Population growth confounds phylogeographic inference in Namaqua sandgrouse

The Namaqua sandgrouse, Pterocles namaqua, is a highly nomadic granivore of semiarid to arid habitats. As a result of nomadic movements in response to rainfall, the size of the breeding population in any one area fluctuates dramatically between breeding seasons. This high mobility in response to spatial and temporal abundance of food resources is expected to result in little population genetic structuring. Namaqua sandgrouse also shows a seasonally predictable partial migration between the southeast and northwest regions of South Africa, and a further possible north-south migration between southwestern South Africa and central Namibia. It is unclear whether birds migrating between these regions breed in only one or both regions. If populations breed in only one region of their migratory range, then population genetic structuring is predicted to occur. This study addresses Namaqua sandgrouse movements with the analysis of mitochondrial DNA control region sequences. In general, little population genetic structure was evident, yet strong signals of population growth were detected. Several populations have private alleles, which is in direct contradiction to the spatial genetic pattern expected under high levels of gene flow. We suggest that the inference of high levels of female gene flow could be an artifact of population growth and that additional loci will allow a greater understanding of Namaqua sandgrouse movements.     

The relationship between outbreaks, rainfall and low density populations of the African armyworm, Spodoptera exempta, in Kenya

This study has demonstrated that low numbers of African armyworms (Spodoptera exempta) (Walker) (Lepidoptera: Noctuidae) can be found throughout the year in parts of the coastal and highland regions of Kenya where there is frequent rainfall. During the study period the numbers of moths caught in pheromone traps in these regions built up during the short and long rains, and decreased dramatically during the intervening dry seasons. There was a lag of one to two months between the peaks for rainfall and moth numbers. This contrasts with the situation at sites of the seasonal outbreaks of armyworm larvae, where a sudden preceding influx of moths coincides with the rainfall. A positive correlation was found, for the long rains seasons only, between the number of armyworm outbreaks throughout the country and the peak numbers of moths in coastal and highland regions.It is suggested therefore that the peak numbers of moths trapped in the eastern highlands and coastal regions during the rainy seasons arise principally from outbreaks in the extensive seasonal grasslands. These grassland areas diminish considerably during the prolonged dry periods between the two rainy seasons, and it is only the eastern highland and coastal regions that are likely to provide suitable habitats for breeding during the dry seasons. The significance of these populations for initiating the first outbreaks of the following season is discussed.     

Gonadal and reproductive patterns in Tilapia leucosticta (Teleostei: Cichlidae) in an equatorial lake, Lake Naivasha (Kenya)

The cichlid fish T. leucosticta (Trewavas) was studied by fortnightly seining an area of an equatorial lake (Lake Naivasha) in Kenya to study their reproductive and gonadal patterns. The study lasted 16 months during which period temperature, rainfall and (for the last 11 months) sunlight patterns were obtained and analysed. Testicular and ovarian development was lowest in the months of July-August. No evidence of breeding was detected during this period. This period is also marked by the lowest rainfall, the lowest temperatures and least sunlight. Slight but fluctuating breeding activity and gonadal development is resumed in October following increased sunlight and temperatures from September. The period December to February is marked by sustained high temperatures, high sustained maximal sunlight and low rainfall. It is also the period of rising gonadal development in most mature males and females reaching a peak in both sexes in February. This is followed shortly after by a sharp peak in reproductive activity. It is concluded that sunlight and temperature are the most important factors influencing gonadal development and subsequent breeding in Tilapia even under equatorial conditions. When preceded by conditions of sustained high temperatures and good sunlight which are conducive to the maximal development of the gonads of both sexes, the onset of rainfall seems to stimulate the initiation of the peak breeding activity. Sustained heavy rainfall on the other hand seems to check breeding.     

