Skeletochronological assessment of age structure and population stability for two threatened frog species

Populations that fluctuate in size may become small at particular times and therefore be at risk of extinction. I used skeletochronology to examine the potential for fluctuations in population size in the threatened frog species Geocrinia alba and Geocrinia vitellina from southwestern Australia. The change in number of lines of arrested growth (LAG) in toe bones of recaptured frogs indicated that most individuals probably gain one LAG annually. Therefore, skeletochronology can be used for age determination. From a combination of skeletochronology and a mark–recapture study, both species appear to live for a maximum of six years. However, the majority of adult males only bred in a single year, which gave rise to a consistently dominant three-year-old age class. A drop in the number of two-year-old frogs and a large increase in the number of three-year-olds (up to 85%) from 1993 to 1994 in both species highlighted the potential for substantial fluctuations in population size. Widespread changes in abundance of between 25 and 50% may therefore be expected over periods as short as one or two years, due to variation in recruitment success.     

Demographic sensitivity and persistence of the threatened white- and orange-bellied frogs of Western Australia

Anurans have been the subject of numerous experimental and monitoring studies, yet their autecology under natural conditions remains poorly understood. Previous work has focussed on the pre-metamorphic life stages, with data suitable for estimating demographic parameters for all life stages collected in few species. This deficiency has almost certainly confounded current views of amphibian autecology and understanding of which mechanisms of decline represent the greatest threats to the persistence of amphibian species. We used a stage-structured metapopulation model, parameterised with comprehensive field data, to explore the sensitivity of two species of myobatrachid frogs from the southwest of Australia, Geocrinia alba and G. vitellina, to changes in demographic parameters at all life stages. The simulation modelling indicated that the population dynamics of both species were consistently most sensitive to changes in juvenile survival, then to fecundity, and third to both adult survival and standard deviation of survival to about equal extent. In practical terms, this indicates that management interventions which attempt to mitigate reductions in juvenile survival are likely to be most successful in arresting anuran metapopulation declines.     

Impact of fuel-reduction burning on the frog Geocrinia lutea in southwest Western Australia

Abstract The four frog species in the Geocrinia rosea complex occur in state forest in southwest Western Australia. Fire management of these forests involves fuel-reduction burning with an average rotation of five to nine years. In this study we examined the impact of fire on Geocrinia lutea by counting calling males in six pairs of burned and control sites from 1992 to 1994. The immediate impact of the fire on G. lutea adults, and the survival of G. lutea eggs and larvae after the fire, were also addressed. We found that fuel-reduction burning in spring was associated with a significant decline in the number of calling males. The populations had not recovered two years after the fire. Up to 29% of the calling males may have been killed in the fire. Egg and larval survival was not significantly different between treatments. However, the treatments did differ in the cause of death, with higher in-situ egg death and lower predation at burned sites. The short-term impact of spring fuel-reduction burns may pose a serious threat of extinction for very small populations. The endangered species Geocrinia alba has many small, isolated populations and frequent fire may therefore be inimical to their survival. However, we do not know if there is a long-term effect. Populations may or may not have time to recover between fires.     

Winter microhabitat selection of a threatened pond amphibian in constructed urban wetlands

Mitigating the threat of habitat loss requires actions such as restoring and creating new habitat. In order to effectively achieve this, species habitat requirements and use patterns need to be understood. While many studies have been conducted on the habitat choice of species, these generally focused on habitat use during periods of high activity and detection probability without considering seasonal shifts in habitat use. Understanding habitat selection by frogs during the winter season of low activity may be crucial since it may differ from that used during the summer and may be overlooked as important for population success. We describe the microhabitat use of the threatened green and golden bell frog (Litoria aurea) using radio tracking methods during winter when detection is low and knowledge is limited. We followed 26 individuals between May and July, 2011 to determine whether they selected specific overwintering microhabitats and related this to levels of individual exposure to predators, distance from the edge of the water and temperature of microhabitats. We found that overwintering bell frogs inhabited reeds and rock gabions more frequently than expected and that females used a reduced subset of microhabitats compared to males. Additionally, microhabitats used were more likely to conceal an individual from view, and the majority of overwintering sites were located within 5 m of the edge of the water which may be important for reducing the risk of predation and desiccation. Rock gabions had significantly warmer (1.2°C–1.8°C) mean temperatures than the other microhabitats used. The information presented here can be used in habitat creation and reintroduction programmes to provide habitat which is suitable during both the breeding and non‐breeding season for the conservation of other populations.     

