Introgressive hybridization between Dolly Varden Salvelinus malma and white‐spotted charr Salvelinus leucomaenis on Hokkaido Island, Japan

Intra‐ and interspecific phylogenetic analysis of Dolly Varden Salvelinus malma and white‐spotted charr Salvelinus leucomaenis throughout Hokkaido Island was conducted using nucleotide sequences of the mitochondrial DNA control region and three nuclear DNA markers [ribosomal DNA (ITS‐1), microsatellite ( u ‐85), SINE ( Fok ‐223)]. A total of 19 mtDNA haplotypes from 271 individuals of Dolly Varden, nine haplotypes from 134 white‐spotted charr were defined, and identified two well‐supported monophyletic clades for each species. Two haplotypes of Dolly Varden, however, were clustered together in the white‐spotted charr clade. Those haplotypes were found only from several rivers in the Shiretoko Peninsula, and a neighbouring river of that region. Analyses of ribosomal DNA and Fok ‐223 loci revealed that both species are characterized by having completely species‐specific diagnostic sequence and fragment patterns. Dolly Varden of the Shiretoko populations are typically allopatric in distribution where white‐spotted charr do not currently occur. In addition, incongruence in genetic relationships between mtDNA and nuclear DNA markers give strong evidence of historical mtDNA introgression between Dolly Varden and white‐spotted charr. Some white‐spotted charr diagnostic alleles in a nuclear microsatellite locus ( u ‐85) were found in some Shiretoko Dolly Varden populations, suggesting that introgressive hybridization might have also occurred in relatively recent contact and potentially ongoing evolutionary event. The present study presents an example of historical hybridization and introgression at the southernmost distribution limits of Dolly Varden.     

Variation in migratory history of Dolly Varden in a stream with an artificial dam in the Shiretoko Peninsula,Hokkaido, Japan

The variation of migration patterns in Dolly Varden Salvelinus malma in a stream with an artificial dam (erosion-control-dam) in the Shiretoko Peninsula, Hokkaido Island, Japan, at the southernmost part of its distribution, was examined by otolith Sr:Ca ratio analysis. All specimens from the above-dam area showed consistently low Sr:Ca ratios (<5.0 × 10⁻³) throughout the otolith. In contrast, many specimens from the below-dam area had higher ratios (>5.0 × 10⁻³) throughout the otolith or only at the outer part than do those from the above-dam area. This higher ratio is probably due to salinity effects. Specimen from the below-dam area could be divided into three types: (1) Consistently low Sr:Ca ratios throughout the otolith, being freshwater residents. (2) Low Sr:Ca ratios around the inner part of the otolith, and thereafter higher ratios toward the outer part, being typically anadromous. (3) Relatively higher Sr:Ca ratios throughout many or in some parts of the otolith, indicating that these have migrated from the freshwater to brackish water or seawater from the early life stage. These findings suggest that Dolly Varden has a high degree of plasticity in its migratory behavior.     

Metapopulation structure of stream-dwelling Dolly Varden charr inferred from patterns of occurrence in the Sorachi River basin, Hokkaido, Japan

1. We investigated the occurrence (presence or absence) of stream‐dwelling Dolly Varden (Salmonidae: Salvelinus malma) in the 78 tributaries of the Sorachi River system, Hokkaido, Japan to examine evidence for metapopulation structure. 2. We hypothesised (i) that the occurrence of Dolly Varden would be less likely in small tributaries because of the higher extinction risk and (ii) that occurrence would be less likely in isolated tributaries because of reduced rates of dispersal and colonisation. In addition to these two general predictions, we hypothesised (iii) that the occurrence would be more likely in spring‐fed tributaries, because Dolly Varden prefer cold water as present in these systems. 3. When all the tributaries were included in a multiple logistic regression analysis, habitat size and habitat type (spring‐fed versus non‐spring‐fed tributaries) were selected for the best model, whereas isolation was not important. However, when tributaries in the lower Sorachi basin (an area likely influenced by human activities) were excluded from the analysis, the effect of isolation became significant. 4. Our results indicate there is a potential metapopulation structure of the Dolly Varden in the Sorachi River basin. Furthermore, it is possible that metapopulation dynamics follow a clinal pattern. In the upper and middle reaches of the Sorachi River basin, an equilibrium between extinction and colonisation of Dolly Varden populations is a possibility. In the lower portion of the basin, isolation appeared to be less important, and extinction rate may be driving patterns of occurrence toward systematic extinction of local populations. 5. Our results illustrate the possibility of spatial variation in the processes influencing occurrence of Dolly Varden within a single river basin, and the potential influences of habitat alteration on extinction and colonisation processes.     

