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.     

Interactive effects of time and vegetation on reproduction of redshanks (Tringa totanus) breeding in Wadden Sea salt marshes

As shown for various species, nesting waders are non-randomly distributed on wetlands and preferentially select riparian nest-sites adjacent to limnic or marine waterbodies. Studying the redshank Tringa totanus, we tested the hypotheses that, in a coastal wader species which conceals its clutch in vegetation, predation and hatching success are affected by vegetation zonation, and that breeding in lower salt marsh areas has negative consequences for reproduction. We further predicted effects of timing of breeding and breeding experience/age of adults potentially reflected by egg biometrics both on nest-site selection and reproduction. Effects of vegetation, space, time and individual quality on hatching success of redshanks were studied in the German part of the Wadden Sea. Dominant plant species, vertical vegetation structure and nest concealment varied significantly between nests. Variation in nest concealment was relatively low: about 90% of clutches were classified as being well concealed. This variation was explainable by vegetation structure but not by vegetation composition at the nest-site, distance to shoreline, and time of clutch initiation. Vertical vegetation structure varied by dominant plant species but not by distance to shoreline and time of clutch initiation. Hatching success of clutches was low (10.6%) due to high predation (daily predation rate: 7.4%). Hatching success and duration of clutch survival were negatively and predation positively related to the date of clutch initiation. Furthermore, negative relationships were found between egg size and predation and duration of survival, respectively. We assume that concealed nests, early breeding and breeding experience diminish predation in salt marsh breeding redshanks. Thus, redshank reproduction appears to be affected by interactive effects of timing of breeding and vegetation facilitating early breeding. In contrast to open-nesting species, breeding in riparian habitats next to waterbodies may be disadvantageous for species breeding concealed in vegetation if these are covered by less structured vegetation.     

Variation in age at first reproduction of male Japanese fluvial sculpin induced by the timing of parental reproduction

The relationships between age and size at reproduction and lifetime reproductive output of male Japanese fluvial sculpin Cottus pollux were estimated by a mark-recapture study. Although all males were physiologically capable of breeding at age 2 years, age at first successful reproduction varied amongst individuals. Males with delayed reproduction had lower net reproductive rate than males that bred at age 2 years on average suggesting that age at first reproduction was a conditional strategy. Males that delayed reproduction were significantly smaller at age 1 and 2 years than males that bred at age 2 years. Despite no significant difference in body size of hatched yolk-sac larvae between the early and late phase of the breeding season, by May of the first year of life, progeny from nests in the early phase had hatched earlier and were larger than those from the nests in the late phase. The results suggested an important effect of timing of reproduction of parents on the growth and subsequent age at first reproduction of their progeny.     

Effects of parental body condition and size on reproductive success in a tenebrionid beetle with biparental care

Abstract. 1. The beetle Parastizopus armaticeps (Coleoptera: Tenebrionidae) inhabits the Kalahari desert of southern Africa, constructs breeding burrows after rainfall, and shows extensive biparental care. Previous work has shown that it is predominantly male size, not female size, that determines breeding success; however, in the field these beetles show size assortative mating. This might obscure or override effects of female size on reproduction. Moreover, the inaccessibility of the breeding burrows makes it impossible to test effects of female and male size on offspring development and survival before adulthood. 2. To disentangle the effects of male and female length, body mass, and body condition on reproductive success, males and females were paired randomly in small breeding cages in the laboratory (n = 887 breeding pairs). The construction of the breeding cages allowed a clear view of the brood chamber contents at each stage in offspring development. Larva, pupa, and imago numbers and development were monitored daily, and imago mass at hatching from the pupa (hatchlings), offspring mass, and offspring body length at complete exoskeleton melanisation (juveniles) were determined. 3. There was a weak positive correlation between body condition and body length for females only. Breeding chronology was related to male body condition: males in better condition were fast to start and finish a breeding bout. Males in better condition produced heavier hatchlings and juveniles, and larger‐sized males produced larger‐sized juveniles. In contrast, numbers of larvae and juveniles produced were determined mainly by female length and body condition: larger females in better condition hatched more larvae and produced more offspring. 4. The results suggest that male size and condition will be the most important determinant of reproductive success under relatively dry conditions, when burrow length is critical for reproductive success. Female size might be more important for the pair's reproductive success under wet breeding conditions, when burrow length is less critical for successful reproduction.     