Dispersal behaviour of individuals in metapopulations of two British butterflies

Dispersal patterns are important in metapopulation ecology because they affect the dynamics and survival of populations. However, because little empirical information exists on dispersal behaviour of individuals, theoretical models usually assume random dispersal. Recent empirical evidence, by contrast, suggests that the butterfly Maniola jurtina uses a non‐random, systematic dispersal strategy, can detect and orient towards habitat from distances of 100–150 m, and prefers a familiar habitat patch over a non‐familiar one (‘homing behaviour’). The present study (1) investigated whether these results generalise to another butterfly species, Pyronia tithonus; and (2) examined the cause of the observed ‘homing behaviour’ in M. jurtina. P. tithonus used a similar non‐random, systematic dispersal strategy to M. jurtina, had a similar perceptual range for habitat detection and preferred a familiar habitat patch over a non‐familiar one. The ‘homing behaviour’ of M. jurtina was found to be context‐dependent: individual M. jurtina translocated within habitat did not return towards their capture point, whereas individuals translocated similar distances out of habitat did return to their ‘home’ patch. We conclude that butterfly ‘homing behaviour’ is not based on an inherent preference for a familiar location, but that familiarity with an area facilitates the recognition of suitable habitat, towards which individuals orient if they find themselves in unsuitable habitat. Contrary to conventional wisdom, we suggest that frequent, short ‘excursions’ over habitat patch boundaries are evolutionarily advantageous to individuals, because increased familiarity with the surrounding environment is likely to increase the ability of a straying animal to return to its natural habitat, and to reduce the rate of mortality experienced by individuals attempting to disperse between habitat patches. We discuss the implications of the non‐random dispersal for existing metapopulation models, including models of the evolution of dispersal rates.     

Do invasive species undergo metapopulation dynamics? A case study of the invasive Caspian gull,Larus cachinnans,in Poland

Aim The mechanisms of initial dispersal and habitat occupancy by invasive alien species are fundamental ecological problems. Most tests of metapopulation theory are performed on local population systems that are stable or in decline. In the current study we were interested in the usefulness of metapopulation theory to study patch occupancy, local colonization, extinction and the abundance of the invasive Caspian gull (Larus cachinnans) in its initial invasion stages. Location Waterbodies in Poland. Methods Characteristics of the habitat patches (waterbodies, 35 in total) occupied by breeding pairs of Caspian gulls and an equal sample of randomly selected unoccupied patches were compared with t‐tests. Based on presence–absence data from 1989 to 2006 we analysed factors affecting the probability of local colonization, extinction and the size of local populations using generalized linear models. Results Occupied habitat patches were significantly larger and less isolated (from other habitat patches and other local populations) and were located closer to rivers than empty patches. The proximity of local food resources (fish ponds, refuse dumps) positively affected the occurrence of breeding pairs. The probability of colonization was positively affected by patch area, and negatively by distances to fish ponds, nearest habitat patch, nearest breeding colony and to a river, and by higher forest cover around the patch boundaries. The probability of extinction was lower in patches with a higher number of breeding pairs and with a greater area of islets. The extinction probability increased with distances to other local populations, other habitat patches, fish ponds and to refuse dumps and with a higher cover of forest around the patch boundaries. The size of the local population decreased with distances to the nearest habitat patch, local population, river, fish pond and refuse dump. Local abundance was also positively affected by the area of islets in the patch. Main conclusions During the initial stages of the invasion of Caspian gulls in Poland the species underwent metapopulation‐like dynamics with frequent extinctions from colonized habitat patches. The results prove that metapopulation theory may be a useful conceptual framework for predicting which habitats are more vulnerable to invasion.     

