Asynchronous hatching in the laughing gull: Cutting losses and reducing rivalry

The adaptive significance of hatching asynchrony was investigated in the laughing gull (Larus atricilla). Staggered hatching of the brood is frequently correlated with reduced survival in the later-hatching chicks, and the phenomenon has therefore been the subject of speculation about adaptive function. The relative timing of hatching within the brood was experimentally manipulated in order to compare the effects of synchrony versus asynchrony on parental reproductive success. The average number of young fledging per nest was significantly higher in the asynchronous group. Staggered hatching of the brood is hypothesized to be a parental strategy offering the option of brood reduction under conditions of food shortage. The possibility is discussed that asynchronous hatching benefits parents independently by reducing sibling rivalry over parental investment and minimizing wasteful competition.     

Does food supplementation really enhance productivity of breeding birds?

Food availability influences multiple stages of the breeding cycle of birds, and supplementary feeding has helped in its understanding. Most supplementation studies have reported advancements of laying, whilst others, albeit less numerous, have also demonstrated fitness benefits such as larger clutches, shorter incubation periods, and greater hatching success. Relatively few studies, however, have investigated the effects of supplementary feeding for protracted periods across multiple stages of the breeding cycle. These effects are important to understand since long-term food supplementation of birds is recommended in urban habitats and is used as a tool to increase reproductive output in endangered species. Here, we compare the breeding phenology and productivity of blue tits Cyanistes caeruleus and great tits Parus major breeding in food-supplemented and non-supplemented blocks in a broadleaf woodland in central England over three seasons (2006–2008). Supplementation was provided continuously from several weeks pre-laying until hatching, and had multiple significant effects. Most notably, supplementation reduced brood size significantly in both species, by half a chick or more at hatching (after controlling for year and hatching date). Reduced brood sizes in supplemented pairs were driven by significantly smaller clutches in both species and, in blue tits, significantly lower hatching success. These are novel and concerning findings of food supplementation. As expected, supplementary feeding advanced laying and shortened incubation periods significantly in both species. We discuss the striking parallels between our findings and patterns in blue and great tit reproduction in urban habitats, and conclude that supplementary feeding may not always enhance the breeding productivity of birds.     

Effects of Shiny Cowbird Molothrus bonariensis parasitism on different components of House Wren Troglodytes aedon reproductive success

Avian brood parasites, including cuckoos and cowbirds, have multiple negative effects on their hosts. We analysed the effects of Shiny Cowbird Molothrus bonariensis parasitism on different components (e.g. egg losses, hatching success, chick survival and nest abandonment) of House Wren Troglodytes aedon reproductive success. We also conducted an experiment to discriminate between two mechanisms that may reduce hatching success in parasitized clutches: lower efficiency of incubation due to the increase in clutch volume and disruption of host incubation by the early hatching of Cowbirds. Egg puncturing by Shiny Cowbirds reduced host clutch size at hatching by 10–20%, and parasitized nests had a decrease in hatching success of 40–80%. Egg losses and hatching failures were positively associated with the intensity of parasitism. Brood reduction was greater in parasitized nests, but the growth rate of the chicks that fledged was similar to that in unparasitized nests. The combined effects of egg losses, hatching failures and brood reduction decreased the number of fledged chicks by 80%. In addition, egg puncturing increased the likelihood of nest abandonment by Wrens. Experimental data showed that hatching failures occurred when there was a combination of: (1) an increase in the volume of the clutch by the addition of the Cowbird egg without removal of host eggs, and (2) the addition of the Cowbird egg before the onset of incubation. This was relatively common in House Wren nests, as Cowbirds generally parasitize before the onset of incubation. Our results indicate that Shiny Cowbird parasitism imposes a major impact on House Wrens, as it affects all components of the Wren's reproductive success.     

Overproduction in the Lesser Black‐backed Gull – can marginal chicks overcome the initial handicap of hatching asynchrony?