Space use by the Japanese fluvial sculpin, Cottus pollux, related to spatio-temporal limitations in nest resources

Space use by the Japanese fluvial sculpin, Cottus pollux, as related to nest availability was studied in a Japanese mountain stream. Males had larger home ranges than females before breeding. A measure of nest resource limitation was L < 1 and the same nest rock was used by two males successively in three cases, indicating a limited supply of suitable nest rocks for males. All males included the nest rocks in their home ranges, but the timing of nest occupation was apparently dependent on male size, suggesting the presence of male-male competition for the nest rocks. Females, on the other hand, exhibited two different patterns in space use, independent of body size: (1) they spawned in their home ranges, or (2) they spawned out of their home ranges and returned. Nest cavity area did not affect male reproductive success. Spatial separation between female home ranges and nest location was the main reason for the females' movement out of their home ranges. The limitation in the male mating phase (5.7 days on average) might also have forced the females to move out of their home range to spawn. The reason behind such temporal limitation in the male mating interval was discussed from the viewpoint of the energy cost accompanying male parental care.     

Effects of size at metamorphosis on stonefly fecundity, longevity, and reproductive success

Many organisms with complex life cycles show considerable variation in size and timing at metamorphosis. Adult males of Megarcyssignata (Plecoptera: Perlodidae) are significantly smaller than females and emerge before females (protandry) from two western Colorado streams. During summer 1992 stoneflies from a trout stream emerged earlier in the season and at larger sizes than those from a colder fishless stream, and size at metamorphosis did not change over the emergence period in either stream. We performed two experiments to determine whether variation in size at metamorphosis affected the fecundity, reproductive success and longevity of individuals of this stonefly species and if total lifetime fecundity was affected by the number of matings. In the first experiment, total lifetime fecundity (eggs oviposited) was determined for adult females held in small plastic cages in the field. Males were removed after one copulation, or pairs were left together for life and allowed to multiply mate. Most copulations occurred in the first few days of the experiment. Females in treatments allowing multiple matings had significantly lower total lifetime fecundity and shorter adult longevity than females that only mated once. Multiple matings also reduced longevity of males. Fecundity increased significantly with female body mass at emergence, but only for females that mated once. While multiple matings eliminated the fecundity advantage of large female body size, number of matings did not affect the significant positive relationship between body mass at metamorphosis and longevity of males or females. In a second experiment designed to determine if body mass at emergence affected male mating success, we placed one large and one small male Megarcys in an observation arena containing one female and recorded which male obtained the first mating. The large and the small male had equal probabilities of copulating with the female. Copulations usually lasted all night, and the unmated male made frequent, but unsuccessful attempts to take over the copulating female. Our data suggest that selection pressures determining body size at metamorphosis may operate independently on males and females, resulting in evolution of sexual size dimorphism, protandry, and mating early in the adult stage. We emphasize the importance of interpreting the fitness consequences of larval growth and development on the timing of and size at metamorphosis in the context of the complete life cycle. Received: 1 July 1997 / Accepted: 12 November 1997     

Tactic changes in dusky frillgoby Bathygobius fuscus sneaker males: effects of body size and nest availability

Field and laboratory studies were conducted to examine the effects of nest availability and body size on changes in male mating tactics from sneaking to nest‐holding in the dusky frillgoby Bathygobius fuscus. In the field, the body size of nest‐holding males decreased from early to mid‐breeding season, suggesting the possibility of a change in the tactics of sneaker males to nest‐holding. Many sneaker males did not use vacant spawning nests even when size‐matched nests were available, but they continued to reproduce as sneakers. Similarly, in aquarium experiments with available vacant nests, some sneaker males became nest‐holders irrespective of their body size, but some did not. These results showed that nest availability is not a limiting factor for changes in tactics by sneaker males in this species. Because tactic‐unchanged sneaker males were co‐housed with larger nest‐holding males in the tanks, the body size of nearby nest‐holding males may have affected the decision to change tactics for sneaker males. Moreover, smaller individuals among tactic‐changed males tended to spend more time until spawning, probably because they had relatively larger costs and smaller benefits of reproduction as nest‐holding males compared to larger males.     

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.     