Patch occupancy and abundance of local populations in landscapes differing in degree of habitat fragmentation: a case study of the colonial black‐headed gull,Chroicocephalus ridibundus

Aim This study investigated whether habitat fragmentation at the landscape level influences patch occupancy and abundance of the black‐headed gull, Chroicocephalus ridibundus, and whether the response of the species to environmental factors is consistent across replicated landscape plots. Location Water bodies (habitat patches) in southern Poland. Methods Surveys were conducted in two landscape types (four plots in each): (1) more‐fragmented landscape, in which habitat patches were small (mean size 2.2–6.2 ha) and far apart (mean distance 2.5–3.1 km); and (2) less‐fragmented landscape, in which habitat patches were large (mean size 9.2–16.5 ha) and separated by short distances (mean 0.9–1.4 km). Observations were performed twice in 284 potential habitat patches during the 2007 breeding season. Results Colonies were significantly more frequent and larger in the less‐fragmented landscapes than in the more‐fragmented ones. Probability of patch occupancy and number of breeding birds were positively related with patch size and these relationships were especially strong in the more‐fragmented landscapes. In the less‐fragmented landscapes, the occurrence of black‐headed gulls was negatively related to the distance to the nearest local population, but in the more‐fragmented landscapes such a relationship was not detected. As distance to the nearest habitat patch increased, the probability of the patch occupancy decreased in the more‐fragmented landscapes. Moreover, abundance was negatively influenced by distance to the nearest habitat patch, especially strongly in more‐fragmented landscapes. Proximity of corridors (rivers) positively influenced the occupation of patches regardless of landscape type. The number of islets positively influenced occupancy and abundance of local populations, and this relationship was stronger in the more‐fragmented landscapes. Main conclusions Our results are in agreement with predictions from metapopulation theory and are the first evidence that populations of black‐headed gulls may have a metapopulation structure. However, patch occupancy and abundance were differentially affected by explanatory variables in the more‐fragmented landscapes than in the less‐fragmented ones. This implies that it is impossible to derive, a priori, predictions about presence/abundance patterns based on only a single landscape.     

Reproductive strategy and carry-over effects for species with complex life histories

We investigated the influences of natal-pool and metamorph characteristics on juvenile survival, age-specific breeding probabilities, and dispersal of wood frogs (Lithobates sylvatica) and used this information to infer how life history strategies of short-lived species may offset risks associated with breeding in highly variable habitats. We used multistate mark-recapture data from eight wood frog populations in Maryland, USA, to investigate the influences of natal-pond and metamorph characteristics on post-metamorphic demographics. We found that post-metamorphic juvenile survival was highly variable and negatively influenced by abiotic conditions experienced during development but showed little relationship to larval density or size at metamorphosis. Estimates of recruitment and dispersal probabilities indicated that males mature earlier than females, and a small percentage of each sex disperse to non-natal pools. Survival probabilities for adults during the non-breeding season were less variable than juvenile rates, lower for females, and negatively related to mean monthly precipitation. Survival of adults during the breeding season was generally very high. We provide the first robust estimates of post-metamorphic vital rates of wood frogs that allow for variation in capture probabilities. We found little evidence for an effect of metamorph traits on juvenile survival, suggesting that wood frogs may be able to overcome initial disadvantages to have similar post-metamorphic performance. Our study suggests that variation in the age of maturity for wood frogs may mitigate risks associated with breeding in a highly variable habitat to maximize lifetime fitness without increasing lifespan, and this strategy is minimally affected by carry-over effects from the larval stage.     

Dispersal and connectivity in metapopulations

This paper reviews characteristics of dispersal that influence metapopulation functioning, such as releasing factors, density dependence, timing and types and health of dispersers. Economic thresholds, intraspecific conflicts and avoidance of inbreeding arc often regarded as the key ultimate or proximate (or both) causes of dispersal, but there is no consensus about the most important mechanisms. Dispersing individuals arc often considered to differ genetically from the residents but good supporting evidence has only been presented for some insect species. Sex and age differences in dispersal rates are most common in polygamous species and in long-lived species with many litters per female. A bimodal distribution of dispersal distances, earlier thought to be a common pattern, is probably an artifact, caused by habitat heterogeneity and varying survival of settled individuals. Dispersal distances are longer in poor environments. Habitat specialists are more affected by boundaries during dispersal than generalists. Dispersal just before or during the early reproductive season is common in certain species occupying early successional habitats. Dispersal increased both population and metapopulation size and persistence in plants, insects and small mammals.     

