Changes in serum thyroxine levels during smolting in captive and wild Atlantic salmon, Salmo salar L.

On a commercial fish farm in Western Scotland, smolting Atlantic salmon, Salmo salar L., showed no major changes in serum thyroxine (T4) levels in the spring. In contrast, serum T4 levels in smolting, wild salmon, trapped as they left the Girnock Burn, Aberdeenshire, varied markedly over this period. Serum T4 levels were correlated with the stream discharge rate pertaining as the fish left the stream. Levels in migrating, smolting fish were higher than those in fish captured above the fish-trap by electro-fishing. In a stream channel, serum T4 levels in smolting fish caught previously in the fish-trap were rapidly elevated by exposure to increased water-flow, whereas levels in fish previously captured above the trap remained substantially unchanged. Serum T4 levels in salmon parr leaving the stream in autumn were generally lower than those in smolting fish leaving in spring and were not demonstrably affected by stream discharge rate.     

BIRD MIGRATION AT ELAT, ISRAEL

Several years' observations at the head of the Gulf of Aqaba are summarized. Migrants occur the whole year round, but most numerously in March-April and September. Only a few of the 185 species recorded pass the winter at Elat. About 25 species are recorded more in autumn, whereas about 50 species are commoner in spring. The causes of these disparities are discussed. Thousands of soaring raptors pass through, mainly in spring. In spring only, thousands of Lesser Black-backed Gulls stream through Elat, and many rest there for a short period. Mass migration of storks occurs too, more conspicuously in spring than autumn. An attempt is made to construct the routes of these birds between Eurasia and Africa, by analysing published sight records of raptors and storks and ringing recoveries of gulls. It is suggested that many of these birds move in autumn on a wide front, which may include Arabia, but that the core of the spring passage is shifted westward and part of it is channelled through the Rift Valley north of the northern end of the Red Sea and in the areas between the rift and the Mediterranean (Fig. 4). Supporting evidence is still needed from Arabia and the coasts of the Red Sea, especially from its southern end, where birds may be concentrated at the straits of Bab-el-Mandeb, as they are over the Bosphorus. Pelicans and a few other species perform very late southerly movements. These movements involve small numbers of birds, which may belong to late-breeding populations. About 45 other species of water and shore birds have been recorded, many of which occur in winter. With the expansion of areas of artificial water, some of them have become very common. About 75 passerines, near-passerines and other migrants pass through. The numbers involved suggest that the movement is on a broad front. Out of about 50 species whose passage is adequately recorded for seasonal comparison, 30 are more common in spring. Most of these are also commoner in other eastern Mediterranean countries and in Iraq in spring, md are presumed to perform a continuous overhead flight in autumn. Cases of “loop-migration” among these species are rare.     

Spatial distribution and temporal dynamic of the seed pool in a Danish lowland stream

In this paper we quantify the seed pools in four locations down through a small stream in Denmark. The objectives of the study were (1) to test if the number of plant taxa in the seed pool increases downstream in the stream system; (2) to test if the seed density in the pool is highest in autumn; and (3) to analyze to what extent the seed pool contributes to re-establishment of vegetation in spring. Counting method showed no increase in species taxa in the seed pool down through the stream system. Neither did we find clear seasonal patterns in seed abundance in the seed pool. By concurrent field and laboratory observations in spring we found that seed pools can function as a source for re-establishing the vegetation in spring. However, the actual germination rate in the field was <100 and <10 times the germination rate in the laboratory for aquatic and terrestrial plants, respectively. The results suggest that small lowland streams contain large seed pools in deposition areas throughout the year, and that seed pools are much more important for the re-establishment of riparian stream vegetation in spring than submerged vegetation.     

Composition and intra-annual variation of the macroinfauna in the estuarine zone of the Pando Stream (Uruguay).

We studied the temporal patterns of macroinfaunal distribution in the estuarine portion of Pando Stream, Uruguay. We found a very low number of species and a high dominance of the polychaete Heteromastus similis. There were clear seasonal patterns in abundance with almost all species peaking in autumn, coinciding with the periods of higher salinity. Minimal values were found in winter and spring when salinity was at a minimum. We suggest that the seasonal pattern is a consequence of the combined effects of recruitment, and of high osmotic stress and disturbance during the periods of freshwater discharge that are common in winter and spring.     

