Dynamics of space use and male vigour amongst wood mice,Apodemus sylvaticus,in the cereal ecosystem

The spatial organisation of male and female wood mice,Apodemus sylvaticus, was investigated in a large-scale radio-tracking study on arable farmland near Oxford, United Kingdom, during the breeding season. Both sexes had significantly larger home ranges in the breeding season than at other times, and the breeding season home ranges of male (X = 1.44 ha) were significantly larger than those of females (X = 0.49 ha). Home range overlap was significantly greater between males, and between males and females, than it was between females. Overlap between males tended to be greatest in heavily utilised areas. Except during sexual consortship, there was minimal evidence of dynamic interaction among individuals. Home range sizes of breeding males varied widely, as did their body weights. There was no relationship between male body weight and home range size or any other movement parameter. However, males with the largest home ranges had the highest scores on all other movement parameters, indicating that they expended more energy in movement. These more vigorous males had access to the home ranges of more females than did males with small home ranges.     

Lifetime reproductive success in the spotted sandpiper (Actitis macularia) : sex differences and variance components

Summary Lifetime reproductive success (LRS) of male (N=103) and female (N= 66) spotted sandpipers (Actitis macularia) was studied for 13 years of a 17-year study at Little Pelican Island, Leech Lake, Minnesota. There was no sex difference in longevity, but females had significantly more mates, eggs, chicks, fledged young, and young returning in subsequent years than did males. Variance in LRS was partitioned into five life-history components: longevity (L), mates per year (M), eggs per mate (E), proportion eggs hatched (H), and proportion of chicks fledged (F). For both sexes, F accounted for the greatest proportion of variance in LRS (males, 43%; females, 47%), followed by L (males, 26%; females, 43%) and H (males, 21%; females, 28%). Positive covariance between H and F was consistent with predator-caused clutch and brood loss. Contrary to our expectations, males had a higher coefficient of variation in reproductive success than did females. This was because males were relatively more likely than females to produce no young.Offprint requests to: L.W. Oring at the current address     

Vertical distribution, life cycle, and developmental characteristics of the mesopelagic calanoid copepod Gaidius variabilis (Aetideidae) in the Oyashio region, western North Pacific Ocean

Vertical distribution, life cycle, and developmental characteristics of the mesopelagic copepod Gaidius variabilis Brodsky in the Oyashio region were investigated by combining analyses of field copepodite populations with laboratory-rearing data of egg hatching and naupliar development. Field samplings from five discrete depths between the surface and ≤2000 m were made approximately every month for 1 year. Most populations of G. variabilis occurred between 600 and 1000 m depth. A modest degree of reversed diel vertical migration behavior and some stage-specific depth-distribution patterns were noted. All copepodite stages were observed throughout the year, suggesting a year-round spawning of G. variabilis. From a prominent abundance peak of Copepodite Stage 1 (C1) seen in June to August, together with development times of eggs and nauplii obtained in laboratory-rearing experiments, the major spawning season was extrapolated to be April to June, the phytoplankton bloom season. Tracing the peak abundance of each copepodite stage (distinguishing males and females for C4 to C6), the generation times of males and females were deduced as 2 and 1 year, respectively. All between-stage increments in terms of wet-, dry-, and ash-free dry weights were greatest in C3/C4, and least in C5/C6 for both males and females. The increments in C3/C4 and C4/C5 were greater for males than for females, reflecting a longer stage duration of the males. These weights did not increase in C5/C6 males, possibly because feeding ceased in C6 males. These results for G. variabilis are compared with those for some mesopelagic copepods previously reported from other regions. Received: 25 October 1999 / Accepted: 20 March 2000     

Size-related reproductive biology of the infaunal bivalve<Emphasis Type="Italic"> Abra alba</Emphasis> (Bivalvia) in Kinsale Harbour (south coast of Ireland)

