Comparison of larval duration and pre- and post-settlement growth in two species of damselfish,Chromis atripectoralis andPomacentrus coelestis (Pisces: Pomacentridae), from the Great Barrier Reef

Light traps were used to capture larval fishes, immediately before settlement, at two localities 500 km apart on the Great Barrier Reef (GBR) in December, 1987. Samples from Lizard Island, in the northern GBR, and Davies Reef, in the central GBR, were dominated by two species of damselfish:Chromis atripectralis andPomacentrus coelestis. Analysis of otoliths revealed significant differences in both size and age at settlement between the two localities forP. coelestis, but not forC. atripectoralis. Growth rates determined for pre- and post-settlementP. coelestis suggested a sigmoidal growth trajectory through the larval life, with growth slowing as fishes approached the time of settlement. Post-settlement growth rates were faster than growth prior to settlement in both species. Growth in both species was, however, similar between localities. The relationship between fish size and otolith size was complex, varying both between pre- and post-settlement fishes, and among localities. This emphasizes the need to validate the relationship between fish size and otolith size before otoliths may be used to back-calculate individual growth trajectories.Contribution No. 500 from the Australian Institute of Marine Science     

Age-specific filial cannibalism in a paternal mouthbrooding fish

Entire-brood cannibalism by mouthbrooding males of the cardinal fish Apogon doederleini was investigated in temperate waters of southern Japan during two breeding seasons. The rate of cannibalism was 17–18% in each season and did not differ among age-groups. However, the seasonal pattern of cannibalism differed markedly among age-groups: young (1- and 2-year-old) males frequently cannibalized early broods, especially the first brood, of the season, whereas cannibalism by middle-aged (3- and 4-year-old) and old (5- and 6-year-old) males mainly occurred late in the breeding season. We explain this difference in terms of trade-offs between current and future reproduction. Young males, whose future reproductive success is enhanced by the growth increment, may allocate more time and energy to growth by cannibalizing early broods. In contrast, for older males who have had more breeding cycles and grow little, cannibalism could be a way to reverse the deterioration in their somatic condition that occurs as the breeding season progresses. It is also likely that the current reproductive loss entailed by the cannibalism is effectively compensated by quick re-mating with another female. Received: 24 February 1997 / Accepted after revision: 20 July 1997     

Seasonal decline in reproductive performance varies with colony size in the fairy martin, <Emphasis Type="Italic">Petrochelidon ariel</Emphasis>

Reproductive success within populations often varies with the timing of breeding, typically declining over the season. This variation is usually attributed to seasonal changes in resource availability and/or differences in the quality or experience of breeders. In colonial species, the timing of breeding may be of particular importance because the costs and benefits of colonial breeding are likely to vary over the season and also with colony size. In this study, we examine the relationship between timing of breeding and reproductive performance (clutch size and nest success) both within and between variable sized colonies (n = 18) of fairy martins, Petrochelidon ariel. In four of these colonies, we also experimentally delayed laying in selected nests to disentangle the effects of laying date and individual quality/experience on reproductive success. Within colonies, later laying birds produced smaller clutches, but only in larger colonies. The general seasonal decline in nest success was also more pronounced in larger colonies. Late laying birds were generally smaller than earlier laying birds, but morphological differences were also related to colony size, suggesting optimal colony size also varies with phenotype. Experimentally delayed clutches were larger than concurrently produced non-delayed clutches, but only in larger colonies. Similarly, delayed clutches were more likely to produce fledglings, particularly later in the season and in larger colonies. We suggest that the reduced performance of late breeding pairs in larger colonies resulted primarily from inexperienced/low quality birds preferring to settle in larger colonies, possibly exacerbated by an increase in the costs of coloniality (e.g., resource depletion and ectoparasite infestations) with date and colony size. These findings highlight the importance of phenotype-related differences in settlement decisions and reproductive performance to an improved understanding of colonial breeding and variation in colony size.     

Early nutritional conditions, growth trajectories and mate choice: does compensatory growth lead to a reduction in adult sexual attractiveness?

It is well documented that poor early nutrition can have profound negative effects on adult life-history traits. However, it has also been demonstrated that organisms can undergo compensatory resource allocation strategies (such as an accelerated growth rate) if food availability improves, so as to mitigate the effects of the poor early conditions. Previous research has indicated that elevated growth rates can incur costs in the longer term, such as an increased rate of senescence and shorter lifespan. We tested whether a phase of compensatory growth after a period of reduced food availability earlier in life affected the sexual attractiveness of adult male green swordtails Xiphophorus helleri, a species in which it has previously been documented that females prefer larger-bodied and longer-tailed males. The experiment compared the attractiveness of size-matched brothers that had experienced contrasting growth trajectories as juveniles; the experiments were initially conducted in the middle of a male’s sexually reproductive life and were then repeated towards the very end of life. At both ages, males that had undergone compensatory growth were equally as attractive as their brothers that had grown normally. These results suggest that the growth compensation benefits males through an increase in their attractiveness over that which they would have had if they had remained on their original growth trajectory. The lack of change in relative attractiveness with age indicates that the compensation does not cause greater deterioration in secondary sexual characters at older ages than in continuously well-fed males.     

