Male Nest Choice in Sand Gobies, Pomatoschistus minutus

Although material resources can have a direct bearing on the fitness of both sexes, few studies have actually examined resource-based preferences from a male choice perspective. In sand gobies, Pomatoschistus minutus , the size of a male's nest influences his attractiveness to females and also dictates the number of eggs he can receive. Thus, one might expect males to prefer larger nests. However, an earlier study of marine sand gobies from a population with a surplus of nest sites and high nest predation found that males exhibited size-assortative nest preferences. Here, we investigated male nest preferences from a brackish population characterised by a chronic nest shortage but lower predation risk. A survey of naturally settled nests in the field (shells and rocks) showed a pattern of size-assortative nest occupancy consistent with the previously studied population, with larger males occupying larger (i.e. rock) nests. However, when offered a choice of potential nests in the absence of male competition, we found that all male gobies in our population, irrespective of their own body size, actually preferred larger nests. Moreover, a predilection towards large nests superseded any preferences based on nest colour. Our results not only indicate the existence of male preferences for material resources but, considered in the light of previous work, also suggest that such preferences may vary among populations and, importantly, may not necessarily be realised in a competitive setting.     

Reproductive success in male sunbleak, a recent invasive fish species in the U.K.

The spawning behaviour of male nest guarding sunbleak Leucaspius delineatus , a recent invasive species in southern England, was studied and quantified for the first time. In the absence of physical differences between territorial and non‐territorial males ( i.e . colour, size, etc .), the reproductive behaviour of territorial males was analysed and related to reproductive success. The results showed that females preferred high‐courting and highly aggressive males. The initial cue in female mate choice, however, was based on courtship, while aggression was the decisive behavioural trait in influencing mate choice, providing a direct signal of physical condition and 'paternal competence'. Some males picked nest sites which were subsequently preferred by other males taking over the nest of a previous male ('communal nest'), with the new territorial male adopting the eggs already present at the nest. It appears that either due to female preference for nests already containing eggs or lower rates of sired egg predation by dilution among unrelated eggs, sunbleak males have adopted the mating strategy of allopaternal care.     

Reproductive-tactic-specific variation in sperm swimming speeds in a shell-brooding cichlid

Theory predicts that males experiencing elevated levels of sperm competition will invest more in gonads and produce faster-swimming sperm. Although there is ample evidence in support of the first prediction, few studies have examined sperm swimming speed in relation to sperm competition. In this study, we tested these predictions from sperm competition theory by examining sperm characteristics in Telmatochromis vittatus, a small shell-brooding cichlid fish endemic to Lake Tanganyika. Males exhibit four different reproductive tactics: pirate, territorial, satellite, and sneaker. Pirate males temporarily displace all other competing males from a shell nest, whereas sneaker males always release sperm in the presence of territorial and satellite males. Due to the fact that sneakers spawn in the presence of another male, sneakers face the highest levels of sperm competition and pirates the lowest, whereas satellites and territorials experience intermediate levels. In accordance with predictions, sperm from sneakers swam faster than sperm from males adopting the other reproductive tactics, whereas sperm from pirates was slowest. Interestingly, we were unable to detect any variation in sperm tail length among these reproductive tactics. Thus, sperm competition appears to have influenced sperm energetics in this species without having any influence on sperm size.     

Nest use by territorial males in a shell-brooding cichlid: the effect of reproductive parasitism

Male Telmatochromis vittatus, a substrate-brooding Tanganyikan cichlid, exhibit two parasitic reproductive tactics: takeover of spawning by larger males (pirates) and sneaking by smaller males (sneakers). Medium-sized males are territorial and pair-spawn within nests of clumped shells that harbor several resident females that are potential mates of the territory owner. To study nest use by territorial males, we analyzed the relationship between the body size of territorial males and nest quality, the number of females per nest, the distance between nests, and the frequency and intensity of reproductive parasitism. The size of males was not correlated with nest quality, the number of females, or the number of sneakers, but was negatively correlated with the frequency of intrusion by pirates and was positively correlated with the distance between nests. Territorial males effectively defended nests against sneakers but failed to defend against pirates. These results suggest that larger territorial males selected nests that have a lower risk of usurpation of spawning. We hypothesize that the risk of intrusion by pirate males affects the selection of nests by territorial males in this species.     