The reproductive behaviour of 248 specimens comprising four size classes of the infaunal bivalve Abra alba (Wood) was investigated between September 1994 and November 1995 in Kinsale Harbour. Small males returned a very small proportion of sexually mature and spawning animals. By contrast, no spawning females were found in this size class, suggesting that males develop and mature at a smaller size than females. Mean gonad maturity indices indicated that, except for the smallest size class, A. alba exhibits two spawning events in 1 year, i.e. a lesser spring and a major autumn spawning event. The spring event was well synchronised between males and females, while the autumn spawning was more broadly synchronised between the sexes. Only one spawning event was recorded for very large males and females, also occurring in spring. Spawning stages were recorded for both sexes over all size classes between February and November, indicating only a short period of sexual repose, if any. Oocyte diameters did not show a discrete size range from developing to spawning stages. All stages were seen to contain varying proportions of smaller, less mature oocytes, suggesting that large A. alba females do not develop and spawn en bloc, but release their gametes periodically. Maturity indices showed only a broad similarity with off-bottom temperatures.Communicated by O. Kinne, Oldendorf/Luhe     

Bigamy and mate choice in the biparental cichlid fish Cichlasoma nigrofasciatum

Summary Sixty percent of C. nigrofasciatum males showed bigamous behaviour when held in small, outdoor ponds at an adult sex ratio of 5 males to 10 females. No male held more than two pair bonds simultaneously. Bigamy had reproductive benefits; bigamous males, on average, fathered more broods that survived well into the free-swimming fry stage than did monogamous males. In ponds where adult size varied, no positive correlations were found, for either sex, between adult size and frequency of spawning or success of broodrearing, despite predictions that larger size would confer reproductive benefits on both sexes. A significant positive correlation was found between the sizes of mated individuals in the pond where both sexes varied in size. On theoretical grounds this relationship is likely to be more strongly influenced by female than by male choice of spawning partner.     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Cephalopod hatchling growth: the effects of initial size and seasonal temperatures

Temperature is known to have a strong influence on cephalopod growth during the early exponential growth phase. Most captive growth studies have used constant temperature regimes and assumed that populations are composed of identically sized individuals at hatching, overlooking the effects of seasonal temperature variation and individual hatchling size heterogeneity. This study investigated the relative roles of initial hatchling size and simulated natural seasonal temperature regimes on the growth of 64 captive Octopus pallidus over a 4-month period. Initial weights were recorded, and daily food consumption and fortnightly growth monitored. Two temperature treatments were applied replicating local seasonal water temperatures: spring/summer (14–18°C) and summer/autumn (18–14°C). Overall octopuses in the spring/summer treatment grew at a rate of 1.42% bwd⁻¹ (% body weight per day) compared to 1.72% bwd⁻¹ in the summer/autumn treatment. Initial size influenced growth rate in the summer/autumn treatment with smaller octopuses (<0.25 g) growing faster at 1.82% bwd⁻¹ compared to larger octopuses at 1.68% bwd⁻¹. This was opposite to individuals in the spring/summer treatment where smaller octopuses grew slower at 1.29% bwd⁻¹ compared to larger octopuses at 1.60% bwd⁻¹. Initial size influenced subsequent growth, however, this was dependent on feeding rate and appears to be secondary to the effects of temperature.     

Growth of Chaetodon larvatus (Chaetodontidae: Pisces) in the southern Red Sea

Growth and age of Chaetodon larvatus were studied using growth bands in otoliths and length-frequency analyses. Otoliths of 180 C. larvatus were extracted and measured. Polished sections of sagittae revealed alternating opaque and translucent bands corresponding with a seasonal growth pattern. Both mass and size of the otoliths continue to grow steadily throughout life. Length-at-age data revealed very fast growth during the first year. Growth proceeded at a decreasing rate during the second and the third year; fishes older than 3 years did not grow noticeably. No difference in growth patterns between males and females could be detected. The growth parameters obtained for the whole population are: the asymptotic length (L _∞)=10.64 cm, growth constant (K)=1.14 year⁻¹ and the theoretical age at length zero (t ₀)=−0.30 year. The maximum age recorded was 14 years. Length frequency data collected at a recruitment site confirmed the fast growth of juveniles.     