Dispersal, interpatch movements, and survival in a shrubland breeding bird community

ABSTRACT Dispersal events can affect the distribution, abundance, population structure, and gene flow of animal populations, but little is known about long‐distance movements due to the difficulty of tracking individuals across space. We documented the natal and breeding dispersal of shrubland birds among 13 study sites in a 1000 km² area in southeastern Ohio. In addition, we radio‐marked and tracked 37 adult males of one shrubland specialist, the Yellow‐breasted Chat (Icteria virens). We banded 1925 juveniles and 2112 adults of nine shrubland species from 2002 to 2005. Of these, 33 (1.7%) juveniles were encountered in subsequent years (2003–2006) as adults (natal dispersal) and 442 (20.9%) birds initially banded as breeding adults were re‐encountered in subsequent years (breeding dispersal). Apparent survival of juvenile shrubland birds on their natal patches was 0.024 (95% CI 0.016–0.036). After accounting for the probability of detection, we found that 21% of birds banded as juveniles and recaptured as adults returned to their natal patches, whereas 78% of adult birds showed fidelity to the patch where they were originally captured. Moreover, natal dispersers tended to move farther than breeding dispersers (corrected natal median = 1.7 km ± 0.37; corrected breeding median = 0.23 km ± 0.10). We used our estimates of natal dispersal and annual apparent survival to estimate true survival at 0.11 (95% CI 0.07–0.18) for juveniles in their first year. However, this estimate was only applicable for birds dispersing within 7 km of their natal patches. Interpatch movements of radio‐marked Yellow‐breasted Chats were not uncommon, with 13 of 37 males located in more than one habitat patch. Overall, we observed low natal philopatry, but high adult site fidelity for shrubland birds in our study area. Considering the frequency of short‐distance movements observed (median = 531 m, range = 88–1045 m), clustering of patches within 1 km might facilitate use of shrubland habitat.     

Demographic consequences of population subdivision on the long-furred woolly mouse opossum (Micoureus paraguayanus) from the Atlantic Forest

Habitat destruction and fragmentation severely affected the Atlantic Forest. Formerly contiguous populations may become subdivided into a larger number of smaller populations, threatening their long-term persistence. The computer package VORTEX was used to simulate the consequences of habitat fragmentation and population subdivision on Micoureus paraguayanus, an endemic arboreal marsupial of the Atlantic Forest. Scenarios simulated hypothetical populations of 100 and 2000 animals being partitioned into 1–10 populations, linked by varying rates of inter-patch dispersal, and also evaluated male-biased dispersal. Results demonstrated that a single population was more stable than an ensemble of populations of equal size, irrespective of dispersal rate. Small populations (10–20 individuals) exhibited high instability due to demographic stochasticity, and were characterized by high rates of extinction, smaller values for metapopulation growth and larger fluctuations in population size and growth rate. Dispersal effects on metapopulation persistence were related to the size of the populations and to the sexes that were capable of dispersing. Male-biased dispersal had no noticeable effects on metapopulation extinction dynamics, whereas scenarios modelling dispersal by both sexes positively affected metapopulation dynamics through higher growth rates, smaller fluctuations in growth rate, larger final metapopulation sizes and lower probabilities of extinction. The present study highlights the complex relationships between metapopulation size, population subdivision, habitat fragmentation, rate of inter-patch dispersal and sex-biased dispersal and indicates the importance of gaining a better understanding of dispersal and its interactions with correlations between disturbance events.     

Consequences of dispersal limitation and habitat fragmentation for the Brazilian heart‐tongued frogs (Phyllodytes spp.)

Poorly known species may be cryptically endangered, especially when they inhabit fragmented and threatened habitats. Heart‐tongued frogs (genus Phyllodytes, family Hylidae, Lophyohylinae) comprise 17 species of poorly known frogs that have obligatory associations with tank bromeliads. The distributions of all species are restricted to a small, extremely fragmented, region of Atlantic Forest in eastern Brazil. We model climate and tank bromeliad distributions to better understand frog distribution limits. Using records from several sources for frogs and bromeliads with climate data from WorldClim, we modelled the distribution of Phyllodytes using maximum entropy. We compared climate and altitude within the distribution and nearby to test how climate may limit distribution. Climate together with bromeliad distributions provided the best model and predicted the smallest suitable area for Phyllodytes that was larger than that occupied, from the state of Paraíba in the north to Rio Grande do Sul in the south. Phyllodytes occurs in lower elevations that are warmer, wetter and less variable than the surrounding regions where it does not occur, and dispersal is apparently limited by the surrounding, inhospitable, region. Dispersal limitation and habitat fragmentation have relegated Phyllodytes to many very small habitat fragments. With many species in this genus being known from a single or few samples, this unfortunate combination of limitation and fragmentation suggests that some or all species of Phyllodytes may be threatened with extinction, especially if habitat fragmentation continues at its present pace in eastern Brazil.     