The amphipod Corophium multisetosum (Corophiidae) in Ria de Aveiro (NW Portugal). I. Life history and aspects of reproductive biology

The population of Corophium multisetosum Stock, 1952 in Areão displayed a semiannual, iteroparous life history. Mean longevity was ～6?mo, with the estimated life span longer for overwintering individuals born in autumn than for individuals born in spring. Length-frequency data indicated that the length increment per moult is probably higher in males than females; however females moulted more frequently and achieved a larger body size. Preliminary growth rates were 100?μm?d^?1 for juveniles and 19 to 29?μm?d^?1 for mature females, with the lower values occurring during the winter. It was estimated that under favourable conditions females may attain reproductive size and mature within 1?mo. Although incubating females were present all year round, recruitment occurred in spring, almost ceased during the summer, peaked in autumn, and decreased again during the winter. Extreme temperatures and very low salinities during winter and summer may have deterred breeding, while moderate temperatures (15 to 20?°C) and salinities > 1?psu in spring and autumn were apparently favourable for reproduction. The unfavourable summer conditions constrained breeding and synchronised the timing of reproduction. In late-autumn and during the winter, as temperature decreased and brooding time increased, synchrony was progressively lost. Brood size varied as a function of embryonic developmental stage, size of incubating females, and season. The life-history pattern and reproductive features of C. multisetosum in Areão are closely related to temperature and salinity; other environmental conditions such as oxygen content of the water and food availability may also be relevant.     

Variation in plasma levels of insulin-like growth factor-I (IGF-I) in lingcod: relationships among season,size, and gonadal steroids

The physiological response of a fish to its environment is mediated through the endocrine axis controlling growth. Therefore, growth-regulating hormone levels can serve as ecologically relevant indicators of fish growth rate. We quantified variation in plasma insulin-like growth factor-I (IGF-I) to evaluate its potential as an indicator of growth in lingcod, an economically and ecologically important bottomfish species in the northeast Pacific. An information-theoretic model selection approach was used to test the hypothesis that variation in lingcod IGF-I is related to season, body size, and gonadal steroid concentration. Season and a length × season interaction were the most important predictors of plasma IGF-I among the variables we evaluated, suggesting that season and body size should be explicitly accounted for when interpreting endocrine patterns in wild fish populations. This is among the few studies that have measured and interpreted patterns of IGF-I in wild fish and the first to describe seasonal endocrine profiles in lingcod.     

Longitudinal monitoring of neutral and adaptive genomic diversity in a reintroduction

Restoration programs in the form of ex-situ breeding combined with reintroductions are becoming critical to counteract demographic declines and species losses. Such programs are increasingly using genetic management to improve conservation outcomes. However, the lack of long-term monitoring of genetic indicators following reintroduction prevents assessments of the trajectory and persistence of reintroduced populations. We carried out an extensive monitoring program in the wild for a threatened small-bodied fish (southern pygmy perch, Nannoperca australis) to assess the long-term genomic effects of its captive breeding and reintroduction. The species was rescued prior to its extirpation from the terminal lakes of Australia's Murray-Darling Basin, and then used for genetically informed captive breeding and reintroductions. Subsequent annual or biannual monitoring of abundance, fitness, and occupancy over a period of 11 years, combined with postreintroduction genetic sampling, revealed survival and recruitment of reintroduced fish. Genomic analyses based on data from the original wild rescued, captive born, and reintroduced cohorts revealed low inbreeding and strong maintenance of neutral and candidate adaptive genomic diversity across multiple generations. An increasing trend in the effective population size of the reintroduced population was consistent with field monitoring data in demonstrating successful re-establishment of the species. This provides a rare empirical example that the adaptive potential of a locally extinct population can be maintained during genetically informed ex-situ conservation breeding and reintroduction into the wild. Strategies to improve biodiversity restoration via ex-situ conservation should include genetic-based captive breeding and longitudinal monitoring of standing genomic variation in reintroduced populations.     

