Individual differences in reproductive costs examined using multi-state methods

1.?Trade-offs among life-history traits are common because individuals have to partition limited resources between multiple traits. Reproductive costs are generally assumed to be high, resulting in reduced survival and fecundity in the following year. However, it is common to find positive rather than negative correlations between life-history traits. 2.?Here, we use a data set from the individual-based study of red deer on the Isle of Rum to examine how these costs vary between individuals and at different ages, using multi-state mark-recapture methodology. 3.?Females that had reproduced frequently in the past incurred lower costs of reproduction in terms of survival in the following year and were more likely to reproduce in two consecutive years. Older individuals and those that had not reproduced frequently exhibited higher costs. 4.?These results highlight the importance of considering heterogeneity and individual quality when examining trade-offs and demonstrate the effectiveness of using detailed long-term data sets to explore life-history strategies using multi-state mark-recapture models.     

Effects of primary seed dormancy on lifetime fitness of Arabidopsis thaliana in the field

Background and AimsSeed dormancy determines the environmental niche of plants in seasonal environments, and has consequences for plant performance that potentially go far beyond the seed and seedling stages. In this study, we examined the cascading effects of seed dormancy on the expression of subsequent life-history traits and fitness in the annual herb Arabidopsis thaliana.MethodsWe planted seeds of >200 recombinant inbred lines (RILs) derived from a cross between two locally adapted populations (Italy and Sweden), and both parental genotypes at the native site of the Swedish population in three consecutive years. We quantified the relationship between primary seed dormancy and the expression of subsequent life-history traits and fitness in the RIL population with path analysis. To examine the effects of differences in dormancy on the relative fitness of the two parental genotypes, we planted dormant seeds during the seed dispersal period and non-dormant seeds during the germination period of the local population.Key ResultsIn the RIL population, strong primary dormancy was associated with high seedling survival, but with low adult survival and fecundity, and path analysis indicated that this could be explained by effects on germination timing, rosette size and flowering start. The relationship between primary seed dormancy and germination proportion varied among years, and this was associated with differences in seasonal changes in soil moisture. The planting of dormant and non-dormant seeds indicated that the lower primary dormancy of the local Swedish genotype contributed to its higher germination proportion in two years and to its higher fecundity in one year.ConclusionsOur results show that seed dormancy affects trait expression and fitness components across the life cycle, and suggest that among-year variation in the incidence of drought during the germination period should be considered when predicting the consequences of climatic change for population growth and evolution.     

Ecological,Evolutionary and Social Constraints on Reproductive Effort: Are Hoary Marmots Really Biennial Breeders?

Biennial breeding is a rare life-history trait observed in animal species living in harsh, unproductive environments. This reproductive pattern is thought to occur in 10 of 14 species in the genus Marmota, making marmots useful model organisms for studying its ecological and evolutionary implications. Biennial breeding in marmots has been described as an obligate pattern which evolved as a mechanism to mitigate the energetic costs of reproduction (Evolved Constraint hypothesis). However, recent anecdotal evidence suggests that it is a facultative pattern controlled by annual variation in climate and food availability (Environmental Constraint hypothesis). Finally, in social animals like marmots, biennial breeding could result from reproductive competition between females within social groups (Social Constraint hypothesis). We evaluated these three hypotheses using mark-recapture data from an 8-year study of hoary marmot (Marmota caligata) population dynamics in the Yukon. Annual variation in breeding probability was modeled using multi-state mark-recapture models, while other reproductive life-history traits were modeled with generalized linear mixed models. Hoary marmots were neither obligate nor facultative biennial breeders, and breeding probability was insensitive to evolved, environmental, or social factors. However, newly mature females were significantly less likely to breed than older individuals. Annual breeding did not result in increased mortality. Female survival and, to a lesser extent, average fecundity were correlated with winter climate, as indexed by the Pacific Decadal Oscillation. Hoary marmots are less conservative breeders than previously believed, and the evidence for biennial breeding throughout Marmota, and in other arctic/alpine/antarctic animals, should be re-examined. Prediction of future population dynamics requires an accurate understanding of life history strategies, and of how life history traits allow animals to cope with changes in weather and other demographic influences.     