Geographical differences in marine feeding of Atlantic salmon post‐smolts in Norwegian fjords

Stomach content analyses were conducted on Atlantic salmon Salmo salar post‐smolt (average size, 119–154 mm fork length, L _F) caught in eight large Norwegian fjord systems along a north–south geographical axis during 1998–2001. In general, post‐smolts from southern Norway showed low feeding intensity in the fjords, whereas extensive feeding was observed in fjords in the northern and middle parts of Norway. The marine diet mainly included different crustaceans and in particular marine pelagic fish larvae (sand-eels Ammodytes spp., herring Clupea harengus and gadoids), but with a substantial spatial and annual variation in prey diversity and feeding intensity. Insects were most frequently taken in the estuary, although fishes often made a large contribution in mass. In contrast, fishes, and to some extent various crustaceans (particularly Hyperiidae, Gammaridae, Euphausiacea and Copepoda) dominated the diet in the middle and outer parts of the fjords, where post‐smolts also fed more extensively than in the inner part. The results indicate that extensive feeding immediately after sea entrance may be more common for post‐smolts in the northern and middle parts of Norway, than in the southern fjords. The observed differences in post‐smolt feeding may be due to spatial and temporal differences in prey availability within and between the different types of fjord systems, and this might influence post‐smolt growth and survival.     

Flexible seaward migration of Dolly Varden <Emphasis Type="Italic">Salvelinus malma</Emphasis> in the Shiretoko Peninsula,Hokkaido, Japan

The migratory history of Dolly Varden Salvelinus malma was investigated in 30 streams in the Shiretoko Peninsula, Hokkaido, Japan, which is in the southern part of their distribution. Captured Dolly Varden were classified into two categories by their body coloration: silvery fish were thought to be an anadromous form, and non-silvery fish a fluvial form that remains in freshwater throughout its life. Forty-seven of 1,739 Dolly Varden collected in the spring and ten of 2,126 Dolly Varden collected in the autumn were silvery fish. Twenty silvery fish and 120 non-silvery fish were examined by analysis of otolith Sr:Ca ratio. Three silvery fish from autumn and 15 non-silvery fish had migrated to salt water or brackish water. The other specimens of silvery fish were regarded as smolt or pre-smolt before seaward migration. The present study suggests that Dolly Varden in the Shiretoko Peninsula have a flexible migratory history, where some of the non-silvery fish utilize marine environments while some individuals follow an anadromous life history.     

Patterns of migration and residency in coastal cutthroat trout Oncorhynchus clarkii clarkii from two tributaries of the lower Columbia River

Coastal cutthroat trout Onchorhynchus clarkii clarkii life-history variants, migration and freshwater residency were monitored using stationary passive integrated transponder (PIT) tag arrays in two tributaries of the Columbia River from 2001 to 2005 (Abernathy Creek, river kilometre, rkm 76) and from 2002 to 2005 (Chinook River, rkm 6). In 2001–2003 and 2002–2003 (Abernathy and Chinook, respectively), 300–500 coastal O. c. clarkii were captured in each tributary by electrofishing and implanted with 23 mm PIT tags. PIT arrays monitored movements from the initiation of tagging through the spring of 2005. Rotary screw traps were also operated on both tributaries. In Abernathy Creek, 28% of tagged individuals were observed through either active capture or passive interrogation. Of these, 32% were identified as migrants and 68% were identified as residents. In the Chinook River, 48% of tagged fish were observed subsequent to tagging; 92% of these fish were migrants and only 8% were resident. In both tributaries, a greater proportion of resident fish were in the upper reaches. The majority of migrants (78–93%) moved the spring following tagging. Migrants leaving at age 2+ years tended to grow faster than those that migrated at age 3+ years or residents. Patterns of growth or growth opportunities may influence both patterns of life-history expression and the timing of migration.     