SEXUAL SELECTION AND FITNESS VARIATION IN A POPULATION OF SMALLMOUTH BASS,MICROPTERUS DOLOMIEUI (PISCES: CENTRARCHIDAE)

Monogamy is often presumed to constrain mating variance and restrict the action of sexual selection. We examined the reproductive patterns of a monogamous population of smallmouth bass (Micropterus dolomieui), and attempted to identify sources of within-season fitness variation among females and known-age males. Many males did not acquire a nest site, and many territorial males were unsuccessful in acquiring a mate. The likelihood that territorial males mated depended on several aspects of nest sites. Mated males of age three were larger than the average size of age-three males in the population. The mean sizes of age-four and age-five mated males were not different from the average of same-age males in the population. Thus, selection resulting from the acquisition of a mate favored large size among only age-three males. Timing of nest construction and breeding among territorial males was negatively related to male size and did not depend on male age after taking male size into account. Indirect evidence (numbers of eggs deposited in nests) suggests that the timing of spawning among females was also negatively related to female size. Fertility selection favored early reproduction within the season by males of all ages, but large male size was favored among only age-four males. The combined early breeding of fecund females and female mate choice of large males may explain the positive correlation between the size of age-four males and the number of eggs acquired. Despite large differences of female fecundity, however, the variance of relative mate number contributed about two times more than the variance of relative fertility among females to the total variance of relative fitness within each sex.     

Size-assortative nest choice in the absence of competition in males of the sand goby,Pomatoschistus minutus

Synopsis Size-assortative choice of nest site is commonly found among fish with paternal care. This pattern can be explained either by the superior competitive ability of large males for preferred large nests, or by a choice of nest size that is related to the male's own body size. Choice of nest size was studied in males of a small marine fish, the sand goby Pomatoschistus minutus, by providing artificial nest sites in laboratory and field experiments. A positive correlation between nest size and male size, indicating a size-assortative nest choice process, was found in both aquarium and field experiments. This is argued to be the outcome of a preference for an optimal nest size and not to be due to competition, as (a) a surplus of nest sites was available to the individuals in nature; (b) in the aquarium experiments, single males of all sizes, when offered a choice between a smaller and a larger nest site, showed a preference for nests in proportion to their body size and not for the largest available nests; and (c) in the aquaria, when single males were offered only a small or only a large nest site, small males abstained from building nests in the large sites, whereas in the small nest sites males of all sizes built nests. My conclusion is that size assortative nest choice occurred in the absence of competition, which thus is not a necessary condition for size-assortative nesting, contrary to what is commonly assumed in many studies. This preference may represent a compromise (or trade-off) between the advantage of a nest large enough to accommodate many eggs and a smaller one with lower maintenance and defence costs.     

Cooperative nest defence in red-winged blackbirds: reciprocal altruism,kinship or by-product mutualism?

Male red-winged blackbirds (Agelaius phoeniceus) often cooperate with their neighbours in defending nests against predators. Some studies have suggested that this is an example of by-product mutualism, whereas others have suggested the possibility of reciprocal altruism. No study has addressed the possibility of kin-selected cooperation in nest defence in this species. Reciprocal altruism, kin selection and by-product mutualism are not mutually exclusive alternatives, but few studies of territorial neighbours have tested for multiple mechanisms simultaneously. We test for these three possibilities in a population of red-winged blackbirds. We used simulated defections to test for reciprocal altruism. We used analysis of microsatellite loci to test for kin selection between adult male neighbours. We also used microsatellite loci to test for by-product mutualism resulting from nest defence of offspring sired on neighbouring territories. We found that male red-winged blackbirds cooperate in nest defence primarily as a form of reciprocal altruism. Experimental males reduced their level of nest defence relative to controls following simulated defection by a neighbour. In contrast to some earlier studies, we found no evidence for by-product mutualism: males did not defend nests where they had sired extra-pair offspring. We also found no evidence for kin selection: males were no more cooperative with more closely related neighbours. Considered alongside the results from other studies, our study suggests that mechanisms stabilizing cooperation in red-winged blackbirds may vary among populations.     