In response to unpredictability of both food availability and core offspring failure, parents of many avian species initially produce more offspring than they commonly rear (overproduction). When parental investment is insufficient to raise the whole brood the handicap of hatching last means ‘marginal’ chicks are less likely to survive if brood reduction occurs. Conversely, if marginal offspring are required as replacements for failed ‘core’ chicks, or parental investment is sufficient to rear the whole brood, the handicap imposed on marginal chicks must be reversible if overproduction is to be a viable strategy. I investigated the ability of marginal offspring to overcome the handicap imposed by hatching asynchrony using a combination of a field experiment, designed to manipulate both the amount of total competition and the relative competitive ability of chicks within a brood, and data on the growth and survival of unmanipulated, three‐chick broods from three consecutive years. The results indicate that, even when resources are abundant, marginal offspring do not begin to overcome the competitive handicap imposed by hatching asynchrony until the period of growth when energetic requirements reach their peak, and subsequent survival to fledging is almost assured. This is apparently a consequence of parents controlling allocation of early parental investment, so that any brood reduction ‘decisions’ can be left as late as possible. Marginal chicks initially channel resources into maintaining mass, relative to skeletal size, as a buffer against starvation. However this also means competitiveness is reduced, so if conditions are poor marginal chicks are rapidly out‐competed, lose condition and die. Conversely, when food availability is good marginal offspring devote more resources to skeletal growth and quickly close the gap on their core siblings, meaning the handicap is reversible. The benefits of overproduction and hatching asynchrony as reproductive strategies to maximise success in Lesser Black‐backed Gulls are discussed in relation to the reproductive alternatives.     

The social system of the Pale Chanting Goshawk Melierax canorus; monogamy v polyandry and delayed dispersal

The social organization and reproductive strategy of the Pale Chanting Goshawk Melierax canorus were investigated in four southern African study areas. Territories supporting adults additional to the breeding pair were observed in two study areas. Within one of these areas, these additional adults were either non-breeders staying in their natal territory but actively excluded from the nesting area during the breeding season or a male cobreeder participating fully in reproductive activities with the breeding pair. Polyandrous trios were, however, observed in only one vegetation type in this area, Broken Veld. The annual number of offspring fledged per group did not differ significantly between vegetation types nor between polyandrous trios and monogamous pairs within Broken Veld. Pale Chanting Goshawks lay predominantly two-egg clutches. In 2 out of 5 years, breeding groups succeeded in laying, hatching and even fledging a second brood of young after successfully fledging their first brood. Double brooding occurred more frequently in Broken Veld and most frequently in polyandrous trios in this vegetation type. To accommodate their relatively long breeding cycle (>115 days) in this temperate study area with its limiting summer breeding period, double-brooding Pale Chanting Goshawks laid the first clutch in midwinter and the second, on average, 24 days after the offspring from the first brood left the nest.     

Hatching asynchrony that maintains egg viability also reduces brood reduction in a subtropical bird

In birds, hatching failure is pervasive and incurs an energetic and reproductive cost to breeding individuals. The egg viability hypothesis posits that exposure to warm temperatures prior to incubation decreases viability of early laid eggs and predicts that females in warm environments minimize hatching failure by beginning incubation earlier in the laying period, laying smaller clutches, or both. However, beginning incubation prior to clutch completion may incur a cost by increasing hatching asynchrony and possibly brood reduction. We examined whether Florida scrub jays (Aphelocoma coerulescens) began incubation earlier relative to clutch completion when laying larger clutches or when ambient temperatures increased, and whether variation in incubation onset influenced subsequent patterns of hatching asynchrony and brood reduction. We compared these patterns between a suburban and wildland site because site-specific differences in hatching failure match a priori predictions of the egg viability hypothesis. Females at both sites began incubation earlier relative to clutch completion when laying larger clutches and as ambient temperatures increased. Incubation onset was correlated with patterns of hatching asynchrony at both sites; however, brood reduction increased only in the suburbs, where nestling food is limiting, and only during the late nestling period. Hatching asynchrony may be an unintended consequence of beginning incubation early to minimize hatching failure of early laid eggs. Food limitation in the suburbs appears to result in increased brood reduction in large clutches that hatch asynchronously. Therefore, site-specific rates of brood reduction may be a consequence of asynchronous hatching patterns that result from parental effort to minimize hatching failure in first-laid eggs. This illustrates how anthropogenic change, such as urbanization, can lead to loss of fitness when animals use behavioral strategies intended to maximize fitness in natural landscapes.     

