Cryptic multiple hypotheses testing in linear models: overestimated effect sizes and the winner's curse

Fitting generalised linear models (GLMs) with more than one predictor has become the standard method of analysis in evolutionary and behavioural research. Often, GLMs are used for exploratory data analysis, where one starts with a complex full model including interaction terms and then simplifies by removing non-significant terms. While this approach can be useful, it is problematic if significant effects are interpreted as if they arose from a single a priori hypothesis test. This is because model selection involves cryptic multiple hypothesis testing, a fact that has only rarely been acknowledged or quantified. We show that the probability of finding at least one ‘significant’ effect is high, even if all null hypotheses are true (e.g. 40% when starting with four predictors and their two-way interactions). This probability is close to theoretical expectations when the sample size (N) is large relative to the number of predictors including interactions (k). In contrast, type I error rates strongly exceed even those expectations when model simplification is applied to models that are over-fitted before simplification (low N/k ratio). The increase in false-positive results arises primarily from an overestimation of effect sizes among significant predictors, leading to upward-biased effect sizes that often cannot be reproduced in follow-up studies (‘the winner's curse’). Despite having their own problems, full model tests and P value adjustments can be used as a guide to how frequently type I errors arise by sampling variation alone. We favour the presentation of full models, since they best reflect the range of predictors investigated and ensure a balanced representation also of non-significant results.     

Order restricted randomized designs and two sample inference

This paper develops a new design that relies on subjective judgment ranking to compare subsets of experimental units. This judgment ranking is used along with restricted randomization to improve statistical inference for the contrast between two levels of a treatment. The new design assigns the judgment ranked units in a subset to different treatments. Such an assignment translates the positive dependence among units within each subset into negative dependence for the estimators of treatment means, and hence leads to a reduction in variance for the contrast. For the proposed design, a test for the difference in means of two treatment levels is developed along with an associated confidence interval. It is shown that the null distribution of the proposed test is approximated reasonably well with the Student’s t-distribution for sample sizes as small as 6. A simulation study indicates that the proposed design is advantageous compared to its competitors in the literature for both high and low quality rankings. The new design’s advantage increases with the quality of rankings.     

Division of labor between scouts and recruits: genetic influence and mechanisms

Every recruitment system in social insects requires some individuals that serve as scouts, foragers that search independently for food sources. It is not well understood which factors influence whether an individual becomes a scout or a recruit, nor how the division of labor between the two forager groups is regulated. It is shown here for honeybees (Apis mellifera), using two different molecular techniques, that there is a genetically based difference in the probability that individuals will scout independently for food. In contrast to earlier suggestions, experimental tests showed that the age of a bee does not seem to influence its probability of becoming a scout or a recruit. Furthermore, scout bees do not search opportunistically for either pollen or nectar but, rather, individuals have preferences that are genetically based. These findings are discussed in the framework of foraging regulation by specialization in honeybees and the adaptive significance of polyandry. Received: 23 October 1997 / Accepted after revision: 10 April 1998     

Variation in foraging effort by lactating Antarctic fur seals: response to simulated increased foraging costs

Seasonally breeding predators, which are limited in the time available for provisioning young at a central location, and by the fasting abilities of the young, are likely to maximize energy delivery to the young by maximizing the rate of energy delivery averaged over the whole period of investment. Reduction in food availability or increased foraging costs will alter the optimal behavior of individuals. This study examined the behavioral adaptations of a diving predator, the Antarctic fur seal, to increased foraging costs during lactation. One group of mothers (n=5, treatment) was fitted with additional drag to increase the cost of transport in comparison with a control group (n=8). At the scales of the individual dives, the treatment group made more shorter, shallower (< 30 m) dives. Compensation for slower swimming speeds was achieved by diving at a steeper angle. Overall, diving behavior conformed to several specific theoretical predictions but there were also departures from theory, particularly concerning swimming speed during diving. Diving behavior appears to be adjusted to maximize the proportion of time spent at the bottom of dives. At the scale of diving bouts, no difference was observed between the treatment and control groups in terms of the frequency and duration of bouts and there was also no difference between the two groups in terms of the proportion of time spent diving. At the scale of complete foraging cycles, time taken to return to the pup was significantly longer in the treatment group but there was no difference in the rate of delivery of energy (measured from pup growth rate) to the pups in each group. Since mothers in the treatment group did not use significantly more body reserves, we conclude that behavioral adjustments at the scale of individual dives allowed mothers in the treatment group to compensate for the additional foraging costs. Pup growth rate appears to be less sensitive to the foraging conditions experienced by mothers than foraging trip duration. Received: 14 June 1996 / Accepted after revision: 16 November 1996     

