Estimates of Natural Selection in a Salmon Population in Captive and Natural Environments

Abstract:  Captive breeding is a commonly used strategy for species conservation. One risk of captive breeding is domestication selection—selection for traits that are advantageous in captivity but deleterious in the wild. Domestication selection is of particular concern for species that are bred in captivity for many generations and that have a high potential to interbreed with wild populations. Domestication is understood conceptually at a broad level, but relatively little is known about how natural selection differs empirically between wild and captive environments. We used genetic parentage analysis to measure natural selection on time of migration, weight, and morphology for a coho salmon ( Oncorhynchus kisutch ) population that was subdivided into captive and natural components. Our goal was to determine whether natural selection acting on the traits we measured differed significantly between the captive and natural environments. For males, larger individuals were favored in both the captive and natural environments in all years of the study, indicating that selection on these traits in captivity was similar to that in the wild. For females, selection on weight was significantly stronger in the natural environment than in the captive environment in 1 year and similar in the 2 environments in 2 other years. In both environments, there was evidence of selection for later time of return for both males and females. Selection on measured traits other than weight and run timing was relatively weak. Our results are a concrete example of how estimates of natural selection during captivity can be used to evaluate this common risk of captive breeding programs.     

Reconstruction of Pacific salmon abundance from riparian tree-ring growth.

We use relationships between modern Pacific salmon (Oncorhynchus spp.) escapement (migrating adults counted at weirs or dams) and riparian tree-ring growth to reconstruct the abundance of stream-spawning salmon over 150-350 years. After examining nine sites, we produced reconstructions for five mid-order rivers and four salmon species over a large geographic range in the Pacific Northwest: chinook (O. tschwatcha) in the Umpqua River, Oregon, USA; sockeye (O. nerka) in Drinkwater Creek, British Columbia, Canada; pink (O. gorbuscha) in Sashin Creek, southeastern Alaska, USA; chum (O. keta) in Disappearance Creek, southeastern Alaska, USA; and pink and chum in the Kadashan River, southeastern Alaska, USA. We first derived stand-level, non-climatic growth chronologies from riparian trees using standard dendroecology methods and differencing. When the chronologies were compared to 18-55 years of adult salmon escapement we detected positive, significant correlations at five of the nine sites. Regression models relating escapement to tree-ring growth at the five sites were applied to the differenced chronologies to reconstruct salmon abundance. Each reconstruction contains unique patterns characteristic of the site and salmon species. Reconstructions were validated by comparison to local histories (e.g., construction of dams and salmon canneries) and regional fisheries data such as salmon landings and aerial surveys and the Pacific Decadal Oscillation climate index. The reconstructions capture lower-frequency cycles better than extremes and are most useful for determination and comparison of relative abundance, cycles, and the effects of interventions. Reconstructions show lower population cycle maxima in both Umpqua River chinook and Sashin Creek pink salmon in recent decades. The Drinkwater Creek reconstruction suggests that sockeye abundance since the mid-1990s has been 15-25% higher than at any time since 1850, while no long-term deviations from natural cycles are detected for salmon in the Kadashan River or in Disappearance Creek. Decadal-scale cycles in salmon abundance with periods of 25-68 years were detected in all of the reconstructions. This novel approach provides river-specific, long-term perspectives on salmon abundance and cycles. Additionally, it provides a new frame of reference for maintaining and rebuilding individual stocks and for striking a balance between societal demands and the limited, always-changing salmon resource.     

Nonrandom, Size- and Timing-Biased Breeding in a Hatchery Population of Steelhead Trout

Abstract:  Hatcheries have been built and operated to buffer salmon and trout populations from overfishing and to compensate for habitat lost or degraded by human activities. These facilities are now so prevalent that in some cases hatchery-produced salmon outnumber salmon produced in the wild. By default, this makes them an important component in the current ecology and evolution of salmonids. Hatcheries differ from natural environments in many ways, and among the most fundamental is the necessity that humans select fish for breeding instead of allowing natural processes of mate choice and competition. We examined the mating system for steelhead trout ( Oncorhynchus mykiss ) at Forks Creek Hatchery in southwest Washington and investigated factors affecting selection of individual steelhead for spawning by the hatchery staff. Despite efforts by the staff to not spawn selectively, data on steelhead spawned over 7 years revealed selection for large adult body size and early reproductive timing and a tendency for size-assortative mating (i.e., large with large). Selection on size was related to selection on reproductive timing because early returning fish tended to be larger than those returning later. To improve the fitness of both hatchery fish destined to spawn in the wild and hatchery fish designated to be spawned in the hatchery, a better understanding of factors associated with the range of reproductive success and mate-choice mechanisms in the wild is vital. This knowledge may then be applied to artificial propagation programs designed for conservation or enhancement.     

