for measuring maternal investment because mothers distribute food sources to each brood cell for each offspring separately. Generally in aculeate Hymenoptera, larger females produce larger offspring and invest more in female offspring than in male offspring. 2. This study investigated patterns of maternal investment in Ceratina chalcites, which has an uncommon type of sexual size dimorphism in Hymenoptera: on average, males are heavier than females. It was found that larger females produce a significantly higher proportion of male offspring, as males are the costlier sex in this species. 3. Facultative nest guarding by females was observed. Females can guard offspring until adulthood, as is typical for bees of genus Ceratina (34.43% of nests); however, in the majority of cases (65.56% of nests), females plug and abandon the nest. Significant differences were found in the amount of investment between guarded and unguarded nests. Guarded nests had a greater number of provisioned brood cells and a higher proportion of male offspring. It is suggested that mothers have two facultative strategies – either she makes a large investment in the offspring of one nest or she abandons the first nest and carries out a second nesting elsewhere.     

The effect of partial brood loss on male desertion in a cichlid fish: an experimental test

There is little experimental evidence testing whether currentbrood size and past brood mortality influence mate desertion.In the cichlid Aequidens coeruleopunctatus both parents initiallydefend offspring. In a field study, all experimental broods,irrespective of initial brood size (222.9 ± 60.4, mean± SD), were manipulated to a size of 100 fry. Neitherthe duration nor investment of females in parental care differed between control and brood reduced pairs, even though care seemedcostly. On average, females lost 5.1 ± 4.8% of initialweight while guarding a brood until independence. In contrast,males with experimentally reduced broods guarded fry for significantlyfewer days before deserting their mate than did males fromcontrol pairs with natural-sized broods (20.5 ± 7.5 vs. 14.2 ± 6.2 days). In at least 20% of cases (n = 9/45),the deserting male immediately mated with another female. Maleswith experimentally reduced broods also spent less time guardingfry before deserting and attacked fewer brood predators thandid males with control broods. For broods manipulated to have100 fry, there was a significant negative relationship betweenthe days until male desertion and the proportion of the initialbrood removed. This indicates that male assessment of the futuresuccess of the current brood (hence its reproductive value)is based on past mortality and/or that there is variation amongmales in the expected size of future broods. Both current broodsize and brood size relative to initial brood size are thereforepredictors of male, but not female, parental behavior and matedesertion. Female care may be unaffected by brood reductiondue to limited breeding opportunities and partial compensationfor reduced male care.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

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.     

The Costs of Confronting Infanticidal Intruders in a Burying Beetle

Infanticide by unrelated adults is a complex behavior in burying beetles (Nicrophorus spp.) serving multiple functions (resource competition, access to mates, cannibalism). The costs of confronting an infanticidal intruder are likely to vary with context. To assess these costs for a single female parent (Nicrophorus pustulatus), we systematically manipulated the sex of a conspecific intruder and the timing of the intrusion. Male intruders were a greater takeover threat than female intruders, and infanticidal takeovers were more common earlier in the breeding cycle. Even though a male intruder posed a greater threat to the brood, a female intruder was a greater threat to the reproductive success of the resident female. Female intruders that took over a carcass excluded the resident female from the resource. When a male intruder took over a carcass, the resident female was able to recover much of her loss by producing a replacement brood. Even when females successfully defended their broods, they suffered decreased reproductive output relative to control females that never had to confront an intruder (expt 1), but the mechanisms underlying this cost were unclear. To test the hypothesis that defeated female intruders attempt to parasitize late‐stage broods, female intruders whose eggs could be identified by a fat‐soluble dye were introduced to resident females caring for larvae (expt 2). Fifteen of 20 intruders oviposited eggs and the number of eggs were related to intrusion pressure. Because resident females rarely produced eggs while caring for larvae, it is uncertain whether the behavior of the defeated female should be characterized as brood parasitism, a failed takeover attempt, or an attempt to use the remains of the depleted resource. This study provides the most complete picture of the changing costs of confronting an infanticidal threat throughout the vulnerable period of offspring development.     

Delayed copulation as a means of female choice by the hermit crab Pagurus filholi

Male hermit crabs perform precopulatory mate-guarding behavior during their reproductive season. As females generally cannot reject guarding attempts by males, male guarding prevents females from inspecting and choosing other male mates. However, as guarding males are often replaced by other males through competition for females during the guarding phase, females may be able to select males by delaying their copulation. To examine the possibility of female choice by hermit crabs, we investigated whether female Pagurus filholi that were being guarded in the field were ready to copulate and spawn. We found that about 30% of females guarded in the field were ready to spawn, indicating that guarded females delayed copulation with their current male. Our results suggest that by delaying copulation females may exploit male–male competition to choose dominant males. However, delaying copulation reduced the spawning potential of females. Hence, there is a trade-off between waiting for the opportunity to mate with a dominant male and decreased spawning success if females exploit male–male competition.     

