Effects of resource holding potential and resource value on tenure at nest sites in sand gobies

Over a broad range of animal systems, male reproductive successdepends on resource holding potential (RHP) and resource quality.In a field study, we randomly combined males of different sizeswith nests of different sizes to investigate the relative roleof resource holding potential and resource quality in determininga sand goby, Pomatoschistus minutus, male's nest tenure. Individuallymarked small and large males were given either small or largeflowerpots for nests in isolation and were exposed to intrudersafter they had built nests. We found that nest tenure was longerfor big males and owners of big nests. In most cases (34 of51) the original nest owner was replaced by a bigger male. Thesereplacements by larger males were probably due to takeoversby stronger intruders. Replacement males were larger at bignests. Our results support resource defense theory, as individualswith higher RHP and more valuable resources defended their nestfor longer. On nine occasions males abandoned their nests. Ownersof these nests were larger than the nest owners that were replaced.Hence, our results may provide an example of a situation inwhich sand goby males are able to judge the reproductive valueof their current situation and act accordingly.     

Food affects the potential reproductive rates of sand goby females but not of males

The interspawning interval of female sand gobies, Pomatoschistusminutus, a batch-spawning fish with paternal care, was significantlyshorter when the fish were fed daily than when they were fedevery fourth day. The incubation time of males was not affectedby feeding, nor was the interbrood interval Males have an equalor higher potential reproductive rate than females. As femalesreproduce more slowly when food is scarce than when it is abundant,and males do not, the difference between the sexes in potentialreproductive rate increases when there is food shortage. Becauseof this difference, both male bias in operational sex ratioand intensity in male-male competition for mates are predictedto increase as food availability decreases. Furthermore, a tradeoffbetween current and future reproduction is demonstrated to operateonly when resources are limited, because the correlation betweenegg number of the first and second clutch was positive amonghigh-food females but negative among low-food females. The numberof eggs per female clutch did, however, not differ between treatmentsin first or second dutch. I conclude that operational sex ratioand sexual selection are expected to vary within and betweensand goby populations in accordance with prey availability     

Monogamy when there is potential for polygyny: tests of multiple hypotheses in a group-living fish

Monogamy within social groups where there exists a high potentialfor polygyny poses a challenge to our understanding of matingsystem evolution. Specifically, the traditional explanationthat monogamy evolves due to wide female dispersion, affordingmales little opportunity to defend multiple females, cannotapply. Instead, monogamy in groups potentially arises becausefemales compete for breeding resources such as breeding sites,food, and paternal care. We conducted manipulative experimentsto determine whether females compete over limiting resourceswithin groups of the obligate coral-dwelling goby, Paragobiodonxanthosomus (Gobiidae). Breeding females behaved aggressivelytoward individuals of their own sex and evicted subordinatefemales that were large and mature from the group. Experimentalremoval of nest sites caused breeding partners to breed in alternativenest sites, demonstrating that nest site limitation was notthe cause of female competition. Supplemental feeding resultedin an increase in the fecundity of breeding females but no maturationof subordinate females, demonstrating that food-limited femalefecundity was a likely cause of female competition. Finally,supplemental feeding of breeding pairs demonstrated that thedifference in eggs hatched by fed versus unfed males was lessthan the difference in eggs laid by fed versus unfed females,suggesting that paternal care limitation might also drive femalecompetition. These results suggest that competition over foodand possibly paternal care selects for dominant, breeding femalesto suppress the maturation of subordinate females to minimizecompetition. Monogamy in association with group living is thereforelikely to have evolved because female competition prevents malesfrom utilizing the potential for polygyny.     