Mangroves as nursery sites: comparisons of the abundance and species composition of fish and crustaceans in mangroves and other nearshore habitats in tropical Australia

Daytime sampling of mangrove and seagrass (Halophila/Halodule community) habitats every 7 wk at Alligator Creek, Queensland, Australia, over a period of 13 mo (February 1985–February 1986) using two types of seine net, revealed distinct mangrove and seagrass fish and crustacean faunas. Total abundance of fish and relative abundance of small and large fish also varied between habitats and seasonally. Post-larval, juvenile and small adult fish captured with a small seine-net (3 mm mesh) were significantly more abundant (4 to 10 times) in the mangrove habitat throughout the 13 mo of sampling. Mangrove fish abundance showed significant seasonality, greatest catches being recorded in the warm, wet-season months of the year. Relative abundances of larger fish (captured in a seine net with 18 mm mesh) in the two habitats varied throughout the year, but did not show a seasonal pattern. At the same site, small crustaceans were significantly more abundant in the mangroves in all but one dryseason sample. Similar comparisons for three riverine sites, sampled less frequently, in the dry and wet seasons of 1985 and 1986, respectively, showed that in general mangrove habitats had significantly more fish per sample, although the relative abundance of fish in mangroves and other habitats changed with season. Crustacean catches showed a similar pattern, except that densities among sites changed with season. Fish and crustacean abundance in mangroves varied among sites, indicating that estuaries differ in their nursery-ground value. The juveniles of two commercially important penaeid prawn species (Penaeus merguiensis and Metapenaeus ensis) were amongst the top three species of crustaceans captured in the study, and both were significantly more abundant in the mangrove habitat. By contrast, mangroves could not be considered an important nursery for juveniles of commercially important fish species in northern Australia. However, based on comparisons of fish catches in other regions, the results of the present study indicate the importance of mangroves as nursery sites for commercially exploited fish stocks elsewhere in South-East Asia. Contribution No. 378 from the Australian Institute of Marine Science     

Polygyny and male parental care in Savannah sparrows: effects on female fitness

Summary To determine the effects of male mating status on female fitness, we compared the reproductive success, survival, and future fecundity of female Savannah sparrows (Passerculus sandwichensis) mated to monogamous vs. polygynous males in a 5-year study on Kent Island, New Brunswick, Canada. The proportion of males with more than one mate varied from 15 to 43% between years and sites. Polygynous and monogamous males fledged young of equal size in every year of the study. Females who shared paternal care with other females laid as many eggs per clutch and clutches per season as monogamously mated females. In most years polygynously mated females showed no delay in laying a second clutch, and they suffered no reduction in fecundity the following year. Recruitment of a female's offspring into the breeding population was generally independent of her mating status. Fitness costs of being mated to a polygynous male were only apparent in one year of the study, during which females mated to polygynous males had higher over-winter mortality than those mated to monogamous males. That same year, young raised by polygynous males were only one-third as likely to survive to reproductive maturity (as inferred by returns) as those raised by monogamous males. A male's mating status had no effect on his own survivorship. A male's mating status did not necessarily reflect his contributions to raising nestlings, which may partially explain why monogamously and polygynously mated females had equal fitness. At 35 nests the proportion of food deliveries brought by individual males varied from 0 to 75%; on average, males brought fewer than 30% of all food deliveries. Yet parental care by polygynous males was no less than that of monogamous males, at least at the nests of their primary females. Secondary females tended to receive less male assistance during the nestling stage, but their reproductive success was indistinguishable from that of primary females. Females feeding young without male assistance made as many food deliveries/h as did pairs in which males brought at least 30% of all food deliveries. Unassisted females did not suffer diminished fledging success or produce smaller fledglings. The benefits of polygyny for male Savannah sparrows are clear: polygynous males recruit more surviving offspring into the breeding population than monogamous males. The fitness of females, on the other hand, appears to be unaffected by whether their mate was monogamous or polygynous except in occasional years. Polygyny may be maintained in this population by the constraints of a female-biased sex ratio, the inability of females to predict a male's paternal care based on his morphology or behavior, the poor correlation between a male's mating status and his assistance at the nest, and inconsistent natural selection against mating with a polygynous male. Correspondence to: N.T. Wheelwright     

Tardy females,impatient males: protandry and divergent selection on arrival date in the two sexes of the barn swallow