Type of intruder and reproductive phase influence male territorial defence in wild-caught Siamese fighting fish

This study investigated how parental care increases with offspring age by examining the level of male aggressiveness toward potential intruders during egg guarding in a natural population of Siamese fighting fish (Betta splendens Regan). The degree of aggressiveness was measured at two reproductive phases in response to three types of intruders: male, female and females that have laid eggs. The nest-holding males became more aggressive after their eggs hatched than after the eggs were laid. Nest-holding males displayed gill cover erection, biting, tail beating and attacking at the highest rate towards male intruders, intermediate towards female intruders and the least aggressive towards females, which have laid eggs.     

Parental behaviour in the Lapwing Vanellus vanellus

Parental behaviour of monogamous and polygynous Lapwings was studied during incubation and brood care. Both parents attended the nest in 86% of monogamous pairs ( n = 29 pairs). In 14% of pairs, only the male parent continued incubation until the eggs hatched, whereas the female deserted the clutch before or at the end of incubation. There was a clear division of parental roles during incubation. Females spent more time incubating (64% of time) than their mates (27%), whereas males spent more time defending the nest (3%) than females (>1%). Time spent incubating did not differ between monogamous and polygynous males. However, polygynous females spent more time incubating (primary females: 95%; secondary females: 97%) than monogamous females. Biparental care was the most common pattern of post-hatching care, although in some broods either the male or the female parent deserted before the chicks fledged. Division of sex roles was less pronounced in brood care than during incubation. Females spent more time brooding (21%) than males (3%), and females attended their chicks more closely than males. Nevertheless, males and females spent similar amounts of time defending the brood from predators and conspecifics. We suggest that the apparent division of parental roles may be explained by sexual selection, i.e. the remating opportunities for male Lapwings might be reduced if they increase their share in incubation. However, the different efficiency of care provision, for example in ability to defend offspring, may also influence the roles of the sexes in parental care.     

Competing for last place: Mating behaviour in a pill-box crab, Halicarcinus cookii (Brachyura: Hymenosomatidae)

The mating strategy of Halicarcinus cookii was investigated to ascertain how males maximised their fitness through mate choice. An intertidal population at Kaikoura, New Zealand, was dominated by mature crabs of both sexes in summer and by immature crabs in the colder months. More than 95% of mature females were ovigerous with early stage and late stage broods found in almost every month, indicating that egg production and larval release is continuous. The operational sex ratio was less than 1 male/female in summer, but often more than 1.0 in the colder months. The gonosomatic index increased along with brood development so that as soon as zoeae were released, the next clutch of eggs was ready to be fertilised. Males searched for receptive females and began pre-copulatory mate guarding without any courtship display. They mated preferentially with late stage or non-ovigerous females: copulation duration was longest for stage 5 females as was post-copulatory guarding (mean 18.3 h). Late stage females were up to 14% of the female population. Mate attraction seems to be the result of an ovarian signal rather than from the developing brood. Manipulation of the sex ratio had effects upon copulation duration and post-copulatory guarding: presence of a rival male increased duration of guarding. Females showed precocious mating in the penultimate instar and were able to lay fertilised eggs after their pubertal moult in the absence of males. H. cookii females have many mates, but males attempt to ensure paternity by preferentially pursuing mature females close to egg laying and by guarding these females after copulation. These behaviours are all elements of a competitive strategy to ensure that a male loses (not wins) the race to copulate because females have a ventral seminal receptacle, giving sperm precedence to the last male to mate. Male mating behaviour is a consequence and evolutionary response to female morphology.     

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.     

The evolution of breeding strategies in the flightless cormorant (Nannopterum harrisi) of the Galapagos

The evolution of breeding strategies in male and female flightless cormorants is thought to have been shaped by three principal factors: Firstly, an unpredictable food supply; secondly, a prolonged breeding season; and thirdly, the inbred nature of colonies.
These factors have selected for opportunistic breeding and for females (but not males) to attempt to raise two broods per season by deserting their offspring to the further care of their mates, while they re-breed with new males. That this strategy maximizes the inclusive fitness of both sexes is examined in terms of genetic pay-off and the relative food-providing efficiency of male and female parents. Parental investment is examined in relation to female desertion. Mate selection is discussed and some theoretical aspects of breeding strategy evolution are considered.     