Population biology and life strategies of Chlorophthalmus agassizii Bonaparte, 1840 (Pisces: Osteichthyes) in the Mediterranean Sea

The population biology and life strategies of Chlorophthalmus agassizii were studied in the Ionian Sea (eastern–central Mediterranean) using the data collected during the experimental trawl surveys carried out from 1995 to 2000. Depth-related trends of both density and size were found. With depth the former decreased while the latter increased. A typical bigger–deeper phenomenon was detected: young-of-the-year individuals occur on the shelf during autumn–winter months and move towards bathyal bottoms as they grow. The sampled population was made up of several size–age groups. The maximum age of 8 years was identified by means of otolith readings. The Von Bertalanffy growth parameters were estimated from the age–length key (L _∞=189.04±5.401 mm; k=0.24±0.021; t _o=−1.20±0.132; φ′=3.94) and modal progression analysis (L _∞=218.33±18.397 mm; k=0.164±0.028; t _o=−1.694±0.171; φ′=3.89). Reproduction of this monoecious fish was observed during summer–early autumn. Considering the female portion of the gonad, the size at attainment of 50% maturity was 115 mm TL. The corresponding age is within the third year of life. The simultaneous occurrence of oocytes in different development stages was shown in the ovary. Both the asynchronous ovary and oocyte size distribution indicate that C. agassizii spawns more than once during a reproductive season (batch spawner). Functional fecundity (on average 3,018 hydrated oocytes) was between 37 and 69% of the absolute fecundity and increased significantly with the individual size. Since adult specimens are mostly distributed on the slope, eggs and larvae develop in epipelagic waters and migrate in-shore where juvenile forms recruit on the shelf. Juveniles migrate ontogenetically towards deeper bottoms and after 2–3 years start to reproduce annually within a life span greater than 10 years.     

Present-day genetic composition suggests contrasting demographic histories of two dominant chaetognaths of the North-East Atlantic, <Emphasis Type="Italic">Sagitta elegans</Emphasis> and <Emphasis Type="Italic">S. setosa</Emphasis>

Sagitta elegans and S. setosa are the two dominant chaetognaths in the North-East (NE) Atlantic. They are closely related and have a similar ecology and life history, but differ in distributional ranges. Sagitta setosa is a typical neritic species occurring exclusively above shelf regions, whereas S. elegans is a more oceanic species with a widespread distribution. We hypothesised that neritic species, because of smaller and more fragmented populations, would have been more vulnerable to population bottlenecks resulting from range contractions during Pleistocene glaciations than oceanic species. To test this hypothesis we compared mitochondrial Cytochrome Oxidase II DNA sequences of S. elegans and S. setosa from sampling locations across the NE Atlantic. Both species displayed very high levels of genetic diversity with unique haplotypes for every sequenced individual and an approximately three times higher level of nucleotide diversity in S. elegans (0.061) compared to S. setosa (0.021). Sagitta setosa mitochondrial DNA (mtDNA) haplotypes produced a star-like phylogeny and a uni-modal mismatch distribution indicative of a bottleneck followed by population expansion. In contrast, S. elegans had a deeper mtDNA phylogeny and a multi-modal mismatch distribution as would be expected from a more stable population. Neutrality tests indicated that assumptions of the standard neutral model were violated for both species and results from the McDonald-Kreitman test suggested that selection played a role in the evolution of their mitochondrial DNA. Congruent with these results, both species had much smaller effective population sizes estimated from genetic data when compared to census population sizes estimated from abundance data, with a factor of ~10⁸–10⁹ difference. Assuming that selective effects are comparable for the two species, we conclude that the difference in genetic signature can only be explained by contrasting demographic histories. Our data are consistent with the hypothesis that in the NE Atlantic, the neritic S. setosa has been more severely affected by population bottlenecks resulting from Pleistocene range shifts than the more oceanic S. elegans.     