Allocation in reproduction is not tailored to the probable number of matings in common toad (<Emphasis Type="Italic">Bufo bufo</Emphasis>) males

The theory of life history evolution assumes trade-offs between competing fitness traits such as reproduction, somatic growth, and maintenance. One prediction of this theory is that if large individuals have a higher reproductive success, small/young individuals should invest less in reproduction and allocate more resources in growth than large/old individuals. We tested this prediction using the common toad (Bufo bufo), a species where mating success of males is positively related to their body size. We measured testes mass, soma mass, and sperm stock size in males of varying sizes that were either (1) re-hibernated at the start of the breeding season, (2) kept without females throughout the breeding season, or (3) repeatedly provided with gravid females. In the latter group, we also estimated fertilization success and readiness to re-mate. Contrary to our predictions, the relationship between testes mass and soma mass was isometric, sperm stock size relative to testes mass was unrelated to male size, fertilization success was not higher in matings with larger males, and smaller males were not less likely to engage in repeated matings than larger males. These results consistently suggest that smaller males did not invest less in reproduction to be able to allocate more in growth than larger males. Causes for this unexpected result may include relatively low year-to-year survival, unpredictable between-year variation in the strength of sexual selection and low return rates of lowered reproductive investment.     

Carotenoid-based status signalling in red-shouldered widowbirds (<Emphasis Type="Italic">Euplectes axillaris</Emphasis>): epaulet size and redness affect captive and territorial competition

In breeding plumage, the African male red-shouldered widowbirds (Euplectes axillaris) are black except for red carotenoid-based epaulets ('shoulder patches'), similar to the well-studied American red-winged blackbirds (Agelaius phoeniceus). To experimentally test the signal function of the red epaulets in male red-shouldered widowbirds, we manipulated epaulet colour and size (within natural variation), and observed the effects in two competitive contexts. First, in captivity, unfamiliar males with epaulets of different size or redness were staged in dyadic contests over food. Only epaulet manipulations significantly affected contest outcome; red epaulet males out-competed orange and control males, which in turn dominated blackened (no epaulet) males. Epaulet size manipulations had particularly strong effects; enlarged epaulet males vigorously defeated smaller epaulet males within colour treatments, whereas the independent effect of 'redness' (i.e., within size treatments) was much weaker. Second, in a field experiment, epaulets were manipulated prior to territory establishment. Males with enlarged epaulets acquired territories, whereas most males with blackened or reduced signals failed to obtain territories. Furthermore, among the territorial males, birds with enlarged epaulets defended larger territories and spent less time in boundary signalling, yet they received fewer intrusions from conspecifics. In contrast, epaulet manipulations had no effect on either female visitation rates or subsequent female settlement. These results concur with recent studies and suggest that the carotenoid-based epaulets of red-shouldered widowbirds are status signals, sexually selected through male contest competition for territories.     

Sex-specific patterns of lifetime reproductive success in single and repeat breeding steelhead trout (Oncorhynchus mykiss)

All other things being equal, the lifetime reproductive success (LRS) of iteroparous and semelparous individuals should scale with the number of breeding seasons. Deviations from this relationship may occur for many reasons, including age- or size-related fecundity or life history trade-offs, which may differ between sexes. We used 19 brood years of DNA parentage analysis in a small (N = 4–143 year⁻¹) wild, unexploited population of steelhead trout (Oncorhynchus mykiss) to compare the LRS of individuals that spawned only once [“one time spawners” (OTS), N = 355 male, 371 female] to those spawning twice [“repeat spawners” (RPS), N = 13 male, 49 female]. Female RPS had nearly twice the LRS of female OTS (1.17 offspring per female vs 0.91 offspring per female), whereas male RPS had nearly three times the LRS of male OTS (1.54 offspring per male vs 0.57 offspring per male). Female RPS produced slightly more adult offspring during their second breeding season than their first (0.78 vs 0.82 offspring per female); however, male RPS produced all of their adult offspring in their second breeding season (0 vs 1.54 offspring per male). The additional growth in body size of males between breeding seasons may give them an advantage in their second breeding season, but the lack of offspring produced in their first season suggests a trade-off between survival and future reproduction that was not expressed in females.     