Colonial Nesting and the Importance of the Brood Size in Male Parasitic Reproduction of the Mediterranean Damselfish Chromis chromis (Pisces: Pomacentridae)

We investigated male parasitic spawning in a protected natural population of Mediterranean damselfish. Chromis chromis nested in colonies, inside which males showed a high variability in mating success. Our field observations indicate that the egg batches obtained by the most successful fish were five times bigger than the ones obtained by the less successful fish and many males never received ovipositions. On the other hand, reproductive parasitism was a common tactic within the colony. Successful nesting males sneaked into their neighbours' nests depending on the amount of eggs in their nest, with small clutch size inducing the males to parasitic reproduction. Males failing to receive egg depositions on their nests showed a significantly higher parasitism rate than successful males. Non-territorial males occupied stations in the water column above the breeding grounds and whenever the opportunity arose, they disrupted spawning in progress, stealing copulation with females. We observed that the likelihood of males being parasitized by sneakers was not correlated with the size of their own clutch; on the contrary, it depended both on the number of neighbouring nests and on the number of neighbouring males with barren nests (i.e. unsuccessful males). No correlation was found between parasitic behaviour and male size, suggesting males may switch between spawning in their own and in their neighbour's nests depending on mating opportunity. The hypothesis that colonial nesting facilitates parasitic reproduction is here discussed.     

Correlates of male reproductive success in Padogobius martensi (Gobiidae)

Factors affecting male reproductive success were investigated in a natural population of Padogobius martensi , a small freshwater goby showing paternal care. Males were found to be polygynous and their mating success was related both to their body length and to the size of the nest occupied. Body size was a good predictor for the presence of eggs in the nest. The number of egg batches guarded by a male was highly correlated with the area of his nest. Mature males with a total length of 48 mm or less (1-year-old individuals) occupied nest sites in breeding areas but were never found with eggs in their nests. The components of sexual selection which may determine the variance in mating success are discussed.     

Handicapped females receive more feedings during incubation from their mates: support for the female nutrition hypothesis

The female nutrition hypothesis posits that provisioning intensity of incubating females by their mates may depend on female needs and ensure proper incubation and a corresponding high hatching and breeding success of breeding pairs. Here, we have handicapped female pied flycatchers Ficedula hypoleuca at the beginning of incubation by clipping two primaries on each wing and filmed nests during incubation and later nestling provisioning to estimate male involvement in incubation feeding at the nest and in offspring care. Incubation feeding was more frequent at late nests. Correcting for this seasonal effect, incubation feeding was significantly affected by treatment and twice as high at experimental as at control nests. There was no effect of the experiment on female incubation attendance. The handicap did not result in any effect on hatching and breeding success, nestling growth and male or female provisioning and mass at the end of the nestling period. Males adjust their incubation feeding activity at the nest to female energetic requirements during incubation.     

Timing of current reproduction directly affects future reproductive output in European coots

Life-history theory suggests that the variation in the seasonal timing of reproduction within populations may be explained on the basis of individual optimization. Optimal breeding times would vary between individuals as a result of trade-offs between fitness components. The existence of such trade-offs has seldom been tested empirically. We experimentally investigated the consequences of altered timing of current reproduction for future reproductive output in the European coot (Fulica atra). First clutches of different laying date were cross-fostered between nests, and parents thereby experienced a delay or an advance in the hatching date. The probability and success of a second brood, adult survival until and reproduction in the next season were then compared to the natural variation among control pairs. Among control pairs the probability of a second brood declined with the progress of season. Delayed pairs were less likely and advanced pairs were more likely to produce a second brood. These changes were quantitatively as predicted from the natural seasonal decline. The number of eggs in the second clutch was positively related to egg number in the first clutch and negatively related to laying date. Compared to the natural variation, delayed females had more and advanced females had fewer eggs in their second clutch. The size of the second brood declined with season, but there was no significant effect of delay or advance. Local adult survival was higher following a delay and reduced following an advance. The effect of the experiment on adult survival was independent of sex. Laying date and clutch size of females breeding in the next year were not affected by treatment. The study demonstrates the existence of a trade-off between increased probability of a second brood and decreased parental survival for early breeding. Timing-dependent effects of current reproduction on future reproductive output may thus play an important role in the evolution of the seasonal timing of reproduction.     

栗头鹟莺繁殖生态的初步观察

鹟莺属(Seicercus)鸟类的繁殖资料十分缺乏。2015年4~8月,采用录像全程监控的方法,在贵州宽阔水国家级自然保护区,对4巢栗头鹟莺(S.castaniceps)的繁殖过程进行了完整观察。栗头鹟莺的繁殖期主要集中在5~7月,巢址选择专一性较强,均筑巢于公路边的土坎内壁,距路(1.3±1.2)m,巢为球状侧开口,巢材主要为新鲜苔藓及细草根,巢高(2.2±0.6)m,巢宽(10.9±1.5)cm,杯宽(3.3±0.5)cm,巢深(9.5±1.9)cm,杯深(5.5±1.0)cm。窝卵数(4.5±0.6)枚(4~5枚),卵重(0.92±0.04)g,卵长(14.30±0.30)mm,卵宽(11.22±0.23)mm,卵体积(0.92±0.05)mm3(n=18)。亲鸟在产满窝卵后开始孵卵,孵卵期12~13 d,在孵卵中期(第5~9天)亲鸟的孵卵时间开始增加,翻卵次数增多,在孵卵后期(第10~13天)亲鸟的孵卵时间和翻卵次数基本保持不变。育雏期13~14 d。雌、雄共同育雏,雏鸟在3日龄时,体重和跗跖开始显著增长,在7日龄时,增长速度变缓。孵化率为88.9%(16/18),营巢成效为100%,出飞成效为3.3只/巢。     