Individual heterogeneity in life-history trade-offs with age at first reproduction in capital breeding elephant seals

Recruitment age plays a key role in life-history evolution. Because individuals allocate limited resources among competing life-history functions, theory predicts trade-offs between current reproduction and future growth, survival and/or reproduction. Reproductive costs tend to vary with recruitment age, but may also be overridden by fixed individual differences leading to persistent demographic heterogeneity and positive covariation among demographic traits at the population level. We tested for evidence of intra- and inter-generational trade-offs and individual heterogeneity relating to age at first reproduction using three decades of detailed individual life-history data of 6,439 capital breeding female southern elephant seals. Contrary to the predictions from trade-off hypotheses, we found that recruitment at an early age was associated with higher population level survival and subsequent breeding probabilities. Nonetheless, a survival cost of first reproduction was evident at the population level, as first-time breeders always had lower survival probabilities than prebreeders and experienced breeders of the same age. However, models accounting for hidden persistent demographic heterogeneity revealed that the trade-off between first reproduction and survival was only expressed in “low quality” individuals, comprising 35% of the population. The short-term somatic costs associated with breeding at an early age had no effect on the ability of females to allocate resources to offspring in the next breeding season. Our results provide strong evidence for individual heterogeneity in the life-history trajectories of female elephant seals. By explicitly modeling hidden persistent demographic heterogeneity we show that individual heterogeneity governs the expression of trade-offs with first reproduction in elephant seals.     

Survival costs of reproduction in a short-lived perennial plant: Live hard, die young

According to life-history theory, reproductive investments involve costs in terms of growth, future fecundity, and/or survival. However, studies to date have often failed to detect costs of reproduction, with survival costs among the less documented. We investigated the cost of reproduction in Helianthemum squamatum (Cistaceae), a short-lived perennial of semiarid Mediterranean environments. After experimental flower removal, we evaluated next season's growth, reproduction, and survival of the plants. We also monitored an indicator of plant physiological status (F(v)/F(m)) and leaf nutrient concentration at key phenological stages during reproduction. Survival rate in deblossomed plants was significantly higher than in control plants. As far as we know, this is the first experimental evidence of a survival cost of reproduction in a perennial plant. In contrast, no cost to growth or reproduction was found during the next season, and no significant differences in F(v)/F(m) or leaf nutrients were found between control and deblossomed plants. Helianthemum squamatum's success in semiarid Mediterranean ecosystems seems to rely on a persistent seed bank, combined with a sustained high reproductive output at the expense of survival. We conclude that this strategy might be more common than previously thought among short-lived shrubby plants growing in stressful Mediterranean areas.     

SPATIAL AND TEMPORAL VARIATION IN SELECTION ON CORRELATED LIFE-HISTORY TRAITS AND PLANT SIZE IN CHAMAECRISTA FASCICULATA

Intraspecific studies of selection on multiple traits of a plant's life history provide insight as to how the composite life history of an organism evolves. Current understanding of selection on plant life-history traits is deficient in three important areas: 1) the effects of selection through correlated traits, 2) the effects of selection on a trait throughout the plant's lifetime, and 3) spatial and temporal variation in selection on plant life-history traits among populations and years. This study documents spatial and temporal variation in selection on three life-history and two morphological traits for two natural populations of Chamaecrista fasciculata, a native summer annual. Life-history and morphological traits (date of seedling emergence, size at establishment, size prior to reproduction, date of initial flowering, and date of initial fruit maturation) varied significantly between sites and/or years. Selection on traits varied either spatially, between sites and among transects within one site, or temporally, between years. In addition, life-history traits were phenotypically correlated among themselves and with morphological traits; correlations were generally constant over time and space. Indirect selection caused changes in means and variances in traits not under direct selection, but which were correlated with traits under selection. Selection on date of emergence varied in direction and magnitude among different life-cycle stages, while selection on other traits varied only in magnitude among life stages of the plant. This study documents the complexity of the selective process and the importance of considering multiple life stages and traits when studying the evolution of life-history traits.     