Some aspects of the breeding biology of the equatorial land snail Limicolaria martensiana (Achatinidae: Pulmonata)

A study of the breeding season of the polymorphic equatorial land snail Limicolaria martensiana Smith (Achatinidae: Pulmonata) lasting six years and six months in populations in the Kampala District (lat. 0 20') and the Equator line shows that the snail breeds continuously throughout the year but has bimodal peaks in its breeding which occurs during the dry seasons, while during the rainy seasons breeding is low. It is the decrease in the amount of rainfall which induces peak breeding and as a result there is an inverse relationship between the amount of rainfall and the breeding intensity. The evolution of ovoviviparity with the ability to retain eggs and/or young in utero during aestivation is the phenomenon that enables L. martensiana to have peak breeding during the dry seasons. Moisture which enables young to feed and thus grow is an ultimate factor in determining breeding in L. martensiana . The gestation period is 10 to 14 days during the rainy seasons but the snail retains its eggs and/or young in utero during the dry seasons and only deposits them when rain returns. The eggs take 18.6±5.43 days ( N= 130) to hatch after they are laid. Hatching in utero and outside occurs by the young eating the egg shells. Oogenesis in the snails occurs at all times. L. martensiana stores viable allosperms which it can use for further reproduction.     

Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.)

Climatic and biogeographical associations of southern African dung beetles (Scarabaeinae, Coprinae) were analysed from a collection amassed between 1971 and 1986. Endemism to Africa south of 15°S was much greater in southwesterly climates (winter rainfall, bimodal spring/autumn rainfall, arid late summer rainfall) than to the north‐east in the moist, mid‐summer rainfall region. Major biogeographical groups centred to the south‐west comprised predominantly southern African endemics, Western/Eastern Cape coast, Karoo, Karoo/Namib, northern Namibia and the south‐western Kalahari. Biogeographical groups centred on the south‐eastern highlands and the subtropical east coast (mid‐summer rainfall) also comprised predominantly southern African endemics. All other major groups centred to the north‐east in the mid‐summer rainfall region comprised predominantly species with widespread tropical biogeographical affiliations, pan‐southern Africa but centred in the east, pan‐mid‐summer rainfall region, eastern mid‐summer rainfall region, tropical east Zimbabwe/central Moçambique, subtropical/tropical game reserves (non‐ruminant dung specialists). There were cross‐climatic differences in taxonomic composition of the fauna. Within the winter rainfall region, percentage species composition of Scarabaeinae was greater whereas that of the coprine tribe, Onthophagini, was lower than within the other three climatic regions. Percentage species richness of most other tribes of Coprinae differed little between most climates but that of Scarabaeinae declined from west to east (Canthonini, Scarabaeini), east to west (Sisyphini) or to either side of the late summer rainfall region (Gymnopleurini).     

Effect of abiotic factors on reproduction in the centre and periphery of breeding ranges: a comparative analysis in sympatric harriers

Variables such as weather or other abiotic factors should have a higher influence on demographic rates in border areas than in central areas, given that climatic adaptation might be important in determining range borders. Similarly, for a given area, the relationship between weather and reproduction should be dissimilar for species which are in the centre of their breeding range and those that are near the edge. We tested this hypothesis on two sympatric ground‐nesting raptors, the hen harrier Circus cyaneus and the Montagu's harrier Circus pygargus in Madrid, central Spain, where the hen harrier is at the southern edge of its breeding range in the western Palearctic and the Montagu's harrier is central in its distribution. We examined the reproductive success of both species during an 8‐yr period, and looked at the influence of the most stressful abiotic factors in the study area (between‐year variation in rainfall and within‐year variation in temperature) on reproductive parameters. In the hen harrier, low levels of rainfall during the breeding season had a negative influence on annual fledging success and thus on population fledgling production. The relationship between rainfall and reproduction was probably mediated through food abundance, which in Mediterranean habitat depends directly on rainfall levels. In the Montagu's harrier, no negative effect of dry seasons on productivity was found. Additionally, in the hen harrier, the proportion of eggs that did not hatch in each clutch increased with higher temperatures during the incubation period. No such relationship was found in the Montagu's harrier. We interpret these between‐species differences in terms of differences of breeding range and adaptations to the average conditions existing there. Hen harriers, commonest at northern latitudes, are probably best adapted to the most typical conditions at those latitudes, and have probably not developed thermoregulatory or behavioural mechanisms to cope with drought and high temperatures in Mediterranean habitats, in contrast to Montagu's harrier. Thus hen harrier distribution might be constrained by these variables, due to lower reproductive success or higher reproductive costs. Accordingly, a logistic regression analysis of the presence or absence of both species in 289 random points throughout the western Palearctic showed that the distribution of both species was related to temperature, but the relationship was in opposite directions for the two species: hen harriers had lower probability of breeding in areas with higher temperature (as expected in a species with a more northerly distribution).     