Closed population estimation models and their extensions in Program MARK

Program MARK provides > 65 data types in a common configuration for the estimation of population parameters from mark-encounter data. Encounter information from live captures, live resightings, and dead recoveries can be incorporated to estimate demographic parameters. Available estimates include survival (S or ϕ), rate of population change (λ), transition rates between strata (Ψ), emigration and immigration rates, and population size (N). Although N is the parameter most often desired by biologists, N is one of the most difficult parameters to estimate precisely without bias for a geographically and demographically closed population. The set of closed population estimation models available in Program MARK incorporate time (t) and behavioral (b) variation, and individual heterogeneity (h) in the estimation of capture and recapture probabilities in a likelihood framework. The full range of models from M ₀ (null model with all capture and recapture probabilities equal) to M _tbh are possible, including the ability to include temporal, group, and individual covariates to model capture and recapture probabilities. Both the full likelihood formulation of Otis et al. (1978) and the conditional model formulation of Huggins (1989, 1991) and Alho (1990) are provided in Program MARK, and all of these models are incorporated into the robust design (Kendall et al. 1995, 1997; Kendall and Nichols 1995) and robust-design multistrata (Hestbeck et al. 1991, Brownie et al. 1993) data types. Model selection is performed with AICc (Burnham and Anderson 2002) and model averaging (Burnham and Anderson 2002) is available in Program MARK to provide estimates of N with standard error that reflect model selection uncertainty.     

Determining prey availability for rock lobsters, Jasus lalandii : diver-sampling versus monochrome video photography

Accurate determination of diet is important in ecological studies. Many macrobenthic predators fragment their prey so finely that identification of component organisms is difficult. Knowledge of the food available to masticatory predators such as lobsters can help in determining potential prey and enhance the accuracy of dietary assessments. With SCUBA divers limited by depth and submersion time, benthic analyses are becoming increasingly reliant on camera systems. Here, based on paired samples, we assess the relative merits of using removal-sampling by divers versus monochrome video photography to determine the prey spectrum available to macrobenthic predators. We also relate the results to the gut contents of rock lobsters (Jasus lalandii) that were collected simultaneously. Diver samples took on average four times longer to collect and process than video-image samples. No significant differences were evident between the number of species identified in diver samples and video-image samples (n = 21 paired samples, Student's t = 0.233, P > 0.1), although an analysis of similarity (ANOSIM) test revealed that the results obtained from the two sampling methods were significantly different in terms of species composition (global R = 0.203, P < 0.01). Cluster analysis based on Bray–Curtis similarity to compare diver and video-image samples revealed four clusters of samples with <35% similarity, confirming that the two techniques do sample different assemblages of species. Video images detected and allowed identification of all species of prey recorded in the rock lobsters' stomach contents, probably because rock lobsters prey mainly on relatively large prey which are readily detected by video. Diver-sampling underestimated or failed to detect two important prey types, namely small barnacles and encrusting coralline algae. We concluded that the camera system employed was adequate for assessing the prey assemblages available to macrobenthic predators such as lobsters, and that it was considerably more economical than using divers. Received: 10 September 1998 / Accepted: 6 May 1999     