The annual pattern of reproduction of Talitrus saltator (Crustacea: Amphipoda: Talitridae)

The reproductive cycle and breeding biology of Talitrus saltator (Montagu), a supralittoral talitrid amphipod, were investigated over a 15 month period on a sandy beach in the Isle of Man. The species is shown to have an annual, univoltine reproductive cycle with a period of high reproductive activity between May and late August. Two generations are present throughout most of the year and the life spans of female and male T. saltator were calculated to be approximately 18 months and 21 months respectively. Initial juvenile growth was rapid, juveniles attaining sexual differentiation within three to four months of hatching. Juvenile growth rates averaged 5–5 mm in 100 days decreasing to 1–3 mm in 100 days after sexual differentiation at around 8.0 mm.
The relative brood size of incubating females was shown to be dependent on at least two factors, the length of the female and the stage of embryological development of the brood.
It is suggested that photoperiod is the major exogenous factor controlling gonadal maturation and the duration of reproductive activity in T. saltator.     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.     

Variation in Predispersal Losses in Reproductive Potential in Rowan (<Emphasis Type="Italic">Sorbus aucuparia</Emphasis> L. Rosaceae) in the NW Iberian Peninsula

We identified the principal factors causing losses in reproductive potential (i.e. quantity of potentially dispersable seeds) over the predispersal period of the reproductive cycle of Sorbus aucuparia L. (Rosaceae) in various populations in the NW Iberian Peninsula. Over the period 1999–2002 we examined the magnitude and variability of losses observed in each stage from bud production to seed production (i.e. bud to flower, flower to initiated fruit, initiated to full-size fruit, and full-size to mature fruit). We also investigated spatial and temporal variation in these losses. Reproductive potential showed very significant losses over the predispersal period in all 4 years: in no case did the number of mature fruits available for dispersal exceed 20% of the initial number of flower buds. In all 4 years of study, we detected statistically significant variation among populations in total losses. The principal cause of loss of reproductive potential over the predispersal period in our Sorbus aucuparia populations was non-initiation of fruit. The plant’s interactions with predispersal predators of buds and seeds show high spatial and temporal variability. Also, the proportions of loss due to the different components did not show within-individual consistency over time. It thus seems unlikely that the different factors contributing to predispersal losses in rowan are currently generating significant selective pressures in our study populations.     

Annual and seasonal reproductive trends in the Lesser Spotted Woodpecker Dendrocopos minor

Annual and seasonal variation in reproductive timing and performance were studied in a population of the Lesser Spotted Woodpecker Dendrocopos minor over 10 years in southern Sweden. The median laying date of the first egg varied by up to 17 days between years, being generally larger than the variation of laying dates within years. Neither clutch size, brood size in successful nests, fledging success in successful nests nor mean nestling weight differed significantly between years. There was no trend for mean clutch size to vary between early and late years. In spite of a more than threefold variation in population size, no reproductive variable demonstrated an apparent density-dependence. Within the season, clutch size declined steeply with increasing clutch initiation date, whereas fledging success and nesting success did not, leading to a trend in brood size almost identical to the trend in clutch size. The survival prospects of fledged young declined with increasing clutch initiation date, and it is argued that the clutch size laid is a strategic adjustment to laying date. Out of 124 breeding attempts, 34% did not produce fledged young. In 9% of the breeding attempts, pairs laid no eggs. At least 20% of the breeding attempts failed after egg-laying. The most common cause of breeding failure was loss of the breeding partner followed by nest abandonment (40% of the failures). Only 16–28% of the failures were due to predation on the nest. Most complete failures, and also partial losses from nests, occurred at the early breeding stages. It is argued that the early nestling phase may be a critical stage, which the woodpeckers adjust to coincide with the seasonal food peak, explaining the strikingly late breeding season compared with other non-migrant species.     