Helpers in colonial cooperatively breeding sociable weavers <Emphasis Type="Italic">Philetairus socius</Emphasis> contribute to buffer the effects of adverse breeding conditions

Some studies on the effects of helpers in cooperatively breeding vertebrates show a positive effect of helper presence on reproductive output whereas others find no effect. One possibility for this discrepancy is that helpers may have a positive effect when breeding conditions are adverse, while their effect might go unnoticed under good conditions. We investigate this hypothesis on sociable weavers Philetairus socius, a colonial cooperatively breeding passerine that inhabits a semi-arid region where breeding conditions vary markedly. We used multivariate mixed models to analyse the effect of helpers on reproduction under contrasting environmental and social conditions while controlling for parental and colony identity. We found that reproductive success in sociable weavers was primarily influenced by nest predation and rainfall. In addition, colony size was negatively associated with hatching and fledging success and number of young fledged per season. Helpers had a less prominent but significant influence on feeding rates and reproductive outcome. In agreement with expectations, the presence of helpers counteracted some of the negative effects of breeding in periods of low rainfall or in large colonies and was also associated with an increased number of young fledged per season. Our results illustrate that the effect of helpers might be detectable mostly under unfavourable conditions, but can contribute to improve reproductive performance in those situations.     

Patterns of shelter usage and social aggregation by the vocal Lusitanian toadfish

In some marine fishes, males migrate from deeper to littoral water to breed, where they defend nests and provide parental care. In this study, we describe seasonal patterns of shelter occupation in estuarine shallow waters and assess social aggregation in the Lusitanian toadfish (Batrachoididae), a vocal species with male parental care. Occupation of intertidal shelters was restricted to the breeding season but adult fish remained in subtidal regions during the non-reproductive season and they produced sounds throughout the year. Intertidal shelters were aggregated with up to five shelters found per 2-m segment of an 80-m transect. This aggregation probably resulted from social attraction since many segments were found unoccupied. Moreover, shelters contained on average two fish (maximum of nine). Based on size, shelters with multiple fish seemed to contain females, or possibly sneakers, but also more than one type I male inside, indicating a high social tolerance for conspecifics.     

Is reduced clutch size a cost of parental care in Eastern Phoebes (Sayornis phoebe)?

What is the cost of parental care in birds? Previous studies using observational and experimental techniques on nest building and clutch sizes in a small migrant flycatcher, the Eastern Phoebe (Sayornis phoebe), led to contradictory results that did not show a consistent cost of current reproductive effort on residual reproductive output. The data presented here indicate that different elements of parental behaviors are indeed costly because they reduce various aspects of phoebes' subsequent reproductive performance. Experimental removal of old nesting structures at previously used breeding sites reduced but did not eliminate the chance of phoebes' settlement in the subsequent year. Comparing sites at which phoebes did and did not build new nests showed that nest builders completed their first clutches later, had lower probabilities of second breeding attempts, and more often lost their nesting attempt due to fallen nest structures than nest reusers. There was, however, no significant effect of nest building on the clutch sizes and rates of cowbird parasitism of first nesting attempts. Overall, sites with newly built nests had lower seasonal reproductive effort than sites with reused nests. I also examined phoebes' relative residual reproductive output in a separate breeding season when nest building was not experimentally manipulated. When controlled for confounding variables this analysis indicated that in those phoebes that did breed for a second time, the relative decrease of the sizes of first to presumed second clutches was greater at sites where first breeding attempts consisted of more total nestlings. These data are consistent with the hypothesis that parental care is costly in Eastern Phoebes and support predictions of trade-offs between the nest building, brood care, and residual egg-investment components of reproduction.     

Trace metals in mussels (Mytilus edulis) from southwest Iceland

The temperate population of the tropical anemonefish Amphiprion clarkii has a seasonality of reproduction, larval settlement and growth. There was a considerable difference in size (15–47 mm) among 0-year olds in December, after the season of larval settlement and growth. This difference could be attributed mainly to a difference in the duration of the first growing season spent after settlement and a social inhibition of growth of the subordinate late settlers by frequent attacks by the dominant early settlers. The growth equation which was used to describe the growth of tagged individuals estimated that an anemonefish would get an opportunity to spawn within four of five years after settlement. Breeding adult pairs occupied almost all sea anemones essential to breeding within a 50×50 m study area, and all 17 new adults that matured from non-breeding juveniles to breeding adults between June 1983 and August 1985 were found in the territories from which either or both of the mated adults had disappeared. The new adults were large juveniles who had resided near those vacant territories before. These results suggest that the onset of breeding by an individual is not only determined by his age, but also by his ranking in the dominance hierarchy. Therefore, the difference in size among 0-year olds in December might give rise to the difference in age of the earliest breeding and might be a factor in making a female apply more reproductive effort at the beginning of the season.     