Sulfur isotopes link overwinter habitat use and breeding condition in Double-crested Cormorants.

North American Double-crested Cormorant (Phalacrocorax auritus) populations have increased greatly. Both breeding and overwintering ground factors have likely contributed to these increases. However, demonstrating how overwintering conditions may affect breeding birds has not been possible because of the difficulty in linking breeding birds to their wintering grounds. Here, we demonstrate the utility of stable sulfur isotopes to elucidate overwintering habitat use by cormorants breeding on Lake Erie. Sulfur isotopes in feathers grown on overwintering grounds provided insights into the degree to which birds used freshwater vs. marine environments. The proportion of birds utilizing freshwater habitats increased through time. This change may have reflected increases in freshwater aquaculture (i.e., catfish) in the U.S. south. Examination of body condition in birds returning to breed on Lake Erie indicated that those individuals that solely used marine habitats for at least a portion of the overwintering period were in poorer condition than birds using freshwater. Enhanced foraging opportunities at aquaculture facilities may improve the fitness of individuals that have returned to breed after overwintering at such locations. This study is the first to demonstrate a linkage between overwinter habitat use and breeding ground parameters in Double-crested Cormorants. These results underscore that factors throughout the Mississippi flyway are likely acting together to regulate cormorant populations.     

Directional selection by fisheries and the timing of sockeye salmon (Oncorhynchus nerka) migrations.

The timing of migration from feeding to breeding areas is a critical link between the growth and survival of adult animals, their reproduction, and the fitness of their progeny. Commercial fisheries often catch a large fraction of the migrants (e.g., salmon), and exploitation rates can vary systematically over the fishing season. We examined daily records of sockeye salmon (Oncorhynchus nerka) in the Egegik and Ugashik management districts in Bristol Bay, Alaska (USA), for evidence of such temporally selective fishing. In recent years, the early migrants have experienced lower fishing rates than later migrants, especially in the Egegik district, and the median migration date of the fish escaping the fisheries has been getting progressively earlier in both districts. Moreover, the overall runs (catch and escapement) in the Egegik district and, to a lesser extent the Ugashik district, have been getting earlier, as predicted in response to the selection on timing. The trends in timing were not correlated with sea surface temperature in the region of the North Pacific Ocean where the salmon tend to concentrate, but the trends in the two districts were correlated with each other, indicating that there may be some common environmental influence in addition to the effect of selection. Despite the selection, both groups of salmon have remained productive. We hypothesize that this resilience may result from representation of all component populations among the early and late migrants, so that the fisheries have not eliminated entire populations, and from density-dependent processes that may have helped maintain the productivity of these salmon populations.     

Long-term demographic and genetic effects of releasing captive-born individuals into the wild

Because of continued habitat destruction and species extirpations, the need to use captive breeding for conservation purposes has been increasing steadily. However, the long-term demographic and genetic effects associated with releasing captive-born individuals with varied life histories into the wild remain largely unknown. To address this question, we developed forward-time, agent-based models for 4 species with long-running captive-breeding and release programs: coho salmon (Oncorhynchus kisutch), golden lion tamarin (Leontopithecus rosalia), western toad (Anaxyrus boreas), and Whooping Crane (Grus americana). We measured the effects of supplementation by comparing population size and neutral genetic diversity in supplemented populations to the same characteristics in unaltered populations 100 years after supplementation ended. Releasing even slightly less fit captive-born individuals to supplement wild populations typically resulted in reductions in population sizes and genetic diversity over the long term when the fitness reductions were heritable (i.e., due to genetic adaptation to captivity) and populations continued to be regulated by density-dependent mechanisms over time. Negative effects for species with longer life spans and lower rates of population replacement were smaller than for species with shorter life spans and higher rates of population replacement. Programs that released captive-born individuals over fewer years or that avoided breeding individuals with captive ancestry had smaller reductions in population size and genetic diversity over the long term. Relying on selection in the wild to remove individuals with reduced fitness mitigated some negative demographic effects, but at a substantial cost to neutral genetic diversity. Our results suggest that conservation-focused captive-breeding programs should take measures to prevent even small amounts of genetic adaptation to captivity, quantitatively determine the minimum number of captive-born individuals to release each year, and fully account for the interactions among genetic adaptation to captivity, population regulation, and life-history variation.     