Sea‐run migratory behaviour in the Japan Sea form of three‐spined stickleback Gasterosteus aculeatus in the tidal pool of eastern Hokkaido Island,Japan

Juveniles in the Japan Sea form of three‐spined stickleback Gasterosteus aculeatus collected from a tidal pool in eastern Hokkaido Island, Japan, had a unimodal standard length (L_S) frequency distribution in each period, ranging from 11·9 to 31·6 mm, and those of sea‐run migrating individuals collected from the outlet of this tidal pool ranged from 17·0 to 36·0 mm, suggesting juveniles of this form would migrate towards the sea when they reached >17·0 mm. In general linear models (GLM) for testing the best model, there was a relationship between the number of sea‐run migratory individual and precipitation, which suggests that precipitation may trigger migration.     

Return migration of one‐sea‐winter Atlantic salmon in the River Tana

Fifty‐three one‐sea‐winter Atlantic salmon Salmo salar (45–63 cm L _T) were radio‐tagged in the Tana fjord, Barents Sea, in 1995. Thirty‐seven fish (70%) entered the freshwater zone of the River Tana in an average of 3 days after release in the fjord. The migration speeds in the lowest river section below the first riffle area were significantly higher than in the subsequent river section below the second riffle area. Similarly, the observed time spent in the first riffle area was significantly lower than in the next riffle area. The majority of Atlantic salmon entered the river during the hours of high tide and the subsequent ebb tide. In addition, most river entries were recorded around midnight. No effects of river flow on the river entry or migration speed were detected, but the migration speed of Atlantic salmon in both river sections examined was greater at lower temperatures. Twenty‐eight fish (72%) were recaptured in the river, 71% of them with weirs and gillnets, and 29% by rod and line. Over half of the Atlantic salmon (54%) were recaptured within 3 weeks following river entry, and within the first 100 km of the river (56%). The results are discussed in relation to earlier studies on multi‐sea‐winter Atlantic salmon in the River Tana.     

Strongly female‐biased sex allocation in a trivoltine population of Trypoxylon (Trypargilum) opacum Brèthes (Hymenoptera,Crabronidae)

Buschini, M.L.T. and Bergamaschi, A.C.B. 2009. Strongly female‐biased sex allocation in a trivoltine population of Trypoxylon (Trypargilum) opacum Brèthes (Hymenoptera, Crabronidae). —Acta Zoologica (Stockholm) 91 : 433–439. This study was carried in southern Brazil from December 2001 to December 2004. The aim of this paper is to provide additional information on the life‐history and sex allocation of this little known species. This wasp’s species has two alternative life histories: either they pupated immediately and emerged as adults later in the same season (direct development) or they entered diapause, overwintering and pupating the following spring (delayed development). The numerical sex ratio of overwintering and of direct developing wasps were strongly female biased in 2002, 2003 and 2004.     

Female‐biased size dimorphism in a diapausing caddisfly,Mesophylax aspersus: effect of fecundity and natural and sexual selection

1. The effect of mating success, female fecundity and survival probability associated with intra‐sex variation in body size was studied in Mesophylax aspersus, a caddisfly species with female‐biased sexual size dimorphism, which inhabits temporary streams and aestivates in caves. Adults of this species do not feed and females have to mature eggs during aestivation. 2. Thus, females of larger size should have a fitness advantage because they can harbour more energy reserves that could influence fecundity and probability of survival until reproduction. In contrast, males of smaller size might have competitive advantages over others in mating success. 3. These hypotheses were tested by comparing the sex ratio and body size of individuals captured before and after the aestivation period. The associations between body size and female fecundity, and between mating success and body size of males, were explored under laboratory conditions. 4. During the aestivation period, the sex ratio changed from 1 : 1 to male biased (4 : 1), and a directional selection on body size was detected for females but not for males. Moreover, larger clutches were laid by females of larger size. Finally, differences in mating success between small and large males were not detected. These results suggest that natural selection (i.e. the differential mortality of females associated with body size) together with possible fecundity advantages, are important factors responsible of the sexual size dimorphism of M. aspersus. 5. These results highlight the importance of taking into account mechanisms other than those traditionally used to explain sexual dimorphism. Natural selection acting on sources of variation, such as survival, may be as important as fecundity and sexual selection in driving the evolution of sexual size dimorphism.     