Inbreeding in a lek-mating ant species, Pogonomyrmex occidentalis

In this paper we have two goals. First, we examine the effects of sample size on the statistical power to detect a given amount of inbreeding in social insect populations. The statistical power to detect a given level of inbreeding is largely a function of the number of colonies sampled. We explore two sampling schemes, one in which a single individual per colony is sampled for different sample sizes and a second sampling scheme in which constant sampling effort is maintained (the product of the number of colonies and the number of workers per colony is constant). We find that adding additional workers to a sample from a colony makes it easier to detect inbreeding in samples from given number of colonies; however, adding more colonies rather than more workers per colony always gives greater power to detect inbreeding. Because even relatively large amounts of sib-mating generate relatively small inbreeding coefficients, detection of even substantial deviations from random mating will require very large samples. Second, we look at the amount of inbreeding in a large population of the western harvest ant, Pogonomyrmex occidentalis. We find deviations from Hardy-Weinberg equilibrium equivalent to approximately 27% sib-mating in our population ( f = 0.09). Review of past studies on the population structure of other Pogonomyrmex species suggests that inbreeding may be a regular feature of the mating system of these ants. Although P. occidentalisis a swarm-mating species, there are a number of features of its population biology which suggest that the effective population size may be small. These include topographical variation that potentially breaks the population into demes, variation in the reproductive output of colonies, and variation in the size of reproductives produced by colonies. Received: 6 May 1996 / Accepted after revision: 6 October 1996     

Population ecology of the green/black turtle (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) in Bahía Magdalena,Mexico

The mangrove channels of Bahía Magdalena, Mexico, are important developmental areas for juvenile green, or black turtles (Chelonia mydas), but incidental bycatch and illegal hunting threaten population persistence. We studied size distribution, condition index (CI), growth rates, and mortality of black turtles in Estero Banderitas, the largest mangrove channel in Bahía Magdalena, to supply information for the development of effective conservation strategies. A total of 213 black turtles (including 88 recaptures) were caught in entanglement nets between July 2000 and July 2003. Average yearly catch per unit of effort (CPUE, 1 unit: 100 m of net fishing for 12 h) dropped during the study from 2.19 to 0.76. About 97% of all turtles were considered juveniles, average size was 54.6 ± 9.5 cm. Turtles were significantly smaller at the head of Estero Banderitas than in the central part of the Estero and in the open bay, indicating size-based habitat segregation. Average growth rate was 1.62 cm/year and declined with increasing size. Growth was seasonal and three times higher in summer (0.28 cm/month) than in winter (0.09 cm/month), body CI was also significantly higher during the summer months. A seasonalized von Bertalanffy growth function (VBGF) was used to model growth for the size range studied (43–73 cm SCL), with the parameters: L _∞ = 101 cm SCL; K = 0.04 year⁻¹; t ₀ = 0; C = 0.4 and t _s = 0.75. Growth data indicate that black turtles may spend up to 20 years in Bahía Magdalena before they reach maturity at about 77 cm SCL. The total mortality estimate (Z) from the length converted catch curve was 0.16, corresponding to a yearly survival probability of 0.85.     

The power of simulating experiments

Addressing complex ecological research questions often requires complex empirical experiments. However, due to the logistic constraints of empirical studies there is a trade-off between the complexity of experimental designs and sample size. Here, we explore if the simulation of complex ecological experiments including stochasticity-induced variation can aid in alleviating the sample size limitation of empirical studies. One area where sample size limitations constrain empirical approaches is in studies of the above- and belowground controls of trophic structure. Based on a rule- and individual-based simulation model on the effect of above- and belowground herbivores and their enemies on plant biomass, we evaluate the reliability of biomass estimates, the probability of experimental failure in terms of missing values, and the statistical power of biomass comparisons for a range of sample sizes. As expected, we observed superior performance of setups with sample sizes typical of simulations (n = 1000) as compared to empirical experiments (n = 10). At low sample sizes, simulated standard errors were smaller than expected from statistical theory, indicating that stochastic simulation models may be required in those cases where it is not possible to perform pilot studies for determining sample sizes. To avoid experimental failure, a sample size of n = 30 was required. In conclusion, we propose that the standard tool box of any ecologist should comprise a combination of simulation and empirical approaches to benefit from the realism of empirical experiments as well as the statistical power of simulations.     