Body Size Correlates with Fertilization Success but not Gonad Size in Grass Goby Territorial Males

In fish species with alternative male mating tactics, sperm competition typically occurs when small males that are unsuccessful in direct contests steal fertilization opportunities from large dominant males. In the grass goby Zosterisessor ophiocephalus, large territorial males defend and court females from nest sites, while small sneaker males obtain matings by sneaking into nests. Parentage assignment of 688 eggs from 8 different nests sampled in the 2003–2004 breeding season revealed a high level of sperm competition. Fertilization success of territorial males was very high but in all nests sneakers also contributed to the progeny. In territorial males, fertilization success correlated positively with male body size. Gonadal investment was explored in a sample of 126 grass gobies collected during the period 1995–1996 in the same area (61 territorial males and 65 sneakers). Correlation between body weight and testis weight was positive and significant for sneaker males, while correlation was virtually equal to zero in territorial males. That body size in territorial males is correlated with fertilization success but not gonad size suggests that males allocate much more energy into growth and relatively little into sperm production once the needed size to become territorial is attained. The increased paternity of larger territorial males might be due to a more effective defense of the nest in comparison with smaller territorial males.     

Male and female preferences for nest characteristics under paternal care

Nests play a critical role for offspring development across the animal kingdom. Nest quality may contribute to the builder's extended phenotype and serve as an ornament during mate choice. We examined male and female nest choice in the common goby (Pomatoschistus microps), a benthic fish with male‐only parental care where females deposit eggs in male‐built nests. Using prebuilt nest models, we independently manipulated two candidate nest quality traits: (a) nest entrance width with a role in oxygen ventilation, and (b) extent of sand cover with a role in camouflage. In simultaneous choice trials, male gobies exhibited no preference for any nest model type. This suggests that initial characteristics of a nesting substrate have minor importance for males, which usually remodel the nest. Females were given a choice between two males occupying either entrance‐ or cover‐manipulated nests. The same pair of males was then exposed to a second female but now with alternated nest types assigned. Most females were consistent in choosing the same, typically the heavier male of the two regardless of nest properties. However, the females that chose the same nest regardless of the male preferred low over high sand coverage and narrow over wide nest entrance. Our results indicate that females base their mating decision on a combination of male phenotype and nest traits. While we found no indication that females are attracted to highly decorated nests, our study is the first in fishes to disentangle a preference for narrow (and thus more protective) nest entrances independent of nest coverage.     

Size-assortative nest preference in a paternal brooding blenny Rhabdoblennius ellipes (Jordan & Starks)

Field and laboratory studies were conducted to clarify the processes contributing to the size-assortative nest use by males of a paternal brooding blenny Rhabdoblennius ellipes . The use of natural and artificial nests by males in the field was highly size-assortative; males used smaller nests even when larger nests were available. Aquarium nest choice experiments without a competitive male strongly suggested that the size-assortative nest use by males is not the result of male–male competition for larger nests, but male preference for size-matched nests. Males were likely to choose nests on the basis of the nest entrance size rather than nest length and volume, suggesting that the size-assortative nest preference in this species is an adaptation to predation pressure against eggs and resident males.     

Elaborate Nests in a Weaverbird: A Role for Female Choice?

Studies on sexual selection have focused on behaviour and morphology, but several groups of animals build elaborate structures associated with acquiring a mate. I investigated female choice for nests built by male baya weavers (Ploceus philippinus). Nest choice by females should be strong, as nests are obvious direct benefits provided by males. I used a field experiment supplemented with correlational information to ask whether females appear to base mate choice decisions on male behaviour, nest architecture, and nest location. When the nests of highly visited males were exchanged with those of poorly visited males, female visits remained highest at the original male and location. I found no relationship between female choice and male display or other behaviour. Correlational analyses show that nest location was a better predictor of female choice than was nest architecture. These data suggest that current female choice is driven more by access to safe nesting sites rather than to well‐built nests, possibly because all males are able to build nests of adequate quality. However, nest architecture is unlikely to be irrelevant to females, and its role deserves further investigation.     