Seed yield,seed size and germination behaviour in the annualPogogyne abramsii

Summary Models of the evolution of seed dormancy reveal that dormancy is favoured either when opportunities for establishment vary over time and when there is wide variation in the probability of success, or when the probability of success is limited by frequency dependence. Empirical evidence supporting the temporal heterogeneity hypothesis exists, but there is scant evidence for dormancy being favoured by frequency dependent competition among seedlings. We test the hypothesis that the intensity of between-sib competition should favour a positive relationship between maternal fecundity and seed dormancy. This hypothesis is supported for the rare, vernal pool annual,Pogogyne abramsii: the proportion of dormant offspring was significantly higher among high fecundity mothers than among other mothers. Dormancy inP. abramsii is controlled by the seed coat, a maternal tissue, so delaying germination favours the inclusive fitness of mothers by reducing the potential for competition among siblings. Seed weight and time to first germination varied significantly amongP. abramsii plants and mean seed weight increased linearly with plant biomass. Seed weight and seed number are independently regulated by plant size. Overall, seed weight varied 10-fold and variability in seed weight within mothers was not explained by plant biomass, seed yield or mean seed weight. GerminableP. abramsii seeds were significantly heavier than dormant seeds, and germinable seeds heavier than 0.20 mg germinated more rapidly than those smaller than 0.20 mg.     

Reductions in abscisic acid are linked with viviparous reproduction in mangroves

We investigate physiological mechanisms behind the convergent evolutionary loss of seed dormancy in plant lineages, focusing on mangroves as a model system. More than 60 angiosperm families, including several mangrove taxa, contain species with seeds that are intolerant of drying and do not undergo dormancy. These desiccation-intolerant species occur with disproportionate frequency in wet or coastal tropical habitats. In plants, the hormone abscisic acid (ABA) coordinates both the development of desiccation tolerance during the onset of seed dormancy and whole-organism responses to flooding. Thus, changes in ABA levels and/or modes of action in different plant compartments are implicated in the repeated evolutionary loss of seed dormancy among species of wet habitats. We compare ontogenetic dynamics of ABA levels in embryonic, maternal, and mature vegetative tissue of four phylogenetically independent pairs of related viviparous mangroves and nonviviparous nonmangroves. We demonstrate that ABA levels are consistently lower in embryos of viviparous mangrove taxa than embryos of nonmangrove, nonviviparous sister taxa. In contrast, elevated tissue concentrations of ABA characterize leaves of all mangrove species tested, while ABA levels in maternal tissues vary among mangrove species. These commonalities suggest a functionally important trade-off between the maintenance of embryonic development and the adjustment of the parent tree to salinity stress. This study yields comparative data on seed physiology in naturally occurring desiccation-intolerant species, for which these data are currently scarce, and demonstrates a potentially significant role of phytohormones in the evolution of plant life histories.     

Egg Size Effects across Multiple Life-History Stages in the Marine Annelid Hydroides diramphus

The optimal balance of reproductive effort between offspring size and number depends on the fitness of offspring size in a particular environment. The variable environments offspring experience, both among and within life-history stages, are likely to alter the offspring size/fitness relationship and favor different offspring sizes. Hence, the many environments experienced throughout complex life-histories present mothers with a significant challenge to optimally allocate their reproductive effort. In a marine annelid, we tested the relationship between egg size and performance across multiple life-history stages, including: fertilization, larval development, and post-metamorphosis survival and size in the field. We found evidence of conflicting effects of egg size on performance: larger eggs had higher fertilization under sperm-limited conditions, were slightly faster to develop pre-feeding, and were larger post-metamorphosis; however, smaller eggs had higher fertilization when sperm was abundant, and faster planktonic development; and egg size did not affect post-metamorphic survival. The results indicate that egg size effects are conflicting in H. diramphus depending on the environments within and among life-history stages. We suggest that offspring size in this species may be a compromise between the overall costs and benefits of egg sizes in each stage and that performance in any one stage is not maximized.     