Mate preference in males of the cabbage butterfly, Pieris rapae crucivora, changes seasonally with the change in female UV color

We initially investigated whether females of the cabbage butterfly, Pieris rapae crucivora, exhibit a seasonal change in ultraviolet wing color, which is a key stimulus for mate recognition by conspecific males, and whether and how a seasonal change affects the mating behavior of the males. We found that female UV wing color changes seasonally, the color being more pronounced in summer than in spring or autumn. We also demonstrated that male mate preference changes seasonally, concomitantly with the change in female UV color. Specifically, males appearing in summer exhibit a mating preference for summer-form females over spring- or autumn-form females, while those appearing in spring or autumn exhibit no seasonal preference, thereby facilitating more effective mate location. Our results suggest that this field of study will require more strictly controlled experimental investigation in which the seasonal change in UV color is considered when UV-influenced mating behaviors such as mate choice are investigated.     

The downstream migration of precociously mature Atlantic salmon, Salmo salar L, parr in autumn; its relation to the spawning migration of mature adult fish

The autumn emigration of juvenile Atlantic salmon, Sulmo salur L. from the Girnock Burn, Aberdeenshire, is shown to be comprised of two components. Precociously mature males and sexually immature fish differ in their dependence on seasonal and biological cues for the release of migratory behaviour. The emigration of ripe mature male parr is shown to follow the entry of mature adult salmon to the stream; when adult fish are prevented from entering the stream, the emigration is induced by seasonal factors. Spent male parr tend not to participate in the autumn emigration. The emigration is considered to be principally an appetitive phenomenon which maximizes the number of precociously mature males attaining sexual contact with adults.     

Macroinvertebrate drift in a Rocky Mountain stream

An extensive series of drift collections from a Rocky Mountain stream was used to investigate quantitative patterns in the taxonomic composition of drift throughout spring, summer and fall for 1975–1978. Drift was estimated by drift rate, the number of organisms drifting past a point per 24 h; and by drift density, the numbers of organisms collected per 100 m³ of water sampled.Drift densities were up to ten times greater by night than by day, and 24 h drift densities for the total fauna approached 2000 per 100 m³ in June–July, declining to <500 by autumn. Ephemeroptera, and especially Baetis, dominated the drift. Drift rates were greatest in late spring, around 10⁶ per 24 h, which are among the highest values reported for small trout streams. Drift rates declined to <10⁵ during the summer, and shifts in the taxonomic composition are described.Multiple regression analysis of the relationship between drift rate and density, and the independent variables discharge, benthic density and temperature, showed that discharge typically was a significant predictor of 24 h drift rate, usually the best single predictor. In contrast, 24 h drift density most frequently was independent of discharge, indicating that this measure tends to correct for seasonal variation in discharge, as suggested in the literature. However, this was not invariably true. Drift density significantly correlated with benthic density in five of eight taxa inspected, thus seasonal declines in the benthos probably accounted for parallel declines in drift density.     

Brain aromatase, 5 alpha-reductase,and 5 beta-reductase change seasonally in wild male song sparrows: relationship to aggressive and sexual behavior

In many species, territoriality is expressed only during the breeding season, when plasma testosterone (T) is elevated. In contrast, in song sparrows (Melospiza melodia morphna), males are highly territorial during the breeding (spring) and nonbreeding (autumn) seasons, but not during molt (late summer). In autumn, plasma sex steroids are basal, and castration has no effect on aggression. However, inhibition of aromatase reduces nonbreeding aggression, suggesting that neural steroid metabolism may regulate aggressive behavior. In wild male song sparrows, we examined the neural distribution of aromatase mRNA and seasonal changes in the activities of aromatase, 5 alpha-, and 5 beta-reductase, enzymes that convert T to 17 beta-estradiol, 5 alpha-dihydrotestosterone (5 alpha-DHT, a potent androgen), or 5 beta-DHT (an inactive metabolite), respectively. Enzyme activities were measured in the diencephalon, ventromedial telencephalon (vmTEL, which includes avian amygdala), caudomedial neostriatum (NCM), and the hippocampus of birds captured during spring, molt, or autumn. Aromatase and 5 beta-reductase changed seasonally in a region-specific manner. Aromatase in the diencephalon was higher in spring than in molt and autumn, similar to seasonal changes in male sexual behavior. Aromatase activity in the vmTEL was high in both spring and autumn but significantly reduced at molt, similar to seasonal changes in aggression. 5 beta-Reductase was not elevated during molt, suggesting that low aggression during molt is not a result of increased inactivation of androgens. These data highlight the relevance of neural steroid metabolism to the expression of natural behaviors by free-living animals.     