Interannual, seasonal, local and body size variations in reproduction of the prawn Penaeus (Marsupenaeus) japonicus (Crustacea: Decapoda: Penaeidae) in the Ariake Sea and Tachibana Bay, Japan

 The reproductive biology of female Penaeus japonicus Bate was investigated in the Ariake Sea and Tachibana Bay (located outside the Ariake Sea), Japan from 1994 to 1996. Interannual, seasonal, individual female body size and spatial influences on the incidence of spawning were examined. The proportion of inseminated females, increased with increasing body size up to 170 mm body length (BL) and decreased thereafter. The minimum size at maturity was similar between years; 130 to 140 mm BL. The minimum size of individuals with developing ovaries or spermatophores differed from that of ripe females by 20 to 25 mm. Spawning occurred in the central part of the Ariake Sea and Tachibana Bay but rarely in the inner part. P. japonicus had a clear reproductive cycle. Spawning started earlier and ended later, occurring from mid-May to mid-October, in the eastern central part of and outside of the Ariake Sea compared with the western central part, where spawning occurred from mid-June to mid-September. Spawning dynamics differed across seasons, body sizes and areas but not across years. The seasonal peak in the proportion of ripe females varied with body size. In small individuals (130 to 169 mm BL), no peak was observed, whereas in large individuals (>170 mm BL) the proportion of ripe individuals peaked in June. The proportion of ripe individuals increased with increasing body size and was high outside the Ariake Sea. Factors causing the variation in spawning dynamics are discussed. Received: 30 November 1998 / Accepted: 12 October 1999     

Scarred limpets at hydrothermal vents: evidence of predation by deep-sea whelks

The chaotic physical and chemical environment at deep-sea hydrothermal vents has been associated with an ecosystem with few predators, arguably allowing the habitat to provide refuge for vulnerable species. The dominance of endemic limpets with thin, open-coiled shells at north Pacific vents may support this view. To test their vulnerability to predation, the incidence of healed repair scars, which are argued to reflect non-lethal encounters with predators, were examined on the shells of over 5,800 vent limpets of Lepetodrilus fucensis McLean (1988) that were collected from 13 to 18 August 1996. Three vent fields on the Juan de Fuca Ridge at ca. 2,200 m depth were sampled, two within 70 m of 47°56.87′N 129°05.91′W, and one at 47°57.85′N 129°05.15′W with the conspicuous potential limpet predators, the zoarcid fish Pachycara gymninium Anderson and Peden (1988), the galatheid crab Munidopsis alvisca Williams (1988), and the buccinid snail Buccinum thermophilum Harasewych and Kantor (2002). Limpets from the predator-rich vent were most often scarred, a significant difference created by the high incidence of scars on small (<4 mm long) limpets in this sample. Collected with the limpets were small (median shell diameter 4.4 mm) buccinids. They, rather than the larger, more conspicuous mobile fishes and crabs are argued to be the shell-damaging predator. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Mechanistic understanding of human–wildlife conflict through a novel application of dynamic occupancy models

Crop and livestock depredation by wildlife is a primary driver of human–wildlife conflict, a problem that threatens the coexistence of people and wildlife globally. Understanding mechanisms that underlie depredation patterns holds the key to mitigating conflicts across time and space. However, most studies do not consider imperfect detection and reporting of conflicts, which may lead to incorrect inference regarding its spatiotemporal drivers. We applied dynamic occupancy models to elephant crop depredation data from India between 2005 and 2011 to estimate crop depredation occurrence and model its underlying dynamics as a function of spatiotemporal covariates while accounting for imperfect detection of conflicts. The probability of detecting conflicts was consistently <1.0 and was negatively influenced by distance to roads and elevation gradient, averaging 0.08–0.56 across primary periods (distinct agricultural seasons within each year). The probability of crop depredation occurrence ranged from 0.29 (SE 0.09) to 0.96 (SE 0.04). The probability that sites raided by elephants in primary period t would not be raided in primary period t + 1 varied with elevation gradient in different seasons and was influenced negatively by mean rainfall and village density and positively by distance to forests. Negative effects of rainfall variation and distance to forests best explained variation in the probability that sites not raided by elephants in primary period t would be raided in primary period t + 1. With our novel application of occupancy models, we teased apart the spatiotemporal drivers of conflicts from factors that influence how they are observed, thereby allowing more reliable inference on mechanisms underlying observed conflict patterns. We found that factors associated with increased crop accessibility and availability (e.g., distance to forests and rainfall patterns) were key drivers of elephant crop depredation dynamics. Such an understanding is essential for rigorous prediction of future conflicts, a critical requirement for effective conflict management in the context of increasing human–wildlife interactions.     