Indirect female choice mediated by sex pheromones in the hermit crab Pagurus filholi

Males of the hermit crab Pagurus filholi perform precopulatory guarding behavior, and solitary males often show aggressive behavior to take away guarded females. Males behave coercively while guarding females, so direct mate choice by females seems difficult in such a situation. By performing several experiments we examined possible indirect female choice of hermit crab. Males were attached to a shell by their left cheliped to look like guarding pairs (fake guarding pairs). The shells were filled with cotton containing either seawater or pheromone water. The fake guarding pair with only seawater caused male–male combat in 60% of trials whereas with pheromone water combats occurred in 88% of trials. Mean duration of male–male combat was significantly longer in trials with drops of seawater containing pheromones than in those without pheromones. These results suggest guarding pairs themselves cause male–male combat by visual stimulation, that female sex pheromones have further significant function in the recognition of guarding pairs and intensification of male–male combat, and that females release sex pheromones while they are guarded. As a result of the combat, the larger male ended up guarding a female. This strongly suggests that females choose males indirectly by exploiting male–male competition induced by sex pheromones under male coercive behavior.     

Sexual conflict and parental care in magnificent frigatebirds: full compensation by deserted females

Parental care is a cooperative venture between a male and a female in many socially monogamous birds. Care is costly, and thus, sexual conflict arises between the parents about how much effort they should invest into rearing their young. The sexual conflict over care is most apparent when one parent abandons the brood before the offspring are independent. The deserted parent has three options: (1) desert the brood because a single parent is unable to raise the young on its own; (2) continue care provision at the same level as during biparental care, and thus do not compensate for the absence of mate; or (3) increase care and compensate partially or totally. We investigated these options in the magnificent frigatebird, Fregata magnificens, a species in which the male deserts his mate and brood before the chick is independent. During biparental care, females fed the chick more often than the males. After their mate deserted, the females nearly doubled their feeding rate and thus, fully compensated for the lost care. Consistent with these observations, growth rates of chicks provided with biparental and female-only care did not differ. These results support recent theoretical models of parental care, and suggest that females may withhold care during biparental care to manoeuvre their mates into prolonged care provision. A female only provides at her full capacity once her mate has deserted.     

Part-Time Mate Guarding Affects Paternity in Male Reed Buntings (Emberiza schoeniclus)

Male mate guarding by close following has been reported in many socially monogamous bird species and is generally believed to function as a paternity guard. Many aspects of the dynamics and effectiveness of this behavior are still however poorly understood. Here, we describe the temporal variation in mate guarding behavior in male reed buntings (Emberiza schoeniclus) with a particular focus on how males allocate their mating effort between mate guarding and extrapair mating in a context of intense sperm competition. In our highly synchronous study population most males have to balance the simultaneous and mutually exclusive demands of mate guarding and seeking extrapair copulations (EPCs). We found that males frequently switched between guarding their mates and performing intrusions to neighboring territories. Both activities seemed to have significant fitness payoffs, as male mate guarding effort had a positive effect on paternity, and a large fraction of extrapair fertilizations occurred during the days when the sire guarded its own female. The reed bunting is thus an example of how discontinuous or part‐time mate guarding can still be effective in securing paternity. Female reed buntings were not particularly active in initiating EPCs as they never were observed performing extraterritorial forays. We argue that the absence of female‐initiated EPCs is a prerequisite for males to trade mate guarding against seeking EPCs. Otherwise, if females circumvent male mate guarding by timing their EPCs to periods of male absence, males should guard their mates almost continuously or rely on alternative paternity guards.     

Female Reproductive Cycle and Sexual Conflict over Precopulatory Mate-guarding in Thermosphaeroma (Crustacea, Isopoda)

In species with time-limited opportunities for insemination, precopulatory mate-guarding is expected to coevolve with the duration of female reproductive cycles. Despite this adaptation to female characteristics, it may also be advantageous for males to adjust the duration of guarding with respect to sex ratio because the benefits of guarding are dependent on the availability of females. If female fitness is reduced because of guarding, male guarding behavior leads to intersexual conflict. We studied these aspects of male mate-guarding behavior in two closely related, thermal-spring isopods (Thermosphaeroma). First, guarding duration showed species specificity which was related to the duration of reproductive cycle; cycle length for females and duration of guarding by males in T. milleri were twice as long as in T. thermophilum. Second, males in both species adjusted their guarding duration with sex ratio, guarding longer when a competing male was present. Third, in T. thermophilum, ovarian development began immediately after the birth of the previous brood and continued through guarding, sexual molt and post-molt periods until oviposition, whereas in T. milleri, ovarian development was largely postponed until the post-molt period. Because guarding during ovary provisioning periods may be costly for females, we tested the existence of intersexual conflict over guarding duration in T. thermophilum. We compared the duration of guarding of control pairs with those of pairs in which either male guarding ability or female ability to resist guarding was reduced experimentally. Guarding durations for manipulated and control males were equal, but manipulated females were guarded longer, suggesting that conflict exists and that females can effectively shorten guarding duration by their behavior. Moreover, we suggest that selection in the context of intersexual conflict may play an important role in the evolution of delayed oviposition and sperm-storage organs in mate-guarding crustaceans.     