Behaviour and success of sneaker males in the sand goby, Pomatoschistus minutus

In this study we investigated in laboratory conditions the presence of alternative reproductive tactics in the sand goby and describe proximate factors affecting their expression and success. We describe the reciprocal interactions of resident males, females, and sneaking males. The pre-spawning phase proved to be important for successful nest intrusions by sneakers. The number of sneakers had no effect on the frequency of successful intrusions. When small males had exclusive access to nest sites, they built a nest and courted females, showing a full behavioural repertoire. The intensity of courtship was, however, strongly positively correlated with body size. Using microsatellite DNA markers we assessed paternity shares of territorial and sneaker males in a subset of all replicates. Following successful nest intrusion sneaker males fertilised 5–10% of the eggs. Our interpretation of the results is that sneaking in the sand goby is a conditional tactic, one that is less successful than the normal nest guarding behaviour, at least for one spawning event. Received in revised form: 9 March 2001 Electronic Publication     

Sand goby (Pomatoschistus minutus) males exposed to an endocrine disrupting chemical fail in nest and mate competition

Endocrine disrupting chemicals (EDCs) are a widely studied group of chemicals that interfere with the endocrinology of organisms. So far, few studies have demonstrated the effect of EDCs on the reproductive behavior of aquatic wildlife. Here we show that sand goby males' (Pomatoschistus minutus) success in mating competition greatly decreases after an exposure for 7 to 24 days to 17α-ethinyl estradiol (EE2, measured concentration 4 ng L^− 1). The sand goby exhibits a polygynous mating system with male parental care, in which males compete for nest sites and females. The aim of this study was to test how EE2 exposure affects the ability of males to compete for breeding resources, i.e. nest sites and mates. First, EE2 exposed males competed over a nest site against a non-exposed, control male of the same size. Secondly, we examined male courtship behavior and female mate preferences for EE2 exposed males and similar-sized non-exposed, control males. In addition to the behavioral experiments we determined the zona radiata protein (Zrp) mRNA gene expression and measured morphometric indicators of sexual maturation. Our study revealed that EE2 treated males were not able to acquire or defend a nest site. Additionally, EE2 treated males spent significantly less time in active courtship and nest leading behavior than control males. As a result, females clearly preferred to mate with control males. However, we found no significant differences in Zrp mRNA expression or the morphometric indicators between treatments. Our study illustrates that exposure to this EDC can greatly reduce the chances of an individual reproducing successfully. Moreover, it demonstrates that severe behavioral effects can be seen before any effects are detectable at the molecular or morphometric level.     

The effect of food supplementation on the reproductive success of Black-billed Magpies Pica pica

The effect of supplementary food on the reproductive success of Black-billed Magpies Pica pica was studied in an urban habitat in the city of Edmonton, Alberta, Canada. We provided supplementary food (pelleted dog food) regularly from August 1986 to June 1988 in a feeder situated outside all known magpie territories. Magpies using the supplementary food showed (i) an advancement of 7 days in the initiation of laying, (ii) no increase in clutch-size, (iii) higher survival of nest contents (eggs and young) during a spring snow storm, (iv) an increase in rate of nestling weight gain, and (v) an increase in fledging success. Supplementary food did not prompt the fed pairs to select nest sites nearer the feeder in 1988 than in 1987. The feeder did not affect the breeding density because the nearest neighbour distances of fed magpies did not differ from unfed magpies in either year, nor did they differ between the two years for either fed or unfed magpies. This study provided evidence that magpies frequently renest even if their initial nests fail after the eggs hatch—a practice that has been reported to be absent or rare in other studies. Although brood reduction was equally frequent in the nests of food-supplemented and control pairs, the actual number of nestlings disappearing from the former (1.56 per nest) was significantly smaller than that from the latter (2.60 per nest), suggesting that the availability of food plays a major role in magpie reproductive success.     

Effects of a low oxygen environment on parental effort and filial cannibalism in the male sand goby, Pomatoschistus minutus