The influence of temperature,sexual condition,and season on the metabolic rate of the lizardPsammodromus hispanicus

Summary Male and femalePsammodromus hispanicus from southern Europe were acclimated to four seasonal conditions of photoperiod and night time temperature. During the dark period, the lizards' body temperatures fell to ambient air temperature but during the light period the lizards were allowed to thermoregulate behaviourally and at such times the lizards' mean body temperature varied from 29.0°C to 32.6°C. The resting metabolic rate of these lizards was measured in 5°C steps from 5°C to 30°C or 35°C. Sexual condition had little effect on resting metabolic rate, but at low temperatures lizards acclimated to winter or spring seasonal conditions had lower resting metabolic rates than those acclimated to summer or autumn conditions. At temperatures above 20°C seasonal acclimation had no effect on resting metabolic rate. It is considered that the reduction in low temperature metabolic rate in spring and winter is induced by low night time temperatures and serves to conserve energy during those seasons when lizards must spend long periods at low temperature without being able to feed.     

Temporal Patterns and Environmental Correlates of Macroinvertebrate Communities in Temporary Streams

Temporary streams are characterised by short periods of seasonal or annual stream flow after which streams contract into waterholes or pools of varying hydrological connectivity and permanence. Although these streams are widespread globally, temporal variability of their ecology is understudied, and understanding the processes that structure community composition in these systems is vital for predicting and managing the consequences of anthropogenic impacts. We used multivariate and univariate approaches to investigate temporal variability in macroinvertebrate compositional data from 13 years of sampling across multiple sites from autumn and spring, in South Australia, the driest state in the driest inhabited continent in the world. We examined the potential of land-use, geographic and environmental variables to predict the temporal variability in macroinvertebrate assemblages, and also identified indicator taxa, that is, those highly correlated with the most significantly associated physical variables. Temporal trajectories of macroinvertebrate communities varied within site in both seasons and across years. A combination of land-use, geographic and environmental variables accounted for 24% of the variation in community structure in autumn and 27% in spring. In autumn, community composition among sites were more closely clustered together relative to spring suggesting that communities were more similar in autumn than in spring. In both seasons, community structure was most strongly correlated with conductivity and latitude, and community structure was more associated with cover by agriculture than urban land-use. Maintaining temporary streams will require improved catchment management aimed at sustaining seasonal flows and critical refuge habitats, while also limiting the damaging effects from increased agriculture and urban developments.     

Epidemiological studies on horses infected with nematodes of the family Trichonematidae (Witenberg, 1925)

Observations are reported on seasonal changes in the age structure of populations of nematodes of the family Trichonematidae recovered at regular intervals from the lumens of the large intestines of horses slaughtered in S.W. England. The results show that changes in the size of parasitic populations of Trichonema nassatum follow seasonal variations in the rate of infection, more individuals maturing during summer/autumn than during winter/spring because of proportional differences in the numbers of infective larvae ingested from the pasture. In contrast, larvae of T. longibursatum, T. catinatum and T. goldi ingested by grazing horses during summer accumulate in the gut wall rather than promptly returning to the lumen and developing to maturity. Their development is apparently inhibited until the following spring when 4th-stage larvae emerge en masse and quickly reach the adult stage. The relationship between fluctuations in the size of the adult populations of the 4 species studied and the characteristic seasonal variations in horse faecal egg counts is discussed.     

The significance of storms for the concentration and export of dissolved organic carbon from two Precambrian Shield catchments

Dissolved organic carbon (DOC) concentrations and DOC export arestudied during storms to examine the relationship between DOCconcentration and stream discharge and to assess the importance of stormson DOC export. Storms were monitored in seven subcatchments within twosmall watersheds (Harp 4--21 and Harp 3A) on the Precambrian Shield inCentral Ontario, Canada. Stream DOC concentrations increase during stormsby as much as 100% and 410% in Harp3A and Harp 4--21 respectively. The seasonal regression between DOC andstream discharge is significant in subcatchments without wetlands(r² > 0.7) but is not significant in thetwo subcatchments with small wetland areas (r² <0.06). On average, regressions based on weekly data yield accurate estimatesof DOC export but the variation in regressions among individual storms andthe small number of high DOC samples result in uncertainties of more than30% in DOC export. The period-weighted calculation ofDOC export from weekly data underestimates export by 14%and 22% in Harp 3A and Harp 4--21 respectively. Stormswere responsible for 57% to 68% of theDOC export in the autumn and 29% to 40%of the DOC export in the spring. A single large storm accounted for31% of the autumn DOC export in Harp 3A. The importanceof individual storms for DOC export and the variation in the relationshipbetween DOC and stream discharge among storms make it difficult to predictthe effects of climate change on DOC export and DOC concentrations.     