Protandry reflects the earlier arrival of males than females to the site of reproduction. Such protandry is hypothesised to arise from sex differences in costs and benefits of early arrival. I investigated temporal patterns of arrival date of male and female barn swallows Hirundo rustica and temporal patterns of selection to test the hypothesis that sex differences in selection account for sex differences in arrival date. Mean arrival date of male barn swallows but not of females advanced during the last 33 years, giving rise to an increasing sex difference in arrival date. Early arrival was favoured by increasingly better survival in males, while females showed an opposite pattern that did not reach significance, although the effect differed between sexes. Early arrival increased fecundity in both sexes equally.The sex difference in viability selection in relation to arrival date changed from positive to negative as the degree of protandry increased in recent years, although there was no similar significant relationship for fecundity selection. Furthermore, sex differences in viability selection in a given year affected the degree of protandry in the following year through differential survival of certain phenotypes over others. Finally, temporal changes in sex difference in viability selection and protandry were related to an increase in the interval between first and second clutches, as the duration of the breeding season increased because of climatic amelioration. These findings suggest that arrival date is under divergent selection in the two sexes, providing a mechanism for the evolution of protandry.     

Macroalgae as habitat patch islands for Scutellidium lamellipes (Copepoda: Harpacticoida) and Ampithoe tea (Amphipoda: Gammaridae)

Scutellidium lamellipes and Ampithoe tea live in patches formed by the brown alga Pelvetia fastigiata. The distributions and abundance of both crustaceans were examined on P. fastigiata obtained during 1975 and 1976 from 7 collecting and 6 experimental sites in the mid-intertidal zone at La Jolla, California (USA). At each collecting site, the abundance of each crustacean generally increases with increasing plant size. Following defaunation and transplantation of algae, abundances of both crustaceans on individual plants appear to be governed by equilibria determined by in situ reproduction, immigration and emigration. Immigration rates of both animal species decrease significantly with increasing distance from areas occupied by P. fastigiata. During short-term experiments, neither crustacean was able to establish colonies on defaunated plants placed across a 30 m P. fastigiata-free zone and both species would have to reproduce to persitst on plants transplanted to that site. More frequent immigration and reproduction (and, for S. lamellipes at least, decreasing emigration rates with increasing patch size) would increase abundances and decrease probabilities of local extinction on larger and/or less distant plants. However, even within sites with isolated plants, the smallest plants often are not inhabited by either crustacean. The densitics of both species differ with plant size and distribution. Densities of S. lamellipes are greatest within a large aggregation of P. fastigiata, whereas those of A. tea are greatest on moderate-sized isolated plants at mid-tide levels.     

Biology and population dynamics of the intertidal isopod Cirolana harfordi

The distribution of Cirolana harfordi (Lockington) populations is determined largely by the availability of loose boulders on sandy beaches. The isopods swim out from under the rocks at high tide to feed. Their diet consists primarily of minute polychaetes and crustaceans. In addition, the isopods locate and utilize any available dead animal matter in the surf zone. Breeding occurs throughout most of the year, except for a brief lull in the fall. Females produce 1 or 2 broods of 18 to 68 young during their 2-year life-span. Marsupial incubation lasts 3 to 4 months. The population size and structure remained relatively constant from one year to the next. Mortality rates estimated for juveniles, males and females showed that newly emergent young and post-reproductive females suffer the greatest losses (up to 75% mortality per month).     

Lek structure and territoriality in the chryxus arctic butterfly,Oeneis chryxus (Satyridae)

Summary The territorial and mating systems of the satyrid butterfly Oeneis chryxus were studied for 3 years on individually marked populations in algonquin Provincial Park, Ontario. Males defended territories in a dry open forest clearing by means of chases and spiral flights with conspecifies. Defended areas did not contain food or oviposition sites required by females. Territories were not uniformly distributed over the study area; the highest density of territories was clustered over bare ground covered with sand and small stones rather than areas of vegetation. This pattern of distribution of territories occurred during each year of the study. The patterns of occupancy of territories showed a high degree of consistency from one day to the next, some males defending a given territory for as many as 11 consecutive days. Removal experiments showed that a surplus of non-territorial males did not exist. Vacant territories in the high-density area were absorbed by neighboring males; those in the low-density area remained empty. Male size was not correlated with territory position in the study are. Wing patterning (i.e. size of spots) differed between males occupying territories on sandy/stony versus vegetated areas. The territorial and mating systems of O. chryxus resemble lek polygyny, and are more similar to some vertebrate lek systems than to hilltopping or landmark territoriality.     