Interspecific competition affects evolutionary links between cavity nesting,migration and clutch size in Old World flycatchers (Muscicapdae)

The ecology of cavity nesting in passerine birds has been studied extensively, yet there are no phylogenetic comparative studies that quantify differences in life history traits between cavity‐ and open‐nesting birds within a passerine family. We test existing hypotheses regarding the evolutionary significance of cavity nesting in the Old World flycatchers (Muscicapidae). We used a multi‐locus phylogeny of 252 species to reconstruct the evolutionary history of cavity nesting and to quantify correlations between nest types and life history traits. Within a phylogenetic generalized linear model framework, we found that cavity‐nesting species are larger than open‐nesting species and that maximum clutch sizes are larger in cavity‐nesting lineages. In addition to differences in life history traits between nest types, species that breed at higher latitudes have larger average and maximum clutch sizes and begin to breed later in the year. Gains and losses of migratory behaviour have occurred far more often in cavity‐nesting lineages than in open‐nesting taxa, suggesting that cavity nesting may have played a crucial role in the evolution of migratory behaviour. These findings identify important macro‐evolutionary links between the evolution of cavity nesting, clutch size, interspecific competition and migratory behaviour in a large clade of Old World songbirds.     

Seasonal migration patterns of female sika deer in eastern Hokkaido,Japan

Fifty-seven female sika deer (Cervus nippon yesoensis), captured at the wintering area in the Shiranuka Hills in eastern Hokkaido, Japan, were radio-tracked during 1997–2001 to examine the factors affecting seasonal migration at the individual-landscape level. Ten of the 57 deers migrated between low-altitude summer home ranges and intermediate-altitude winter home ranges (the upward migrants). Twenty-nine migrated between high-altitude summer home ranges and intermediate-altitude winter home ranges (the downward migrants). Twelve used the intermediate-altitude home ranges all year round (the non-migrants). The remaining six were unknown. The summer home ranges of deer were widely scattered over an area of 5734km². Migration distances ranged between 7.2 and 101.7km. Deer showed high site fidelities to their seasonal home ranges. The upward migrants wintered in areas of less snow, higher quality of bamboo grass, and more coniferous cover than their summer home ranges. The downward migrants wintered in areas of less snow, higher quality of bamboo grass, higher winter temperature, and more southern slopes, but less coniferous cover than their summer home ranges. The non-migrants used year-round ranges with little snow, high quality of bamboo grass, and sufficient coniferous cover. We suggest that snow cover and bamboo grass are the factors affecting seasonal migration of the population and that coniferous cover is another factor for the upward migration.     

Proximate causes of sexual size dimorphism in Colorado pikeminnow, a long‐lived cyprinid

Evidence for sexual size dimorphism (SSD) and its possible causes were examined in the endangered Colorado pikeminnow Ptychocheilus lucius, a large, piscivorous, cyprinid endemic to the Colorado River system of North America. Individuals representing 18–24% of the upper Colorado River population were captured, measured, sexed and released in 1999 and 2000. Differing male and female total length‐(L_T) frequency distributions revealed SSD with females having greater mean and maximum sizes than males. Although both sexes exhibit indeterminate post‐maturity growth, growth trajectories differed. The point of trajectory divergence was not established, but slowed male growth might coincide with the onset of maturation. Differing growth rate was the dominant proximate cause of SSD, accounting for an estimated 61% of the observed difference in mean adult L_T. The degree of SSD in adults, however, was also related to two other factors. Evidence suggests males become sexually active at a smaller size and earlier age than females; a 2 year difference, suggested here, accounted for an estimated 12% of the between‐sex difference in mean adult L_T. Temporal shifts in gender‐specific survival accounted for an additional 27% of the observed between‐sex difference in mean adult L_T. Estimated age distributions indicated a higher number of older females than older males and more younger males than younger females in the population during the period of sampling. Dissimilarity of age distributions was an unexpected result because the male : female population sex ratio was 1 : 1 and estimates of long‐term annual survival for adult males and females were equal (88%). Future assessments of SSD in this population are apt to vary depending on the prior history of short‐term gender‐specific survival. Without recognizing SSD, non‐gender‐specific growth curves overestimate mean age of adult females and underestimate mean age of adult males of given L_T. Assuming age 8 years for first reproduction in males and age 10 years for females, the adult male : female ratio was estimated as 1·1 : 1 and mean adult age, or generation time, was estimated as 16·4 years for males and 18·4 years for females.     