Synchronization of laying by great spotted cuckoos and recognition ability of magpies

Brood parasites rely entirely on the parental care of host species to raise the parasitic nestlings until independence. The reproductive success of avian brood parasites depends on finding host nests at a suitable stage (i.e. during egg laying) for parasitism and weakly defensive (i.e. non‐ejector) hosts. Finding appropriate nests for parasitism may, however, vary depending on ecological conditions, including parasite abundance in the area, which also varies from one year to another and therefore may influence coevolutionary relationships between brood parasites and their hosts. In this scenario, we explored: 1) the degree of laying synchronization between great spotted cuckoos Clamator glandarius and magpies Pica pica during two breeding seasons, which varied in the level of selection pressure due to brood parasitism (i.e. parasitism rate); 2) magpie responses to natural parasitism in the pre‐laying period and successfulness of parasitic eggs laid at this stage; and 3) magpie responses to experimental parasitism performed at different breeding stages. We found that, during the year of higher parasitism rate, there was an increase in the percentage of parasitic eggs laid before magpies started laying. However, the synchronization of laying was poor both years regardless of the differences in the parasitism rate. The ejection rate was significantly higher during the pre‐egg‐laying and the post‐hatching stages than during the laying stage, and hatching success of parasitic eggs laid during the pre‐egg‐laying stage was zero. Thus, non‐synchronized parasitic eggs are wasted and therefore poor synchronization should be penalized by natural selection. We discuss four different hypotheses explaining poor synchronization.     

Flexibility of reproductive tactics and their consequences in the brood parasitic fish Pungtungia herzi (Teleostei: Cyprinidae)

Although previous studies have claimed that Pungtungia herzi (Cyprinidae) is an obligate brood parasite, in this study a significant proportion of P. herzi egg masses was found in crevices formed by stones and rocks, which was not guarded by hosts. Egg masses were also found in almost all nests of a goby Odontobutis obscura , suggesting that P. herzi has two alternative reproductive tactics: brood parasitic spawning and non-parasitic crevice spawning, the latter of which may occur when a shortage of reproductive resources ( i.e . host nests) exists. The hatching rates of parasitic egg masses were significantly higher than those of non-parasitic egg masses, and the hatching rate of P. herzi eggs exhibited a strong positive correlation with continuous egg guarding by a reproductive O. obscura male. These results indicate that brood parasitism effectively improves reproductive success in P. herzi . The alternative reproductive tactic of this species suggests that brood parasitism has evolved directly from crevice spawning via the sharing of spawning habitat with host species.     

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi‐precocial Herring Gulls

Intraclutch egg size variation may non‐adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size‐mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c‐eggs may represent a brood‐reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three‐egg clutches over a 3‐yr period (2010–2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date (β = ?0.18 ± SE 0.06, P = 0.002) on c‐egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C‐chicks suffered from increased mortality risk during the first 12 d, identified as the brood‐reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood‐reduction period. Thus, the value of relatively small c‐eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three‐egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.     

Asynchronous hatching in the burying beetle,Nicrophorus quadripunctatus,maxmizes parental fitness

Life history theory predicts that natural selection favours parents who balance investment across offspring to maximize fitness. Theoretical studies have shown that the optimal level of parental investment from the offspring's perspective exceeds that of its parents, and the disparity between the two generates evolutionary conflict for the allocation of parental investment. In various species, the offspring hatch asynchronously. The age hierarchy of the offspring usually establishes competitive asymmetries within the brood and determines the allocation of parental investment among offspring. However, it is not clear whether the allocation of parental investment determined by hatching pattern is optimal for parent or offspring. Here, we manipulated the hatching pattern of the burying beetle Nicrophorus quadripunctatus to demonstrate the influence of hatching pattern on the allocation of parental investment. We found that the total weight of a brood was largest in the group that mimicked the natural hatching pattern, with the offspring skewed towards early hatchers. This increases parental fitness. However, hatching patterns with more later hatchers had heavier individual offspring weights, which increases offspring fitness, but this hatching pattern is not observed in the wild. Thus, our study suggests that the natural hatching pattern optimizes parental fitness, rather than offspring fitness.     