Habitat preferences of sympatric sandgrouse during the breeding season in Spain: a multi-scale approach

Predictive species’ distribution models may answer ecological questions about habitat selection, co-occurrence of species and competition between them. We studied the habitat preferences and segregation of two sympatric species of declining sandgrouse, the black-bellied sandgrouse (Pterocles orientalis) and the pin-tailed sandgrouse (Pterocles alchata), during the breeding season. We developed predictive models that related sandgrouse presence to environmental variables at three different spatial levels: large geographical, landscape and microhabitat scales. At the large geographical scale, differences between sandgrouse distributions, in the Iberian Peninsula, seem to be explained mainly in terms of bioclimatology: pin-tailed sandgrouse appear to be a more thermophilous species and occupy warmer sites usually located in flatter areas. At the landscape spatial level, in those areas that exhibit environmental conditions allowing for both species’ co-existence at a large geographical scale, black-bellied sandgrouse appear to be more tolerant to environmental variation than pin-tailed sandgrouse. At the microhabitat level, however, differences between species could be related to different flocking behaviour as a consequence of different sensitivities to vegetation structure and predators. Thus, the observed spatial distribution patterns are the result of different ecological factors that operate at different spatial levels. Conservation guidelines for these species should therefore consider their habitat preferences at large geographical, landscape and microhabitat scales.     

The influence of climate variability on numbers of three waterbird species in Western Port, Victoria, 1973–2002

Seasonal and annual movements of Australian waterbirds are generally more complex than those of their Northern Hemisphere counterparts, and long-term data are needed to understand their relationships with climatic variables. This paper explores a long-term (1973–2002) set of waterbird counts from coastal Victoria and relates them to climatic data at local and continental scales. Three species (Black Swan Cygnus atratus, White-faced Heron Egretta novaehollandiae and Grey Teal Anas gracilis) were chosen for this analysis. Black Swans have large local breeding populations near the study region; White-faced Herons have smaller local breeding populations and Grey Teal breed extensively in ephemeral inland floodplains, such as those in the Murray-Darling Basin. All showed significant relationships with streamflow, regional rainfall and the Southern Oscillation Index (SOI) at appropriate scales and time-lags, with streamflow explaining the most variance. Black Swans showed a strong seasonal cycle in abundance and local climate variables had the greatest influence on the counts. Numbers were positively correlated with streamflow in southern Victoria three to six seasons before each count. Broader-scale climatic patterns were more important for the other two species. Numbers of White-faced Herons were positively correlated with streamflow or rainfall over various parts of Australia seven to nine seasons before each count. Numbers of Grey Teal showed weak seasonal cycles, and were negatively correlated with rainfall in Victoria or the Murray-Darling Basin in the seasons before or during each count, and positively with streamflow in the Murray-Darling Basin 15–18 months before each count.     

Relationship between breeding activity and rainfall for Swainson's Spurfowl,Pternistis swainsonii,within southern Africa,with specific reference to the Springbok Flats,Limpopo Province,South Africa

We collated the literature available on the breeding activity of the Swainson's Spurfowl Pternistis swainsonii and made use of reliable unpublished reports, nest record cards and field observations within the Springbok Flats, Limpopo Province, South Africa to establish breeding seasons and pairing behaviour. The onset of breeding (egg laying) is closely associated with rainfall, with male gonad development, population density and covey size (pairing behaviour), all correlated with rainfall. Peak breeding activity is from January–April in South Africa, February–May in Zimbabwe and March–June in Botswana. Egg laying has been recorded in all months and sporadic egg laying in the winter months is most likely the result of isolated rainfall. Mean clutch size is 5.2 eggs/hen (n = 140) with an incubation period of 23 days and brood hatching success and chick survival of 69.4% over the southern African sub-region. Current hunting seasons within Limpopo Province are in line with the recommended hunting season for this region and should remain unchanged: 15 June–30 September. The success of this phasianid can be attributed to its extended breeding season, high survival rate of hatchlings and the potential of birds to breed within their first year.     