The influence of group size on natal dispersal in the communally rearing and semifossorial rodent, Octodon degus

In social or group living species, members of groups are expected to be affected differentially by competition through the effect of group size (i.e., the “social competition hypothesis”). This hypothesis predicts an increase in the probability of dispersal with increasing size of social groups. At a more mechanistic level and based on the known effects of competition on stress hormone levels, a positive relationship between group size and glucocorticoids of juveniles should be observed. We used a demographic approach to test these predictions on a natural population of the communally rearing and semifossorial rodent—Octodon degus. Burrow systems provide degus with places to rear offspring and to evade stressful thermal conditions and predators. Thus, we predicted dispersal to increase with increasing number of degus per main burrow system used, a measure of habitat saturation in degus. The probability of dispersal increased with increasing number of degus per main burrow system used. Mean fecal metabolites of cortisol in offspring increased, yet not statistically significantly, with the number of juveniles in groups. These results were consistent with a scenario in which competition drives natal dispersal in juveniles in social degus. In particular, competition would be the consequence of high degu abundance in relation to the abundance of burrow systems available at the time of offspring emergence.     

Organics in wheat and rye straw pulp bleaching and combined mill effluents (II) ecotoxicological screening

Potential environmental effects of the discharge of industrial wastewater from manufacturing of bleached straw pulp have been studied. Bleached neutral sulphite straw pulping results in discharges of both black cooking and spent bleaching liquors. Time proportional samples were taken from the combined bleaching effluent and the combined mill effluent, (i) Mutagenic activity of the two samples in an Ames’ test, and (ii) acute toxicity of the combined mill effluent sample towards the photosynthetic activity of natural marine phytoplankton from the receiving waters were determined. Also, acute toxicity was determined of the slowly biodegradable or persistent organics remaining after aerobic stabilization of the sample towards (ii) photosynthetic activity of natural phytoplankton, (iii) mussels (Mytilus edulis), (iv) eels (Anguilla anguilla) and, (v) crustacean (Nitocra spinipes); and furthermore, (vi) reproduction test of the crustacean and (vii) growth inhibition test of the marine diatom (Phaeodactylum tricornutum) were performed. Mutagenicity emission factors (MEF's) and toxicity emission factors (TEF's) were calculated to classify the effluents. Combined mill effluent sample showed a mutagenic activity per t₉₀ of one fourth the activity of a kraft pulp bleaching effluent, and the inhibition of the photosynthesis of natural phytoplankton was significant up to 2600 times dilution. After aerobic stabilization, inhibition was observed up to 400 times dilution. Other effects of the combined mill effluent sample were only significant below 40 times dilution. The bleaching effluent showed a mutagenic activity per t₉₀ comparable to kraft bleaching effluents. Other effects were only significant below 20 times dilution.     

UV absorption by mycosporine-like amino acids in Phaeocystis antarctica Karsten induced by photosynthetically available radiation