Relationships between egg size, chick size at hatching, and chick survival in the Whimbrel Numenius phaeopus

The relationships between egg size, chick size at hatching and chick survival in Whimbrels Numenius phaeopus were studied over a three-year period in the Shetland Isles. Three measurements of chick size at hatching were all positively correlated with egg volume, though the relationship was strongest with hatchling body-weight. In two of the three years the proportion of chicks from a brood which survived to fledging increased significantly with the mean hatching weight of chicks in the brood. Within broods, a significant effect of hatching weight on survival was detectable only up to 7 days after hatching. Between years the egg volumes and hatchling weights of individual female Whimbrels showed relatively little variability, indicating that these attributes could be controlled to a large extent by inheritance.     

Parental investment and brood defence by made and female great skuas Catharacts skua: the influence of food supply, laying date, body size and body condition

This paper tests the predictions of parental investment theory and other hypotheses relating to variation in brood defence by examining aggression displayed by great skuas Catharacta skua towards intruders within their territories. Aggression serves the function of nest defence; hatching success of adults breeding in Shetland increased with aggression displayed during incubation, and the correlation between aggression and hatching success was apparent in three separate age classes. Adults displayed higher levels of aggression and greater parental investment in reproduction in years of poor food supply. This was not due to an increase in adults' expectations of future benefits of brood defence with increased investment. since hatching success was unaffected by food supply, and breeding success was lower in years of poor food supply. The observed increase in aggression therefore supports parental investment theory. Aggression increased with body condition index (in terms of mass corrected for body size) for females, but decreased with increasing body condition index for males. This probably reflects size-specific differences in the relative benefits of weight and manoeuvrability as means of promoting effective brood defence and reducing the risk of injury to parents. Aggression may also reflect adult quality, with body condition reflecting quality in opposite ways in males and females, as a result of their different roles during the breeding season.     

Reproductive success and failure: the role of winter body mass in reproductive allocation in Norwegian moose

A life history strategy that favours somatic growth over reproduction is well known for long-lived iteroparous species, especially in unpredictable environments. Risk-sensitive female reproductive allocation can be achieved by a reduced reproductive effort at conception, or the subsequent adjustment of investment during gestation or lactation in response to unexpected environmental conditions or resource availability. We investigated the relative importance of reduced investment at conception compared with later in the reproductive cycle (i.e. prenatal, perinatal or neonatal mortality) in explaining reproductive failure in two high-density moose (Alces alces) populations in southern Norway. We followed 65 multiparous, global positioning system (GPS)-collared females throughout the reproductive cycle and focused on the role of maternal nutrition during gestation in determining reproductive success using a quasi-experimental approach to manipulate winter forage availability. Pregnancy rates in early winter were normal (≥0.8) in all years while spring calving rates ranged from 0.4 to 0.83, with prenatal mortality accounting for most of the difference. Further losses over summer reduced autumn recruitment rates to 0.23–0.69, despite negligible predation. Over-winter mass loss explained variation in both spring calving and autumn recruitment success better than absolute body mass in early or late winter. Although pregnancy was related to body mass in early winter, overall reproductive success was unrelated to pre-winter body condition. We therefore concluded that reproductive success was limited by winter nutritional conditions. However, we could not determine whether the observed reproductive allocation adjustment was a bet-hedging strategy to maximise reproduction without compromising survival or whether females were simply unable to invest more resources in their offspring.     

Prolactin stress response does not predict brood desertion in a polyandrous shorebird