Survival,density and population composition of bellbirds (Anthornis melanura) on the Poor Knights Islands,New Zealand

Abstract

Annual survival by age, sex and population size and structure were estimated for bellbirds (Anthornis melanura) from a banding study undertaken on 66 ha Aorangi Island, the Poor Knights Islands, during the period 1978–86. The most parsimonious model showed that survival varied little with age and between years, and survival of males was significantly greater than that of females. Recapture probabilities of males and females were similar, but did vary with age and among years. There was a greater proportion of adult males than adult females and sub‐adult males and females in the population, and among adults the male bias in the population was greater during the breeding season than at other times of the year. The high density of bellbirds on the Poor Knights and larger size of males may explain the reduced survival of females relative to that of males.     

Seeds photoblastism and its relationship with some plant traits in 136 cacti taxa

Seed germination triggered by light exposure (positive photoblastism) has been determined in quantitative studies for numerous plant families and species. For Cactaceae, positive photoblastism is thought to be associated with life form and seed mass, but this association has never been evaluated. To explore hypotheses on associations between seed mass, seed dispersal, seed dormancy, life form, taxa and plant height with Relative Light Germination (RLG) in Cactaceae, we evaluated the effect of light on seed germination of 136 taxa. The taxa studied are native to several countries: México, Chile, Argentina, Brazil, Perú, USA, and Venezuela. Seed traits contrasted with RLG were life form, seed mass, seed dispersal, seed dormancy, adult plant height and taxon. We found some differences between RLG among taxa; Cacteae, Pachycereeae and Trichocereeae had higher RLG than Notocacteae. RLG was lower for seeds from taller than for shorter taxa, and lower for taxa with heavier seeds than for taxa with lighter seeds. Dispersal syndrome groups varied with RLG. RLG did not differ between cylindrical and globose taxa. Trends found here were in agreement with expectations for small-seeded species to have a light requirement to germinate more often than large-seeded species. This is the first time that cactus height is related to photoblastism. It is possible that seeds from tall plants are larger and thus have the capacity to produce taller seedlings than those from small plants, and that seedlings from large seeds with more resources have the ability to emerge from greater soil depths than those from small seeds.     

Effects of plant hybridization on herbivore-parasitoid interactions

We studied the effects of host plant hybridization on the survival and mortality of the leaf-mining moth Phyllonorycter salicifoliella on hybrid and parental willow plants in the field and in a common garden experiment. P. salicifoliella survival differed significantly among three willow taxa in the field in 1994 but not in the field in 1995 or in the common garden. Parasitism by eulophid wasps differed significantly among taxa in 1994 and appeared to account for the variation in their survival. In the field in 1995, host feeding predation varied significant among taxa. The theory of tritrophic interactions predicts that plant genotype can affect natural enemy impact, and this study supports this prediction. Significant variation in survival and eulophid parasitism was also found among genotypes within taxa in the field in both years and in the common garden experiment. The common garden results show that genetic differences in plants affect the herbivore-parasitoid interaction. Variation among years in the patterns of survival and causes of mortality among field plants suggest that genotype by environment interactions may be important. Received: 1 March 1996 / Accepted: 4 November 1996     