Niche-habitat mechanisms and biotic interactions explain the coexistence and abundance of congeneric sandgrouse species

Ascertaining which niche processes allow coexistence between closely related species is of special interest in ecology. We quantified variations in the environmental niches and densities of two congeneric species, the pin-tailed and the black-bellied sandgrouse (Pterocles alchata and Pterocles orientalis) in allopatry and sympatry under similar abiotic, habitat and dispersal contexts to understand their coexistence. Using principal component analysis, we defined environmental gradients (niche dimensions) including abiotic, habitat and anthropogenic variables, and calculated niche breadth, position and overlap of both species in sympatry and allopatry. Additionally, sandgrouse density was modelled as a function of the niche dimensions and the density of the other species. We found evidence that each species occupies distinct environmental niches in sympatry and in allopatry. The black-bellied sandgrouse exploits a broader range of environmental conditions (wider niche breadth) while the pin-tailed sandgrouse reaches high densities where conditions seem to match its optimum. In sympatry, both species shift their niches to intermediate positions, indicating the importance of abiotic factors in setting coexistence areas. Environmental conditions determine regional densities of pin-tailed sandgrouse whereas biotic interactions explain the density of the black-bellied sandgrouse in areas with abiotic conditions similarly conducive for both species. Highly suitable areas for the pin-tailed sandgrouse fall beyond the upper thermal limit of the black-bellied sandgrouse, leading to niche segregation and low densities for the latter. Finally, local niche shift and expansion plus possible heterospecific aggregation allow the pin-tailed sandgrouse to thrive in a priori less favourable environments. This work provides insight into how different mechanisms allow species coexistence and how species densities vary in sympatry compared to allopatry as a result of environmental filtering and biotic interactions.     

Influence of rainfall on timing and success of reproduction in Marabou Storks Leptoptilos crumeniferus

A decline in breeding success with later laying dates throughout a nesting season is a widespread phenomenon in species living in environments with distinct seasonality, with evidence that some environmental correlate of timing is at least partly responsible in many species. This correlate is often thought to be food availability, which is often related to climatic factors; however, few studies have examined the role of climate. We studied a breeding colony of Marabou Storks Leptoptilos crumeniferus in Southern Africa over five breeding seasons. Timing of breeding was related to rainfall preceding the breeding season. Fecundity (chicks fledged per nest) declined through each season. The probability of an individual hatchling fledging was influenced by rainfall during the hatchling period, temperature during the hatchling period and laying date, three variables that were strongly intercorrelated. To disentangle the three effects, inter‐annual variation in each was compared with the large inter‐annual variation in breeding success, with rainfall providing the greatest explanatory power. Rainfall, which tends to increase through the breeding season, seems to be at least partly responsible for the seasonal decline in breeding success. We were unable to find evidence for the influence of other factors, such as colony size and nest re‐use, known to affect nest success in this and other colonial breeding storks.     

Lacustrine spawning: is this a new reproductive strategy among ‘large’ African cyprinid fishes?

Changes in the gonado‐somatic index and abundance of the different Labeobarbus species in the mouths of four major afferent rivers of Lake Tana, Ethiopia, were monitored monthly during 1999 and 2000. Riverine spawning was characteristic for seven of Lake Tana's 15 contemporary Labeobarbus species. These seven did not show spatial segregation among afferent rivers but significant temporal segregation occurred in aggregating in the river mouths and migrating towards the upstream spawning areas during the breeding season (June–October). Among the eight other species, peak gonad development occurred generally in the same period as in the riverine spawners. These species, however, did not aggregate in the river mouths during the breeding period and were absent from the upstream spawning areas. A derived, novel strategy, lacustrine spawning was hypothesized for these eight Labeobarbus species. This hypothesis was further supported by observations of running female fishes in the littoral zones distant from any of the afferent rivers. This derived strategy is only common among the littoral‐dwelling Labeobarbus species with restricted distribution patterns. At present it is thought that sequential waves of speciation and habitat divergence followed by trophic specialization, shaped the diversity of Lake Tana labeobarbs.     