DISPERSAL,GENE FLOW,AND ALLELIC DIVERSITY BETWEEN LOCAL POPULATIONS OF THOMOMYS BOTTAE POCKET GOPHERS IN THE COASTAL RANGES OF CALIFORNIA

We studied Thomomys bottae pocket gophers from 1979 to 1982 to determine if the amount of gene flow between local populations was sufficient to reduce allele frequency differences between them. Dispersal was quantified using three different trap techniques, and genetic changes in the population were monitored using protein variants. Additional allele frequency data were available for 1976 and 1977. We observed dispersal to be common in pre-reproductive juvenile females throughout the breeding season of their birth. Males on the other hand tended to disperse only from the end of the breeding season. Although dispersal was common, 63% of adults appeared to be recruited within 40 m of where they were born. Gene flow occurred into both established populations and into vacant habitat, but it was too low to reduce the differences in gene frequencies between the fields over seven years. We conclude that allelic diversity in T. bottae populations is a balance between random drift due to small effective population size and gene flow.     

Dispersal, distribution, patch network and metapopulation dynamics of the dingy skipper butterfly (Erynnis tages)

Two general approaches have usually been taken towards understanding the distributions and dynamics of localised species in heterogeneous landscapes, namely habitat characterisation and metapopulation dynamics. We show how habitat and metapopulation dynamics interact to generate a highly localised distribution of a butterfly, despite the extremely widespread nature of the butterfly’s host plant. Egg placement, macro-habitat requirements and dispersal were studied for the butterfly Erynnis tages, in North Wales, where it shows a restricted distribution relative to that of its host plant, Lotus corniculatus. Females laid eggs disproportionately on large plants growing in hollows, with intermediate cover of bare ground and high cover of L. corniculatus. Ideal macro-habitat, studied at 100-m grid resolution, consisted of areas with high host plant densities, sheltered from wind, with light or no grazing or cutting. These specialised conditions are represented as localised patches in the landscape, and define the potential habitat network, within which metapopulation dynamics take place. Although there was a moderate (22%) level of exchange of individual E. tages among local populations, the total number of potential colonists in the whole system was low because source population sizes were small (≤200 individuals at peak in any site in 1997 and 1998). Four unoccupied but apparently suitable 500-m grid squares were colonised between 1997 and 1998, and isolated habitat was less likely to be occupied. Overall, our study suggests that long-term regional persistence of E. tages is very likely to depend on metapopulation processes within the restricted patch network, rather than on the long-term survival of local populations. Received: 25 May 1999 / Accepted: 9 August 1999     

Effects of environmental factors on the ecology and survival of a widespread,endemic Cerrado frog

Understanding the mechanisms that affect habitat use by vertebrates is critical for understanding how species are distributed across landscapes and how they cope with habitat change. The Brazilian Savanna (the Cerrado) has vegetation ranging from grassland to woodland savannas and harbors a rich and diverse amphibian fauna impacted by accelerated habitat loss. Here, we test the influence of vegetation type (from grassy scrubland to woodland) and distance from breeding sites (ephemeral water bodies) on body size, abundance, and survival of the frog Physalaemus nattereri in a natural metapopulation system of south-central Brazil. We also test whether body size is a significant predictor of population abundance. We found that the abundance of P. nattereri varies according to the mean snout–vent length of each metapopulation (sampling unit), as well as a higher estimated mortality rate in woodlands compared with typical Cerrado. Furthermore, we found no difference in estimated mortality among sampling units located far or close to ephemeral water bodies. Thus, our results highlight variable responses of P. nattereri metapopulations to environmental factors, despite the observed high heterogeneity among sampled habitats and the importance of ephemeral water bodies for reproduction. These findings highlight that land cover and availability of breeding sites might not always interact to explain population persistence of Cerrado frogs.     