Species traits and shoot-root biomass allocation in 20 dry-grassland species

Aims A plant has a limited amount of resources at any time and it allocates them to different structures. In spite of the large number of previous studies on allocation patterns within single species, knowledge of general patterns in species allocation is still very limited. This is because each study was done in different conditions using different methodology, making generalization difficult. We investigate intraspecific above- versus below-ground biomass allocation among individuals across a spectrum of dry-grassland plant species at two different developmental stages and ask whether allocation is age- and species specific, and whether differences among species can be explained by their life-history traits and phylogeny.Methods We collected data on above- and below-ground biomass of seedlings and adult plants of 20 species from a common garden experiment. We analysed data on shoot–root biomass allocation allometrically and studied the relationship between the allometric exponents (slopes on log–log scale), species life-history traits and phylogenetic distances.Important findings We found isometric as well as allometric patterns of biomass allocation in the studied species. Seedlings and adult individuals of more than half of the species differed in their above- versus below-ground biomass allometric exponents. Seedlings and adult individuals of the remaining species differed in their allometric coefficients (intercepts). Annual species generally allocated proportionally more to above- than below-ground biomass as seedlings than as adults, whereas perennial species showed the opposite pattern. Plant life-history traits, such as plant life span, age of first flowering, month in which the species begin flowering and specific leaf area were much more important in explaining differences in shoot–root allometry among species than were phylogenetic relationships. This suggests that allocation patterns vary greatly among closely related species but can be predicted based on species life-history traits.     

Connecting fruit production to seedling establishment in two co-occurring <Emphasis Type="Italic">Miconia</Emphasis> species: consequences of seed dispersal by birds in upper Amazonia

This study investigated links between seed production by two species of Miconia (Melastomataceae), whose seeds are dispersed by birds, and later stages of recruitment in lowland forests of eastern Ecuador. Seed dispersal and survival in later stages are crucial for understanding and predicting patterns of plant population dynamics as well as for understanding patterns of diversity in tropical forests. A major goal was to determine if the spatial template of seed deposition established by birds predicted probability of recruitment. We used observational and experimental approaches to compare patterns of recruitment in Miconia fosteri and M. serrulata. We calculated probabilities of transition between successive stages of recruitment for each species in three habitats. The number of plants with fruit, number of fruits removed, and, to a lesser extent, patterns of seed deposition varied between species and among habitats, whereas seed survival, germination, and establishment showed little variation among habitats. The location of seed deposition directly influenced the cumulative probabilities of survival. Among-habitat differences in the probabilities of recruitment set by seed deposition were not modified by later stages, although probability of recruitment was 2.5 times higher for M. serrulata than for M. fosteri after 1 year. The more critical stages for recruitment were seed removal and deposition. Our results from multiple life-cycle stages suggest that habitat associations among plants that reach reproductive maturity become established at early life stages and were mostly a consequence of seed dispersal by birds. These results differ from those obtained in temperate zones and suggest fundamental differences in the importance of recruitment processes. Dispersers, such as manakins, play significant roles in recruitment and population dynamics of M. fosteri, M. serrulata and numerous other understory plants of Neotropical forests. Their role in plant recruitment could be much greater than previously considered in megadiverse tropical forests. Thus, loss of dispersers could have long-term and far-reaching implications for maintenance of diversity.     

Selection on offspring size varies within and among families in relation to host nutritional quality

Life history theory often assumes a positive relationship between offspring size and fitness, although the strength and form of this relationship is expected to vary with environmental conditions. In arthropods, surprisingly few studies have examined the influence of larval environment on the offspring size–fitness relation. In phytophagous insects, the few studies that have examined variation in larval host plants have found a negative correlation between host plant nutritional quality and the strength of selection favoring larger offspring size, suggesting that this pattern might be general. I present experimental evidence for such a relationship in a population of the moth Rothschildia lebeau feeding on its three primary host plant species. Unlike previous studies, I consider the effect of offspring size on growth and survival at two levels, both among families and among siblings within families. Neonate caterpillar mass had a significant effect on growth and survival. The effect on growth, however, was weak, resulted primarily from variation among families, and did not differ among host plant diets. The effect on survival was stronger and varied among host plant diets, among families, and within families on different host plants in a manner that was generally consistent with the hypothesized negative correlation between host plant nutritional quality and the strength of selection favoring larger offspring size. Overall, results suggest that the consequences of variation in offspring size for survival within and among families are host plant-dependent in this system.     