Quantifying and comparing size selectivity among Alaskan sockeye salmon fisheries

Quantifying long-term size-selective harvest patterns is necessary for understanding the potential evolutionary effects on exploited species. The comparison of fishery selection patterns on the same species subject to different gear types, in different areas, and over multi-decadal periods can reveal the factors influencing selection. In this study we quantified and compared size-selective harvest by nine Alaskan sockeye salmon (Oncorhynchus nerka) fisheries to understand overall patterns. We calculated length-specific linear selection differentials (the difference in average length of fish before vs. after fishing), which are produced by different combinations of exploitation rates and length-selectivity values, and nonlinear standardized differentials, describing disruptive selection, across all years for each fishery. Selection differentials varied among years, but larger fish were caught in 73% of years for males and 84% of years for females, leaving smaller fish to spawn. Disruptive selection was observed on female and male fish in 84% and 92% of years, respectively. Linear selection was stronger on females than males in 77% of years examined, and disruptive selection was stronger on males in 71% of years. Selection pressure was influenced by a combination of factors under and beyond management control; analyses using mixed-effects models indicated that fisheries were less size selective in years when fish were larger than average and had lower exploitation rates. The observed harvest of larger than average sockeye salmon is consistent with the hypothesis that size-selective fishing contributes to decreasing age and length at maturation trends over time, but temporal variability in selection and strong disruptive selection suggests that the overall directional pressure is weaker than is often assumed in evolutionary models.     

Escape migration decisions in Eurasian Woodcocks: insights from survival analyses using large-scale recovery data

During unpredictable adverse conditions, endotherms can engage in emergency behaviors (movement, torpor, hyperphagia) to maintain energy balance and reduce mortality hazards. Bird “escape migration” is one of the most visible of these behaviors. In this study, we focus on a Eurasian Woodcock Scolopax rusticola population. Seasonal migrations bring this population from its breeding grounds in Eastern and Northern Europe to its wintering grounds in France. A varying number of these birds are also regularly reported from Spain, supposedly during additional escape movements that occur in winter. Using models that account for the imperfect detection rate of individuals and a large (>44,000 individuals) dataset combining information from the wintering and breeding ranges, we show that severe winters significantly reduced survival probability, but that migration to Spain increased only during the most intense cold spell that occurred over the 20-year study period. This suggests that the decision to resume migration during the winter is submitted to a threshold mechanism, which we discuss in the light of current models of migratory behavior.     

Breeding Periodicity for Male Sea Turtles,Operational Sex Ratios,and Implications in the Face of Climate Change

Abstract: Species that have temperature‐dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long‐standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single‐sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in‐water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female‐biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.     

Arrival timing and seasonal reproductive performance in a long-distance migratory landbird

The date when a landbird migrant arrives on its breeding grounds may have reproductive consequences. Generally, early arriving individuals begin breeding earlier and consequently experience greater seasonal reproductive performance. Here, we describe relationships between arrival timing and seasonal reproductive performance in the American redstart (Setophaga ruticilla), a long-distance passerine migrant, arriving at northerly breeding grounds in Michigans eastern Upper Peninsula. Evidence suggests that both males and females benefited from early arrival at the breeding grounds. Early males appeared to settle on higher quality territories and hatched nestlings sooner than later arrivals. Early females began their clutches early, produced heavier nestlings and possibly laid more eggs than later arrivals. Larger clutches and heavier offspring increase the likelihood of offspring recruiting into the breeding population. The findings of this study point to fitness consequences arising from when a bird arrives at its breeding grounds. These results also have implications for understanding how events occurring during spring migration influence reproductive performance as migratory delays likely influence arrival timing.Communicated by: W. Wiltschko     

Biotic control of stream fluxes: spawning salmon drive nutrient and matter export