Risk of predation and the feeding behavior of juvenile coho salmon (Oncorhynchus kisutch)

Summary During their first 1–2 years of life, juvenile coho salmon (Oncorhynchus kisutch) are stream-dwelling, and feed upon drifting invertebrates. They move upstream from a holding position to intercept individual prey items; the distance moved (attack distance) is an increasing, but decelerating, function of prey size. Since the fish are presumably more visible to predators during such feeding excursions, prey size and risk are associated variables.The effect on attack distance of the presentation of a model predator (a photograph of a rainbow trout) was examined in the laboratory. Attack distances are shortened following presentation of a predator; this is particularly true when the prey are large (Fig. 1). The extent of the reduction of attack distance is directly related to predator presentation frequency, although there appears to be a minimum level to which it will decline (Fig. 2). Hungry fish and fish in the presence of a competitor (simulated by a mirror) are less responsive to the predator, suggesting a trade-off of energetic requirements and risk (Fig. 3 and Table 3). The effect of predation risk should be to reduce the relative proportion of large prey in a juvenile coho's diet, and its net rate of energy intake.     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

An experimental study of male adornment in the scarlet-tufted malachite sunbird: II. The role of the elongated tail in mate choice and experimental evidence for a handicap

Summary Scarlet-tufted malachite sunbirds (Neetarinia johnstoni) are endemic to the high altitude zones of East African mountains. On Mount Kenya the males are resident and territorial all year. The females arrive over an extended period at the beginning of the breeding season, after spending the non-breeding season in nomadic flocks at slightly lower elevations. Males are bright iridescent green with elongated central tail feathers, which are displayed during courtship. We examined the role of the tail in mate choice using natural variation and investigated the influence of tail length on male time budgets by experimental manipulation. Territorial males that paired had longer tails than territorial males which remained unpaired. Males with naturally long tails started breeding earlier, resulting in fledglings being produced earlier in the season when the weather was more clement. Birds which bred together in one year were paired together in the next year if they both survived. Thus the male's tail may only be involved as a cue in the formation of new pairs. Males which were already paired were subjected to one of three experimental treatments — having their tail elongated, shortened, or manipulated but kept at the same length. Males with experimentally shortened tails spent more time in flight and hawked for flying insects with a higher efficiency than control males. Both control and elongated tail males reduced the amount of time in flight and had a lower hawking efficiency after manipulation. These results suggest that the long tail of male scarlet-tufted malachite sunbirds is a handicap. The factors influencing the reproductive success of a pair are discussed. Offprint requests to: M.R. Evans     

Age,size, dominance and reproduction among male kudus: mating enhancement by attrition of rivals

Summary Size dimorphism with males larger than females has been related to the benefits for males of enhanced dominance and hence reproductive success. However, mating gains must outweigh the fitness costs of deferred reproduction and the mortality associated with further growth. The relationships between male age, size and reproduction were assessed for greater kudus (Tragelaphus strepsiceros) in the Kruger National Park in South Africa. Individually identifiable animals were monitored over 10 years, with detailed observations made during six breeding seasons. In the non-breeding season males formed loose all-male groups. Horn grappling and low intensity agonistic interactions fostered dominance rankings. Dominance was age-graded, until males reached full weight at 6 years of age. Males aged 6 and 7 years monopolized courtship and mating, but 5-year-old males secured about 10% of mating opportunities. Few males survived beyond 7 years. Male mortality rate rose steeply with age, so that the functional sex ratio of fertile females per mature male was about 14:1. During the breeding season many female groups remained unattended by a mature male. Reproductive sorting among males occurred largely through variation in survival to full size and maturity. Increased size enhances fighting success and hence dominance. Further growth ceases when the functional sex ratio exceeds the number of mating opportunities that males can effectively achieve during a breeding season. Predation amplifies the mortality cost of continued growth. In the absence of large predators, male-male interactions may be atypically exaggerated.     

Seasonal variation in territorial response,and other aspects of the ecology of the Australian temperate pomacentrid fish Parma microlepis

The Australian temperate pomacentrid fish Parma microlepis Gunther maintains permanent, interspecifically defended territories. Breeding occurs in late spring (October to December), and juveniles after a pre-settling period of 4 to 6 weeks take up territories in the adult habitat. P. microlepis feed in their territories on benthic algae and invertebrates. Density of fish is significantly correlated with the availability of suitable shelter. There is a variation in the territorial response directed at different species, and a significant correlation exists between intensity of response to a species and its similarity to P. microlepis in use of living space. No significant correlation exists between intensity of response and dietary similarity. P. microlepis defends larger areas in the breeding than in the non-breeding season against those species which represent a possible threat to its breeding success. It shows a reduced tendency to attack juvenile conspecifics at the time of year when they would be settling from the plankton. Responses to species not likely to interfere with breeding success do not vary through the year. P. microlepis is thus seen to have complex territorial responses, specific to the species of intruder and time of year. In this it is quite similar to other territorial pomacentrids (chiefly tropical) for which information exists.