Mycosporine-like amino acids (MAAs), which occur in diverse taxonomic groups, exhibit in vivo absorption maxima between 310 nm and 360 nm and may play a photoprotective role against ultraviolet (UV) exposure. Using cultures of colonial Phaeocystis antarctica, we examined the relationship between MAA concentration, in vivo UV absorption, photoprotective (carotenoid) and photosynthetic pigments, and photosynthetically available radiation (PAR, 350–700 nm). UV absorption was high; chlorophyll-specific absorption, a ^* _ph, at 330 nm ranged from 0.06 to 0.41 m²/mg chlorophyll a. Values of a ^* _ph (330) were 4–13 times greater than a ^* _ph (676). Mycosporine-glycine, shinorine, and mycosporine-glycine valine are responsible for the strong in vivo UV absorption. The sum of all MAAs increased with irradiance when normalized to chlorophyll a or carbon concentrations, whereas individual MAAs varied independently from each other. Mycosporine-glycine concentrations showed no statistically significant change over the range of light intensities, whereas mycosporine-glycine and shinorine concentrations increased at higher irradiances. The relative fluorescence yield for chlorophyll a was low in the UV region compared to the visible region, implying that absorbed UV radiation (<375 nm) is transferred inefficiently to chlorophyll a in the reaction center. Quantitative estimates of UV screening by MAAs are attributed to elevated MAA concentrations and increased diameter at high light. Received: 31 March 1999 / Accepted: 13 July 2000     

A game model for dominance relations among group-living animals

We present here an attempt to understand behaviors of dominant individuals and of subordinate individuals as behavior strategies in an asymmetric “hawk-dove” game. We assume that contestants have perfect information about relative fighting ability and the value of the resource. Any type of asymmetry, both relevant to and irrelevant to the fighting ability, can be considered. It is concluded that evolutionarily stable strategies (ESSs) depend on the resource value (V), the cost of injury (D), and the probability that the individual in one role will win (x). Different ESSs can exist even when values of V, D, and x are the same. The characteristics of dominance relations detected by observers may result from the ESSs that the individuals are adopting. The model explains some characteristics of dominance relations, for example, the consistent outcome of contests, the rare occurrence of escalated fights, and the discrepancy between resource holding potential (RHP) and dominance relations, from the viewpoint of individual selection. Received: 7 May 1997 / Accepted after revision: 17 October 1997     

Individual dispersal status influences space use of conspecific residents in the common lizard, Lacerta vivipara

The effects of immigration on the behaviour of residents may have important implications for the local population characteristics. A manipulative laboratory experiment with yearlings of the common lizard (Lacerta vivipara) was performed to test whether the introduction of dispersing or philopatric individuals influences the short-term spacing behaviour of resident individuals. Staged encounters were carried out to induce interactions within dyads. The home cage of each responding individual was connected by a corridor to an unfamiliar “arrival cage” to measure the latency to leave their own home cage after each encounter. Our results showed that the time that pairs spent in close proximity was longer when a dispersing individual was introduced in the home cage. The latency to leave the home cage was longer after the introduction of a dispersing individual. These response variables were not influenced by the relative body sizes of contestants nor by the level of aggression towards each other. In contrast, the aggressive response was significantly influenced by the residency asymmetry established experimentally (“owner” of the home cage vs introduced individual). Our results suggest that the space use by resident individuals is influenced by the dispersal status of conspecifics. The potential ultimate causes driving this effect are discussed.     

Bayesian analysis of abundance for binomial sighting data with unknown number of marked individuals

The mark-resight method for estimating the size of a closed population can in many circumstances be a less expensive and less invasive alternative to traditional mark-recapture. Despite its potential advantages, one major drawback of traditional mark-resight methodology is that the number of marked individuals in the population available for resighting needs to be known exactly. In real field studies, this can be quite difficult to accomplish. Here we develop a Bayesian model for estimating abundance when sighting data are acquired from distinct sampling occasions without replacement, but the exact number of marked individuals is unknown. By first augmenting the data with some fixed number of individuals comprising a marked “super population,” the problem may then be reformulated in terms of estimating the proportion of this marked super population that was actually available for resighting. This then allows the data for the marked population available for resighting to be modeled as random realizations from a binomial logit-normal distribution. We demonstrate the use of our model to estimate the New Zealand robin (Petroica australis) population size in a region of Fiordland National Park, New Zealand. We then evaluate the performance of the proposed model relative to other estimators via a series of simulation experiments. We generally found our model to have advantages over other models when sample sizes are smaller with individually heterogeneous resighting probabilities. Due to limited budgets and the inherent variability between individuals, this is a common occurrence in mark-resight population studies. WinBUGS and R code to carry out these analyses is available from .