A multi-scale approach for identifying conservation needs of two threatened sympatric steppe birds

Habitat selection is an inherently scale-sensitive process in which detected selection patterns frequently depend on the scale of analysis employed. We used a multi-scale modelling approach to identify how the distributions of two sympatric birds are shaped by differential selection at the landscape, land use and microhabitat scales and by human infrastructures as possible sources of disturbance. We studied two threatened steppe birds, the pin-tailed sandgrouse (PTS) and black-bellied sandgrouse (BBS) in central Spain. Land use gradients explained most of the variation in PTS and BBS occurrence, but there was cross-scale interdependence between the lower (microhabitat) and upper (landscape) spatial scales for the PTS. Synergies between the three scales highlighted the importance of integrating habitat scales in a single modelling framework. The process of habitat selection was also modulated by human disturbance. Both species selected ploughs of large size distant from houses, tracks and other infrastructures, although BBS exhibited broader habitat tolerance than the PTS, and was more sensitive to human disturbance. At microhabitat scale, PTS selected ploughs with greater green vegetation cover and insect abundance and fallows with lower dry vegetation cover and height but greater stone cover. This might reflect a trade-off between camouflage (vegetation and stone cover for concealment) and visibility for predator detection and escape. Ploughs and fallows should be maintained by means of traditional 2-year rotations and low management during the breeding season. Ongoing urbanization trends and infrastructure development inside protected areas should be limited. Multi-scale models were key to identify scale-specific factors that determine sandgrouse habitat preferences and conservation requirements at appropriate levels, and are recommended to better guide regional and local conservation efforts of threatened species.     

Characterization and utilization of microsatellites in the <Emphasis Type="Italic">Coffea canephora</Emphasis> genome to assess genetic association between wild species in Kenya and cultivated coffee

Coffee is an important beverage crop in the world and has a significant contribution to Kenya’s economy. Here, we analyzed the genome-wide distribution of microsatellites in the Coffea canephora genome. A total of 159,041 SSRs were identified, with an overall density of 308 SSRs per Mb. Tetra-nucleotide repeats are the most abundant, accounting for 32 % of the total SSRs. AT-rich motifs are dominant across all SSR repeat units, while GC-rich motifs were generally rare. A set of 100 SSRs was selected to amplify 96 coffee accessions, including 10 wild accessions collected from Mt. Marsabit (Kenya). Of these SSRs, 33 % generated clear polymorphic bands among all tested accessions, with an average of 3.9 alleles per SSR locus. Wild coffee species from Mt. Marsabit showed a close genetic similarity with cultivated accessions in Kenya, suggesting that the wild species in Mt. Marsabit played an important role in the domestication of cultivated coffee in Kenya. Significantly low pairwise genetic divergence was observed between cultivated and wild accessions in Kenya, suggesting a relatively narrow level of genetic basis among coffee germplasm in Kenya. In addition, cultivated and wild coffee accessions in Kenya show a great divergence from those in other countries. Our results not only provide molecular tools for genetic studies in coffee but are also helpful for conservation and coffee breeding programs in Kenya.     

BREEDING SEASONS OF BIRDS IN DECIDUOUS WOODLAND AT ZIMBABWE,RHODESIA, FROM 1970 TO 1974

Vernon, C. J. 1978. Breeding seasons of birds in deciduous woodland at Zimbabwe, Rhodesia, from 1970 to 1974. Ostrich 49:102-115.

The birds in Brachystegia woodland bred in the spring and 98% of 1645 breeding records occurred between September and November. 90% of the records, and 43 of the 71 breeding species, were of insectivorous passerines. The breeding seasons of individual species lasted for between less than two months to more than three months, and their peak breeding periods varied between early September and the middle of October. The breeding season started earliest in 1974 and latest in 1973, and the start of two species was related to the timing of leaf emergence of the dominant tree in the woodland and to rainfall the previous rainy season. Food supply may be the proximal factor influencing the onset of breeding.     

Do females adjust the sex of their offspring in relation to the breeding sex ratio?

When the adult sex ratio differs between years in local populations, but still is predictable between adjacent years, it has been proposed that the best strategy would be to bias the offspring sex ratio in favour of the rare sex. We tested this hypothesis using a data set of great reed warbler offspring, sexed by molecular techniques, that were collected over 11 breeding seasons at two adjacent reed marshes. Three important assumptions for this hypothesis are fulfilled in the studied great reed warbler population. First, a substantial proportion of great reed warblers are living in small local populations where sex ratio distortions would be sufficiently large and common. Second, breeding adults and their offspring return to breed in the local population to a high degree. Third, females have a possibility to assess the breeding sex ratio before laying their eggs. At our study site, the breeding sex ratio was positively correlated between successive years. However, contrary to our prediction, female great reed warblers seemed not to adjust their offspring sex ratio in relation to the local breeding sex ratio.     