Short‐term dynamics of nest occupancy in an allopaternal species,the tessellated darter Etheostoma olmstedi

To inquire how male size interacts with alloparental behaviour and mating success in the tessellated darter Etheostoma olmstedi, males were given a choice of nests with or without eggs; subsequent nest occupancy, takeovers and egg deposits were monitored. Subordinate males readily occupied available nests with eggs but were often evicted by dominant males, suggesting that males of all sizes compete for the opportunity to provide allopaternal care in this species.     

Size-assortative nest choice by the Japanese fluvial sculpin in the presence of male-male competition

Nest choice by the Japanese fluvial sculpin Cottus pollux and its relationship to male reproductive success, was examined in a natural habitat, central Japan. The larger males tended to occupy the deeper nest cavities, suggesting size-assortative nest choice by the species. Stepwise regression analysis showed that cavity area was correlated positively to male reproductive success. Nest choice experiments without a competitive male indicated an absence of size-assortative nest choice, but larger males tended to choose a large nest when another male was present, suggesting that size-assortative nest choice by the species is attributable to male-male competition for large nests, such being related to a resource-defence polygynous mating system.     

Breeding pattern and nest guarding in Sicyopterus lagocephalus, a widespread amphidromous Gobiidae

Amphidromous gobies are usually nest spawners. Females lay a large number of small eggs under stones or onto plant stems, leaves or roots while males take care of the clutch until hatching. This study investigates the breeding pattern and paternal investment of Sicyopterus lagocephalus in a stream on Reunion Island. In February 2007 and January 2010, a total of 170 nests were found and the presence of a goby was recorded at 61 of them. The number of eggs in the nests ranged from 5,424 to 112,000 with an average number of 28,629. We showed that males accepted a single female spawning in the nest and cared for the eggs until hatching. The probability for a nest to be guarded increased with the number of eggs within it, suggesting that paternal investment depends on a trade-off between the reproductive value of the current reproduction and future nesting events. We showed that large nest stones were occupied by large males (TL >80 mm), whereas smaller males (TL <50 mm) were found under smaller cobbles, probably because of male–male competition for available nests. Our results suggest that the male’s choice relies upon a similarity to the female size, while the female’s choice was based on both body and nest stone sizes.     

Have your cake and eat it too: male sand gobies show more parental care in the presence of female partners

Traditionally, male parental effort and mate attraction effortare expected to be in conflict as they compete for the sameresource budget. However, the quality of care provided by themale may be of a direct benefit to females and may provide animportant mate choice cue. In a laboratory experiment, we examinedhow males modified their parental behavior with respect to matingopportunity by allowing male sand gobies to mate with a singlefemale either in a big or small nest (a constraint on futuremating potential). We then exposed half of these males to thevisual stimulus from additional females and recorded male eggfanning and nest building (two components of care), courtshipbehavior, and reproductive success through out the brood cycle.We found that males fanned longer and more frequently and didmore nest construction in the presence of females and in bignests. Males guarding large nests courted females more thandid males guarding small nests. All males consumed eggs duringthe brood cycle, but complete clutch cannibalism was most frequentwhen males were guarding small nests in the absence of females.The pattern of filial cannibalism that we observed suggeststhat males prematurely terminated care when their reproductivepotential was low, that is, when there was little nest spacefor additional mating and no mates present. We found no supportfor a trade-off between mate attraction and parental care. Indeed,taken together our results suggest that males may use parentalcare as a courtship strategy and that males who invest in mateattraction also have higher parental effort.     

Males call more from wetter nests: effects of substrate water potential on reproductive behaviours of terrestrial toadlets

Laboratory studies of terrestrial-breeding frogs have demonstrated that wetter substrates produce fitter offspring but the relevance of substrate wetness to adult reproductive strategies is unknown. I hypothesized that male toadlets (Pseudophryne bibronii) would select wetter areas for nesting and would advertise wet nests strongly, and tested these predictions by manipulating water potentials at a breeding site. Males preferred to nest in the wettest areas, and called at greater rates on almost twice as many nights as males occupying drier nests. Overall, males that mated called on significantly more nights than unmated males. Hence, because males occupying wet nests called more, they also mated more and in 19 out of 20 cases, oviposition occurred in wet nests that were suitable for embryonic development. Males occupying drier nests may have risked dehydration by calling, and so were less able to signal to females. Hydration states therefore have the potential to influence the reproductive success of terrestrial male frogs.     