Hatch date‐dependent differences in early growth and development recorded in the otolith microstructure of Trachurus japonicus

Otolith microstructure of 308 juvenile jack mackerel Trachurus japonicus sampled in the southern Sea of Japan between June and September 2002 were examined by light microscopy. Hatch‐date distribution indicated a protracted spawning season from 16 January to 30 May 2002. The first secondary primordium (SP) was formed at ages 19–54 days, with a mean ±  s . d . of 30·4 ± 6·1. The number of SP in an otolith ranged from two to 15, with a mean of 6·1 ± 1·7. Age of the first SP formation, number of SP and increment width varied with hatch date. In general, late‐hatched fish were younger when the first SP formed, and had a higher number of SP and wider increments, than early‐hatched fish. As increment width is a linear function of somatic growth and formation of the first SP is associated with metamorphosis in jack mackerel, these variations in otolith microstructure indicated that rates of growth (as inferred from wider increments) and development (as inferred by younger age of first SP formation) were higher in late‐hatched fish relative to early‐hatched fish. Higher temperature experienced by late‐hatched fish during early life stages may contribute critically to their higher growth and development rates compared to early‐hatched fish.     

Sex‐biased dispersal: a review of the theory

Dispersal is ubiquitous throughout the tree of life: factors selecting for dispersal include kin competition, inbreeding avoidance and spatiotemporal variation in resources or habitat suitability. These factors differ in whether they promote male and female dispersal equally strongly, and often selection on dispersal of one sex depends on how much the other disperses. For example, for inbreeding avoidance it can be sufficient that one sex disperses away from the natal site. Attempts to understand sex‐specific dispersal evolution have created a rich body of theoretical literature, which we review here. We highlight an interesting gap between empirical and theoretical literature. The former associates different patterns of sex‐biased dispersal with mating systems, such as female‐biased dispersal in monogamous birds and male‐biased dispersal in polygynous mammals. The predominant explanation is traceable back to Greenwood's ( 1980 ) ideas of how successful philopatric or dispersing individuals are at gaining mates or the resources required to attract them. Theory, however, has developed surprisingly independently of these ideas: models typically track how immigration and emigration change relatedness patterns and alter competition for limiting resources. The limiting resources are often considered sexually distinct, with breeding sites and fertilizable females limiting reproductive success for females and males, respectively. We show that the link between mating system and sex‐biased dispersal is far from resolved: there are studies showing that mating systems matter, but the oft‐stated association between polygyny and male‐biased dispersal is not a straightforward theoretical expectation. Here, an important understudied factor is the extent to which movement is interpretable as an extension of mate‐searching (e.g. are matings possible en route or do they only happen after settling in new habitat – or can females perhaps move with stored sperm). We also point out other new directions for bridging the gap between empirical and theoretical studies: there is a need to build Greenwood's influential yet verbal explanation into formal models, which also includes the possibility that an individual benefits from mobility as it leads to fitness gains in more than one final breeding location (a possibility not present in models with a very rigid deme structure). The order of life‐cycle events is likewise important, as this impacts whether a departing individual leaves behind important resources for its female or male kin, or perhaps both, in the case of partially overlapping resource use.     