Habitat use by a Midwestern U.S.A. riverine fish assemblage: effects of season, water temperature and river discharge

The hypothesis that temperate stream fishes alter habitat use in response to changing water temperature and stream discharge was evaluated over a 1 year period in the Neosho River, Kansas, U.S.A. at two spatial scales. Winter patterns differed from those of all other seasons, with shallower water used less frequently, and low‐flow habitat more frequently, than at other times. Non‐random habitat use was more frequent at the point scale (4·5 m²) than at the larger reach scale (20–40 m), although patterns at both scales were similar. Relative to available habitats, assemblages used shallower, swifter‐flowing water as temperature increased, and shallower, slower‐flowing water as river discharge increased. River discharge had a stronger effect on assemblage habitat use than water temperature. Proportion of juveniles in the assemblage did not have a significant effect. This study suggests that many riverine fishes shift habitats in response to changing environmental conditions, and supports, at the assemblage level, the paradigm of lotic fishes switching from shallower, high‐velocity habitats in summer to deeper, low‐velocity habitats in winter, and of using shallower, low‐velocity habitats during periods of high discharge. Results also indicate that different species within temperate river fish assemblages show similar habitat use patterns at multiple scales in response to environmental gradients, but that non‐random use of available habitats is more frequent at small scales.     

Diet of Black-legged Kittiwakes (Rissa tridactyla) feeding chicks at two Irish colonies highlights the importance of clupeids

Capsule We examined regurgitates from Black-legged Kittiwakes during the early chick-rearing period over two breeding seasons at two colonies in Ireland where diet has not been studied previously. Clupeids were the dominant food source at both colonies, which contrasts with other studies throughout the British Isles that suggest Kittiwakes feed mainly on sandeels during this period. Our study is limited by sample size and restricted to the early breeding season, but suggests a link between reproductive success and dietary composition which should be investigated further.     

Association of seasonal reproductive patterns with changing food availability in an equatorial carnivore, the spotted hyaena (Crocuta crocuta).

Reproductive seasonality was examined in an equatorial population of free-living spotted hyaenas (Crocuta crocuta) in Kenya. The study population was observed continuously for 10 years, during which time the dates of all births, conceptions, weanings, and cub deaths were recorded. Local prey abundance was estimated two to four times per month, and rainfall was recorded daily throughout the study period. Births occurred during every month of the year, but a distinct trough in births occurred from February to May. This trough occurred approximately one gestation period after the phase of the annual cycle during which prey animals were least abundant in the home range of the hyaenas, and conceptions occurred most frequently when food abundance was greatest. Neither rainfall nor cub mortality were correlated with births or conceptions. Thus, although spotted hyaenas are capable of breeding throughout the year, they exhibit a moderate degree of seasonality that most likely reflects responses to seasonal variation in energy availability.     

Seasonal change in the spatial utilization of host mussels in relation to ovipositor length by female rosy bitterling Rhodeus ocellatus kurumeus

The seasonal changes in ovipositor length and utilization patterns of mussels for oviposition in the rosy bitterling Rhodeus ocellatus kurumeus (Cyprinidae) were investigated in a field experiment and field surveys during the breeding period (April to August). The mean length of ovipositors at oviposition was short at the start (early April) and end (July) of the breeding period. Females with long ovipositor at oviposition were collected between mid‐April and June. Mark‐and‐recapture data showed that ovipositor length at oviposition changed rhythmically throughout the breeding period, shortening and lengthening as the female entered the spawning and resting phase. The density of rosy bitterling embryos in mussels increased between April and June, peaking in May, but decreasing in July. The position of eggs on mussel gills varied from close to the exhalant siphon to deeper inside the gill during April, and periodically thereafter. There was a positive correlation between ovipositor length at oviposition and the distance from exhalant siphon of mussels to eggs deposited by females, suggesting that ovipositor length at oviposition determined the position of eggs deposited on a mussel gill. Because dissolved oxygen in mussel gills decreased with the density of bitterling embryos, suitable positions for embryo survival in gills changed with embryo density. By changing ovipositor length at oviposition, females might be able to spawn their eggs in a position that maximizes embryo survival. Thus, plasticity in ovipositor length at oviposition may play an important role as an adaptation of rosy bitterling in utilizing mussels when their quality as a spawning substratum fluctuates seasonally.