Social organization of woodchucks (Marmota monax)

Summary The social organization of woodchucks (Marmota monax) in southeast Ohio was studied at two sites, at one for two 2 years (1979–1980) and the other for 3 years (1981–1983). Spatial organization was determined by trapping and radio tracking. The home ranges of adult females did not overlap in the early spring but during late spring and summer there was some overlap (<10%) as females expanded their home ranges. Adult females tended to occupy the same home range in consecutive years. Some adult males occupied well-defined home ranges that did not overlap the home ranges of other males but did overlap extensively the home range of one to three adult females. These males tended to occupy the same home range in consecutive years. Infants used the same home range of their dam until about 2–3 months of age when most males and females apparently dispersed. About 35% of the juvenile females did not disperse until their second spring, just before their mother's new litter first emerged from their burrow. The average social group consisted of an adult male with two female kin groups comprising an adult female, an offspring (usually female) of the previous year, and the young of the year. Interactions within the kin group and with the adult male were relatively frequent and generally amicable. Interactions between kin groups both within and between different social groups were relatively rare and agonistic. The social organization of woodchucks in Ohio differs from that described in previous studies of woodchucks elsewhere and from that predicted by current models proposed by others on the evolution of social organization of marmots.     

Spatial and temporal variation in the size at maturity of the western rock lobster Panulirus cygnus George

The sizes at which female and male western rock lobster Panulirus cygnus become mature were examined over 32 years from records at six localities along the coast of Western Australia. The size of males at maturity was estimated from a subset of these data by a morphometric and a physiological method, with both producing statistically similar results. Males were larger at first maturity than females at the same sites. For both sexes, the sizes at first maturity at each location correlated (P<0.05) with the mean annual water temperature at that location, decreasing from south to north along the mainland coast and being smallest at the offshore Abrolhos Islands. Smaller sizes at maturity were recorded for both sexes than have been published previously. One certain explanation for these differences is that management measures protecting females with ovigerous setae have distorted size compositions and the ratio of immature to mature females, thereby increasing the likelihood of capturing small mature females. However, these fishing effects cannot fully account for the progressive decline in CL₅₀ observed over the past 20 years. Other possible hypotheses considered include increases in water temperature over this period, as well as whether this change could be consistent with a genotypic response caused by the selective removal of large lobsters combined with high exploitation rates.     

Distribution,growth and life cycle of the mesopelagic amphipod Primno abyssalis (Hyperiidea: Phrosinidae) in the southern Japan sea

The vertical distribution, growth, maturation, brood size and life cycle of the hyperiid amphipod Primno abyssalis (formerly P. macropa) were investigated using seasonal samples collected from Toyama Bay, southern Japan Sea, during the period June 1986 to September 1992. Over four different seasons of the year, P. abyssalis was most abundant in the 200 to 350 m strata at night and the 350 to 400 m strata during the day, indicating 100 to 150 m as the general distance of diel vertical movement. Some differences in vertical migrating behavior were noted among juveniles, adult males and females. Population-structure analysis revealed the occurrence of three cohorts aged 0+, 1+ and 2+ yr. Growth as body length in this species is linear with time. Estimated time to complete one life cycle is 1.8 to 2.5 yr for females, but only 0.8 yr for males. Maximum longevity is 2.8 yr. Instar analysis based on the segment number of pleopod rami indicated that newly hatched juveniles molt ten times to reach adult male, and four more times to reach adult female. Adult instar number was found to be only 1 for males and 5 for females. Ovigerous females occurred throughout the year, but the annual peak of release of juveniles from the female's marsupium is estimated to be arly March. Brood size was not correlated with female size, a maximum brood size of 214 eggs was recorded. The dry and ash-free dry weights of instars suggested that juveniles in the female marsupium, adult males, and older adult females are less active feeding or non-feeding stages. Except for the reduced growth rate and the occurrence of small, short-lived males, most characteristics of P. abyssalis are consistent with the present view of the life modes of mesopelagic animals, including linear growth in length, aseasonal reproduction, and smaller brood size coupled with larger eggs.     