Ancestral ecological regime shapes reaction to food limitation in the Least Killifish,Heterandria formosa

Populations with different densities often show genetically based differences in life histories. The divergent life histories could be driven by several agents of selection, one of which is variation in per‐capita food levels. Its relationship with population density is complex, as it depends on overall food availability, individual metabolic demand, and food‐independent factors potentially affecting density, such as predation intensity. Here, we present a case study of two populations of a small live‐bearing freshwater fish, one characterized by high density, low predation risk, low overall food availability, and presumably low per‐capita food levels, and the other by low density, high predation risk, high overall food availability, and presumably high per‐capita food levels. Using a laboratory experiment, we examined whether fish from these populations respond differently to food limitation, and whether size at birth, a key trait with respect to density variation in this species, is associated with any such differential responses. While at the lower food level growth was slower, body size smaller, maturation delayed, and survival reduced in both populations, these fitness costs were smaller in fish from the high‐density population. At low food, only 15% of high‐density fish died, compared to 75% of low‐density fish. This difference was much smaller at high food (0% vs. 15% mortality). The increased survival of high‐density fish may, at least partly, be due to their larger size at birth. Moreover, being larger at birth enabled fish to mature relatively early even at the lower food level. We demonstrate that sensitivities to food limitation differ between study populations, consistent with selection for a greater ability to tolerate low per‐capita food availability in the high‐density population. While we cannot preclude other agents of selection from operating in these populations simultaneously, our results suggest that variation in per‐capita food levels is one of those agents.     

Upstream migration rates of radio‐tagged adult Chinook salmon in riverine habitats of the Columbia River basin

Upstream migration rates were assessed for 1801 radio‐tagged adult spring–summer Chinook salmon Oncorhynchus tshawytscha through 12 unimpounded river reaches in the Columbia River basin from 1997 to 2002. Reaches were 36 to 241 km long (mean = 130 km) and included sections of the large Columbia and Snake Rivers and smaller free‐flowing tributaries. Median Chinook salmon migration rates ranged from <10 km day⁻¹ in the Deschutes and Clearwater Rivers to >35 km day⁻¹ in the Columbia and Snake Rivers. Using multivariate analyses, migration date explained the most variance in Chinook salmon migration rates while river discharge, migration year and migration reach were secondary. Both within and between years, Chinook salmon migrated more rapidly as migration date increased and more slowly when discharge was high. Arrival at high elevation spawning grounds at appropriate times and increased metabolic activity and reproductive maturation may explain the greater power of migration date, relative to river discharge, in predicting migration rates of Columbia basin spring–summer Chinook salmon.     

Stream-dwelling Dolly Varden (<Emphasis Type="Italic">Salvelinus malma</Emphasis>) density and habitat characteristics in stream sections installed with low-head dams in the Shiretoko Peninsula,Hokkaido, Japan

Numerous low-head dams have been constructed for erosion control and installed in almost all mountainous streams in Japan. Shiretoko Peninsula, Hokkaido, is considered as one of the most intact areas in Japan, however, more than 331 low-head dams have been installed in 39 streams of the peninsula since the 1960s. In this study, the density of stream-dwelling Dolly Varden (Salvelinus malma) and the habitat characteristics in dam-installed sections were investigated in Shiretoko streams and compared with unaltered sections in which no dams have been installed as a reference. Fish density in dam-installed sections was significantly lower than reference sections. Canopy cover, maximum temperature, wetted width, frequencies of pebble, bedrock and boulder, and areas of rapid, glide and riffle was significantly differed between dam-installed and reference sections. High stream temperature, occurrence of glides, and loss of boulder in dam-installed sections may reduce the suitability of habitats for fish and minimize the population size. We suggest that negative effects of dam installations on Dolly Varden populations have occurred in Shiretoko streams.     

Home range size and distribution in a wild population of Cape pangolins, Manis temminckii, in north‐west Zimbabwe

The aim of this study was to estimate the size of the home range used by individual Cape pangolins ( Manis temminckii ). The study was conducted in 1991–95 at Sengwa Wildlife Research Institute, Gokwe, Zimbabwe. Radio‐telemetry was used to repeatedly locate study pangolins. Home range area was estimated from the location of the burrows used by each pangolin. It was established that pangolins must be tracked for 85+ days to generate a reliable estimate of home range. The size of home ranges, determined from 1141 tracking days of data for 10 pangolins (3·0–15·8 kg body mass) that were each tracked for at least 85 days, was from 0·17 to 11·07 km². Larger (older) pangolins used more burrows and had larger home ranges than smaller (younger) pangolins. The data indicate that large adult males had larger home ranges than large adult females. Within each sex, the home ranges were adjacent to each other with only slight overlap at the boundaries. There was clear overlap of home ranges between males and females.