In fish, brood cycling parental males sometimes eat some orall of their eggs, a behavior termed filial cannibalism. Wetested predictions of filial cannibalism models related to thecost of parental care in the male sand goby, Pomatoschistusminutus, by increasing the parental effort (fanning expenditure)through reduced levels of dissolved oxygen to 39% in an experimentalgroup, whereas a control group had fully saturated water. Malesshowed both full-clutch cannibalism and partial-clutch cannibalismin both treatments. Giving the males one to three females tospawn with, we found that small clutches were completely eatenmore often than were larger ones, whereas partial-clutch cannibalismwas not affected by clutch size. Although treatment did notaffect filial cannibalism, it did affect a male's energy statesuch that males in the low oxygen treatment lost more body fat,indicating a greater fanning effort. This shows that males inthe low oxygen treatment allocated more energy to the presentbrood, potentially at the expense of future reproductive success.Our study strongly suggests that filial cannibalism in malesand gobies represents a strategic life-history decision asan investment in future reproductive success, and is not triggeredby a proximate need for food necessary for the male's own survival.Furthermore, males in the low oxygen treatment built nests withlarger entrances, and were less likely to rebuild their nestsafter destruction. Presumably, this makes fanning easier butthe nest more vulnerable to predators, suggesting a trade-offbetween fanning and nest defense.     

Reproductive energetics of the role reversing bushcricket,Kawanaphila nartee (Orthoptera: Tettigoniidae: Zaprochilinae)

Theory predicts that sexual differences in reproductive strategies arise because of differences in the magnitude of investment made by males and females in reproduction. In some bushcrickets, the typical sex role of competitive male and choosy female is reversed when populations are subject to nutrient stress. Here I present an energetic analysis of reproduction for the role reversing bushcricket, Kawanaphila nartee, that supports the contention that this sex role reversal is a consequence of reversal in the pattern of relative reproductive investment. When fed ad libitum, males spent 16% of their daily energy reserves on the spermatophore compared with 26% spent on calling to attract a mate. Females spent 29% of their daily energy reserves in producing and laying eggs. However, when allowed only limited access to food, female expenditure in eggs was reduced to 23% of daily reserves while male expenditure remained unchanged. After accounting for the incorporation of male nutrients into eggs, female energy expenditure in reproduction exceeded male expenditure when animals were fed ad libitum, but male expenditure exceeded female expenditure when diet was limited. This role reversal in relative energy expenditure that is associated with courtship role reversal supports classical and contemporary theories on the control of sexual selection.     

Daily foraging routines and feeding effort of a small bird feeding on a predictable resource

According to theory, a small diurnal bird living in a predictable environment should have the highest feeding effort as late as possible in the day in order to minimize the time it carries large and costly reserves. The feeding effort should also decline with increasing food availability. We tested both these ideas with the lesser spotted woodpecker (Dendrocopos minor). For most of the year, this bird feeds on wood-living insects in dead tree branches. This food supply is likely to be highly predictable on a daily scale. Our results corroborated the theory. We found that the proportion of time spent actively feeding was lower in the mornings (before noon) than in the afternoons. We also found that woodpeckers spent less time feeding the higher their food availability However, for a given food availability they spent more time feeding in the afternoons. This supports the idea that feeding is less and other activities are more valuable in the mornings given a predictable food resource. This is the first demonstration of daily routines in small birds concordant with a predictable environment. In spring, males but not females reduced their feeding time. This difference between the sexes may be related to their sex-specific reproductive effort.     

Does the reproductive status of male round gobies (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) influence their response to conspecific odours?

While pheromone communication has been well documented in the female round goby, male responses have not been examined. We determined if male round gobies responded to odours of their conspecifics, and whether that response varied with their reproductive status (reproductive and non-reproductive). No significant differences were observed for any treatment (dechlorinated water, or water conditioned by reproductive males (RM) and non-reproductive males (NRM) and females), although RM spent more time inside shelter than NRM (p < 0.05). Morphologically, length and weight did not vary, but gonadosomatic index, seminal vesicle somatic index and relative head width did differ between the two male groups (p = 0.000). Round goby males guard nests in cavities, and so a preference for shelter with increasing reproductive status is expected. Also, a lack of male responses to conspecific odours may be adaptive in this species, since males nest together in high densities, where increased aggression might prove detrimental for nesting males.     