Sex ratio manipulation and decreased growth of male offspring of undernourished golden hamsters (Mesocricetus auratus)

Summary The theory that female mammals in poor condition may increase individual fitness by skewing the sex ratio of their offspring toward daughters and by investing more resources in daughters than in sons was tested in hamsters. Newly mated experimental females were food-restricted during pregnancy and lactation (RR) or during lactation only (AR). Controls received food ad libitum. Maternal body weights were assessed daily from mating to 25 days postpartum. Litter survival (% litters with at least one pup surviving on any day), litter size, offspring sex ratios (=% males), and pup weights were monitored daily from birth (Day 1) to Day 25. All control and AR dams gave birth 16 days after mating. Gestation was extended by 1–3 days for 35.4% of RR dams. RR dams weighed significantly less at parturition than controls and AR females. During lactation, AR females showed the greatest weight loss and control females the least. AR weight loss exceeded that of RR females, possibly because the former maintained larger litters. Survival was highest for controls, intermediate for AR, and lowest for RR litters. Mean sex ratio at birth was significantly less for RR (40.7%) than for control (49.6%) and AR (48.8%) litters. RR sex ratio did not change significantly postnatally. Sex ratios of control and AR litters never differed statistically from 50%. Control male pups were significantly heavier than their sisters throughout the experiment. No significant gender differences were observed for AR pup weights after Day 2 postpartum. RR female offspring weighed more than their brothers throughout the experiment, and this difference was statistically significant immediately prior to the time that pups began to feed independently (Days 14–17). RR female pup weights were similar to, and sometimes significantly greater than, weights of control daughters during the period of postnatal maternal investment. Control males were always heavier than males from the other treatments. Patterns of weight gain by AR and RR males varied with age. We conclude that underfed female hamsters are able to adjust the sex ratio of offspring prenatally and parental investment postnatally to favor daughters.     

Reproduction of a Northumberland population of the polychaete Cirratulus cirratus

A population of Cirratulus cirratus (O. F. Müller) was sampled from September 1966 to September 1968. The population is found intertidally in crevices at Cullercoats Bay, Northumberland (UK). Measurements of the sexual maturity and size of the animals have been made. The breeding season is not restricted, spawning occurs in the population asynchronously throughout the year. Ripe worms and worms that have just spawned are invariably present, but the majority of females are always at intermediate stages of oogenesis. Adult males, however, always contain active spermatozoa. Females, but not males, are devoid of coelomic gametes after spawning; a new generation of coelomic oocytes begin to accumulate in the coelom after 30 days. The growth of the coelomic oocytes is relatively slow, the coelomic phase of oocyte development takes between 1 and 2 years for completion. The coelomoducts which serve for the emission of the gametes are described. An interpretation of the female population structure based upon length and sexual maturity data suggests that most females survive to spawn 2 or 3 times, and some to spawn 4 or 5 times. They grow in length by about 2 cm in the period between successive spawnings. The reproduction of Cirratulus cirratus is compared with that of other annelid species breeding throughout the year, and this unusual mode of reproduction is discussed.     

Diet and respiration of the small planktonic marine copepod Oithona nana

The functional responses of Oithona nana (Giesbr.) to various phytoplankton and zooplankton food species are described. The food species were divided into three size categories, the seasonal abundances of which were measured in Loch Turnaig, a Scottish sea loch in 1977. The seasonal variations in feeding rates in the sea for each size class were derived. The seasonal variation in respiration rate of O. nana was measured, and metabolic requirements were claculated as between 6 and 40% of the food material estimated as being eaten. O. nana differs from other common copepods in having a wide food-particle size spectrum and a low metabolic rate. It is suggested that these adaptations constitute the strategy whereby O. nana maintains its population levels throughout the year.