Costs of reproduction in male lizards, Sceloporus virgatus

Models of life history evolution typically assume a balance between the benefits of current reproductive activity and the costs to future reproductive success or survivorship, but empirical studies often find positive correlations between such components of fitness in undisturbed animal populations. I examined possible survivorship costs of reproduction in free-ranging male lizards, Sceloporus virgatus , and found that males with low levels of mating success were less likely to survive to the following breeding season. I also investigated two possible indicators of reproductive effort, increase in ectoparasite load and decrease in body weight during the breeding season. Levels of parasitism with trombiculid mites at the end of the breeding season were not associated with any measure of fitness or body condition (mating success, survivorship to the following year, relative weight loss). Yearling males (which have low levels of mating success) usually gained weight during the breeding season, while older males generally lost weight during this period. This suggests that young males may have postponed reproduction in favor of body growth and that seasonal weight loss of older males might reflect reproductive effort. Within the group of older males, individuals with the highest levels of mating success did not have high levels of either weight loss or mortality. Mate guarding behavior, an alternative to the aggressive territorial behavior typical of many lizard species, may allow certain males to obtain mates without expending large amounts of energy or exposing themselves to great mortality risks.     

Seasonal clutch size decline and individual variation in the timing of breeding are related to female body condition in a non-migratory species, the Upland Goose Chloephaga picta leucoptera

Several studies have shown that waterfowl females in good condition lay larger clutches and start egg-laying earlier in a breeding season. However, most of these studies lack corresponding data on individual condition and timing of breeding in different years. We analysed data on clutch size, egg size, hatching date and female body condition of a non-migratory species, the Upland Goose Chloephaga picta leucoptera, recorded in three seasons on New Island, Falkand Islands. We found a strong seasonal decline in both egg and clutch size. Egg size increased with clutch size. The mean egg volume and total clutch volume increased with female body condition and hatching date was earlier for females in higher body condition. Chicks hatched from nests with higher mean egg volume had higher early body condition. We also compared individual females between different study years and found that individual body condition showed a significant repeatability between years, whereas hatching date did not. While female aging did not systematically affect hatching date, females produced clutches that hatched earlier in years of higher body condition. We could thus show for the first time that the frequently reported negative relationship between female condition and hatching date also holds true on an individual basis. To our knowledge, this is the first study on seasonal clutch size decline in sheldgeese and one of a few in waterfowl comparing condition and timing of breeding within individuals between different years.     

Sexual size dimorphism in a landlocked Pacific salmon in relation to breeding habitat features

Many animal species exhibit size dimorphism between sexes. Sexual selection, whereby male–male competition favors larger body sizes, has been considered a likely cause of sexual size dimorphism. Habitat features in breeding areas could affect the outcome of male–male competition, yet few attempts have been made to relate breeding habitat features with interpopulation variation in sexual size dimorphism. In this study, we examined interpopulation variation in sexual size dimorphism by studying the landlocked amago salmon (Oncorhynchus masou ishikawae) at a microgeographic scale. We found that female body size was independent of stream size but that male body size decreased with smaller stream sizes. A likely explanation is that the relationship between reproductive success and the size of males is influenced by the availability of refuges that are only available to small-bodied males. Sexual differences in body size increased with decreasing stream sizes, supporting the hypothesis that the reproductive success of larger males is reduced in smaller streams. In contrast, the maturation-length threshold increased with stream size for both sexes. The stream-size-based interpopulation variation in sexual size dimorphism and size at maturity in landlocked amago salmon may therefore have arisen through a combination of sexual and natural selection.     