Organisms can control movements of nutrients and matter by physically modifying habitat. We examined how an ecosystem engineer, sockeye salmon (Oncorhynchus nerka), influences seasonal fluxes of sediments, nitrogen (N), and phosphorus (P) in streams of southwestern Alaska. The purpose of this study was to investigate whether salmon act as net importers or net exporters of matter and nutrients from streams and how these roles change as a function of salmon population density. We measured discharge and concentrations of suspended sediments and total N and P every 7-14 days for up to four summers in 10 streams spanning a gradient in salmon densities. We statistically allocated whole-season fluxes to salmon activities, such as excretion and bioturbation, and to export by hydrologic discharge. In addition, we used counts of spawning salmon to estimate nutrient and matter imports by salmon to streams. Large seasonal pulses of suspended sediments, P, and N were associated with salmon spawning activities, often increasing export an order of magnitude higher than during pre-salmon levels. Years and streams with more salmon had significantly higher levels of export of sediments and nutrients. In addition, years with higher precipitation had higher background export of P and N. Salmon exported an average of the equivalent of 189%, 60%, and 55% of total matter, P, and N that salmon imported in their bodies. The relative magnitude of export varied; salmon exported more than their bodies imported in 80%, 20%, and 16% across all streams and years for sediments, P, and N, respectively. A bioassay experiment indicated that the P exported by salmon is directly available for use by primary producers in the downstream lake. These results demonstrate that salmon not only move nutrients upstream on large spatial scales via their migration from the ocean and subsequent death, but also redistribute matter and nutrients on finer spatial scales through their spawning activities.     

Selection of both habitat and genes in specialized and endangered caribou

Genetic mechanisms determining habitat selection and specialization of individuals within species have been hypothesized, but not tested at the appropriate individual level in nature. In this work, we analyzed habitat selection for 139 GPS-collared caribou belonging to 3 declining ecotypes sampled throughout Northwestern Canada. We used Resource Selection Functions comparing resources at used and available locations. We found that the 3 caribou ecotypes differed in their use of habitat suggesting specialization. On expected grounds, we also found differences in habitat selection between summer and winter, but also, originally, among the individuals within an ecotype. We next obtained Single Nucleotide Polymorphisms (SNPs) for the same caribou individuals, we detected those associated to habitat selection, and then identified genes linked to these SNPs. These genes had functions related in other organisms to habitat and dietary specializations, and climatic adaptations. We therefore suggest that individual variation in habitat selection was based on genotypic variation in the SNPs of individual caribou, indicating that genetic forces underlie habitat and diet selection in the species. We also suggest that the associations between habitat and genes that we detected may lead to lack of resilience in the species, thus contributing to caribou endangerment. Our work emphasizes that similar mechanisms may exist for other specialized, endangered species.     

Genetic Analyses to Establish Captive Breeding Priorities for Endangered Snake River Sockeye Salmon

The plight of the Snake River sockeye salmon presents a challenging genetic puzzle for conservation biologists. Although the Snake River sockeye has been declared endangered, assessment of captive breeding strategies are complicated because a healthy non-migratory strain of O. nerka (kokanee) resides in Redfish Lake, Idaho—the same lake the anadromous fish return to for spawning. The migration of 1-year-old fish (outmigrants) from the lake each year and the observation of non-migratory fish spawning in areas previously used by the sockeye further complicate the issue. We estimated the relatedness of these strains by direct examination of their genetic similarity. Mixed DNA fingerprint analyses suggested that the outmigrants were more closely related to the anadromous sockeye than to the kokanee. Closer analysis using a probe, One-HO.8, to examine allele frequencies at a single locus revealed polymorphism in a number of O. nerka populations. Within the Redfish Lake populations this probe detected an allele present at 0.21 frequency in the kokanee (n = 43), 0.01 (n = 324) in the outmigrants, and absent among 13 anadromous fish. These results support a close genetic relationship of the outmigrants to the anadromous sockeye and the probable utility of the outmigrants in a captive breeding program to restore the anadromous strain.     

Climate change threatens polar bear populations: a stochastic demographic analysis