Nitrate retention in a sand plains stream and the importance of groundwater discharge

We measured net nitrate retention by mass balance in a 700-m upwelling reach of a third-order sand plains stream, Emmons Creek, from January 2007 to November 2008. Surface water and groundwater fluxes of nitrate were determined from continuous records of discharge and from nitrate concentrations based on weekly and biweekly sampling at three surface water stations and in 23 in-stream piezometers, respectively. Surface water nitrate concentration in Emmons Creek was relatively high (mean of 2.25 mg NO₃?CN l^?1) and exhibited strong seasonal variation. Net nitrate retention averaged 429 mg NO₃?CN m^?2 d^?1 and about 2% of nitrate inputs to the reach. Net nitrate retention was highest during the spring and autumn when groundwater discharge was elevated. Groundwater discharge explained 57?C65% of the variation in areal net nitrate retention. Specific discharge and groundwater nitrate concentration varied spatially. Weighting groundwater solute concentrations by specific discharge improved the water balance and resulted in higher estimates of nitrate retention. Our results suggest that groundwater inputs of nitrate can drive nitrate retention in streams with high groundwater discharge.     

Redundancy, synergism, and active inhibitory range of non-host volatiles in reducing pheromone attraction in European spruce bark beetle Ips typographus

Brandt's voles ( Microtus brandti ) are small native mammals that inhabit the grasslands of Inner Mongolia, China. The species is considered a pest, particularly during population outbreaks, which have increased in frequency since 1970 from 1 every 7 years to 1 every 3 to 5 years. Using historical records taken between 1948 and 1998, we found that there was significant correlation between years for which the monthly averages of the Southern Oscillation Index (SOI) were consistently high, and years in which outbreaks occurred. Also there was a tendency for outbreaks not to occur in years with precipitation above or below average. For some years at some sites, annual or biannual estimates are available for the density of Brandt's vole populations. We found that the seasonal rate of increase over the non-breeding season from autumn to spring was negatively correlated with the density in autumn. The rate of increase over the breeding season from spring to autumn was independent of the density in spring but instead, reflects the species' preference for habitat with short, sparse grass; populations do not persist in highly degraded grasslands or in tall, dense grassland. The link between outbreaks and climatic indices, and the numerical response of Brandt's voles to particular habitat characteristics, suggest that current grazing intensity tends to maintain grass at low height in years with average precipitation. The substantial increase in livestock numbers over the last 50 years appears to have increased the number of years when the balance of grazing and plant growth favours high rates of increase in Brandt's vole populations, and hence, there has been an increase in the frequency of outbreaks.     

Contrasting nutrient exports from a forested and an agricultural catchment in south-eastern Australia

Dissolved organic carbon (DOC) and total and inorganic nitrogen and phosphorus concentrations were determined over 3 years in headwater streams draining two adjacent catchments. The catchments are currently under different land use; pasture/grazing vs plantation forestry. The objectives of the work were to quantify C and nutrient export from these landuses and elucidate the factors regulating export. In both catchments, stream water dissolved inorganic nutrient concentrations exhibited strong seasonal variations. Concentrations were highest during runoff events in late summer and autumn and rapidly declined as discharge increased during winter and spring. The annual variation of stream water N and P concentrations indicated that these nutrients accumulated in the catchments during dry summer periods and were flushed to the streams during autumn storm events. By contrast, stream water DOC concentrations did not exhibit seasonal variation. Higher DOC and NO₃ ⁻ concentrations were observed in the stream of the forest catchment, reflecting greater input and subsequent breakdown of leaf-litter in the forest catchment. Annual export of DOC was lower from the forested catchment due to the reduced discharge from this catchment. In contrast however, annual export of nitrate was higher from the forest catchment suggesting that there was an additional NO₃ ⁻ source or reduction of a NO₃ ⁻ sink. We hypothesize that the denitrification capacity of the forested catchment has been significantly reduced as a consequence of increased evapotranspiration and subsequent decrease in streamflow and associated reduction in the near stream saturated area.     