BODY SIZE,SPERM COMPETITION,AND DETERMINANTS OF REPRODUCTIVE SUCCESS IN MALE SAVANNA BABOONS

One component of sexual selection is sperm competition. It has been reasoned that the intensity of sperm competition may be reflected in the relative testicular sizes of animals. Among males residing in multimale breeding systems, testicular size is relatively larger than among males residing in unimale mating systems. Information on whether differences in testicular size within a species can account for differences in male reproductive success is unavailable for natural populations of primates. A population of six troops of savanna baboons in Kenya was surveyed for morphometric analysis, and one of these troops was the subject of extensive behavioral observations afterwards. Testicular weights could not be obtained, but measurements of linear dimensions were transformed into volumetric estimates. Male weight accounted for 30% of the variance in testicular volume. Neither body size nor testicular volume was associated with differences in male reproductive activity. The outcome of fights over access to females could not be related to male body size, and ejaculatory patterns of males were independent of testicle size. Both sperm competition and aggressive competition intensified during the four-day optimum conception period, but fights over access to consort females were infrequent. Among savanna baboons, the probability of an ejaculation resulting in a conception is fairly low, which may account for the infrequency of injurious fights. Although testicle size influences sperm production, it does not influence either the timing of mating or the fertilizing capacity of spermatozoa, and both of these factors probably account for a substantial fraction of the variance in male baboon paternity. Sperm competition is an adjunct to agonistic competition as a mechanism affecting male baboon reproductive success. It is concluded that male reproductive success in baboons is affected more by social factors than by morphological traits associated with size.     

Mate competition and resource competition are inter‐related in sexual selection

Sexual selection can be affected by the competition for limited breeding resources and/or the competition for limited mates. Although there is ample evidence for each type of competition by itself, little is known about their relative importance and interaction. To address these questions, we established 48 experimental breeding populations of the two‐spotted goby (Gobiusculus flavescens), a substrate‐breeding fish with paternal care. In three experimental treatments, males were limited in the access to either nest sites or mates or were provided with both nests and mates in excess. We quantified male competition behaviour (agonistic and courtship), the opportunity for selection and selection on male body size. Limited access to nests and mates produced similar opportunities for selection, but only limited access to mates increased male competitive behaviours and caused positive selection on male body size. Selection on body size in the mate‐limited treatment was due both to larger males being more likely to take up nests and to larger males being more likely to mate once they had a nest. These findings demonstrate that resource and mate limitation can differ in their effects on sexual selection. The results also reveal that resource and mating competition can be highly inter‐related and not always separated in time, implying that methods to disentangle the two processes must be chosen with care. Future research should consider experimental and analytical approaches similar to those of the present study in attempts to elucidate the interaction of resource and mating competition in animals.     

Male–male competition for large nests as a determinant of male mating success in a Japanese stream goby, Rhinogobius sp. DA

Males of the stream goby Rhinogobius sp. DA (dark type) court females in deep pools and care for the eggs under stones in shallow riffles. We studied male–male competition for access to females and nest sites to understand how male size influences the mating success of this species. In field observations, larger males won in fighting with other males. However, large males did not tend to monopolize courtship opportunities, and the frequency of successful courtships, after which males led the females to the nests, was not related to male body size. The fact that courted females always escaped from the fighting sites once males began fighting likely explains why male size was not positively related to courtship success. Large males occupied large nest stones, and the number of eggs received in the nest was correlated positively with nest size. In aquarium experiments with two tiles of different sizes provided as nesting materials, males always chose the larger nest and, when two males were introduced simultaneously, the larger one occupied the larger nest. These results suggested that male mating success of this goby is determined by male–male competition for large nests rather than for access to females. Received: June 9, 2000 / Revised: September 2, 2000 / Accepted: October 4, 2000     

Relationship between clutch size,egg volume and hatching success in a Yellow-legged Gull Larus michahellis colony in south-eastern Tunisia

This study aimed to test the hypothesis that clutch size covaries with egg volume and hatching success in the Yellow-legged Gull Larus michahellis. We determined clutch size and egg volume in a sample of 131 nests, and we used the data to check whether egg volume varied among nests according to clutch size, while taking into account the effects of egg laying order. We also estimated hatching success rate and investigated the relationship between hatching success and clutch size. Egg volume varied among clutches according to clutch size, with eggs being larger in three-egg clutches than in two-egg clutches. Moreover, three-egg clutches showed higher daily survival rates, and hence hatching success, than two-egg clutches. Overall, our results suggest that in the Yellow-legged Gull clutch size covaries with egg volume and hatching success, which could possibly reflect an age effect through different mechanisms. Indeed, older females could be hypothesised to exhibit greater breeding performance than younger females because of their higher experience in tapping energy resources for egg formation and defending nests from dangers. Moreover, due to their age, older females are likely to have lower residual reproductive potential and should invest more heavily in current breeding attempts.