Evaluation of reproductive costs for Weddell seals in Erebus Bay, Antarctica

1. Organisms balance current reproduction against future survival and reproduction, which results in life-history trade-offs. These trade-offs are also known as reproductive costs and may represent significant factors shaping life-history strategy for many species. 2. Using multistate mark-resight models and 26 years of mark-resight data (1979-2004), we estimated the costs of reproduction to survival and reproductive probabilities for Weddell seals in Erebus Bay, Antarctica and evaluated whether this species either conformed to the 'prudent parent' reproductive strategy predicted by life-history theory for long-lived mammals or alternatively, incurred costs to survival in order to reproduce in a variable environment (flexible-strategy hypothesis). 3. Results strongly supported the presence of reproductive costs to survival (mean annual survival probability was 0.91 for breeders vs. 0.94 for nonbreeders), a notable difference for a long-lived mammal, demonstrating that investment in reproduction does result in a cost to survival for Weddell seals, contrary to the prudent parent hypothesis. 4. Reproductive costs to subsequent reproductive probabilities were also present for first-time breeders (mean probability of breeding the next year was 31.3% lower for first-time breeders than for experienced breeders), thus supporting our prediction of the influence of breeding experience. 5. We detected substantial annual variation in survival and breeding probabilities. Breeding probabilities were negatively influenced by summer sea-ice extent, whereas weak evidence suggested that survival probabilities were affected more by winter sea-ice extent, and the direction of this effect was negative. However, a model with annual variation unrelated to any of our climate or sea-ice covariates performed best, indicating that further study will be needed to determine the appropriate mechanism or combination of mechanisms underlying this annual variation.     

It is risky out there: the costs of emergence and the benefits of prolonged dormancy

Prolonged dormancy is a stage in herbaceous perennial plants in which some individuals remain alive below ground for one or more growing seasons instead of emerging. Prolonged dormancy is puzzling, because foregoing opportunities for growth and reproduction seems costly. However, studies have shown that it buffers plants from the negative consequences associated with environmental stochasticity, suggesting that dormancy is a beneficial strategy to avoid the risks of stress above ground. If so, emergence during unfavorable conditions should have significant costs. Here, we test the hypothesis that emergence during times of stress has negative demographic consequences in a native perennial forb, Astragalus scaphoides, and investigate the potential underlying physiological mechanisms. We measured plant responses to a severe seasonal drought and an experimental defoliation to ask: (1) How do emergent plants respond to above-ground stress? (2) Do these responses have negative demographic consequences? and (3) Based on these responses, does stress increase the risk of emergence? Plants showed remarkable physiological tolerance to stress in the short term: high temperatures and low moisture did not have a strong effect on photosynthesis rates, and neither drought nor defoliation significantly impacted stored resources. However, stress did result in demographic costs for emergent plants relative to plants experiencing more favorable conditions. Drought resulted in decreased flowering probabilities relative to the long-term average and defoliation significantly increased mortality rates. These results demonstrate that the risk of emerging and experiencing stress entails considerable costs, supporting the hypothesis that prolonged dormancy is a beneficial strategy to avoid such risk.     

Differential seed germination responses to the ratio of red to far-red light in temperate and tropical species

Variation in vegetation density creates a range of red to far-red ratios of irradiance (R:FR) potentially permitting fine-scale discrimination of light conditions for seed germination. However, remarkably few studies have explored whether R:FR responses of germination vary among species that differ in distribution and life-history traits. In this study, we explored the relationships between R:FR requirements and four species characteristics: seed mass, latitudinal distribution (tropical vs. temperate), seed dormancy (dormant vs. nondormant), and plant growth form (woody vs. nonwoody). We obtained data on germination response to R:FR of 62 species from published literature and added new data for ten species from aseasonal tropical forests in Borneo. First, we analyzed whether species characteristics influenced overall light dependency of germination using phylogenetic logistic regression. We found that seed mass had a strong negative effect on light dependency, but that the seed mass at which tropical taxa had a 50 % probability of light dependency was 40 times that of temperate taxa. For light-dependent species, we found that the threshold R:FR that stimulates 50 % of maximum germination (R:FR₅₀) was also related to seed mass and latitudinal distribution. In agreement with an earlier study, we found that for temperate taxa, the R:FR₅₀ was significantly negatively correlated with seed mass. In contrast, for 22 tropical taxa, we found a significant positive correlation. These opposing relationships suggest contrasting selection pressures on germination responses of tropical taxa (mostly trees) and temperate herbaceous plants, and which are likely related to differences in seed longevity, seed burial rates, and reproductive output.     