The polar bear (Ursus maritimus) depends on sea ice for feeding, breeding, and movement. Significant reductions in Arctic sea ice are forecast to continue because of climate warming. We evaluated the impacts of climate change on polar bears in the southern Beaufort Sea by means of a demographic analysis, combining deterministic, stochastic, environment-dependent matrix population models with forecasts of future sea ice conditions from IPCC general circulation models (GCMs). The matrix population models classified individuals by age and breeding status; mothers and dependent cubs were treated as units. Parameter estimates were obtained from a capture-recapture study conducted from 2001 to 2006. Candidate statistical models allowed vital rates to vary with time and as functions of a sea ice covariate. Model averaging was used to produce the vital rate estimates, and a parametric bootstrap procedure was used to quantify model selection and parameter estimation uncertainty. Deterministic models projected population growth in years with more extensive ice coverage (2001-2003) and population decline in years with less ice coverage (2004-2005). LTRE (life table response experiment) analysis showed that the reduction in lambda in years with low sea ice was due primarily to reduced adult female survival, and secondarily to reduced breeding. A stochastic model with two environmental states, good and poor sea ice conditions, projected a declining stochastic growth rate, log lambdas, as the frequency of poor ice years increased. The observed frequency of poor ice years since 1979 would imply log lambdas approximately - 0.01, which agrees with available (albeit crude) observations of population size. The stochastic model was linked to a set of 10 GCMs compiled by the IPCC; the models were chosen for their ability to reproduce historical observations of sea ice and were forced with "business as usual" (A1B) greenhouse gas emissions. The resulting stochastic population projections showed drastic declines in the polar bear population by the end of the 21st century. These projections were instrumental in the decision to list the polar bear as a threatened species under the U.S. Endangered Species Act.     

Analysis of lipid components for determining the condition of anchovy larvae,Engraulis mordax

It has been presumed that intertidal spawning by Limulus polyphemus minimizes the loss of egges to subtidal predators; however, this strategy involves considerable risks. Massive beach strandings of adults accompany seasonal spawning migrations of crabs along Cape May in Delaware Bay, (USA). At least 190000 horseshoe crabs, approximating 10% of the adult population, died from beach stranding along the New Jersey shore of Delaware Bay during the 1986 (May to June) spawning season. Abnormalities of the telson (which is used in righting behavior) were significantly more common among stranded crabs than among individuals actively spawning on the intertidal beach. The number of stranded crabs per day was not correlated with tidal height or environmental variables (wind speed, wave height) which characterized the conditions at spawning. A complex suite of factors, including the size of the available spawning population, tidal and weather conditions, and beach slope, influence the number stranded during the breeding season. Horseshoe crab stranding results in a large loss of gravid females from the population, and may represent a major input of organic matter to intertidal sandy beaches in certain regions of Delaware Bay.     

Behaviour and physiology of sockeye salmon homing through coastal waters to a natal river

Adult sockeye salmon (Oncorhynchus nerka, N = 179) from six Fraser River populations (British Columbia) were intercepted in continental shelf waters ∼215 km from the Fraser River mouth, gastrically implanted with acoustic transmitters, non-lethally biopsied for blood biochemistry, gill Na⁺,K⁺-ATPase activity and somatic energy density and then released. Migration behaviour and travel times to the river mouth and into the river were monitored by underwater telemetry receivers positioned at the river mouth and in the river. Post-release survival of salmon was excellent, with 84% (N = 150) of fish reaching the furthest receiving station ∼85 km upriver. Fish from Late-summer run populations (Adams and Weaver Creek) averaged a migration rate of ∼20 km day⁻¹ through the marine area and held at the river mouth and adjacent areas for 7–9 days before entering the river. Summer-run populations (Birkenhead, Chilko, Horsefly and Stellako) had a migration rate ∼33 km day⁻¹ and held for only 1–3 days. Once in river, similar patterns were observed: Late-run populations migrated at ∼28 km day⁻¹ and Summer-run populations at ∼40 km day⁻¹. From point of release to the river mouth, males migrated faster than females, but once in river migration rates did not differ between sexes. Among all females, a correlation was discovered between levels of circulating testosterone and river entry timing. This correlation was not observed among males. Plasma K⁺, Cl⁻, glucose, lactate and osmolality were also correlated with entry timing in both sexes.     

Evolution of a stream ecosystem in recently deglaciated terrain

Climate change and associated glacial recession create new stream habitat that leads to the assembly of new riverine communities through primary succession. However, there are still very few studies of the patterns and processes of community assembly during primary succession for stream ecosystems. We illustrate the rapidity with which biotic communities can colonize and establish in recently formed streams by examining Stonefly Creek in Glacier Bay, Alaska (USA), which began to emerge from a remnant glacial ice mass between 1976 and 1979. By 2002, 57 macroinvertebrate and 27 microcrustacea species had become established. Within 10 years of the stream's formation, pink salmon and Dolly Varden charr colonized, followed by other fish species, including juvenile red and silver salmon, Coast Range sculpin, and sticklebacks. Stable-isotope analyses indicate that marine-derived nitrogen from the decay of salmon carcasses was substantially assimilated within the aquatic food web by 2004. The findings from Stonefly Creek are compared with those from a long-term study of a similarly formed but older stream (12 km to the northeast) to examine possible similarities in macroinvertebrate community and biological trait composition between streams at similar stages of development. Macroinvertebrate community assembly appears to have been initially strongly deterministic owing to low water temperature associated with remnant ice masses. In contrast, microcrustacean community assembly appears to have been more stochastic. However, as stream age and water temperature increased, macroinvertebrate colonization was also more stochastic, and taxonomic similarity between Stonefly Creek and a stream at the same stage of development was <50%. However the most abundant taxa were similar, and functional diversity of the two communities was almost identical. Tolerance is suggested as the major mechanism of community assembly. The rapidity with which salmonids and invertebrate communities have become established across an entire watershed has implications for the conservation of biodiversity in freshwater habitats.     