The Role of Agonistic Sounds in Male Nest Defence in the Painted Goby Pomatoschistus pictus

Animals often vocalize during territorial challenges as acoustic signals may indicate motivation and fighting ability and contribute to reduce aggressive escalation. Here, we tested the function of agonistic sounds in territorial defence in the painted goby. Pomatoschistus pictus, a small vocal marine fish that defends nests during the breeding season. We first measured the number of times a male approached, avoided, explored, entered and exited two unattended nests associated with either conspecific agonistic sounds or a control: silence or white noise. Acoustic stimuli were played back when the male approached a nest. In a second experimental set, we added visual stimuli, consisting of a conspecific male in a small confinement aquarium near each nest. Even though we found no effect of the visual stimuli, the sound playbacks induced similar effects in both experimental conditions. In the sound vs. silence treatment, we found that when males approached a nest, the playback of conspecific sounds usually triggered avoidance. However, this behaviour did not last as in longer periods males visited nests associated with agonistic sounds more often than silent ones. When the control was white noise, we found no significant effect of the playback treatment in male behaviour. Although we cannot exclude the possibility that other sounds may dissuade nest occupation, our results suggest that agonistic sounds act as territorial intrusion deterrents but are insufficient to prevent nest intrusion on their own. Further studies are needed to test the significance of sound production rate, spectral content and temporal patterns to deter territorial intrusion in fish.     

Female preference for nest size in the stream goby Rhinogobius sp DA

Females of the stream goby Rhinogobius sp. DA with paternal care favor males courting in fast water currents, whereby they mate males of high parental ability. Here we examined female choice of male nest size of this goby in laboratory. The dichotomous choice experiment clearly indicated that females prefer large nests. Spawning at large nests seems to improve egg survival rates in natural habitats in this goby. We discuss the possibility of multiple criteria in mate choice of this goby.     

Big is better: the importance of size for reproductive success in male Pomatoschistus minutus (Pallas) (Pisces, Gobiidae)

The sand goby, Pomatoschistus minutus , was used in aquarium experiments to study the importance of body size for the ability of males to gain access to nest sites and mates. When several male and female P. minutus were allowed to reproduce together, on average, half of the males built nests, and half of those males obtained eggs in their nests. Males with nests were significantly larger than males without nests, and nests with eggs belonged to males larger than the males with empty nests. In another experiment, when two males were competing for one nest, the largest male occupied the nest when both fish were put into the aquarium simultaneously. However, when the smaller male had been allowed to establish a nest before the larger male was introduced, the small male could usually retain the nest. Males with a body length < 50 mm did not build nests at all in the early part of the breeding season. In female choice experiments, no preference for larger males was found. Thus, male-male competition for nest sites and behavioural differences between different sized males seem to be the main factors influencing the non-random mating success in male P. minutus.     

Territorial responses of male blue tits, Cyanistes caeruleus , to UV-manipulated neighbours

Multiple phenotypic traits can affect the outcome of interactions among territorial animals. Individuals may use current and previously acquired information on phenotypic traits to assess the competitive ability of opponents and adjust the strength of their response depending on the threat the opponent poses. In birds, colourful plumage and song are widespread phenotypic traits. Recent work has shown that ultraviolet (UV) plumage reflectance may be used by males in assessing an opponent’s strength and by females in mate choice. In the present study, we investigated whether and how territorial male blue tits, Cyanistes caeruleus, use previously acquired information from UV reflectance of the crown feathers of neighbours to adjust their response to playback of song of these neighbours simulated to intrude their territory. We compared responses to neighbours with those to unfamiliar strangers with unknown plumage features. We found that subject males with UV-enhanced neighbours responded more strongly to these neighbours than to strangers, i.e. showed more flights, used songs without trill and tended to overlap more songs. Subject males with UV-reduced neighbours gave a lower or similar response to neighbours compared to strangers. This indicates that male blue tits combine previously acquired information about an intruder’s plumage with familiarity of its song, and that their response depends on the perceived quality of the neighbour. This study provides evidence that familiarity in combination with multiple signals of quality may influence territorial relations among neighbours.