The effects of supplemental food on reproduction in parental male smallmouth bass

Synopsis We provided supplemental food for parental male smallmouth bass,Micropterus dolomieu, to determine if food supply limits the reproductive performance of nesting males as measured by care duration, reproductive success and survival of adult males. Although supplementing the diet of parental males had a positive effect on all three reproductive measures, the experiment generated contrasting results in different years. In the first year, supplemental feeding only improved the survival of fed males versus unfed males. In the second year, supplemental feeding increased care duration and reproductive success of fed males but reduced their survival relative to unfed males. Our supplemental feeding improved current or future measures of reproductive performance, but not both simultaneously. The results appear to demonstrate phenotypic plasticity in the extent to which energy from supplemental feeding can be allocated to present or future reproduction.     

Effects of sex ratio on male behaviour and reproductive success in a field population of threespine sticklebacks (Gasterosteus aculeatus) (Pisces: Gasterosteidae)

Biased sex ratios of breeding threespine sticklebacks (Gasterosteus aculeatus L.) occur naturally in tide-pools of a Quibec saltmarsh. We experimentally manipulated sex ratios in certain pools to evaluate the effects on male behaviour and reproductive success (RS). Sticklebacks were stocked at male: female ratios of 1: I, 1: 2 and 2: 1 and observed for a 23-day inter-tidal period. In male-biased pools, only half of the males built nests, compared to nearly 100% in unbiased and female-biased pools. Males in male-biased pools also were less likely to rebuild after losing a nest, visited their nests less often, were more likely to abandon or destroy their nests, had lower RS (measured as the proportion of males hatching fry), but hatched fry sooner, than males in other pools. In female-biased pools, males built nests sooner, lost more nests due to nest-raiding by females, spent more time in aggression (proportion of time spent fighting and threatening), spent more time attacking female conspecifics than male conspecifics or heterospecifics, and courted more frequently, than males in other pools. Habituation to conspecific males, but not to females, occurred in all pools. These findings are discussed with respect to sexual selection theory.     

Spatial and temporal patterns of nest distribution influence sexual selection in a marine fish

In many species, the natural distribution of material resources important for reproduction can profoundly impact reproductive success among individuals and, hence, the opportunity and intensity of sexual selection. Here, we report on a field‐based experiment investigating the effects of nest aggregation on sexual selection in a fish, the sand goby Pomatoschistus minutus. We found that the distribution of potential nests (sparse versus aggregated nest treatments) affected patterns of nest colonization and reproductive success. Specifically, in the treatment with aggregated nesting resources, a greater proportion of nests remained unoccupied by sand goby males. Although the size of nesting males did not differ between treatments, eggs accumulated more rapidly when nests were sparsely distributed. We found that the opportunity for selection decreased over time with the accumulation of eggs in the nests in both the aggregated and sparse treatments. Moreover, the effect of male size on reproductive success was influenced by an interaction between nest distribution and time, with the selection gradient being highest right after nest colonization when nests were aggregated, while the opposite pattern was observed in the sparse nest treatment. Such findings highlight the vital role that environmental and social factors can play in determining the importance of male phenotypic traits (in this case, male size). More broadly, our results also underscore how the natural distribution of resources, both in space and time, can impact the strength of sexual selection acting on wild animal populations.     

Courtship Sounds Advertise Species Identity and Male Quality in Sympatric Pomatoschistus spp. Gobies

Acoustic signals can encode crucial information about species identity and individual quality. We recorded and compared male courtship drum sounds of the sand goby Pomatoschistus minutus and the painted goby P. pictus and examined if they can function in species recognition within sympatric populations. We also examined which acoustic features are related to male quality and the factors that affect female courtship in the sand goby, to determine whether vocalisations potentially play a role in mate assessment. Drums produced by the painted goby showed significantly higher dominant frequencies, higher sound pulse repetition rates and longer intervals between sounds than those of the sand goby. In the sand goby, male quality was predicted by visual and acoustic courtship signals. Regression analyses showed that sound amplitude was a good predictor of male length, whereas the duration of nest behaviour and active calling rate (i.e. excluding silent periods) were good predictors of male condition factor and fat reserves respectively. In addition, the level of female courtship was predicted by male nest behaviour. The results suggest that the frequency and temporal patterns of sounds can encode species identity, whereas sound amplitude and calling activity reflects male size and fat reserves. Visual courtship duration (nest-related behaviour) also seems relevant to mate choice, since it reflects male condition and is related to female courtship. Our work suggests that acoustic communication can contribute to mate choice in the sand goby group, and invites further study.     