Inbreeding Depression in the Speke's Gazelle Captive Breeding Program

Abstract: The Speke's gazelle ( Gazella spekei ) captive breeding program has been presented as one of the few examples of selection reducing the genetic load of a population and as a potential model for the captive breeding of endangered species founded from a small number of individuals. In this breeding program, three generations of mate selection apparently increased the viability of inbred individuals. We reanalyzed the Speke's gazelle studbook and examined potential causes for the reduction of inbreeding depression. Our analysis indicates that the decrease in inbreeding depression is not consistent with any model of genetic improvement in the herd. Instead, we found that the effect of inbreeding decreased from severe to moderate during the first generation of inbreeding, and that this change is responsible for almost all of the decline in inbreeding depression observed during the breeding program. This eliminates selection as a potential explanation for the decrease in inbreeding depression and suggests that inbreeding depression may be more sensitive to environmental influences than is usually thought.     

Cooperation in breeding by nonreproductive wrens: kinship,reciprocity, and demography

Summary Stripe-backed wrens (Campylorhynchus nuchalis) often live as adults in large groups on permanent, communally defended territories. Nonbreeding adults cooperate in rearing the young of a single breeding pair; this aid substantially increases the reproductive success of the breeders. In a 6-year study in Venezuela of a completely colorbanded population of 25–30 groups, most adults participated in breeding only as helpers and priority to breeding status was strictly age-determined. Detailed behavioral observations at breeding nests with nestlings showed that, in a sample of 100, helpers nearly always contributed as much to the care of young as breeders. Further, aid-giving does not vary systematically with relatedness of ycung to helpers or with probability of future reciprocation by young. Young being raised are most often at least half siblings of helpers, but seldom return aid to adults that helped raise them. Even adopted helpers collaborate fully. Patterns of demography and dispersal show slow turnover of breeders, delayed reproduction, and a viscous population structure.Application of Hamilton's condition for selection for aid-giving reveals that most individuals in this population can maximize inclusive fitness in the first 2 years by helping instead of breeding. Variation in helping effort and in age of first breeding is related to variation in natal group size and competition resulting from variable demographic neighborhoods in different years or in different parts of the population. Because reciprocation in the form of specific alliance formation among nonreproductives is uncommon, nonspecific reciprocity between cohorts and kin selection account well for the observed pattern of age-dependence in first breeding. Nondiscriminating helping in this population is associated with stable monogamous pair boncs, stable territory boundaries and group membership, strict seniortiy for breeding position, high viscosity and consistent effectiveness of aid. Under these circumstances, very simple behavioral rules amounting to nearly automatic helping seem sufficient to confer critical inclusive fitness gain on helpers.     

Environmental conditions of Chilika Lake during pre and post hydrological intervention: an overview

Chilika Lake, the largest brackish water lagoon in Asia, is a prominent biodiversity hotspot along the Indian east coast. The geomorphology, water quality and biological productivity of the lake had undergone significant changes over the years under the influence of natural events and anthropogenic interventions. Decades of research have shown that the ecological changes of the lake system and its fisheries were influenced mainly by the water exchange between the lake and the sea that controls the salinity, siltation, macrophyte infestation and recruitment of marine forms. Depletion of fisheries and loss of biodiversity in 1980s and 1990s have been linked to the northward shifting of lake’s inlet and silting up of the outer channel. In order to improve the health of the lagoon and restoration of its biodiversity, a new mouth was dredged open in September 2000. Opening of the new mouth has resulted in some improvements with substantial increase in capture fishery, reduction in weed infestation, growth of seagrasses, appearance of dolphins and increase in the population of migratory birds. But, many stake holders claim that the problems associated with the lake ecology and its contribution to the socio-economic development of the stake holders still persist which could escalate in future.