Biparental mouthbrooding and guarding in a Tanganyikan CichlidHaplotaxodon microlepis

Differences between parental roles of males and females inHaplotaxodon microlepis (Cichlidae) were investigated in Lake Tanganyika, and the early ontogeny and growth of the species were studied in the aquarium. Eggs were mouthbrooded by the female, and it is suggested that small larvae (<9 mm in total length) were also mouthbrooded by females though such samples were not collected. Above this size the larvae began to feed, and parents jointly performed mouthbrooding and guarding until the young grew to 25–30 mm, nearly 2 months after spawning. Males and females mouthbrooded to the same extent, but when a part of the brood was released, females mainly guarded the released brood and males took the mouthbrooding role. Differences in parentalcare patterns betweenH. microlepis and other monogamous mouthbrooding cichlids are discussed.     

Size, operational sex ratio, and mate-guarding success of the carrion beetle, Necrophila americana

When male insects guard females until oviposition, the benefitsfrom last-male sperm precedence must outweigh the costs of relinquishingadditional fertilizations. The profitability of guarding isincreased when males guard large, fecund females and when femalesare scarce because fewer fertilizations are sacrificed. However,the male reproductive success is not only determined by theprofitability of guarding but also by his ability to maintainguarding. In this study, we used male carrion beetles (Necrophilaamericana) to examine the effects of sex ratio, male relativesize, and female quality on the ability to guard. First, wepresent a model of mate guarding that explores factors, suchas sperm precedence, sex ratio, male size, and female quality,that influence the profitability of postcopulatory riding. Ourmodel predicts that large N. americana males should preferentiallyguard the largest female only when the sex ratio is male biasedand sperm precedence is above 80%. In contrast, small malesgain little from guarding because they are not likely to maintainit and be the last male to mate. Then, we tested these predictionsby manipulating sex ratio, relative male size, and female quality.All males in equal sex ratio and large males in male-biasedsex ratio guarded females significantly longer than did malesin female-biased sex ratio. In male-biased sex ratio, largemales guarded significantly longer and achieved more takeoversthan small males. Large females were guarded longer. The successof guarding males in this beetle depends on their size relativeto other males and the operational sex ratio.     

Multiple Copulations and Post-Copulatory Guarding in a Free-Living Population of Sika Deer (Cervus nippon)

Multiple copulations by females and post‐copulatory guarding by males, were studied in a free‐living sika deer population, on Nozaki, an island of the Goto Group, off the coast of Kyushu, Japan. In 1990, 1991, and 1993, observations were carried out mainly on open grassland in the central section of the island. It was found, that 10 of 22 females in estrus copulated more than once and five of them copulated with several males (multi‐male copulations), while the remainder copulated with a single male (repeated copulation). Almost all copulations were followed by guarding behavior. Dominant males (DMs) guarded significantly longer and more effectively than subordinate males (SMs). Guarding was interrupted in four of the SMs, when the guarding male was forcibly driven away from a female by higher ranked males (‘take‐over’) and in three cases when the female left the guarding male spontaneously (‘escape’). The only interruption of guarding of a DM was caused when a lower ranked male sneaked towards the guarded female and mated with her briefly (sneak). Interruptions of guarding initiated by females (escapes) occurred more when they were guarded by SMs than by DMs. Our results suggest that female sika deer indulge in multiple copulations to seek better genetic quality, rather than insuring fertility, enhancing genetic diversity, or avoiding harassment. The post‐copulatory guarding by DMs appears to be more effective in the prevention of additional copulations by females with other males than guarding by SMs. Moreover, SMs decrease the duration of the pre‐copulatory phase to achieve copulation before having to give way to DMs.     

Indirect evidence that guarded quiescent deutonymph females invest energy to attract conspecific males in the Kanzawa spider mite (Acari: Tetranychidae)?

Because only the first mating results in fertilization in Tetranychus kanzawai (Acari: Tetranychidae), adult males guard quiescent deutonymph females (i.e., precopulatory mate guarding). A previous study reported that quiescent deutonymph females guarded by a male attract more conspecific males than solitary females and then hypothesized that guarded females release more chemical signals than solitary ones to attract males. Quiescent deutonymph females do not feed. If the hypothesis is appropriate, guarded females should invest energy in attracting males at the expense of investment in other activities, such as egg production. Therefore, we compared oviposition rates immediately after adult emergence between guarded females and solitary females. On the first day, the oviposition rate of guarded females was lower than that of solitary females. On the second day, however, there was no significant difference between female groups. These results suggest that guarded females invest energy in activities other than egg production before adult emergence and that the energetic cost is easily recoverable. We believe that our finding indirectly supports the hypothesis that guarded females release more chemical signals than solitary females to attract conspecific males.