Predation risk affects trade-off between nest guarding and foraging in Seychelles warblers

The fitness costs of egg loss for Seychelles warblers (Acrocephalus sechellensis)on Cousin Island are considerable because warblers have a single-eggclutch and no time to lay a successful replacement clutch. Onthe islands of Cousin and Cousine, with equal densities of Seychellesfodies (Foudia sechellarum), nearly 75% of artificial eggs placedin artificial nests were predated by fodies after 3 days. OnAride Island with no fodies present, loss of artificial eggswas not observed. Female warblers incubate the clutch, and malewarblers guard the clutch when females are absent. Deterrenceof fodies by male warblers is efficient: loss rate of eggs fromunattended warbler nests was seven times as high as from attendednests, and the more nest guarding, the lower the egg loss andthe higher the hatching success. Egg loss is independent ofthe amount of incubation by females. There is no trade-off betweenincubating and foraging by females. Nest guarding competes withforaging by males, and this trade-off has a more pronounced effecton egg loss when food availability is low. The transfer of breeding pairsfrom Cousin to either Cousine with egg-predating fodies or toAride without fodies allowed us to experimentally investigatethe presumed trade-off between nest guarding and foraging. OnCousine, individual males spent the same amount of time nestguarding and foraging as on Cousin, and egg loss was similarand inversely related to time spent nest guarding as on Cousin.Males that guarded their clutch on Cousin did not guard theclutch on Aride but allocated significantly more time to foragingand gained better body condition. Loss of warbler eggs on Aridewas not observed. Time allocation to incubating and foragingby individual females before and after both translocations remainedthe same.     

Size Dependent Male Reproductive Tactic in the Two-Spotted Goby (Gobiusculus flavescens)

Male investment in testes and sperm duct gland in the polygamous nest breeding two-spotted goby Gobiusculus flavescens (Fabricius) was investigated in relation to time in reproductive season and individual physical parameters. This small teleost fish is most likely the most abundant species found along the rocky shores of the North East Atlantic. The two-spotted goby has a single reproductive season, during which nest-caring males can raise several clutches of offspring. According to the literature the males are on average larger than the females. Here we report for the first time a population showing a reversal of this trend, with males on average being smaller than females, a difference likely caused by a large proportion of small males. Early in the breeding season these small males have typical sneaker characters, with relatively large testes and small seminal duct glands compared to the larger dominant territorial males. The presence of these two alternative male reproductive tactics is confirmed by histological studies, which shows the presence of sperm in the sperm duct glands (SDG) of smaller males, but not in the SDG of intermediate and larger males. To our knowledge, males with typical sneaker characters have not been reported in earlier studied populations of two-spotted goby. Interestingly we found that testes investment declined significantly over the course of the breeding season, and that this reduction was significantly more pronounced in small compared to the large males. Further, a significant increase in seminal duct gland (SDG) mass was observed for the smaller males over the breeding season. We propose that this indicates a possible shift in mating tactic by smaller males from a parasitic to a nest-holding tactic over the course of the breeding season. Thus, the observed size dependent plasticity in investment in SDG over time suggests that the reproductive tactic of G. flavescens is conditional, and possibly influenced by mate availability and male—male competition.     