Estimating Detection Probabilities of Waterfowl Broods From Ground-Based Surveys

ABSTRACT Brood:pair ratios could provide an economical method for assessing spatial or temporal variation in waterfowl productivity, but such estimators are severely biased by incomplete detection of broods. We conducted 3 sequential counts of 1,357 waterfowl broods in northeastern North Dakota, USA, and used closed-population mark-recapture models to estimate total brood abundance while controlling for variation in detection probabilities (p). Blue-winged teal (Anas discors) broods had the lowest average detection probability (p = 0.305), whereas diving-duck broods had the highest average detectability (p = 0.571). Detection was generally highest in morning or evening, but temporal patterns varied among species and there was no survey window that maximized detection probabilities for all species. Detection probabilities averaged 0.108 (SD = 0.056) higher for an experienced observer versus an inexperienced observer. Detection probabilities were 0.044 higher for roadside versus walk-up surveys and increased with increasing brood size, total brood abundance, survey date, wind speed, temperature, cloud cover, and amount of time spent surveying each wetland. Detection probabilities declined with increasing wetland size and amount of tall peripheral vegetation. Our mark-recapture results indicated that a traditional unreplicated brood survey would have missed 67.5% of estimated broods, summed over all species. Use of closed-population mark-recapture techniques provided an effective method for reducing this bias and identifying and quantifying factors that reduce detection probabilities of waterfowl broods. We recommend that future brood surveys incorporate 2 or 3 temporally segregated replicate counts to allow for formal estimation of detection probabilities.     

Mark-recapture and Moran curve estimates of the survival probabilities of an island population of tsetse flies Glossina morsitans morsitans (Diptera: Glossinidae)

A study on populations of Glossina morsitans morsitans Westwood on Antelope Island, Lake Kariba, Zimbabwe provided Jolly-Seber (J-S) mark-recapture estimates of adult survival and Moran curve estimates of the overall survival of all developmental stages. For females, Moran survival estimates derived using ox fly-round catches showed similar trends to, but were more variable than, those calculated from J-S population estimates. Regression of one set on the other removed only 26% of the variance. Undue emphasis should not be placed on small changes in Moran survival estimates based on sequences of catches. Catch data cannot provide Moran estimates of male survival probabilities and no comparison is thus possible with estimates from the J-S data. The J-S and the Moran approaches were combined to estimate survival probabilities of the immature stages of male and female tsetse. The overall survival per three-week period averaged 45% for males and 59% for females, comprising mature adult survivals of 27 and 46%, and pooled survivals of immature stages of 59 and 77%, respectively. The high survival of immature flies is due to the sheltered, inactive nature of pupal life. Adult and overall survival probabilities were highly correlated in males (r(2) = 0.61) but less so in females (r(2) = 0.24) where capture rates were lower and variance in the results greater. Immature and overall survival was more highly correlated for both sexes, with r(2) = 0.77 and 0.53 for males and females respectively. When a fixed pupal mortality of 1% per day was assumed, estimates of the survival of young adult males suggested that these were even lower than the survival of mature flies at the harshest times of the year, but were not markedly different when overall survival was high. Assuming equal mortality in all adults enabled the estimation of pupal survival. These had high variances but there was no evidence of any difference between the survival probabilities of male and female pupae.