One of the fundamental principles of the life-history theory is that parents need to balance their resources between current and future offspring. Deserting the dependent young is a radical life-history decision that saves resources for future reproduction but that may cause the current brood to fail. Despite the importance of desertion for reproductive success, and thus fitness, the neuroendocrine mechanisms of brood desertion are largely unknown. We investigated two candidate hormones that may influence brood desertion in the Kentish plover Charadrius alexandrinus: prolactin ('parental hormone') and corticosterone ('stress hormone'). Kentish plovers exhibit an unusually diverse mating and parental care system: brood desertion occurs naturally since either parent (the male or the female) may desert the brood after the chicks hatch and mate with a new partner shortly after. We measured the hormone levels of parents at hatching using the standard capture and restraint protocol. We subsequently followed the broods to determine whether a parent deserted the chicks. We found no evidence that either baseline or stress-induced prolactin levels of male or female parents predicted brood desertion. Although stress-induced corticosterone levels were generally higher in females compared with males, individual corticosterone levels did not explain the probability of brood desertion. We suggest that, in this species, low prolactin levels do not trigger brood desertion. In general, we propose that the prolactin stress response does not reflect overall parental investment in a species where different parts of the breeding cycle are characterized by contrasting individual investment strategies.     

Reproductive cycle in female Waigieu seaperch (Psammoperca waigiensis) reared under different salinity levels and the effects of dopamine antagonist on steroid hormone levels

Waigieu seaperch (Psammoperca waigiensis) is a tropical marine finfish species, which may inhabit wide range of salinity during the entire life cycle. Regardless of the wide salinity tolerance, little is known about how salinity may influence the reproductive endocrinology of this important tropical species. In the present study, we investigated the seasonal variations in steroid hormone levels, oocyte maturation (OM) and ovulation in fish reared under different salinity levels. In addition, we investigated the effects a dopamine antagonist (domperidone: DOM) during the peak spawning period. Mature brood fish at 3 years old were cultured in four different salinities of 5, 10, 20 and 30‰ (part per thousand, ppt) from March to December 2007. Blood samples were collected monthly and key steroid hormones (testosterone (T), 11-ketotestosterone (11-KT), 17β-estradiol (E2) and progesterone (P)) levels in female fish plasma were analyzed using enzyme immunoassay method. OM, ovulation and several reproductive performance indexes were evaluated twice per month during the breeding season. Plasma hormone analysis showed significant differences in fish groups cultured in different salinities during the seasonal cycle. The gonadosomatic index (GSI) gradually increased from March, peaking in July at 10 and 30‰, and then decreased thereafter until December. Plasma steroid levels (E2, T, 11-KT and P) were observed in variable concentrations during the spawning period and showed monthly fluctuations that were apparently salinity dependent during on and off-seasons. Interestingly, 11-KT levels were found in small amounts in female fish and peaked in September at salinities of 10 and 20‰. DOM exposure produced significant differences in steroid hormone levels that were apparently dependent on dose, exposure time and the individual hormone. The present study indicated that holding Waigieu seaperch brood fish in different salinities during the breeding season produced significant effects on gonadal development and spawning incidences. The first spawning was observed on 8th of April for all groups after 100% water exchange at night. No significant differences of absolute and relative fecundity were observed between groups. Fish held at 10 ppt salinity during the breeding season spawned naturally, but maturity and spawning rates were reduced and no hatching was observed at the same salinity. Fish holding at 5‰ resulted to 100% mortality in brood fish. This indicated that salinity limits for oocyte maturation or ovulation and subsequent reduction in spawning, but embryonic development requires a higher salinity. The effect of salinity on embryo development and hatching rate suggests that in breeding season the brood fish spawned in areas where the salinity and other parameters are favorable.     

Temporal and sex-specific patterns of breeding territory defense in a color-polymorphic cichlid fish

In biparental species, the costs and benefits of parental investment can vary between the sexes and shift over time. However, such sex-specific and temporal changes in territory defense are not well understood. Here, we experimentally investigated parental investment in breeding territory defense in a feral population of the color-polymorphic, biparental cichlid fish, the red devil (Amphilophus labiatus). We presented either gold- or dark-colored conspecific intruder models (i.e., dummy models) to A. labiatus pairs at three key stages during the breeding cycle (i.e., after pair formation, after eggs have been laid, and when fry were free-swimming). We found that males were more aggressive when the pair first formed, whereas females significantly increased their territory defense with time, and were most aggressive when fry were free-swimming. These results show that parental roles in territory defense can markedly shift over key stages of the breeding cycle. Our results demonstrate that parental behaviors may not only vary between the sexes, but can also shift dramatically over the course of the brood cycle.