Plastic sex-roles in the common goby – the effect of nest availability

One of the most important factors shaping animal sex-roles is the operational sex ratio, since a skew of this ratio promotes mating competition in the more abundant sex. In this study, we wanted to see if a change in sex-roles, related to an environmental induced change in operational sex ratio, could be demonstrated in a field situation. Common goby ( Pomatoschistus microps ) males build nests under bivalve shells and provide exclusive parental care. Therefore, available nest sites are crucial for successful breeding in this species and should also affect the operational sex ratio. We addressed the question of whether the availability of nest sites, through an effect on the operational sex ratio, affects courtship and mating competition in the common goby. We increased nest site availability in the field by adding potential nest material (bivalve shells) to one location. We then compared the reproductive behaviour of the gobies at this and another location with lower nest site abundance. In general, the reproductive behaviour differed as predicted. Under nest shortage, males occupied new nests and received eggs faster than under nest excess. Behavioural observations of nests revealed that males initiated courtship more often than females where nest sites were abundant, whereas female courtship dominated where nests were scarce. Males more often rejected females attempting to enter their nests under nest shortage, male sneaking attempts were also more common in this bay. Male-male aggression was frequent at both locations, whereas agonistic interactions between females only occurred in the nest shortage bay. Thus, males should be subject to sexual selection in both bays even though courtship roles differed. These results suggest that nest site availability affects mating competition and courtship roles in the common goby.     

The importance of sperm competition risk and nest appearance for male behavior and female choice in the sand goby, Pomatoschistus minutus

To test if an increased sperm competition risk affects malebehavior and mating decisions of both sexes, we performed twoexperiments using the sand goby, Pomatoschistus minutus, a nest-buildingfish with exclusive paternal care. In our first experiment,a nest-holding male, with a confined female, was sequentiallyexposed to a vial with a sneaker male or an empty vial. Whilemale courtship, nest building, displacement fanning, and timeoutside the nest were unaffected, individual males showed ahigher mucus preparation effort inside the nest in the presenceof a sneaker male than when alone. We found such mucus to containsperm, thus clearly suggesting an importance in sperm competition.In our second experiment, a female was free to spawn with twodifferent males, one of which was exposed to a confined sneakermale. Male mating success was not affected by the presence ofa sneaker male. However, the volume of sand the male had puton his nest was positively associated with female spawning decision,while nest-opening width was not. In a partial correlation offive traits thought to attract females (nest-opening width,sand volume, male courtship display, displacement fanning, andmale size), males that fanned well were found to also buildlarge nests or display intensely, but not both. This indicatesthat rather than being jacks-of-all-trades, individual malesfocus on a subset of traits for attracting females.     

Parent birds assess nest predation risk and adjust their reproductive strategies

Avian life history theory has long assumed that nest predation plays a minor role in shaping reproductive strategies. Yet, this assumption remains conspicuously untested by broad experiments that alter environmental risk of nest predation, despite the fact that nest predation is a major source of reproductive failure. Here, we examined whether parents can assess experimentally reduced nest predation risk and alter their reproductive strategies. We experimentally reduced nest predation risk and show that in safer environments parents increased investment in young through increased egg size, clutch mass, and the rate they fed nestlings. Parents also increased investment in female condition by increasing the rates that males fed incubating females at the nest, and decreasing the time that females spent incubating. These results demonstrate that birds can assess nest predation risk at large and that nest predation plays a key role in the expression of avian reproductive strategies.     

The effect of adult feeding on the fecundity,weight loss and survival of the pine beauty moth,Panolis flammea (D&S)

Summary Adult females of the pine beauty moth, Panolis flammea, fed on a saturated honey solution, laid significantly more eggs than those individuals given water only or not fed at all.At 20°, the longevity of both male and female moths was significantly increased when they were supplied with a food source. At 10° fed male months were not significantly longer lived than unfed moths. Fed female moths at 10° were however, significantly longer lived than unfed individuals. At 20°, fed male moths lived longer than fed females (8.17 days as compared with 5.95 days). At 10°, fed female moths lived longer than fed males (16.00 days compared with 11.41 days).At 20°, weight loss in unfed females over the reproductive period was proportional to the number of eggs laid and the survival time of the individual. In fed females, no significant weight loss occurred over the reproductive period, irrespective of the number of eggs laid or the longevity of the individual.Egg production, even in fed females, over the life span of an individual was only a fifth of the total eggs present in the reproductive tract at eclosion.It is concluded that adult females of P. flammea are likely to require an extra external food source to achieve their full reproductive potential.