The trans‐equatorial loop migration system of Eleonora’s falcon: differences in migration patterns between age classes,regions and seasons

Internal factors such as experience (e.g. age) and motivation for breeding, and external ones such as environmental conditions (e.g. meteorology and landscape characteristics) can promote differences in migratory behaviour and routes among seasons, regions and populations. Using satellite telemetry we investigated whether such differences occur and which factors promote them among migrating Eleonora’s falcons breeding in the Mediterranean area (Spain and Croatia) and wintering in Madagascar. We found that during autumn migration no age differences occur when crossing the Sahara desert, but in the remaining African regions, juveniles were more prone than adults to fly at a slower and more tortuous rate, as well as exhibiting longer stop‐overs, particularly in the Sahel region. Such differences might be promoted by a lower foraging and pre‐migratory fattening efficiency in juveniles. During spring, routes were significantly more eastern than during autumn, resulting in a loop migration occurring in all studied populations. This could be accounted by seasonal variation in the distribution of trophic resources. Our results show that Eleonora’s falcons integrate spatially seasonal changing resources on a continental scale throughout their annual cycle, changing their movement patterns in response to internal (age) and external (habitat) factors. This loop migration pattern may prove to be widespread among other Palearctic trans‐continental migratory bird species.     

Seasonal changes in semen quality and freezability in the Warmblood stallion

The objective of this study was to investigate seasonal changes in stallion semen quality and to determine the best time for semen cryopreservation. Experiments were performed using 10 Warmblood stallions from the National Stud Farm in Avenches (Switzerland). Ejaculates were collected and frozen every other week during 1 year from January to December 1999. Volume, concentration, and motility, and the number of morphologically normal sperm and sperm with major defects (abnormal heads, acrosome defects, nuclear vacuoles, proximal droplets, abnormal midpieces) were evaluated. For all frozen-thawed semen samples motility as well as viability (SYBR-14/PI) was tested, and the hypoosmotic swelling test (HOS) was performed. To analyze seasonal differences 4 periods of 3 months each were defined: autumn (September, October, November), winter (December, January, February), spring (March, April, May) and summer (June, July, August). During the 1 year experiment all semen quality parameters showed a clear seasonal pattern. The volume, total sperm count and motility in fresh semen were significantly higher (P<0.05) in summer than in winter, while sperm concentration was significantly lower in summer compared to the other seasons. Regarding morphology, normal sperm was significantly lower (P<0.05) in summer than at any other time of the year and higher values (P<0.05) were found for major defects in summer than in spring and autumn. In frozen-thawed semen motility was significantly (P<0.05) improved in autumn when compared to spring and summer. Viability was lowest in summer and differed significantly (P<0.05) from other seasons. The HOS test revealed significantly more (P<0.05) membrane damaged spermatozoa in winter than in spring, summer and autumn. Our results demonstrate that in our climatic conditions clear seasonal differences occur in semen quality of fresh and frozen-thawed semen and that cryopreservation of stallion semen should preferably be performed in autumn.     

Brain aromatase, 5α‐reductase,and 5β‐reductase change seasonally in wild male song sparrows: Relationship to aggressive and sexual behavior

In many species, territoriality is expressed only during the breeding season, when plasma testosterone (T) is elevated. In contrast, in song sparrows (Melospiza melodia morphna), males are highly territorial during the breeding (spring) and nonbreeding (autumn) seasons, but not during molt (late summer). In autumn, plasma sex steroids are basal, and castration has no effect on aggression. However, inhibition of aromatase reduces nonbreeding aggression, suggesting that neural steroid metabolism may regulate aggressive behavior. In wild male song sparrows, we examined the neural distribution of aromatase mRNA and seasonal changes in the activities of aromatase, 5α‐, and 5β‐reductase, enzymes that convert T to 17β‐estradiol, 5α‐dihydrotestosterone (5α‐DHT, a potent androgen), or 5β‐DHT (an inactive metabolite), respectively. Enzyme activities were measured in the diencephalon, ventromedial telencephalon (vmTEL, which includes avian amygdala), caudomedial neostriatum (NCM), and the hippocampus of birds captured during spring, molt, or autumn. Aromatase and 5β‐reductase changed seasonally in a region‐specific manner. Aromatase in the diencephalon was higher in spring than in molt and autumn, similar to seasonal changes in male sexual behavior. Aromatase activity in the vmTEL was high in both spring and autumn but significantly reduced at molt, similar to seasonal changes in aggression. 5β‐Reductase was not elevated during molt, suggesting that low aggression during molt is not a result of increased inactivation of androgens. These data highlight the relevance of neural steroid metabolism to the expression of natural behaviors by free‐living animals. © 2003 Wiley Periodicals, Inc. J Neurobiol 56: 209–221, 2003     