Potential metapopulation structure and the effects of habitat quality on population size of the endangered False Ringlet butterfly

The False Ringlet (Coenonympha oedippus) is a European butterfly species, endangered due to the severe loss and fragmentation of its habitat. In Hungary, two remaining populations of the butterfly occur in lowland Purple Moorgrass meadows. We studied a metapopulation occupying twelve habitat patches in Central Hungary. Our aim was to reveal what measures of habitat quality affect population size and density of this metapopulation, estimate dispersal parameters and describe phenology of subpopulations. Local population sizes and dispersal parameters were estimated from an extensive mark–release–recapture dataset, while habitat quality was characterized by groundwater level, cover of grass tussocks, bush cover, height of vegetation and grass litter at each habitat patch. The estimated size of the metapopulation was more than 3,000 individuals. We estimated a low dispersal capacity, especially for females, indicating a very low probability of (re)colonization. Butterfly abundance and density in local populations increased with higher grass litter, lower groundwater level and larger area covered by tussocks. We suppose that these environmental factors affect butterfly abundance by determining the microclimatic conditions for both larvae and adult butterflies. Our results suggest that the long-term preservation of the studied metapopulation needs the maintenance of high quality habitat patches by appropriate mowing regime and water regulation. Management also should facilitate dispersal to strengthen metapopulation structure with creating stepping-stones or gradually increase habitat quality in present matrix.     

Dynamic occupancy models reveal within‐breeding season movement up a habitat quality gradient by a migratory songbird

The timing of settlement decisions likely influences the quality of breeding site choices.This is particularly the case in migratory birds, because the conditions that enhance breeding success are often not apparent upon arrival after migration. A strategy that addresses this problem is to adjust settlement decisions when reliable information becomes available. We used a new indirect method – dynamic site occupancy modeling – to estimate apparent movement of black‐throated blue warblers Dendroica caerulescens among sites within a breeding season. Because individuals should disperse to sites that maximize their fitness, we hypothesized that warblers would move up a habitat quality gradient when opportunities arose. For our study species, that would involve moving into sites with greater shrub density and at higher elevation within northern hardwoods forest, as these two features are positively correlated with reproduction and apparent survival in this species. Although the probability of site occupancy in our study landscape remained consistent throughout the breeding season (range: 0.66–0.69), occupancy models revealed substantial support for apparent movement of individuals within the breeding season. The mean probability of emigration from a point count site was 0.21 (±0.03 SE), and the mean probability of immigration to a site not previously occupied was 0.51 (±0.05 SE). The spatial distribution of this movement was a function of habitat quality. A portion of the black‐throated blue warbler population appears to arrive on the breeding grounds and settle initially in sub‐optimal habitat, moving subsequently into high quality densely shrubbed habitat at higher elevations. This modeling approach provides a new means to test hypotheses about habitat selection and movement by using presence–non‐detection data.     

Migration of moth species in a network of small islands

Rapidly increasing fragmentation of natural landscapes decreases the ability of many species to reach the smaller and more isolated patches of habitat in a metapopulation. The densities of local populations of several moth species and the butterfly Hipparchia semele in a network of small islands, and the rates of inter-island movement and movement patterns, were investigated, to determine the factors affecting the rate and pattern of movements. The estimated population densities ranged from 0.001 to 0.2 individuals/m². The observed emigration and immigration rates depended on island isolation and various traits of the species, with great variability in migration rates among species. Thin-bodied, slow-flying species did not move among the islands, whereas many robust, fast-flying species moved among the islands relatively frequently. Migration rate increased significantly with body size and was significantly higher in oligophagous than in polyphagous species, suggesting that these factors are important determinants of the migration rate of the species. Migration rate was low when the surface temperature of the sea was low, and a greater proportion of individuals emigrated from small than large patches of habitat. The migration distances of female noctuids were shorter than those of males and those of both sexes of the butterfly H. semele. The observed movement patterns are consistent with a metapopulation structure in most of the moth species.     

Year-round local movements of the japanese fluvial sculpin,Cottus pollux (large egg type), with special reference to the distribution of spawning nests

Year-round local movements of adult Japanese fluvial sculpin.Cottus pollux (large egg type), were investigated by a capture-mark-and-recapture method from July 1989 to July 1990 in the upper reaches of the Inabe River, central Japan. In the pre-breeding (July to January) and post-breeding seasons (June to July), the mean distance of movements in males and females was less than 20 m, and there was no apparent tendency to move into a particular channel-unit habitat, suggesting residential tendency in both sexes. In the breeding season (February to May), males tended to move into the raceways where most of the spawning nests were found, but females did not do likewise, indicating a sexual difference in movements in that season. Such sexual difference in movements was also confirmed by the records of individuals captured and recaptured more than three times: six (42.9%) of 14 males moved into the raceways in the breeding season, whereas only one (7.1%) of 14 females did so in that season. The reason behind this sexual difference in movements observed in the breeding season is discussed from the viewpoint of the patterns of spatial distribution between sexes during the pre-breeding and breeding seasons.