Reproductive biology of the endemic goby, Lentipes concolor, from Makamaka'ole Stream, Maui and Waikolu Stream, Moloka'i

Constant pressure in Hawai'i to use limited freshwater resources has resulted in increasing concern for the future of the native stream fauna. Hawaiian freshwater gobies have an amphidromous life cycle with a marine larva period and require streams which flow continuously to the ocean for the critical reproductive periods and during recruitment. As such, the stream fauna is particularly sensitive to any anthropogenic perturbations which disrupt the continuity of stream flows. The objective of this 2-year study was to compare the life cycles of the goby, Lentipes concolor, from a heavily diverted stream on Moloka'i and a relatively undisturbed stream on Maui. In Makamaka'ole Stream, Maui, the population of L. concolor was reproductively active all year with females potentially spawning 2–3 times annually. The timing of spawning did not occur consistently during the wet or dry season but coincided with high stream flow conditions regardless of time-of-year. In Waikolu Stream, Moloka'i, the reproductive pattern was more variable with the number of reproductively active females ranging from 0% to 100%. In general the number of eggs was greater and egg size smaller for female L. concolor in Waikolu Stream than in Makamaka'ole Stream. However, female reproductive condition of L. concolor from Maui was consistently higher than from fish on Moloka'i. Reproduction of L. concolor in Makamaka'ole Stream was correlated with the seasonal pattern of flow rates with peaks in female reproductive condition associated with periods of elevated discharge. No correlation between reproduction and discharge occurred in Waikolu Stream. There were considerable differences between the magnitude of discharge in the two streams. Waikolu Stream experienced prolonged periods of extremely low flows which have become common since the Moloka'i Irrigation System began diverting water from the stream in 1960. In Makamaka'ole Stream, L. concolor was capable of reproducing throughout the year and adjusting fecundity in response to stream flow conditions. In contrast, the population in Waikolu Stream appeared to have a ‘boom or bust’ reproductive pattern; the population had reduced or no reproduction when stream flow conditions reached extreme low levels, but the population succesfully reproduced during higher flow months. The diversion structure in Waikolu Stream has dampened the natural seasonal discharge cycle, exacerbated natural low flow conditions, and increased the likelihood of prolonged periods of extremely low flow. Stream management practices in the Hawaiian Islands must take into account the complex life cycles and sensitivity to variable stream flow conditions of the native fauna. This revised version was published online in July 2006 with corrections to the Cover Date.     

Seasonal patterns in feeding ecology of three small fishes in the Biandantang Lake, China

Feeding ecology of three small fish species, Hypseleotris swinhonis, Ctenogobius giurinus and Pseudorasbora parva was studied seasonally in the Biandantang Lake, a small, shallow lake in central China. Gut length, adjusted for total body length, was significantly higher in spring than in other seasons for all the three species. Seasonal changes in gut length were not associated with changes in food quality. Weight of fore-gut contents, adjusted for body weight, was significantly higher in winter and spring than in summer and autumn in H. swinhonis and C. giurinus , and significantly higher in autumn than in spring and summer for P. parva. Percentage of empty fore-guts was highest in summer and lowest in spring for H. swinhonis and C. giurinus , and highest in winter and lowest in autumn for P. parva. Diet of the three small fishes showed apparent seasonal changes, and these changes reflected partly the seasonal fluctuations of food resources in environment. Diet breadth was high in winter and low in autumn for H. swinhonis , high in winter and low in spring and summer for C. giurinus , and high in autumn and low in spring for P. parva. Diet overlaps between pairs of species were biologically significant in most cases, except between H. swinhonis and P. parva in summer and autumn and between C. giurinus and P. parva in autumn.